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https://openalex.org/W2150088027
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https://dash.harvard.edu/bitstream/1/4553340/1/2857650.pdf
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English
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Alternative Strategies to Reduce Maternal Mortality in India: A Cost-Effectiveness Analysis
|
PLoS medicine
| 2,010
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cc-by
| 15,907
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Terms of Use This article was downloaded from Harvard University’s DASH repository, and is made available
under the terms and conditions applicable to Other Posted Material, as set forth at http://
nrs.harvard.edu/urn-3:HUL.InstRepos:dash.current.terms-of-use#LAA Permanent link http://nrs.harvard.edu/urn-3:HUL.InstRepos:4553340 Published Version
doi:10.1371/journal.pmed.1000264 Published Version
doi:10.1371/journal.pmed.1000264 Abstract Background: Approximately one-quarter of all pregnancy- and delivery-related maternal deaths worldwide occur in India. Taking into account the costs, feasibility, and operational complexity of alternative interventions, we estimate the clinical
and population-level benefits associated with strategies to improve the safety of pregnancy and childbirth in India. Methods and Findings: Country- and region-specific data were synthesized using a computer-based model that simulates
the natural history of pregnancy (both planned and unintended) and pregnancy- and childbirth-associated complications in
individual women; and considers delivery location, attendant, and facility level. Model outcomes included clinical events,
population measures, costs, and cost-effectiveness ratios. Separate models were adapted to urban and rural India using
survey-based data (e.g., unmet need for birth spacing/limiting, facility births, skilled birth attendants). Model validation
compared projected maternal indicators with empiric data. Strategies consisted of improving coverage of effective
interventions that could be provided individually or packaged as integrated services, could reduce the incidence of a
complication or its case fatality rate, and could include improved logistics such as reliable transport to an appropriate
referral facility as well as recognition of referral need and quality of care. Increasing family planning was the most effective
individual intervention to reduce pregnancy-related mortality. If over the next 5 y the unmet need for spacing and limiting
births was met, more than 150,000 maternal deaths would be prevented; more than US$1 billion saved; and at least one of
every two abortion-related deaths averted. Still, reductions in maternal mortality reached a threshold (,23%–35%) without
including strategies that ensured reliable access to intrapartum and emergency obstetrical care (EmOC). An integrated and
stepwise approach was identified that would ultimately prevent four of five maternal deaths; this approach coupled
stepwise improvements in family planning and safe abortion with consecutively implemented strategies that incrementally
increased skilled attendants, improved antenatal/postpartum care, shifted births away from home, and improved
recognition of referral need, transport, and availability/quality of EmOC. The strategies in this approach ranged from being
cost-saving to having incremental cost-effectiveness ratios less than US$500 per year of life saved (YLS), well below India’s
per capita gross domestic product (GDP), a common benchmark for cost-effectiveness. Share Your Story The Harvard community has made this article openly available. Please share how this access benefits you. Submit a story . Accessibility eve Sweet1, Natalie Carvalho1, Uma Chandra Mouli Natchu2,4, Delphine Hu1,3 1 Center for Health Decision Science, Harvard School of Public Health, Boston, Massachusetts, United States of America, 2 Department of Nutrition, Harvard School of
Public Health, Boston, Massachusetts, United States of America, 3 Department of Health Policy and Management, Harvard School of Public Health, Boston, Massachusetts,
United States of America, 4 Pediatric Biology Centre, Translational Health Science and Technology Institute, Delhi, India Citation: Goldie SJ, Sweet S, Carvalho N, Natchu UCM, Hu D (2010) Alternative Strategies to Reduce Maternal Mortality in India: A Cost-Effectiveness Analysis. PLoS
Med 7(4): e1000264. doi:10.1371/journal.pmed.1000264
Academic Editor: Zulfiqar A. Bhutta, Aga Khan University, Pakistan
Received September 14, 2009; Accepted March 12, 2010; Published April 20, 2010
Copyright: 2010 Goldie et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: John D. and Catherine T. MacArthur Foundation #07-89007-0 00GSS. The funder had no role in study design, data collection and analysis, decision to
publish, or preparation of the manuscript.
Competing Interests: The authors have declared that no competing interests exist.
Abbreviations: bEmOC, basic emergency obstetrical care; cEmOC, comprehensive emergency obstetrical care; EmOC, emergency obstetrical care; GDP, gross
domestic product; MDG, Millennium Development Goal; MMR, maternal mortality ratio; PPH, postpartum hemorrhage; SBA, skilled birth attendant; TFR, total
fertility rate; YLS, year of life saved
* E-mail: sue_goldie@harvard.edu Sue J. Goldie1,3*, Steve Sweet1, Natalie Carvalho1, Uma Chandra Mouli Natchu2,4, D Sue J. Goldie1,3*, Steve Sweet1, Natalie Carvalho1, Uma Chandra Mouli Natchu2,4, Delphine Hu1,3 Introduction relevant comorbidities, aggregates individual outcomes to the
population
level,
and
reflects
setting-specific
epidemiology. Separate models were adapted to urban and rural India using
data on antenatal care, family planning, facility births, and skilled
birth attendants (SBAs), and information about access to transport,
referral facilities, and quality of care. Model outcomes include
clinical events (e.g., pregnancies, live births, maternal complica-
tions), measures of maternal mortality (e.g., MMR, proportionate
mortality ratio [i.e., proportion of deaths that are pregnancy-
related among women aged 15–45 y], and lifetime risk of maternal
death), population outcomes (e.g., life expectancy), and economic
costs. Approximately one-quarter of all pregnancy- and delivery-related
maternal deaths worldwide occur in India, which has the highest
burden of maternal mortality for any single country [1,2]. Although the
inclusion of maternal mortality reduction in the United Nations’
Millennium Development Goals (MDGs) reflects the importance of
improving maternal health as a key mechanism in reducing poverty
and promoting social and economic growth, global progress has been
suboptimal [2–4]. Several factors may be changing the landscape for
maternal health in India in particular [5]. These factors include more
information on maternal mortality measures [6,7], an increasing
number of studies evaluating interventions [8], renewed determination
on the part of the maternal health and public health communities [9],
and, most importantly, the emergence of maternal mortality reduction
as a clear priority on the Indian national political agenda [5,10–12]. We evaluated alternative approaches to reducing maternal
mortality in settings in India that differ according to underlying
maternal risk, health, and socioeconomic status. Interventions can
be provided individually or packaged into integrated services. Following standard recommendations for economic evaluation
[25], strategies are first ranked in terms of increasing costs and
benefits; those that are less effective and more costly than an
alternative strategy are considered inefficient, and those that cost
less than the status quo are considered ‘‘cost saving.’’ For all other
strategies, we calculate an incremental cost-effectiveness ratio,
defined as the additional cost of a specific strategy divided by its
additional clinical benefit, compared with the next least expensive
strategy. We considered interventions with cost-effectiveness ratios
of less than the per capita gross domestic product (GDP)
(US$1,068)
to
be
very
cost-effective
as
suggested
by
the
Commission on Macroeconomics and Health. Sensitivity analyses
are conducted to assess the impact of parameter uncertainty. The Model The computer-based Global Maternal Health Policy Model
simulates the natural history of pregnancy (both planned and
unintended) and pregnancy- and childbirth-associated complica-
tions (Figure 1). This model defines health states to reflect
important characteristics that affect prognosis, quality of life, and
resource use. The time horizon incorporates a woman’s lifetime
and is divided into equal time increments during which women
transition from one health state to another. Nonpregnant girls
enter the model and in each time period may become pregnant
depending on age, use of contraception, and clinical history
(Figure 1, upper panel). Once pregnant, women have a chance of
spontaneous abortion (i.e., miscarriage), induced abortion, or
continued pregnancy. A proportion of induced abortions will be
unsafe (i.e., surgical or medical abortion conducted by untrained
personnel). Labor and delivery may be associated with a direct
complication
of
pregnancy
(e.g.,
hypertensive
disorders
of
pregnancy, obstructed labor, hemorrhage, sepsis). Case fatality
rates are conditional on the type and severity of complication (e.g.,
moderate sepsis requiring antibiotics versus severe hemorrhage
requiring blood transfusion) and underlying comorbidity (e.g.,
anemia). Nonfatal complications include neurological sequelae,
rectovaginal fistula, severe anemia, and infertility (Figure 1, upper
panel). In addition to death from maternal complications, women
face an annual risk of death from age-specific all-cause mortality. Previous model-based studies have provided important insights into
the potential high public health value of reducing maternal deaths,
however, many of these have not considered the full range of
interventions to reduce maternal mortality, such as family planning,
safe abortion, and intrapartum care [7,17,18]. Some have only focused
on single interventions [19–21], others have not included costs [22],
and recent analyses that did assess multiple strategies did not explicitly
model critical barriers to life-saving referral, such as recognition of
referral need and accessible transport [23,24]. Taking into account the
costs, feasibility, and operational complexity of alternative interven-
tions, we extend this body of work to estimate the clinical and
population-level benefits associated with a comprehensive set of
strategies to improve the safety of pregnancy and childbirth in India. Strategies in the model to reduce maternal mortality consist of
improving coverage of effective interventions, which may be
provided individually or packaged as integrated services. Introduction Despite consensus on the need for universal access to high-quality
intrapartum and emergency obstetrical care (EmOC), uncertainties
remain about how to adapt ‘‘ideal recommendations’’ to specific
situations [8,13]. The need for an adequate supply of skilled providers,
functional referral and transport, and well-equipped facilities for
EmOC will prove a formidable barrier in the near-term for countries
with weak health systems, as well as for states with inadequate health
delivery infrastructure and for communities in predominantly rural
areas [13]. This challenge will be particularly relevant for India, with its
largely rural population, and striking disparities between states. For
example, while Kerala reports a maternal mortality ratio (MMR) of
fewer than 100 maternal deaths per 100,000 live births, rural Uttar
Pradesh and Rajasthan report MMRs of more than 400 [14,15]. To most effectively leverage India’s national commitment to
reducing maternal mortality, identifying evidence-based strategies
that consider the local context is imperative. Our analysis is
motivated by questions that include: What are the fundamental
drivers of the effectiveness, cost-effectiveness, and affordability of a
package of interventions to reduce maternal mortality? Because
adequate facilities, health infrastructure, and skilled human
resources will not be readily available in all settings, can we provide
interim guidance to policy makers? While no single empirical study
can provide clear answers to these questions, a modeling approach
within a decision-analytic framework can extend empiric informa-
tion by extrapolating outcomes beyond the time horizon of a single
study, can facilitate synthesis of multiple data sources in an
internally consistent and epidemiologically plausible way [16], and
may be adapted to specific settings so that existing infrastructure,
resources, and political realities can be considered. Abstract Conclusions: Early intensive efforts to improve family planning and control of fertility choices and to provide safe abortion,
accompanied by a paced systematic and stepwise effort to scale up capacity for integrated maternal health services over
several years, is as cost-effective as childhood immunization or treatment of malaria, tuberculosis, or HIV. In just 5 y, more
than 150,000 maternal deaths would be averted through increasing contraception rates to meet women’s needs for spacing
and limiting births; nearly US$1.5 billion would be saved by coupling safe abortion to aggressive family planning efforts; and
with stepwise investments to improve access to pregnancy-related health services and to high-quality facility-based
intrapartum care, more than 75% of maternal deaths could be prevented. If accomplished over the next decade, the lives of
more than one million women would be saved. Please see later in the article for the Editors’ Summary. Citation: Goldie SJ, Sweet S, Carvalho N, Natchu UCM, Hu D (2010) Alternative Strategies to Reduce Maternal Mortality in India: A Cost-Effectiveness Analysis. PLoS
Med 7(4): e1000264. doi:10.1371/journal.pmed.1000264
Academic Editor: Zulfiqar A. Bhutta, Aga Khan University, Pakistan
Received September 14, 2009; Accepted March 12, 2010; Published April 20, 2010 Copyright: 2010 Goldie et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: John D. and Catherine T. MacArthur Foundation #07-89007-0 00GSS. The funder had no role in study design, data collection and analysis, decision to
publish, or preparation of the manuscript. Abbreviations: bEmOC, basic emergency obstetrical care; cEmOC, comprehensive emergency obstetrical care; EmOC, emergency obstetrical care; GDP, gross
domestic product; MDG, Millennium Development Goal; MMR, maternal mortality ratio; PPH, postpartum hemorrhage; SBA, skilled birth attendant; TFR, total
fertility rate; YLS, year of life saved * E-mail: sue_goldie@harvard.edu PLoS Medicine | www.plosmedicine.org 1 April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 PLoS Medicine | www.plosmedicine.org Reducing Maternal Mortality in India The Model In
addition to family planning, antenatal care (i.e., prenatal care)
and treatment of anemia, safe abortion, and postpartum care, the
model includes both intrapartum interventions that reduce the
incidence of a complication (e.g., misoprostol for postpartum
hemorrhage [PPH], clean delivery for sepsis), as well as those that PLoS Medicine | www.plosmedicine.org Analytic Overview The effectiveness of
interventions to reduce the incidence of complications (e.g.,
active management of labor) was estimated from published
studies using methods detailed in Text S1. The effectiveness of
interventions to reduce case fatality rates was from published
studies and assumed treatment in an appropriate facility; a
wide plausible range was used for sensitivity analyses (Table 1;
Text S1). Data on facility births, SBAs, family planning for
spacing or limiting births, and antenatal care were from country-
specific surveys [28]. The effectiveness of interventions to either reduce the incidence
of complications or to reduce case fatality rates associated with
complications depends, in part, on access to specific services (e.g.,
trained SBA) and to specific levels of facilities (e.g., comprehensive
EmOC [cEmOC] with capacity for blood transfusion). Accord-
ingly, the ultimate impact of interventions depends on several
setting-specific factors. These include delivery site, presence of
birth attendant, quality and type of referral facility, as well as
successful referral when necessary. The model therefore explicitly
considers the location of delivery, type of assistance, access to basic
or comprehensive obstetrical care, and the ability to overcome a
series of barriers around the timing of delivery (e.g., recognition of
referral need, reliable transport, timely treatment at an appropri-
ate referral facility); these factors collectively determine the health
services a woman can access and the specific interventions that
would be included (Figure 1, lower panel). For women delivering at home or in a birthing center, the
probability of successful referral depended on overcoming three
potential barriers [74]: (1) delay in recognizing referral need; (2)
delay in transfer to referral facility (means of transport and interim
care en route); (3) delay in receiving appropriate care at the
appropriate EmOC facility such as inadequate staffing and
supplies, inexpedient attention (e.g., delay to collect fees), and/or
non-evidence-based or substandard care. Assumptions about the
latter two delays were based on survey data (e.g., National Family
Health Survey [NFHS-3] [28,75], District Level Household
Survey [DLHS] [76], and Facility Survey [26]), state-level facility
surveys [77,78], government reports [27], and published studies
(Text S1) [14,15,79–83]. (
g
p
)
Delivery setting is differentiated by provider (e.g., family
member, traditional birth attendant [TBA], or SBA) and by site
(e.g., home versus facility). Analytic Overview The best available data were synthesized using a computer-
based model that simulates the natural history of pregnancy and April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 2 Reducing Maternal Mortality in India Figure 1. Schematic of the model. Upper panel: Model simulates the natural history of pregnancy (both planned and unintende
pregnancy- and childbirth-associated complications. Case fatality rates for complications depend on severity and comorbidity. General interv
categories (open red boxes) include family planning for spacing or limiting births, antenatal or prenatal care (and treatment of anemia), safe ab
Reducing Maternal Mortality i Figure 1. Schematic of the model. Upper panel: Model simulates the natural history of pregnancy (both planned and unintended) and
pregnancy- and childbirth-associated complications. Case fatality rates for complications depend on severity and comorbidity. General intervention
categories (open red boxes) include family planning for spacing or limiting births, antenatal or prenatal care (and treatment of anemia), safe abortion,
intrapartum care (e.g., active management of labor), basic and comprehensive EmOC, and postpartum care. Interventions can reduce the incidence or
severity of a complication or can reduce the case fatality rate through appropriate treatment. Lower panel: Model reflects the intervention pathway
during labor and delivery, including location (home, birthing or health center, bEmOC, cEmOC), attendant (family member, traditional birth attendant
[TBA], or SBA), and three potential barriers to effective treatment in the event of a complication, including recognition of referral need, transfer (e.g., PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 3 Reducing Maternal Mortality in India Reducing Maternal Mortality in India transport), and timely quality care in an appropriate EmOC facility. Management of labor and delivery depends on attendant (e.g., SBA, clean delivery)
and site (e.g., expectant management in birthing center, active management in EmOC facility), as does access to specific levels of treatment (e.g.,
blood transfusion only available in cEmOC). doi:10.1371/journal.pmed.1000264.g001 reduce the case fatality rate through appropriate management in a
referral facility (Figure 1, upper panel). tions were obtained from published data, and a plausible range
for sensitivity analysis was based on review of the literature. Case
fatality rates were adjusted based on complication severity (e.g.,
life-threatening complications requiring cEmOC) and underlying
severity of anemia (Table 1; Text S1). Analytic Overview Facility levels are categorized as (1)
birthing centers or health centers, which cannot provide all
services necessary to qualify as a basic emergency obstetrical care
(bEmOC) facility, but are staffed with SBA who provide expectant
management of labor and more reliable referral when necessary
than with delivery at home; (2) facilities with bEmOC capacity
(e.g., first referral units); and (3) facilities with cEmOC capacity
(e.g., district hospitals) [26,27]. Facilities capable of bEmOC are
assumed to be capable of administering injectable antibiotics,
oxytocics, and sedatives or anticonvulsants, and performing
manual removal of placenta, removal of retained products, and
assisted vaginal delivery. Facilities capable of cEmOC also are able
to provide blood transfusion, cesarean section, and management of
advanced shock. Model performance was assessed by comparing the distribution
of direct causes of maternal mortality, life expectancy, propor-
tionate mortality ratio, MMR, and total fertility rate (TFR) to
empiric data [2,14,15,28,29,32,38,84–87] in rural and urban
India. Model validation was assessed by using state-specific data
from Rajasthan and Uttar Pradesh as model inputs, and
comparing model-projected indicators of maternal mortality with
survey-reported outcomes [14,15]. In addition, a secondary
analysis assessing all strategies evaluated in the base case was
conducted in Uttar Pradesh. Details of this process are included in
Text S1. This model also allows us to evaluate phased approaches that
involve scaling up access to services over time; the stepwise
investments in infrastructure required to assure high-quality
intrapartum care are designated as ‘‘upgrades.’’ In addition to
reducing unmet need for family planning and unsafe abortion,
four consecutively implemented strategies increased skilled atten-
dants, improved antenatal/postpartum care, incrementally shifted
births away from home, and improved the availability and quality
of EmOC. For women delivering at home or in birthing centers,
these ‘‘upgrades’’ also improved recognition of referral need,
access to transport, and expedient referral to an appropriate
facility (Figure 2). PLoS Medicine | www.plosmedicine.org Costs Selected costs used in the model are provided in Table 3
[18,25,80,88–94]. Details are provided in Text S1. With the
exception of facility costs, salaries, and transport costs, resource
requirements to deliver interventions and the costs of maternal
complications were estimated from the United Nations Population
Fund’s (UNFPA) Reproductive Health Costing Tools Model
(RHCTM) [88]. The RHCTM uses an ingredients approach to
estimate direct costs (including drugs, supplies, and personnel
requirements) of 45 reproductive health interventions, as well as
investments required for scale-up. We obtained personnel costs
(salaries) and facility costs from public access country-specific
databases [25,89], and drugs and supply costs from the UNICEF
Supply Catalogue and Management Sciences for Health (MSH)
International Drug Price Indicator Guide [90,91]. We leveraged
public access sources and published studies to inform assumptions
about the financial requirements for improving transport and
scaling up facility- and human-resource capacity [7,30,80,92,93]. We assessed the face validity of model input values and established
a plausible range for sensitivity analysis by comparing estimates to All models were built using TreeAge Pro 2008 (TreeAge
Software Inc.) and analyzed using IBM/Lenovo Dual-Core VT
Pro Desktop computers running Microsoft Windows XP, using
Microsoft Excel 2007 and Visual Basic for Applications 6.5
(Microsoft Corp.). We used Monte Carlo simulation to generate
the number of per woman events such as pregnancies, live births,
facility-based births, and maternal complications. Model Validation Model-estimated life expectancy for a 15-y-old female was 55 y
compared to the World Health Organization’s (WHO) estimate of
55.1 y [85]. The distribution of maternal deaths by cause closely
approximated published regional estimates (Table 1) [29]. Model-
generated estimates of TFR and MMR for India, stratified for
rural and urban status, closely approximated survey-reported
values [1,2,28,38], as did the state-specific models for Rajasthan
and Uttar Pradesh (Table 2) [14,15]. Model predicted deaths for
2005, taking into account direct and indirect causes of maternal
mortality, were 117,657, compared to 117,000 estimated by
UNICEF, WHO, and UNFPA [86,87]. Increased family planning to reduce the unmet need also reduced
the number of deaths attributable to unsafe abortion (Figure 3). For
example, in rural India increasing contraceptive rates to 67.6% cut
abortion-related deaths by more than 50%—even with no change in
rates of unsafe abortion. Adding improved access to safe abortion and
postabortion care for three out of four women pursuing elective
termination of pregnancy prevented an additional 22% to 50% of
abortion-related deaths, depending on the underlying level of unmet
need; similarly, the additional cost savings ranged from 22% more, to
more than double the savings expected from family planning alone. Data Alternatives evaluated in sensitivity analysis (Results and Text S1). doi:10 1371/journal pmed 1000264 g002 Figure 2. Stepwise improvements in scaling up maternal services. Four strategies that scale up access to critical maternal health services in
consecutive phases are designated as upgrade 1, upgrade 2, upgrade 3, and upgrade 4. Shown are the percent increases in facility-based delivery,
SBAs, recognition of referral need (by SBA at birthing/health center), transport (to appropriate referral facility), and availability/quality of EmOC
(including adequate staff/supplies, appropriate clinical treatment, immediate attention), for rural and urban India. Shifts from home births assume a
70% shift to health centers/birthing centers and a 30% shift to EmOC; for routine births in EmOC, we assume 90% bEmOC and 10% cEmOC. Alternatives evaluated in sensitivity analysis (Results and Text S1). doi:10 1371/journal pmed 1000264 g002 Figure 2. Stepwise improvements in scaling up maternal services. Four strategies that scale up access to critical maternal health services in
consecutive phases are designated as upgrade 1, upgrade 2, upgrade 3, and upgrade 4. Shown are the percent increases in facility-based delivery,
SBAs, recognition of referral need (by SBA at birthing/health center), transport (to appropriate referral facility), and availability/quality of EmOC
(including adequate staff/supplies, appropriate clinical treatment, immediate attention), for rural and urban India. Shifts from home births assume a
70% shift to health centers/birthing centers and a 30% shift to EmOC; for routine births in EmOC, we assume 90% bEmOC and 10% cEmOC. Alternatives evaluated in sensitivity analysis (Results and Text S1). doi:10.1371/journal.pmed.1000264.g002 the cost savings for a single birth cohort of 15-y-old girls (2010)
that would be expected to accrue over their reproductive lifespans
ranged from US$111.4 million to US$448.2 million. Reducing the
unmet need, coupled with provision of safe abortion, provided
synergistic benefits and saved additional costs (Table 4). Results
were similar in urban India, although the amount of deaths
averted and costs saved were smaller, reflecting both the lower
initial TFR and the smaller population size (Table 4). those in published studies (Text S1). All costs were converted to
2006 US$. PLoS Medicine | www.plosmedicine.org Data Selected parameters and assumptions used in the model are
provided in Tables 1 and 2 [7,14,15,17,22,23,28–73]. Additional
details are provided in Text S1. Initial estimates of incidence and
case fatality rates associated with pregnancy-related complica- April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 4 Reducing Maternal Mortality in India Figure 2. Stepwise improvements in scaling up maternal services. Four strategies that scale up access to critical maternal health services in
consecutive phases are designated as upgrade 1, upgrade 2, upgrade 3, and upgrade 4. Shown are the percent increases in facility-based delivery,
SBAs, recognition of referral need (by SBA at birthing/health center), transport (to appropriate referral facility), and availability/quality of EmOC
(including adequate staff/supplies, appropriate clinical treatment, immediate attention), for rural and urban India. Shifts from home births assume a
70% shift to health centers/birthing centers and a 30% shift to EmOC; for routine births in EmOC, we assume 90% bEmOC and 10% cEmOC. Alternatives evaluated in sensitivity analysis (Results and Text S1). doi:10.1371/journal.pmed.1000264.g002 Figure 2. Stepwise improvements in scaling up maternal services. Four strategies that scale up access to critical maternal health services in
consecutive phases are designated as upgrade 1, upgrade 2, upgrade 3, and upgrade 4. Shown are the percent increases in facility-based delivery,
SBAs, recognition of referral need (by SBA at birthing/health center), transport (to appropriate referral facility), and availability/quality of EmOC
(including adequate staff/supplies, appropriate clinical treatment, immediate attention), for rural and urban India. Shifts from home births assume a
70% shift to health centers/birthing centers and a 30% shift to EmOC; for routine births in EmOC, we assume 90% bEmOC and 10% cEmOC. Alternatives evaluated in sensitivity analysis (Results and Text S1). doi:10.1371/journal.pmed.1000264.g002 Figure 2. Stepwise improvements in scaling up maternal services. Four strategies that scale up access to critical maternal health services in
consecutive phases are designated as upgrade 1, upgrade 2, upgrade 3, and upgrade 4. Shown are the percent increases in facility-based delivery,
SBAs, recognition of referral need (by SBA at birthing/health center), transport (to appropriate referral facility), and availability/quality of EmOC
(including adequate staff/supplies, appropriate clinical treatment, immediate attention), for rural and urban India. Shifts from home births assume a
70% shift to health centers/birthing centers and a 30% shift to EmOC; for routine births in EmOC, we assume 90% bEmOC and 10% cEmOC. Interventions Packaged as Integrated Services Increased family planning to reduce the unmet need (for spacing
and limiting births) by amounts ranging from 25% to 100%,
reduced maternal deaths by amounts ranging from 7.0% to 28.1%
in rural India and 5.8% to 23.5% in urban India (Table 4). In
rural India, eliminating the unmet need for family planning
decreased the TFR from 2.97 to 2.14, the proportion of deaths
that are pregnancy related from 16.4% to 12.3%, and the lifetime
risk of maternal death from 1 in 65 to 1 in 90. In rural India alone, Our results suggest that reaching the MDG 5 goal of a 75%
reduction in maternal mortality would require investments
targeting the intrapartum period, in addition to family planning
and safer abortion. Without these additional strategies, the model
predicts a ceiling on the level of maternal mortality reduction
achievable, ranging from 32% in urban India to 34% in rural
India. PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 5 Reducing Maternal Mortality in India Table 1. Selected model parameters: Incidence and mortality of pregnancy and delivery-related complications, and impact of
interventions. Parameter
Hemorrhage
Obstructed Labor
Hypertensive
Disorders
Sepsis
Unsafe Abortion
Incidence and mortality
Probability of event [15,29,31–44]
0.114
0.047
0.035
0.050
0.128a
Range
0.051–0.228
0.030–0.074
0.025–0.050
0.043–0.060
0.050–0.250
Probability of morbidity [28,30,45–48]b
0.008
0.022
0.001
0.400
0.120
Range
0.006–0.010
0.018–0.026
0.001–0.001
0.320–0.480
0.096–0.144
CFR [7,49–51]
0.010
0.007
0.017
0.013
0.003
Adjusted CFRc
0.023
0.019
0.021
0.028
0.009
Range
0.007–0.030
0.005–0.025
0.012–0.027
0.009–0.036
0.002–0.012
Attributable mortality [29]d
46.2% (9%–73%)
14.1% (3%–52%)
13.7% (0%–18%)
17.4% (0%–20%)
8.6% (0%–20%)
Model-projected attributable mortality
40.6%
16.8%
12.3%
20.4%
9.8%
Impact of interventions
Decreased incidence [34,37]e
50%,75%e
—
—
25%, 50%f
—
Rangef
25%–91%
—
25%–50%
0%–60%
0%–100%
Decreased CFR [7,17,50,52–64]g
75%
95%
59%
90%
98%a
Rangef
60%–90%
76%–100%
45%–95%
63%–93%
50%–100%
See [7,14,15,17,22,23,28–64,69–73]. aIncidence of elective abortion is 0.170, and 75% are assumed to be unsafe in the base case [15,29,39–44]. Case fatality rate (CFR) of safe abortion is 0.000006,
representing a 98% reduction in mortality [50,62–64]. For more details on abortion-related assumptions, see Text S1. Incidence of miscarriage (not shown) is 0.150
[69,70]. bSpecific examples of nonfatal complications include Sheehan’s Syndrome following maternal hemorrhage, fistula resulting from obstructed labor, neurological
sequelae from eclampsia, pelvic inflammatory disease (PID). Interventions Packaged as Integrated Services [29], based on the entire Asia region, as well as other
data to establish a range for sensitivity analysis [14,15]. Cause-specific proportions sum to 66%, reflecting approximately 33% indirect causes, although this varies from
15% to 35% in different studies. Estimates shown reflect adjustment of data from Khan et al. [29] such that a distribution is shown for the 66% of direct causes, to
compare to model output. Further, anemia was reported to be responsible for 15% of deaths and was assumed to exert mortality impact on direct causes through
severity of PPH, sepsis, and unsafe abortion. eIncidence of sepsis reduced by 50% with SBA and clean delivery in birthing center, bEmOC, and cEmOC; and reduced by 25% with SBA and clean delivery at home [37]. Incidence of maternal hemorrhage reduced by 50%–75% depending on expectant versus active management of labor; we assume for the status quo, all cEmOC
facilities provide active management, 50% of bEmOC facilities provide active management, and birthing centers/health centers provide expectant management only
[34]. Exploratory analyses that estimate the impact of community-based provision of oral misoprostol in birthing centers and at home assume a 25% to 50% reduction
in PPH [22,72]. For each baseline estimate, sensitivity analysis was conducted across a plausible range based on literature review; references are documented in the
Text S1. fFor each baseline estimate, sensitivity analysis was conducted across a plausible range based on literature review; references and assumptions are documented in the
Text S1. gEstimates shown represent average reduction in case fatality rate provided complications necessitating surgery (e.g., cesarean section), blood transfusion, intensive
hemodynamic support are treated in cEmOC. Obstructed labor is managed using assisted vaginal delivery with forceps or vacuum and, if necessary, cesarean section;
severe pre-eclampsia and eclampsia treated with intravenous hydralazine and magnesium sulfate, in addition to induction of labor or emergency cesarean section
when required; sepsis treated with ampicillin, gentamycin, and metronidazole or equivalent regimen followed by an 8-d course of intramuscular gentamycin and oral
metronidazole (see Text S1 for details) [7,73]. doi:10.1371/journal.pmed.1000264.t001 mortality ratio by 14% to 4%, and lifetime risk of maternal
mortality from one in 78 to one in 282. PLoS Medicine | www.plosmedicine.org Interventions Packaged as Integrated Services Not shown but included are the risk of infertility from PID (0.086) and the risk of severe anemia following
maternal hemorrhage (0.09) [23,45,46]. cCFRs were adjusted based on complication severity (e.g., life threatening complications requiring cEmOC) and underlying severity of anemia [71]. See Text S1. dEstimates for distribution of causes of maternal mortality for India are from India overall estimates from Khan et al. [29], based on the entire Asia region, as well as other
data to establish a range for sensitivity analysis [14,15]. Cause-specific proportions sum to 66%, reflecting approximately 33% indirect causes, although this varies from
15% to 35% in different studies. Estimates shown reflect adjustment of data from Khan et al. [29] such that a distribution is shown for the 66% of direct causes, to
compare to model output Further anemia was reported to be responsible for 15% of deaths and was assumed to exert mortality impact on direct causes through Table 1. Selected model parameters: Incidence and mortality of pregnancy and delivery-related complications, and impact of
interventions. Table 1. Selected model parameters: Incidence and mortality of pregnancy and delivery-related complications, and impact of
interventions. See [7,14,15,17,22,23,28 64,69 73]. aIncidence of elective abortion is 0.170, and 75% are assumed to be unsafe in the base case [15,29,39–44]. Case fatality rate (CFR) of safe abortion is 0.000006,
representing a 98% reduction in mortality [50,62–64]. For more details on abortion-related assumptions, see Text S1. Incidence of miscarriage (not shown) is 0.150 [69,70]. bSpecific examples of nonfatal complications include Sheehan’s Syndrome following maternal hemorrhage, fistula resulting from obstructed labor, neurological
sequelae from eclampsia, pelvic inflammatory disease (PID). Not shown but included are the risk of infertility from PID (0.086) and the risk of severe anemia following
maternal hemorrhage (0 09) [23 45 46] [
,
]
bSpecific examples of nonfatal complications include Sheehan’s Syndrome following maternal hemorrhage, fistula resulting from obstructed labor, neurological
sequelae from eclampsia, pelvic inflammatory disease (PID). Not shown but included are the risk of infertility from PID (0.086) and the risk of severe anemia following
maternal hemorrhage (0.09) [23,45,46]. cCFRs were adjusted based on complication severity (e.g., life threatening complications requiring cEmOC) and underlying severity of anemia [71]. See Text S1. dEstimates for distribution of causes of maternal mortality for India are from India overall estimates from Khan et al. April 2010 | Volume 7 | Issue 4 | e1000264 Interventions Packaged as Integrated Services Compared to the status
quo in urban India (Table 4, lower section), similar reductions in
maternal deaths were predicted; although the number of absolute
lives saved would be lower, the MMR, lifetime risk, and
proportionate mortality ratio would be expected to decline to
113, one in 553, and 2.3%, respectively, with the most intensive
strategy (upgrade 4). Table 5 shows the health and economic outcomes associated
with
interventions
packaged
as
integrated
services;
these
included phased approaches that scale up access to intrapartum
services over time in rural (upper section) and urban India (lower
section), coupled with incremental improvements in family
planning and safe abortion. The four stepwise ‘‘upgrades’’
incorporated improvements in available SBAs for home births,
recognition of referral need, transport, and availability/quality
of EmOC, as well as shifts from home- to facility-based
delivery. Because the stepwise improvements in each component of the
integrated package (intrapartum care, family planning, and safe
abortion) were assumed to occur in consecutive phases, the
incremental cost-effectiveness ratio for each ‘‘upgrade’’ strategy
was calculated as the difference in costs relative to the difference in
effects, compared with the preceding next best strategy. While the
initial strategy was cost saving in both urban and rural India, Compared to the status quo in rural India (upper section,
Table 5), our model predicted that integrated strategies coupling
family
planning
and
safe
abortion
with
four
consecutive
‘‘upgrades’’ would be expected to reduce maternal deaths by
17.3% to 77.1%, the MMR to less than 200, proportionate April 2010 | Volume 7 | Issue 4 | e1000264 6 Reducing Maternal Mortality in India Table 2. Selected model parameters and assumptions: Coverage of interventions and maternal health indicators by setting. Table 2. Selected model parameters and assumptions: Coverage of interventions and maternal health indicators by setting. Interventions Packaged as Integrated Services p
p
g
y
g
Parameter
India
India, Urban
India, Rural
Rajasthan
Uttar Pradesh, Rural
Coverage of contraception (%) [28]
Family planning (any method)
56.3
64.0
53.0
47.2
39.7
Modern methods
48.5
55.8
45.3
44.4
25.2
Pill
6.4
7.0
6.2
4.5
5.2
IUD
3.7
6.1
2.4
3.6
3.2
TOL
76.9
67.7
81.9
77.0
66.7
Condom
10.9
17.9
7.3
12.8
24.6
Unmet need
13.2
10.0
14.6
14.6
23.8
Coverage of prenatal care (%) [28]
Prenatal care
50.7
73.8
42.8
41.2
22.6
Treatment for anemiaa
22.3
34.5
18.1
13.1
6.7
Delivery location (%) [28]
Total skilled delivery
48.3
75.2
39.1
41.0
23.8
Facility deliveryb
40.7
69.4
31.1
29.6
17.5
Home delivery with SBAb
12.8
19.0
11.6
16.2
7.6
Assumptions for available transport/interim
care to appropriate facility (%) [65,66]
From home to EmOC
30.4
44.4
24.4
24.4
18.1
Range
20–40
35–55
15–35
20–40
15–30
From HC or BC to EmOC
54.8
68.8
48.8
48.8
36.1
Range
40–65
60–80
40–60
30–55
25–45
From bEmOC to cEmOC
67.0
81.0
61.0
61.0
45.1
Range
55–80
70–90
50–70
50–70
35–55
Assumptions for available facility,
staff/supplies, quality of care (%) [67,68]
EmOC
50.0
67.5
42.5
42.5
31.5
Range
40–60
55–80
30–55
30–55
20–40
Maternal health indicators
TFR [28]
2.68
2.07
2.98
3.21
4.13
Model-projected TFRc
2.70
2.07
2.97
3.24
4.13
Model-projected MMRc
440
407
520
524
633
See [14,15,28,65–68,71]. aCase fatality rates (CFRs) were adjusted based on complication severity (e.g., life threatening complications requiring cEmOC) and underlying severity of anemia [71]. See Text S1. bRoutine deliveries in EmOC facilities assume that 90% would be in bEmOC and 10% in cEmOC. Alternative assumptions explored in sensitivity analysis (Text S1). We
calculated the percentage of births with skilled attendance at home by subtracting the percentage delivered in facilities (which we assume are with skilled attendance)
from the total of births with skilled attendance: (total skilled delivery2facility based births)/home births; for rural India: (0.39120.311)/(120.311) = 0.116 or 11.6%. cUsing the empirically calibrated India model, we parameterized the state-level models for Rajasthan and Uttar Pradesh and adjusted for the TFR as reported in NFHS 3
[28]. To provide comparison, reported MMRs for Uttar Pradesh include an estimate of 707 from Mills [14], and prior data from SRS including the SRS 2001–2003 estimate
of 517 (confidence interval [CI] 461–573), SRS 1999–2001 estimate 539 (481–596), and the SRS 1997–98 (606, CI 544–668) [15]. Interventions Packaged as Integrated Services For Rajasthan, we used the 2001–2003,
special survey of deaths using RHIME, which reported 445 (371–519), and SRS 1999–2001, which reported 501 (423–580) [15]. BC, birthing center; HC, health center; IUD, intrauterine device; TOL, female sterilization. doi:10.1371/journal.pmed.1000264.t002 PLoS Medicine | www.plosmedicine.org Integrated Safe Motherhood Interventions in Rural Uttar
Pradesh Costing details and methods for converting costs to 2006 US$ are
id d i
h
T
S1 provided in the Text S1. aRanges for sensitivity analyses established on the basis of assumptions and other published literature documented in the Text S1. b p
aRanges for sensitivity analyses established on the basis of assumptions and other published literature documented in the Text S1. bAntenatal care includes tetanus vaccination syphilis gonorrhea chlamydia screening (and treatment) urinalysis blood tests treatment for anemia counseling (e g p
aRanges for sensitivity analyses established on the basis of assumptions and other published literature documented in the Text S1. bAntenatal care includes tetanus vaccination, syphilis, gonorrhea, chlamydia screening (and treatment), urinalysis, blood tests, treatm
family planning, spacing, intrapartum care). y p
g, p
g,
p
)
cPostabortion complications assumed to require manual vacuum aspiration, treatment of sepsis in 25%, surgical repair in 25% [92]. d p
q
p
,
p
,
g
p
[
]
dTotal costs reflect skill level of attendant, level of facility, and drugs and supplies. For example, delivery at birthing center (US$14.4
facility (US$4.52), and drugs and supplies (US$3.50). Other assumptions documented in the Text S1. f attendant, level of facility, and drugs and supplies. For example, delivery at birthing center (US$14.46) includes personnel (US$6.44)
nd supplies (US$3.50). Other assumptions documented in the Text S1. eCommunity-based interventions evaluated in sensitivity analysis included SBA-administered misoprostol to reduce incidence of PPH in deliveries at home and in
birthing centers. Costs for misoprostol (US$0.99) and training (upper bound, US$3.40) based on assumptions presented in Sutherland and Bishai [18]; these costs
represent the incremental costs above routine SBA delivery. y
fTransport costs include those incurred from home to a referral facility (bEmoc or cEmOC), and those incurred between facilities when necessary (e.g., bEmOC to
cEmOC). Assumptions based on literature [80,93,94] and public access data described in the Text S1. gEstimates shown represent average total costs using case-specific unit costs weighted by severity. Complications requiring surgery (e.g., cesarean section), blood
transfusion, intensive hemodynamic support assumed to require cEmOC. Details of unit cost assumptions for facility-specific treatment documented in Text S1. hPostpartum care includes examination, iron/folate supplementation, and counseling. doi:10.1371/journal.pmed.1000264.t003 In Figure 4 (upper panel) each cell is also color-coded to
reflect the cost-effectiveness profile associated with the partic-
ular strategy. PLoS Medicine | www.plosmedicine.org Integrated Safe Motherhood Interventions in Rural Uttar
Pradesh incremental cost-effectiveness ratios ranged from US$150 to
US$300 per YLS in rural India and from US$150 to US$350
per YLS in urban India. Cost-effectiveness ratios are also
expressed as percent of the per capita GDP (US$1,068). Even
the most intensive and effective strategic package was well below
50% of the per capita GDP. Figure 4 (upper panel) displays the health outcomes associated
with stepwise approaches to improve maternal health in rural Uttar
Pradesh. The vertical axis (from bottom to top) shows outcomes
associated with increased access to family planning and safe
abortion; the horizontal axis (from left to right) displays outcomes
associated with investments in high-quality health-center–based
intrapartum care (e.g., facility, attendance, referral, transport,
EmOC). In Figure 4 (upper panel) the cell located in the far left
lower corner represents current conditions in rural Uttar Pradesh. In contrast to these integrated strategies, implementing only the
stepwise intrapartum care upgrades—without family planning and
safe abortion—was less effective and less cost-effective. The
incremental
cost-effectiveness
ratios
ranged
from
US$490–
US$1,060 in rural India and US$200–US$990 per YLS in urban
India (Text S1). April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 7 Reducing Maternal Mortality in India Table 3. Selected model input costs. Cost Components
Base Case
Rangea
Family planning [25,88]
Oral contraceptives
10.64
6.03–15.96
Injectable contraceptives
10.20
4.92–15.30
Condoms
8.40
3.79–12.60
Intrauterine device
9.17
2.58–13.76
Female sterilization
18.98
9.49–28.47
Male sterilization
12.67
6.34–19.01
Antenatal careb [25,88]
Four visits
17.82
8.54–25.61
Abortionc [25,88,94]
Incomplete abortion
8.90
4.45–17.80
Elective abortion
21.87
10.94–43.74
Postabortion complications
43.40
21.70–86.80
Deliveryd [25,88]
Home (TBA, SBA)
4.52, 6.44
0–9.66
Facility (birthing center, bEmOC, cEmOC)
14.46, 24.58, 32.54
7.23–48.81
Community-based interventionse
Misoprostol (home, birthing center)
0.99
0.75–2.00
SBA training
3.40
0.62–5.00
Transportation costsf
Home to facility
3.62–8.13
1.81–12.20
Birthing/health center/bEmOC to referral facility
4.88–7.14
2.44–10.71
Management of complicationsg [25,88]
Obstructed labor
70.16
12.76–139.38
Maternal hemorrhage
67.99
18.40–212.51
Puerperal sepsis
47.92
23.15–111.02
Severe pre-eclampsia/eclampsia
65.85
33.50–153.62
Postpartum careh [25,88]
One visit
4.99
1.04–7.49
See [18 25 80 88–94] Table 3. Selected model input costs. See [18,25,80,88 94]. Estimates of costs under current standard of care (2006 US$). Estimates for the base case were country-specific and from UNFPA’s Reproductive Health Costing Tools
Model (RHCTM) [88] and WHO CHOICE/public databases [25,89–91], unless otherwise specified. April 2010 | Volume 7 | Issue 4 | e1000264 Integrated Safe Motherhood Interventions in Rural Uttar
Pradesh Strategies that only employed family planning and
safe abortion (vertical axis, from bottom to top) were generally
cost saving, but reduced mortality by a maximum of 40%. Strategies that only invested in intrapartum care improvements Each of the other cells represents a unique strategy; the reduction in
maternal deaths expected with each strategy, relative to current
conditions, is shown. For example, a strategy that reduced the
unmet need by 75%, increased safe abortion to 60%, and
implemented improvements to intrapartum care consistent with
upgrade 3, reduced maternal deaths by 57%. April 2010 | Volume 7 | Issue 4 | e1000264 8 Reducing Maternal Mortality in India Table 4. Health and economic outcomes of family planning to reduce the unmet need for limiting and spacing births, and safe
abortion, in rural and urban India. Table 4. Health and economic outcomes of family planning to reduce the unmet need for limiting and spacing births, and safe
abortion, in rural and urban India. Table 4. Health and economic outcomes of family planning to reduce the unmet need for limiting and spacing births, and safe
abortion, in rural and urban India. Integrated Safe Motherhood Interventions in Rural Uttar
Pradesh Strategy
Lifetime
Deaths per
100,000
Women
Reduction
in Maternal
Deaths
Proportionate
Mortality Ratio
Lifetime Risk
of Death Due
to Maternal
Complications
Model-projected
Savings for
a Single Birth
Cohort of 15 y
olds (US$)a
Cost Savings
for a Single
Year (Current
Distribution of
15–45 y Olds in
India) (US$)b
Rural India, current conditions (TFR, 2.97)c
1,543
—
16.4%
1 in 65
NA
—
Family planning
Reduce unmet need 25% (56.7%)
1,435
7.0%
15.4%
1 in 70
111,357,615
60,200,655
Reduce unmet need 50% (60.3%)
1,327
14.0%
14.4%
1 in 75
223,221,615
120,611,563
Reduce unmet need 75% (64.0%)
1,218
21.1%
13.4%
1 in 82
335,439,615
181,233,496
Reduce unmet need 100% (67.6%)
1,109
28.1%
12.3%
1 in 90
448,188,615
242,067,230
Safe abortion
Increase safe abortion 50%
1,517
1.7%
16.2%
1 in 66
48,080,115
42,078,125
Increase safe abortion 75%
1,473
4.5%
15.8%
1 in 68
130,739,115
114,289,234
Increase safe abortion 95%
1,433
7.1%
15.4%
1 in 70
214,460,115
167,790,590
Family planning and safe abortion
Reduce unmet need (56.7%), safe abortion 75%
1,369
11.3%
14.8%
1 in 73
233,930,115
166,870,014
Reduce unmet need (60.3%), safe abortion 75%
1,265
18.0%
13.8%
1 in 79
337,386,615
219,568,526
Reduce unmet need (64.0%), safe abortion 75%
1,160
24.8%
12.8%
1 in 86
441,108,615
272,385,038
Reduce unmet need (67.6%), safe abortion 95%
1,026
33.5%
11.5%
1 in 98
580,230,615
362,579,472
Urban India, current conditions (TFR, 2.07)d
842
—
9.6%
1 in 119
NA
—
Family planning
Reduce unmet need 25% (66.5%)
793
5.8%
9.1%
1 in 126
22,089,305
12,838,532
Reduce unmet need 50% (69.0%)
743
11.7%
8.6%
1 in 135
44,214,305
25,696,437
Reduce unmet need 75% (71.5%)
694
17.6%
8.1%
1 in 144
66,398,305
38,578,054
Reduce unmet need 100% (74.0%)
644
23.5%
7.5%
1 in 155
88,611,805
51,483,279
Safe abortion
Increase safe abortion 50%
822
2.4%
9.4%
1 in 122
11,351,305
9,742,159
Increase safe abortion 75%
788
6.4%
9.1%
1 in 127
30,821,305
26,413,778
Increase safe abortion 95%
758
9.9%
8.7%
1 in 133
50,438,805
36,823,443
Family planning and safe abortion
Reduce unmet need (66.5%), safe abortion 75%
741
11.9%
8.6%
1 in 135
51,235,305
37,766,193
Reduce unmet need (69%), safe abortion 75%
695
17.5%
8.1%
1 in 144
71,649,305
49,126,308
Reduce unmet need (71.5%), safe abortion 75%
648
23.0%
7.6%
1 in 154
92,122,305
60,499,204
Reduce unmet need (74%), safe abortion 95%
580
31.2%
6.8%
1 in 173
119,557,305
79,270,520
See [87]. Integrated Safe Motherhood Interventions in Rural Uttar
Pradesh Reduction in direct causes of maternal mortality, including abortion-related complications, postpartum hemorrhage, hypertensive disorders, sepsis, and
obstructed labor. aModel-projected cost savings reflect net costs averted over a woman’s reproductive lifespan (ages 15–45 y) applied to the current population of 15 y olds in India
stratified by rural (75%) and urban (25%) settings [87]. Future costs discounted 3% annually. bCost savings for a single representative year of a successfully implemented strategy were calculated using population-level data from India [87] stratified by rural (75%)
and urban (25%) settings, for the current distribution of reproductive age women (ages 15–45 y). cIn rural India model-projected TFR is 2 76 2 56 2 36 2 14 with reductions in unmet need of 25% 50% 75% 100% respectively (horizontal axis, left to right) were generally associated with the
highest cost-effectiveness ratios (i.e., least attractive), reflecting
the higher costs required for infrastructure improvements. An
overarching strategic approach that moves along the diagonal,
from the lower left corner to the upper right corner, was most
effective and cost-effective; the cost savings from enhanced
family planning and safe abortion offset the resources required
to improve intrapartum care. PLoS Medicine | www.plosmedicine.org Sensitivity Analyses For deliveries at home and in birthing centers in rural Uttar
Pradesh, removing only one ‘‘delay’’ in accessing EmOC had
minimal impact (,5%) on lowering maternal mortality and was
not cost-effective (e.g., US$700–US$4,900 per YLS) (Figure 4,
lower panel). In contrast, an integrated strategy that made modest
improvements in all components (e.g., SBA, referral, transport,
and quality) reduced mortality by 22%. Cost-effectiveness of an April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 9 Reducing Maternal Mortality in India Figure 3. Averted deaths with family planning and safe abortion. Averted deaths attributable to unsafe abortion in rural India by addressing
need for family planning (green shading) and providing 75% safe abortion (blue shading). Magnitude of additional averted abortion-related deaths
with improved access to safe abortion depends on the amount of unmet need for contraception. doi:10.1371/journal.pmed.1000264.g003 Figure 3. Averted deaths with family planning and safe abortion. Averted deaths attributable to unsafe abortion in rural India by addressing
need for family planning (green shading) and providing 75% safe abortion (blue shading). Magnitude of additional averted abortion-related deaths
with improved access to safe abortion depends on the amount of unmet need for contraception. doi:10.1371/journal.pmed.1000264.g003 integrated strategy ranged from cost saving to US$170 per YLS
(Text S1). uncertainty in data and assumptions used in the analysis, four
critical themes emerge as robust. Universal antenatal care by itself averted fewer than 2% of
maternal deaths; however, if enhanced antenatal care increased
the probability of either facility-based delivery or SBA-attended
birth (linked with accurate referral and transport) from 31% to
60%, health benefits increased 5-fold (Text S1). First, increasing effective family planning is the most effective
individual intervention to reduce pregnancy-related mortality. If
the unmet need was met in rural and urban India by 2012, our
results imply that the lives of 168,000 women would be saved by
the end of 2015. The cost savings over that time period would
exceed US$1 billion. Because strategies to increase contraceptive
options for limiting and spacing do not require the same level of
infrastructure as improving intrapartum care, targeting these
strategies toward rural areas with high TFRs is a promising way to
initiate equitable improvements in maternal health. Sensitivity Analyses As a greater proportion of routine deliveries shifted from home
to facilities, we assumed 70% would shift to birthing centers or
health centers staffed by SBA and 30% to facilities with full
EmOC capacity. Although the differential benefits of routine
delivery in birthing/health centers versus bEmOC was dependent
on expedient transfer from a center to referral EmOC if needed,
provided this was assured, both approaches were cost-effective. In
contrast, when we varied assumptions about the proportion of
routine deliveries in cEmOC versus bEmOC, cost-effectiveness
results changed drastically; as routine deliveries shifted to cEmOC,
the
incremental
cost-effectiveness
ratios
became
much
less
attractive, ranging from US$8,300 to US$27,000 per YLS. Second,
two
distinct—yet
synergistic—approaches,
family
planning and safe abortion, can reduce deaths from unsafe
abortion. Enhanced access to family planning by itself reduces
demand for elective abortion and consequently reduces deaths
attributable to unsafe abortion. In fact, reducing the unmet need
for contraception can prevent one of every two abortion-related
deaths. Furthermore, just a fraction of the cost savings from family
planning would fully fund an intervention to provide safe abortion
and postabortion care. Table 6 shows the potential incremental benefits and cost-
effectiveness of adjunctive community-based SBA-administered
misoprostol for births at home and birthing centers/health centers
in rural India. For all four ‘‘upgrade’’ strategies, additional lives
could be saved; depending on the phase of improvements in
intrapartum care, an additional 7%–13% of maternal deaths were
prevented. Cost-savings for a single birth cohort of 15-y-old girls
(2010) expected to accrue over their reproductive lifespan (age 15–
45 y) ranged from US$128 million to US$190 million. Third, despite the substantial health and economic benefits
associated with family planning and safe abortion, there is a
threshold above which further reductions in mortality are
impossible. MDG 5 will therefore not be achievable without
involving integrated interventions that ensure reliable access to
high-quality intrapartum and EmOC. These interventions could
be implemented, however, in a staged, scale-up fashion. While formidable effort and financial investment would be
required to scale up maternal health services over time, we
identified a number of phased approaches that would ultimately
prevent four out of five maternal deaths. Coupled with stepwise
improvements in family planning and safe abortion, these
approaches incrementally shifted home births to birthing centers
or facilities with EmOC, and improved both access to SBAs as well
as accurate recognition of referral need, transport, and availabil-
ity/quality
of
EmOC. Sensitivity Analyses Successful
implementation
of
these PLoS Medicine | www.plosmedicine.org Discussion Cost-effectiveness ratios are also expressed as percent of the per capita GDP (US$1,068), shown in the farthest right column, as
interventions with cost-effectiveness ratios of less than the per capita GDP are considered very cost-effective according to criteria proposed by the Commission on
Macroeconomics and Health [98]. p
g
gy
p
p
p
p
g
interventions with cost-effectiveness ratios of less than the per capita GDP are considered very cost-effective according to criteria proposed by the Commission on
Macroeconomics and Health [98]. facility births; 11.6% SBA (home births); transport from home (24.4%), primary-level health center (48.8%), bEmOC (61%); recognition o
primary-level health center (40%); availability and quality of EmOC (42.5%); 53% family planning. eStatus quo (rural India): 31.1% facility births; 11.6% SBA (home births); transport from home (24.4%), primary-level health center (48.8%), bEmOC (61%); recognition of
referral need at home (20%), primary-level health center (40%); availability and quality of EmOC (42.5%); 53% family planning. fStatus quo (urban India): 69.4% facility births; 19% SBA (home births); transport from home (44%), primary-level health center (69%), bEmOC (81%); recognition of
referral need at home (20%), primary-level health center (40%); availability and quality of EmOC (67.5%); 64% family planning. CS, cost saving; ICER, incremental cost-effectiveness ratio. eStatus quo (rural India): 31.1% facility births; 11.6% SBA (home births); transport from home (24.4%), primary-level health center (48.8%), bEmOC (61%); recognition of
referral need at home (20%), primary-level health center (40%); availability and quality of EmOC (42.5%); 53% family planning. fSt t
(
b
I di ) 69 4% f
ilit
bi th
19% SBA (h
bi th ) t
t f
h
(44%)
i
l
l h
lth
t
(69%) bE
OC (81%)
iti
f % facility births; 19% SBA (home births); transport from home (44%), primary-level health center (69%), bEmOC (81%); recognition of
primary-level health center (40%); availability and quality of EmOC (67.5%); 64% family planning. ntal cost-effectiveness ratio. doi:10.1371/journal.pmed.1000264.t005 includes SBA-administered oral misoprostol in homes and birthing
centers is likely to be cost-effective. PLoS Medicine | www.plosmedicine.org Discussion Accuracy of referral need recognition at home and in health center with SBA increase, on average, to 60%, 75%, 90%, and 95% (not shown) with upgrade 1, 2, 3, and 4
in both rural and urban India aTransport encompasses the expedient availability of means of transport (e.g., vehicle, cart), fuel (if needed), driver, and interim attendant care. Facility transport
represents a weighted average of transport availability from a health center or birthing center to an EmOC facility and from a bEmOC facility to a cEmOC if indicated. Accuracy of referral need recognition at home and in health center with SBA increase, on average, to 60%, 75%, 90%, and 95% (not shown) with upgrade 1, 2, 3, and 4
in both rural and urban India. in both rural and urban India. bQuality refers to the availability and quality of services at EmOC facilities, including adequate staffing and supplies, expedient attention (e.g., without delay to collect
fees or requirement for family to bring supplies), and evidence-based clinical practices. in both rural and urban India. bQuality refers to the availability and quality of services at EmOC facilities, including adequate staffing and supplies, expedient attention (e.g., without delay to collect
fees or requirement for family to bring supplies), and evidence-based clinical practices. y
g
pp
),
p
traceptive use for limiting and spacing; shown are values representing the reduction in unmet need by 25%, 50%, 75%, and 100% wit
ctively, for both rural and urban India. dStepwise improvements in maternal health services are assumed to occur in consecutive phases (e.g., first upgrade 1, then upgrade 2, etc.). Therefore, the incremental
cost-effectiveness ratio (US$ per YLS) for each upgrade is calculated as the difference in lifetime costs relative to the difference in lifetime effects, compared with the
preceding next best strategy. Cost-effectiveness ratios are also expressed as percent of the per capita GDP (US$1,068), shown in the farthest right column, as dStepwise improvements in maternal health services are assumed to occur in consecutive phases (e.g., first upgrade 1, then upgrade 2, etc.). Therefore, the incremental
cost-effectiveness ratio (US$ per YLS) for each upgrade is calculated as the difference in lifetime costs relative to the difference in lifetime effects, compared with the
preceding next best strategy. April 2010 | Volume 7 | Issue 4 | e1000264 Discussion We have identified several strategic options that would cost-
effectively reduce maternal mortality in both rural and urban
India. Our principal findings are that early intensive efforts to
improve family planning and provide safe abortion, accompanied
by a systematic stepwise effort to scale up intrapartum and EmOC,
could reduce maternal mortality by 75%. Despite the inherent April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 10 Reducing Maternal Mortality in India Table 5. Health and economic outcomes of integrated packages of services: intrapartum care, family pla Table 5. Health and economic outcomes of integrated packages of services: intrapartum care, family planning, and safe abortion. Region
Facility
Birth
(%)
Transport-
Homea
(%)
Transport-
Facilitya
(%)
Quality
of Careb
(%)
Family
Planningc
(%)
Safe
Abortion
(%)
Decrease
in
Maternal
Deaths
(%)
MMR
(Deaths
per
100,000
Live
Births)
Maternal
Deaths as
Percent
of Deaths
Ages
15–45 y
Lifetime
Risk of
Maternal
Death
Lifetime
Costsd
(US$)
Cost-
Effectivenessd
ICER (US$/
YLS)
ICER
(% per
capita
GDP)
Rural Indiae
—
—
—
—
—
—
—
520
16.4
1 in 65
218.38
—
—
Upgrade 1
45
50
65
70
56.7
50
17.3
460
14.0
1 in 78
212.41
CS
CS
Upgrade 2
60
60
75
80
60.3
60
33.7
397
11.5
1 in 98
218.51
150
14
Upgrade 3
75
70
85
90
64.0
75
53.4
302
8.3
1 in 139
226.18
160
15
Upgrade 4
80
75
95
95
67.6
95
77.1
162
4.3
1 in 282
243.46
300
28
Urban Indiaf
—
—
—
—
—
—
—
407
9.6
1 in 119
184.00
—
—
Upgrade 1
75
60
85
70
66.5
50
15.8
363
8.2
1 in 141
174.91
CS
CS
Upgrade 2
80
75
90
80
69.0
60
33.3
305
6.6
1 in 178
178.30
150
14
Upgrade 3
90
80
92.5
90
71.5
75
54.1
225
4.7
1 in 259
183.44
220
21
Upgrade 4
95
85
95
95
74.0
95
78.5
113
2.3
1 in 553
194.97
350
33 labor. aTransport encompasses the expedient availability of means of transport (e.g., vehicle, cart), fuel (if needed), driver, and interim attendant care. Facility transport
represents a weighted average of transport availability from a health center or birthing center to an EmOC facility and from a bEmOC facility to a cEmOC if indicated. Discussion With the
exception of anemia, we focused on direct causes of maternal
mortality; a priority for future analyses is to include interventions
to reduce the indirect causes of pregnancy-related mortality. The
analysis would be strengthened by availability of indicators that
reflect safe motherhood externalities including measures of
enhanced household well-being, increased school attendance,
decreased numbers of orphans, and reduced impoverishment
resulting from catastrophic expenses [94,99]. Other limitations in our analysis stem from its inherent reliance
on high-quality data about maternal mortality specifically. While
our calibration of setting-specific models allows us to better
represent within-country differences in baseline risk, coverage, and
capacity than previous studies, high-quality empiric evidence for Shiffman and Smith [101] have described the importance of
framing priority public health issues in a manner that resonates
with both the ‘‘internal’’ community and other ‘‘external’’
decision makers. With regard to the internal community, the Table 6. Incremental benefits of community-based misoprostol in rural India. Rural India
Family
Planninga
(%)
Safe
Abortion
(%)
Facility
Birth (%)
Transport-
Homeb (%)
Transport-
Facilityb (%)
Quality of
Carec (%)
Incremental Benefits and Cost-Effectiveness
of Community-Based Misoprostold
Decrease in
Maternal
Deaths
Lives Saved with
Addition of
Community-
based
Misoprostole
Cost-
Effectiveness
Upgrade 1
56.7
50
45
50
65
70
—
—
—
Plus community
interventiond
—
—
—
—
—
—
12.3%
16,992
Cost savinge
Upgrade 2
60.3
60
60
60
75
80
—
—
—
Plus community
interventiond
—
—
—
—
—
—
13.0%
17,612
Cost savinge
Upgrade 3
64.0
75
75
70
85
90
—
—
—
Plus community
interventiond
—
—
—
—
—
—
10.2%
13,983
Cost savinge
Upgrade 4
67.6
95
80
75
95
95
—
—
—
Plus community
interventiond
—
—
—
—
—
—
6.9%
9,470
Cost savinge
aFamily planning refers to contraceptive use for limiting and spacing; shown are values representing the reduction in unmet need by 25%, 50%, 75%, and 100% with
upgrades 1, 2, 3, and 4, respectively. bTransport encompasses the expedient availability of means of transport (e.g., vehicle, cart), fuel (if needed), driver, and interim attendant care. cQuality refers to the availability and quality of services at EmOC facilities, including adequate staffing and supplies, and evidence-based clinical practices. dCommunity-based interventions assume SBA-administered misoprostol for births at home and birthing centers/health centers with a 50% (25%–60%) reduction in PPH
[72]. Discussion Although alone it cannot
substitute for reliable intrapartum care and EmOC, if added to a
long-term plan that increases facility-based intrapartum care, it
will save additional lives and will reduce costs; (4) Because settings
with the highest TFRs and worst maternal health indicators also
tend to be those with the greatest need for enhanced health
delivery infrastructure, early consistent commitments to provide
family planning and safe abortion reduce the total resources
required. To place the synergistic benefits of enhanced family
planning and safe abortion in context, the magnitude of cost
savings from eliminating unmet need and ensuring access to safe
abortion is approximately 25% of the required 10-y investment
estimated by Johns et al. for scaling up maternal services in India
[30]. strategies would be expected to dramatically reduce the MMR,
proportionate mortality ratio, and lifetime risk of maternal death. Fourth, despite the possible variation in pace associated with
scaling up maternal health services in India, systematic and
consecutive phases will be cost-effective. Our results showed
that—when coupled with family planning and safe abortion—both
early initial strategies and late intensive strategies resulted in cost-
effectiveness ratios that were just a fraction of India’s per capita
GDP; these would unarguably be considered very cost-effective
[95–98]. Although our general findings are consistent with earlier
suggestions that interventions to reduce maternal mortality are
good public health investments [7,17], our analytic approach also
allowed us to identify more and less efficient ways to achieve this. In this regard, we highlight four robust insights: (1) In settings with
limited infrastructure, investing in ‘‘intermediate’’ facilities (e.g.,
birthing centers) is very cost-effective, provided there is reliable
referral capacity and transport to an appropriate EmOC facility if
necessary; (2) A strategy of routine hospital-based delivery (i.e.,
cEmOC) is not cost-effective; (3) A community-based strategy that We may have underestimated both effectiveness and cost-
effectiveness by excluding effects of certain indirect indicators and
interventions. Although
we purposely
focused on
maternal
mortality, if we included neonatal health and survival, for
example, most strategies would be even more cost-effective due April 2010 | Volume 7 | Issue 4 | e1000264 11 Reducing Maternal Mortality in India PLoS Medicine | www.plosmedicine.org PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 PLoS Medicine | www.plosmedicine.org 12 Reducing Maternal Mortality in India Figure 4. Health and economic outcomes in rural Uttar Pradesh. Upper panel. Discussion Reduction in maternal deaths and cost-effectiveness with
stepwise approaches to improve maternal health in rural Uttar Pradesh. The vertical axis (from bottom to top) shows outcomes associated with
increased access to family planning and safe abortion. The horizontal axis (from left to right) displays outcomes associated with investments in high-
quality health-center–based intrapartum care, which involved stepwise improvements in SBAs, recognition of referral need, and antenatal/
postpartum care, incrementally shifted births away from home, and improved transport, availability, and quality of EmOC. Each cell represents a
unique strategy; the reduction in maternal deaths shown is relative to current conditions (far lower left corner). Shading reflects cost-effectiveness
ratios, compared to status quo (pink, cost saving; blue, ,US$250/YLS; green, ,US$500/YLS; purple, ,US$750/YLS; orange, US$890/YLS). See text for
details. Lower panel. Sensitivity analysis depicting the impact of improving only one component of intrapartum care services in rural Uttar Pradesh. Base case and improvements for each component are shown in numbers below the component name (e.g., quality of EmOC is 31.5% in the base
case, improvements of 50%–100% were assessed). In settings where most deliveries occur at home, linkage of services in multiple domains is a critical
determinant of reduction in maternal deaths. Even large improvements in skilled birth attendants for nonfacility births (yellow bar), recognition of
referral need (pink bar), transport (green bar), or quality of EmOC (purple bar) fail to reduce mortality more than 2%–5% if other interdependent
components are not improved as well. In contrast, even modest improvements in all four components, as shown in upgrade 1 (blue bar), reduce
mortality by 22%. Cost-effectiveness of the integrated strategy ranged from cost saving to US$170 per YLS compared with US$700 to US$4,900 per
YLS for single unlinked improvements. doi:10.1371/journal.pmed.1000264.g004 the effectiveness of comprehensive strategies to reduce maternal
mortality and morbidity is often either lacking or inconsistent. More studies quantifying the benefits of community-level inter-
ventions on preventing maternal morbidity remain a priority
[100]. The additional costs that we assumed would be required to
scale up interventions and build infrastructure are, at best, gross
estimates. That being said, our assumption of 2- to 3-fold increases
in the per woman costs to reflect the additional resources required
to improve capacity is consistent with those implied by recent
analyses assessing global resource needs for maternal health
[30,93]. to associations between place of birth and presence of a skilled
attendant, with neonatal and maternal deaths [99,100]. Supporting Information Text S1
Supplemental material accompanying the article. Part I,
overview of model; part II, overview of model parameterization,
calibration, performance; part III, overview of costs and estimates;
part IV, supplemental results; part V, references. Finally, by placing and prioritizing safe motherhood in the
context of other global health priorities [101,102], our results can
also be effectively framed for policymakers who must allocate
limited resources, by providing comparative and contextual
information about the relative benefits and cost-effectiveness of
investments in maternal health measured against other public
health priorities. One of the robust findings of our analysis, for
example, is that there are integrated strategies that involve
improvements in family planning, safe abortion, and intrapartum
care that are equally or more cost-effective or attractive than
childhood immunization or treatment of malaria, tuberculosis, or
HIV [95]. Found at: doi:10.1371/journal.pmed.1000264.s001 (0.53 MB
PDF) Author Contributions ICMJE criteria for authorship read and met: SG SS NC UCMN DH. Agree with the manuscript’s results and conclusions: SG SS NC UCMN
DH. Designed the experiments/the study: SG SS NC DH. Analyzed the
data: SG SS NC UCMN DH. Collected data/did experiments for the
study: SG DH. Wrote the first draft of the paper: SG. Contributed to the
writing of the paper: SG SS NC UCMN DH. Wrote first draft of Text S1:
DH. Acknowledgments We are appreciative of the administrative assistance from Meredith
O’Shea, and constructive feedback on the model assumptions (Dan
Grossman, Kelly Blanchard, Carol Levin). The Indian government has initiated several policies to improve
maternal health [5], particularly in rural areas [27,103], and
efforts to both implement and evaluate new strategies are ongoing
[83,94,104–106]. Although our analysis is intended to catalyze
actionable steps, we recognize that decisions in India about the
choice of strategies and rate of stepwise investments to reduce
maternal mortality will be a function not only of cost-effectiveness
and affordability, but also of political will and local circumstances. Identifying approaches that can be tailored to local situations, but Reducing Maternal Mortality in India that rely on firm core principles and are cost-effective, holds
considerable promise as a way to mobilize further political support
and convince stakeholders that MDG 5 is within reach. strategies we identified as most effective support three crucial
elements already recognized as essential to achieve MDG 5:
family planning and control of fertility choices, provision of safe
abortion,
and
assurance
that
all
women
have
access
to
intrapartum care and EmOC. Our results reinforce this message,
and extend it by quantifying the cost savings of family planning
and safe abortion, and identifying efficient and cost-effective
approaches to scaling-up capacity for integrated maternal health
services. In particular, it is clear from our analysis that an initial focus on
family planning, especially in rural poor areas, will significantly
prevent pregnancy-related deaths, reduce deaths from unsafe
abortion, and save resources. Providing universal access to safe
abortion will further augment these benefits. The cost savings from
these two strategies will partially offset the resources required to
invest in the necessary infrastructure that would assure every
woman access to high-quality intrapartum care and EmOC. While
MDG 5 is unlikely to be met without assuring access to health-
center–based intrapartum care, implementation of a phased
stepwise approach, designed to reach this goal while reflecting
the current realities and most feasible initial approaches in
different settings, is absolutely within reach. With regard to the external community, we have tried to
provide a range of outcomes that can be used to create effective
‘‘take home’’ messages for different target audiences. For example,
in only 5 y, more than 150,000 lives could be saved just from
increasing contraception rates by a few percentage points; nearly
US$1.5 billion could be saved by adding safe abortion to family
planning efforts; and finally, with stepwise investments to provide
facility-based intrapartum care, the majority of maternal deaths
could be prevented. In the next decade, this accomplishment
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community-based misoprostol intervention). These were calculated by applying model-projected outcomes to population-level data from rural India [87]. Cost-savings
for a single birth cohort of 15-y-old girls (2010) expected to accrue over their reproductive lifespan (age 15–45) ranged from US$128 million to US$190 million. doi:10.1371/journal.pmed.1000264.t006 Table 6. Incremental benefits of community-based misoprostol in rural India. aFamily planning refers to contraceptive use for limiting and spacing; shown are values representing the reduction in unmet need by 25%, 50%, 75%, and 100% with
upgrades 1, 2, 3, and 4, respectively. bTransport encompasses the expedient availability of means of transport (e.g., vehicle, cart), fuel (if needed), driver, and interim attendant care. cQuality refers to the availability and quality of services at EmOC facilities, including adequate staffing and supplies, and evidence-based clinical practices. dCommunity-based interventions assume SBA-administered misoprostol for births at home and birthing centers/health centers with a 50% (25%–60%) reduction in PPH
[72]. ePopulation-level incremental benefits (lives saved) associated with the community-based misoprostol intervention (compared to the same strategy without the upgrades 1, 2, 3, and 4, respectively. bTransport encompasses the expedient availability of means of transport (e.g., vehicle, cart), fuel (if needed), driver, and interim attendant care. cQuality refers to the availability and quality of services at EmOC facilities, including adequate staffing and supplies, and evidence-based clinical practices. dCommunity-based interventions assume SBA-administered misoprostol for births at home and birthing centers/health centers with a 50% (25%–60%) reduction in PPH
[72]. ePopulation-level incremental benefits (lives saved) associated with the community-based misoprostol intervention (compared to the same strategy without the
community-based misoprostol intervention). These were calculated by applying model-projected outcomes to population-level data from rural India [87]. Cost-savings
for a single birth cohort of 15-y-old girls (2010) expected to accrue over their reproductive lifespan (age 15–45) ranged from US$128 million to US$190 million. doi:10.1371/journal.pmed.1000264.t006 [72]. ePopulation-level incremental benefits (lives saved) associated with the community-based misoprostol intervention (compared to the same strategy without the
community-based misoprostol intervention). These were calculated by applying model-projected outcomes to population-level data from rural India [87]. Cost-savings
for a single birth cohort of 15-y-old girls (2010) expected to accrue over their reproductive lifespan (age 15–45) ranged from US$128 million to US$190 million. doi:10.1371/journal.pmed.1000264.t006 PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 13 Reducing Maternal Mortality in India Reducing Maternal Mortality in India Reducing Maternal Mortality in India 17. Adam T, Lim SS, Mehta S, Bhutta ZA, Fogstad H, et al. (2005) Cost
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deaths in rural Rajasthan, India: exploring causes, context, and care-seeking
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36. Dolea C, AbouZahr C (2003) Global burden of hypertensive disorders
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hypertensivedisordersofpregnancy.pdf. Accessed 23 August 2009. 65. World Bank (2002) India’s transport sector: the challenges ahead. Vol 1. Washington (D.C.): World Bank Group. Available: http://www-wds. worldbank.org/external/default/main?pagePK = 64193027&piPK=64187937
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2009. 66. PMGSY (Pradhan Mantri Gram Sadak Yojana) (2006) Briefing book. Available: http://pmgsy.nic.in/. Accessed 27 July 27 2009. 38. AbouZahr C, Wardlaw T (2004) Maternal mortality in 2000: estimates developed
by WHO, UNICEF and UNFPA. Geneva: World Health Organization. Available: http://www.who.int/reproductivehealth/publications/monitoring/
9241562706/en/index.html. Accessed 23 August 2009. 67. Katrak H (2008) Measuring the shortage of medical practitioners in rural and
urban areas in developing countries: a simple framework and simulation
exercise with data from India. Int J Health Plann Manage 23: 93–105. 9241562706/en/index.html. Accessed 23 August 2009. 39. Henshaw SK, Singh S, Haas T (1999) The incidence of abortion worldwide. Int Fam Plann Persp 25: S30–S38. 68. Ma S, Sood N A comparison of the health systems in China and India. Rand
Corporation. Available: http://www.rand.org/pubs/occasional_papers/2008/
RAND_OP212.pdf. Accessed 27 July 2009. 40. Shah I, Ahman E (2004) Age patterns of unsafe abortion in developing country
regions. Reprod Health Matters 12: S9–S17. PLoS Medicine | www.plosmedicine.org PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 April 2010 | Volume 7 | Issue 4 | e1000264 15 Editors’ Summary Background. Every year, more than half a million women—
most of them living in developing countries—die from
pregnancy- or childbirth-related complications. About a
quarter of these ‘‘maternal’’ deaths occur in India. In 2005,
a woman’s lifetime risk of maternal death in India was 1 in
70; in the UK, it was only one in 8,200. Similarly, the maternal
mortality ratio (MMR; number of maternal deaths per
100,000 live births) in India was 450, whereas in the UK it
was eight. Faced with the enormous maternal death toll in
India and other developing countries, in September 2000,
the
United
Nations
pledged,
as
its
fifth
Millennium
Development Goal (MDG 5), that the global MMR would
be reduced to a quarter of its 1990 level by 2015. Currently, it
seems unlikely that this target will be met. Between 1990
and 2005, global maternal deaths decreased by only 1% per
annum instead of the 5% needed to reach MDG 5; in India,
the decrease in maternal deaths between 1990 and 2005 was
about 1.8% per annum. alone would reduce maternal deaths by 35% at most, so the
researchers also used their model to test the effect of
combinations of strategies on maternal death. They found
that an integrated and stepwise approach (increased family
planning and safe abortion combined with consecutively
increased skilled birth attendants, improved care before and
after birth, reduced home births, and improved emergency
obstetric care) could eventually prevent nearly 80% of
maternal deaths. All the steps in this strategy either saved
money or involved an additional cost per year of life saved of
less than US$500; given one suggested threshold for cost-
effectiveness in India of the per capita GDP (US$1,068) per
year of life saved, these strategies would be considered very
cost-effective. What Do These Findings Mean? The accuracy of these
findings depends on the assumptions used to build the
model and the quality of the data fed into it. Nevertheless,
these findings suggest that early intensive efforts to improve
family planning and to provide safe abortion accompanied
by a systematic, stepwise effort to improve integrated
maternal health services could reduce maternal deaths in
India by more than 75% in less than a decade. Furthermore,
such a strategy would be cost-effective. Editors’ Summary Indeed, note the
researchers, the cost savings from an initial focus on family
planning and safe abortion provision would partly offset the
resources needed to assure that every woman had access to
high quality routine and emergency obstetric care. Thus,
overall, these findings suggest that MDG 5 may be within
reach in India, a conclusion that should help to mobilize
political support for this worthy goal. Why Was This Study Done? Most maternal deaths in
developing countries are caused by severe bleeding after
childbirth, infections soon after delivery, blood pressure
disorders during pregnancy, and obstructed (difficult) labors. Consequently, experts agree that universal access to high-
quality routine care during labor (‘‘obstetric’’ care) and to
emergency obstetrical care is needed to reduce maternal
deaths. However, there is less agreement about how to
adapt these ‘‘ideal recommendations’’ to specific situations. In developing countries with weak health systems and
predominantly rural populations, it is unlikely that all women
will have access to emergency obstetric care in the near
future—so would beginning with improved access to family
planning and to safe abortions (unsafe abortion is another
major cause of maternal death) be a more achievable, more
cost-effective way of reducing maternal deaths? How would
family planning and safe abortion be coupled efficiently and
cost-effectively with improved access to intrapartum care? In
this study, the researchers investigate these questions by
estimating the health and economic outcomes of various
strategies to reduce maternal mortality in India. Additional Information. Please access these Web sites via
the online version of this summary at http://dx.doi.org/10. 1371/journal.pmed.1000264. N UNICEF (the United Nations Children’s Fund) provides
information on maternal mortality, including the WHO/
UNICEF/UNFPA/The World Bank 2005 country estimates of
maternal mortality N The World Health Organization also provides information on
maternal health and about MDG 5 (in several languages) What Did the Researchers Do and Find? The researchers
used a computer-based model that simulates women through
pregnancy and childbirth to estimate the effect of different
strategies (for example, increased family planning or increased
access to obstetric care) on clinical outcomes (pregnancies,
live births, or deaths), costs, and cost-effectiveness (the cost of
saving one year of life) in India. Increased family planning was
the most effective single intervention for the reduction of
pregnancy-related mortality. Reducing Maternal Mortality in India Shiffman J, Smith S (2007) Generation of political priority for global health
initiatives: a framework and case study of maternal mortality. Lancet 370:
1370–1379. 83. Mavalankar DV, Vora KS, Ramani KV, Raman P, Sharma B, et al. (2009)
Maternal health in Gujarat, India: a case study. J Health Popul Nutr 27:
235–248. 102. Filippi V, Ronsmans C, Campbell OM, Graham WJ, Mills A, et al. (2006)
Maternal health in poor countries: the broader context and a call for action. Lancet 368: 1535–1541. 84. AMDD Working Group on Indicators (2002) Program note: using UN process
indicators to assess needs in emergency obstetric services: Bhutan, Cameroon
and Rajasthan, India. Int J Gynaecol Obstet 77: 277–284. 103. National Rural Health Mission (2005–2012) Mission document. Available:
http://india.gov.in/outerwin.php?id = http://mohfw.nic.in/NRHM/
NRHM%28National%20Rural%20Health%20Mission%29.htm. Accessed 1
September 2009. j
J
y
85. World Health Organization (WHO) (2006) World Health Report: working
together for health. Geneva: World Health Organization. Available: http://
www.who.int/whr/2006/en/index.html. Accessed 23 August 2009. 104. Padmanaban P, Raman PS, Mavalankar DV (2009) Innovations and
challenges in reducing maternal mortality in Tamil Nadu, India. J Health
Popul Nutr 27: 202–219. 86. UNICEF, WHO, UNFPA (2007) Maternal mortality declining in middle-
income countries; women still die in pregnancy and childbirth in low-income
countries. UNICEF, WHO, UNFPA Joint Press Release. Available: http://
www.unicef.org/media/media_41208.html. Accessed 27 July 2009. 105. Prakasamma M (2009) Maternal mortality-reduction programme in Andhra
Pradesh. J Health Popul Nutr 27: 220–234. 87. United Nations (UN), Department of Economic and Social Affairs, Population
Division (2007) World Population Prospects: the 2006 revision. CD-ROM 106. McPake B, Koblinsky M (2009) Improving maternal survival in South Asia-
what can we learn from case studies? J Health Popul Nutr 27: 93–107. PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 16 Reducing Maternal Mortality in India Editors’ Summary Editors’ Summary If the current unmet need for
family planning in India could be fulfilled over the next 5 years,
more than 150,000 maternal deaths would be prevented,
more than US$1 billion saved, and at least half of abortion-
related deaths averted. However, increased family planning N The United Nations Millennium Development Goals Web site
provides detailed information about the Millennium Decla-
ration, the MDGs, their targets and their indicators, and about
MDG 5 N The Millennium Development Goals Report 2009
and its
progress chart provide an up-to-date assessment of progress
toward all the MDGs N Computer simulation modeling as applied to health is further
discussed at the Center for Health Decision Science at Harvard
University PLoS Medicine | www.plosmedicine.org April 2010 | Volume 7 | Issue 4 | e1000264 PLoS Medicine | www.plosmedicine.org 17
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https://www.frontiersin.org/articles/10.3389/fncel.2023.1277729/pdf?isPublishedV2=False
|
English
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Live-cell imaging of endocytosed synaptophysin around individual hippocampal presynaptic active zones
|
Frontiers in cellular neuroscience
| 2,023
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cc-by
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OPEN ACCESS OPEN ACCESS
EDITED BY
Chung Yu (Keith) Chan,
National Institute of Health, United States
REVIEWED BY
Lisi Wei,
National Institutes of Health (NIH),
United States
Kohgaku Eguchi,
Okinawa Institute of Science and Technology
Graduate University, Japan
*CORRESPONDENCE
Hiromitsu Tanaka
hiromitsu.tanaka@chiba-u.jp
RECEIVED 15 August 2023
ACCEPTED 06 October 2023
PUBLISHED 19 October 2023
CITATION
Tanaka H, Funahashi J and Hirano T (2023)
Live-cell imaging of endocytosed
synaptophysin around individual hippocampal
presynaptic active zones. Front. Cell. Neurosci. 17:1277729. doi: 10.3389/fncel.2023.1277729 EDITED BY
Chung Yu (Keith) Chan,
National Institute of Health, United States Hiromitsu Tanaka 1,2*, Junichiro Funahashi 1 and Tomoo Hirano 1
1 Department of Biophysics, Graduate School of Science, Kyoto University, Kyoto, Japan, 2 Department
of Developmental Biology, Graduate School of Medicine, Chiba University, Chiba, Japan Hiromitsu Tanaka 1,2*, Junichiro Funahashi 1 and Tomoo Hirano 1 Hiromitsu Tanaka 1,2*, Junichiro Funahashi 1 and Tomoo Hirano 1 Hiromitsu Tanaka 1,2*, Junichiro Funahashi 1 and Tomoo Hirano 1
1 Department of Biophysics, Graduate School of Science, Kyoto University, Kyoto, Japan, 2 Department
of Developmental Biology, Graduate School of Medicine, Chiba University, Chiba, Japan 1 Department of Biophysics, Graduate School of Science, Kyoto University, Kyoto, Japan, 2 Department
of Developmental Biology, Graduate School of Medicine, Chiba University, Chiba, Japan In presynaptic terminals 4 types of endocytosis, kiss-and-run, clathrin-mediated,
bulk and ultrafast endocytosis have been reported to maintain repetitive
exocytosis of neurotransmitter. However, detailed characteristics and relative
contribution of each type of endocytosis still need to be determined. Our
previous live-cell imaging study demonstrated individual exocytosis events of
synaptic vesicle within an active-zone-like membrane (AZLM) formed on glass
using synaptophysin tagged with a pH-sensitive fluorescent protein. On the other
hand, individual endocytosis events of postsynaptic receptors were recorded
with a rapid extracellular pH exchange method. Combining these methods, here
we live-cell imaged endocytosed synaptophysin with total internal reflection
fluorescence microscopy in rat hippocampal culture preparations. Clathrin-
dependent and -independent endocytosis, which was seemingly bulk endocytosis,
occurred within several seconds after electrical stimulation at multiple locations
around AZLM at room temperature, with the locations varying trial to trial. The
contribution of clathrin-independent endocytosis was more prominent when
the number of stimulation pulses was large. The skewness of synaptophysin
distribution in intracellular vesicles became smaller after addition of a clathrin
inhibitor, which suggests that clathrin-dependent endocytosis concentrates
synaptophysin. OPEN ACCESS Ultrafast endocytosis was evident immediately after stimulation
only at near physiological temperature and was the predominant endocytosis
when the number of stimulation pulses was small. COPYRIGHT
© 2023 Tanaka, Funahashi and Hirano. This is
an open-access article distributed under the
terms of the Creative Commons Attribution
License (CC BY). The use, distribution or
reproduction in other forums is permitted,
provided the original author(s) and the
copyright owner(s) are credited and that the
original publication in this journal is cited, in
accordance with accepted academic practice. No use, distribution or reproduction is
permitted which does not comply with these
terms. synaptic vesicle, endocytosis, clathrin, TIRFM, synaptophysin (SYN), cytomatrix at the
active zone-associated structural protein (CAST), active zone (AZ) synaptic vesicle, endocytosis, clathrin, TIRFM, synaptophysin (SYN), cytomatrix at the
active zone-associated structural protein (CAST), active zone (AZ) TYPE Original Research
PUBLISHED 19 October 2023
DOI 10.3389/fncel.2023.1277729 TYPE Original Research
PUBLISHED 19 October 2023
DOI 10.3389/fncel.2023.1277729 frontiersin.org Primary cell culture and transfection The methods for preparing the primary culture of
hippocampal neurons and transfection of cDNA were described
previously (Tanaka and Hirano, 2012; Tanaka et al., 2014). Briefly,
hippocampi were dissected out from E18-P0 Wistar rat embryos,
treated with 0.25% trypsin (Thermo Scientific, 15090-046) and
dissociated by trituration with a fire-polished Pasteur pipette. Dissociated cells were seeded on poly-D-lysine-(Merck, P7280
and P6407) and NLG-coated glass (Figure 1A) in Neurobasal
medium (Thermo Scientific, 21103-049) containing 1%
penicillin–streptomycin (Thermo Scientific, 15140-122), 500 μM
glutamine (Merck, G6392) and 2% B27 Supplement (Thermo
Scientific, 17504-044). In the present study, we used total internal reflection fluorescence
microscopy (TIRFM), which allows the observation of fluorescent
molecules with a high signal-to-noise ratio by limiting the depth of
the excitation field to approximately 100–200 nm (Axelrod, 2001;
Toomre and Manstein, 2001), to improve the quality of the fluorescent
images of endocytosed presynaptic proteins. In our previous studies,
to efficiently visualize fluorescent molecules at active zones with
TIRFM, we induced the formation of active zone-like membranes
(AZLM) parallel to the glass surface by coating with neuroligin
(NLG). This synaptic adhesion molecule induces presynaptic
differentiation through binding with presynaptic neurexin (NRX)
(Funahashi et al., 2018). With this setup, we recorded single
exocytosis events using a synaptic vesicle protein, synaptophysin
(Syp), tagged with a pH-sensitive variant of green fluorescent protein
(GFP) called super-ecliptic pHluorin (SEP). SEP is non-fluorescent
in the low pH intra-vesicular solution but becomes fluorescent after
exposure to the neutral pH of the extracellular solution (Miesenböck
et al., 1998; Kavalali and Jorgensen, 2013). We also visualized
endocytosis events of glutamate receptors labeled with SEP in the
postsynaptic membrane (Fujii et al., 2017, 2018). In those studies, the
extracellular pH was changed intermittently and locally to an acidic
condition using a U-tube system. At pH 6.0, cell-surface SEP signals
were quenched, and only signals could be detected from recently
endocytosed vesicles in which acidification had not been completed. pH-sensitive fluorophores and extracellular pH exchange have been
used to study endocytic processes (Merrifield et al., 2005; Rosendale
et al., 2017). Combining TIRFM, AZLM formation and rapid pH
exchange using U-tube, in the present study, we observed
endocytosed Syp-SEP signals after an electrical field stimulation to
trigger the exocytosis of synaptic vesicles and analyzed the
distribution patterns of the endocytosed Syp-SEP signals around
AZLM at several time points. DNA constructs Expression vectors for rat synaptophysin labeled with SEP (Syp-
SEP), cytomatrix at the active zone-associated protein (CAST) labeled
with TagRFPt (CAST-RFP), neuroligin1 with splice insertion A
labeled with human immunoglobulin-Fc region (NLG-Fc) and
neurexin1β without splice insertion 4 labeled with HA tag (NRX) were
prepared as described previously (Funahashi et al., 2018). Animals All experimental procedures were carried out in accordance with
the National Institute of Health guide for the care and use of laboratory
animals and the ethical guidelines on animal experimentation of
Kyoto University, and were approved by the local committee for
handling experimental animals in the Graduate School of Science,
Kyoto University. Each of the above methods to study presynaptic endocytosis has
merits and demerits. EM captures images with ultra-high resolution
but at only one-time point for each image (Watanabe et al., 2013,
2014; Jin et al., 2019). Recording the membrane capacitance
measures the change of the surface area of the presynaptic terminal
with high temporal resolution (Delvendahl et al., 2016; Jin et al.,
2019), but it does not provide information about the location or
morphological properties of each endocytosis type. Live-cell
fluorescence imaging records the movement and location of
released synaptic vesicle proteins even at a single synapse, and
imaging trials can be repeated in the same cell, but this approach
has limited spatial resolution compared with EM and low time
resolution compared with electrophysiological capacitance
measurements. Overall, the distribution of each endocytosis type
around individual active zones has not been demonstrated at
multiple time points in living neurons.ll Primary cell culture and transfection We show the distributions of Syp-SEP
during and after exo- and endocytosis around AZLM and analyze the
characteristics of each type of endocytosis. Materials and methods Materials and methods activation of a presynaptic neuron, and UFE occurs immediately after
the presynaptic exocytosis (Cheung and Cousin, 2013; Watanabe
et al., 2013; Soykan et al., 2017). Kiss-and-run endocytosis (K&R), in
which the vesicle and cell membrane fuse transiently has also been
reported (Zhang et al., 2009; Ge et al., 2022), although its existence in
neuronal presynaptic terminals is controversial (Granseth et al., 2006;
Wu et al., 2014). Materials and methods
Animals Materials and methods Introduction In presynaptic terminals, the rapid and repetitive exocytosis of synaptic vesicles occurs. Because the number of synaptic vesicles is limited, efficient synaptic vesicle retrieval processes
including endocytosis are necessary. Morphological observations using electron microscopy
(EM), electrophysiological capacitance measurements of the cell-surface area and live-cell
imaging have been performed (Jin et al., 2019), and four types of endocytosis have been
reported. After the full-fusion type of exocytosis, clathrin-mediated endocytosis (CME) and
clathrin-independent one occurs in the vicinity of an active zone (Granseth et al., 2006;
Chanaday et al., 2019). The latter include activity-dependent bulk endocytosis (ADBE) and
ultrafast-type endocytosis (UFE). In ADBE a large endosome is produced after repetitive Frontiers in Cellular Neuroscience Frontiers in Cellular Neuroscience 01 Tanaka et al. 10.3389/fncel.2023.1277729 Glass coating and AZLM formation FIGURE 1
Experimental scheme and system. (A) Schematic diagram of AZLM and pH exchange method. Arrows around top right U-tube indicate directions of
solution flow. (B) Experimental design. An electrical field stimulation (50 pulses, 50 Hz) was applied to induce the exocytosis of synaptic vesicles
containing Syp-SEP [① in (A,B)]. The endocytosed Syp-SEP was observed at each time point by repeating a pH exchange from 7.4 to 6.0 [② in (A,B)]. The
green two-way arrow corresponds to the amount of endocytosed Syp-SEP. (C) The pH-exchanging speed. The right graph shows an enlargement of
the region enclosed by the dotted yellow line. (D) Observation area of TIRFM in the Z-axis. The tip of a glass electrode containing CF488 dye (left) was
observed under TIRFM. The middle and right TIRFM images are enlargements of the yellow-dotted square region in the left bright field + fluorescence
i
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d 250
f
th
l
f
A
i di
t th
l
t
d ti
Th l
th
t
t
f th
i
li
ti
d
th
f TIRFM FIGURE 1
Experimental scheme and system. (A) Schematic diagram of AZLM and pH exchange method. Arrows around top right U-tube indicate directions of
solution flow. (B) Experimental design. An electrical field stimulation (50 pulses, 50 Hz) was applied to induce the exocytosis of synaptic vesicles
containing Syp-SEP [① in (A,B)]. The endocytosed Syp-SEP was observed at each time point by repeating a pH exchange from 7.4 to 6.0 [② in (A,B)]. The
green two-way arrow corresponds to the amount of endocytosed Syp-SEP. (C) The pH-exchanging speed. The right graph shows an enlargement of
the region enclosed by the dotted yellow line. (D) Observation area of TIRFM in the Z-axis. The tip of a glass electrode containing CF488 dye (left) was
observed under TIRFM. The middle and right TIRFM images are enlargements of the yellow-dotted square region in the left bright field + fluorescence
image at 0 and 250 nm from the glass surface. Arrows indicate the electrode tip. The length constant of the visualization depth of TIRFM was
calculated to be 151 nm (right). (E) Single molecule analysis of SEP. Sequential images of the SEP signal recorded every 120 ms with TIRFM (top). The
time course of the SEP fluorescence intensity, showing single-step photobleaching characteristic of a single molecule (bottom left). Distribution of
single SEP fluorescence intensities at PT (bottom middle) and RT (bottom right). Glass coating and AZLM formation NLG-Fc was prepared from transfected HEK293 cells using
nProtein A Sepharose (GE Healthcare, 17-5280-01). A piece of
cover glass was first incubated with 43 μM biotinylated bovine
serum albumin (Thermo Scientific, 29130) in buffer A (100 mM
KCl, 5 mM MgCl2, 25 mM HEPES-KOH, pH7.4) at 4°C overnight
(Tanaka et al., 2014). Then, the glass was incubated in buffer A
containing 17 mM streptavidin (Wako, 191-12851) for 1 h, followed
by incubation in buffer A containing biotin-conjugated anti-human
IgG (1:100; Jackson ImmunoResearch, 109-065-098) for 1 h at
RT. Then, the glass was washed and further incubated in buffer A
containing 3–5 mg/mL NLG-Fc for 5 h. Expression vectors for
Syp-SEP, CAST-RFPt and NRX were transfected into neurons after
10–15 days in vitro (DIV) with Lipofectamine 2000 (Thermo
Scientific, 11668-019) (Funahashi et al., 2018). Half of the medium
was replaced with fresh prewarmed medium 4–6 h after the
transfection. AZLM was formed, and all imaging experiments were
carried out 2–3 days after the transfection. Frontiers in Cellular Neuroscience 02 frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 1
Experimental scheme and system. (A) Schematic diagram of AZLM and pH exchange method. Arrows around top right U-tube indicate directions of
solution flow. (B) Experimental design. An electrical field stimulation (50 pulses, 50 Hz) was applied to induce the exocytosis of synaptic vesicles
containing Syp-SEP [① in (A,B)]. The endocytosed Syp-SEP was observed at each time point by repeating a pH exchange from 7.4 to 6.0 [② in (A,B)]. The
green two-way arrow corresponds to the amount of endocytosed Syp-SEP. (C) The pH-exchanging speed. The right graph shows an enlargement of
the region enclosed by the dotted yellow line. (D) Observation area of TIRFM in the Z-axis. The tip of a glass electrode containing CF488 dye (left) was
observed under TIRFM. The middle and right TIRFM images are enlargements of the yellow-dotted square region in the left bright field + fluorescence
image at 0 and 250 nm from the glass surface. Arrows indicate the electrode tip. The length constant of the visualization depth of TIRFM was
calculated to be 151 nm (right). (E) Single molecule analysis of SEP. Sequential images of the SEP signal recorded every 120 ms with TIRFM (top). The
time course of the SEP fluorescence intensity, showing single-step photobleaching characteristic of a single molecule (bottom left). Distribution of
single SEP fluorescence intensities at PT (bottom middle) and RT (bottom right). Image analysis Acquired images were analyzed using MetaMorph (Molecular
Devices), R (The R Project) and Excel (Microsoft). AZLM area was
determined as follows. First, the CAST-RFP image was compensated
for mechanical drifts of the stage. Then, the mean plus 3 times SD
of the background signal intensity in each image was set as the
threshold. The CAST-labeled area ranging from 0.098 to 0.38 μm2
was defined as AZLM. The active zone area in the hippocampus
estimated by previous EM studies is <0.18 μm2 (mean 0.04 μm2)
(Schikorski and Stevens, 1997; Holderith et al., 2012). However,
we recorded a fluorescence signal from a single SEP molecule in
several pixels due to diffraction and scattering (Figure 1E). Therefore, we set the AZLM area slightly larger. Analysis was
performed on AZLMs which showed significant Syp-SEP signal
increase by 5 or 50 pulses stimulation.hl The Syp-SEP area reflecting exocytosis or endocytosis was
determined as follows. After drift compensation (Tanaka et al.,
2014), the fluorescence values of Syp-SEP were compensated for
bleaching. The bleaching rate was computed by fitting the bleach
curve with a double exponential function obtained from
experiments without stimulation. Then, the mean plus 3 times SD
of the background signal in each image was set as the threshold. Areas (> 0.098 μm2) of contiguous pixels above the threshold were
defined as the Syp-SEP positive area. The centroid of the
fluorescence intensity of the Syp-SEP-positive area was regarded as
the center of Syp-SEP-containing vesicles. For a detailed analysis of the signal intensity, area and position
of endocytosed Syp-SEP, the average of 4 consecutive high-
magnification images taken from the time when the extracellular
pH was changed to 6.0 was used. For example, when the pH
exchange to 6.0 started 3 s after the stimulation, Syp-SEP images
taken at 3.24, 3.36, 3.48 and 3.60 s were averaged and used for the
quantitative analyses at 3.24 s. To exclude the Syp-SEP signal from
that of organelles such as the endoplasmic reticulum, each image
was subtracted from the image recorded at pH 6.0 at 3.88 s before
the stimulation. If an increase in the signal was observed from the
background level while the extracellular pH was 6.0, it was assumed
that Syp migrated from outside the TIRFM observation area within
the presynaptic terminal, and the corresponding data was excluded
from the analysis. When multiple Syp-SEP containing vesicles were
in close proximity, it was impossible to estimate each center. Glass coating and AZLM formation 03 Frontiers in Cellular Neuroscience frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 10.3389/fncel.2023.1277729 Image analysis However, we tried to separate a contiguous Syp-SEP positive area
containing multiple intensity peaks using watershed lines
(Broadhead et al., 2016; Imig et al., 2020). Watershed lines were
drawn to connect local minima that existed between peaks
using MetaMorph. To measure the observation depth of TIRFM, the tip of the glass
electrode containing CF488 (Nacalai testque, 20 μM) was raised in
the Z-axis direction at 125 nm steps from the glass surface using a
micro manipulator (HEKA, MP-285). The fluorescence signal
decreased as the glass tip was raised with a length constant of
151 nm (Figure 1D). The same laser intensity and illumination angle were used to
record the fluorescence of single SEP molecules. SEP molecules
were prepared from SEP-transfected HEK293 cells by supersonic
treatment and diluted with PBS. A single SEP molecule
on the glass was identified by single-step photobleaching
(Figure 1E). Live-cell imaging and electrical field
stimulation The intra-U-tube solution was the same as the extracellular solution,
except that the pH was adjusted to 6.0 with 2-(N-morpholino)
ethanesulfonic acid and KOH. The intra-U-tube solution was the same as the extracellular solution,
except that the pH was adjusted to 6.0 with 2-(N-morpholino)
ethanesulfonic acid and KOH. The TIRFM imaging system was composed of an inverted
fluorescence microscope (Olympus, IX71) equipped with a 100×
(NA 1.49) or 150× (NA 1.45) TIRFM objective lens and 1.6×
intermediate lens, EM-CCD camera (Andor, iXonEM+ DU-897),
488 nm laser (Melles Griot, 85-BCD-020) and 561 nm laser
(Coherent, Sapphire 561LP). Changes in Syp-SEP signals before and
after the electrical stimulation were recorded with the camera using
a 40 ms exposure time and the 100× objective lens. When we wanted
higher resolution images, Syp-SEP and CAST-RFP images were
acquired for 120 ms with the 150× objective lens and 1.6×
intermediate lens using a dual-color filter (Semrock, FF01-523/610-
25). Imaging was performed in extracellular solution (120 mM
NaCl, 4 mM CaCl2, 1 mM MgCl2, 10 mM glucose, 10 mM
HEPES-KOH, pH 7.3) at RT or PT. The temperature of the
experimental room was controlled by air conditioning. The pH was
changed intermittently, and the duration of the pH 6.0 condition
was 700 ms. Within this pH 6.0 period we recorded 5 high resolution
images and the later 4 were used to generate an averaged image. An
electrical field stimulation (duration 1 ms, 20–26 V/cm, 5, 20, 50 or
125 pulses at 50 Hz) was applied to the cultured neurons between
platinum electrodes (Tanaka et al., 2014). To observe clathrin-
independent endocytosis, pitstop 2 (pit2) was added at a final
concentration of 10 μM, and imaging was performed 10 min later. Pit2 was used here to inhibit clathrin-dependent processes, because
it was possible to record Syp-SEP images in a same preparation
before and after its application. Previous studies also used
knockdown of clathrin by RNAi or overexpression of a dominant
negative form of clathrin-related protein (Granseth et al., 2006), but
these manipulations take much longer time and recording
endocytosis before and after the manipulation in a same cell is very
difficult if not impossible. Frontiers in Cellular Neuroscience Location of endocytosed Syp-SEP signals Next, we imaged endocytosed Syp-SEP after the rapid
extracellular pH change to 6.0 at a higher magnification using 150×
objective lens to examine where endocytosed Syp-SEP was localized
in relation to AZLM (Figure 3A). Line scans across CAST-RFP and
Syp-SEP signals from a representative case showed that the position
of endocytosed Syp-SEP signal peaks at 3.24 and 7.24 s after the
stimulation were about 170 nm from the CAST-RFP signal peak
(Figures 3A,B). However, Syp-SEP signals at pH 6.0 were scattered,
and there were multiple intensity peaks in several trials. In such
cases, we attempted to divide the Syp-SEP areas by watershed lines
(Figure 3C). The centroid of the Syp-SEP signal after the stimulation
at pH7.4, which reflects the exocytosis of a large number of synaptic
vesicles, was about 86 ± 13 nm (n = 45 clusters) inside of the edge of
the CAST-RFP signal, whereas that at pH 6.0, which reflects
endocytosis, was about 100 ± 28 nm (n = 69 clusters) and 73 ± 27 nm
(n = 62 clusters) outside of the edge of the CAST-RFP signal at 3.24 s
and 7.24 s after the stimulation (Figure 3D). The latter two values
were significantly different from the former (p < 0.001, Steel-Dwass’s
test). Thus, Syp-SEP signals at pH 6.0, which reflect endocytosis-
related vesicles, were found in the periphery of AZLM. The centroid
of the Syp-SEP signal at pH 6.0 does not always correspond to the
center of an endocytosed vesicle, because the Syp-SEP signal might
have come from multiple vesicles that were not separated by the
watershed lines. On the other hand, we previously reported that
Syp-SEP is released inside AZLM by an electric field stimulation
(Funahashi et al., 2018), which is consistent with the present results. Additionally, the CAST-RFP signal was stable and moved little An electric field stimulation (50 Hz, 50 pulses) was applied to
cultured neurons so that a large number of synaptic vesicles were
fused to AZLM at RT, and an apparent increase in the Syp-SEP signal
was detected (Figure 2A) (Funahashi et al., 2018). When the number
of stimulation pulses was reduced from 50 to 20 or 5, the increase in
the Syp-SEP signal decreased in proportion to the number of
stimulation pulses (p < 0.05 and 0.001, Steel’s test). U-tube system The U-tube system was prepared as described previously
(Bretschneider and Markwardt, 1999; Fujii et al., 2017). To improve
the speed of the external fluid exchange (Figure 1C), the inner
diameter of the U-tube hole was made smaller (6–8 μm), and the
hole was placed 25–50 μm above the glass surface. Using the U-tube
system, the pH of extracellular solution was changed from 7.4 to
6.0 in about 100 ms and from 6.0 to 7.4 in about 200 ms (Figure 1C). For the analysis of exocytosed Syp-SEP signal intensity, images
recorded at 120 ms after the end of the stimulation at pH 7.4 were
used. The position of exocytosis indicates the center of the entire
Syp-SEP positive area, where multiple vesicles were fused to the
plasma membrane, 120 ms after the onset of the stimulation. 04 frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 Frontiers in Cellular Neuroscience Visualization of Syp-SEP around AZLM after
presynaptic activation To visualize Syp-SEP translocated to the plasma membrane
upon the arrival of action potentials and also endocytosed Syp-SEP
in presynaptic terminals, we applied TIRFM (Figure 1A). We cultured rat hippocampal neurons on NLG-coated glass and
induced AZLM formation through NLG binding to presynaptic
NRX at DIV 12–18 (Funahashi et al., 2018). Neurons were
transfected with CAST labeled with tagRFPt (CAST-RFP), Syp-SEP
and NRX. CAST-RFP was used as an AZLM marker (Ohtsuka et al.,
2002). Then, the U-tube system was applied to this experimental
system (Fujii et al., 2017) to transiently and locally change the
extracellular pH from 7.4 to 6.0 around AZLM, thus quenching SEP
signals on the cell surface, and internalized Syp-SEP signals by
endocytosis were detected (Figures 1A–C). The extracellular pH was
changed from 7.4 to 6.0 in about 100 ms and from 6.0 to 7.4 in about
200 ms (Figure 1C). The length constant of the visualization depth
of the TIRFM was 151 nm (Figure 1D). Therefore, the internalized
Syp-SEP signals should have come from a region about 0–200 nm
above the glass surface. We also evaluated the sensitivity of the
recording system by recording fluorescence from a single SEP
molecule on the glass surface (Figure 1E). A single SEP molecule was
identified by one-step bleaching, and the fluorescence intensity was
2,900 ± 77 (n = 114 points) at room temperature (RT, 21–23°C) and
2,200 ± 71 (n = 102 points) at near physiological temperature (PT,
31–32°C) on an arbitrary scale of our experimental system.i Experimental design and statistical
analysis exocytosis at 50 pulses, an observation consistent with a previous
report (Sakamoto et al., 2018). p
Endocytosis was likely to occur during the decay of the Syp-SEP
signal after exocytosis. To precisely visualize the endocytosed
Syp-SEP around AZLM, the pH of the extracellular solution was
changed to 6.0 repeatedly so that the fluorescence from cell-surface
Syp-SEP was quenched and that only the internalized Syp-SEP
signals were recorded (Figures 1A,B, 2B,C). The fluorescence signal
observed at pH 6.0 was more intense at 3.12, 7.12, 15.12 and 27.12 s
after the stimulation onset than before the stimulation (Figures 2C,D,
p < 0.001 and 0.01, Steel’s test), indicating that endocytosis primarily
took place at these times after exocytosis. About 40 s after the
stimulation, when the Syp-SEP fluorescence signal at pH 7.4
returned to the pre-stimulation level, the signal observed at pH 6.0
almost disappeared. The Syp-SEP signal at pH 7.4 increased and
decayed similarly, irrespective of the pH exchange (Figure 2C),
suggesting that the pH exchange did not significantly affect the
release and uptake of vesicles containing Syp-SEP. The SEP signal
intensity at pH 6.0 relative to that at pH 7.4 at 1 s after the stimulation
showed a near-linear relationship (Figure 2E). The intensity of
endocytosed Syp-SEP was positively correlated with the intensity of
exocytosed Syp-SEP. These results confirmed that the more Syp-SEP
is exocytosed, the more it is endocytosed. There was weak Syp-SEP
signal at pH 7.4 before the stimulation (Figure 2C, −4.04 and
−4.00 s). Thus, a small portion of the Syp-SEP signal might have
been derived from Syp-SEP present on the plasma membrane before
the stimulation. The sample size in each experiment was determined based on
previous publications dealing with live-cell imaging (Fujii et al.,
2018; Sposini et al., 2020). In each experiment, N indicates the
number of Syp-SEP or CAST-RFP signal clusters and/or cells except
in the single molecule fluorescence analysis. When we compared
means of two groups, Mann–Whitney U test or Wilcoxon signed-
rank test was used in a case the normality of data distribution and/
or equality of deviations were denied by Shapiro–Wilk test and/or
F-test, otherwise we performed Student’s paired t-test. In multiple
comparison of means, we used Steel’s test or Steel-Dwass’s test. All
values are presented as the mean ± standard error of the mean. Statistical tests were performed using Excel and Kyplot (KyensLab,
ver. 5.0.3). frontiersin.org Location of endocytosed Syp-SEP signals Conversely, the
Syp-SEP signal intensity did not increase proportionally when the
number of pulses was increased to 125 (Figure 2A), suggesting that
the amount of release was nearly saturated at 125 and that most of
the synaptic vesicles in a readily releasable pool underwent 05 Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 2
Visualization of endocytosed Syp-SEP. (A) Release of Syp-SEP induced by electrical field stimulation. In the left, averaged traces of Syp-SEP
fluorescence intensity (means ± standard errors) before and after 5 (11 cells), 20 (12 cells), 50 (16 cells) or 125 (23 cells) pulses stimulation are presented
with representative images of CAST-RFP (magenta) and Syp-SEP (green) before and after 50 pulses stimulation. In the right, the peak Syp-SEP signal
ntensities after 5, 20, 50 or 125 pulses stimulation normalized by the intensities before the stimulation are presented. The amount of exocytosed Syp-
SEP increased almost proportionally to the number of stimulation pulses (up to 50 pulses). (B) Representative images of Syp-SEP and CAST-RFP before
and after the 50 pulses stimulation at pH 7.4 and 6.0. Arrows indicate Syp-SEP signal at pH 6.0 observed only after the stimulation. (C) An example of
the signal intensity of Syp-SEP before and after the 50 pulses stimulation with (black) or without (yellow) pH 6.0 exchange (left graph). Kymograph
showing Syp-SEP with pH exchange is presented on the top. We set the timing of pH change to 6.0 so that whole time course of endocytosis
occurrence could be clarified and that later pH change to 6.0 would show the Syp-SEP endocytosed after the previous pH exchange, although
we know that the Syp-SEP signal at pH 6.0 detected at 7 s include some signal endocytosed before 3 s as described in results. The light blue trace
shows the Syp-SEP signal intensity without any stimulation. Right images show Syp-SEP fluorescence before (−) and after (+) the stimulation at pH 7.4
or 6.0. (D) Quantification of the Syp-SEP signal intensities at pH 6.0 at various times normalized by the intensity at pH 7.4 before the stimulation (3.12 s,
n = 43 clusters, 10 cells, 7.12 s, n = 43 clusters, 10 cells, 15.12 s, 40 clusters, 10 cells, 27.12 s, 39 clusters, 9 cells, 39.12 s, n = 41 clusters, 10 cells). Location of endocytosed Syp-SEP signals (E) Correlation of Syp-SEP signal intensities at pH 7.4 at 1 s after the stimulation and at pH 6.0 at 3.12, 7.12 or 15.12 s after the stimulation (3.12 s, r = 0.54,
7.12 s, r = 0.56, 15.12 s, r = 0.58), *, **, and *** indicates p < 0.05, 0.01, and 0.001 respectively, n.s., not significant. FIGURE 2
Visualization of endocytosed Syp-SEP. (A) Release of Syp-SEP induced by electrical field stimulation. In the left, averaged traces of Syp-SEP
fluorescence intensity (means ± standard errors) before and after 5 (11 cells), 20 (12 cells), 50 (16 cells) or 125 (23 cells) pulses stimulation are presented
with representative images of CAST-RFP (magenta) and Syp-SEP (green) before and after 50 pulses stimulation. In the right, the peak Syp-SEP signal
intensities after 5, 20, 50 or 125 pulses stimulation normalized by the intensities before the stimulation are presented. The amount of exocytosed Syp-
SEP increased almost proportionally to the number of stimulation pulses (up to 50 pulses). (B) Representative images of Syp-SEP and CAST-RFP before
and after the 50 pulses stimulation at pH 7.4 and 6.0. Arrows indicate Syp-SEP signal at pH 6.0 observed only after the stimulation. (C) An example of
the signal intensity of Syp-SEP before and after the 50 pulses stimulation with (black) or without (yellow) pH 6.0 exchange (left graph). Kymograph
showing Syp-SEP with pH exchange is presented on the top. We set the timing of pH change to 6.0 so that whole time course of endocytosis
occurrence could be clarified and that later pH change to 6.0 would show the Syp-SEP endocytosed after the previous pH exchange, although
we know that the Syp-SEP signal at pH 6.0 detected at 7 s include some signal endocytosed before 3 s as described in results. The light blue trace
shows the Syp-SEP signal intensity without any stimulation. Right images show Syp-SEP fluorescence before (−) and after (+) the stimulation at pH 7.4
or 6.0. (D) Quantification of the Syp-SEP signal intensities at pH 6.0 at various times normalized by the intensity at pH 7.4 before the stimulation (3.12 s,
n = 43 clusters, 10 cells, 7.12 s, n = 43 clusters, 10 cells, 15.12 s, 40 clusters, 10 cells, 27.12 s, 39 clusters, 9 cells, 39.12 s, n = 41 clusters, 10 cells). Frontiers in Cellular Neuroscience Location of endocytosed Syp-SEP signals (E) Correlation of Syp-SEP signal intensities at pH 7.4 at 1 s after the stimulation and at pH 6.0 at 3.12, 7.12 or 15.12 s after the stimulation (3.12 s, r = 0.54,
7.12 s, r = 0.56, 15.12 s, r = 0.58), *, **, and *** indicates p < 0.05, 0.01, and 0.001 respectively, n.s., not significant. 06 Frontiers in Cellular Neuroscience frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 3
Location of endocytosed Syp-SEP signal. (A) Example of endocytosed Syp-SEP visualized at high magnification using 150× objective lens. The right
images are averages of the images enclosed by the yellow lines and were merged with an image of CAST-RFP signals. (B) Line-scan of Syp-SEP or
CAST-RFP signal intensities on the yellow broken lines shown in (A). (C) Examples of binarized Syp-SEP signals and those separated by watershed
segmentation. Right most images show raw data with yellow lines indicating Syp-SEP positive areas after watershed segmentation. (D) Location of
centroids of the Syp-SEP signal positive areas at pH 7.4 at 0.12 s after the stimulation and at pH 6.0 at 3.24 or 7.24 s after the stimulation in relation to
the edge of CAST-RFP (0.12 s, n = 45 clusters, 3.24 s, 69 clusters, 7.24 s, 62 clusters, 13 cells). (E) Stability of CAST-RFP signal. Representative images of
CAST-RFP before and after the stimulation (top). Distribution of averaged CAST-RFP centroids at 10–14 s after the stimulation (black points) compared
with that at 1–5 s before the stimulation (0.0, red point) (bottom left) (n = 11 clusters, 4 cells). Displacement of CAST-RFP centroids after the stimulation
(bottom right). *** indicates p < 0.001. FIGURE 3
Location of endocytosed Syp-SEP signal. (A) Example of endocytosed Syp-SEP visualized at high magnification using 150× objective lens. The right
images are averages of the images enclosed by the yellow lines and were merged with an image of CAST-RFP signals. (B) Line-scan of Syp-SEP or
CAST-RFP signal intensities on the yellow broken lines shown in (A). (C) Examples of binarized Syp-SEP signals and those separated by watershed
segmentation. Right most images show raw data with yellow lines indicating Syp-SEP positive areas after watershed segmentation. frontiersin.org Location of endocytosed Syp-SEP signals (D) Location of
centroids of the Syp-SEP signal positive areas at pH 7.4 at 0.12 s after the stimulation and at pH 6.0 at 3.24 or 7.24 s after the stimulation in relation to
the edge of CAST-RFP (0.12 s, n = 45 clusters, 3.24 s, 69 clusters, 7.24 s, 62 clusters, 13 cells). (E) Stability of CAST-RFP signal. Representative images of
CAST-RFP before and after the stimulation (top). Distribution of averaged CAST-RFP centroids at 10–14 s after the stimulation (black points) compared
with that at 1–5 s before the stimulation (0.0, red point) (bottom left) (n = 11 clusters, 4 cells). Displacement of CAST-RFP centroids after the stimulation
(bottom right). *** indicates p < 0.001. found at different locations in different trials. The distance between
the centroid of the Syp-SEP signals in the first trial and the nearest
centroid in the second trial was 310 ± 42 nm (n = 21 clusters) at 3.24 s
and 340 ± 50 nm (n = 19 clusters) at 7.24 s (Figure 4B), although all of
the centroids were distributed in the periphery of AZLM. The average
distance between the centroids at 3.24 s and 7.24 s in the same trial
was 140 ± 18 nm (n = 29 clusters), which was smaller than the distance during the recording: the distance between the centroid of the
CAST-RFP signal before and after the stimulation was 30.8 ± 4.2 nm
(Figure 3E). It remains unclear whether synaptic vesicle proteins were
endocytosed in specialized regions. To examine if there were such
endocytic zones around an active zone, the same neuron was
repeatedly stimulated (Figure 4A). Syp-SEP signals at pH 6.0 were 07 frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 4
Locations of endocytosed Syp-SEP signals in different trials. (A) Representative high magnification images of Syp-SEP at pH 6.0 after the 50 pulses
stimulation in different trials using the same sample. The right images are averages of the images enclosed by the yellow lines and were merged with
an image of CAST-RFP signals. Arrows indicate positions of Syp-SEP around an AZLM. The signal locations were different between 1st and 2nd trials. (B) Displacement of centroids of Syp-SEP signals at pH 6.0 in different trials (3.24 s, 7.24 s) or over a time range (3.24–7.24 s) in a trial (3.24 s, n = 21
clusters, 7.24 s, 19 clusters, 3.24–7.24 s, 29 clusters, 11 cells). Location of endocytosed Syp-SEP signals ** indicate p < 0.01. GURE 4 Locations of endocytosed Syp-SEP signals in different trials. (A) Representative high magnification images of Syp-SEP at pH 6.0 after the 50 pulses
stimulation in different trials using the same sample. The right images are averages of the images enclosed by the yellow lines and were merged with
an image of CAST-RFP signals. Arrows indicate positions of Syp-SEP around an AZLM. The signal locations were different between 1st and 2nd trials. (B) Displacement of centroids of Syp-SEP signals at pH 6.0 in different trials (3.24 s, 7.24 s) or over a time range (3.24–7.24 s) in a trial (3.24 s, n = 21
clusters, 7.24 s, 19 clusters, 3.24–7.24 s, 29 clusters, 11 cells). ** indicate p < 0.01. for different trials (p < 0.01, Steel-Dwass’s test). These results suggest
that endocytosis locations change in different trials and that a part of
the Syp-SEP signal recorded at 7.24 s might have been derived from
Syp-SEP endocytosed before 3.24 s. pitstop2 (pit2) at RT (Figures 5A, 6A). When pit2 was added to the
extracellular solution, the Syp-SEP signal intensity at pH 6.0 after the
stimulation became smaller: at 3.24 s, the intensity was 20,700 ± 3,600
(n = 26 clusters) without pit2 and 12,200 ± 1700 with pit2 (n = 25
clusters, p < 0.05, Mann–Whitney U test); at 7.24 s, it was 16,300 ± 2,200
without pit2 (n = 26 clusters) and 10,100 ± 1,300 with pit2 (n = 26
clusters, p < 0.01, Mann–Whitney U test). The Syp-SEP signal intensity
at pH 7.4 did not change significantly by pit2 application (Figure 5B,
p = 0.67, Mann–Whitney U test). The pit2-sensitive component of the
Syp-SEP signal at pH 6.0 presumably related to CME, whereas
endocytosis that was not inhibited by pit2 was clathrin-independent. Frontiers in Cellular Neuroscience frontiersin.org Clathrin-dependent and -independent
endocytosis It has been proposed that ADBE is triggered by a high
load of membrane addition into the presynaptic plasma membrane
through the exocytosis of a large number of synaptic vesicles and that
the elevated concentration of cytoplasmic calcium in presynaptic
terminals plays a role in this process. A field stimulation of 50 pulses at
50 Hz released most vesicles in a readily releasable pool. Therefore, the
clathrin-independent endocytosis we recorded here is likely to
correspond to ADBE. To address whether the ADBE-dependent and CME-dependent
intracellular Syp-SEP spatial distribution patterns were different,
we compared the Syp-SEP signal distribution characteristics before and
after pit2 application. We found that the maximum Syp-SEP signal
intensity and that divided by the Syp-SEP signal-positive area at 7.24 s
after the stimulation were significantly larger than those before pit2
application (Figure 6E, n = 15 clusters before application and n = 16
clusters after, signal intensity, p < 0.05 Mann–Whitney U test; signal
intensity / area, p < 0.01 Student’s paired t test). The Syp-SEP signal-
positive areas were not significantly different with or without pit2. These results suggest that the intracellular Syp-SEP cluster formed
through CME might be more concentrated than the cluster formed
through ADBE. However, it might be also possible that some
CME-mediated and ADBE-mediated Syp-SEP signals were not
spatially separated and that in such cases the recorded maximum
Syp-SEP signal intensity without pit2 was the sum of the CME- and
ADBE-mediated signals. Next, we took higher magnification images of Syp-SEP
(Figures 6A,B). The centroids of the Syp-SEP signal at pH 6.0
detected in the presence of pit2, which presumably reflected ADBE,
were located 139 ± 49 nm (n = 33 clusters) outside of the edge of the
CAST-RFP signal at 3.24 s and 97 ± 44 nm (n = 31 clusters) at 7.24 s,
but at pH 7.4 and 1 s after the stimulation, the centroid was
115 ± 16 nm (n = 24 clusters) inside of the edge (Figure 6C, p < 0.001
for both, Steel-Dwass’s test). Thus, ADBE are also likely to occur at
the periphery of the CAST-RFP signal.i To examine whether the region where ADBE occur is fixed, the
stimulation was applied repeatedly (Figure 6B). Clathrin-dependent and -independent
endocytosis The distance between the
centroid of the Syp-SEP signal in the first trial and the nearest centroid
in the second trial was 378 ± 94 nm (n = 11 clusters) at 3.24 s and
351 ± 65 nm (n = 12 clusters) at 7.24 s (Figure 6D), which are significantly
smaller than the distances recorded at 3.24 s and 7.24 s in a single trial
(158 ± 23 nm, n = 25 clusters, p < 0.01 and 0.05, Steel-Dwass’s test). These Frontiers in Cellular Neuroscience Clathrin-dependent and -independent
endocytosis Next, we tried to determine whether the endocytosis we studied
was clathrin-dependent or -independent using the clathrin inhibitor 08 frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 5
Clathrin-dependent and -independent endocytosis. (A) Representative images of Syp-SEP and CAST-RFP signals at pH 7.4 or 6.0 before and after the
50 pulses stimulation in the presence of pit2. While endocytosis was inhibited in some AZLM (arrowhead, pH 6.0, 3.12 s) after the exocytosis of Syp-SEP
(pH 7.4, 1.0 s), endocytosed Syp-SEP was detected in other AZLM (arrow, pH 6.0, 3.12 s). (B) Quantification of Syp-SEP at pH 7.4 at 1 s after the
stimulation (blue) and at pH 6.0 at 3.24 or 7.24 s after the stimulation (red) without or with pit2 (−pit2, n = 26 clusters; +pit2, 25 clusters, 7 cells). * and
** indicates p < 0.05 and 0.01 respectively, n.s., not significant. Clathrin-dependent and -independent endocytosis. (A) Representative images of Syp-SEP and CAST-RFP signals at pH 7.4 or 6.0 before and after the
50 pulses stimulation in the presence of pit2. While endocytosis was inhibited in some AZLM (arrowhead, pH 6.0, 3.12 s) after the exocytosis of Syp-SEP
(pH 7.4, 1.0 s), endocytosed Syp-SEP was detected in other AZLM (arrow, pH 6.0, 3.12 s). (B) Quantification of Syp-SEP at pH 7.4 at 1 s after the
stimulation (blue) and at pH 6.0 at 3.24 or 7.24 s after the stimulation (red) without or with pit2 (−pit2, n = 26 clusters; +pit2, 25 clusters, 7 cells). * and
** indicates p < 0.05 and 0.01 respectively, n.s., not significant. Previous studies reported that a high-frequency stimulation triggers
ADBE (Cheung and Cousin, 2013; Kokotos et al., 2018; Renard and
Boucrot, 2021). It has been proposed that ADBE is triggered by a high
load of membrane addition into the presynaptic plasma membrane
through the exocytosis of a large number of synaptic vesicles and that
the elevated concentration of cytoplasmic calcium in presynaptic
terminals plays a role in this process. A field stimulation of 50 pulses at
50 Hz released most vesicles in a readily releasable pool. Therefore, the
clathrin-independent endocytosis we recorded here is likely to
correspond to ADBE. results suggest that there are multiple ADBE zones around an active zone
and which zone is used for ADBE varies trial to trial. Previous studies reported that a high-frequency stimulation triggers
ADBE (Cheung and Cousin, 2013; Kokotos et al., 2018; Renard and
Boucrot, 2021). Temperature-sensitive endocytosis occurs
immediately after the stimulation Next, we examined whether endocytosis could be detected
immediately after the stimulation (Figure 7). UFE, which is faster 09 frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 6
Pit2-resistant endocytosis. (A) Representative high magnification images of Syp-SEP signals at pH 7.4 and 6.0 before and after the 50 pulses stimulation
without pit2 (upper images) and after application of pit2 (lower images) in the same sample. The right images are averages of the images enclosed by
the yellow lines and were merged with an image of CAST-RFP signals. The Syp-SEP signal indicated by an arrow in the top image was clearly
diminished after addition of pit2 (bottom image). (B) Representative images of Syp-SEP signals at pH 7.4 and 6.0 before and after the stimulation in the
presence of pit2 in different trials using the same sample. The right images are averages of the images enclosed by the yellow lines and were merged
with an image of CAST-RFP signals. Arrows indicated Syp-SEP in different locations in different trials. (C) Location of centroids of Syp-SEP signals at pH
7.4 at 0.12 s after the stimulation and at pH 6.0 at 3.24 s or 7.24 s after the stimulation in relation to the edge of the CAST-RFP signal (0.12 s, n = 24
clusters, 3.24 s, 33 clusters, 7.24 s, 31 clusters, 8 cells). (D) Displacement of centroids of Syp-SEP signal at pH 6.0 in different trials (3.24 s, 7.24 s) or over
a time range (3.24–7.24 s) in a trial (3.24 s, n = 11 clusters, 7.24 s, 12 clusters, 3.24–7.24 s, 25 clusters, 8 cells). (E) Quantification of the maximum
intensity, the area and the maximum intensity divided by the area of Syp-SEP signals at pH 6.0 at 7.24 s after the stimulation (Before n = 15 clusters;
After, 16 clusters, 6 cells). *, **, and *** indicate p < 0.05, 0.01 and 0.001, respectively. n.s., not significant. FIGURE 6
Pit2-resistant endocytosis. (A) Representative high magnification images of Syp-SEP signals at pH 7.4 and 6.0 before and after the 50 pulses stimulation
without pit2 (upper images) and after application of pit2 (lower images) in the same sample. The right images are averages of the images enclosed by
the yellow lines and were merged with an image of CAST-RFP signals. The Syp-SEP signal indicated by an arrow in the top image was clearly
diminished after addition of pit2 (bottom image). Temperature-sensitive endocytosis occurs
immediately after the stimulation (B) Quantification of Syp-SEP at pH 6.0 without
stimulation (n = 29 clusters, 10 cells), and at 0.36 s (25 clusters, 7 cells),
0.84 s (13 clusters, 6 cells) and 3.24 s (55 clusters, 20 cells) after the 5
pulses stimulation. The intensity value (F – Fbefore) was calculated by
subtracting the value 3.88 s before the stimulation (in no stimulation,
the value 4.24 s before the timing of stimulation onset was subtracted). Thus, when the intensity value before the stimulation was comparably
large due to the signal noise and that after was comparably small, the
subtracted and presented value took small negative value. * and ***
indicates p < 0.05 and 0.001 respectively; n.s., not significant. FIGURE 7
Absence of ultrafast endocytosis at RT. (A) Representative images of
Syp-SEP at pH 7.4 at 4 s before and at 0.12 s (top) or 0.60 s (bottom)
after the 5 pulses stimulation, and at pH 6.0 at 3.88 s before and at
0.36 s (top) or 0.84 s (bottom) after the stimulation. All Syp-SEP signals
were merged with CAST-RFP signals. Endocytosed Syp-SEP signals
were detected not at 0.36 s (arrowheads) but at 0.84 s (arrows) after
the stimulation. (B) Quantification of Syp-SEP at pH 6.0 without
stimulation (n = 29 clusters, 10 cells), and at 0.36 s (25 clusters, 7 cells),
0.84 s (13 clusters, 6 cells) and 3.24 s (55 clusters, 20 cells) after the 5
pulses stimulation. The intensity value (F – Fbefore) was calculated by
subtracting the value 3.88 s before the stimulation (in no stimulation,
the value 4.24 s before the timing of stimulation onset was subtracted). Thus, when the intensity value before the stimulation was comparably
large due to the signal noise and that after was comparably small, the
subtracted and presented value took small negative value. * and ***
indicates p < 0.05 and 0.001 respectively; n.s., not significant. Lastly, the intracellular Syp-SEP signal spatial distribution after
UFE was analyzed. The maximum Syp-SEP signal intensity divided
by the Syp-SEP signal-positive area at 0.36 s was 8.3 ± 0.6 and at
3.24 s it was 9.6 ± 0.9 (Figure 8G, p = 0.30, Mann–Whitney U test). Temperature-sensitive endocytosis occurs
immediately after the stimulation Thus, when the intensity value before the stimulation was comparably
large due to the signal noise and that after was comparably small, the than other types of endocytosis, has been reported to occur
around active zones at 34°C (Watanabe et al., 2013). After a single
stimulus, the synaptic vesicle membrane is recovered at sites in the
vicinity of the active zone within 100 ms independently of clathrin. It is not known whether UFE mediates the recovery of synaptic
vesicle proteins, which should be an important function of
endocytosis at presynaptic terminals (Watanabe and Boucrot,
2017). We attempted to determine whether UFE retrieves laterally
diffused Syp-SEP after exocytosis (Gimber et al., 2015; Funahashi
et al., 2018) in a living neuron. At RT, we detected the Syp-SEP
signal increases at pH 6.0 at 0.84 s after the stimulus but not at
0.36 s after (Figure 7B, p < 0.001 and 0.05, Steel-Dwass’s test). Thus, very fast endocytosis was not detected at RT, suggesting that
UFE was unlikely to occur at RT, which is consistent with previous
reports (Watanabe et al., 2013). than other types of endocytosis, has been reported to occur
around active zones at 34°C (Watanabe et al., 2013). After a single
stimulus, the synaptic vesicle membrane is recovered at sites in the
vicinity of the active zone within 100 ms independently of clathrin. It is not known whether UFE mediates the recovery of synaptic
vesicle proteins, which should be an important function of
endocytosis at presynaptic terminals (Watanabe and Boucrot,
2017). We attempted to determine whether UFE retrieves laterally
diffused Syp-SEP after exocytosis (Gimber et al., 2015; Funahashi
et al., 2018) in a living neuron. At RT, we detected the Syp-SEP
signal increases at pH 6.0 at 0.84 s after the stimulus but not at
0.36 s after (Figure 7B, p < 0.001 and 0.05, Steel-Dwass’s test). Thus, very fast endocytosis was not detected at RT, suggesting that
UFE was unlikely to occur at RT, which is consistent with previous
reports (Watanabe et al., 2013). To make UFE more likely to occur, the temperature was
raised to near physiological temperature (PT, 31–32°C). We also
reduced the number of stimulation pulses to 5. At PT, the
Syp-SEP signal was recorded when pH was changed at 120 ms
after the onset of the stimulation (Figure 8A). Temperature-sensitive endocytosis occurs
immediately after the stimulation (B) Representative images of Syp-SEP signals at pH 7.4 and 6.0 before and after the stimulation in the
presence of pit2 in different trials using the same sample. The right images are averages of the images enclosed by the yellow lines and were merged
with an image of CAST-RFP signals. Arrows indicated Syp-SEP in different locations in different trials. (C) Location of centroids of Syp-SEP signals at pH
7.4 at 0.12 s after the stimulation and at pH 6.0 at 3.24 s or 7.24 s after the stimulation in relation to the edge of the CAST-RFP signal (0.12 s, n = 24
clusters, 3.24 s, 33 clusters, 7.24 s, 31 clusters, 8 cells). (D) Displacement of centroids of Syp-SEP signal at pH 6.0 in different trials (3.24 s, 7.24 s) or over
a time range (3.24–7.24 s) in a trial (3.24 s, n = 11 clusters, 7.24 s, 12 clusters, 3.24–7.24 s, 25 clusters, 8 cells). (E) Quantification of the maximum
intensity, the area and the maximum intensity divided by the area of Syp-SEP signals at pH 6.0 at 7.24 s after the stimulation (Before n = 15 clusters;
After, 16 clusters, 6 cells). *, **, and *** indicate p < 0.05, 0.01 and 0.001, respectively. n.s., not significant. 10 Frontiers in Cellular Neuroscience frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 FIGURE 7
Absence of ultrafast endocytosis at RT. (A) Representative images of
Syp-SEP at pH 7.4 at 4 s before and at 0.12 s (top) or 0.60 s (bottom)
after the 5 pulses stimulation, and at pH 6.0 at 3.88 s before and at
0.36 s (top) or 0.84 s (bottom) after the stimulation. All Syp-SEP signals
were merged with CAST-RFP signals. Endocytosed Syp-SEP signals
were detected not at 0.36 s (arrowheads) but at 0.84 s (arrows) after
the stimulation. (B) Quantification of Syp-SEP at pH 6.0 without
stimulation (n = 29 clusters, 10 cells), and at 0.36 s (25 clusters, 7 cells),
0.84 s (13 clusters, 6 cells) and 3.24 s (55 clusters, 20 cells) after the 5
pulses stimulation. The intensity value (F – Fbefore) was calculated by
subtracting the value 3.88 s before the stimulation (in no stimulation,
the value 4.24 s before the timing of stimulation onset was subtracted). frontiersin.org Temperature-sensitive endocytosis occurs
immediately after the stimulation Line scans across
CAST-RFP and Syp-SEP images at high magnification show
the position of the endocytosed Syp-SEP signal peak at 0.36 s
after the stimulation was about 500 nm from the position of the
CAST-RFP signal peak (Figure 8B). The Syp-SEP signal centroid
at pH 6.0 was 20 ± 21 nm (n = 17 clusters) outside the edge of
the CAST-RFP signal at 0.36 s, but 110 ± 25 nm (n = 17 clusters)
inside the CAST-RFP signal edge at pH 7.4 at 0.12 s after the
stimulation (Figure 8C, p < 0.001, Wilcoxon signed-rank test). Importantly, the Syp-SEP signal intensity at pH 6.0 was larger
at 0.36 s after the stimulation than that at 3.24 s (Figure 8D,
p < 0.05, 0.01, and 0.001, Steel-Dwass’s test). These results
suggest that temperature-dependent UFE occurs immediately
after the stimulation, and it dominates when the number of
stimulation pulses is small at PT. We did not observe a clear
Syp-SEP signal at pH 6.0 inside of AZLM at 0.36 s after the onset
of stimulation. Next, we addressed whether the region where UFE occurred
was fixed by applying the stimulation repeatedly (Figure 8E). The
distance between the centroid of the Syp-SEP signal in the first trial
and the nearest centroid in the second trial at 0.36 s was 369 ± 46 nm
(n = 11 clusters), which was significantly larger than the distance
between the centroids of the Syp-SEP signal at 0.36 s and 3.24 s in
a single trial (Figure 8F, 157 ± 27 nm, n = 14 clusters, p < 0.01, Steel-
Dwass’s test). These results suggest that there are multiple UFE
zones around an active zone and that the Syp-SEP signal recorded
at 3.24 s included Syp-SEP endocytosed before 0.36 s. FIGURE 7
Absence of ultrafast endocytosis at RT. (A) Representative images of
Syp-SEP at pH 7.4 at 4 s before and at 0.12 s (top) or 0.60 s (bottom)
after the 5 pulses stimulation, and at pH 6.0 at 3.88 s before and at
0.36 s (top) or 0.84 s (bottom) after the stimulation. All Syp-SEP signals
were merged with CAST-RFP signals. Endocytosed Syp-SEP signals
were detected not at 0.36 s (arrowheads) but at 0.84 s (arrows) after
the stimulation. Temperature-sensitive endocytosis occurs
immediately after the stimulation Considering that Syp-SEP is about 32% brighter at RT than at PT,
the UFE-dependent maximum Syp-SEP signal intensity divided by
the signal-positive area at RT at 0.36 s was 10.9 ± 0.8, which was not
significantly different from the value at 7.24 s after 50 pulses
stimulation at RT with pit2 (11.5 ± 1.6, p = 0.98, Steel-Dwass’s test),
but smaller than that without pit2 (21.6 ± 3.6, p < 0.01, Steel-
Dwass’s test). We could not separate the Syp-SEP signal-positive
area at 0.36 s using watershed lines. 11 Frontiers in Cellular Neuroscience frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 GURE 8
ltrafast endocytosis at PT. (A) Representative high magnification images of Syp-SEP at pH 7.4 and 6.0 before and immediately after the 5 pulses
timulation at PT. The right images are averages of the images enclosed by the yellow lines and were merged with an image of CAST-RFP signals. B) Line-scan of Syp-SEP and CAST-RFP signal intensities on the yellow broken line shown in (A). (C) Location of centroids of Syp-SEP signals at pH 7.4
t 0.12 s after the stimulation and at pH 6.0 immediately after the stimulation (the centroid calculated from an averaged image between 0.36 s and
.72 s) in relation to the edge of CAST-RFP signal (0.12 s, n = 17 clusters, 0.36 s, 17 clusters, 7 cells). The data obtained in a trial were connected by a line. D) Quantification of Syp-SEP signal intensities at pH 6.0 without stimulation and at 0.36 s or 3.24 s after the stimulation (No stim., n = 7 clusters; 0.36 s,
6 clusters; 3.24 s, 15 clusters, 8 cells). The intensity value (F – Fbefore) was calculated by subtracting the value 3.76 s before the stimulation (in no
timulation, the value 3.76 s before the timing of stimulation onset was subtracted). Thus, when the intensity value before the stimulation was
omparably large due to the signal noise and that after was comparably small, the subtracted and presented value took small negative value. E) Representative images of Syp-SEP at pH 7.4 and 6.0 before and immediately after the stimulation in different trials using the same sample. The right
mages are averages of the images enclosed by the yellow lines and were merged with an image of CAST-RFP. Arrows indicate Syp-SEP signal. The
gnal locations were different between 1st and 2nd trials. Temperature-sensitive endocytosis occurs
immediately after the stimulation (F) Displacement of centroids of Syp-SEP signals at pH 6.0 in different trials (0.36 s, 3.24 s) or
ver a time range (0.36–3.24 s) in a trial (0.36 s, n = 11 clusters, 3.24 s, 11 clusters, 0.36–3.24 s, 14 clusters, 4 cells). (G) Quantification of the maximum
ntensity, the area and the maximum intensity divided by the area of Syp-SEP signals at pH 6.0 at 0.36 s and 3.24 s after the stimulation (0.36 s, n = 37
lusters, 3.24 s, 29 clusters, 9 cells). *, **, and *** indicate p < 0.05, 0.01 and 0.001, respectively. n.s., not significant. FIGURE 8
Ultrafast endocytosis at PT. (A) Representative high magnification images of Syp-SEP at pH 7.4 and 6.0 before and immediately after the 5 pulses
stimulation at PT. The right images are averages of the images enclosed by the yellow lines and were merged with an image of CAST-RFP signals. (B) Line-scan of Syp-SEP and CAST-RFP signal intensities on the yellow broken line shown in (A). (C) Location of centroids of Syp-SEP signals at pH 7.4
at 0.12 s after the stimulation and at pH 6.0 immediately after the stimulation (the centroid calculated from an averaged image between 0.36 s and
0.72 s) in relation to the edge of CAST-RFP signal (0.12 s, n = 17 clusters, 0.36 s, 17 clusters, 7 cells). The data obtained in a trial were connected by a line. (D) Quantification of Syp-SEP signal intensities at pH 6.0 without stimulation and at 0.36 s or 3.24 s after the stimulation (No stim., n = 7 clusters; 0.36 s,
16 clusters; 3.24 s, 15 clusters, 8 cells). The intensity value (F – Fbefore) was calculated by subtracting the value 3.76 s before the stimulation (in no
stimulation, the value 3.76 s before the timing of stimulation onset was subtracted). Thus, when the intensity value before the stimulation was
comparably large due to the signal noise and that after was comparably small, the subtracted and presented value took small negative value. (E) Representative images of Syp-SEP at pH 7.4 and 6.0 before and immediately after the stimulation in different trials using the same sample. The right
images are averages of the images enclosed by the yellow lines and were merged with an image of CAST-RFP. Arrows indicate Syp-SEP signal. The
signal locations were different between 1st and 2nd trials. CME and ADBE Various types of fluorescence imaging techniques have been used to
study exocytotic and/or endocytic processes in presynaptic terminals
(Welzel et al., 2013; Egashira et al., 2015; Midorikawa and Sakaba, 2015;
Okamoto et al., 2016; Guillaud et al., 2017; Soykan et al., 2017; Mori et al.,
2021). Our method is an advancement of these studies in the following
two aspects. First, the signal-to-noise ratio and z-axis spatial resolution
were improved by applying TIRFM to AZLM. As described, using
TIRFM, we could record signals from single SEP molecules (Figure 1E). In addition, a near two-dimensional analysis of the distribution of
endocytosed synaptic proteins was performed by forming AZLM
parallelly on the glass surface. Second, extracellular pH exchange using
U-tube enabled fast extracellular pH switching and the recording of
endocytosed synaptic vesicle proteins at different times repetitively. Previous studies have demonstrated the presence of CME and
clathrin-independent endocytosis at mammalian central synapses. CME was originally thought to be the predominant endocytic
mechanism for recycling synaptic vesicles because of the sensitivity
to perturbation of clathrin or clathrin-associated proteins, and its
time constant was reported to be 15–30 s at RT (Granseth et al.,
2006). Here, we showed that the exocytosed Syp-SEP signal after 50
pulses at 50 Hz decreased with a similar time constant of about 20 s
(Figure 2A). The speed of CME is thought to be limited by the
selection and gathering processes of clathrin and its associated
proteins including cargo receptors (Ehrlich et al., 2004; Taylor et al.,
2011; Watanabe and Boucrot, 2017). Therefore, not all endocytosis
detected at 3.24 and 7.24 s after the 50 pulses stimulation was
unlikely to be CME (Figures 3A, 4A). We used a 150× optical lens with a high numerical aperture (1.45)
to obtain a high XY spatial resolution; nevertheless, the resolution was
restricted by the diffraction limit. Super resolution techniques, such
as stochastic optical reconstruction microscopy (STORM),
photoactivated localization microscopy (PALM), stimulated emission
depletion microscopy (STED) and/or structured illumination
microscopy (SIM), have been used to study exo- and endocytic
processes in presynaptic terminals and endocrine cells at higher
spatial resolution (Schermelleh et al., 2010; Maglione and Sigrist, 2013;
Sigal et al., 2018; Schermelleh et al., 2019). Each of these methods has
merits and demerits. Future studies combining our method and super-
resolution imaging should provide higher quality data and more
detailed information about the recycling processes of synaptic vesicles. CME and ADBE When a strong stimulation such as 50 Hz or high K+ solution is
applied to neurons, a large area of the plasma membrane is
internalized through the clathrin-independent mechanism ADBE
(Clayton and Cousin, 2009). Synaptic vesicles are then regenerated
from endosomes through a clathrin-dependent process (Kononenko
et al., 2014; Watanabe et al., 2014). Recent studies have suggested
that ADBE is the dominant endocytosis mechanism during and
after high frequency presynaptic activation at PT. The major role of
ADBE is not the immediate restoration of synaptic vesicles but it
may involve the clearance of fusion sites for later exocytosis
(Watanabe et al., 2014; Soykan et al., 2017; Chanaday et al., 2019;
Ivanova and Cousin, 2022). On the other hand, CME might
contribute to the reformation of synaptic vesicles more directly by
gathering membrane proteins of synaptic vesicles through clathrin-
associated adaptor proteins over a slower time course. Synaptophysin is diffused out of AZLM following exocytosis Discussion In order not to include such
signals in the analysis, we excluded signals which appeared after the
pH change to 6.0. It has been reported that the acidification of
endocytosed vesicles takes 3–15 s (Atluri and Ryan, 2006; Granseth
et al., 2006; Egashira et al., 2015). It has also been shown that
synaptic vesicles can enter and exit the TIRFM visualization zone
in about 100 ms (Midorikawa and Sakaba, 2015). Our observations
indicated that the Syp-SEP signal recorded at pH 6.0 at 7.24 s
includes partial signals from Syp-SEP endocytosed before 3.24 s. Methods to study endocytosis in
presynaptic terminals We have established a new experimental method for the live-cell
imaging of endocytosed synaptic vesicle membrane proteins at multiple
time points after the exocytotic fusion of vesicles triggered by electrical
stimulation. By combining the rapid extracellular pH exchange method
and TIRFM observation around AZLM formed on a glass surface,
we recorded clathrin-dependent and -independent endocytosis-related
signals at RT and also temperature-sensitive ultrafast endocytosis-
related signals. This method enabled us to directly observe the spatial
pattern of the endocytosed Syp-SEP signal around AZLM repeatedly,
which is in principle inaccessible with EM and electrophysiological
capacitance measurements. Morphological observations by EM need
fixation of the preparation and thus only provide data at one time point
for a preparation (Watanabe et al., 2013). Electrophysiological
recordings provide continuous data from a live preparation with a high
temporal resolution, but they cannot provide information about the
localization of synaptic vesicle proteins or the morphological properties
of endocytic events (Delvendahl et al., 2016). frontiersin.org Discussion within about 200 nm from the glass surface, which corresponds to
the TIRFM visualization zone. Considering the thickness of the
NLG coating and the extracellular space and thickness of plasma
membrane, we estimate that the presynaptic intracellular space was
about >50 nm above the glass surface. Most of the Syp-SEP signal
observed at pH 6.0 likely came from one or multiple vesicles
endocytosed just before the pH exchange, before intraluminal
acidification and before the vesicles moved out of the TIRFM
visualization zone. The Syp-SEP signal could have also come from
endosomes having near neutral intraluminal pH and/or
endoplasmic reticulum. Therefore, we subtracted the Syp-SEP
image recorded before the stimulation from that recorded after the
stimulation to exclude intracellular Syp-SEP signals insensitive to
the stimulation. The Syp-SEP signal may also have come from
vesicles endocytosed sometime before but moved into the
visualization zone from the outside. In order not to include such
signals in the analysis, we excluded signals which appeared after the
pH change to 6.0. It has been reported that the acidification of
endocytosed vesicles takes 3–15 s (Atluri and Ryan, 2006; Granseth
et al., 2006; Egashira et al., 2015). It has also been shown that
synaptic vesicles can enter and exit the TIRFM visualization zone
in about 100 ms (Midorikawa and Sakaba, 2015). Our observations
indicated that the Syp-SEP signal recorded at pH 6.0 at 7.24 s
includes partial signals from Syp-SEP endocytosed before 3.24 s. within about 200 nm from the glass surface, which corresponds to
the TIRFM visualization zone. Considering the thickness of the
NLG coating and the extracellular space and thickness of plasma
membrane, we estimate that the presynaptic intracellular space was
about >50 nm above the glass surface. Most of the Syp-SEP signal
observed at pH 6.0 likely came from one or multiple vesicles
endocytosed just before the pH exchange, before intraluminal
acidification and before the vesicles moved out of the TIRFM
visualization zone. The Syp-SEP signal could have also come from
endosomes having near neutral intraluminal pH and/or
endoplasmic reticulum. Therefore, we subtracted the Syp-SEP
image recorded before the stimulation from that recorded after the
stimulation to exclude intracellular Syp-SEP signals insensitive to
the stimulation. The Syp-SEP signal may also have come from
vesicles endocytosed sometime before but moved into the
visualization zone from the outside. Frontiers in Cellular Neuroscience Temperature-sensitive endocytosis occurs
immediately after the stimulation (F) Displacement of centroids of Syp-SEP signals at pH 6.0 in different trials (0.36 s, 3.24 s) or
over a time range (0.36–3.24 s) in a trial (0.36 s, n = 11 clusters, 3.24 s, 11 clusters, 0.36–3.24 s, 14 clusters, 4 cells). (G) Quantification of the maximum
intensity, the area and the maximum intensity divided by the area of Syp-SEP signals at pH 6.0 at 0.36 s and 3.24 s after the stimulation (0.36 s, n = 37
clusters, 3.24 s, 29 clusters, 9 cells). *, **, and *** indicate p < 0.05, 0.01 and 0.001, respectively. n.s., not significant. 12 Frontiers in Cellular Neuroscience frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 UFE An innovative flash-and-freeze approach using optogenetics
and freezing neurons at defined time points after the stimulation
revealed the existence of UFE. UFE occurs at sites lateral to fusion
sites (typically <200 nm) within 100 ms after a stimulation, does
not require clathrin, and is temperature dependent (Watanabe
et al., 2013, 2014). The amount of membrane internalized by UFE
equals the membrane exocytosed (Watanabe et al., 2013). It was
suggested that any excess membrane must be removed rapidly
from the plasma membrane to restore fusion sites and keep the
membrane area and tension constant during a high load of
membrane turnover (Rosenmund et al., 1993), something UFE
seems to manage (Watanabe and Boucrot, 2017). However, it was
unknown whether UFE contributes to the recovery of vesicle
proteins. We found that endocytic vesicles contained Syp-SEP at
0.36 s after the onset of a stimulation at PT (Figure 8A). The
maximum Syp-SEP signal intensity divided by the signal-positive
area of UFE was comparable to that of ADBE after compensating
for temperature effects on SEP fluorescence, suggesting that
Syp-SEP densities were similar in UFE-mediated and ADBE-
mediated intracellular vesicles (Figures 6E, 8G). Caveat, remaining questions and future
advancement One caveat of this study is that AZLM is an artificial structure and
may have characteristics that are different from normal presynaptic
active zones. However, our previous study demonstrated the
accumulation of various active zone proteins in AZLM and that the
electrical stimulation triggered fusion of synaptic vesicles in AZLM,
suggesting that AZLM shows essential properties of active zones and
is a reliable model (Funahashi et al., 2018). Future study comparing
the results obtained in AZLM with those in normal synapses in detail
might provide useful information. It might be also possible to induce
AZLM with somewhat different characteristics such as that in large or
inhibitory presynaptic terminals using other synaptic adhesion
molecules (Südhof, 2021). Comparison of properties among different
types of AZLM could be interesting. We would like to note that Syp-SEP signal recorded at 0.36 s
might have come from single vesicle in some cases because of the
following reason. UFE forms a vesicle of about 100 nm diameter
(Watanabe et al., 2013) and the diameter of synaptic vesicle is
about 50 nm. Thus, UFE internalizes membrane area
corresponding to 4 synaptic vesicles. On the other hand, according
to our previous study on synaptic vesicle exocytosis in AZLM, 5
pulses stimulation at 50 Hz seems to induce fusion of around 6
synaptic vesicles (Funahashi et al., 2018). Another limitation of the present method is XY spatial resolution
is restricted by the diffraction limit and is inferior to super resolution
techniques such as STED, STORM, PALM and SIM (Schermelleh
et al., 2010, 2019; Maglione and Sigrist, 2013; Sigal et al., 2018). Future
studies combining the present method and one or some of super-
resolution imaging techniques would provide more detailed
information about the recycling processes of synaptic vesicle proteins. Here, we have examined only synaptophysin as a synaptic vesicle
protein. Other synaptic vesicle proteins such as synaptotagmin,
synaptobrevin or vglut1 might show different intracellular distribution
after endocytosis, because each might be trapped on the membrane of
endocytosed vesicles through different adaptor proteins (Cousin,
2017). Thus, extending the analyses to other types of synaptic vesicle
protein would contribute to better understanding of retrieval
processes of synaptic vesicles. Identity of Syp-SEP signal recorded at pH 6.0 The Syp-SEP signal we recorded at pH 6.0 should have come
from intracellular vesicles with an intraluminal neutral pH located Frontiers in Cellular Neuroscience 13 Tanaka et al. 10.3389/fncel.2023.1277729 (Gimber et al., 2015; Funahashi et al., 2018). We showed that the
maximum Syp-SEP signal intensity at pH 6.0 divided by the
Syp-SEP-positive area was significantly larger before pit2 application
than after, which suggests that CME-dependent intracellular
vesicles concentrate Syp more than ADBE-dependent ones. This
different effect by the two types of vesicles might be caused by
trapping Syp through adaptor proteins in CME but not in
ADBE. Thus, the density of Syp-SEP in CME-derived vesicles might
be higher than that in ADBE-derived vesicles. In addition, a part of
ADBE-derived large vesicle could be far from the bottom glass
surface and out of TIRFM visualization zone, which might have
made the Syp-SEP signal intensity weaker. It is to be noted that
there is a possibility that in some cases CME-mediated and ADBE-
mediated Syp-SEP signals were closely localized, such that both
contributed to the signal intensity of a particular pixel. intense stimulation by high K+ solution was reported using quantum
dots (Zhang et al., 2009). However, the large size of quantum dots
might have affected fusion pore openings and vesicle collapse
(Dittman and Ryan, 2009). K&R should take place at exocytosis sites
within an active zone immediately after the exocytosis. However,
we did not record events that unequivocally reflected K&R in AZLM,
suggesting K&R was rare in our preparations. Frontiers in Cellular Neuroscience Endocytosis sites One merit of our experimental system is that AZLM can
be repeatedly stimulated and endocytosed SEP-tagged synaptic
proteins can be imaged after each stimulation. The results show that
under different conditions (5 or 50 pulses at RT or PT), the majority
of endocytosed Syp-SEP signals was located <200 nm away from the
edge of AZLM, but the positions varied trial to trial. Proteins involved
in endocytosis might not be immobilized. However, the existence of
hotspots, where a particular type of endocytosis such as UFE, ADBE
and/or CME preferentially occurs, cannot be excluded, since the
number of trials in our experimental protocol was limited due to the
bleaching of Syp-SEP and CAST-RFP. As for the exocytosis of synaptic
vesicles, the existence of several hot spots within an active zone has
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10.1111/j.1471-4159.2009.06384.x Granseth, B., Odermatt, B., Royle, S. J. Data availability statement supported by grants 18H02526 to TH and 21 K06393 to HT from the
Japan Society for the Promotion of Science, and grants to HT from the
Takeda Science Foundation, SPIRITS 2020 of Kyoto University, and
FY 2021 Kusunoki 125 of Kyoto University 125th Anniversary Fund
in Japan. supported by grants 18H02526 to TH and 21 K06393 to HT from the
Japan Society for the Promotion of Science, and grants to HT from the
Takeda Science Foundation, SPIRITS 2020 of Kyoto University, and
FY 2021 Kusunoki 125 of Kyoto University 125th Anniversary Fund
in Japan. The original contributions presented in the study are included in
the article/supplementary material, further inquiries can be directed
to the corresponding author. Author contributions The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could
be construed as a potential conflict of interest. HT: Conceptualization, Formal analysis, Funding acquisition,
Investigation, Methodology, Software, Validation, Writing – original
draft, Writing – review & editing. JF: Methodology, Writing – review
& editing. TH: Conceptualization, Funding acquisition, Supervision,
Writing – review & editing. K & R An alternative fast endocytosis pathway, K&R, was reported at
neuronal synapses and in neuroendocrine cells (Gandhi and Stevens,
2003; Zhang et al., 2009), although whether K&R occurs in the
mammalian central nervous system is controversial (He and Wu,
2007). The transient opening and closing of fusion pores during 14 frontiersin.org Tanaka et al. 10.3389/fncel.2023.1277729 10.3389/fncel.2023.1277729 Acknowledgments We thank S. Kawaguchi and P. Karagiannis for comments on
the manuscript. The animal study was approved by Local committee for handling
experimental animals in the Graduate School of Science, Kyoto
University. The study was conducted in accordance with the local
legislation and institutional requirements. Funding The author(s) declare financial support was received for the
research, authorship, and/or publication of this article. This work was Publisher’s note All claims expressed in this article are solely those of the authors
and do not necessarily represent those of their affiliated
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reviewers. Any product that may be evaluated in this article, or
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Tradução e adaptação brasileira do Questionnaire D'Alimentation
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Revista CEFAC
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cc-by
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(1)
Universidade Estadual de Campinas, Piracicaba, São
Paulo, Brasil
(2)
McGill University, Montreal, Quebec, Canada
(3)
Universidade Cidade de São Paulo, São Paulo, São Paulo,
Brasil
(4)
Universidade Federal de São Paulo, Diadema, São Paulo,
Brasil
Conflict of interest: non-existent ABSTRACT Purpose: to translate the French language instrument “Questionnaire D’Alimentation” to Brazilian
Portuguese and culturally adapt it for adolescents. Methods: the translation was based on a protocol
that consisted of translation to Brazilian Portuguese (by a professor with a Literary Arts degree and
a Doctor of Dental Surgery, both fluent in French and Brazilian Portuguese); back-translation into
French; revision by a Committee of specialists (two translators, one native and one sworn translator,
and two university professors, one being a Doctor of Dental Surgery and a Speech Therapist) and
cultural equivalence (pre-test). The version used in the pre-test consisted of 26 questions with five
possible responses (5-Likert), distributed in five domains (Food-Mastication, Habits, Meats, Fruits and
Vegetables). The pre-test and test-retest was performed with a sample of 20 adolescents (10 boys/10
girls) from public schools of Piracicaba (Brazil). At this stage, the alternative “I did not understand” was
added to each question in order to identify those that were not understood. Test-retest reliability was
assessed for each domain using intra-class correlation coefficients (ICCs). Results: in the pre-test,
an excellent comprehension of the instrument was observed; in test-retest, ICCs ranged from 0.45 to
0.81 (moderate to excellent agreement). Conclusion: the Portuguese version of the Questionnaire
D’Alimentation has shown to be easy to understand by Brazilian adolescents and useful in the
evaluation of the masticatory function and feeding or swallowing disorders that may affect food intake. KEYWORDS: Translating; Mastication; Food; Adolescent KEYWORDS: Translating; Mastication; Food; Adolescent 1929 1929 KEYWORDS: Translating; Mastication; Food; Adolescent
INTRODUCTION degree of malocclusion, as well as the masticatory
muscles functioning, may influence the quality of an
individual’s masticatory function3,4. Moreover, eating
behavior may influence the amount of food taken
per mouthful, then altering the rate of digestion and
absorption5. Mastication is an essential part of the digestive
process, because this is when food is fragmented
into smaller particles and mixed with saliva, making
it possible to swallow and digest1. During the masti
cation process, contraction of various muscle groups
occurs, generating pressure between the tooth
cusps, which breaks down the food2. The condition
of the dentition, such as the number of teeth present
and in contact, the size of functional areas, and the With the aim of evaluating quality of masticatory
function, the majority of studies found in the liter
ature used objective evaluation methods, such as
measurements of performance and efficiency6,7. It is
believed that one type of method does not exclude
the other, and an overall evaluation of the subject is
desirable, particularly when it concerns the evalu
ation of individuals who are undergoing dental and/
or orofacial treatment or who present feeding and/
or swallowing disorders, as a way of verifying the
impact of certain dental conditions or orofacial
motricity have on masticatory quality. Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1930 Hilasaca-Mamani M, Barbosa TS, Feine J, Ferreira RI, Boni RC, Castelo PM Previous studies aimed to evaluate masticatory
ability by means of questionnaires related to his/
her masticatory capacity, food preference and
consistency8-10. Some have used the Visual Analog
Scale (VAS) to make a more generalized evalu
ation, in which the subject attributes a visual score
that varies from 0 to 10 to questions such as “How
well do you chew?”, or “Do you feel any discomfort
when you chew?”6,8,11. A simple evaluation has
also been found, such as “Are you satisfied with
your chewing capacity?”, with dichotomous YES/
NO responses12. A questionnaire (Questionnaire
D’Alimentation) was developed for French speaking
subjects who wore partial/complete dentures living
in Montreal, Quebec13,14. This questionnaire consists
of 38 questions, with 29 questions being specifically
related to the frequency of and difficulty chewing
different types of foods during the two weeks before
the evaluation. adolescents, thereby obtaining a questionnaire that
can be used to evaluate the quality of masticatory
function in Brazilian Portuguese language. Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938
METHODS This study was approved by the Research
Ethics Committee of the Piracicaba Dental School,
University of Campinas (UNICAMP), Protocol
Number 108/2012. Those responsible for the
subjects signed the Informed Consent. It is important
to point out that before the translation processes
began, the authors of Questionnaire D’Alimentation
were contacted (JF). The Questionnaire D’Alimentation consists of
38 questions, and the domains “Food-Mastication”,
“Habits”,
“Meats”,
“Fruits”
and
“Vegetables”
comprise 29 questions specifically related to the
frequency of and difficulty with mastication of foods
of different types of consistency, in the two weeks
preceding the subject’s evaluation. The remaining
nine questions relate to diet specificities (for
example, appetite, allergies, reflux, nausea, and
other). Each question offers five possible response
indicators (5-Likert), according to the content of
the question, whether it is about the intensity of
the difficulty, (domain “Food-Mastication”) or about
the frequency of consuming a certain food (other
domains). In addition, the domains “Meats”, “Fruits”
and “Vegetables” also present an alternative to be
checked (not applicable - N/A) if the subject does
not usually eat these foods. Difficulty with mastication is the most probable
mechanism by which poor dental health conditions
or feeding/swallowing disorders may affect food
intake, particularly foods with greater consistency,
thereby leading to inadequate intake and having
a negative impact on nutritional status. A previous
study has shown that the higher the number of teeth
affected by caries, the greater the chance of dissat
isfaction with mastication12. It was also observed
that subjects who have masticatory problems
also present a higher DMFT (number of decayed,
missing and filled teeth), fewer functional teeth,
and orthodontic treatment need, both in adults and
children9,12,15. To measure chewing ability in relation
to diet, it is important to gain an understanding of
how the subject prepares and chooses their foods. Translation and cross-cultural adaptation of the
Questionnaire D’Alimentation were performed in
the following steps, proposed by Guillemin, et al.16:
initial translation, back-translation, revision by a
committee of specialists, and cultural adaptation
(Figure 1). To our knowledge, there are very few instru
ments with the aim of evaluating chewing ability
specifically. Thus, the aim of the present study was
to translate and cross-culturally adapt the instrument
Questionnaire
D’Alimentation
for
Brazilian Rev. CEFAC.
METHODS 2015 Nov-Dez; 17(6):1929-1938 1931 Quality of masticatory function questionnaire Original Version
Final Version
V5
Back-Translation
V4
Back-Translation
V3
Pre-test (n=20)
V5 in Brazilian Portuguese
Revision by Committee
Translation V2
Translation V1
Figure 1 – Stages of the processes of translation
and cultural adaptation of the instrument
Questionnaire D’Alimentation Cultural Equivalence of the Instrument Finally, to evaluate the cultural equivalence
of the instrument, the version V5 in Brazilian
Portuguese
(Quality
of
Masticatory
Function
Questionnaire - QMFQ) was self-applied by 20
adolescents18 from public schools of Piracicaba
(SP, Brazil), 13 (n=12) and 14 years of age (n=8),
of both genders (10♂/10♀), selected by means of
a draw from all authorizations received, under the
supervision of two researchers (MHM and TSB). In version V5 the option “I didn’t understand” was
added as an alternative response to all questions,
as a way of identifying questions that had not been
adequately understood. The percentage of “I didn’t
understand” responses had to be lower than 15%
so that the instrument could be considered culturally
adapted19. If the established limit was exceeded,
the instrument would have to be submitted to a new
cultural adaptation process, until the item “I didn’t
understand” had not been chosen in any question
by 85% or more of the adolescents. Revision by a Committee of Specialists and
Cultural Adaptation Original Version The versions V1 and V2 in Brazilian Portuguese
and the Versions V3 and V4 in French, as well
as the original instrument, were submitted to a
Reviser Committee formed by two French language
professors (one being native and the other a
certified translator), a Speech Therapist – University
Professor (RCB) and a Doctor of Dental Surgery -
University Professor (PMC). This stage consisted of
the following aspects17: •
Semantic equivalence: This refers to the meaning
of words; words that do not have a literal trans
lation with a similar meaning were translated
into terms in Brazilian Portuguese that had an
equivalent meaning; V5 in Brazilian Portuguese •
Idiomatic equivalence: The formulation of collo
quial expressions equivalent to those in the
original language; •
Cultural
equivalence
of
each
question:
experiences undergone within the cultural
context of the society. In this stage, the cultural adaptation was
performed, based on the target population of the
research; that is, Brazilian adolescents who were
not denture wearers. Therefore, three questions
related to denture wearing were excluded from the
domain “Habits” (questions 19, 23 and 24). Thus
version V5 in Brazilian Portuguese was obtained. Final Version
V5 Figure 1 – Stages of the processes of translation
and cultural adaptation of the instrument
Questionnaire D’Alimentation Initial Translation The version in French (original questionnaire)
was initially translated to Brazilian Portuguese
by a Professor with a degree in Literary Arts, as
well as a Doctor of Dental Surgery and University
Professor (RIF), both fluent in French and Brazilian
Portuguese, and aware of the aim of this study,
emphasizing conceptual translation rather than
literal translation (versions in Brazilian Portuguese
V1 and V2). Back-translation The versions in Brazilian Portuguese V1 and V2
went through back-translation into French, done by
two native French teachers, who did not participate
in the first stage of translation, and who had no
access to the original instrument, thus obtaining
the translations in French V3 and V4. The purpose
of back-translation is to compare the back transla
tioned version with the original instrument. For assessment of test-retest reliability, the same
volunteers were invited to fill out a second copy
of the questionnaire one week later for Intraclass
Correlation Coefficient (ICC) determination using
the BioEstat 5.3 (Mamirauá, Belém, PA, Brazil)
statistical software package. Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1932 Hilasaca-Mamani M, Barbosa TS, Feine J, Ferreira RI, Boni RC, Castelo PM Initial Translation and back-translation Each version in Brazilian Portuguese (V1 and
V2) was independently re-translated into French
(back-translation) by two native French teachers,
who were not aware of the purpose of the work,
thereby giving rise to the origin of versions V3 and
V4 in French. Revision by a Committee of Specialists
RESULTS original instrument were submitted to a Reviewing
Committee. Figure 2 presents the questions in the
original version and their translations, as well as a
synopsis of the decision making process relative
to specification of the first and second version in
Portuguese (V1 and V2) made by the committee. For some questions, the translations made by both
translators were identical, or practically identical;
for others, one or other version was prioritized;
moreover, in other questions, the option taken was
to combine the two versions, generating a version
of consensus, with the purpose of obtaining greater
clarity of the item. Initial Translation and back-translation Figure 2 – Synopsis of the decision-making processes regards specification of the versions V1 and
V2 in Portuguese for construction of the final instrument V1, choice of the first translator’s version; V2, choice of the second translator’s version; V3, choice of a modified version; V1≈V2,
similarity between the two versions with specificity for the first translator; V1≈V2, similarity between the two versions with specificity for
the second translator. Revision by a Committee of Specialists The versions V1 and V2 in Brazilian Portuguese
and the versions V3 and V4 in French, as well as the Original Version
Version
Committee
Question
Domain
Terms
10
Food-mastication
La grosseur d´un dé à coudre
V2
Beef/small pieces
11
Food-mastication
La grosseur d´un dé à coudre
V2
Small pieces
12
Food-mastication
Viande hachée
V1≈V2
13
Food-mastication
Croquer
V3
Bite
14
Food-mastication
Croquer
V3
Bite
15
Food-mastication
Croquer
V3
Bite
16
Food-mastication
La pelure des fruits
V1≈V2
17
Food-mastication
Pain croûté
V3
Bread with a hard crust
18
Food-mastication
Noix et des graines
V1
19
Habits
L´une ou l´autre de vos prothèses
V1
Your denture
20
Habits
Boire en mangeant
V3
Drink while eating
21
Habits
La sauce a vos aliments
V1
22
Habits
Trempé
V1
23
Habits
Prothèses
V1
Denture
24
Habits
Prothèses
V1
Denture
25
Habits
Bien màchés
V1
26
Meats
La grosseur d´un dé à coudre
V3
Small pieces
27
Meats
Hacher
V3
Shred
28
Meats
La grosseur d´un dé à coudre
V3
Small pieces
29
Meats
Hacher
V3
Shred
30
Meats
Metre la viande em purée
V3
Cook it until it falls apart
31
Fruits
Croqué
V1
Bit
32
Fruits
D´enlever la pelure
V1≈V2
Peel
33
Fruits
Couper em quartiers
V1≈V2
34
Fruits
La grosseur d´un dé à coudre
V2
Small pieces
35
Fruits
Mettre em purée
V1
Mash or grate
36
Vegetables
Croqué
V3
Bit
37
Vegetables
La grosseur d´un dé à coudre
V2
Small pieces
38
Vegetables
Mettre em purée
V2
Make puré
V1 choice of the first translator’s version; V2 choice of the second translator’s version; V3 choice of a modified version; V1≈V2 Figure 2 – Synopsis of the decision-making processes regards specification of the versions V1 and
V2 in Portuguese for construction of the final instrument Figure 2 – Synopsis of the decision-making processes regards specification of the versions V1 and
V2 in Portuguese for construction of the final instrument Figure 2 – Synopsis of the decision-making processes regards specification of the versions V1 and
V2 in Portuguese for construction of the final instrument Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1933 Quality of masticatory function questionnaire At this stage, replacements were made of terms
presented in V1 and V2 by synonyms, so that the
terms would be better understood by the target
population. Questions that sought to specify the
size of the food (meats, fruits and vegetables) were
those that generated the greatest difficulty. Revision by a Committee of Specialists For
example, the literal translation of the expression
“la grosseur d´un dé à coudre” would be “the
thickness of a thumb”. The second translator had
also suggested the term “inch”, which is commonly
used by the Brazilian adult population. However, the
target population (adolescents) may not know the
size of an “inch” and so the Committee suggested
the use of the expression “small pieces”. of the difficulty (extreme, very, moderate, hardly, no
difficulty) and frequency (never, rarely, sometimes,
frequently, always, not applicable). For the first
item, the option was to choose the modified version
(V3). For the frequency scale, the option was for
the V1; however, the term “à l´occasion”, which
was translated as ”occasionally”, was changed to
“sometimes” by the Committee, which is the word
more routinely used. After conclusion of the process of translation and
revision by the Committee, three questions related
to denture wearing were excluded from the domain
“Habits” (questions 19, 23 and 24). In questions 13, 14, 15, 31, and 36 the term
“croquer” was used, which was translated as “grind”
in V1 and as “chew” in V2. The Committee therefore
suggested the use of “bite”, because this would
linguistically be more accept and easier to under
stand, since “chew” would be translated into French
as “mastiquer”. Cultural Equivalence of the Instrument Mash or grate hard raw fruits to eat them
N/A
N/A
Vegetables
24. Bite into whole raw carrots
X
25. Cut raw carrots into small pieces to chew them
X
N/A
26. Make a puré of hard vegetables to eat them
X
N/A
Figure 3 – Pre-Test Synopsis: Description of the questions with alternative responses “not applicable”
(N/A) or left blank (X) checked by six subjects
Este questionário visa avaliar sua escolha por alimentos em função de sua capacidade de mastigar
nas duas últimas semanas. ALIMENTAÇÃO-MASTIGAÇÃO
Extrema
Muita
Moderada
Pouca
Nenhuma
dificuldade
1. Você tem dificuldade para mastigar carne de vaca cortada em
pedaços pequenos? □ (Assinale aqui se você não come carne de vaca)
□
□
□
□
□
2. Você tem dificuldade para mastigar frango cortado em pedaços
pequenos? □ (Assinale aqui se você não come frango)
□
□
□
□
□
3. Você tem dificuldade para mastigar carne moída? □ (Assinale aqui se você não come carne moída)
□
□
□
□
□
4. Você tem dificuldade para morder legumes duros, crus, inteiros
(exemplo: cenouras)? □
□
□
□
□
5. Você tem dificuldade para morder frutas duras, cruas, inteiras
(exemplo: maçãs)? □
□
□
□
□
6. Você tem dificuldade para morder frutas duras, cruas, cortadas
em quatro (exemplo: maçãs)? □
□
□
□
□
7. Você tem dificuldade para comer a casca de frutas duras, cruas? □
□
□
□
□
8. Você tem dificuldade de mastigar pão com casca dura? □
□
□
□
□
9. Você tem dificuldade de mastigar nozes e grãos? □
□
□
□
□
HÁBITOS
Nas duas últimas semanas:
Nunca
Raramente
Às vezes
Frequentemente
Sempre Individual
(gender, age)
Domain
Question
4
(♀,13)
7
(♀,13)
8
(♂,13)
9
(♂,13)
10
(♂,13)
16
(♂,14)
Meats
15. Shred beef before eating it
N/A
16. Cut chicken into small pieces
N/A
17. Shred chicken before eating it
N/A
18. Cook meat until it was tender before eating it
N/A
Fruits
19. Bite into whole raw apples
N/A
N/A
X
N/A
20. Peel apples before eating them
N/A
N/A
N/A
21. Cut apples into quarters to chew them
N/A
N/A
N/A
22. Cut apples into small pieces to chew them
N/A
N/A
N/A
23. Mash or grate hard raw fruits to eat them
N/A
N/A
Vegetables
24. Bite into whole raw carrots
X
25. Cultural Equivalence of the Instrument The use of the “Quality of Masticatory Function
Questionnaire” was considered easy, fast and the
questions were well understood by the subjects,
since the alternative “I didn’t understand” was not
checked by any of the participants. On the other
hand, of the 20 adolescents who participated in
pre-test, six checked the alternative response N/A
(not applicable) to at least one question in the
domains “Meats”, “Fruits” or “Vegetables” (Figure
3). Of these six participants, only one handed in the
questionnaire with the responses left blank, being
one question with reference to the domain “Fruits”
(19: Bite into whole raw apples) and three questions
of the domain “Vegetables”. Question 17 asks whether the subject has difficult
with chewing “pain croûté”. This expression was
translated by both translators as “toasted bread”;
whereas for the Committee this translation was not
acceptable, and it was replaced by “bread with a
hard crust”. The term “hacher” (questions 27 and 29)
was translated as ”chop” (V1) and ”mince” (V2), and
it was afterwards changed by the Committee, which
proposed the term ”shred” (the meat). Question 30
also generated doubts because of the expression
“metre la viande em purée”, and it was translated as
“Boiling the meat till it was tender”. The test-retest reliability of the questionnaire
was assessed for each domain, and the ICCs found
were: 0.79 (Food-mastication), 0.45 (Habits), 0.62
(Meats), 0.74 (Fruits) and 0.81 (Vegetables), showing
moderate to excellent agreement (p<0.01)20. The
final version of the instrument is shown in Figure 4. As regards the alternative scales to be chosen,
there were two to be translated: the scale of intensity Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1934 Hilasaca-Mamani M, Barbosa TS, Feine J, Ferreira RI, Boni RC, Castelo PM 1934 Hilasaca Mamani M, Barbosa TS, Feine J, Ferreira RI, Boni RC, Castelo PM
Individual
(gender, age)
Domain
Question
4
(♀,13)
7
(♀,13)
8
(♂,13)
9
(♂,13)
10
(♂,13)
16
(♂,14)
Meats
15. Shred beef before eating it
N/A
16. Cut chicken into small pieces
N/A
17. Shred chicken before eating it
N/A
18. Cook meat until it was tender before eating it
N/A
Fruits
19. Bite into whole raw apples
N/A
N/A
X
N/A
20. Peel apples before eating them
N/A
N/A
N/A
21. Cut apples into quarters to chew them
N/A
N/A
N/A
22. Cut apples into small pieces to chew them
N/A
N/A
N/A
23. FRUTAS FRUTAS
Nas duas últimas semanas:
Nota: Se você não comeu essa fruta, assinale a alternativa N/A (não aplicável). Nunca
Raramente
Às vezes
Frequentemente
Sempre
N/A
19. Você mordeu maçãs cruas, inteiras? □
□
□
□
□
□
20. Foi necessário descascar as maçãs antes
de comê-las? □
□
□
□
□
□
21. Foi necessário cortar as maçãs em quatro
para mastigá-las? □
□
□
□
□
□
22. Foi necessário cortar as maçãs em pedaços
pequenos para mastigá-las? □
□
□
□
□
□
23. Foi necessário amassar ou ralar as frutas
duras cruas para comê-las? □
□
□
□
□
□ CARNES CARNES
Nas duas últimas semanas:
Nota: Se você não comeu carne, assinale a alternativa N/A (não aplicável). Nunca
Raramente
Às vezes
Frequentemente
Sempre
N/A
14. Foi necessário cortar a carne de vaca em
pedaços pequenos? □
□
□
□
□
□
15. Foi necessário desfiar a carne de vaca
antes de comê-la? □
□
□
□
□
□
16. Foi necessário cortar o frango em pedaços
pequenos? □
□
□
□
□
□
17. Foi necessário desfiar o frango antes de
comê-lo? □
□
□
□
□
□
18. Foi necessário cozinhar a carne até des
manchar antes de comê-la? □
□
□
□
□
□ Se você não comeu carne, assinale a alternativa N/A (não aplicável). 1935 Quality of masticatory function questionnaire Nunca
Raramente
Às vezes
Frequentemente
Sempre
11. Você adicionou molho aos seus alimentos para
engolir melhor? □
□
□
□
□
12. Você molhou os alimentos em líquidos para masti
gar e engolir melhor? □
□
□
□
□
13. Em geral, os alimentos que você engole são bem
mastigados? □
□
□
□
□ Cultural Equivalence of the Instrument Cut raw carrots into small pieces to chew them
X
N/A
26. Make a puré of hard vegetables to eat them
X
N/A
Figure 3 – Pre-Test Synopsis: Description of the questions with alternative responses “not applicable”
(N/A) or left blank (X) checked by six subjects Este questionário visa avaliar sua escolha por alimentos em função de sua capacidade de mastigar
nas duas últimas semanas. ALIMENTAÇÃO-MASTIGAÇÃO
Extrema
Muita
Moderada
Pouca
Nenhuma
dificuldade
1. Você tem dificuldade para mastigar carne de vaca cortada em
pedaços pequenos? □ (Assinale aqui se você não come carne de vaca)
□
□
□
□
□
2. Você tem dificuldade para mastigar frango cortado em pedaços
pequenos? □ (Assinale aqui se você não come frango)
□
□
□
□
□
3. Você tem dificuldade para mastigar carne moída? □ (Assinale aqui se você não come carne moída)
□
□
□
□
□
4. Você tem dificuldade para morder legumes duros, crus, inteiros
(exemplo: cenouras)? □
□
□
□
□
5. Você tem dificuldade para morder frutas duras, cruas, inteiras
(exemplo: maçãs)? □
□
□
□
□
6. Você tem dificuldade para morder frutas duras, cruas, cortadas
em quatro (exemplo: maçãs)? □
□
□
□
□
7. Você tem dificuldade para comer a casca de frutas duras, cruas? □
□
□
□
□
8. Você tem dificuldade de mastigar pão com casca dura? □
□
□
□
□
9. Você tem dificuldade de mastigar nozes e grãos? □
□
□
□
□ nário visa avaliar sua escolha por alimentos em função de sua capacidade de mastig
mas semanas. Este questionário visa avaliar sua escolha por alimentos em função de sua capacidade de mastigar
nas duas últimas semanas. 1935
DISCUSSION Originally it was proposed that this instrument
should be self-applied, being necessary for the
respondent to read the instructions carefully before
answering the questions; therefore, future studies
will be able to show whether there is equivalence. In
this study, it was considered feasible for researchers
to provide initial instructions to the subjects, with
the aim to ensure that the rules for filling out the
questionnaire were understood by the studied
sample. For responses left blank or for alternative
response N/A, the missing value may be replaced
by the mean/median scores for each domain or for
each subject28,29. An instrument may only be considered valid
if it is capable of accurately capturing a certain
concept21. Furthermore, in the culture for which it
is being adapted, a translated instrument must be
capable of obtaining the same effect as the original
instrument has in the context in which it was created. The lack of cultural equivalence compromises
the validity of the information collected, making it
impossible to use the instrument to study a concept
correctly22. This is why there are standardized
instructions that are intended to minimize the loss
of the original instrument characteristics that could
result from the change in language16,22-24. Therefore,
the methodology of the present study followed the
protocol suggested by Guillemim, et al.16, by which
mistakes and misinterpretations in the initial transla
tions are revealed. During these stages, semantic
equivalence was appreciated, and not the literal
interpretation, between the terms, since the literal
interpretation is not always shown to be more
advantageous to express concepts or situations of
the new population one wishes to study16,24-26. The
process
of
translation
and
cultural
adaptation of health-related questionnaires must
keep a conceptual basis and follow standardized
methodologies, since differences in subject’s
habits and culture may lead to misunderstandings,
thereby altering the psychometric properties of
the instrument. Therefore, in order to fully achieve
cultural adaptation, it is also necessary to conduct a
study of equivalence measurement, evaluating the
reliability and validity of this new version in different
populations and conditions (e.g. subjects with
dental caries and/or periodontal disease, patients
presenting feeding and/or swallowing disorders,
denture users, patients undergoing orthognathic or
bariatric surgery, and others). In this process, using and comparing more than
one version is relevant. LEGUMES Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938
Nas duas últimas semanas:
Nota: Se você não comeu esse legume, assinale a alternativa N/A (não aplicável). Nunca
Raramente
Às vezes
Frequentemente
Sempre
N/A
24. Você mordeu cenouras cruas inteiras? □
□
□
□
□
□
25. Foi necessário cortar as cenouras cruas em
pedaços pequenos para mastigá-las? □
□
□
□
□
□
26. Foi necessário fazer purê com os legumes
duros para comê-los? □
□
□
□
□
□
OBRIGADO POR SUA COLABORAÇÃO! The English-version of this questionnaire is available at: Muller K, Morais J, Feine J. Nutritional and anthropometric analysis of eden
tulous patients wearing implant overdentures or conventional dentures. Braz Dent J. 2008;19:145-50. Figure 4 – QUESTIONÁRIO DE AVALIAÇÃO DA QUALIDADE DA MASTIGAÇÃO (QAQM) Nas duas últimas semanas:
Nota: Se você não comeu esse legume, assinale a alternativa N/A (não aplicável). Nunca
Raramente
Às vezes
Frequentemente
Sempre
N/A
24. Você mordeu cenouras cruas inteiras? □
□
□
□
□
□
25. Foi necessário cortar as cenouras cruas em
pedaços pequenos para mastigá-las? □
□
□
□
□
□
26. Foi necessário fazer purê com os legumes
duros para comê-los? □
□
□
□
□
□
OBRIGADO POR SUA COLABORAÇÃO! Se você não comeu esse legume, assinale a alternativa N/A (não aplicável). OBRIGADO POR SUA COLABORAÇÃO! The English-version of this questionnaire is available at: Muller K, Morais J, Feine J. Nutritional and anthropometric analysis of eden
tulous patients wearing implant overdentures or conventional dentures. Braz Dent J. 2008;19:145-50. Figure 4 – QUESTIONÁRIO DE AVALIAÇÃO DA QUALIDADE DA MASTIGAÇÃO (QAQ Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1936 Hilasaca-Mamani M, Barbosa TS, Feine J, Ferreira RI, Boni RC, Castelo PM
DISCUSSION In addition to making it
possible to choose items to incorporate, or to allow the
junction of items arising from different versions, this
strategy allows to carefully examine the sequence
of the procedures, including the translations
themselves, their re-translation and the appreciation
that follows. The importance of a general criticism
made by the committee of specialists must also be
pointed out; among the members of the committee
there must be individuals who are specialists in the
disease investigated, in the measure used and in
the concept explored, and they should preferably be
bilingual16.
CONCLUSION The Portuguese version of the Questionnaire
D’Alimentation has shown to be easy to understand
by Brazilian adolescents and useful in the evalu
ation of the masticatory function and feeding or
swallowing disorders that may affect food intake. The differences between definitions, beliefs and
behaviors demand that the use of an instrument
drawn up in other cultural contexts should, in addition
to reliable translation, be preceded by cross-cultural
adaptation to the country in which it will be applied,
in order to keep the same concepts as those of the
original27. At the stage of cross-cultural adaptation,
the number and characteristics of volunteers
involved in pre-test were chosen in agreement with
methodological criteria used for cultural adaptation
of questionnaires16,18. RESUMO Objetivo: traduzir o instrumento em língua francesa “Questionnaire D’Alimentation” para o português
brasileiro e realizar a adaptação transcultural em adolescentes brasileiros. Métodos: a tradução foi
realizada com base em protocolo que consistiu na tradução para o português brasileiro (por um
professor formado em Letras e uma Cirurgiã Dentista fluentes em francês), tradução reversa para
o francês (realizada por dois professores de francês), revisão por Comitê de especialistas (duas
tradutoras, uma nativa e uma juramentada, e duas professoras universitárias, sendo uma Cirurgiã
Dentista e uma Fonoaudióloga) e equivalência cultural (pré-teste). A versão utilizada no pré- teste
consistiu de 26 questões com cinco respostas possíveis (5-Likert), distribuídas em cinco domínios
(Alimentação- mastigação, Hábitos, Carnes, Frutas e Legumes). O pré-teste e teste-reteste foram
realizados com uma amostra de 20 adolescentes (10 meninos/10 meninas) de escolas públicas de
Piracicaba (Brasil). Nesta fase, a alternativa “não entendi” foi incluída a cada questão, a fim de iden
tificar aquelas não compreendidas. A confiabilidade teste-reteste foi avaliada para cada domínio utili
zando os coeficientes de correlação intra-classe (CCI). Resultados: no pré-teste, foi observada uma
excelente compreensão do instrumento; no teste-reteste, os CCIs variaram entre 0,45-0,81 (mode
rada a excelente concordância). Conclusão: a versão em Português do Questionnaire D’Alimentation
mostrou ser de fácil compreensão por parte dos adolescentes brasileiros e útil na avaliação da função
mastigatória e dos distúrbios de alimentação e deglutição que podem alterar a ingestão de alimentos. DESCRITORES: Tradução; Mastigação; Alimentos; Adolescente
ACKNOWLEDGEMENTS The authors would like to thank Prof. Fábio Nunes
da Silva for his help in the translation process. This study was supported by the State of São
Paulo Research Foundation (FAPESP, SP, Brazil, n. 2012/04492-2). Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1937 Quality of masticatory function questionnaire RESUMO
Objetivo: traduzir o instrumento em língua francesa “Questionnaire D’Alimentation”
brasileiro e realizar a adaptação transcultural em adolescentes brasileiros. Método
realizada com base em protocolo que consistiu na tradução para o português b
professor formado em Letras e uma Cirurgiã Dentista fluentes em francês), tradu
o francês (realizada por dois professores de francês), revisão por Comitê de es
tradutoras, uma nativa e uma juramentada, e duas professoras universitárias, sen
Dentista e uma Fonoaudióloga) e equivalência cultural (pré-teste). A versão utiliza
consistiu de 26 questões com cinco respostas possíveis (5-Likert), distribuídas em
(Alimentação- mastigação, Hábitos, Carnes, Frutas e Legumes). O pré-teste e te
realizados com uma amostra de 20 adolescentes (10 meninos/10 meninas) de esc
Piracicaba (Brasil). Nesta fase, a alternativa “não entendi” foi incluída a cada questã
tificar aquelas não compreendidas. A confiabilidade teste-reteste foi avaliada para c
zando os coeficientes de correlação intra-classe (CCI). Resultados: no pré-teste, fo
excelente compreensão do instrumento; no teste-reteste, os CCIs variaram entre 0
rada a excelente concordância). Conclusão: a versão em Português do Questionnai
mostrou ser de fácil compreensão por parte dos adolescentes brasileiros e útil na ava
mastigatória e dos distúrbios de alimentação e deglutição que podem alterar a inges
DESCRITORES: Tradução; Mastigação; Alimentos; Adolescente RESUMO
Objetivo: traduzir o instrumento em língua francesa “Qu
brasileiro e realizar a adaptação transcultural em adoles
realizada com base em protocolo que consistiu na tra
professor formado em Letras e uma Cirurgiã Dentista f
o francês (realizada por dois professores de francês),
tradutoras, uma nativa e uma juramentada, e duas pro
Dentista e uma Fonoaudióloga) e equivalência cultural
consistiu de 26 questões com cinco respostas possíve
(Alimentação- mastigação, Hábitos, Carnes, Frutas e L
realizados com uma amostra de 20 adolescentes (10 m
Piracicaba (Brasil). Nesta fase, a alternativa “não entend
tificar aquelas não compreendidas. A confiabilidade teste
zando os coeficientes de correlação intra-classe (CCI). R
excelente compreensão do instrumento; no teste-reteste
rada a excelente concordância). Conclusão: a versão em
mostrou ser de fácil compreensão por parte dos adolesce
mastigatória e dos distúrbios de alimentação e deglutição
DESCRITORES: Tradução; Mastigação; Alimentos; Ado DESCRITORES: Tradução; Mastigação; Alimentos; Adolescente
REFERENCES comparisons of implant–supported mandibular
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A. Mastication and swallowing: influence of fluid
addition to foods. J Appl Oral Sci. 2007;15:55-60. 9. Hassan AH, Amin Hel-S. Association of orthodontic
treatment needs and oral health-related quality of
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RMS. Masticatory performance in adults related to
temporomandibular disorder and dental occlusion. Pró-Fono R Atual Cient. 2007;19:151-8. 10. Souza JF, Grechi TH, Anselmo-Lima WT,
Trawitzki LV, Valera FC. Mastication and deglutition
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Care Dentist. 1998;18:33-9. 11. Prado MMS, Borges TF, Prado CJ, Gomes
VL, Neves FD. Masticatoy function of subjects
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Gavião MB, Castelo PM. Assessment of masticatory
performance, bite force, orthodontic treatment
need and orofacial dysfunction in children and
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Traebert J, Panizzi M. Impact of dental caries and
dental fluorosis on 12-year-old schoolchildren’s
self-perception of appearance and chewing. Cad
Saúde Pública. 2003;19:323-30. 5. Sun L, Ranawana DV, Tan WJ, Quek YC, Henry
CJ. The impact of eating methods on eating rate and
glycemic response in healthy adults. Physiol Behav. 2014;139C:505-10. 6. Katsuhiko K, Takahiro O, Garrett NR, Minoru
T. Assessment of masticatory performance –
methodologies and their application. Prosthodont
Res Pract. 2004;3:33-45. 13. Muller K.The mandibular implant overdenture
versus the mandibular conventional denture:impact
on the nutritional status [dissertation] Montreal
(Quebec, Canada): Faculty of Dentistry; McGill
University; 1999. 7. Magalhães IB, Pereira LJ, Marques LS, Gameiro
GH. The influence of malocclusion on masticatory
performance. A systematic review. Angle Orthod. 2010;80:981-7. 14. Muller K, Morais J, Feine J. Nutritional and
anthropometric analysis of edentulous patients
wearing implant overdentures or conventional
dentures. Braz Dent J. 2008;19:145-50. 8. de Grandmont P, Feine JS, Taché R, Boudrias
P, Donohue WB, Tanguay R et al. Within–subject Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 1938 Hilasaca-Mamani M, Barbosa TS, Feine J, Ferreira RI, Boni RC, Castelo PM screening of violence against pregnant women. Rev
Saude Publ. 2000;34:610-6. 15. Cushing AM, Sheiham A, Maizels J. Developing
socio-dental indicators the social impact of dental
disease. Community Dent Health. 1986;3:3-17. 23.
REFERENCES da Mota Falcão D, Ciconelli RM, Ferraz
MB.Translation and cultural adaptation of quality of
life questionnaires:an evaluation of methodology. J
Rheumatol. 2003;30:379-85. 16. Guillemin F, Bombardier C, Beaton D. Cross-
cultural adaptation of health-related quality of
life measures: literature review and proposed
guidelines. J Clin Epidemiol. 1993;46:1417-32. 24. Herdman M, Fox-Rushby J, Badia X. “Equivalence” and the translation and adaptation of
Health-Related Quality of Life Questionnaires. Qual
Life Res. 1997;6:237-47. 17. Beaton DE, Bombardier C, Guillemin F,
Ferraz MB.Guidelines for the process of cross-
cultural adaptation of self-report measures. Spine. 2000;25:3186-91. 25. Reichenheim ME, Moraes CL.Pillars for
assessing validity in epidemiological studies. Rev
Bras Epidemiol. 1998;1:131-48. 18. Yusuf H, Gherunpong S, Sheiham A, Tsakos G. Validation of an English version of the Child-OIDP
index, an oral health-related quality of life measure
for children. Health Qual Life Outcomes. 2006;4:38. 26. Tesch FC, Oliveira BH, Leão A. Semantic
equivalence of the Brazilian version of the Early
Childhood Oral Health Impact Scale. Cad Saude
Publica. 2008;24:1897-909. 19. Ciconelli RM, Ferraz MB, Santos W, Meinão I,
Quaresma MR. Brazilian-Portuguese version of the
SF-36. A reliable and valid quality of life outcome
measure. Rev Bras Reumatol. 1999;39:143-50. 27. Barbosa TS, Gavião MB.Quality of life and oral
health in children - Part II: Brazilian version of the
Child Perceptions Questionnaire 11-14. Cien Saude
Colet. 2011;16:3267-76. 20. Fleiss JL. Reliability of Measurement. In: Fleiss
JL. The design and analysis of clinical experiments. New York: Wiley; 1986. pp. 1-32. 28. Shrive FM, Stuart H, Quan H, Ghali WA.Dealing
with missing data in a multi-question depression
scale:a comparison of imputation methods. BMC
Med Res Methodol. 2006;6,57. 21. Barbosa TS, Serra MD, Gavião MBD.Quality
of life and oral health in children - Part I: Brazilian
version of the Child Perceptions Questionnaire
8-10. Cien Saude Colet. 2011;16:4077-85. 21. Barbosa TS, Serra MD, Gavião MBD.Quality
of life and oral health in children - Part I: Brazilian
version of the Child Perceptions Questionnaire
8-10. Cien Saude Colet. 2011;16:4077-85. 22. Reichenheim ME, Moraes CL, Hasselmann MH. Semantic equivalence of the Portuguese version
of the Abuse Assessment Screen tool used for the 29. Eekhout I, de Boer RM, Twisk JW, de Vet HC,
Heymans MW. Missing data: a systematic review of
how they are reported and handled. Epidemiology. 2012;23:729-32. 22. Reichenheim ME, Moraes CL, Hasselmann MH. Semantic equivalence of the Portuguese version
of the Abuse Assessment Screen tool used for the http://dx.doi.org/10.1590/1982-021620151764715
Received on: April 13, 2015
Accepted on: September 11, 2015 Mailing address:
Paula Midori Castelo
Depto. Ciências Biológicas - Universidade Federal
de São Paulo (UNIFESP)
R. São Nicolau, 210 – 1. Andar
Diadema – SP – Brasil
CEP: 09913-030
E-mail: pcastelo@yahoo.com Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938 Rev. CEFAC. 2015 Nov-Dez; 17(6):1929-1938
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W3023666651.txt
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https://zenodo.org/records/1669673/files/article.pdf
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en
|
Optical Activity and Chemical Composition
|
Science
| 1,900
|
public-domain
| 660
|
SCIENCE.
[N.S. VOL.XI. NO. 282.
taken up isomeric compounds, including stereoisomers, homologous series, influence of the
mode of linkage of the carbon atoms, summation of the rotatory actions of several asymmetric groups, optical superposition and the
dependence of the rotatory power of ao active
atomic grouping on the masses of the four radicles united to the asymmetric carbon atom, the
hypothesis of Guye.
The translation is well done and the subject
is brought up to date by notes and additions by
Optical Activity and Chemical Composition. By the translator. The subject is presented in a
DR. H . LANDOLT,
Professor of Chemistry in very attractive and readable form and the book
the University of Berlin. Translated, with can be heartily recommended to anyone, who
MCCRAE, desires to know the present state of our knowlthe author's permission, by JOHN
Ph.D. Whittaker and Co., London, and the edge regarding the relation existing between
Macmillan Co., 66 Fifth Ave., New York. optical activity and chemical composition ;
1899. Small8vo. Pp. 158. Price, $1.00.
though for more detailed information Landolt's
This little book forms a translation of the
Das optische Drehungsvermiigen organischer
eighth chapter of the first volume of Graham- Substanzen und dessen praktische AnwendungOtto's ' Lehrbuch der Chemie ' and is a smaller en ' must be used.
W. R. ORNDORFF.
and condensed edition of the author's wellknown ' Das optische Drehungsvermiigen organischer Substanzen und desseu praktische AnSCIENTIFIC JOURNALS A N D ARTICLES.
wendungen,' published in 1898. The subject
THE Osprey for April, a little belated, opens
is treated under three heads: I. General with the fourth part of 'Birds of the Road,' by
Principles of Optical Activity ; 11. Connection Paul Bartsch. Wm. L. Wells describes the
between the Rotatory Power and the Chemical 'Nesting of some Rare Birds,' including the
Composition of Carbon Compounds, and 111. yellow rail and solitary sandpiper, and TheoConnection between Degree of Rotation and dore Gill presents the second part of ' William
Chemical Constitution. Under the first head Swainson and his Times ' which carries Swainare discussed such subjects as crystal rotation, son through his journey to Brazil. I n editorial
liquid rotation, molecular rotation, measure- comments under ' Birds and Women ' the situment of rotation, specific rotation, variations ation is summed up in a few words ' L If the deof specific rotation with concentration and mand exists for anything, that demand will be
change of rotatory power of dissolved sub- supplied if it can be done with aprofit." Under
stances with time, multirotation. Under the Notes is to be found a n extraordinary account
second head are treated optical modifications, of ' How Two Lions stopped an African Railthe investigations of Pasteur, the van't Hoff road,' and other matters of interest.
and Le Be1 theory, calculation of the number
A Bulletin of Mathematics and of the Physical
of optically active isomers of a compound from and Natural Sciences, to be published semithe number of asymmetric carbon atoms which monthly in the interest of teachers in Italian
it contains, the formation and properties of schools, has been established by Professor
racemic compounds, resolution of racemic Alberto Conti, of Bologne.
substances into the antipodes, formation and
properties of the active modifications, transSOCIETIES AND A CADEJIIES.
formation of one antipode into the other, the
GEOLOGICAL
SOCIETY O F WASHINGTON.
configurationally inactive non-decomposable
THE
lOlst
meeting
of the Society was held a t
modifications and their differences from racemic
the
Cosmos
Club
April
11, 1900.
inactive isomers. Under the third head are
beginners. A student with some knowledge of
organic chemistry could use it as a text-book if
i t were possible for him to resolutely confine his
attention to the coarse print.' But it is as a
reference book for the student who wishes to refresh his memory not merely of one compound,
but of the complete chemistry of a group of
compounds, that the work is of peculiar value,
and may be cordially recommended.
EDWARDRENOUF.
|
|
https://openalex.org/W2010653101
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https://europepmc.org/articles/pmc3027619?pdf=render
|
English
| null |
Prioritizing Conservation of Ungulate Calving Resources in Multiple-Use Landscapes
|
PloS one
| 2,011
|
cc-by
| 15,183
|
Received June 23, 2010; Accepted December 15, 2010; Published January 26, 2011 Copyright: 2011 Dzialak et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: Pioneer Natural Resources Company funded this work. Hayden-Wing Associates, LLC provided in-kind contributions including travel costs associated
with dissemination of this work, and materials associated with data collection and analysis including radio-telemetry equipment, GPS devices, and statistical
analysis software. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: This work was funded by a commercial source, Pioneer Natural Resources Company. Hayden-Wing Associates, LLC provided in-kind
contributions including travel costs associated with dissemination of this work, and materials associated with data collection and analysis including radio-
telemetry equipment, GPS devices, and statistical analysis software. Having received funding from this source, the consultancy could reasonably be perceived as a
financial competing interest. This does not alter the authors’ adherence to all the PLoS ONE policies on sharing data and materials. ¤ Current address: New Mexico Game and Fish Department, Santa Fe, New Mexico, United States of America Matthew R. Dzialak*, Seth M. Harju, Robert G. Osborn¤, John J. Wondzell, Larry D. Hayden-Wing,
Jeffrey B. Winstead, Stephen L. Webb Hayden-Wing Associates, LLC, Natural Resource Consultants, Laramie, Wyoming, United States of America Hayden-Wing Associates, LLC, Natural Resource Consultants, Laramie, Wyoming, United States of America Hayden-Wing Associates, LLC, Natural Resource Consultants, Laramie, Wyoming, United States of America PLoS ONE | www.plosone.org Abstract Background: Conserving animal populations in places where human activity is increasing is an ongoing challenge in many
parts of the world. We investigated how human activity interacted with maternal status and individual variation in behavior
to affect reliability of spatially-explicit models intended to guide conservation of critical ungulate calving resources. We
studied Rocky Mountain elk (Cervus elaphus) that occupy a region where 2900 natural gas wells have been drilled. Methodology/Principal Findings: We present novel applications of generalized additive modeling to predict maternal
status based on movement, and of random-effects resource selection models to provide population and individual-based
inference on the effects of maternal status and human activity. We used a 262 factorial design (treatment vs. control) that
included elk that were either parturient or non-parturient and in areas either with or without industrial development. Generalized additive models predicted maternal status (parturiency) correctly 93% of the time based on movement. Human
activity played a larger role than maternal status in shaping resource use; elk showed strong spatiotemporal patterns of
selection or avoidance and marked individual variation in developed areas, but no such pattern in undeveloped areas. This
difference had direct consequences for landscape-level conservation planning. When relative probability of use was
calculated across the study area, there was disparity throughout 72–88% of the landscape in terms of where conservation
intervention should be prioritized depending on whether models were based on behavior in developed areas or
undeveloped areas. Model validation showed that models based on behavior in developed areas had poor predictive
accuracy, whereas the model based on behavior in undeveloped areas had high predictive accuracy. Conclusions/Significance: By directly testing for differences between developed and undeveloped areas, and by modeling
resource selection in a random-effects framework that provided individual-based inference, we conclude that: 1) amplified
selection or avoidance behavior and individual variation, as responses to increasing human activity, complicate conservation
planning in multiple-use landscapes, and 2) resource selection behavior in places where human activity is predictable or less
dynamic may provide a more reliable basis from which to prioritize conservation action. Citation: Dzialak MR, Harju SM, Osborn RG, Wondzell JJ, Hayden-Wing LD, et al. (2011) Prioritizing Conservation of Ungulate Calving Resources in Multiple-Use
Landscapes. PLoS ONE 6(1): e14597. doi:10.1371/journal.pone.0014597 Editor: Michael Somers, University of Pretoria, South Africa Received June 23, 2010; Accepted December 15, 2010; Published January 26, 2011 Prioritizing Conservation of Ungulate Calving Resources
in Multiple-Use Landscapes Matthew R. Dzialak*, Seth M. Harju, Robert G. Osborn¤, John J. Wondzell, Larry D. Hayden-Wing,
Jeffrey B. Winstead, Stephen L. Webb January 2011 | Volume 6 | Issue 1 | e14597 Capturing elk p
g
In February and March 2006–2009, helicopter net-gunning was
used to capture yearling (1.5 years) and adult ($2.5 years) female
elk throughout and adjacent to the gas field. Elk were fitted with
Global Positioning Systems (GPS) collars (TGW-3590, Telonics,
Inc., Mesa, AZ 85204) configured with store-on-board and Very
High Frequency (VHF) beacon options. Twenty-five female elk
were fitted with GPS collars in 2006, 40 in 2007, 50 in 2008, and
50 in 2009. GPS collars attempted to record location information
every 3 h resulting in a maximum of 8 locations/elk/day. Age of
elk was estimated based on dental eruption and wear patterns [26]. Blood samples were collected from captured elk in 2008 and 2009
to determine pregnancy, but not from elk captured in previous
years. Animal capture and handling protocols were approved by
the Colorado Division of Wildlife (Permit #s 06TR1083,
07TR1083, 08TR1083 and 09TR1083A001). Areas associated with parturition are important because female
ungulates make resource-related choices that affect offspring
development during gestation and provisioning of recently born
calves that are susceptible to malnutrition and predation [12], [13]. While there is little evidence of consistent parturition site fidelity in
many ungulates, strong fidelity among females to seasonal ranges,
particularly around calving time, has been demonstrated [2], [14]. New light has been shed on the adaptive significance of resource
selection during the period that encompasses reproductive activity
in many vertebrates through the study of maternal effects –
developmental mechanisms by which parents translate their
environmental experience into adaptive variation in their offspring
[15], [16], [17]. The adaptive significance of resource selection
during reproductive periods suggests that conservation strategies
designed around ungulate parturition areas might be most reliable
when based on parturient females rather than samples including
both parturient and non-parturient individuals. Study area The 1845 km2 study area encompassed northern portions of the
Raton basin in south-central Colorado, USA. Topography is
rugged with steep slopes, rocky outcrops, ridges, and valleys
ranging in elevation from 2000–3000 m. Mean annual precipita-
tion is about 40–53 cm depending on elevation [23]. Vegetation
includes conifer forest, montane shrub, and grassland. Dominant
species include ponderosa pine (Pinus ponderosa), one-seed juniper
(Juniperus monosperma), two-needle pinyon (Pinus edulis), Gambel oak
(Quercus
gambelii)
which
commonly
forms
shrub-thickets
on
southern aspects, antelope bitterbrush (Purshia tridentata), skunkbush
sumac (Rhus trilobata), and willow (Salix spp.) in riparian areas. Predators of elk (including neonates) include black bear (Ursus
americanus), mountain lion (Felis concolor), and coyote (Canis latrans);
no wolf (Canis lupus) pack occurred in the study area. The study
area encompassed historic and ongoing energy development. Bituminous coal mining was a dominant land use during 1873–
1970. Coal-bed methane development was initiated in Raton
basin in 1982 and accelerated in the late 1990s [24], [25]. In 2009,
there were about 2900 wells associated with methane development
in the Basin (Figure 1). A key approach in studying wildlife-human interaction is
resource selection modeling [4]. Resource selection is a funda-
mental ecological process that structures animal movement and
distribution [5]. The choices animals make as they move
throughout the landscape reflect trade-offs between selecting
resources that meet their needs for survival and reproduction, and
minimizing perceived risk of harm – such risk often is a function of
interaction with predators or humans [6], [7], [8]. A set of
analytical methods used with increasing frequency for investigating
resource selection in animals is the estimation of RSFs [9]. RSFs
describe the relative probability of occurrence of animals as a
function of behavioral responses to features of the environment. The probability of occurrence is described as relative because
RSFs are estimated in a use-versus-availability framework in which
selection is quantified relative to available but presumed non-used
features. Environmental features can include a wide range of
variables such as vegetation, terrain, group/herd size, risk of
predation, or human-modifications of the landscape [10], [11]. RSFs have strong application in conservation planning where
wildlife-human interaction is a concern; specifically, in establishing
a spatially explicit basis from which to prioritize conservation
action such as reclamation, mitigation, or minimizing human
activity in particular habitats. Introduction Model-based methods to designate maternal
status were necessary because data bearing directly on maternal
status were unavailable for most elk. Second, we estimated group-
dependent random-effects resource selection functions (RSFs) to
identify how resource selection patterns differed relative to
maternal status and human activity, and how behavior varied
among individuals. We applied
results by developing and
validating group-dependent predictive maps of critical calving
resources, and quantifying discrepancies among maps relative to
predictive accuracy. Introduction wildlife and their habitats [3]. Understanding how human activity
such as resource extraction interacts with wildlife, and developing
tools to guide conservation planning in areas where human activity
is widespread or increasing are ongoing challenges in conservation
science. In this paper, we investigated interactions between human
activity associated with energy development and resource selection
by female Rocky Mountain elk (Cervus elaphus) during calving
season with the larger goal of informing conservation planning for
ungulates in places where human activity is widespread or
increasing. We asked: 1) what are the relative influences of
maternal status (parturiency) and human activity on resource
selection, 2) to what extent does behavior vary among individuals, Identifying resources associated with critical life-history phases
in ungulates is a conservation priority. Winter range, parturition
areas, and migration routes are important seasonal habitats in
North America that provide resources necessary for survival and
reproduction such as high-quality forage, reduced exposure to
inclement conditions, and reduced risk of predation [1], [2]. In the
Intermountain West USA, these habitats have become increas-
ingly
fragmented. Here,
energy
development
is
of
broad
conservation interest because its prominence has increased in
recent decades along with concern about its potential impact on PLoS ONE | www.plosone.org January 2011 | Volume 6 | Issue 1 | e14597 January 2011 | Volume 6 | Issue 1 | e14597 1 Ungulate Conservation situation-specific basis [22] making it difficult to generalize
behavior
across
the
population
and
thus
effectively
guide
conservation planning. 3) how does individual variation interact with human activity, and
4) how can our findings be applied in conservation planning and
decision making? First, we describe a novel application of
generalized additive models for designating maternal status using
movement data. Model-based methods to designate maternal
status were necessary because data bearing directly on maternal
status were unavailable for most elk. Second, we estimated group-
dependent random-effects resource selection functions (RSFs) to
identify how resource selection patterns differed relative to
maternal status and human activity, and how behavior varied
among individuals. We applied
results by developing and
validating group-dependent predictive maps of critical calving
resources, and quantifying discrepancies among maps relative to
predictive accuracy. 3) how does individual variation interact with human activity, and
4) how can our findings be applied in conservation planning and
decision making? First, we describe a novel application of
generalized additive models for designating maternal status using
movement data. PLoS ONE | www.plosone.org Grouping Elk Relative to Maternal Status and Human
Activity We predicted maternal status (parturient versus non-parturient)
by using generalized additive models (GAMs) to parameterize
response curves depicting daily movement of elk during calving
season. GAMs are semi-parametric extensions of generalized
linear models [27]. The central concept is that the function of a
covariate is estimated nonparametrically from the data by means
of scatterplot smoothers. The functional form of the relationship
between the response and covariate(s) is therefore determined by
the data rather than being restricted to a parametric form [28]. Formally, the linear regression model Another key feature of animal ecology that warrants further
attention as part of conservation planning is variation in behavior
among individuals. Individual variation is widespread and well-
known in many animal species [18] and can reflect long-term
selection for a given trait or learned behavior [19], [20]. Fitness is
influenced by the choices individuals make in terms of resource
selection because each resource type has particular costs and
benefits to the individual [21]. In risky or rapidly changing
environments such as those in which human activity is increasing,
optimal behavioral strategies may vary among individuals on a y~az
X
I
i~1
bi(Xi)ze is generalized by modeling y as being related to covariates
additively by is generalized by modeling y as being related to covariates
additively by January 2011 | Volume 6 | Issue 1 | e14597 2 Ungulate Conservation Figure 1. The Raton Basin gas field and adjacent areas in south-central Colorado, USA. Each natural gas well is encircled by a 1-km buffer
(shaded region). We designated elk locations occurring within the buffered region to be ‘‘inside of the gas field’’ whereas locations adjacent but
external to the buffered region were designated ‘‘outside of the gas field’’. doi:10.1371/journal.pone.0014597.g001 Figure 1. The Raton Basin gas field and adjacent areas in south-central Colorado, USA. Each natural gas well is encircled by a 1-km buffer
(shaded region). We designated elk locations occurring within the buffered region to be ‘‘inside of the gas field’’ whereas locations adjacent but
external to the buffered region were designated ‘‘outside of the gas field’’. doi:10.1371/journal.pone.0014597.g001 the effective number of parameters of a smoother) is described by
the quantity equivalent degrees of freedom (df). Span is related
inversely to df so as span increases df decreases [27]. Data that are
best described by a straight line (a single parameter comprises the
smoother) correspond to a span of 100% and thus 1 df. PLoS ONE | www.plosone.org Grouping Elk Relative to Maternal Status and Human
Activity This data summary (i.e., PLoS ONE | www.plosone.org January 2011 | Volume 6 | Issue 1 | e14597 January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org 3 Ungulate Conservation [35] [36]) W h
th i
d th t d
i
l i
ti
t
d
i ti
d il
h
i
th di t
d b
Figure 2. Movement in female ungulates during calving time. Generalized from the literature [34], [35], [36], parturient females often will
make long-distance movements associated with pre-parturient ‘‘restlessness’’ within days of parturition and then exhibit reduced movement
associated with provisioning the neonate (a). This pattern may provide a quantifiable distinction between parturient and non-parturient females. Also
shown (b) is parameterization (equivalent degrees of freedom) as a function of span based on generalized additive models of distance moved
between consecutive locations in 103 female Rocky Mountain elk. Loess smoothing and automated calculation of degrees of freedom using the
generalized cross validation method were specified. doi:10.1371/journal.pone.0014597.g002
Ungulate Conservation [35], [36]). We hypothesized that during calving time, movement
i
d
i h
l
d
li
d
ki
i
i
lk
response curves depicting dai
lk b
i
l
i
Figure 2. Movement in female ungulates during calving time. Generalized from the literature [34],
make long-distance movements associated with pre-parturient ‘‘restlessness’’ within days of parturition
associated with provisioning the neonate (a). This pattern may provide a quantifiable distinction between par
shown (b) is parameterization (equivalent degrees of freedom) as a function of span based on generaliz
between consecutive locations in 103 female Rocky Mountain elk. Loess smoothing and automated calcu
generalized cross validation method were specified. doi:10.1371/journal.pone.0014597.g002 Figure 2. Movement in female ungulates during calving time. Generalized from the literature [34], [35], [36], parturient females often will
make long-distance movements associated with pre-parturient ‘‘restlessness’’ within days of parturition and then exhibit reduced movement
associated with provisioning the neonate (a). This pattern may provide a quantifiable distinction between parturient and non-parturient females. Also
shown (b) is parameterization (equivalent degrees of freedom) as a function of span based on generalized additive models of distance moved
between consecutive locations in 103 female Rocky Mountain elk. Loess smoothing and automated calculation of degrees of freedom using the
generalized cross validation method were specified. doi:10.1371/journal.pone.0014597.g002 [35], [36]). Grouping Elk Relative to Maternal Status and Human
Activity Conversely,
data that are best described by gradients or turning points (several
or many parameters comprise the smoother) correspond to a
smaller span and thus to a larger number of df [27], [30]. The
analyst may set the span by specifying df based on visual
examination of the data, or implement generalized cross validation
methods in which appropriate df are identified automatically given
the data. From a GAM perspective, movement of female ungulates
during the calving season likely contains information on maternal
status. Restlessness and the seeking of solitude characterize
imminent parturition [31], [32], [33] and establish a general
pattern of increased daily movement pre-partum and decreased
movement post-partum relative to barren females (Figure 2a; [34], g(y)~az
X
I
i~1
fi(Xi)ze where g is the link function and e is a random error term. Functions fi may be linear or nonparametric functions defined by
smoothers
such
as
smoothing
splines
or
locally
estimated
scatterplot smoothers (loess; also referred to as locally estimated,
or weighted, polynomial regression). Smoothers provide a series of
data summaries of the response that are specific to regions of the
covariates; a well known smoother is the moving average [29]. The
amount of smoothing is calibrated by the size of the neighborhood,
or percentage of the data points, over which averaging is done; a
quantity known as span. A larger span yields a smoother data
summary (less curvature) whereas a smaller span yields a less
smooth data summary (more curvature). This data summary (i.e., where g is the link function and e is a random error term. Functions fi may be linear or nonparametric functions defined by
smoothers
such
as
smoothing
splines
or
locally
estimated
scatterplot smoothers (loess; also referred to as locally estimated,
or weighted, polynomial regression). Smoothers provide a series of
data summaries of the response that are specific to regions of the
covariates; a well known smoother is the moving average [29]. The
amount of smoothing is calibrated by the size of the neighborhood,
or percentage of the data points, over which averaging is done; a
quantity known as span. A larger span yields a smoother data
summary (less curvature) whereas a smaller span yields a less
smooth data summary (more curvature). PLoS ONE | www.plosone.org Grouping Elk Relative to Maternal Status and Human
Activity We hypothesized that during calving time, movement
associated with restlessness and solitude seeking in parturient elk
provides a general pattern of complexity that is not observed in
non-parturient females, and we predicted that smoothers associ-
ated with movement data on parturient elk consistently would be
comprised of more parameters and thus more df than smoothers
associated with non-parturient elk. response curves depicting daily changes in the distance moved by
elk between successive locations. We used PROC GAM in SASH
(SAS Institute, Inc., Cary, North Carolina, USA), specifying
automated calculation of df using the generalized cross validation
method and loess smoothing, to assign df to response curves
depicting calving season movement patterns in each elk. We
established the following prediction: movement described by ,3 df
in GAMs would depict non-parturiency whereas movement
described by $3 df would depict parturiency. This prediction
was based on two observations. First, the shape of the response
curve depicting relatively simple movement, as in non-parturient
individuals, appears to correspond with #2 regions that differ in We used GAMs to regress distance traveled within a 24 h
period (using locations recorded at 1200 h on consecutive days)
against date. Date encompassed the 15 May – 1 July calving
season [33] in each of the 4 years comprising the study period. We used the generalized cross validation option to assign df to PLoS ONE | www.plosone.org January 2011 | Volume 6 | Issue 1 | e14597 4 Ungulate Conservation slope, each of which being calibrated over a span of $50% of the
data (Figure 2a; [34], [35]). Second, by plotting df as a function of
span, our data show that a span of 50% corresponds to ,3 df
(Figure 2b). Random intercepts can account for unbalanced data, correlation
among observations, and provide improved model fit and
parameter estimation [39] (but see [11]). Random-effects (or
mixed-effects) RSF models provide information on individual
behavior, how individuals contribute to population-level observa-
tions, and how their responses to a resource may change as a
function of its availability – a process known as a functional
response [40], [41]. Analytical approaches to model functional
responses in resource selection are particularly important when
there is a trade-off in selection for a particular resource [4], [40],
which would be the case if human activity is perceived as a risk of
harm [7]. Grouping Elk Relative to Maternal Status and Human
Activity We tested the GAM approach using blood samples obtained
from elk during capture in 2008 and 2009, and field observation of
females and calves in those same years. Blood sera were tested for
presence of pregnancy-specific protein-B (PSPB; BioTracking,
LLC, Moscow, Idaho, USA; [37]). We conducted field observation
from dawn to 0900 and from 1800 until dusk using binoculars and
spotting scopes to watch for behavior that suggested a maternal
bond between a female and calf. This behavior included nursing
and licking bouts, traveling as a female/calf unit, and heightened
attentiveness between a female and calf [13], [38]. Females for
which PSPB testing indicated pregnancy and for which field
observation suggested a strong female-calf bond were designated
as parturient. Determining non-parturiency is never definitive;
however, we designated females as non-parturient with negative
PSPB results and for which field observation was unable to
associate the female with a calf [13]. We thus established two
groups of elk relative to maternal status by which we analyzed
resource selection. We incorporated random effects into the use versus availability
design [9] in which covariates representing important resources
are compared at used and available (but presumed non-used)
locations using ^w(x)~exp(^b0z^b1x1z^b2x2z:::z^bnxn) where ^w(x) is the relative probability of use as a function of
covariates xn with coefficients ^bn estimated from logistic regression. Availability was defined for each elk by including random
locations within 100% minimum convex polygon seasonal use
area estimates; the number of random locations generated was 3
times the number of used locations for each elk. We examined
resource selection within seasonal use areas (i.e., 3rd order
selection). We modeled resource selection separately during day
and night because we expected behavior of elk to differ between
day and night. We assigned time of day at random to available
locations for day versus night comparisons; times assigned to
available locations corresponded to times associated with used
locations (e.g., every 3 hours on the sampled hour). Day models
included the times 0900, 1200, 1500, and 1800 h whereas night
models included 0000 and 0300 h. Using a Geographic Informa-
tion System (GIS; ArcGIS 9.2), we calculated 7 covariates at used
and random locations (Table 1). Four of these covariates including
cover type, slope, elevation, and habitat edge density were
calculated at locations both within and outside of the gas field. Grouping Elk Relative to Maternal Status and Human
Activity Three covariates including road density, distance to a human-built
structure, and industrial development footprint (Table 1) were
calculated only within the gas field because human activity in areas
adjacent but outside of the gas field was less intense and not
associated with industrial development. Raster data for cover type
were developed from annual aerial photography of the study area,
terrain covariates were calculated from a 30-m resolution digital
elevation model, and human activity covariates including roads,
structures, and industrial development footprint were heads-up-
digitized from aerial photography and analyzed as year-specific
covariates (Table 1). We used Spatial Analyst in ArcGIS to extract
values from raster data for all covariates. where ^w(x) is the relative probability of use as a function of
covariates xn with coefficients ^bn estimated from logistic regression. Similarly, we established two groups of elk relative to human
activity: elk occupying developed areas versus elk occupying
undeveloped areas. Elk locations occurring within 1 km of a gas
well were considered to be in the developed area (Figure 1). Human activity was apparent in the areas we called undeveloped
including some ranching and residences; however, no industrial
development occurred in undeveloped areas and human activity
was limited relative to developed areas. Given the short temporal
window within which we conducted these analyses, calving season
use areas (100% minimum convex polygons) generally occurred
wholly within developed areas or wholly within undeveloped
areas. Only a small number of elk occupied both areas during the
calving season – we discuss these elk separately. We estimated
minimum convex polygons because the temporal window of the
study was relatively short, calving season use areas comprised a
portion of the annual use areas, and we wanted to err on the side
of inclusiveness rather than potentially omitting a portion of
critical calving range from the analysis. In comparing resource
selection between elk occupying developed versus undeveloped
areas, a key assumption is that human activity associated with
energy development is the primary difference between areas and
that other factors to which elk respond were similar between areas. We compared landscape and habitat covariate values between
areas to inform this assumption. January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org Random-effects Resource Selection Modeling Modeling
variables
as
random
effects
can
improve
our
understanding of resource selection, which will enhance the
practical application of RSFs in management decisions [4], [39]. Random effects models assume that sample units are drawn at
random from a larger population and that the data are structured
hierarchically (i.e., within subject responses are more similar than
between-subject responses). Mechanistically, assumptions of ran-
dom-effects models are: (1) the random effects are distributed
normally with mean equal to 0 and unknown variance, (2) within-
group correlation is constant through time, and (3) the analyst has
correctly specified the variance-covariance structure (see [39] for a
review of the application of random-effects models in resource
selection analysis). In RSF models, it is appropriate to model
intercepts and/or covariates as random effects when variance
among sample units is of interest, animal response to gradients in a
resource is suspected, or population-level inference is of interest. We estimated a three-level random-effects model in which
locations i = 1…I occurred within strata representing individual
elk j = 1…J [39]. Considering a random intercept and random
coefficients, the RSF is estimated by g(x)~b0zb1x1ijz:::zbkxkijzckjxkjzc0j g(x)~b0zb1x1ijz:::zbkxkijzckjxkjzc0j where covariates k (k = 1…K) have values x, c0j is the random
intercept and ckj is the random coefficient of xk for elk j, which is
the difference in the intercept and coefficient for elk j from the
mean population-level intercept b0 and coefficient bkj, respectively. We estimated models using the GLIMMIX procedure in SASH. We specified the conditional probability distribution of the data as where covariates k (k = 1…K) have values x, c0j is the random
intercept and ckj is the random coefficient of xk for elk j, which is where covariates k (k = 1…K) have values x, c0j is the random
intercept and ckj is the random coefficient of xk for elk j, which is
the difference in the intercept and coefficient for elk j from the
mean population-level intercept b0 and coefficient bkj, respectively. We estimated models using the GLIMMIX procedure in SASH. We specified the conditional probability distribution of the data as PLoS ONE | www.plosone.or January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org PLoS ONE | www.plosone.org 5 Ungulate Conservation Table 1. Covariates used in random-effects resource selection models and their descriptions. Covariate
Description
Slope
Digital elevation model (DEM) provided at a resolution of 1.52 m and re-sampled for covariate calculation to a resolution of
30 m and measured in degrees. Values for slope calculated at point locations in our study ranged from 0.02 (flat ground) to 41.8
degrees (very steep). Elevation
DEM re-sampled for covariate calculation to a resolution of 30 m and measured in m. Elevation at sample points in our study
ranged from 1918.7–2970.0 m. Cover type
Raster dataset compiled from 1-foot resolution aerial imagery of the study area using Image AnalysisTM and re-sampled to 30 m
resolution for covariate calculation. From an elk-centric perspective, habitat in the Raton basin functioned in one of two ways:
as security cover or as forage resources. Raster cells were assigned one of two values (binary covariate) representing habitat
that functioned to provide cover versus habitat that did not function to provide cover. All tree or oak-thicket dominated
habitats were considered security cover, whereas all shrub and grassland dominated habitats were considered non-cover. Edge density
Density of line features depicting the interface of cover and non-cover habitat calculated for the central grid cell within a
990 m2 moving window. Values in our study were 22.2–136.0 km/km2. Mapping Responses We applied the results from random-effects models to map
relative probability of use across the study area which encom-
passed developed and undeveloped areas. We used marginal
coefficients from logistic regression to derive an RSF at a
resolution of 30 m using w(x)~exp
X
K
k~1
b
kxk
! w(x)~exp
X
K
k~1
b
kxk
! where covariates k (k = 1…K) have values x [9]. We were interested
in differences in resource selection patterns depending on maternal
status and between elk that occupied developed versus undevel-
oped areas. It is important to note that, although we analyzed an
independent group of field-observed parturient elk to facilitate
comparison of marginal estimates, these elk were withheld from
final RSF development as a validation sample. Thus, we examined
differences in how the relative probability of use was assigned
throughout the landscape depending on whether this probability
was based on models of: 1) all sample elk inside the gas field, 2) elk
predicted to be parturient based on GAMs inside the gas field, and
3) all sample elk outside of the gas field (groups 1,2, and 5 above). We conducted this examination using the following methods
separately for day and night. We developed annual RSF maps for
each of the 3 groups identified above. We estimated quantiles in
SASH
(PROC
RANK,
PROC
MEANS)
by
which
pixels
comprising the raster surface were partitioned into 5 equal-sized
subsets based on pixel value. In GIS we reclassified RSF values
based on quantiles establishing 5 ranks of the relative probability
of use (1 = low probability, 5 = high probability). We summed
within-year maps across all years and ranked relative probability of
use as described above yielding 6 multi-year predictive maps (day
map and night map for each group) with relative probability of use
ranging from 1 (low) to 5 (high). We validated predictive maps
using locations from 24 elk that were observed in the field to be ^bbk~ 1
J
X
J
j~1
^bkj where ^bkj was the estimate of coefficient k for elk j [43], [44], and
we estimated variance using where ^bkj was the estimate of coefficient k for elk j [43], [44], and
we estimated variance using var ^bk
~
1
J{1
X
J
j~1
(^bkj{^bbk)2 Population-level coefficient estimates (^bbk) are similar to the
average of the conditional estimates because the conditional
coefficient estimates are constrained to have means ^bk [44]. g(x)~b0zb1x1ijz:::zbkxkijzckjxkjzc0j Density of line features depicting the interface of cover and non-cover habitat calculated for the central grid cell within a
990 m2 moving window. Values in our study were 22.2–136.0 km/km2. Density of line features depicting roads calculated for grid cells at the center of a 990 m2 moving window. Road density was
calculated as a year-specific variable – that is, new road features were added to the data set as annual aerial imagery became
available. Values in our study, calculated within the gas field only, were 0.0–7.6 km/km2. Linear distance from a sample location to a human-built structure including houses, agricultural facilities, and industrial
facilities. Distance to structure was a year-specific variable and was analyzed as natural log transformed distance +0.1 to allow
its magnitude to decrease with increasing distance. Values in our study, calculated within the gas field only, were 0.0–5854.9 m. Area (density) of physically modified ground calculated from aerial photography as a year-specific variable. Physically modified
ground primarily reflected industrial development including well pads, pipe yards, pipelines, construction areas, or clearing for
facilities development. Industrial development footprint was calculated for the central grid cell within a 990 m2 moving
window. This covariate excludes physical disturbance associated with roads. Values in our study, calculated only within the gas
field, were 0.0–0.62 km2/km2. Industrial development footprint
Area (density) of physically modified ground calculated from aerial photography as a year-specific variable. Physically modified
ground primarily reflected industrial development including well pads, pipe yards, pipelines, construction areas, or clearing for
facilities development. Industrial development footprint was calculated for the central grid cell within a 990 m2 moving
window. This covariate excludes physical disturbance associated with roads. Values in our study, calculated only within the gas
field, were 0.0–0.62 km2/km2. doi:10.1371/journal.pone.0014597.t001 binomial and used a logit link function. We included ordinal date
as a class variable and specified a variance components covariance
structure in which random intercepts for ordinal date were nested
within each individual to address within-day autocorrelation
among locations. R-side, or marginal, random-effects models
estimated
using
GLIMMIX
provide
conditional
coefficient
estimates for each individual, that is, estimates for individual
animals that are conditional on the distribution of coefficient
estimates across all individuals in the population. g(x)~b0zb1x1ijz:::zbkxkijzckjxkjzc0j To estimate
marginal (i.e., population-level) coefficients we assumed that
conditional coefficients for each elk represented a random sample
from a normal distribution with the mean of that distribution
representing the population-level effects of covariates on the
probability of use [42]. We estimated marginal coefficients using g(x)~b0zb1x1ijz:::zbkxkijzckjxkjzc0j Road density
Density of line features depicting roads calculated for grid cells at the center of a 990 m2 moving window. Road density was
calculated as a year-specific variable – that is, new road features were added to the data set as annual aerial imagery became
available. Values in our study, calculated within the gas field only, were 0.0–7.6 km/km2. Distance to structure
Linear distance from a sample location to a human-built structure including houses, agricultural facilities, and industrial
facilities. Distance to structure was a year-specific variable and was analyzed as natural log transformed distance +0.1 to allow
its magnitude to decrease with increasing distance. Values in our study, calculated within the gas field only, were 0.0–5854.9 m. Industrial development footprint
Area (density) of physically modified ground calculated from aerial photography as a year-specific variable. Physically modified
ground primarily reflected industrial development including well pads, pipe yards, pipelines, construction areas, or clearing for
facilities development. Industrial development footprint was calculated for the central grid cell within a 990 m2 moving
window. This covariate excludes physical disturbance associated with roads. Values in our study, calculated only within the gas
field, were 0.0–0.62 km2/km2. doi:10.1371/journal.pone.0014597.t001 Table 1. Covariates used in random-effects resource selection models and their descriptions. Table 1. Covariates used in random-effects resource selection models and their descriptions. Digital elevation model (DEM) provided at a resolution of 1.52 m and re-sampled for covariate calculation to a resolution of
30 m and measured in degrees. Values for slope calculated at point locations in our study ranged from 0.02 (flat ground) to 41.8
degrees (very steep). DEM re-sampled for covariate calculation to a resolution of 30 m and measured in m. Elevation at sample points in our study
ranged from 1918.7–2970.0 m. Raster dataset compiled from 1-foot resolution aerial imagery of the study area using Image AnalysisTM and re-sampled to 30 m
resolution for covariate calculation. From an elk-centric perspective, habitat in the Raton basin functioned in one of two ways:
as security cover or as forage resources. Raster cells were assigned one of two values (binary covariate) representing habitat
that functioned to provide cover versus habitat that did not function to provide cover. All tree or oak-thicket dominated
habitats were considered security cover, whereas all shrub and grassland dominated habitats were considered non-cover. Status #Elk
#Locations
Year
Groupa
Day Night
Day
Night
2006
Inside – all sample elk
11
10
1,759
879
Inside – GAM-predicted parturient
5
5
917
458
Outside – all sample elk
7
7
122
61
2007
Inside – all sample elk
26
24
3,281
1,641
Inside – GAM-predicted parturient
9
9
1,174
587
Outside – all sample elk
16
14
900
451
2008
Inside – all sample elk
19
19
2,056
1,455
Inside – GAM-predicted parturient
7
7
776
438
Inside – field-observed parturient
12
12
1,598
802
Inside – field-observed non-parturient 5
5
425
227
Outside – all sample elk
19
16
417
230
2009
Inside – all sample elk
15
15
2,735
1,482
Inside – GAM-predicted parturient
8
8
1,381
757
Inside – field-observed parturient
12
12
1,976
1,062
Inside – field-observed non-parturient 5
5
685
496
Outside – all sample elk
16
14
411
215 Random-effects Resource Selection Modeling – Marginal
Inference Mapping Responses In all,
five groups of elk were available for comparison: 1) all sample elk
inside of the gas field, 2) elk predicted to be parturient based on
GAMs inside the gas field, 3) field-observed parturient elk inside of
the gas field, 4) field-observed non-parturient elk inside of the gas
field, and 5) all sample elk outside of the gas field (sample size was
too low to develop maternal groups; see Results). January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org 6 Ungulate Conservation parturient (see below); these elk were withheld from final RSF
development so they represent an independent validation sample. Locations from these elk were plotted on multi-year predictive
maps. We tested whether the number of locations that occurred
within each predicted probability of use rank (1–5) differed from
expectation using a chi-square test for specified proportions
(PROC FREQ, SASH). To provide a measure of the amount to
which a map that validated well differed from a map that validated
poorly, we calculated the number of pixels comprising the raster
surface that differed between maps in terms of relative probability
rank. were minimal. Based on 25,290 GIS-generated random sample
points (12,645 inside and 12,645 outside of the gas field,
respectively), x 6 SD values inside versus outside of the gas field
respectively were 2,347.7 6 146.4 m and 2,382.6 6 243.4 m for
elevation, 10.6 6 6.2 degrees and 9.4 6 7.0 degrees for slope, and
82.0 6 10.2 km/km2 and 77.5 6 12.7 km/km2 for edge density. The proportion of sample points that occurred within security
habitat was 0.84 and 0.80 inside versus outside of the gas field,
respectively. Topographic covariates (elevation and slope) were
not ground truthed; however, vegetation attributes were ground-
truthed based on field-established polygons (n = 1177) of known
vegetation type. Status #Elk
#Locations
Year
Groupa
Day Night
Day
Night
2006
Inside – all sample elk
11
10
1,759
879
Inside – GAM-predicted parturient
5
5
917
458
Outside – all sample elk
7
7
122
61
2007
Inside – all sample elk
26
24
3,281
1,641
Inside – GAM-predicted parturient
9
9
1,174
587
Outside – all sample elk
16
14
900
451
2008
Inside – all sample elk
19
19
2,056
1,455
Inside – GAM-predicted parturient
7
7
776
438
Inside – field-observed parturient
12
12
1,598
802
Inside – field-observed non-parturient 5
5
425
227
Outside – all sample elk
19
16
417
230
2009
Inside – all sample elk
15
15
2,735
1,482
Inside – GAM-predicted parturient
8
8
1,381
757
Inside – field-observed parturient
12
12
1,976
1,062
Inside – field-observed non-parturient 5
5
685
496
Outside – all sample elk
16
14
411
215
aInside refers to areas within the gas field and outside refers to areas adjacent
but external to the gas field. GAM-predicted parturient refers to female elk that
we predicted to be parturient based on results of generalized additive
modeling (GAM) of movement data. Field-observed refers to elk for which we
observed behavior in the field to assess parturition status. Total sample fitted with Global Positioning Systems collars was 25, 40, 50, and
50 in 2006, 2007, 2008, and 2009, respectively. Within-year deviation from the
available sample arises from 3 sources: 1) some collared elk moved out of the
study area during calving, 2) the GAM-predicted sample was nested within the
total sample, and 3) some individuals occupied areas both inside and outside of
the gas field. doi:10.1371/journal.pone.0014597.t002
PLoS ONE | www.plosone.org Table 2. Resource selection by Rocky Mountain elk during
calving time; sample size and number of locations used in
analyses. Table 2. Resource selection by Rocky Mountain elk during
calving time; sample size and number of locations used in
analyses. Results Generalized Additive Modeling to Designate Maternal
Status aInside refers to areas within the gas field and outside refers to areas adjacent
but external to the gas field. GAM-predicted parturient refers to female elk that
we predicted to be parturient based on results of generalized additive
modeling (GAM) of movement data. Field-observed refers to elk for which we
observed behavior in the field to assess parturition status.
Total sample fitted with Global Positioning Systems collars was 25, 40, 50, and
50 in 2006, 2007, 2008, and 2009, respectively. Within-year deviation from the
available sample arises from 3 sources: 1) some collared elk moved out of the
study area during calving, 2) the GAM-predicted sample was nested within the
total sample, and 3) some individuals occupied areas both inside and outside of
the gas field. Random-effects Resource Selection Modeling – Marginal
Inference We found little evidence of lack of fit or overdispersion in RSF
models; generalized chi-square/degrees of freedom was 0.79–0.92
for night time models and 0.87–0.97 for day time models. The
most notable response by elk inside of the gas field was strong
avoidance of the industrial development footprint during the day
among all groups (at the population-level; Figure 3a). Other
notable behavior in field-observed parturient elk included strong
selection for cover and avoidance of high road density during the
day relative to other groups (Figure 3b,c). While field-observed
parturient elk showed some day-time preference for flatter areas
and lower elevation relative to the larger sample, differences
among maternal groups in selection for these and other resources
were small, inconsistent, or not apparent. Table 3. Parameterizing movement using generalized
additive models. Year/Group
df,2a
2,df,3
df$3
2008 Field-observed parturient
2
1
9
2009 Field-observed parturient
5
3
4
2008 Field-observed non-parturient
2
2
1
2009 Field-observed non-parturient
3
2
0
aAs part of generalized additive modeling we specified the generalized cross
validation option to assign parameters (df) to polynomials depicting daily
changes in the distance moved by elk between successive locations. Maternal status was assigned to 34 female Rocky Mountain elk (10 non-
parturient and 24 parturient) using information from blood samples
(pregnancy-specific protein-B) and field observation (788 h). We assessed how
generalized additive models of location data parameterized movement, and
how parameterization corresponded with calf status as determined in the field. Results show that among 14 elk for which movement was parameterized by $3
df, 13 were observed in the field to be with calf. doi:10.1371/journal.pone.0014597.t003 Table 3. Parameterizing movement using generalized
additive models. aAs part of generalized additive modeling we specified the generalized cross
validation option to assign parameters (df) to polynomials depicting daily
changes in the distance moved by elk between successive locations. Maternal status was assigned to 34 female Rocky Mountain elk (10 non-
parturient and 24 parturient) using information from blood samples
(pregnancy-specific protein-B) and field observation (788 h). We assessed how
generalized additive models of location data parameterized movement, and
how parameterization corresponded with calf status as determined in the field. Results show that among 14 elk for which movement was parameterized by $3
df, 13 were observed in the field to be with calf. January 2011 | Volume 6 | Issue 1 | e14597 Status We applied results from PSPB and field observation of female
elk in 2008 and 2009 to test the GAM approach, in which
movement patterns parameterized by ,3df were predicted to
depict non-parturiency whereas movement patterns parameterized
by $3 df were predicted to depict parturiency. Based on PSPB
results and 788 hours of field observation, we assigned maternal
status to 34 elk; 24 were identified as parturient and 10 as non-
parturient. Movement during calving time was highly variable,
especially among parturient elk, with parameterization ranging
from 1.24–16.9 df. However, among elk observed to be parturient
in the field, 93% (13/14) were parameterized by $3 df (Table 3). We fitted 25 female elk with GPS collars in 2006, 40 in 2007, 50
in 2008, and 50 in 2009. The entire within-year sample was
unavailable for these analyses because the analyses spanned a
relatively short period during calving (Table 2), and in each year
several elk moved from the study area to alpine habitat for
parturition. Inadequate sample size (i.e., #2 elk comprising
maternal
groups)
in
undeveloped
areas
restricted
a
more
comprehensive assessment. Differences in elevation, slope, edge
density, and cover type between developed and undeveloped areas parturition. Inadequate sample size (i.e., #2 elk comprising
maternal
groups)
in
undeveloped
areas
restricted
a
more
comprehensive assessment. Differences in elevation, slope, edge
density, and cover type between developed and undeveloped areas
Table 2. Resource selection by Rocky Mountain elk during
calving time; sample size and number of locations used in
analyses. Random-effects Resource Selection Modeling – Marginal
Inference doi:10.1371/journal.pone.0014597.t003 Total sample fitted with Global Positioning Systems collars was 25, 40, 50, and
50 in 2006, 2007, 2008, and 2009, respectively. Within-year deviation from the
available sample arises from 3 sources: 1) some collared elk moved out of the
study area during calving, 2) the GAM-predicted sample was nested within the
total sample, and 3) some individuals occupied areas both inside and outside of
the gas field. g
doi:10.1371/journal.pone.0014597.t002 January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org 7 Ungulate Conservation Figure 3. Group-dependent population-level responses: inside of the gas field. Marginal coefficient estimates 6 95% CL of selection for
human disturbance (a), security cover (b), and road density (c) by female elk during 2006-2009 in Raton Basin, Colorado, USA. Day time results are
displayed among maternal statuses for elk inside of the gas field. doi:10.1371/journal.pone.0014597.g003 Figure 3. Group-dependent population-level responses: inside of the gas field. Marginal coefficient estimates 6 95% CL of s
human disturbance (a), security cover (b), and road density (c) by female elk during 2006-2009 in Raton Basin, Colorado, USA. Day time
displayed among maternal statuses for elk inside of the gas field. doi:10.1371/journal.pone.0014597.g003 Figure 3. Group-dependent population-level responses: inside of the gas field. Marginal coefficient estimates 6 95% CL of selection for
human disturbance (a), security cover (b), and road density (c) by female elk during 2006-2009 in Raton Basin, Colorado, USA. Day time results are
displayed among maternal statuses for elk inside of the gas field. doi:10.1371/journal.pone.0014597.g003 We estimated selection for 4 landscape/habitat covariates in
both developed and undeveloped areas including security cover,
slope, elevation, and habitat edge density. Considering population-
level coefficient estimates, selection for resources that functioned to
provide security cover differed markedly between elk that occupied
developed areas versus undeveloped areas (Figure 4). Elk inside of
the gas field consistently showed strong selection for security cover
during the day (Figure 4a) and strong selection for non-cover or
forage habitats during the night (Figure 4b) throughout the study
period whereas elk occupying areas outside of the gas field
generally selected randomly for security cover with coefficient
estimates near zero. Another difference between elk inside versus
outside of the gas field involved selection for elevation. Elk outside
of the gas field generally selected randomly for elevation with coefficient estimates always near zero. Random-effects Resource Selection Modeling – Marginal
Inference Elk inside of the gas field
generally selected for higher elevation during the day (Figure 4c;
however, 95% CL overlapped zero), and for lower elevation at
night (Figure 4d). Diurnal patterns in selection for slope and edge
density were apparent but differences in selection for these features
between elk inside versus outside of the gas field generally were
small (Table 4). PLoS ONE | www.plosone.org Random-effects Resource Selection Modeling –
Conditional Inference Conditional estimates revealed a complex association between
human activity and individual variation in response to environmental
and anthropogenic features (Figure 5). Elk inside the gas field showed
greater heterogeneity among individuals relative to elk outside of the January 2011 | Volume 6 | Issue 1 | e14597 8 Figure 4. Population-level responses: inside versus outside of the gas field. Marginal coefficient estimates 695% CL of day-time (a) and
night-time (b) selection for security cover, and day-time (c) and night-time (d) selection for elevation by female elk during 2006-2009 in Raton Basin,
Colorado, USA. Inside refers to elk occupying an active gas field whereas outside refers to elk occupying adjacent undeveloped areas; maternal status
is not considered (i.e., all sample elk are grouped depending on whether they occurred inside versus outside of the gas field). doi:10.1371/journal.pone.0014597.g004
Ungulate Conservation Ungulate Conservation gas field in their responses to edge density and elevation. Selection for
l
(Fi
5 b)
d d
d
it (Fi
5
d)
ti
t d
Seasonal use areas of 15 e
d
l
d
I di id
l
Figure 4. Population-level responses: inside versus outside of the gas field. Marginal coefficient e
night-time (b) selection for security cover, and day-time (c) and night-time (d) selection for elevation by fema
Colorado, USA. Inside refers to elk occupying an active gas field whereas outside refers to elk occupying adjace
is not considered (i.e., all sample elk are grouped depending on whether they occurred inside versus outsid
doi:10.1371/journal.pone.0014597.g004 Figure 4. Population-level responses: inside versus outside of the gas field. Marginal coefficient estimates 695% CL of day-time (a) and
night-time (b) selection for security cover, and day-time (c) and night-time (d) selection for elevation by female elk during 2006-2009 in Raton Basin,
Colorado, USA. Inside refers to elk occupying an active gas field whereas outside refers to elk occupying adjacent undeveloped areas; maternal status
is not considered (i.e., all sample elk are grouped depending on whether they occurred inside versus outside of the gas field). doi:10.1371/journal.pone.0014597.g004 Seasonal use areas of 15 elk overlapped both developed and
undeveloped areas. Individuals showed notable behavioral differ-
ences, including diurnal variation, relative to selection for security
cover and elevation depending on whether they were in developed
versus undeveloped areas (Figure 6). January 2011 | Volume 6 | Issue 1 | e14597 Random-effects Resource Selection Modeling –
Conditional Inference During the day, 13 of 15 elk
showed stronger selection for security cover when inside the gas
field, but at night this pattern broke down with no apparent
consistent behavioral response. Selection for elevation was variable
inside the gas field during day and night with some elk showing
relatively strong selection for higher or lower elevations. When elk
were outside of the gas field they generally used elevation at
random (Figure 6). gas field in their responses to edge density and elevation. Selection for
slope (Figure 5a,b) and edge density (Figure 5c, d) was estimated more
precisely among elk inside of the gas field; however, elk outside the gas
field selected (randomly) for elevation with high precision (Figure 5e,
f). Also apparent inside the gas field but not outside was a temporal
trend of increasing avoidance of high edge density (Figure 5c). Within-year marginal estimates of day time selection for road density
were positive (i.e., elk selected for higher road density) in 3 of 4 years. However, conditional estimates revealed that, across all years, only
37% of sampled elk selected for relatively high road density; 32%
selected neither for nor against high road density, and 31% avoided
high road density (Figure 5g). gas field in their responses to edge density and elevation. Selection for
slope (Figure 5a,b) and edge density (Figure 5c, d) was estimated more
precisely among elk inside of the gas field; however, elk outside the gas
field selected (randomly) for elevation with high precision (Figure 5e,
f). Also apparent inside the gas field but not outside was a temporal
trend of increasing avoidance of high edge density (Figure 5c). Within-year marginal estimates of day time selection for road density
were positive (i.e., elk selected for higher road density) in 3 of 4 years. However, conditional estimates revealed that, across all years, only
37% of sampled elk selected for relatively high road density; 32%
selected neither for nor against high road density, and 31% avoided
high road density (Figure 5g). PLoS ONE | www.plosone.org January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org 9 Ungulate Conservation Table 4. Marginal coefficient estimates (^bbn[SE]) for two-level random-effects models of resource selection by Rocky Mountain elk
during calving time (15 May - 1 July), 2006–2009. Random-effects Resource Selection Modeling –
Conditional Inference Day
Night
Effect/Year
Inside
Outside
Inside
Outside
Slope 2006
20.001 (0.002)
20.003 (0.034)
20.064 (0.013)
-
Slope 2007
0.011 (0.006)
0.005 (0.013)
20.030 (0.010)
20.040 (0.016)
Slope 2008
20.003 (0.007)
0.003 (0.003)
20.035 (0.007)
20.045 (0.008)
Slope 2009
0.012 (0.005)
0.024 (0.014)
20.023 (0.011)
20.027 (0.013)
Edge Density 2006
0.015 (0.016)
20.022 (0.004)
20.007 (0.021)
-
Edge Density 2007
0.001 (0.013)
20.003 (0.012)
0.006 (0.011)
20.013 (0.004)
Edge Density 2008
20.026 (0.009)
20.008 (0.003)
20.026 (0.008)
20.010 (0.002)
Edge Density 2009
20.027 (0.009)
20.006 (0.010)
20.036 (0.012)
20.044 (0.021)
- Model failed to converge. doi:10.1371/journal.pone.0014597.t004 Table 4. Marginal coefficient estimates (^bbn[SE]) for two-level random-effects models of resource selection by Rocky Mountain elk
during calving time (15 May - 1 July), 2006–2009. Table 4. Marginal coefficient estimates (^bbn[SE]) for two-level random-effects models of resource selection by Rocky Mountain elk
during calving time (15 May - 1 July), 2006–2009. Table 4. Marginal coefficient estimates (^bbn[SE]) for two-level random-effects models of resource selection by Rocky Mountain elk
during calving time (15 May - 1 July), 2006–2009. Figure 5. Population-level and individual responses. Marginal (dash) and conditional (circles) coefficient estimates (SE) of day-time selection
for slope (a, b) edge density (c, d), elevation (e, f), and road density (g) by female elk during 2006-2009 in Raton basin, Colorado, USA. Results are
displayed by year for elk that occupied developed (panels a, c, e, and g) and undeveloped (panels b, d, and f) areas. Orange depicts elk that were
predicted to be parturient using generalized additive modeling; open circles (#) indicate conditional estimates for field-observed parturient elk in
2008 and 2009. doi:10.1371/journal.pone.0014597.g005 Figure 5. Population-level and individual responses. Marginal (dash) and conditional (circles) coefficient estimates (SE) of day-time selection
for slope (a, b) edge density (c, d), elevation (e, f), and road density (g) by female elk during 2006-2009 in Raton basin, Colorado, USA. Results are
displayed by year for elk that occupied developed (panels a, c, e, and g) and undeveloped (panels b, d, and f) areas. Orange depicts elk that were
predicted to be parturient using generalized additive modeling; open circles (#) indicate conditional estimates for field-observed parturient elk in
2008 and 2009. doi:10 1371/journal pone 0014597 g005 Figure 5. Population-level and individual responses. January 2011 | Volume 6 | Issue 1 | e14597 Random-effects Resource Selection Modeling –
Conditional Inference Symbols represent conditional coefficient estimates (y-axes) for each elk in 2007 (squares; n = 6), 2008 (circles; n = 4),
and 2009 (triangles; n = 5); lines show how selection changed within each elk depending on whether it was inside versus outside of the gas field. Solid
lines depict larger selection coefficients inside the gas field whereas dashed lines depict larger coefficients outside of the gas field. Coefficients are
informed by 36-293 location per elk inside the gas field, and 20-248 locations per elk outside the gas field. doi:10.1371/journal.pone.0014597.g006 Functional Responses resources near physical disturbance and, during the day, showed a
stronger functional response to the proportion of cover than did
elk in undeveloped areas (Figure 7b, c). Consistent with a functional response in resource selection, elk
selected randomly for disturbed areas when disturbance was
minimal. As human activity increased (i.e., across individual elk
calving season areas, or through time), elk showed stronger
avoidance of the industrial development footprint during the day
but not at night; this spatiotemporal pattern of avoidance revealed
that elk continued to use physically disturbed areas but modified
their behavior to avoid human activity (Figure 7a). One way elk
avoided human activity in the day was by modifying selection for
security cover. In developed areas elk were constrained to select Random-effects Resource Selection Modeling –
Conditional Inference Marginal (dash) and conditional (circles) coefficient estimates (SE) of day-time selection
for slope (a, b) edge density (c, d), elevation (e, f), and road density (g) by female elk during 2006-2009 in Raton basin, Colorado, USA. Results are
displayed by year for elk that occupied developed (panels a, c, e, and g) and undeveloped (panels b, d, and f) areas. Orange depicts elk that were
predicted to be parturient using generalized additive modeling; open circles (#) indicate conditional estimates for field-observed parturient elk in
2008 and 2009. doi:10 1371/journal pone 0014597 g005 doi:10.1371/journal.pone.0014597.g005 doi:10.1371/journal.pone.0014597.g005 January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org 10 Ungulate Conservation Figure 6. Movement from within the gas field to areas adjacent to the gas field. A subsample of female elk (n = 15) occupied areas within
the gas field as well as areas outside but adjacent to the gas field. This figure shows differences in day-time (a) and night-time (b) selection for habitat
that provides security cover, and day-time (c) and night-time (d) selection for elevation among these elk depending on whether they were inside
versus outside of the gas field. Symbols represent conditional coefficient estimates (y-axes) for each elk in 2007 (squares; n = 6), 2008 (circles; n = 4),
and 2009 (triangles; n = 5); lines show how selection changed within each elk depending on whether it was inside versus outside of the gas field. Solid
lines depict larger selection coefficients inside the gas field whereas dashed lines depict larger coefficients outside of the gas field. Coefficients are
informed by 36-293 location per elk inside the gas field, and 20-248 locations per elk outside the gas field. doi:10.1371/journal.pone.0014597.g006 Figure 6. Movement from within the gas field to areas adjacent to the gas field. A subsample of female elk (n = 15) occupied areas within
the gas field as well as areas outside but adjacent to the gas field. This figure shows differences in day-time (a) and night-time (b) selection for habitat
that provides security cover, and day-time (c) and night-time (d) selection for elevation among these elk depending on whether they were inside
versus outside of the gas field. PLoS ONE | www.plosone.org Mapping Responses Dashes (-) and dashed lines
symbolize night-time resource selection, and circles (N) and solid lines symbolize day-time resource selection. Dashes and circles outlined by open
circles (#) or boxes (%) indicate conditional estimates for field-observed parturient elk in 2008 and 2009. doi:10.1371/journal.pone.0014597.g007 Figure 7. Functional responses in resource selection by female elk. Selection for the human disturbance footprint (a), security cover inside of
the gas field (b), and security cover outside of the gas field (c) change as a function of availability. Availability was calculated at the seasonal use area
level and conditional coefficients were estimated from generalized linear random-effects resource selection models. Dashes (-) and dashed lines
symbolize night-time resource selection, and circles (N) and solid lines symbolize day-time resource selection. Dashes and circles outlined by open
circles (#) or boxes (%) indicate conditional estimates for field-observed parturient elk in 2008 and 2009. doi:10 1371/journal pone 0014597 g007 and 3505 day locations) and then summed day and night values
within the three groups of interest to estimate overall within-group
model performance. For both models based on elk behavior inside
of the gas field (Figure 8a), habitat with low predicted probability
of use was modeled accurately; however, validation elk used
habitat predicted to have high probability of use only as much as
would be expected if elk used all predictive classes equally
(Figure 8c). The model based on elk behavior outside of the gas
field (Figure 8b) validated well with few locations occurring in
lowest ranked areas and many locations occurring in highest
ranked areas (Figure 8d). The proportion of pixels ranked
differently between the model that validated most poorly and
the model that validated best was 0.72–0.88 depending on year. Mapping Responses We developed six models (day and night separately for each of
three groups): the first was based on all sample elk that occurred
inside of the gas field; the second was a subset of the first group for
which GAMs predicted females to be parturient; and the third was
based on all sample elk that occurred outside of the gas field. We
validated day and night models separately (1830 night locations January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org 11 Figure 7. Functional responses in resource selection by female elk. Selection for the human disturbance footprint (a), security cover inside of
the gas field (b), and security cover outside of the gas field (c) change as a function of availability. Availability was calculated at the seasonal use area
level and conditional coefficients were estimated from generalized linear random-effects resource selection models. Dashes (-) and dashed lines
symbolize night-time resource selection, and circles (N) and solid lines symbolize day-time resource selection. Dashes and circles outlined by open
circles (#) or boxes (%) indicate conditional estimates for field-observed parturient elk in 2008 and 2009. doi:10.1371/journal.pone.0014597.g007
Ungulate Conservation Ungulate Conservation Figure 7. Functional responses in resource selection by female elk. Selection for the human disturbance
the gas field (b), and security cover outside of the gas field (c) change as a function of availability. Availability wa
level and conditional coefficients were estimated from generalized linear random-effects resource selection m
symbolize night-time resource selection, and circles (N) and solid lines symbolize day-time resource selection. circles (#) or boxes (%) indicate conditional estimates for field-observed parturient elk in 2008 and 2009. doi:10.1371/journal.pone.0014597.g007 Figure 7. Functional responses in resource selection by female elk. Selection for the human disturbance footprint (a), security cover inside of
the gas field (b), and security cover outside of the gas field (c) change as a function of availability. Availability was calculated at the seasonal use area Figure 7. Functional responses in resource selection by female elk. Selection for the human disturbance footprint (a), security cover inside of
the gas field (b), and security cover outside of the gas field (c) change as a function of availability. Availability was calculated at the seasonal use area
level and conditional coefficients were estimated from generalized linear random-effects resource selection models. Discussion and 3505 day locations) and then summed day and night values
within the three groups of interest to estimate overall within-group
model performance. For both models based on elk behavior inside
of the gas field (Figure 8a), habitat with low predicted probability
of use was modeled accurately; however, validation elk used
habitat predicted to have high probability of use only as much as
would be expected if elk used all predictive classes equally
(Figure 8c). The model based on elk behavior outside of the gas
field (Figure 8b) validated well with few locations occurring in
lowest ranked areas and many locations occurring in highest
ranked areas (Figure 8d). The proportion of pixels ranked
differently between the model that validated most poorly and
the model that validated best was 0.72–0.88 depending on year. We successfully integrated GAMs, field observation, and
random-effects RSF modeling to designate parturiency, describe
individual and population-level resource selection, and determine
the relative influence of maternal status and human activity on the
reliability of spatially explicit models intended to guide conserva-
tion of critical ungulate calving resources. We assumed that
planning tools such as spatially explicit models would be more
reliable if they account for adaptive behavior which, among
parturient elk, should be reflected in movement and resource
selection [45]. This was the basis for using GAMs to designate
parturiency based on movement; our thinking was that ungulate PLoS ONE | www.plosone.org PL January 2011 | Volume 6 | Issue 1 | e14597 PLoS ONE | www.plosone.org January 2011 | Volume 6 | Issue 1 | e14597 12 Ungulate Conservation Figure 8. Assigning the relative probability of use throughout the landscape. Comparison of how the relative predicted probability of use
was assigned throughout the landscape depending on whether RSF mapping was based on resource selection behavior among female elk that
occupied a natural gas field (a) or occupied areas adjacent to but outside of a natural gas field (b). Maps depict day-time probability of use. Wells are
depicted as black dots. Borders of wildlife areas managed by the state of Colorado are depicted in red. Probability of use is scaled from low to high
with each of 5 ordinal bins representing quantiles of the total number of pixels (30-m resolution) comprising the area. Discussion doi:10.1371/journal.pone.0014597.g008 behavior contains information on reproductive status and GAMs
would reveal a difference in the shape and relative complexity of
the response curve between parturient and non-parturient females. Maternal status was evident in elk movement with our application
of GAMs correctly predicting parturiency 93% of the time. We
note, however, the GAM approach as presented would require
further development if improved sensitivity (reduced false negative
rate)
is
desirable. GAMs
have
been
a
standard
tool
in
epidemiologic analyses and have found broad application in
ecology for modeling tolerance thresholds and spatial distributions
[46], [47], [48], [49]. Using movement data to designate
functional groups, seasonality, or behavior in animals has
increased in prevalence as GPS-based research has become more
frequent [50], [51]. Although we found some potentially important
differences in resource selection patterns between parturient and
non-parturient elk (see below), behavior in these groups was more behavior contains information on reproductive status and GAMs
would reveal a difference in the shape and relative complexity of
the response curve between parturient and non-parturient females. Maternal status was evident in elk movement with our application
of GAMs correctly predicting parturiency 93% of the time. We
note, however, the GAM approach as presented would require
further development if improved sensitivity (reduced false negative
rate)
is
desirable. GAMs
have
been
a
standard
tool
in
epidemiologic analyses and have found broad application in
ecology for modeling tolerance thresholds and spatial distributions
[46], [47], [48], [49]. Using movement data to designate
functional groups, seasonality, or behavior in animals has
increased in prevalence as GPS-based research has become more
frequent [50], [51]. Although we found some potentially important
differences in resource selection patterns between parturient and
non-parturient elk (see below), behavior in these groups was more similar than expected. Had maternal status been more apparent in
shaping resource selection, methods to designate status might have
found better application in landscape-level mapping of the relative
predicted probability of use. Here, both parturient and non-
parturient elk were examined inside the gas field where any
behavior distinctly associated with parturiency was obscured by
apparent risk-aversive behavior related to human activity during
the day time. At the population-level parturient elk, while avoiding roads and
selecting for security cover more strongly than other elk,
conformed to a general pattern of avoiding human activity during
the day by occupying upland forest or Q. Discussion Charts (c, d) display map
validation with columns depicting the number of locations from an independent sample of elk that, when plotted on RSF maps (a, b), occurred within
each ordinal bin. The black horizontal line depicts expectation if resources were selected at random. The map based on behavior inside of the gas
field validated poorly (c; x2 = 577.46, df = 4, p,0.001) relative to the map based on behavior outside of the gas field (d; x2 = 849.18, df = 4, p,0.001)
with the distribution of validation locations differing significantly from random in both cases. RSF maps (a, b) are based on all sample elk inside and
outside of the gas field, respectively; not shown is the RSF map based on elk predicted to be parturient using generalized additive models. doi:10.1371/journal.pone.0014597.g008 Figure 8. Assigning the relative probability of use throughout the landscape. Comparison of how the relative predicted probability of use
was assigned throughout the landscape depending on whether RSF mapping was based on resource selection behavior among female elk that
occupied a natural gas field (a) or occupied areas adjacent to but outside of a natural gas field (b). Maps depict day-time probability of use. Wells are
depicted as black dots. Borders of wildlife areas managed by the state of Colorado are depicted in red. Probability of use is scaled from low to high
with each of 5 ordinal bins representing quantiles of the total number of pixels (30-m resolution) comprising the area. Charts (c, d) display map
validation with columns depicting the number of locations from an independent sample of elk that, when plotted on RSF maps (a, b), occurred within
each ordinal bin. The black horizontal line depicts expectation if resources were selected at random. The map based on behavior inside of the gas
field validated poorly (c; x2 = 577.46, df = 4, p,0.001) relative to the map based on behavior outside of the gas field (d; x2 = 849.18, df = 4, p,0.001)
with the distribution of validation locations differing significantly from random in both cases. RSF maps (a, b) are based on all sample elk inside and
outside of the gas field, respectively; not shown is the RSF map based on elk predicted to be parturient using generalized additive models. Discussion gambelii thickets, and
selecting forage resources in valley bottoms at night regardless of
proximity to infrastructure. Trade-offs that structure resource
selection in many ungulates, most notably between forage
requirements and risk avoidance [6], [52], often are amplified in PLoS ONE | www.plosone.org PLoS ONE | www.plosone.org January 2011 | Volume 6 | Issue 1 | e14597 13 January 2011 | Volume 6 | Issue 1 | e14597 Ungulate Conservation parturient females through nutritional demands to support body
condition, lactation, and neonatal defense [53]. Here, responses to
human activity were more apparent than differences in resource
selection as a function of parturiency with elk exhibiting a clear
spatiotemporal avoidance of human activity at the population-
level and modified patterns of selection for environmental features,
notably security cover and elevation, in developed areas relative to
undeveloped areas. Research has shown that ungulates exhibit
avoidance behavior relative to human development, recreation,
hunting, and other activities [54]. Modeling resource selection
separately during day and night offered insight into whether the
physical presence of infrastructure versus the operation and
maintenance of such infrastructure (only occurring during the day)
was more important in avoidance behavior. Elk inside the gas field
used cover and elevation to modulate avoidance of human activity
during the day [55] whereas cover and elevation were less
influential outside of the gas field suggesting that human activity
during the day was the factor to which elk were responding [3]. At
night during calving time female elk showed no aversion to
infrastructure and selected for areas characterized by valley-
bottoms and foraging resources that in many instances were in
close proximity to, or directly associated with, disturbances such as
well pads and roads [56]. Day time refugia characterized by
minimal human disturbance, security cover, and upper slope
positions will be critical for maintenance of calving and perhaps
other seasonal habitats in multiple-use landscapes. drome, individuals have a behavioral type such as risk-aversion
(i.e., more risk-averse versus less risk-averse types; [58]). The
notion that individuals can be more or less risk aversive implies a
limit to their range of behavioral plasticity. From a conservation
perspective, animals exhibiting limited plasticity in environments
undergoing rapid change, such as those affected by industrial
development, may be less able to adapt. Elk clearly show some
ability to adapt to human activity [59]. Discussion Nonetheless, if the effect of
human activity is a threshold phenomenon [60], we might expect
there to exist a limit of physical disturbance corresponding to a
limit in the range of behavioral plasticity in ungulates, beyond
which redistribution, social, or demographic effects may be
observed [61], [62]. Examining only average responses across populations obscures
variability among individuals that may have important implica-
tions for management or provide new ecological insight. For
example, management strategies designed to conserve a resource
that is important, on average, to the population may overlook
resources that are critical to individuals that comprise a smaller
demographic segment that functions disproportionately in popu-
lation persistence. It has been stated ‘‘information on individual
resource use is necessary if we are to make the transition from
phenomenological models of population dynamics to mechanistic
models in which the dynamics of a population are predicted from
the properties of its components’’ [18]. Conditional estimates also
provide information on why marginal estimates may be counter-
intuitive, or how individuals assemble to comprise the marginal
estimate. For example, the marginal estimate of selection for road
density in our study indicated that elk selected for higher road
density in 3 of the 4 years comprising the study period (Figure 5). This observation is counter to most research on roads and
ungulates [63], [64]. Conditional estimates revealed that ,1/3 of
elk showed a positive association with higher road density but just
as many avoided high road density; this observation sets the stage
for examining potential links between a particular behavioral
strategy such as road avoidance and demographic responses. The
increase in the within-year proportion of elk occupying areas with
higher road density from 2007 to 2009 (Figure 5d), concurrent
with an increase in new road development, suggests that elk
maintained relatively consistent calving season use areas during
the study period, but modified their behavior as they became
increasingly constrained to select resources in proximity to
development. Such modification was consistent with a functional
response in resource selection; elk response to human activity
changed as a function of availability and with time of day. During
the day when humans are active elk showed avoidance behavior
that strengthened with increasing disturbance. This response
spilled over to elk selection for security cover which appeared to
have high importance to elk inside of the gas field as a day time
refugium. Discussion Outside of the gas field elk showed relatively weak day
time selection for cover and, as might be expected, selection
weakened as the proportion of cover within the seasonal use area
increased (sensu [40], Figure 7). These observations are consistent
with the hypothesis that ungulates face a trade-off that is mediated
by human activity in multiple-use landscapes – that is, the strength
of the trade-off varies in direct relation to resource availability
which is driven by human activity [41]. To our knowledge, fully random-effects RSF models do not
appear in the published literature. The random-effects framework
provided insight into individual behavior and how elk modified
patterns of calving season resource selection relative to develop-
ment. Individuals that moved between developed and undevel-
oped areas (Figure 6) were spatially aware and showed marked
changes in their behavior that were consistent with the hypothesis
that human activity during the day was perceived as a source of
risk [6]. An observation that warrants further attention in wildlife-
human interaction studies is that heterogeneity among individuals
in response to their environment was apparent, and this variation
was amplified in developed areas relative to undeveloped areas. Associated with this observation was generally more precise
estimation of selection within individuals in developed areas
(except selection for elevation). Among-individual variation can
comprise the majority of a population’s niche width, and only
when the within-individual component of total niche width is
constrained does between-individual variation become prominent. Of particular relevance here is that trade-offs remain among the
most plausible mechanisms for the observation of limited within-
individual variation [18]. If ungulates in developed areas must
make trade-offs associated with avoiding human activity, partic-
ularly during daylight hours, we might expect constrained within-
individual variation and thus more heterogeneity among individ-
uals. Human activity functioning to constrain decision-making in
ungulates is consistent with the notion that risky environments
impose pressures that disallow animals to respond to other features
as they otherwise would [57]. This could make it difficult to
establish or predict general patterns of resource selection during
periods in which ungulates show fidelity to historic ranges, yet
human activity rapidly modifies the landscape. In fact, we showed
that RSF maps based on day-time resource selection behavior in
developed areas had poor predictive accuracy. Spatially explicit models of relative predicted probability of use
validated poorly when they were based on resource selection
behavior inside of the gas field. PLoS ONE | www.plosone.org Discussion The model based on behavior
outside of the gas field validated well. To our knowledge, this
treatment/control concept is a novel approach in RSF-based
conservation planning. Typically, quantifying relative predicted
probability of use in human-modified areas is based on animal The study of behavioral syndromes offers a relevant framework
within which to discuss the conservation implications of individual
variation in human-wildlife interaction studies [58]. A behavioral
syndrome is a suite of correlated behaviors reflecting among-
individual consistency across multiple contexts. Within a syn- January 2011 | Volume 6 | Issue 1 | e14597 January 2011 | Volume 6 | Issue 1 | e14597 14 Ungulate Conservation extensive portions of the landscape, potentially resulting in large
errors in where conservation action would have its greatest impact. In population response research it is always desirable to include
spatial and temporal controls (before-after, control-impact design),
and demographic responses. Availability of such components is
uncommon in large-scale wildlife-human interaction studies. In
such situations the approach we describe, including efforts to
account for possible adaptive behavior among reproductive
groups, attention to the treatment/control concept, and a
random-effects modeling framework, should have general appli-
cation in human-wildlife interaction research particularly among
species that inhabit places where human activity is intense, or
among special status species for which little information on
resource needs exists. behavior in those areas. This approach has been used effectively; it
has been shown that predictive maps based on mule deer
(Odocoileus hemionus) behavior in human-modified areas validated
well [3]. In their study [3], variation among individuals was
present but limited relative to our findings. We suggest that
ungulates in developed areas often respond to human activity in
situation-specific ways. In our study, the local attributes of human
activity varied in space and time within the seasonal use area of
each elk making it difficult to generalize across the population. Situation-specific responses, including individual variation in the
strength of selection or avoidance response, induced heterogeneity
which complicated the application of models based on such
behavior for conservation planning. Protecting
unmodified
habitat
in
multiple-use
landscapes
typically is not an option [65]; therefore, measures to account
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relative probability of use differently from the models based on
behavior in the absence of industrial development throughout Author Contributions Conceived and designed the experiments: MRD SMH RGO LDHW
JBW. Performed the experiments: MRD SMH. Analyzed the data: MRD
SMH SLW. Wrote the paper: MRD SMH SLW. Field observation: MRD
SMH RGO JJW. Data collection: SMH RGO JJW. Conceived and designed the experiments: MRD SMH RGO LDHW
JBW. Performed the experiments: MRD SMH. Analyzed the data: MRD
SMH SLW. Wrote the paper: MRD SMH SLW. Field observation: MRD
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English
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Identifying priority areas for European resource policies: a MRIO-based material footprint assessment
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Journal of economic structures
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cc-by
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*Correspondence:
stefan.giljum@wu.ac.at
1 Institute for Ecological
Economics, Vienna University
of Economics and Business
(WU), Vienna, Austria
Full list of author information
is available at the end of the
article Identifying priority areas for European
resource policies: a MRIO‑based material
footprint assessment Stefan Giljum1*, Hanspeter Wieland1, Stephan Lutter1, Martin Bruckner1, Richard Wood2, Arnold Tukker3,4
and Konstantin Stadler2 Stefan Giljum1*, Hanspeter Wieland1, Stephan Lutter1, Martin Bruckner1, Richard Wood2, Arnold Tukker3,4
and Konstantin Stadler2 Giljum et al. Economic Structures (2016) 5:17
DOI 10.1186/s40008-016-0048-5 Open Access Abstract In the context of the transformation toward a “green economy,” issues related to natural
resource use have rapidly increased in importance in European and international policy
debates. The large number of studies applying economy-wide material flow analysis
so far mostly produced aggregated national indicators, making the results difficult to
connect to policies, which are often designed for single sectors or consumption areas. This paper provides a detailed assessment of the composition of EU’s material foot-
print in its global context, aiming at identifying the main product groups contributing
to overall material consumption and specifying the geographical sources for the raw
materials required to satisfy EU’s final demand. Based on multi-regional input–output
(MRIO) modeling, we apply production layer decomposition to assess supply chains
and their structural changes from 1995 to 2011. The global MRIO database used in this
study is EXIOBASE 3, which disaggregates 200 products and 163 industries, of which 33
represent material extraction sectors. By that means, we increase the level of detail to
a degree where policies can more easily connect to. We find that the generally grow-
ing material footprint of the EU was characterized by a dramatic shift regarding the
origin of raw materials, with the share of materials extracted within the EU territory
falling from 68 % in 1995 to 35 % in 2011. In 2011, raw materials extracted in China to
produce exports to the EU already contributed an equal share to EU’s material footprint
as material extraction within the EU itself. Import dependency is most critical for the
material group of metal ores, with only 13 % of all metals required as inputs to EU final
demand stemming from within the EU. Regarding product composition, construction
was confirmed as the most important sector contributing to the material footprint, fol-
lowed by the group of manufacturing products based on biomass. Materials embodied
in service sector activities together contributed a quarter to the total material footprint
in 2011, making services an important, but currently disregarded area for European
resource policies. We also find that supply chain structures became more complex over
time, with a growing part located outside the EU territory. Keywords: Material footprint, Multi-regional input–output analysis,
Production layer decomposition, Supply chains, Resource policies © 2016 The Author(s). © 2016 The Author(s). This article is distributed under the terms of the Creative Commons Attribution 4.0 International License
(http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium,
provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and
indicate if changes were made. 1.1 Policy background Economic development in the past decades was characterized by steadily increasing
levels of global resource use and rising human pressures on global ecosystems (Giljum
et al. 2014; Krausmann et al. 2009; UNEP 2011a). Driven by rapidly growing levels of
material consumption in emerging economies together with continued high levels of per
capita resource use in the industrialized world, the global economy is also characterized
by increasing competition between world regions regarding the access to raw materials. Against this background issues related to material consumption and resource productiv-
ity have rapidly increased in importance in European and international policy debates in
the past few years. In response to these environmental and economic challenges, numerous policy ini-
tiatives have been introduced aiming at significantly increasing resource efficiency to
reduce negative environmental impacts, while addressing issues related to material sup-
ply security and import dependency. Examples for these strategies on the international
level with a strong focus on resource efficiency include the “Green Economy” concept
promoted by UNEP (2011b) or the “Green Growth Strategy” of the OECD (2011). The most important policy strategy at the European level promoting resource efficiency
is the Commission’s Flagship Initiative “A resource-efficient Europe” (European Commis-
sion 2011b) as well as the subsequent “Roadmap to a resource-efficient Europe” (Euro-
pean Commission 2011c), both part of the implementation of the “Europe 2020” strategy. The overall objective of the Roadmap is to ensure that “by 2050 the EU’s economy has
grown in a way that respects resource constraints and planetary boundaries” and to
identify priority areas and actions “to put us on a path to resource efficient and sustain-
able growth” (p. 3). Three consumption areas receive particular attention in the Road-
map: food, housing and mobility. The food value chains are identified as causing major
global impacts, in particular related to the production of animal-based products. Regard-
ing the housing and building sector, the Commission recognizes the significant material
flows related to construction and maintenance activities of buildings. It is argued that
the aggregated impacts of housing and infrastructure account for around 15–30 % of all
environmental pressures caused by European consumption. The third high-impact area
regarding resource use identified in the Roadmap is mobility with its high demand for
material-intensive transport infrastructure and the high dependency on fossil fuels. In some consumption areas, notably food consumption, the Commission acknowl-
edges that European consumption causes negative global impacts. Abstract This article is distributed under the terms of the Creative Commons Attribution 4.0 International License
(http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium,
provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and
indicate if changes were made. Page 2 of 24 Giljum et al. Economic Structures (2016) 5:17 1.1 Policy background In general, however,
the Commission is rather concerned about the negative impacts on the EU economy
caused by international developments such as restrictions in raw material availability or
increasing raw material and food prices (European Commission 2011a). The most prom-
inent related policy strategy focusing on aspects of import dependency, resource supply
and access to natural resources is the Raw Materials Initiative (European Commission
2008, 2011d). In these documents, the Commission focuses on critical materials, in par-
ticular metal ores with high import dependency rates and high relevance for European
industries, and describes measures to secure and improve access to these raw materials. In the initiative, certain materials with specifically high dependency rates are identified,
such as indium, titanium and niobium, where the EU faces an import dependency rate Giljum et al. Economic Structures (2016) 5:17 Page 3 of 24 of 100 % (European Commission 2008). However, the supply and dependency aspects
are solely tackled from a perspective of direct imports and inputs into the European
economy, not from a supply chain perspective. Issues such as supply chain security and
dependency on indirect supplies via imported manufactured goods are not mentioned. Recently, the EU has also committed itself to the vision of a circular economy and has
presented a package of legislative and voluntary measures. This initiative aims at closing
the loops of materials and thus maintaining the value of resources and products as long
as possible within the economy, while minimizing the amount of generated waste (Euro-
pean Commission 2015). Given the increased demand for robust measures from the policy side, discussions on
the most suitable indicators to measure resource use, productivity and criticality of raw
materials are intensively ongoing. In recent years, awareness of policy makers generally
increased regarding the significance not only of materials and products directly used by
a national economy, or a regional economy such as the EU, but also of indirect resource
use required along supply chains and embodied in internationally traded products. Consideration of all indirect effects leads to a consumption—or footprint—perspective,
allowing illustrating the global impacts related to final demand. As a consequence, there
is increasing research on the differences between production-based versus consump-
tion-based trends of material use and its relation to GDP, in order to evaluate whether
decoupling is being achieved (UNEP 2011a). 1.1 Policy background Recent results suggest that assessments
of decoupling can lead to very different results when measured with different indica-
tors. Industrialized countries and regions, such as the USA or the EU, have in general
achieved a relative decoupling when measured with indicators of direct material use, i.e.,
production-oriented indicators. Some countries, such as Japan, were even able to reduce
direct material use in absolute terms (Giljum et al. 2014; Steinberger et al. 2013). How-
ever, when consumption-based—or footprint-type—indicators are applied, the decou-
pling performance is much weaker or even reversed, with material footprints growing
faster than GDP (Wiedmann et al. 2015). 1.2 Methodological background A rapidly growing body of literature assessing material flows considers indirect mate-
rial flows embodied in internationally traded products and applies consumption-based
or footprint-type indicators with the objective of providing complementary information
for the design of resource policies. Studies include national assessments (for example,
Kovanda and Weinzettel 2013; Schaffartzik et al. 2014), EU-wide studies (Arto et al. 2012; Schoer et al. 2012) as well as international assessments (for example, Bruckner
et al. 2012; Giljum et al. 2015; Wiedmann et al. 2015). Existing studies so far mostly pro-
duced aggregated national indicators on material consumption and resource productiv-
ity, making it difficult to use MFA results for policy making, which is often focused on
specific sectors or consumption areas. The drivers underlying these aggregated indica-
tors as well as the hotspot products and supply chains mostly contributing to the overall
results have rarely been investigated (the study by Acosta-Fernández (2007) on identify-
ing sectoral hotspots of material use in the German economy and related priority areas
of policy action is one of the very few exceptions). A more detailed understanding of
developments on the level of economic sectors and supply chains is essential to identify Giljum et al. Economic Structures (2016) 5:17 Page 4 of 24 areas in the production–consumption system, which needs to be transformed, if abso-
lute levels of material consumption should be reduced. Several methods are available to calculate consumption-based MFA indicators, each
having its advantages and drawbacks (Lutter et al. 2016; Schoer et al. 2013). Three
methodological approaches have been developed and applied so far: coefficient-based
approaches building on process analysis, environmental input–output analysis and
hybrid approaches combining elements from both approaches. Coefficient-based
approaches allow calculating material footprint-type indicators at a very high level of
detail, as coefficients can be calculated for a large number of single products without
any restrictions of pre-defined sectors or product groups. However, material input coef-
ficients are difficult to calculate for higher manufactured products and services with
complex supply chains. Coefficients are often not specific regarding geographical origin
of raw materials—in many cases, the geographical origin of a raw material is different to
the geographical origin of an imported manufactured good. Furthermore, based on coef-
ficient approaches, it is likely that the calculated total of global environmental pressures
related to consumption does not equal the pressures of worldwide production. 1.2 Methodological background Input–output analysis, in particular in the context of multi-regional input–output
(MRIO) models, has the key advantage of depicting the whole global economy and thus
allowing calculating material footprints for all products and sectors independently of the
complexity of the underlying supply chains. MRIO models also avoid double counting
as supply chains are clearly distinguished from each other. Furthermore, these mod-
els allow determining the geographical origin of each raw material related to the final
demand of a country or region. For these reasons, in this paper, we apply a MRIO-based
approach, based on the database EXIOBASE, which was specifically developed for envi-
ronmental-related assessments and distinguishes a large number of different raw materi-
als (see Sect. 2 for more details). 2.1 MRIO database For the material footprint assessments in this paper, we apply the MRIO database
EXIOBASE (Tukker et al. 2013, 2014; Wood et al. 2015), a very detailed global MRIO
database, developed in particular for environmental assessments of the EU. With 200
commodities and 163 industries, of which 33 represent the primary sector of the econ-
omy, EXIOBASE provides the highest consistent level of product and sector detail by
country across all currently available MRIO models (see Additional file 1: Full list of
EXIOBASE products and product groups). In EXIOBASE, agricultural activities are rep-
resented by 15 product groups and thus in a similar detail as in the GTAP model (Andrew
and Peters 2013). In contrast to most available MRIO tables, which aggregate mining and
quarrying of fossil fuels and mineral resources into one or two sectors (see, Giljum et al. 2015), EXIOBASE contains 12 extraction sectors for fossil fuels, 8 metal extraction sec-
tors and three sectors representing the extraction of industrial and construction minerals. It thus represents a major advantage in terms of detail for assessing global material flows. l
On the downside, EXIOBASE only details 44 countries (28 EU member states plus
16 major trade partners of the EU, together adding up to around 95 % of global GDP)
and covers all other countries in the form of five rest-of-the-world regions (see Addi-
tional file 2: Full list of EXIOBASE countries and regions). In contrast, the Eora database
(Lenzen et al. 2013) distinguishes 187 countries/regions, and the latest version of GTAP
(Narayanan et al. 2015) separately models 140 countries and regions. However, for
assessments of EU’s global environmental impacts, EXIOBASE is a very suitable data-
base. In this paper, we aggregated all results of the EU-28 countries into one region, in
order to provide an overall assessment of the EU as a whole. Future research may focus
on specific patterns of single EU countries. i
Furthermore, a high level of regional detail as provided by Eora or GTAP is required
to avoid errors caused by product and price inhomogeneity between countries in a cer-
tain region and becomes relevant when there are large trade flows from those regions. 1.3 Objectivesh This paper provides a detailed assessment of the composition of the EU’s material con-
sumption in a global context, with the aim to identify primary product groups con-
tributing to the EU’s global material footprint. We further aim at assessing the supply
chain structure of various product groups, thus identifying in which parts of the supply
chain major material extractions take place. The temporal and geographical coverage of
EXIOBASE further allows for the assessment of changes over time, for example regard-
ing the importance of raw material inputs stemming from non-European countries. This analysis elucidates whether outsourcing of environmental pressures through inter-
national trade occurred and illustrates which world regions serve as major suppliers of
raw materials directly and indirectly required to satisfy EU’s final demand. Based on the
empirical assessment, we aim at reflecting current priority areas of EU resource policy
making and at deriving recommendations for the priority areas and topics the EU should
focus in its future policy efforts. f
The remainder of the paper is structured as follows: Section 2 starts with a descrip-
tion of the MRIO and material flow data as well as of the applied analytical methods. Section 3 presents the various results from the MRIO assessment both on the level
of overall EU material consumption and on the level of different product groups and Page 5 of 24 Giljum et al. Economic Structures (2016) 5:17 their underlying supply chains. It also discusses the results in the context of current EU
resource policy priorities. The final Sect. 4 concludes. 2.2 Material extraction data and indicatorsh The material extensions in EXIOBASE were compiled using the WU Global Material
Flow database (WU 2015, available at www.materialflows.net). This database contains
material extraction data for all countries worldwide, for all raw material categories and
in annual time series from 1980 to 2013. The database is compiled following the stand-
ards of economy-wide material flow accounting (EW-MFA) provided by Eurostat and
the OECD (EUROSTAT 2013; OECD 2007). Four main data sources are used for the compilation of the database—the British Geo-
logical Survey (BGS) and the US Geological Survey (USGS) for metal and mineral data;
the International Energy Agency (IEA) for the data on fossil fuels; and the FAO for the
data on biomass extraction. The quality of the material extension data varies across the
different material categories, depending on the data source and necessary estimation or
manipulation steps. In general, the primary data from BGS, USGS, IEA and the FAO are
of good quality. However, for some types of material extraction, estimation techniques
need to be applied due to a lack of official statistics. This holds especially true for the
cases of the extraction of construction minerals, biomass uptake by rooming animals
(grazing) and the generation of by-products from agricultural harvest. For more infor-
mation on the compilation of the underlying material flow database, for example, regard-
ing the treatment of coupled production in metal mining, see Additional file 3: Technical
details on the compilation of the global material flow database and WU (2015). l
In order to generate the physical satellite to the MRIO system, the around 380 catego-
ries of primary raw material extraction data are aggregated into the 33 extraction sectors
of EXIOBASE (see Additional file 4: Correspondence table between raw materials and
EXIOBASE sectors). All MFA data are fed into the MRIO system in the unit of 1000 t. The main indicator generated with the MRIO system is the economy-wide indicator
raw material consumption (RMC), also known as material footprint (Giljum et al. 2015;
Wiedmann et al. 2015). The RMC or material footprint indicator comprises all domes-
tically and foreign raw materials that were directly and indirectly required to produce
goods and services of the EU final demand. The material footprint thus expresses the
amounts of materials embodied in final demand and aims at understanding the socio-
economic drivers, such as private or public consumption, of raw material extraction. 2.1 MRIO database We argue that for the analysis of the flows of raw materials, limited trade flows to the
EU come directly from the aggregate regions, and because those materials traded have
prices being mainly determined on global markets, price differences between countries
within an aggregate region will be smaller compared to price differences between raw
materials. Consequently, for this study a high level of commodity detail is regarded as
more relevant. In this paper, we apply EXIOBASE in its version 3.1, covering the time period of 1995
to 2011. In order to adequately represent the global economic structure over time,
EXIOBASE 3.1 combines several data sources to capture economic changes. Principle
data include macroeconomic accounts (GDP, total trade, final demand from UN SNA
database) and other detailed data sources (IEA, FAOSTAT, Comtrade, various mining
databases, BACI and others) to estimate product and industry output over time as well
as product-specific bilateral trade. The structural change of domestic economies was
estimated by inter/extrapolate observed changes of national IO tables over time (for 38 Giljum et al. Economic Structures (2016) 5:17 Page 6 of 24 of the 44 EXIOBASE countries, i.e., excluding Mexico, Luxembourg, Croatia, Indone-
sia, Turkey and Canada). Inconsistencies within the database caused by conflicting data
sets were resolved by balancing the systems using a flexible mathematical programming
approach where information gain is minimized (Lenzen et al. 2009). The estimation of
the uncertainty associated with such techniques has not received a great deal of atten-
tion in research, albeit Lenzen et al. (2010) have attempted to capture some of the impact
of these techniques for the case of greenhouse gas emissions. 2.3 Analytical methods The data analysis starts on the aggregated level illustrating the overall material footprint
time series of the EU followed by decompositions into the four main material categories,
into sectors and product groups as well as into source countries for raw materials. The
latter are identified for the total material footprint and separately for each aggregated
material group. This is carried out for the whole time period of 1995–2011 in order to
determine whether and to which regions the EU has outsourced material extraction
via international trade. We also analyze the sector composition, identifying the sec-
tor groups contributing most to the EU’s material footprint, and how this composition
changed over time, thus analyzing structure and temporal dynamic of the EU material
footprint. Apart from these standard analyses, MRIO models representing the entire global
economy and all underlying supply chains allow for the application of additional ana-
lytical tools. These tools assess supply chain structures underlying the total material
footprint of a product or product group and thus visualize the type and complexity of
intermediate deliveries. In this paper, we apply production layer decomposition, PLD
(compare Lenzen and Crawford 2009; Llop and Ponce-Alifonso 2015; Wiedmann et al. 2009), which is an aggregated form of structural path analysis (SPA). SPA allows open-
ing up the black box of national and international supply chains uncovering the larg-
est inter-sector and inter-country pathways of flows of goods and services through the
global economy (Lenzen 2007; Peters and Hertwich 2006). The benefit of the very high
level of detail through identifying all paths in the global economy related to final demand
in a certain country or region entails the disadvantage of producing very large data sets,
in particular when an SPA is performed for various types of raw materials and with time-
series data. Therefore, in order to derive more general conclusions from the MRIO model in terms
of the biophysical dimension of supply chain structures and temporal dynamics, in con-
trast to SPA, production layer decomposition does not calculate the “weight” of single
paths but aggregates all material extractions occurring at discrete supply chain layers. In
its core, PLD can be expressed as: (1)
ek,m = ˆsmAk ˆy ek,m = ˆsmAk ˆy (1) where ek,m equals the material extraction of raw material group m on layer k along the
path of a product or product group, which directly or indirectly serves final demand. 2.2 Material extraction data and indicatorsh It
is important to note that this approach differs from other types of material flow analysis,
which aim at physically tracking actual flows of certain raw materials or substances (for
example, Graedel et al. 2015; Laner et al. 2015). Furthermore, it needs to be emphasized
that the approach applied in this paper focuses on the footprint of primary materials. Integrating secondary raw materials from recycling in this assessment would require
additional material flow data (see Haas et al. 2015). Page 7 of 24 Giljum et al. Economic Structures (2016) 5:17 In MFA, indicators are generally decomposed into four major material categories, i.e.,
biomass, fossil fuels, minerals and metal ores. In this paper, we present indicators both
on the aggregated level across all materials and on the level of the four main material
groups. 2.3 Analytical methods ˆs represents the diagonalized direct intensities of material group m, which results
from dividing the domestic extraction vector of m by the sectors gross output vector
x, and where ˆx = Lˆy, with L being the Leontief inverse matrix and ˆy the diagonalized
final demand vector. Subsequently, ˆse = ˆmˆx−1. A is the technical coefficients matrix. For the case of layer 0, i.e., direct deliveries material extraction sectors to final demand:
ek=0,m = ˆsmIˆy, where I represents the identity matrix. where ek,m equals the material extraction of raw material group m on layer k along the
path of a product or product group, which directly or indirectly serves final demand. ˆs represents the diagonalized direct intensities of material group m, which results
from dividing the domestic extraction vector of m by the sectors gross output vector
x, and where ˆx = Lˆy, with L being the Leontief inverse matrix and ˆy the diagonalized
final demand vector. Subsequently, ˆse = ˆmˆx−1. A is the technical coefficients matrix. For the case of layer 0, i.e., direct deliveries material extraction sectors to final demand:
ek=0,m = ˆsmIˆy, where I represents the identity matrix. Page 8 of 24 Giljum et al. Economic Structures (2016) 5:17 Due to the fact that there are an infinite number of layers, we calculate the residual er+
via via (2)
Lr+ = L −I −
r
k=1
Ak and
(2)
Lr+ = L −I −
r
k=1
Ak
(3)
e
ˆs L
ˆy Lr+ = L −I −
r
k=1
Ak (2) and (3)
er+,m = ˆsmLr+ˆy er+,m = ˆsmLr+ˆy (3) where r is the selected threshold regarding the number of layers separately analyzed. where r is the selected threshold regarding the number of layers separately analyzed. We can then relate the single-layer results ek to the overall material footprint MF
through their summation: (4)
MF =
r−1
k=0
ek
+ er+ (4) Moreover, we can use production layer decomposition to disaggregate the individual
layer results into the amount of raw materials extracted within and outside the EU. 3.1 Resultsh The results section of the paper is structured by starting with aggregated results for
the EU material footprint and subsequently decomposing the overall trends in various
dimensions. The first perspective is the origin of the various raw materials related to EU
final demand, followed by an analysis of the main products and product groups contrib-
uting to the EU material footprint. Based on the PLD, we then illustrate the structure of
supply chains behind each of the aggregated product groups and assess how the layer
structure evolved over time. For selected product groups, we also discuss how the distri-
bution between materials extracted in the EU versus abroad changed on each layer from
1995 to 2011. Finally, we provide a disaggregation of the material footprint layers by the
main material categories, thereby analyzing the structural differences in the use of cer-
tain material groups in the economic system. Figure 1 illustrates the overall development of the material footprint of the EU, disag-
gregated by the four main material groups (top) and by the countries and world regions,
where the raw materials were extracted to directly and indirectly serve final demand in
the EU (bottom). The overall material footprint of the EU rose from 8.4 billion tonnes in 1995 to 12.2
billion tonnes in 2011. After a 5-year period of relatively strong increases between 2002
and 2007, the impact of the economic crisis in 2008 and in particular in 2009 is clearly
visible in the overall trend. In these two years, the EU material footprint dropped by
around 13 % compared to the maximum level of 2007. In 2011, the EU material footprint
regained an absolute level that was equal to the level in 2005/2006. When analyzed by main material groups, it is clearly visible that the category of
industrial and construction minerals contributes the largest share to the material foot-
print (54 % in 2011, up from 43 % in 1995). Minerals were also the group that most Page 9 of 24 Giljum et al. Economic Structures (2016) 5:17 Fig. 1 Overall material footprint of the EU-28, by main material category (top) and main countries of origin of
raw materials (bottom), 1995–2011, in billion tonnes Fig. 3.1 Resultsh 1 Overall material footprint of the EU-28, by main material category (top) and main countries of origin of
raw materials (bottom), 1995–2011, in billion tonnes significantly declined in the period of the economic crisis, contributing around half of
the overall reduction between 2007 and 2009. This indicates the strong impact of the
crisis on the construction sector, in particular in countries such as Spain, Italy or the
UK, but also in some Eastern European countries such as Romania. Direct and indirect
extraction of biomass grew by 35 % over the whole time period, and the required metal
ores to satisfy EU final demand was increased by 116 %, but from a lower absolute level. Fossil fuels were the only category where the EU consumed less in 2011 compared to
1995 (−14 %). From the perspective of the origin of raw materials, dramatic shifts can be observed
over the analyzed time period. The share of raw materials that were extracted within the
territory of the EU fell significantly from 68 % in 1995 to 35 % in 2011. This illustrates
that today around two-thirds of the raw materials that are directly and indirectly needed
to satisfy European final demand is extracted in other world regions. As illustrated in
Fig. 1 (bottom), the most significant shift occurred between the EU and China, which in
2011 already contributed an equal amount of raw materials related to EU’s final demand Page 10 of 24 Giljum et al. Economic Structures (2016) 5:17 for goods and services. This particular pattern is closely related to the current develop-
ment stage of China, which—as a fast-growing emerging market—is in the process of
building up its housing, transport and energy infrastructure. On the one hand, this leads
to significant amounts of raw materials required for investments and capital formation,
which dominate China’s own material footprint (see Giljum et al. 2015). On the other
hand, construction materials also serve as indirect inputs to manufacturing industries,
which produce goods exported to the EU (see also Wang et al. 2014; Wiedmann et al. 2015). This indirect flow of construction materials grew from around 230 million tonnes
in 1995 to more than 3 billion tonnes in 2011 and was thus far above flows of other key
raw materials extracted in China and serving EU final demand, such as coal (394 million
tonnes) or iron ores (157 million tonnes). 3.1 Resultsh As the material footprint calculations can be performed separately for various raw
materials and then be aggregated into the four major material groups, Fig. 2 depicts the
regional origin of raw materials serving the EU final demand for biomass, fossil fuels,
minerals and metal ores. The overall development as well as the regional patterns of origin differs signifi-
cantly across the four main material categories. However, two general trends can be
observed for all four raw material groups. First, extraction of raw materials within the
EU decreased, while the material footprint increased for all four categories except fossil
fuel consumption, which in 2011 was on an equal level as in the mid-1990s. This implies
that the EU is to an increasing extent directly as well as indirectly dependent on imports
from other world regions. Second, the Asia and Pacific region serves as the main source Fig. 2 Biomass, fossil fuels, minerals and metal footprint of the EU-28, by main world region of origin of raw
materials, 1995–2011, in million tonnes Fig. 2 Biomass, fossil fuels, minerals and metal footprint of the EU-28, by main world region of origin of raw
materials, 1995–2011, in million tonnes Giljum et al. Economic Structures (2016) 5:17 Page 11 of 24 of raw materials embodied in European imports across all four major categories, with
China playing an outstanding role. Biomass is the material category, where the highest share of raw materials still stems
from within the EU territory. The share was 76 % in 1995 and decreased to 47 % in 2011. With 35 % in 2011, biomass originated to a significant extent from Asia, up from 15 % in
1995. China played an important role contributing 16 % to biomass extraction in 2011,
in particular related to the production of cattle and other meat. Three percentage of bio-
mass in the EU material footprint was extracted in India with a similar product focus as
in the Chinese case. With 1 %, Indonesia was the third largest Asian contributor, with a
clear focus on the sector of “oil seeds,” i.e., palm oil. Smaller amounts of biomass were
embodied in imports from America and Africa. Fossil fuels extraction within the EU decreased significantly, from around 900 million
tonnes in 1995 to around 570 million tonnes in 2011, in particular due to a decrease in
coal mining. 3.1 Resultsh With 41 %, the Asia and Pacific region made up the largest fraction of fos-
sil fuels embodied in EU final demand in 2011. Fossil fuels embodied in imports from
China were rapidly increasing in the 2000s and made up 23 % of the total fossil fuel foot-
print of the EU-28 in 2011. These fossil fuels included, for example, coal (394 million
tonnes in 2011) that was extracted in China and used in electricity production, which
is then embodied in Chinese exports. Russia also plays an important role as a direct
exporter of particularly natural gas to the EU. The Middle East contributed 15 % to the
fossil fuel footprint in 2011. Minerals extraction within the EU also showed a decreasing trend, with a particu-
larly sharp decrease during the years of the economic crisis, underpinning the argument
above that the construction sector in many EU countries was one of the most effected
parts of the economies. At the same time, the minerals embodied in imports from the
Asia and Pacific region grew almost by a factor of 5. With more than 3 billion tonnes in
2011, China alone contributed 48 % to the total EU mineral footprint, almost exclusively
in the categories of construction materials, such as sand, gravel and stone. From a European policy perspective, the situation regarding metal ores can be regarded
as most critical, as Europe has only a minor production of metals within its territory and
is thus heavily dependent on imports from other world regions. In 2011, only 13 % of all
metals required as direct and indirect inputs for products and services finally consumed
in the EU stemmed from within the European territory. Again, the Asia and Pacific region
plays the most important role as a raw material provider (50 %). Most important extrac-
tion countries are China, which accounts for 27 % of total EU metal footprint, mostly
in the categories of iron ore and copper. Australia delivered 6 % of metals serving as
indirect inputs to EU final demand, with iron ore, copper and precious metals being the
most important commodities. Three percent of the metal footprint originated in Indone-
sia, dominated by lead and zinc. Thirty-two percentage of metals embodied in EU’s final
demand were extracted in the Americas, with Latin American countries providing almost
three quarters, mostly iron ore, copper, lead, zinc and precious metals. In Fig. 3.1 Resultsh 3, we turn to another dimension, which the MRIO model allows to put into
focus, i.e., the product groups contributing to the material footprint of the EU. In order to
facilitate visualization, we aggregated the 200 EXIOBASE products into 11 broad product
groups (see Additional file 1: Full list of EXIOBASE products and product groups). Page 12 of 24 Giljum et al. Economic Structures (2016) 5:17 Fig. 3 Sectoral composition of the EU-28 material footprint, 1995–2011 Fig. 3 Sectoral composition of the EU-28 material footprint, 1995–2011 Figure 3 illustrates that the composition of the EU material footprint in terms of
aggregated product groups remained relatively stable across the observed time period,
despite the overall growing material footprint (compare Fig. 1 above). The main aggre-
gated sector contributing to the material footprint was construction, which held a share
of 25 % in 1995, falling slightly to 23 % in 2011. Construction was followed by the group
of manufacturing products based on biomass, most importantly from food-processing
sectors. Its share was 18 % in 2011, similar to the contribution in 1995. Manufactur-
ing of products mainly based on fossil fuels ranked third in 2011, with a share of 14 %. Most important products were motor gasoline with a material footprint of 189 million
tonnes, followed by diesel oil (92 million tonnes). The fourth biggest contributing sec-
tor was the large group of manufacturing products based on metals and minerals (8 %). With a material footprint of 393 million tonnes, motor vehicles were the most important
product group within this aggregated sector, followed by machinery and equipment (232
million tonnes) and radio, television and communication products (129 million tonnes). The contribution of agriculture and forestry products to the EU material footprint in
2011 was 5 %, than that of mining products 3 %. Although raw materials are extracted
only in these two aggregated sectors (see Fig. 4; Table 2), the overwhelming part of the
extracted raw materials are not directly delivered to final demand, but purchased by
other sectors of the economy for further processing. Materials embodied in service sec-
tor activities together contributed 25 % to the total material footprint, as the group of
“other products” mainly contains service sectors, including the large service sectors of
health, education and public administration and defense (see Table 1 for more details). 3.1 Resultsh As indicated in the introduction, one key strength of EXIOBASE is the very high prod-
uct detail. In Table 1, we further decompose the product groups contributing to the EU
material footprint and show the top 30 EXIOBASE products (the full list with the foot-
prints all EXIOBASE products can be found in the Additional file 5: Detailed material
footprint (MF) results). These 30 products made up around 80 % of the total EU material footprint in 2011. Table 1 indicates that among the top 30 contributing products, almost all of the 11 aggre-
gated product groups are represented. The product “construction work,” which is iden-
tical to the aggregated group “construction” as illustrated above, constitutes by far the Page 13 of 24 Giljum et al. Economic Structures (2016) 5:17 Fig. 4 Production layer decomposition of material footprint of EU-28 aggregated product groups, 2011 largest product group in the overall EU material footprint (23.2 % in 2011). Construction
had both a high share in total EU final demand (9.9 %) as well as a high material intensity
or almost 2.3 kg of direct and indirect material extraction required to produce one Euro
of final demand. Given this huge contribution of just one product, it would be desirable
to further split up this product group. It is interesting to note that on the more detailed sector level, service sectors are also
found among the top contributors. For example, health and social work services ranked
second in the product list, with a contribution of 6.6 % to the total material footprint in
2011. This sector contributed 12.8 % to EU’s monetary final demand in that year; however,
material intensity was significantly lower (half a kg of material input per €). The reason,
why this sector has a significant material footprint is that the provision of services in the
health sector requires a large number of material-intensive products serving as indirect
inputs, including, for example, chemical and pharmaceutical products. Also, food provi-
sion in hospitals and other social institutions induces significant flows of biomass, with
meat and animal products contributing most to the biomass footprint. Apart from health
and social services, also public administration and defense (4.5 %) as well as real-estate
services (3 %) has significant shares in the overall material footprint of the EU-28. 3.1 Resultsh i
Following aggregated product groups such as “food products nec” and “chemicals nec,”
some specific manufacturing product groups are also located among the top contribu-
tors. They include motor vehicles (with a share of 3.2 %) and machinery (1.9 %). Ani-
mal products, which are well known for their high material footprint (Lettenmeier et al. 2014), are not found among the top 10 EXIOBASE products. This fact is a consequence
of the construction principles of EXIOBASE, which further split up environmentally
sensitive products. In fact, EXIOBASE contains 11 products related to production of
meat and dairy products. If the material footprint of these sectors is again aggregated,
animal products together contribute almost 900 million tonnes, or 7.1 %, to the total
material footprint of the EU-28 and rank second after construction. Page 14 of 24 Giljum et al. Economic Structures (2016) 5:17 Table 1 Top 30 products contributing to the EU-28 material footprint (MF), values of final
demand (FD) and material intensities (kg/€), 1995 and 2011
Product
Product group
MF 1995
MF 2011
% change
FD 2011
Intensity
2011
1000 t
1000 t
1995–2011
million €
kg per €
Construction work
Construction
2,135,125
2,834,192
33
1,239,222
2.29
Health and social work
services
Other products
371,974
799,155
115
1,607,816
0.50
Food products nec
Manuf. products
(biomass)
542,018
654,621
21
280,307
2.34
Public administration and
defense services
Other products
357,479
543,163
52
1,036,643
0.52
Chemicals nec
Manuf. products (fos-
sil fuels)
234,641
509,761
117
157,823
3.23
Motor vehicles, trailers
and semi-trailers
Manuf. products
(metals/min)
217,156
393,023
81
348,919
1.13
Real-estate services
Financial services
311,943
362,505
16
1,113,134
0.33
Furniture and other
manufactured products
nec
Manuf. products
(metals/min)
149,487
277,609
86
240,462
1.15
Membership organization
services nec
Other products
128,809
236,885
84
455,624
0.52
Machinery and equip-
ment nec
Manuf. products
(metals/min)
194,722
232,110
19
206,048
1.13
Education services
Other products
145,041
222,463
53
893,586
0.25
Recreational, cultural and
sporting services
Other products
117,288
218,279
86
496,697
0.44
Vegetables, fruit, nuts
Agri. and for. prod-
ucts
170,744
193,087
13
79,544
2.43
Motor gasoline
Manuf. products (fos-
sil fuels)
229,335
188,913
−18
129,213
1.46
Products of meat cattle
Manuf. products
(biomass)
176,350
185,881
5
25,177
7.38
Hotel and restaurant
services
Sales and retail
services
108,763
175,816
62
605,336
0.29
Wearing apparel; furs
Manuf. products
(biomass)
109,155
170,524
56
160,629
1.06
Meat products nec
Manuf. products
(biomass)
93,166
155,681
67
75,501
2.06
Fish products
Manuf. 3.1 Resultsh products
(biomass)
77,426
155,457
101
89,126
1.74
Products of meat poultry
Manuf. products
(biomass)
79,197
136,148
72
72,534
1.88
Radio, television and
communication equip-
ment
Manuf. products
(metals/min)
54,644
129,304
137
128,149
1.01
Dairy products
Manuf. products
(biomass)
80,227
122,533
53
78,554
1.56
Electricity by coal
Electricity and water
supply
107,615
115,171
7
14,537
7.92
Distribution services of
gaseous fuels
Electricity and water
supply
125,924
114,488
−9
67,903
1.69
Fabricated metal prod-
ucts, except machinery
Manuf. products
(metals/min)
66,117
104,590
58
79,618
1.31
Transport services equip-
ment
Manuf. products
(metals/min)
38,244
99,858
161
88,686
1.13
Basic plastics
Manuf. products (fos-
sil fuels)
51,881
99,469
92
89,099
1.12 Table 1 Top 30 products contributing to the EU-28 material footprint (MF), values of final
demand (FD) and material intensities (kg/€), 1995 and 2011 Page 15 of 24 Giljum et al. Economic Structures (2016) 5:17 Table 1 continued
Product
Product group
MF 1995
MF 2011
% change
FD 2011
Intensity
2011
1000 t
1000 t
1995–2011
million €
kg per €
Sand and clay
Mining products
110,445
97,418
−12
1354
71.93
Ceramic goods
Manuf. products
(metals/min)
54,608
96,508
77
8511
11.34
Leather and leather
products
Manuf. products
(biomass)
44,832
95,763
114
59,771
1.60
Other product
groups
1,741,897
2,472,081
42
2,603,563
Total footprint
8,426,255
12,192,456
45
12,533,084 Table 1 continued The material intensities differ significantly across the various products. More than 7 kg
of material extraction served as direct and indirect material input to produce one € of
final demand of cattle meat. For vegetables and fruits, it was almost 2.5 kilograms. To
produce one € of final demand of motor vehicles, around 1.1 kg of raw materials were
required. The highest material intensities can be observed for mining activities, such as
the extraction of sand and clay (almost 72 kilograms of extraction per € of final demand
in 2011). Production layer decomposition allows investigating the material dimension of supply
chains and illustrating the structure and complexity of intermediate deliveries behind
the final use of a certain product or service. The numbering of the layers reflects the
distance to the extraction of the various raw materials required to produce the final
product, i.e., the number of intermediate deliveries before a product or service ends up
in final demand. Figure 4 depicts the composition of all layers of the respective supply
chains to the absolute material footprints of the 11 aggregated product groups. 3.1 Resultsh All layers
together constitute the total footprint of a product group. In order to keep complexity
on a manageable level, we illustrate Layers 0 to 4 separately and add up all higher levels
in the category Layer 5+. Table 2 complements the information in Fig. 4 by detailing the
percentage contributions of each layer to the product group total, both in the year 1995
and in 2011. As explained in Methods section above, Layer 0 only contains values for those product
groups, which are directly related to material extraction activities, i.e., biomass extrac-
tion from agriculture and forestry as well as mining of fossil fuels, minerals and metal
ores. These two aggregated product groups are thus the only ones that have direct deliv-
eries from material extraction to final demand. For agriculture and forestry products,
the share of Layer 0 was 56 % in 2011, for mining products 72 %. Material extractions taking place on Layer 1 are located one step away from the prod-
uct group in focus, i.e., they represent the final intermediate deliveries before a product
is being brought to its final purpose. Taking the example of the group of manufacturing
products from biomass, we observe that Layer 1 has the largest contribution (31 % in
2011). These are mainly deliveries of biotic raw materials from the agricultural sectors to
food-processing sectors as well as of harvested timber to sectors producing wood-based
products. Page 16 of 24 Giljum et al. Economic Structures (2016) 5:17 Table 2 Share of layers in material footprint of product groups, 1995 and 2011
Layer 0
(%)
Layer 1
(%)
Layer 2
(%)
Layer 3
(%)
Layer 4
(%)
Layer 5
+ (%)
Material
footprint (1000 t)
Agri. and for. products
1995
69
12
8
5
3
4
543,263
2011
56
13
11
7
5
8
638,561
Mining products
1995
78
15
3
2
1
1
295,577
2011
72
17
4
2
2
3
334,663
Manuf. products (biomass)
1995
0
44
21
14
8
12
1,521,136
2011
0
31
22
16
11
20
2,169,646
Manuf. products (metals/min)
1995
0
37
25
15
9
14
713,056
2011
0
31
26
16
10
18
1,004,042
Manuf. 3.1 Resultsh products (fossil fuels)
1995
0
17
24
21
14
24
996,043
2011
0
13
20
19
15
33
1,742,524
Electricity and water supply
1995
0
60
20
9
5
7
353,418
2011
0
51
18
11
7
13
370,134
Construction
1995
0
38
36
14
6
7
2,135,125
2011
0
31
33
16
8
12
2,834,192
Sales and retail services
1995
0
9
40
21
12
18
126,164
2011
0
5
33
22
14
26
194,515
Transport services
1995
0
10
28
22
15
24
95,321
2011
0
6
22
20
16
35
178,545
Financial services
1995
0
11
33
25
13
18
458,940
2011
0
7
23
23
16
31
608,436
Other products
1995
0
12
31
22
14
21
1,188,212
2011
0
6
24
23
16
31
2,117,199 Table 2 Share of layers in material footprint of product groups, 1995 and 2011 Layer 2 contains extractions of raw materials, which are two intermediate steps away
from the final product. An example from the construction sector, which is characterized
by the largest share of inputs from Layer 2 across all aggregated product groups (33 % in
2011), is the sequence of limestone being extracted in mining, delivered as an input to
cement production, with cement then purchased by construction activities. With regard to the composition and temporal development of the structure of sup-
ply chains across the various layers, two key observations can be drawn from Table 2. First, the distribution across the 5 analyzed layers changes across sectors, as supply
chains increase in complexity. While products from the primary sectors contain by far
the largest parts of inputs on Layer 0, the manufacturing sectors typically peak on Layers
2 or 3. Services have an even higher share of high-layer extractions in their total material Page 17 of 24 Giljum et al. Economic Structures (2016) 5:17 footprint. The second important conclusion concerns the changes of layer structures
over time. Table 2 illustrates that in the past 20 years, supply chains across all product
groups have become more complex, i.e., percentage contributions of layers with lower
numbers were declining in almost all cases, whereas Layers 3–5 + gained in importance. This illustrates that supply chains are increasingly complex and differentiated in their
organization, and an ever-increasing number of production steps is interlinked for the
production of goods and services delivered to final demand. 3.1 Resultsh i
Figure 5 takes the analysis one level further by illustrating to what extent the changes
in the supply chain structures were linked to the overall trends of outsourcing mate-
rial extraction away from the EU territory to other world regions discussed for the over-
all material footprint in Figs. 1 and 2. For illustrative purposes, we select two product
groups with a major importance in the overall EU material footprint, i.e., construction
and manufactured products based on biomass. Figure 5 illustrates that significant geographical shifts can be observed on all lay-
ers when comparing the years 1995 and 2011, i.e., the contribution of material extrac-
tions within the EU was shrinking, while extractions in the non-EU countries gained in
importance (compare the aggregated results in Figs. 1 and 2 above). Construction is a
pronounced example: While 83 % of extractions as inputs to construction activities—
in particular on Layers 1 and 2—were taking place within the EU borders in 1995, this
value decreased to only 38 % in 2011. In the latter year, significant material extractions
took place in the non-EU countries on all layers. For example, on Layer 1, large material
extractions outside the EU took place regarding construction minerals, such as sand and
clays, which are then further processed into construction materials imported into the
EU. On Layer 2, metal ores used in construction, notably iron ore, as well as fossil fuels
(coal and oil) serving as energy inputs into manufacturing of construction materials play
major roles. Fig. 5 Material extraction per layers in the EU versus the non-EU countries for construction and biomass-
based manufactured products, 1995 and 2011 Fig. 5 Material extraction per layers in the EU versus the non-EU countries for construction and biomass-
based manufactured products, 1995 and 2011 Page 18 of 24 Giljum et al. Economic Structures (2016) 5:17 A similar, but less accentuated development could be observed for the group of manu-
factured products from biomass. Also here the ratio of domestic to foreign extractions
changed from 73:27 % in 1995 to 40:60 % in 2011. However, there are still significant
shares of extraction within the EU on Layer 1, indicating that the EU food- and wood-
processing industries are still to an important extent supplied by agricultural and for-
estry products harvested within the EU. 3.2.1 Priorities for European resource policies The results generated with the global MRIO model with detailed material extensions as
applied in this paper confirm the priorities of EU resource policy initiatives with regard
to product groups, i.e., we find that construction as well as the group of manufactured
biomass-based products are the top 2 product groups constituting EU’s material foot-
print. Transport services as such play only a minor role as a final demand category;
however, construction activities also cover the resources used in developing transport
infrastructure. Our analysis illustrated that there are important product groups in addi-
tion to the commonly known high-impact activities of construction, food and mobil-
ity. Most notably, we found that, with around a quarter of the total, the service sectors
contribute a significant share to the overall material footprint of the EU. Examples for
service sectors with high absolute importance are health and education services, public
administration and defense as well as real-estate services. These activities are character-
ized by very complex supply chains, i.e., the provision of services typically requires a
large variety of highly manufactured products as inputs. As the layer analysis illustrated,
these products are characterized by a significant number of processing stages. The tran-
sition toward a “knowledge-based service economy” is often assumed to be an environ-
mentally positive development. While services do have a lower material intensity per €
of generated added value compared to other parts of the economy (see Table 1 above
and the detailed table in the Additional file 5), the growing absolute importance of the
service sectors in generating European GDP, however, results in high absolute material
consumption of services. This is in line with similar finding regarding the carbon foot-
print of countries (Hertwich and Peters 2009; Piaggio et al. 2015). A service-oriented
economy thus is not automatically dematerialized or decarbonized, when taking a
full supply chain perspective. This fact is currently being overlooked in the European
resource policy context. Production layer decomposition, which revealed the underlying structure of supply
chains for different product groups as well as for aggregated raw material groups, also
provides important insights for EU resource policies. On the one hand, the disaggre-
gation by material groups suggests that resource policies should take into account the
different levels of complexities of supply chains. 3.1 Resultsh However, raw materials for higher processed
inputs to the production processes of biomass-based manufactured products are origi-
nating to a growing extent outside the EU. This trend can also be observed for other
product groups. In the final analytical step, we disaggregate the information from the layer analysis by
the four main material groups of MFA. Figure 6 illustrates on which of the layers each of
the four raw material groups was mainly extracted to serve the overall EU final demand
for products and services in the year 2011. We can observe different patterns across the four aggregated material categories. The
use of minerals shows the steepest distribution across all layers, peaking at Layer 2. This
implies that products being produced from extracted mineral resources have relatively
short supply chains (compare also the layer structure of the construction sector in Fig. 4
and Table 2). This is not surprising, as the major products from minerals are construc-
tion and industrial minerals, which are either directly delivered from the mining sector
(e.g., sand and stones) or delivered from the sectors producing further processed con-
struction materials, such as cement. Biomass also has a curve, which peaks at Layer 1,
indicating that there are only one or two steps of processing before the largest share of
extracted biomass is being delivered to final demand (compare the structure of the agri-
cultural/forestry product group and the biomass-based manufactured products in Fig. 5
above). The distribution of the other two main material categories, i.e., fossil fuels and
metal ores, shows a different pattern. Their slope is much lower, indicating that a larger
fraction of the overall fossil and metal footprint is generated on higher layers, i.e., further
away from the final product. Fossil fuels and metal ores thus mainly serve as inputs to
goods and services with more complex supply chains, indicating their fundamental char-
acter for the production processes of various sectors across the whole economy. Fig. 6 Distribution of main material groups across production layers, EU-28 material footprint, 2011 Page 19 of 24 Giljum et al. Economic Structures (2016) 5:17 3.2.1 Priorities for European resource policies Products based on minerals and bio-
mass are characterized by less complex intermediate delivery structures compared to
fossil fuels and metal ores, which serve as inputs for a large number of different products
across numerous economic sectors. Given the very high levels of import dependency
in these two product groups (compare Fig. 2 above), this puts the EU in a vulnerable
position. On the other hand, the layer analysis illustrated that an increasing share of
materials is extracted outside the EU on each of the single layers, thus pointing to the
importance of considering the overall supply chain, if an absolute reduction in global
material extraction related to EU consumption should be achieved. Secondary materials from recycling activities were not explicitly considered in this
study, which focused on the material footprint of primary extraction. Recent estimates
illustrate that there exist huge potentials to increase the degree of circularity in the
European economy. The aggregated recycling rate, calculated as the amounts of recy-
cled materials as a share of all processed materials, was only 12.6 % in 2005 (Haas et al. Giljum et al. Economic Structures (2016) 5:17 Page 20 of 24 2015). An ambitious implementation of the circular economy package would reduce the
European demand for primary raw materials and thus would also decrease the mate-
rial footprint as presented in this paper. Moving to a circular economy is therefore also
closely connected to economic issues, such as import dependence and stable supply of
raw materials for European industries (European Commission 2015). The global impacts related to material consumption in Europe are mentioned in some
of the EU policy documents, most notably with regard to biomass consumption. How-
ever, the results of the MRIO-based assessments in this paper suggest that a supply
chain perspective is of key importance for the EU economy, which has a higher material
import dependence than all other world regions (see Giljum et al. 2015). The consump-
tion of goods and services in the EU is to a large and rapidly growing extent dependent
on indirect material extractions taking place outside the EU’s territory. This has environ-
mental implications, as raw materials originate to an increasing extent from develop-
ing and emerging economies, which often have lower levels of environmental standards
compared to industrialized countries (for example, Hilson 2012). 3.2.1 Priorities for European resource policies The expansion of raw
material extraction activities also often leads to an increasing number of environmental
conflicts with the local population, which faces, e.g., dislocations due to expansions of
mining areas or accumulations of air and water pollution with severe negative effects on
their daily lives (Martinez-Alier 2014). Thus, if considering pollution and other negative
environmental consequences related to material extraction in impact-oriented material
footprint indicators (van der Voet et al. 2009), the geographical location of raw mate-
rial extraction could play an increasingly important role in determining Europe’s envi-
ronmental performance. Also, labor conditions are often very low in developing country
activities related to agricultural production or mining, and material extraction often fos-
ters enclave types rather than inclusive forms of development (for example, Bird 2014). As a final consumer of products to which these raw materials serve as direct and indirect
inputs, Europe should acknowledge its share in the responsibility for the environmen-
tal and social conditions in other world regions. Europe could more actively engage in
financial, technological and knowledge support in extraction countries to raise environ-
mental and social standards, as underlying, for example, the idea of a Global Marshall
Plan (Yunker 2014). This responsibility is currently not sufficiently assumed in EU poli-
cies related to raw materials. For example, the raw material initiative is strongly focused
on securing access to resources in particular in developing countries, which sometimes
contradicts the objectives of EU development policies, which aim at helping develop-
ing countries combat poverty through investing in the sustainable management of their
natural resources (Heinrich Böll Foundation 2011). 3.2.2 Methodological aspects The MRIO analysis performed in this paper allocates material extraction due to eco-
nomic activities through the global supply chains to the final consumers. As such,
MRIOs foremost rely on reliable data on domestic material extraction, as any errors or
uncertainty in the extraction data are passed through the full analysis and directly affect
the footprint results. Although extraction data are similar across global databases with
regard to categories such as agricultural production or extraction of fossil fuels (Fis-
cher-Kowalski et al. 2011), estimations need to be applied for quantitatively important Giljum et al. Economic Structures (2016) 5:17 Page 21 of 24 categories, such as construction materials (see above and Additional file 3 for details). This calls for further international efforts to align existing estimation approaches,
including the consideration of recycled construction materials (Hashimoto et al. 2009;
Wiedenhofer et al. 2015). Allocating material extraction to those industries or users extracting the material
requires detailed knowledge about the composition of each industry—data that are not
readily available. For example, rock aggregate is a high mass but low-value commodity
that is sometimes traded and sometimes extracted directly. Allocating rock aggregate
extraction to the quarrying industry, as is done in EXIOBASE, assumes that all rock
aggregate is traded, whereas in some countries, quarrying may be limited to higher-
value commodities, and rock aggregate may be extracted directly by the construction
industry (Schaffartzik et al. 2014; Schoer et al. 2013). f
High sector detail can help reducing this uncertainty (de Koning et al. 2015; Lenzen
2011). EXIOBASE provides the highest harmonized sector detail of MRIO models, but
because data are not available at such detail for all countries, a combination of LCA
data, proxy country IO data (from some specific, highly detailed national IO tables) and
sector-specific information (e.g., energy balances from the IEA for the energy sector)
was used for disaggregation (Wood et al. 2015) where regional differences can have an
impact, particularly in assessments of material flows (Stadler et al. 2014). l
A general caveat to current MRIO analyses is that the production of capital goods
including infrastructure is not allocated to the users of the infrastructure (by sector, or
over time), such that results show the annual resource requirement of various activi-
ties. 3.2.2 Methodological aspects This approach provides consistency with national accounting, but could be relaxed
by endogenizing capital goods and infrastructure in the coefficient matrix (see Lenzen
2001) to allocate infrastructure to goods and services consumed by households and
government. 4 Conclusions In this paper, we provided a detailed assessment of the composition of the material foot-
print of the EU and its underlying supply chain structures. We argued that recent devel-
opments in the compilation of multi-regional input–output databases open up a range
of new analytical options for the assessment of global material flows related to final
consumption of a country or world region. A particular strength of the MRIO approach
to analyze material flows is the possibility to identify the most important products and
product groups contributing to the material footprint as well as to investigate the inter-
national supply chains underlying the final goods and services delivered to final demand
through applying production layer decomposition. This type of analysis goes far beyond
the calculation of headline indicators on material consumption and material productiv-
ity, which so far has dominated the MFA literature, and allows connecting much better
to issues related to the development of resource policies, which are currently emerging
in the context of European policy making. The objective of this paper was to provide an illustration of the analytical potentials
starting from the macro-level of the overall EU material footprint and disaggregating it
in several dimensions. However, there is still a lot of room to expand this type of research
further and to derive more specific results. One option is to focus the analysis on single Giljum et al. Economic Structures (2016) 5:17 Page 22 of 24 raw materials instead of aggregated material groups. Full structural path analysis (SPA)
could also be applied for selected products of particular policy relevance, in order to
identify hotspots of material extraction on each layer of the supply chain, for example,
with regard to service sectors with important contributions to the overall material foot-
print. Finally, it is important to continue work on uncertainty analyses in connection
with material footprint models based on various MRIO databases. Author details
1 1 Institute for Ecological Economics, Vienna University of Economics and Business (WU), Vienna, Austria. 2 Industrial Ecol-
ogy Programme, Norwegian University of Science and Technology (NTNU), Trondheim, Norway. 3 Institute of Environ-
mental Sciences (CML), Leiden University, Leiden, The Netherlands. 4 Netherlands Organisation for Applied Scientific
Research (TNO), Delft, The Netherlands. Additional files Additional file 1: Full list of EXIOBASE products and product groups. Additional file 2: Full list of EXIOBASE countries and regions. Additional file 3: Technical details on the compilation of the global material flow database. Additional file 4: Correspondence table between raw materials and EXIOBASE sectors. Additional file 5: Detailed material footprint (MF) results. Acknowledgementsi We acknowledge financial support for this research from the European Commission through the FP7 project DESIRE
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https://www.frontiersin.org/articles/10.3389/fenrg.2022.1003473/pdf
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English
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Synthesis and evaluation of delayed anti-high temperature gel plugging agent
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Frontiers in energy research
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cc-by
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OPEN ACCESS OPEN ACCESS
EDITED BY
Xianbin Huang,
China University of Petroleum (East
China), China
REVIEWED BY
Hanyi Zhong,
China University of Petroleum,
Huadong, China
Hui Mao,
Chengdu University of Technology,
China
Yb He,
China University of Petroleum, China
*CORRESPONDENCE
Yang Bai,
baiyanghyq@foxmail.com
SPECIALTY SECTION
This article was submitted to Advanced
Clean Fuel Technologies,
a section of the journal
Frontiers in Energy Research
RECEIVED 26 July 2022
ACCEPTED 31 October 2022
PUBLISHED 14 November 2022
CITATION
Bai Y, Wu L, Luo P and Li D (2022),
Synthesis and evaluation of delayed
anti-high temperature gel
plugging agent. Front. Energy Res. 10:1003473. doi: 10.3389/fenrg.2022.1003473 Yang Bai*, Lingfeng Wu, Pingya Luo and Daoxiong Li State Key Laboratory of Oil and Gas Reservoir Geology and Exploitation, School of Oil and Natural Gas
Engineering, Southwest Petroleum University, Chengdu, China State Key Laboratory of Oil and Gas Reservoir Geology and Exploitation, School of Oil and Natural Gas
Engineering, Southwest Petroleum University, Chengdu, China The malignant leakage of complex formations seriously restricts the drilling time and
has become a major drilling technical problem that needs to be solved urgently at this
stage. In particular, the malignant leakage of large fractures and fractured formations is
difficultto construct,andthesuccessrateofpluggingis low.In thispaper,accordingto
the above formation conditions, byanalyzing the leakage mechanism of the malignant
leakage formation, the method of plugging the large fractures and broken fractured
formations is summarized. The adiabatic copolymerization co-hydrolysis method was
used as the gel synthesis method, 2-methacrylic acid amide and acrylamide (AM) were
used as the main raw materials, and the redox initiation system composed of sodium
sulfate and sodium bisulfite was used as the initiator. Using its redox initiation
characteristics, high-strength bonding between organic polymer covalent bonds
was achieved by a self-made cross-linking agent; using the thermoplasticity of
hot-melt adhesive, the initiator is wrapped in hot-melt adhesive particles to
achieve high-temperature slow-release initiation, and finally, a gel plugging agent
with good compatibility with drilling fluid, controllable gel formation time, stable gel
formation, and good gel breaking performance was synthesized, which is suitable for
severe lost circulation under high-temperature and high-pressure conditions. A
compatibility study of gel and common drilling fluid systems, the effect of
temperature on gel formation, gel rheological properties under high temperature
and high pressure, gel start-up pressure, and gel breakability tests were evaluated
under laboratory conditions. gel synthesis, delay, anti-high temperature gel, malignant leakage plugging, drilling
engineering TYPE Original Research
PUBLISHED 14 November 2022
DOI 10.3389/fenrg.2022.1003473 TYPE Original Research
PUBLISHED 14 November 2022
DOI 10.3389/fenrg.2022.1003473 TYPE Original Research
PUBLISHED 14 November 2022
DOI 10.3389/fenrg.2022.1003473 Introduction In view of the deficiency of the above gel plugging materials
that need the cooperation of a mud system and are not resistant
to high temperatures, a new gel plugging material, ZND-6, is
developed. The gel is compatible with the general water-based
drilling fluid, and it can be mixed with the drilling fluid on the
ground to maintain the relatively low viscosity. It may enter the
target formation with the drill pipe pump and utilize the
temperature of the formation within a specified time to form
a gel polymer with certain strength so as to block the formation. Its performance is superior to that of the general gel, and it has
features such as a controlled gelling time, high temperature
resistance of 150°C, good pressure bearing capacity, etc. It
solves the problem of the compatibility of the general loss-
stopping
agent
with
the
leakage
fracture
size
and
can
effectively
block
the
malignant
leakage
caused
by
large
fractures and broken formations. Loss of circulation is a phenomenon where various working
fluids (including drilling fluid, cement slurry, completion fluid,
and other fluids) leak into the formation under the action of
pressure differences during various downhole operations such as
drilling, cementing, and testing (Fan et al., 2018). The “vicious
leakage” mainly refers to the phenomenon where the mud
pumped into the wellbore has no circulation and no return
during the drilling process. It has the characteristics of high
leakage speed, huge leakage, difficult leakage processing, and easy
recurrence of leakage. Malignant leakage mainly occurs in karst
formations and fractured formations with large widths (Zhou
et al., 2012). Half of the widths of wide fractures are more than
millimeters, and even some fractures can reach about 10 mm in
width; the underground voids of karst strata are much larger than
wide fractures malignant leakage will seriously restrict the
drilling speed during the drilling process. In particular, the
malignant leakage that occurs in complex formations often
requires long-term, multiple times, multiple methods, and
multiple leakage plugging construction, which is inconsistent
with the drilling goal of “safety, high quality, high efficiency, and
economy” (Li and Xiang, 2020). The malignant leakage in
complex formations has become a major drilling technology
problem that has not been fully solved in the current stage. OPEN ACCESS The experimental results show that the gel has good
compatibility with common drilling fluid systems; with the increase of temperature,
the gel time of the gel plugging agent gradually decreases, and the gel time can be
adjusted from 1 to 3 h at a high temperature of 150°C; when the concentration of the
gel in the experimental slurry exceeds 2%, its rheological properties change abruptly;
under certain conditions, the starting pressure of the gel can reach up to 6.5 MPa; and
under the condition of 150°C, the gel breaking rate was over 98% after 24 h. Various
experiments show that the gel’s performance meets the requirements for plugging
malignant leakage formations, and the effect is good. Compared with traditional gel
materials, the gel plugging agent synthesized in this paper has the characteristics of a
simple configuration process and superior performance with drilling fluid, which
provides a new idea for malignant leakage plugging. COPYRIGHT
© 2022 Bai, Wu, Luo and Li. This is an
open-access article distributed under
the terms of the Creative Commons
Attribution License (CC BY). The use,
distribution or reproduction in other
forums is permitted, provided the
original author(s) and the copyright
owner(s) are credited and that the
original publication in this journal is
cited, in accordance with accepted
academic practice. No use, distribution
or reproduction is permitted which does
not comply with these terms. Frontiers in Energy Research 01 frontiersin.org 10.3389/fenrg.2022.1003473 Bai et al. Reagent and main instruments Acrylamide (AM), analytically pure (Elsharafiand Bai, 2017);
initiator (self-made); 2-methacrylic acid amide, analytically pure;
sodium hydroxide (NaOH), analytically pure; deionized water;
sodium persulfate (Na2S2O8), analytically pure (Zhao et al.,
2018); sodium bisulfite (NaHSO3), analytically pure; sodium
sulfate
(Na2SO4),
analytically
pure;
sodium
formate,
analytically pure; surfactant OP-10, analytically pure. FFD-50 adiabatic reactor; 85-1 magnetic stirrer; HHS-11-
6 electric water bath; GJD-B12K variable-frequency and high-
speed mixer; Granulator. With the continuous emergence of new materials and new
technologies in the petroleum industry, the gel plugging
materials
have
been
improved
and
upgraded
for
the
malignant leakage of the complex formation. Jianshan (2019)
takes the rigid aromatic ring as the main body of the structural
unit and develops the SMSHIELD intelligent gel plugging agent
with the carbon–nitrogen bond, carbon–carbon bond, and
other chemical bonds with high stability and outstanding
rigidity
as
the
main
chain. Such
gel
materials
are
characterized by strong deformation and easy formation, and
they could effectively bond the components in the mud cake
and fully fill all spatial positions of the mud cake to make it
denser. Such intelligent gel can be combined with other treating
agents, but they were not compatible with the drilling fluid of
other systems. Suming et al. (2007) have developed a pressure-
bearing delay plugging agent, OC L-GYDL. Such a plugging
agent has a strong pressure bearing capacity, and the gel time is
adjustable. However, it could not be consistent with high
temperature; therefore, it could not be applied to high-
temperature formations. Introduction The
study believes that for micro and narrow fractures, the use of
irregular rigid and flexible plugging materials has a better effect;
for large fractures and fractured formations, inorganic plugging
materials are difficult to construct, the plugging success rate is
low, it is difficult to significantly improve the pressure bearing
capacity of the formation, the cost of plugging is high, and it is
difficult to deal with the plugging of malignant leakage in
complex formations (Du et al., 2019). Synthesis The synthesis method of gel is the adiabatic copolymerization
co-hydrolysis method. This method does not require a separate
hydrolysis operation, which not only reduces the production cost
but also saves the production cycle. The steps of preparing gel by
adiabatic copolymerization co-hydrolysis are as follows. First, 2-methacrylic acid amide and acrylamide were added to
an adiabatic reactor according to a certain feed ratio. According
to the requirements of gel fluidity, the acrylate monomer was
added, deionized water and instant additives were added, and an
appropriate amount of alkali was added to adjust the pH value of
the system. After stirring,
a monomer
solution with
a
concentration of 35% was formed. Then the solution was
controlled to the polymerization temperature (40°C) (Kumar
et al., 2014), and an initiator was added after removing oxygen by
nitrogen. After adiabatic polymerization for 8 h (Barreira and
Negrão, 2017), the colloid was taken out for granulation, drying,
crushing, and packaging to obtain the gel sample. Frontiers in Energy Research 02 frontiersin.org Bai et al. 10.3389/fenrg.2022.1003473 FIGURE 1
Cementation state of the sample before and after the high-temperature aging. FIGURE 1
Cementation state of the sample before and after the high-temperature aging. other and have a strong structural force. We used a scanning
electron microscope to observe the microstructure of the samples
after aging, as shown in Figure 2. It was found that the gel
particles were completely gelled under the conditions of 10 h and
high temperature, and a dense three-dimensional network
structure is formed between the gel molecules. The reticular
formation of the gel molecular closely combines with other gel
molecules that are independently distributed to form the large gel
mass so that the sample has strong structural strength (Wang
et al., 2020). The initiator is a redox initiation system composed of sodium
persulfate and sodium bisulfite, and its addition amount is 2% of
the total weight of the monomer solution (Quiroz et al., 2022). The instant additive is composed of sodium sulfate, sodium
formate, and surfactant OP-10, and the dosage is 3% of the
total weight of the monomer solution (Liu et al., 2018). Compatibility study of gel and common
drilling fluid systems KPS, analytical reagent (Gregolin et al., 2010); HV-CMC,
analytical reagent; SPNH, analytical reagent; the formula for the
experimental pulp is as follows: H2O + bentonite + barytes, +1%HV-CMC+3%SPNH, the
experimental mud density is 1.5. In order to explore the effect of gel ZND-6 mixed into
drilling fluid on the performance of drilling fluid, the
compatibility experiment of gel with NaCl, polysulfonic
acid, and polymer drilling fluid systems was carried out in
the laboratory. The drilling fluid before and after aging after
adding gel was mainly investigated for changes in rheology
and dehydration wall-building. The experimental results are
shown in Tables 1, 2, 3. M7500 high-temperature and high-pressure rheometer; QD-
2 test device on plugging materials; XGRL-4 aging furnace. Frontiers in Energy Research Frontiers in Energy Research Microstructure characterization According to the experimental pulp formula, we add 3%
gelatin ZND-6 to the configured experimental pulp, stir for
10 min at low speed, and then put it into the aging tank. After aging for 10 h at 150°C, gel particles are gelatinized. The
sample status before and after the experiment is shown in
Figure 1. It could be found that the sample has a strong
thixotropy after adding the gel particles. Before aging, the gel
particles in the sample are distributed independently in the
experimental slurry, and there is no cementation between
them. After aging, the gel particles agglomerate with each It can be seen from Table 1 that adding the gel ZND-6 to
the NaCl drilling fluid can enhance the static shear force of the
system, reduce the filtration loss, and enhance its water loss
and wall-building ability without affecting the fluidity of the
system. Gel ZND-6 has good compatibility with NaCl drilling
fluid. It can be seen from Table 2 that the effect of gel ZND-6 on
the system is generally favorable after adding the polysulfonic
drilling fluid, which can improve the rheological properties of the
system and improve the ability of water loss and wall-building. Frontiers in Energy Research 03 frontiersin.org Bai et al. 10.3389/fenrg.2022.1003473 FIGURE 2
Micromorphology of the gel after aging. FIGURE 2
Micromorphology of the gel after aging. Evaluation of the gel’s rheological
property To determine the rheological data of the gel ZND-6 under the
downhole high-temperature and high-pressure conditions, the
rheological property of the gel ZND-6 is tested for 10H in a high-
temperature
and
high-pressure
environment
with
the
M7500
high-temperature
and
high-pressure
rheometer
produced by Grace Company. The time length includes the
time before and after the gelation of gel ZND-6 and the
period after gelation so as to determine the viscosity change
before gelation and the stability after gelation. can improve the water loss and wall-building properties as well as
the viscosity and static shear force of the drilling fluid. The
compatibility between the gel and the commonly used drilling
fluid system is sufficient to meet the requirements of solving the
malignant loss of large fractures and broken formations. Add the gel ZND-6 with a concentration of 1.5% to the
experimental pulp and increase the temperature to 150°C. Then,
test the sample viscosity every 20 min under the environment at a
pressure of 20 MPa, with the experimental results shown in
Figure 3. The samples are gelatinized at 60 min, and the
highest viscosity reaches 19000 mPa s. Besides, the sample
viscosity
basically
remains
stable
in
the
remaining
experiment time. resulting in a faster release of the initiator from the hot-melt
adhesive and a shortening of the gel formation time (Li, X
et al., 2022). resulting in a faster release of the initiator from the hot-melt
adhesive and a shortening of the gel formation time (Li, X
et al., 2022). TABLE 4 Impact of temperature in clear water on gelling time. Temperature/°C
80
100
120
150
tgelation/h
4.2
3.1
1.5
1.1
TABLE 5 Impact of temperature in experimental pulp on gelling time. Temperature/°C
80
100
120
150
tgelation/h
6.8
5.6
4.1
3.3 Frontiers in Energy Research TABLE 1 Compatibility of ZND and NaCl drilling fluids (150°C for 16 h aging). ZND-6/%
Experimental conditions
AV/mPa·s
PV/mPa·s
YP/Pa
Gel 10 min/Pa
Fl/ml
Filter cake thickness/mm
0
Before ageing
30.0
25
5
5.1
8.5
0.5
1.0
Before ageing
82.5
87
6
12.7
6.1
0.5
2.0
Before ageing
142.2
156
4
75.3
5.3
0.5
3.0
Before ageing
204.8
265
6
164.4
3.7
0.5
0
After ageing
15.0
15
0
2.0
19.0
0.5
1.0
After ageing
44.1
51
4
6.4
14.6
0.5
2.0
After ageing
86.2
97
0
34.8
8.2
0.5
3.0
After ageing
102.2
124
12
99.2
7.7
0.5 ZND-6/%
Experimental conditions
AV/mPa·s
PV/mPa·s
YP/Pa
Gel 10 min/Pa
Fl/ml
Filter c nditions
AV/mPa·s
PV/mPa·s
YP/Pa
Gel 10 min/Pa
Fl/ml
Filter cake thickness/mm TABLE 2 Compatibility of ZND and Polysulfonic Drilling Fluid (150°C for 16 h of aging). TABLE 2 Compatibility of ZND and Polysulfonic Drilling Fluid (150 C for 16 h of aging). ZND-6/%
Experimental conditions
AV/mPa·s
PV/mPa·s
YP/Pa
Gel10 min/Pa
FL/mL
Filter cake thickness/mm
0
Before ageing
34.0
31
3
6.1
10.0
1.0
1.0
Before ageing
66.2
66
6
15.3
8.2
1.0
2.0
Before ageing
131.9
124
4
41.3
6.5
0.8
3.0
Before ageing
184.7
156
0
115.8
4.8
0.5
0
After ageing
31.2
21
3
3.4
12.1
0.5
1.0
After ageing
37.1
24
4
7.6
7.5
0.5
2.0
After ageing
64.1
62
4
22.9
8.8
0.5
3.0
After ageing
122.7
98
3
59.7
6.8
0.5 Gel ZND-6 has good compatibility with polysulfonic drilling
fluid. fluid loss of the drilling fluid is low. The performance after aging
also meets the general requirements for drilling fluids and has
good compatibility with polymer drilling fluids. It can be seen from Table 3 that ZND-6 gel can improve the
fluidity of the drilling fluid after adding the polymer drilling fluid,
while maintaining the static shear force of the system, and the To sum up, the gel ZND-6 has good compatibility with
drilling fluids such as NaCl, polysulfonic acid, and polymers and Frontiers in Energy Research Frontiers in Energy Research 04 frontiersin.org 10.3389/fenrg.2022.1003473 Bai et al. TABLE 3 Compatibility of ZND and Polymer Drilling Fluid (150°C for 16 h aging). nditions
AV/mPa·s
PV/mPa·s
YP/Pa
Gel10 min/Pa
FL/mL
Filter cake thickness/mm TABLE 4 Impact of temperature in clear water on gelling time. Influence of temperature on the gelling
time The effect of temperature on the gel formation time is
obvious: the half-life of the gel polymer initiator decreases
exponentially with the increase in temperature. The gelling
times of gelatin in water and the experimental slurry at
80°C–150°C are studied, respectively, to determine the relevant
time. The experimental results are shown in Tables 4, 5. According to Figures 4, 5, the rheological properties of the
samples only increase linearly with the gel concentration when
the gel concentration is 0.5%, 1.0%, or 1.5%. After the gel
concentration reaches 2%, all rheological data of the sample
show sudden changes. When the gel concentration is relatively
low, the gel particles in the sample are far away; therefore, it is
not easy for them to contact each other and form a three-
dimensional network structure between the particles. Actually,
they just change the rheological property of the samples
through the swelling and gelation of each individual gel
particle. After the gel concentration reaches or exceeds 2%,
not only the individual gel particle has an influence on the
rheological property of the samples. Besides, with the increase It can be seen from the experimental results that the gel-
forming time of the gel ZND-6 gradually decreases with an
increase in the experimental temperature. The reasons and
results
may
be
that
①
the
half-life
of
the
initiator
decreases with the increase of temperature, which shortens
the gel formation time (Li et al., 2015) and ②with the
increase
of
the
experimental
temperature,
the
melting
rate of the hot-melt adhesive capsule in the gel increases, Frontiers in Energy Research 05 frontiersin.org Bai et al. 10.3389/fenrg.2022.1003473 FIGURE 3
Viscosity property change of the samples within 10 h. FIGURE 4
Comparison of shearing stress at different rotating speeds under different concentrations of gel. FIGURE 3
Viscosity property change of the samples within 10 h. FIGURE 3
Viscosity property change of the samples within 10 h. FIGURE 4
Comparison of shearing stress at different rotating speeds under different concentrations of gel. FIGURE 4
Comparison of shearing stress at different rotating speeds under different concentrations of gel. property of the slurry will change suddenly when the
concentration of the gel ZND-6 exceeds 2%. Hence, the
optimal dosage concentration of gel ZND-6 should not be
less than 2%. Influence of temperature on the gelling
time of gel particles per unit volume, the distances between the gel
particles are reduced so as to form a network structure among
the particles and create a larger gel mass. Such large gel masses
could obviously affect the rheological properties of samples
more than single gel particles. It could be judged from the above experiments that
the rheological property of gel could basically maintain
stability under the downhole high-temperature and high-
pressure
conditions,
and
it
could
block
the
broken
formation stably for a long time, which could prolong the
safe operation time effectively and strengthen the safety of
well control (Li, Z et al., 2022). Besides, the rheological Frontiers in Energy Research Starting pressure test of gel After reaching the position of lost circulation and
penetrating into large fractures and fractured formations,
the gel swells into glue in the well slurry to block large
pores and fractures. After the gel plugging, there is a Frontiers in Energy Research 06 frontiersin.org frontiersin.org Bai et al. 10.3389/fenrg.2022.1003473 FIGURE 5
Comparison of viscosity at different rotating speeds under different concentrations of gel. TABLE 6 ZND-6 start-up experiment. No. Sample dosage/%
Measuring temperature/°C
Holding time/h
Start-up pressure/MPa
1
3
80
5
1.5
2
3
100
5
2.0
3
3
120
5
2.0
4
3
150
5
4.0
5
5
150
5
6.5
6
5
170
5
4.0
6
5
180
5
3.5 FIGURE 5
Comparison of viscosity at different rotating speeds under different concentrations of gel. Comparison of viscosity at different rotating speeds under different concentrations of gel. TABLE 6 ZND-6 start-up experiment. No. Sample dosage/%
Measuring temperature/°C
Holding time/h
Start-up pressure/MPa
1
3
80
5
1.5
2
3
100
5
2.0
3
3
120
5
2.0
4
3
150
5
4.0
5
5
150
5
6.5
6
5
170
5
4.0
6
5
180
5
3.5 TABLE 6 ZND-6 start-up experiment. No. Sample dosage/%
Measuring temperature/°C
Holding time/h
Start-up pressure/MPa
1
3
80
5
1.5
2
3
100
5
2.0
3
3
120
5
2.0
4
3
150
5
4.0
5
5
150
5
6.5
6
5
170
5
4.0
6
5
180
5
3.5 pressure with 0.5 MPa as the gradient to observe whether there is
a sample extrusion. After the samples are extruded, stop the
experiment and record the pressure, which is the start-up
pressure of the gel. The start-up pressures vary at different
temperatures,
gel
concentrations,
and
high-preservation
standing times. The experimental results are as shown in Table 6. pressure difference inside and outside the wellbore. After the
gelatinization, the pressure difference that the plugging
material can withstand is the start-up pressure of the gel,
and it is a key metric to evaluate its plugging performance
(Yokoi et al., 2022). The QD-2 test device on plugging materials is adopted to
simulate leakage layer cracks with different seam widths (1, 2, 3,
4, and 5 mm). We add the gel to the experimental pulp and
investigate the plugging pressure-bearing capacity of the gel
plugging materials on the large pores and fractures causing
the malignant leakage under the high-temperature and high-
pressure conditions. Frontiers in Energy Research Well Shuangmiao 1 Well
Shuangmiao
1
in
Dazhou
is
a
key
vertical
exploration well in southwest China. There are six leakage
points in the 2083–3,573 m well section. A high-pressure gas
layer was drilled at a depth of 3,446–3,448 m, and the bottom
hole overflowed. In order to seal the leakage layer and protect
the gas layer, the on-site technicians have successively
adopted
bridge
plug
plugging,
nine
times
of
cement
plugging, and bridge plug and cement composite plugging
methods, all of which have not been effective, taking up to
2 months, and the phenomenon of simultaneous leakage and
leakage has not been controlled. Wellhead two single ram
blowout preventers in the upper ram damage failure, four
anti-nozzle lines in three damaged, throttling blowout, and
casing pressure are extremely difficult to reduce. The leakage
layer of 3,446–3,448 m is the main factor to induce complex
downhole conditions. air blockage but also cements with the tank externally. With the
rise in temperature, the strength of the network structure will
increase. Once it exceeds the limiting temperature of gel, the
strength of the structure will decrease. Therefore, the start-up
pressure of gel will increase with the rise in temperature in the
range from 80°C to 150°C. After it exceeds 150°C, the start-up
pressure will decrease with the rise in temperature. Similarly, the
strength of the gel’s network structure will be enhanced with the
concentration of gel in the experimental pulp. Starting pressure test of gel Data in Table 6 indicates that under the same concentration,
the start-up pressure increases with the rise in temperature, and
the maximum start-up pressure is 6.5 MPa in the temperature
range of 80°C–150°C; when the temperature exceeds 150°C, the
sample’s start-up pressure decreases with the rise in temperature. Under the same temperature, the ZND-6 start-up pressure
increases as the gel concentration increases. At 150°C and a
concentration of 5%, the gel will form a tight and elastic air
blockage, as shown in Figure 6. The specific experimental method is as follows: add the gel
ZND-6 to the experimental pulp and put it into the plugging
material test device after mixing evenly, pressurize the upper
valve rod by 0.5 MPa, close the lower valve rod, and let it stand
for several hours at a certain temperature. After the gel in the
samples seals the joint board, open the foot valve and increase the This phenomenon occurs mainly because the hydrophobic
group in gel ZND-6 will form a network structure in the slurry. This network structure not only makes the gel combine into an Frontiers in Energy Research 07 frontiersin.org 10.3389/fenrg.2022.1003473 Bai et al. 10.3389/fenrg.2022.1003473 D is the gel breaking rate; m is the gel quality before gel
breaking, g; and m0 is the remaining gel quality after gel
breaking, g. FIGURE 6
Air Blockage formed by examples. Add 3% gel to the experimental pulp, and then add 1% KPS. After mixing evenly, pour it in the aging tank and take out the
sample after rolling it in the roller furnace for a certain time at
150°C. Then, screen out the remaining gel with the screen and
weigh it to calculate the gel breaking rate. The experimental
results are as shown in Figure 7. It can be seen from Figure 7 that the gel breaking rate could
reach about 50% after 18 h at 150°C; after 24 h, the gel breaking
rate could reach 98%. At this time, the gel breaking of gel is
basically complete, meeting the requirements of the reverse
drainage of the general reservoir. Author contributions 2. The gel plugging materials ZND-6 could penetrate the broken
formation to form the air blockage. When the gel concentration
is 1.5% at 150°C, the sample viscosity reaches 19,000 mPa s and
the stable time is greater than 10 h. YB: ideas and suggestions; LW: data collection and
drafting of papers; PL: approval of the final version of the
paper for publication; and DL: made important revisions to
the paper. 3. It could be found from the experiment that after the concentration
of gel reaches 2% in the slurry, it is easier to form the three-
dimensional network structure and become the larger gel mass,
and the sample performance may change suddenly. In view of the
actual construction and economic benefits, it is suggested to
control the gel concentration at 3%–5%. Gel breaking test In order to effectively plug the three leakage layers at a
depth of 3,483, 3,446, and 3,436 m, the special gel plugging
technology is tried. A total of 70 m3 of gel was injected into
the three layers, and then 25 m3 of cement was added. The
construction was successful. After pressure relief, the casing
pressure drops to 12 MPa and then shuts down. The
standpipe
pressure
is
zero,
and
the
casing
pressure
remains unchanged. Shut-in observation riser pressure is
still zero, the casing pressure is still zero, to achieve the
purpose of blocking the annulus, cut off the bottom hole
high-pressure gas layer. The logging results show that the
cement sheath is in the well depth of 2,250–3,200 m, the
cement return height is in the well depth of 2,355 m, the
ZND-6 gel column in the annulus of the leakage layer is
nearly 100 m, and the plugging is successful. The core material of the oxidation breaker is potassium
persulfate or ammonium persulfate. The principle is that the
breaker can be dissolved into a highly active, solid strong oxidant
in contact with water (Guo et al., 2016). The advantage of
oxidative gel breakers is that they reduce the effect of gel
breakers on the rheological properties of fracturing fluid. After the reservoir leakage plugging, it is necessary to reverse
the gel breaking in the lost circulation plugging slurry to ensure
the downhole production capacity. Hence, the gel-breaking test
shall be carried out. The calculation formula of the gel breaking
rate is: D m −m0
m
× 100%. m Frontiers in Energy Research 08 frontiersin.org Bai et al. 10.3389/fenrg.2022.1003473 FIGURE 7
Change of gel breaking rate of samples with time. FIGURE 7
Change of gel breaking rate of samples with time. Well Liu 67–72 5. Under the condition of 150°C, the gel breaking rate is over
98% after 24 h, which meets the requirement of general
reservoir reverse flow. 5. Under the condition of 150°C, the gel breaking rate is over
98% after 24 h, which meets the requirement of general
reservoir reverse flow. 5. Under the condition of 150°C, the gel breaking rate is over
98% after 24 h, which meets the requirement of general
reservoir reverse flow. Well Liu 67–72 in Changqing leakage occurred at a depth of
1,629 m. The bridge slurry (the particle size of the bridging material is
3 ~ 5 mm) and the cement plug failed nine times. In order to plug the
leakage point, ZND-6 gel plugging technology is adopted. Drill down
the drill pipe a depth of 1,500 m, pump 34 m3 concentration of 1.2%
gel ZND-6, and then inject 12 m3 water; next, pull out the drill, and
allow it to stand for 6 h. Down to the bottom of the well, open the
pump return, normal displacement. After circulating the drilling fluid
for 15 min, drilling 8 m at 150 kN and 65 r/min, the pump pressure
displacement is normal and the plugging is successful. 6. According to field application, the innovation of gel
plugging agent ZND-6 is that it can be used alone with
drilling fluid and has a good effect on plugging malignant
leakage layers. It can well solve the leakage problems that
cannot be solved by methods such as fractured, porous, and
broken formations; bridge plugging; and plugging while
drilling. It is unique in dealing with the leakage problems of
aquifers and the same layer of leakage. 6. According to field application, the innovation of gel
plugging agent ZND-6 is that it can be used alone with
drilling fluid and has a good effect on plugging malignant
leakage layers. It can well solve the leakage problems that
cannot be solved by methods such as fractured, porous, and
broken formations; bridge plugging; and plugging while
drilling. It is unique in dealing with the leakage problems of
aquifers and the same layer of leakage. Conclusion The original contributions presented in the study are
included
in
the
article/supplementary
material;
further
inquiries can be directed to the corresponding author. 1. The
high-temperature
gel
plugging
agent
ZND-6
is
synthesized with a controlled gelling time, and the free
radical is aggregated to form the high-strength gel with
dense cross-linking. Conflict of interest The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could
be construed as a potential conflict of interest. 4. The temperature resistance of the gel ZND-6 could reach
150°C. When the temperature exceeds 150°C, the gel strength
would decrease. Frontiers in Energy Research 09 frontiersin.org frontiersin.org 10.3389/fenrg.2022.1003473 10.3389/fenrg.2022.1003473 Bai et al. References Li, Z., and Xiang, C. (2020). Experimental investigation of a new weak-gel-
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Spleen Spleen Spleen A
E
H
N
O
F
I
J
K
G
P
B
C
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
25
Total Splenocyte Viable Cells (x 10*7)
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = 0.9108
p = 0.0002
Total Splenocyte Viable Cells (x 10*7)
CD3+
CD4+
CD8+
Treg
CD19+
CD11b+
NK
Macro
Neut
DC
0
20
40
60
80
ns
ns
ns
ns
Percent of Live CD45+(%)
Veh-D9
Veh-D21
Anlo-D9
Anlo-D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
100
GzmB+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
IFN- +CD4+/CD4+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
IFN- +CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD8+/CD8+(%)
ns
0
2
4
6
8
0
10
20
30
40
50
r = 0.9690
p<0.0001
CD11b+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
r = -0.6849
p = 0.0289
CD8+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
20
25
r = -0.7522
p = 0.0121
CD4+CD45+/CD45+(%)
Spleen
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
PD1+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
PD1+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
Ki67+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
Ki67+CD8+/CD8+(%)
ns
CD3+
CD4+
CD8+
Treg
CD19+
CD11b+
NK
Macro
Neut
DC
0
10
20
30
40
50
ns
ns
Percent of Live CD45+(%)
ns
ns
D
L
M
ns
ns
ns
ns
ns
ns
ns
Blood
Spleen A
B
C
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
25
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = 0.9108
p = 0.0002
Total Splenocyte Viable Cells (x 10*7)
CD3+
CD4+
CD8+
Treg
CD19+
CD11b+
NK
Macro
Neut
DC
0
20
40
60
80
ns
ns
ns
ns
Percent of Live CD45+(%)
Veh-D9
Veh-D21
Anlo-D9
Anlo-D21
ns
Spleen B
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = 0.9108
p = 0.0002
Total Splenocyte Viable Cells (x 10*7) A
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
25
Total Splenocyte Viable Cells (x 10*7)
ns C A B Total Splenocyte Viable Cells (x 10*7) E
H
N
Q
O
R
F
I
J
K
G
P
S
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
Total Splenocyt
0
2
4
6
8
0
5
Tumor weight(g)
p
0 000
Total Splenocyt
CD3+
CD4+
CD8+
Treg
CD19+
CD11b+
NK
Macro
Neut
DC
0
20
ns
ns
ns
Percent of
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
100
GzmB+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
IFN- +CD4+/CD4+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
IFN- +CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD8+/CD8+(%)
ns
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
r = 0.9690
p<0.0001
CD11b+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.6849
p = 0.0289
CD8+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = -0.7522
p = 0.0121
CD4+CD45+/CD45+(%)
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
CD11b+CD45+/CD45+(%)
r = 0.8857
p = 0.0006
0
2
4
6
8
0
2
4
6
8
10
Tumor weight(g)
r = -0.7536
p = 0.0118
CD8+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.8712
p = 0.0010
CD4+CD45+/CD45(%)
Tumor
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
PD1+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
PD1+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
Ki67+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
Ki67+CD8+/CD8+(%)
ns
CD3+
CD4+
CD8+
Treg
CD19+
CD11b+
NK
Macro
Neut
DC
0
10
20
30
40
50
ns
ns
Percent of Live CD45+(%)
ns
ns
D
L
M
ns
ns
ns
ns
ns
ns
Blood E
F
G
Tumor weight(g)
C
C
C
C
CD
M
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
100
GzmB+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
IFN- +CD4+/CD4+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD4+/CD4+(%)
ns
CD3+
CD4+
CD8+
Treg
CD19+
D11b+
NK
Macro
Neut
DC
0
10
20
30
40
50
ns
ns
Percent of Live CD45+(%)
ns
ns
D
ns
ns
Blood G
M
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
100
GzmB+CD8+/CD8+(%)
ns 5
D G
100 G
100 Percent of Live CD45+(%)
D H
N
Q
O
R
I
J
K
P
S
D9 D21 D9 D21
D9 D21 D9 D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
IFN- +CD8+/CD8+(%)
D9 D21 D9 D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD8+/CD8+(%)
ns
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
r = 0.9690
p<0.0001
CD11b+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.6849
p = 0.0289
CD8+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = -0.7522
p = 0.0121
CD4+CD45+/CD45+(%)
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
CD11b+CD45+/CD45+(%)
r = 0.8857
p = 0.0006
0
2
4
6
8
0
2
4
6
8
10
Tumor weight(g)
r = -0.7536
p = 0.0118
CD8+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.8712
p = 0.0010
CD4+CD45+/CD45(%)
Tumor
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
PD1+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
PD1+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
Ki67+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
Ki67+CD8+/CD8+(%)
ns
CD
CD
CD
Tr
CD1
CD11
Mac
Ne
L
M
ns
ns
ns
ns H
I
J
K
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
IFN- +CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
20
40
60
80
TNF- +CD8+/CD8+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
PD1+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
PD1+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
5
10
15
20
Ki67+CD4+/CD4+(%)
ns
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
Ki67+CD8+/CD8+(%)
ns
C
C
C
CD
CD
M
L
M
ns
ns
ns
ns Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
PD1+CD8+/CD8+(%)
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
Ki67+CD8+/CD8+(%)
ns
L
M
ns
ns H
N
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
10
20
30
40
50
IFN- +CD8+/CD8+(%)
C
C
C
ns I
8 H
50 J
20 Ki67+CD8+/CD8+(%) N
O
P
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
r = 0.9690
p<0.0001
CD11b+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.6849
p = 0.0289
CD8+CD45+/CD45+(%)
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = -0.7522
p = 0.0121
CD4+CD45+/CD45+(%)
Tumor
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21
Veh
D9
Veh
D21
Anlo
D9
Anlo
D21 N
D9 D21 D9 D21
D9 D21 D9 D21
0
2
4
6
8
0
5
10
15
20
25
Tumor weight(g)
r = -0.7522
p = 0.0121
CD4+CD45+/CD45+(%) N O O
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.6849
p = 0.0289
CD8+CD45+/CD45+(%)
D9 D21 D9 D21
9
9 P
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
r = 0.9690
p<0.0001
CD11b+CD45+/CD45+(%) P
CD11b+CD45+/CD45+(%) CD4+CD45+/CD45+(%) CD8+CD45+/CD45+(%) Tumor weight(g) Q
0
2
4
6
8
0
5
10
15
Tumor weight(g)
r = -0.8712
p = 0.0010
CD4+CD45+/CD45(%) R
0
2
4
6
8
0
2
4
6
8
10
Tumor weight(g)
r = -0.7536
p = 0.0118
CD8+CD45+/CD45+(%) S
0
2
4
6
8
0
10
20
30
40
50
Tumor weight(g)
CD11b+CD45+/CD45+(%)
r = 0.8857
p = 0.0006 Q
1 R S
CD11b+CD45+/CD45+(%) R
CD8+CD45+/CD45+(%) CD4+CD45+/CD45(%)
|
https://openalex.org/W2025131735
|
http://medpet.journal.ipb.ac.id/index.php/mediapeternakan/article/download/8795/6870
|
English
| null |
Impacts of Domestic Maize Price Changes on the Performance of Small-scale Broiler Farming in Indonesia
|
Media Peternakan
| 2,014
|
cc-by-sa
| 7,226
|
ISSN 0126-0472 EISSN 2087-4634
Accredited by DGHE No: 66b/DIKTI/Kep/2011 ISSN 0126-0472 EISSN 2087-4634
Accredited by DGHE No: 66b/DIKTI/Kep/2011 ISSN 0126-0472 EISSN 2087-4634
Accredited by DGHE No: 66b/DIKTI/Kep/2011 Media Peternakan, December 2014, 37(3):198-205
DOI: 10.5398/medpet.2014.37.3.198
Available online at http://medpet.journal.ipb.ac.id/ Media Peternakan, December 2014, 37(3):198-205
DOI: 10.5398/medpet.2014.37.3.198
Available online at http://medpet.journal.ipb.ac.id/ Impacts of Domestic Maize Price Changes on the Performance of Small-scale
Broiler Farming in Indonesia S. J. K. Umboha,b,*, D. B. Hakimc, B. M. Sinagac, & I K. Kariyasad
aStudy Program of Agricultural Economics, Faculty of Economics and Management, Graduate School,
Bogor Agricultural University
bFaculty of Animal Husbandry, Sam Ratulangi University
Jln. Kampus Unsrat, Bahu Manado-North Sulawesi 95115, Indonesia
cFaculty of Economics and Management, Bogor Agricultural University
Jln. Kamper, Kampus IPB Darmaga Bogor-West Java 16680, Indonesia
dIndonesian Center for Food Crops Research and Development
Jln. Merdeka No 147 Bogor-West Java 16111, Indonesia
(Received 09-06-2014; Reviewed 01-09-2014; Accepted 03-11-2014) S. J. K. Umboha,b,*, D. B. Hakimc, B. M. Sinagac, & I K. Kariyasad
aStudy Program of Agricultural Economics, Faculty of Economics and Management, Graduate School,
Bogor Agricultural University
bFaculty of Animal Husbandry, Sam Ratulangi University
Jln. Kampus Unsrat, Bahu Manado-North Sulawesi 95115, Indonesia
cFaculty of Economics and Management, Bogor Agricultural University
Jln. Kamper, Kampus IPB Darmaga Bogor-West Java 16680, Indonesia
dIndonesian Center for Food Crops Research and Development
Jln. Merdeka No 147 Bogor-West Java 16111, Indonesia
(Received 09-06-2014; Reviewed 01-09-2014; Accepted 03-11-2014) ABSTRACT This research aimed at analyzing the impact of maize price changes on the performance of
small-scale broiler farming in Indonesia using a multimarket model analysis. The multimarket
model analysis is partial equilibrium analysis that contains six blocks of equations: prices, supply,
input demand, consumption, income and equilibrium. This model analysis was originally designed
in General Algebric Modelling System (GAMS) using the Path NLP solver. Employed data in this
study were classified into 3 types, namely: (1) production and input, consumption, and household
income; (2) inputs and outputs, and (3) elasticities. Decreased domestic maize price was responded
by farmer through reducing maize planted area and fertilizers uses. It further had undesired impact
on the maize production and maize farmer’s income. Whereas, this policy had positive impact on
meat production and small-scale broiler farming income. The opposite impact will happen on those
variables, if government increases domestic maize price. This policy caused the maize demand for
feed industry decreased. As a result of this condition, it decreased the chicken meat production and
small-scale broiler farming income. Key words: maize, broiler, price, multimarket *Corresponding author:
E-mail: sintyajkumboh@yahoo.co.id Kata kunci: jagung, ayam ras pedaging, price, multimarket Key words: maize, broiler, price, multimarket INTRODUCTION This research excluded feed market into model,
therefore feed demand was approximated by a proxy
from maize demand. This proxy was reliable as maize
proportion in feed ingridient was constant about 66%. During 2000-2010, the feed price of broiler and maize
price tended to increase (Table 1). The fluctuation of
maize price was being concern of feed manufacturer
to decide production capacity. This finding was in line
with Kariyasa & Sinaga (2007) that concluded that feed
manufacturer concerned more to maize price than g p
p
y
g
The main components of chicken feed are maize,
soybean meal, and fish flour. The composition of maize
in the feed ranges from 55%-65%. In 2010, maize demand
for broiler feed was 1.11 million tons and 1.65 million
tons for layer. The average increase in maize demand
for broiler and layer feeds during the period 2000-2010
increased by 6.85% and 8.31%, respectively (Swastika et
al., 2011; Directorate General of Livestock and Animal
Health, 2012, processed). Some factors caused the use
of maize for feed is high, among others are its prices is
relatively cheap, easy to produce, as well as palatable
to poultry. In addition, maize contains high calorie
and protein with complete amino acids are also as
consideration or reasons why the use of it is still higher
compared to others (Swastika et al., 2011). Attempts have
been done to replace maize with other grains are not
succesfully. Therefore maize remains as the main raw
material for livestock feed. INTRODUCTION therefore could affect the sustainability of small-scale
broiler farming. Broiler farming in Indonesia has become an indus
try with complete components from upstream to down
stream. The contribution of Gross Domestic Product
(GDP) of livestock sub-sector to the agriculture sector
GDP based on market prices reached 12%. Of this fig
ure, poultry meat contribution was 64%, and of the total
poultry meat contribution, it is about 70.24 percent com
ing from chicken meat (Saptana & Sumaryanto, 2009;
Directorate General of Livestock and Animal Health,
2012). This figure has shown that the chicken meat has
an important role in providing meat for Indonesian
(Ilham, 2009). It also explained that chicken meat has
been the main driver on the development of the poultry
business in Indonesia. Related to feed, maize, and chicken meat price data,
the increasing of average feed price have the same trend
with increasing of average maize price of by 6.78% and
7.96%, respectively, while chicken meat price incereased
around 5.90% (Pusdatin, 2012).fi Viewed from efficiency as a result of business
integration, breeding farms and feed mills which belong
to foreign investors sell DOC and feed to independent
farmers in high prices, making their business inefficient. Indarsih et al. (2010) revealed that there was a disparity
in feed prices to be paid by large-scale and small-scale
broiler farming, which was around 30%-50%. No wonder
the production costs of small-scale broiler farming are
much higher that those of broiler farming large-scale. If
all chicken meat production goes to traditional markets,
independent farmers will lose money because they can
not keep up with the production cost of large-scale. In cost structure side, feed placed the biggest por
tion in broiler and egg production cost. It was account
ing to 70% of the total production cost, while other costs
such as DOC (Day Old Chick) was only about 13 percent
(Yusdja & Pasandaran, 1998). Feed demand for broilers
and layers during the period 2000-2010 increased by
6.85% and 8.31%, respectively (Swastika et al., 2011). In
2010, its demand for broiler reached 3.51 million tons
and 2.06 million tons for layers (Directorate General of
Livestock and Animal Health, 2012). This indicated that
feed available plays a significant role in broilers and
eggs production. Therefore, if there is shock happening
in feed prices, it brings a great influence on broiler farm
ing performance, especially small-scale broiler farming. ABSTRAK Penelitian ini bertujuan untuk menganalisis dampak perubahan harga jagung domestik
terhadap kinerja usaha peternakan rakyat ayam ras pedaging di Indonesia dengan menggunakan
analisis model multimarket. Analisis model multimarket merupakan model keseimbangan parsial
yang terdiri atas enam blok persamaan: harga, penawaran, permintaan input, konsumsi, pendapatan,
dan kondisi keseimbangan. Model ini secara original dibangun dalam software General Algebric
Modelling System (GAMS) dengan metode solusi Path NLP. Data terdiri atas: (1) data produksi
dan input, konsumsi, dan pendapatan rumahtangga, (2) harga input dan output, dan (3) elastisitas. Untuk data produksi, penggunaan input, konsumsi, pendapatan dan harga menggunakan data dari
Badan Pusat Statistik dan Kementerian Pertanian, sedangkan untuk elastisitas menggunakan data
hasil penelitian sebelumnya. Penurunan harga jagung domestik direspon petani dengan mengurangi
luas pertanaman jagung dan penggunaan input pupuk sehingga menyebabkan turunnya produksi
dan pendapatan dari usahatani jagung. Sebaliknya kebijakan ini berdampak pada meningkatnya
permintaan jagung untuk pakan sehingga meningkatkan produksi daging ayam ras dan pendapatan
peternak rakyat ayam ras pedaging. Kondisi berbeda jika terjadi peningkatan harga jagung domestik. Kebijakan ini menyebabkan permintaan jagung untuk pakan menurun, akibatnya produksi daging
ayam ras dan pendapatan peternak rakyat ayam ras pedaging menurun. Kata kunci: jagung, ayam ras pedaging, price, multimarket 198 December 2014 UMBOH ET AL. / Media Peternakan 37(3):198-205 Source: BPS (2012), Pusdatin Ministry of Agriculture (2012), processed INTRODUCTION 231
232
233 Price block. The producer prices (PP) are lower than the
consumer prices (PC) due to the presence of the domestic
marketing margin (MARG), hence:
) (
g
y,
)
PCc,h = PMc * (1+RMARG) Multimarket model is a partial equilibrium model
was used to analyze the impact of changes in price and
quantity in a particular market on household income
and expenditures. Multimarket model analysis was used
to measure the impact of policy changes on the poor
households (Dorosh et al., 1995; Minot & Goletti, 1998),
and it was focused on analyzing the impact of food im
ports on the poverty level of poverty in Mozambique. In
Indonesia, Hutabarat et al. (2012) investigate the impact
of climate changes on the production, imports, and con
sumption of horticulture. 234
235
236
237
238 PCc,h= PPc,h * (1 + MARG) ...................................(1)
arge scale(PRSHN) ofbroiler consumer price diff .(1)
e diff where the subscripts c and h refer to commodity and
household type, respectively. The border prices (PM)
for tradable products are linked to the world price
by the exchange rate (er), import tariffs (tm) and the
international marketing margin (RMARG). y an internal marketing margin (INTMARG) (Sugema &
010). PCc,pr = PCc,p * (1 + INTMARG) where the subscripts c and h refer to commodity and
household type, respectively. The border prices (PM)
for tradable products are linked to the world price
by the exchange rate (er), import tariffs (tm) and the
international marketing margin (RMARG). y an internal marketing margin (INTMARG) (Sugema
010). PCc,pr = PCc,p * (1 + INTMARG) PMc= PWc * er *(1 + tmc) ....................................(2)
PCc,h = PMc * (1+RMARG)................................(3)
PCc,rt = PCc,pr
l d d
i
i d
f
h h
h ld Based on the description above, this research ana
lyzed the impact of domestic maize price changes on the
performance of broiler industry in Indonesia with the
problem statement of how the domestic maize price
changes has impacted the performance of maize farm
ing, broiler farming, and income of small-scale broiler
farming household. This study therefore was conducted
with the aims at analyzing the impact of domestic maize
price changes on: (1) the performance of maize farming,
(2) the performance of broiler farming, and (3) small-
scale broiler farming household income. INTRODUCTION Year
Domestic price
of maize (IDR)
Impor price of
maize (IDR)
Feed price of
broiler (IDR)
2000
1 440
1 029
2 345
2001
1 450
1 227
2 470
2002
1 493
1 069
2 577
2003
1 547
1 084
2 687
2004
1 605
1 497
2 796
2005
1 362
1 577
2 905
2006
1 500
1 425
3 194
2007
2 269
1 963
3 250
2008
3 301
3 160
3 693
2009
2 585
2 378
4 461
2010
2 547
2 340
4 450
Trend (%/year)
Domestic price
of maize (%)
Impor price of
maize (%)
Feed price of
broiler (%)
2000-2001
0.69
19.24
5.33
2001-2002
2.96
-12.87
4.33
2002-2003
3.61
1.40
4.27
2003-2004
3.74
38.09
4.06
2004-2005
-15.14
5.34
3.89
2005-2006
10.13
-9.60
9.95
2006-2007
51.27
37.75
1.75
2007-2008
45.48
60.97
13.63
2008-2009
-21.69
-24.74
20.80
2009-2010
-1.47
-1.60
-0.25
7.96
11.40
6.78
Source: BPS (2012), Pusdatin Ministry of Agriculture (2012), processed
Table 1. Maize and feed prices in Indonesia (Period 2000–2010) Table 1. Maize and feed prices in Indonesia (Period 2000–2010) The sustainability of broiler farming was deter
mined by price changes of inputs and outputs. The
change of input and output prices affected to input allo
cation (feed) and the production decision of small-scale
broiler farming. Saptana & Sumaryanto (2011) revealed
that large-scale broiler farming dominated market struc
ture of chicken meat. In addition, the large-scale broiler
farming was dominant in the input market of poultry
industry by making business agreements with feed
companies and such association of integrated firms that
operated from upstream to downstream activities, e.g. PT. Charoen Phokphan Indonesia, PT. Japfa Comfeed,
and PT. Sierad Produce. Feed price was determined by
feed industry that collaborated with poultry companies, December 2014 199 PMc = PWc
230
UMBOH ET AL. / Media Peternakan 37(3):198-205 (9) model definition, (10) fixing variables, (11) model
closure, (12) display of parameters and variables, (13)
solution statement, (14) optional solution, and (15) solu
tion reports. he border price (PM) of the importable productsare link
change rate (er), import tariffs (tm), and the inte feed price when decided the quantity of feed produc
tion. Moreover, the quantity demanded of maize was
determined by maize price itself rather than feed price. The problem of this research was how the change of
domestik maize prices impact on the performance of
small-scale broiler farming. INTRODUCTION 239
240
241
242
243 In addition, the large-scale (PRSHN) of broiler
consumer prices differ from the small-scale (PTRYT)
by an internal marketing margin (INTMARG). The
consumer prices can be defined as:
e included a price index for each household group
eighted by their shares of consumption:
∗
PC 1h i PCc,pr= PCc,p * (1 + INTMARG)..............................(4)
PCc,rt = PCc,pr.............................................................(5)
PINDEX
PCWTh,i PC0h,i
i
w is the budget share for each commodity. The sup We included a price index for each household group
to reflect changes in prices weighted by their shares of
consumption:
periods 0 denote starting prices and 1 end of simulation p Product Categories PINDEX= Si PCWTh,i * (PC1 h,i/PC0 h,i) .................(6
ly Block PINDEX= Si PCWTh,i * (PC1 h,i/PC0 h,i) .................(6)
where w is the budget share for each commodity. The
superscript on the PC terms refers to periods 0 denote
starting prices and 1 end of simulation prices. pp y
ousehold’s supply of rice and maize are determined by
ailable to each household; b) the share of that land alloca There are 6 (six) commodities in the model, namely:
rice (rc), maize (m), broiler (br), eggs (lr), urea (ur), and
triple super phosphate (TSP). The model assumes that
commodities (rice, maize, chicken meat, and eggs) are
used as final consumption by household and maize as
input in feed industry. 246
247
248 Supply block. Household’s supply of food products is
determined by total amount of land available, share of
that land allocated to the specific crops, and yield of the
crops. The share of land (SH) is a function of:
the associated yield for the crops. The share of land (
op by household group h is a function of all crop prices: Data log(SHh,f) = αh,f
s + βh,f,ff
s
ff
log(PPh,ff)
........(7)
rs to crop commodities (rice and maize). rs to crop commodities (rice and maize). D) f
f b h
h ld
h
f
ti
f This research used data of 2011 consisted of: (1)
production, input utilization, consumption, and income,
(2) price of inputs and outputs, and (3) elasticity. The
data source of production, input utilization, consump
tion, and income were Bureau of Statistics Central,
Directorate General of Livestock and Animal Health,
Bureau of Food Security of Ministry of Agriculture. 250 wh
251
Yie
252
as w
253
254
whe
251
Yie
252
as w
253
254 where f refers to crop commodities (rice and maize). (YLD) for crops f by household groups h are a function of out
(YLD) for crops f by household groups h are a function of Yields (YLD) of household groups are determined
by output producer price and input consumer prices. (
)
p
y
g
p
p
l as land. l as land. log YLDh,f = αh,f
y + βh,f
y log PPh,f +
γh,f,iin
y
iin
log(PCh,iin) (8)
log YLDh,f = αh,f + βh,f log PPh,f +
γh,f,iin
iin
log(PCh,iin) 15)
RESULTS AND DISCUSSION h,f,in
h,in
βh,f,in
f
g
h,f
γh,in
g (
h,in)
and for urea and TSP are given by:
and for urea and TSP are given by:
Total demand for urea and TSP are given by:
g HDFEh f = αl
+ βl
l
log PPh l +
γl
ff
l DINin= Sh HDINh,in ..............................................(14)
i
= HDINh i
h
(
,
, ,
,
,
efer to maize for livestock feed Supply and Demand of Maize log HCh,i = αh,i
h + βh,i,j
h
j
log PCh,j + γh,i
h log(YHh)
(17)
log HCh,i = αh,i
h + βh,i,j
h
j
log PCh,j + γh,i
h log(YHh)
consumption is given by :
log (YHh)(17)
where YH is household income, PC are consumer prices. hold income, PC are consumer prices. log HCh,i = αh,i
h + βh,i,j
h
j
log PCh,j + γh,i
h log(YHh)
(17)
log HCh,i = αh,i
h + βh,i,j
h
j
log PCh,j + γh,i
h log(YHh)
consumption is given by :
log (YHh)(17)
where YH is household income, PC are consumer prices. mption is given by :
S = HC
NSi = HCh,i
h
Total consumption is given by : i
h,i
h
ck
ock
CONSi= Sh HCh,i ..............................................(18) .(18) Agricultural income (YHAG) for rural households is the sum of crop an
ock revenue minus input costs:
Agricultural income (YHAG) for rural households is th
Income block. Agricultural income (YHAG) is the sum
of crop and poultry revenue minus input costs: YHAGh=∑f(PPf*SCRh,f)+∑1(PP1*SLVh,1)–(PCin*DINh,in)-(PCfe*DFEh,fe) (1
tal household income (YH) is the sum of agricultural income and the exogenous
ck revenue minus input costs:
YHAGh=∑f(PPf*SCRh f)+∑1(PP1*SLVh 1)–(PCin*DINh in)-(P
YHAGh=∑f(PPf*SCRh,f)+∑1(PP1*SLVh,1)–(PCin*DINh,in)-
(PCfe*DFEh,fe) ........................(9) mined non-agricultural income (YHNAG). The latter component is adjusted by
ndexand the price index is as defined in equation (6). YHh = YHAGh + YHNAG
h ∗PINDEX
(20)
tal household income (YH) is the sum of agricultural income
mined non-agricultural income (YHNAG). The latter compon
ndexand the price index is as defined in equation (6)
and total household income (YH) is the sum of
agricultural income and the exogenously determined
non-agricultural
income
(YHNAG). The
latter
component is adjusted by a price index and the price
index is as defined in equation (6). d the exogenously
t is adjusted by a
(20)
To meet domestic demand for maize, some at
tempts have been done such as increasing productivity,
expanding planted maize areas, improving technical
assistance, empowering agricultural institution, and
providing financial support. As a result, there was a
significant production increase of 7.86% per year during
the period 2005-2011 (BPS, 2012). It is hoped that the
domestically produced maize will be able to close the
domestic demand. h = YHAGh + YHNAG
h ∗PINDEX
YHh= YHAGh + YHNAGh * PINDEX ................(20) brium Conditions
Equilibrium conditions. In this block, it is assumed that
there is “market clearing condition” for all commodity brium Conditions
Equilibrium conditions. Structure of the Model Those are mathematically defined as SCRf= Sh SCRh,f .......................................(10)
livestock supply is modelled as a function of livestock p
livestock feed products, where the subscript ffe refer SCRf= Sh SCRh,f .......................................(10)
livestock supply is modelled as a function of livestock
ivestock feed products, where the subscript ffe ref The poultry production by households is a function
of poultry prices and poultry input prices of feed. of livestock feed products, where the subscript ffe refers to livestoc
cts. cts. log SLVh l = αh l
1 + βh l ll
1
ll
log PPh l +
γh l ffe
1
ffe
log(PCh ffe 1) SCRf + SLV1 + MC = CONSi + DFEC .....................(21) log SLVh,l = αh,l
1 + βh,l.ll
1
ll
log PPh,l +
γh,l,ffe
1
ffe
log(PCh,ffe)
log (PCh,ffe) (11)
livestock supply is given by: 1) where M equals imports and CONS and DFE denote
human and livestock consumption respectively. stock supply is given by:
Total poultry supply is given by:
LVl = SLVh,l
h Model Simulation = SLVh,l
h
t Bl
k
SLV1= Sh SLVh,1 .....................................................(12)
nput Block and
for
(PP) and
mand for
(13)
(14)
Two simulations were conducted to answer the ob
jectives of this study, i.e: (1) Decreased domestic price of
maize by 10%, and (2) Increased domestic price of maize
by 10%. These simulations concerned to the increasing
trend of maize price during 2000-2010 that reached
about 7.96%. In 2011, maize price increased remarkably
from IDR 2547 to IDR 2933 or increased around 15%. nd Input Block
Household demand for input (HDIN) is a function of output prices (P
prices (PC), where the subscript in refers to urea andTSP. Household dem
Demand input block. Household demand for input
(HDIN) is a function of output prices (PP) and input pric
es (PC), where the subscript in refers to Urea and TSP. Household demand for urea and TSP:
Household demand for input (HDIN) is a function of output price
prices (PC), where the subscript in refers to urea andTSP. Household
nd TSP:
DINin = HDINh in
h d TSP:
og HDINh,f,in = αh,in
f
+ βh,f,in
f
f
log PPh,f + γh,in
f
log(PCh,in)
mand for urea and TSP are given by:
log HDINh,f,in = αh,in
f
+ βh,f,in
f
f
log PPh,f + γh,in
f
log(PCh,in)
emand for urea and TSP are given by:
log (PCh,in)(13)
Total demand for urea and TSP are given by:
old demand maize for livestock feed:
log HDFEh f = αh f
l
+ βh l f
l
l
log PPh l +
γh f
ff
l
ff
lo nd TSP:
log HDINh,f,in = αh,in
f
+ βh,f,in
f
f
log PPh,f + γh,in
f
log(PCh,in)
demand for urea and TSP are given by:
log HDINh,f,in = αh,in
f
+ βh,f,in
f
f
log PPh,f + γh,in
f
log(PCh,in)
demand for urea and TSP are given by:
log (PCh,in)(13)
Total demand for urea and TSP are given by:
ehold demand maize for livestock feed:
log HDFEh f = αh f
l
+ βh l f
l
l
log PPh l +
γh f
ff
l
ff
lo nd TSP:
log HDINh,f,in = αh,in
f
+ βh,f,in
f
f
log PPh,f + γh,in
f
log(PCh,in)
log HDINh,f,in = αh,in
f
+ βh,f,in
f
f
log PPh,f + γh,in
f
log(PCh,in)
log (PCh,in)(13)
hold demand maize for livestock feed: Supply and Demand of Maize Household demand maize for poultry feed:
,
demand maize for livestock feed:
nd for maize is given by: (16)
During the period 2005-2011, the development
trend of harvested area, production and productivity of
maize nationally showed an increase by 2.62%, 7.86%
and 5.23%, respectively. In 2011, the national maize
harvested area reached 3.86 million hectares with pro
duction and productivity levels of 17.63 million tons and
4.57 ton/ha, respectively (BPS, 2012). log HDFEh,fe = αh,fe
l
+ βh,l,fe
l
l
log PPh,l +
γh,fe,ffe
l
ffe
log PCh,ffe (15
(15)
g
y
DFEfe = HDFEh fe
h e fe refer to maize for livestock feed. demand for maize is given by:
where fe refer to maize for poultry feed. Total demand for maize is given by:
umption Block fe = HDFEh,fe
h
DFEfe= Sh HDFEh,fe ..............................................(16)
mand for the consumption items (HC) by the househ oups, where the i
and eggs). YH is
(17)
(18)
um of crop and
*DFEh,fe) (19)
p
y (
)
Although increased, the national maize productiv
ity is still low due to most of farmers still have used
seeds local varieties. Some other farmers eventhough
they grow maize hybrids and composites, but they
managed it less intensively. This conditions have caused
maize yield is still far from its potency (Swastika et al.,
2005). Indeed, if farmers are willing to manage their
maize farm intensively or use input according to recom
mended technology, the productivity can reach 6-10 tons
per hectare for maize hybrid and 5-8 tons/ha for maize
composite. This means that farmers have not utilized
the production potential optimally and the possibility in
getting more income through using new superior variet
ies of maize. This condition is caused by several things,
among others are: (1) farmers do not have enough
money to buy superior seeds, fertilizers and pesticides
needed and (2) the seed available is frequent delay, so
that planting activity cannot be done on time. As a
result, the maize productivity level is not as expected
(Yusdja & Agustian, 2003). umption Block
Demand for the consumption items (HC) by the household groups, where th
o commodities households purchase (rice, maize, chicken meat, and eggs). YH
Consumption block. The demand for each commodity
is modeled as the consumption function of household
groups as follow:
o commodities households purchase (rice, maize, chicken me
hold income, PC are consumer prices. hold income, PC are consumer prices. Structure of the Model the coefficients (α, β, γ)represent the price elasticities. where the coefficients (a, b, g) represent the price
elasticities. the coefficients (α, β, γ)represent the price elasticities. tal household supply to the market is then determined as the pr This research employed multimarket model from
Umboh et al. (2014) and it was divided into 6 blocks of
equations, namely: (1) price, (2) supply, (3) input de
mand, (4) consumption, (5) income, and (6) equilibrium
condition. The model was originally designed in General
Algebric Modelling System (GAMS) using the Path NLP
solver consist of fifteen steps, namely: (1) set declaration,
(2) database, (3) parameter declaration, (4) parameter
definition, (5) variable declaration, (6) variable defini
tion, (7) equation declaration, (8) equation definition,
The
256
are
257
Ad
258
con
259
260
The
261
The
256
area
257
Adj
258
con
259
260
The
261 tal household supply to the market is then determined as the produ
under cultivation, the share of land devoted to the crop, an
tmentsare made for losses and use of the output for seed (loss), and f
rsionfactors (conv). The production of food crops by each household
is determined by total land devoted to the crop, the
share of land devoted to the crop, and the yield. After
that, total production available at the market is adjusted
by losses, use of the food crops for seed (loss), and
conversion factors (conv), hence:
tal household supply to the market is then determined as the pr
under cultivation, the share of land devoted to the crop
mentsare made for losses and use of the output for seed (loss), a
sionfactors (conv). SCRh,f = AREA
∗SHh,f ∗YLDh,f ∗(1 −lossf)
∗convf
SCRh,f = AREA
∗SHh,f ∗YLDh,f ∗(1 −lossf)
∗convf
(9) (9) 200 December 2014 onvf
(9)
f h
h ld
l
(10)
UMBOH ET AL. / Media Peternakan 37(3):198-205 The total supply of each of the food crops are:
Rf = SCRh,f
h
livestock supply is modelled as a function of livestock The total supply of each of the food crops are:
Rf = SCRh,f
h
livestock supply is modelled as a function of livestock 0)
nput
feed
nd input
ock feed
markets. In this regard, the sum of quantity supplied (do
mestic production plus net imports) is equal to the quan
tity demanded for both human consumption and feed
demand. Supply and Demand of Maize and Chicken Meat in Table 2. For maize farmers, the reduction in domestic
maize price was responded by reducing the land share
for planted maize land by 3.120% for small-scale broiler
farming households and by 3.133% for other house
holds, respectively. At the same time both two types of
households have decided to reduce in Urea and TSP fer
tilizers uses of by 5.668% and 2.611%, respectively. The
changes of farmer decision for maize planted area size
and fertilizers uses, then had impact on maize produc
tion decrease by 2.615%. But, opposite impact accoured
when the maize price goes down, in which farmers will
be more interesting to increase rice planted area size. In
this condition, planting rice gives more interesting ben
efit than that of planting maize. This was showed by the
increased share of rice planted land for small-scale broil
er farming households and other households by 1.1436%
and 1.140%, respectively. Therefore, when there was a
decline in the maize price, farmers would divert their
planted maize area to rice farming. In this case, the farm
ers also are willing to use more input production for rice
that had positive impact on rice productivity. Therefore,
at level of productivity and land share increase would
lead to the increase in rice production by 1.7481%. Comparing with the national maize production
data, in 2005 national maize production reached 12.52
million ton, then decreased to 11.61 million ton in 2006
and increased again to 17.63 million ton in 2011. Based
on that, in 2006 there was a deficit so for answered the
demand, the government did an import around 1.84
million ton. In 2007-2011, national maize production
reached over the demand. However, maize still im
ported around 414 thousand ton in 2007 and increased
to 3.1 million ton in 2011 (BPS, 2012). Maize import was done by the feed industries with
some reasons such as, the difficulties to find maize from
the farmer, national maize production does not available
in a whole year, and maize import buying mechanism is
more easier than local corn buying, because they do not
need to get in touch with the farmers or producers di
rectly (Malian, 2004; Hakim, 2005; Swastika et al., 2011). In 2005, the total maize requirement reached around
11.86 million ton, then increased to 13.71 million ton in
2008, and became 16.50 million ton in 2011 (BPS, 2012). Impact of Domestic Maize Price Changes on the
Performance of Broiler Farming Domestic maize price decrease and the existing
margin between the large-scale and small-scale broiler
farming by 20%, where by IDR 2933 on the broiler farm
ing of large-scale and IDR 3519.60 on the small-scale,
respectively (Table 2), had an impact on the performance
of broiler farming as shown in Table 3. For small-scale
broiler farming, when the price of maize to be paid by
the small-scale was 20% higher than the price paid by the
broiler large-scale, increasing the maize demand for feed
by 0.5268%, this in turn would increase chicken meat
production by 0.5279%. The same phenomenon also
occured in large-scale production, where the increased
maize demand for broiler feed triggered the increase in
production large-scale of by 0.5285%. Furthermore, the
impact of domestic maize price decrease by 10% leading
to the increase in maize demand for feed by the large
and small-scale broiler farming, increasing the national The most broiler population are located in 14
provinces, namely West Java, East Java, Central Java,
Banten, North Sumatra, Riau, East Kalimantan, South
Kalimantan, West Kalimantan, South Sumatra, South
Sulawesi, Lampung, Riau, and Jambi. Meanwhile, for
layer are located in 12 provinces: East Java, Central
Java, West Java, North Sumatra, West Sumatra, Banten,
South Sulawesi, South Sumatra, Bali, Lampung, and
West Kalimantan (Directorate General of Livestock and
Animal Health, 2012.) Supply and Demand of Maize and Chicken Meat Related to production and price chicken meat, al
though there was broiler feed price increasing because of
maize price increasing, there was still broiler production
increasing. This was because of the input price increas
ing (feed) compensated with the output price increasing
(chicken meat), so that this gave an incentive to the
farmer to increase their production. During 2000-2010,
chicken price and production increased around 5.86%
and% 14.453, respectively (BPS, 2012). p
y
Domestic maize price increased was responded by
farmers by increasing maize planted area by 2.84% and
2.876% on the households of small-scale broiler farming
and other households, respectively. In addition, the im
provement of maize price led to Urea and TSP fertilizers
uses for maize farming increased by 5.371% and 5.383%,
respectively and then improved maize productivity by
2.39971 and 2.39910 on the households of small-scale
and other households. Increased land share for and pro
ductivity of maize would cause the increase in national
maize production by 5.34606%. On the other side, this
condition has a negative impact on rice production by
1.41942%. This indicated that when the price of maize
increases, farmers would use part of their rice land for
planting maize. These results were in line with other
research results, such as Sayaka et al. (2007); Huang et
al. (2009); and Dorosh et al. (2009). Theoritically, the cause of chicken meat price
increasing, besides caused by production cost increas
ing, also caused by the increasing of demand because
of population, nutritional consumption awareness, and
income increasing (Ilham, 2009; Ilham & Yusdja, 2010). By the time, chicken meat contribution to the national
meat consumption has reached around 70.24% (Saptana
& Sumaryanto, 2009). During 2000-2009, chicken meat
consumption was around 2.33 kg per capita per year. The demand for chicken meat continuously increased to
3.10 kg per capita in 2010 and reached around 4.53 kg
per capita per year in 2011 (Bureau of Food Security of
Ministry of Agriculture, 2012). Supply and Demand of Maize In this block, it is assumed that
there is “market clearing condition” for all commodity December 2014 201 UMBOH ET AL. / Media Peternakan 37(3):198-205 202 December 2014 Note: *) increasing domestic maize price by 10%; **) decreasing domestic maize price by 10%. Impact of Domestic Maize Price Changes on the
Performance of Maize Farming The reseach result showed that domestic maize
price decrease by 10% had a negative impact on the
performance of maize farming in Indonesia as shown 202 December 2014 UMBOH ET AL. / Media Peternakan 37(3):198-205 Note: *) increasing domestic maize price by 10%; **) decreasing domestic maize price by 10%. Variables
Base value
(Unit)
Alternative simulation
Simulation 1*
Simulation 2**
(Unit)
(%)
(Unit)
(%)
PMm (Imported Maize Price)
2346.40
0.00
0.00
0.00
0.00
PCm,p (Maize Price of PRSHN)
2933.00
293.30
10.00
-293.30
-10.00
PCm,pr (Maize Price of PTRYT)
3519.60
351.96
10.00
-351.96
-10.00
PMrc (Imported Rice Price)
6150.40
0.00
0.00
0.00
0.00
PCrc (Domestic Rice Price)
7688.00
0.00
0.00
0.00
0.00
SH1,2 (Rice Land Share of PTRYT)
0.078
-0.001
-0.872
0.001
1.144
SH1,2 (Rice Land Share of RTOTH)
0.526
-0.005
-0.880
0.006
1.14
YLD1,2 (Rice Productivity of PTRYT)
4.207
-0.018
-0.419
0.025
0.594
YLD1,2 (Rice Productivity of RTOTH)
5.58
-0.023
-0.419
0.033
0.591
HSCR1 (Rice Production of PTRYT)
2598.32
-36.877
-1.419
45.425
1.748
HSCR2 (Rice Production of RTOTH)
23240.54
-329.885
-1.419
406.30
1.748
SCR1,2(Rice Production of Indonesia)
25838.86
-366.762
-1.419
451.69
1.748
SH3,4 (Maize Land Share of PTRYT)
0.025
0.001
2.84
-0.001
-3.12
SH3,4 (Maize Land Share of RTOTH)
0.160
0.005
2.876
-0.005
-3.133
YLD3,4 (Maize Productivity of PTRYT)
3.50
0.084
2.400
-0.091
-2.611
YLD3,4 (Maize Productivity of RTOTH)
4.56
0.109
2.399
-0.119
-2.610
HSCR3 (Maize Production of PTRYT)
1107.53
59.209
5.346
-62.80
-5.67
HSCR4 (Maize Production of RTOTH)
9211.81
492.47
5.346
-522.24
-5.669
SCR3,4 (Maize Production of Indonesia)
10319.34
551.68
5.346
-585.03
-5.669
HDIN1 (Demand for Urea PTRYT)
1.679
0.09
5.36
-0.095
-5.658
HDIN2 (Demand for Urea RTOTH)
1049.36
56.48
5.382
-59.50
-5.67
DIN1,2 (Demand for Urea Indonesia)
1051.04
56.57
5.371
-59.58
-5.668
HDIN3 (Demand forTSP PTRYT)
0.705
0.038
5.39
-0.04
-5.673
HDIN4 (Demand for TSP RTOTH)
440.73
23.72
5.381
-11.53
-2.616
DIN3,4 (Demand for TSP Indonesia)
441.43
23.763
5.383
-11.53
-2.611
CONS1,2 (Consumption for Rice)
24789.08
42.60
0.172
-90.65
-0.365
CONS3,4 (Consumption for Maize)
7941.02
-52.46
-0.66
124.32
1.565
IMrc (Net Import of Rice)
4499.991
409.36
9.096
-542.339
-12.052
IMm (Net Import of Maize)
3182.356
-607.83
-19.10
709.364
22.29
Table 2. The results of simulation of domestic maize price changes on the performance of maize farming in Indonesia Table 2. The results of simulation of domestic maize price changes on the performance of maize farming in Indonesia ble 3. Impact of Domestic Maize Price Changes on the
Performance of Maize Farming The results of simulation of domestic maize price changes on the performance of broiler farming in Indone esults of simulation of domestic maize price changes on the performance of broiler farming in Indonesia Table 3. The results of simulation of domestic maize price changes on the performance of broiler farming in Indonesia
Variables
Base value
(Unit)
Alternative Simulation
Simulation 1*
Simulation 2**
(Unit)
(%)
(Unit)
(%)
PCbr (Domestic Chicken Meat Price)
27500.00
0.000
0.000
0.000
0.000
PClr (Domestic Egg Price)
18058.00
0.000
0.000
0.000
0.000
HSLV1(Chicken Meat Production of PRSHN)
1137.12
-5.40
-0.474
6.010
0.529
HSLV2(Chicken Meat Production of PTRYT)
181.83
-0.867
-0.477
0.960
0.528
SLV1,2 (Chicken Meat Production of Indonesia )
1318.95
-6.207
-0.470
6.970
0.528
HSLV3( Egg Production of PTRYT)
0.10
-0.000
-0.460
0.001
1.300
HSLV4( Egg Production of RTOTH)
1025.91
-4.879
-0.476
5.420
0.528
SLV3,4 ( Egg Production of Indonesia)
1026.01
-4.880
-0.477
5.427
0.529
HDFE1 (Demand for Maize PRSHN)
1066.37
-5.070
-0.475
5.640
0.529
HDFE2 (Demand Maize for Broiler Feeds PTRYT)
227.00
-1.080
-0.476
1.196
0.527
HDFE3 (Demand Maize for Layer Feeds PTRYT)
0.127
-0.001
-0.472
0.001
0.787
HDFE4 (Demand for Maize RTOTH)
1292.25
-6.140
-0.475
6.834
0.529
DFE1,2,3,4 (Demand for Maize Indonesia)
2585.747
-12.290
-0.475
13.671
0.528
CONSP1,2 (Consumption for Chicken Meat Indonesia)
1137.21
-0.630
-0.055
1.032
0.090
CONSP1,2 (Consumption for Eggs Indonesia)
1027.80
-3.980
-0.387
7.717
0.750
Note: *) increasing domestic maize price by 10%; **) decreasing domestic maize price by 10%. December 2014 203 UMBOH ET AL. / Media Peternakan 37(3):198-205 Table 4. The results of simulation of domestic maize price changes on households income of small-scale broiler farming (Million
Rupiahs) Table 4. The results of simulation of domestic maize price changes on households income of small-scale broiler farming (Million
Rupiahs)
Variables
Base value
(Unit)
Alternative Simulation
Simulation 1*
Simulation 2**
(Unit)
(%)
(Unit)
(%)
a. YHAG1 (Income of PRSHN)
21888976
-415240.45
-1.897
430092.870
1.965
b. YHAG2 (Income of PTRYT)
22286608
-153717.82
-0.690
210125.455
0.943
1. Rice
15968858.530
-226640.48
-1.419
279174.770
1.748
2. Maize
3115440.070
167759.37
5.385
-176713.795
-5.672
3. Broiler
3201210.80
-94787
-2.961
97182.470
3.036
4. Laying hens
997.65
-49.08
-4.920
58.510
5.865
c. YHAG3 (Income of RTOTH)
1049290416
-7219118.062
-0.688
9621993.115
0.917
Total (a+b+c)
1093466000
-10597552.96
-0.969
10262211.120
0.938
d. YHNAG1(Income of PRSHN)
0.00
0.00
0.000
0.000
0.000
e. YHNAG2(Income of PTRYT)
3039080
0.00
0.000
0.000
0.000
f. Impact of Domestic Maize Price Changes on the
Performance of Maize Farming YHNAG3(Income of RTOTH)
6445640124
0.00
0.000
0.000
0.000
Total (d+e+f)
6448679204
0.00
0.000
-3025496.54
-1.878
g. YH1 (Income of. PRSHN)
21888976
-415240.45
-1.897
430092870
1.965
h. YH2 (Income of PTRYT)
25325688
-153718.77
-0.607
210124.51
0.829
i. YH3 (Income of RTOTH)
7494930540
-9768894
-0.130
9621993
1.284
Total (g+h+i)
7542145204
-10337853
-0.137
10262211.17
0.136
Note: *) increasing domestic maize price by 10%; **) decreasing domestic maize price by 10%. Impact of Domestic Maize Price Changes on the
Household Income of Small-scale Broiler Farming Impact of Domestic Maize Price Changes on the
Household Income of Small-scale Broiler Farming The decrease of domestic maize price by 10% and
if the price margin to be paid by small-scale broiler
farming was 20% higher than the price large-scale had
to pay, the household income of small-scale broiler
farming would decrease by 5.572% from maize farm
ing, but it increased their income from rice farming,
broiler business and layer business by 1.748%, 3.036%,
and 5.865%, respectively (Table 4). This policy caused
the increase in agricultural income (YHAG) as well as
national income (YH) of by 0.938% and 0.136%, respec
tively. In contrast, when government issued the policy of
increasing domestic maize price of by 10%, it caused the
household income of small-scale broiler farming from
maize increased by 5.385%, but decrease of the income
of small-scale broiler farming from rice farming, broiler,
and layer business by 1.419%, 2.961%, and 4.920%,
respectively. Furthermore, this policy caused the agri
cultural sector income (YHAG) and the national income
(YH) go down by 0.969% and 0.137%, respectively. CONCLUSION production of by 0.5284%. This condition also had a
positive impact on the improvement of layers produc
tion of by 0.5289%. The price policy to increase maize price had positive
impacts on performance of maize farming and farmer
income by increasing maize cultivated area and input
utilization to increase domestic maize production and to
reduce maize import. On the other hand, the lowering
of domestic maize price was responded by farmers
through reducing maize planted land and fertilizer input
uses, causing maize production and farmer income
from maize farming decline; in addition, price policy to
increase maize price showed a negative impact on the
performance and income of small-scale broiler farming
as it caused the increase of production cost due to the
increasing feed price. In contrast, reduction maize price
would have an impact on increasing maize demand for
feed industry. Aside from that, other impacts from this
policy were increasing broilers production and small-
scale broiler farming income. In general, this policy
could improve the level of agricultural income and
national income. y
In contrast, when the domestic maize price in
creased by 10%, in which maize demand by both large
and small-scale broiler farming went down by 0.4754%
and 0.4757%, respectively. Consequently, the produc
tion for meat also decreased by 0.474% and 0.477%,
respectively, both in large and small-scale broiler farm
ing. These results were in accordance with Kariyasa &
Sinaga (2007). REFERENCES Bureau of Food Security of Ministry of Agriculture. 2012. De
velopment of Food Consumption Directory. Bureau of
Food Security of Ministry of Agriculture, Jakarta. BPS. 2012. Statistic Indonesia. Central Bureau of Statistic, Ja
karta. Directorate General of Livestock and Animal Health. 2012. Livestock and Animal Health Statistics. Directorate Gen
eral of Livestock and Animal Health, Jakarta. Dorosh, P. A., C. del Ninno, & D. E. Sahn. 1995. Poverty allevi
ation in Mozambique: a multimarket analysis of the role of
food aid. Agricultural Economics 13:89-99. http://dx.doi. org/10.1016/0169-5150(95)01156-0 204 December 2014 UMBOH ET AL. / Media Peternakan 37(3):198-205 Malian, A. H. 2004. International trade policy of agricultural
commodities. Analisis Kebijakan Pertanian 2:135-156.t Dorosh, P. A., S. Dradri, & S. Huggblade. 2009. Regional trade,
government policy and food security recent evidence from
Zambia. Food Policy 34:350-366. http://dx.doi.org/10.1016/
j.foodpol.2009.02.001 Minot, N. & F. Goletti. 1998. Export and liberalization and
household welfare: the case of rice in Vietnam. AJAE
80:738-749. http://dx.doi.org/10.2307/1244060 j
p
Hakim, D. B. 2005. Impact of multilateral cooperation on trade
of agricultural product. Jurnal Manajemen dan Agribisnis
2:144-145. t p
g
Pusdatin Ministry of Agriculture. 2012. Statistics Agricultural
Commodity Prices, Jakarta. Huang, J., Y. Liu, W. Martin, & S. Rozelle. 2009. Changes in
trade and domestic distortions affecting China’s agri
culture. Food Policy Journal 34:407-416. http://dx.doi. org/10.1016/j.foodpol.2009.04.001 Saptana & Sumaryanto. 2009. Anticipating Jakarta Provincial
government’s regulation on poultry in 2010. Analisis Kebi
jakan Pertanian 7:319-335. Swastika, D. K. S., M. Manikmas, B. Sayaka, & I. K. Kariyasa. 2005. The behavior of maize supply and demand in Indo
nesia. Jurnal Penelitian Pertanian Tanaman Pangan 24:67-
75. g
j
p
Hutabarat, B., A. Setiyanto, R. Kustiari, & T. B. Sulser. 2012. Conjecturing production, imports and consumption of
horticulture in Indonesia in 2050: A GAMS simulation
through changes in yields induced by climate change. Jur
nal Agro Ekonomi 30:1-23. Swastika, D. K. S, A. Agustian, & T. Sudaryanto. 2011. Gap
analysis of supply and demand of corn forage production
approach sync center, feed plant, animal and population
in Indonesia. Informatika Pertanian 2:65-75. g
Ilham, N. 2009. Shortage of meat production: policy indication
and implication. Analisis Kebijakan Pertanian 7:43-63.l p
Ilham, N. & Y. Yusdja. 2010. Impact of Avian Influenza on pro
duction and contribution of poultry to small-scale farmers
income in Indonesia. Jurnal Agro Ekonomi 28:39-68. Umboh, S. J. K, D. B. Hakim, B. M. Sinaga, & I. K. Kariyasa. 2014. REFERENCES Impact of maize import tariff policy changes on pro
duction and consumption in Indonesia: A multimarket
model analysis. IJFAE 2: 113-126. g
Indarsih, B., M. H. Tamsil, & M. P. Nugroho. 2010. A study of
contract broiler production in Lombok, NTB: An opportu
nity of introducing syariah partnership. Med. Pet. 33:124-
130. http://dx.doi.org/10.5398/medpet.2010.33.2.124 y
Yusdja, Y. & E. Pasandaran. 1998. Restructurization direction
of Indonesia poultry agribusiness. Jurnal Agro Ekonomi
16:21-32. f Yusdja, Y. & A. Agustian. 2003. Maize tariff policy analysis. Analisis Kebijakan Pertanian 1:36-54. t p
g
p
Kariyasa, I. & B. M. Sinaga. 2007. Feed and chicken meat mar
kets behaviour analysis in Indonesia: simultaneous econo
metric model approach. Journal on Socio-Economics of
Agriculture and Agribusiness (SOCA) 7:158-166. December 2014 205
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https://www.nature.com/articles/srep43105.pdf
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Rumination and Default Mode Network Subsystems Connectivity in First-episode, Drug-Naive Young Patients with Major Depressive Disorder
|
Scientific reports
| 2,017
|
cc-by
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Rumination and Default Mode
Network Subsystems Connectivity
in First-episode, Drug-Naive Young
Patients with Major Depressive
Disorder received: 19 August 2016
accepted: 18 January 2017
Published: 22 February 2017 Xueling Zhu1,2,3, Qiuling Zhu4, Huaizhen Shen2, Weihua Liao5 & Fulai Yuan1 Neuroimaging evidence implicates the association between rumination and default mode network
(DMN) in major depressive disorder (MDD). However, the relationship between rumination and DMN
subsystems remains incompletely understood, especially in patients with MDD. Thirty-three first-
episode drug-naive patients with MDD and thirty-three healthy controls (HCs) were enrolled and
underwent resting-sate fMRI scanning. Functional connectivity analysis was performed based on
11 pre-defined regions of interest (ROIs) for three DMN subsystems: the midline core, dorsal medial
prefrontal cortex (dMPFC) and medial temporal lobe (MTL). Compared with HCs group, patients with
MDD exhibited increased within-system connectivity in the dMPFC subsystem and inter-system
connectivity between the dMPFC and MTL subsystems. Decreased inter-system connectivity was
identified between the midline core and dMPFC subsystem in MDD patients. Depressive rumination
was positively correlated with within-system connectivity in the dMPFC subsystem (dMPFC-TempP)
and with inter-system connectivity between the dMPFC and MTL subsystems (LTC-PHC). Our results
suggest MDD may be characterized by abnormal DMN subsystems connectivity, which may contribute
to the pathophysiology of the maladaptive self-focus in MDD patients. Rumination is defined as a mode of responding to distress that involves repetitively and passively focusing on
symptoms of distress and on the possible causes and consequences of these symptoms1,2. It is strongly and con-
sistently related to depressive symptoms3. Evidence from prospective longitudinal and experimental studies have
emphasized the role of rumination in the onset and maintenance of symptoms and the diagnosis of major depres-
sive disorder (MDD)2,4,5, onset of depressive symptomatology in non-depressed people2, and risk of depressive
relapse in remitted patients6. p
p
Although the pathophysiological mechanisms underlying MDD remain unclear, neuroimaging studies have
shown that MDD has been conceptualized as a neural network-level disease7–10. Among the neural networks iden-
tified in MDD, the default mode network (DMN) has received growing attention. It encompasses a specific set of
brain regions, including the medial prefrontal cortex (MPFC), precuneus/posterior cingulate cortex (PCC), and
medial, lateral, and inferior parietal cortex11,12. Researchers have consistently reported that the DMN is involved
in internal mentation, including self-referential processing, the memory retrieval process and autobiographical
memory retrieval13–15, which is closely associated with depressive symptomatology15,16. www.nature.com/scientificreports www.nature.com/scientificreports www.nature.com/scientificreports received: 19 August 2016
accepted: 18 January 2017
Published: 22 February 2017 Rumination and Default Mode
Network Subsystems Connectivity
in First-episode, Drug-Naive Young
Patients with Major Depressive
Disorder MDD: major depressive disorder; HC:
healthy control; Data value are Mean ± SD; F/M: female/male; SD. CES-D: Center for Epidemiologic Studies
Depression; RSQ: Response to Stress Questionnaire. aChi-square test; bTwo sample t-test. Table 1. Demographic and clinical characteristics for subjects. MDD: major depressive disorder; HC:
healthy control; Data value are Mean ± SD; F/M: female/male; SD. CES-D: Center for Epidemiologic Studies
Depression; RSQ: Response to Stress Questionnaire. aChi-square test; bTwo sample t-test. Recent imaging studies have suggested the intuitive association between the self-focused rumination in MDD
and the self-referential operations performed by DMN25,26, which has prompted further interest in examining the
role of DMN in MDD. It was revealed that people with MDD showed stronger functional connectivity with DMN
in the subgenual prefrontal cortex (sgPFC), the degree of activation being modulated by the level of maladaptive
rumination27,28. Similarly, our previous study also reported increased functional connectivity with DMN in ante-
rior medial cortex including the sgPFC in MDD patients29. Furthermore, effective connectivity analysis revealed
mutually propagating activation between the sgPFC and ventral MPFC in people with MDD, which predicted
higher levels of depressive rumination30. Convergent findings from neuroimaging studies have identified the role
of the sgPFC in the DMN and its relationship with rumination in MDD, which suggest that the sgPFC might be
unique to depression and a neural substrate of depressive rumination15,26. However, the association between the
rumination and DMN in MDD still remains unclear and requires further investigation in this field. q
gi
Although the DMN originally was regarded as a large-scale brain system, current studies have noted the
DMN is not as homogenous as previously assumed and suggested that its architecture can be further subdivided
into smaller anatomical–functional subsystems31. Furthermore, the subsystems and the interaction among them
have been shown to provide interesting insights for exploring mental diseases32–35. Recent evidence both from
monkeys and humans have suggested that the DMN comprises a set of interacting subsystems, including a mid-
line core constituted by hub regions in the anterior medial prefrontal and posterior cingulate cortices as well
as two functionally dissociable subsystems: the dorsal medial prefrontal cortex (dMPFC) subsystem involved
in self-referential processes and the medial temporal lobe (MTL) subsystem involved in episodic memory31. Furthermore, altered interaction among the three subsystems within the DMN might be crucial in the psychopa-
thology of some neuropsychiatric disorders, including Alzheimer’s disease32, obsessive-compulsive disorder33 and
schizophrenia34. Rumination and Default Mode
Network Subsystems Connectivity
in First-episode, Drug-Naive Young
Patients with Major Depressive
Disorder Striking differences in
the activity and functional connectivity of the DMN between individuals with MDD and healthy controls (HCs)
have been identified in a large number of studies using either task-based17–19 or resting-state fMRI20–23. Despite
inconsistent findings, the DMN has been considered to play a central role in the physiopathology of MDD24. 1Health Management Center, Xiangya Hospital, Central South University, Changsha, 410008, China. 2School of
Humanities and Social Sciences, National University of Defense Technology, Changsha, 410074, China. 3Medical
Psychological Institute, Second Xiangya Hospital, Central South University, Changsha, 410011, China. 4Obstetrics
Department, Jinan Maternity and Child Care Hospital, Jinan, 250001, China. 5Department of Radiology, Xiangya
Hospital, Central South University, Changsha, 410008, China. Correspondence and requests for materials should be
addressed to F.Y. (email: fulaiyuan2010@163.com) Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 1 www.nature.com/scientificreports/ Characteristic
MDD (n = 31)
HC (n = 32)
P value
Gender (F/M)
17/14
17/15
0.657a
Age (years)
20.53 ± 1.78
20.96 ± 1.28
0.434b
Education (years)
13.62 ± 0.76
13.75 ± 0.82
0.611b
Illness duration (years)
0.29 ± 0.20
NA
CES-D
36.90 ± 6.79
17.97 ± 7.93
<0.001b
RSQ-rumination
7.74 ± 1.18
4.65 ± 1.68
<0.001b
Table 1. Demographic and clinical characteristics for subjects. MDD: major depressive disorder; HC:
healthy control; Data value are Mean ± SD; F/M: female/male; SD. CES-D: Center for Epidemiologic Studies
Depression; RSQ: Response to Stress Questionnaire. aChi-square test; bTwo sample t-test. Characteristic
MDD (n = 31)
HC (n = 32)
P value
Gender (F/M)
17/14
17/15
0.657a
Age (years)
20.53 ± 1.78
20.96 ± 1.28
0.434b
Education (years)
13.62 ± 0.76
13.75 ± 0.82
0.611b
Illness duration (years)
0.29 ± 0.20
NA
CES-D
36.90 ± 6.79
17.97 ± 7.93
<0.001b
RSQ-rumination
7.74 ± 1.18
4.65 ± 1.68
<0.001b
Table 1. Demographic and clinical characteristics for subjects. MDD: major depressive disorder; HC:
healthy control; Data value are Mean ± SD; F/M: female/male; SD. CES-D: Center for Epidemiologic Studies
Depression; RSQ: Response to Stress Questionnaire. aChi-square test; bTwo sample t-test. Characteristic
MDD (n = 31)
HC (n = 32)
P value
Gender (F/M)
17/14
17/15
0.657a
Age (years)
20.53 ± 1.78
20.96 ± 1.28
0.434b
Education (years)
13.62 ± 0.76
13.75 ± 0.82
0.611b
Illness duration (years)
0.29 ± 0.20
NA
CES-D
36.90 ± 6.79
17.97 ± 7.93
<0.001b
RSQ-rumination
7.74 ± 1.18
4.65 ± 1.68
<0.001b Table 1. Demographic and clinical characteristics for subjects. Rumination and Default Mode
Network Subsystems Connectivity
in First-episode, Drug-Naive Young
Patients with Major Depressive
Disorder Previous studies have reported that patients with MDD exhibited abnormal connectivity in pos-
terior, ventral and core DMN subsystems along with reduced interactions from the anterior to the ventral DMN
subsystems35, which suggested the role of these subsystems and the interplay among them in the pathophysiology
of MDD. To date, however, the extent to which altered function and interaction of three DMN subsystems (the
midline core, dMPFC and MTL subsystems) can contribute to the pathophysiology of MDD is not known. y
p
p y
gy
To the best of our knowledge, limited research has investigated neural correlates of rumination and three
DMN subsystems in MDD. In the current study, our first aim was to investigate DMN subsystems connectivity in
first-episode treatment-naïve young adults with MDD. The second goal was to examine the association between
depressive rumination and DMN subsystems connectivity. We hypothesized that MDD patients would exhibit
altered activity in DMN subsystems, which would correlate with depressive rumination. Figure 1. The left panels of (A) and (B) shows half of the averaged connectivity matrices among 11 ROIs of
the DMN for MDD patients (A) and HCs group (B). The color bar indicates the value of correlation strength. The right panels of (A) and (B) showed visualized brain network for the averaged correlation coefficient matrix
threshold by 0.35, where the thickness of each line reflects correlation strength. The left panel of (C) shows the
two-sample t-tests of functional connectivity matrix in MDD compared with HCs group, where the white “*”
denotes significantly altered connectivity (p < 0.05 with false discovery rate (FDR) correction). The color bar
indicates t value. The right panel of (C) shows visualized brain network for the two-sample t-tests, where the red
and blue lines respectively show increased and decreased functional connectivity, and the thickness of each line
reflects t value. Figure 1. The left panels of (A) and (B) shows half of the averaged connectivity matrices among 11 ROIs of
the DMN for MDD patients (A) and HCs group (B). The color bar indicates the value of correlation strength. The right panels of (A) and (B) showed visualized brain network for the averaged correlation coefficient matrix
threshold by 0.35, where the thickness of each line reflects correlation strength. The left panel of (C) shows the
two-sample t-tests of functional connectivity matrix in MDD compared with HCs group, where the white “*”
denotes significantly altered connectivity (p < 0.05 with false discovery rate (FDR) correction). The color bar
indicates t value. The right panel of (C) shows visualized brain network for the two-sample t-tests, where the red
and blue lines respectively show increased and decreased functional connectivity, and the thickness of each line
reflects t value. Figure 1. The left panels of (A) and (B) shows half of the averaged connectivity matrices among 11 ROIs of
the DMN for MDD patients (A) and HCs group (B). The color bar indicates the value of correlation strength. The right panels of (A) and (B) showed visualized brain network for the averaged correlation coefficient matrix
threshold by 0.35, where the thickness of each line reflects correlation strength. The left panel of (C) shows the
two-sample t-tests of functional connectivity matrix in MDD compared with HCs group, where the white “*”
denotes significantly altered connectivity (p < 0.05 with false discovery rate (FDR) correction). The color bar
indicates t value. Results Demographic and clinical data comparisons. The demographic and clinical data were summarized
in Table 1. There was no significant difference between the MDD and HCs groups in gender, age and years of
education (Table 1). Compared with HCs group, MDD patients showed higher levels of Center for Epidemiologic
Studies Depression (CES-D) (t = 10.030, p < 0.001) and rumination of Response to Stress Questionnaire (RSQ-
rumination) (t = 6.980, p < 0.001). Differences in functional connectivity in 11 regions of interest (ROIs). The average half of 11 × 11
functional connectivity matrices is displayed for both MDD (Fig. 1A) and HCs (Fig. 1B) groups with a threshold
of 0.3534. The MDD patients showed similar connectivity patterns among 11 ROIs as HCs group. As illustrated
in Fig. 1C, compared with HCs, MDD patients showed increased connectivity in dMPFC-TempP, TPJ-LTC,
TPJ-PHC, LTC- PHC, TempP-vMPFC, TempP-pIPL, TempP-Rsp and TempP-PHC (p < 0.05 with false discov-
ery rate (FDR) correction). Of all, the dMPFC-TempP and TPJ-LTC were within-system connectivity in the
dMPFC subsystem, whereas the others were inter-system connectivity between the dMPFC and MTL subsystems. Meanwhile, the decreased inter-system interactions were reported in MDD patients between the midline core and
dMPFC subsystems, including aMPFC-dMPFC and PCC-dMPFC (p < 0.05 with FDR correction). Differences in functional connectivity of DMN subsystems. The average within-system and
inter-system connectivity of DMN subsystems were respectively exhibited in Figs 2 and 3 in MDD and HCs
groups. Relative to HCs group, the patients with MDD exhibited significantly increased within-system con-
nectivity in dMPFC subsystem (t = 5.155, p < 0.001), but not in the midline core (t = 0.408, p = 0.685) or MTL
subsystem (t = 1.835, p = 0.072). Additionally, the significantly increased inter-system connectivity of DMN Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 2 www.nature.com/scientificreports/ www.nature.com/scientificreports/ The right panel of (C) shows visualized brain network for the two-sample t-tests, where the red
and blue lines respectively show increased and decreased functional connectivity, and the thickness of each line
reflects t value. Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 3 www.nature.com/scientificreports/ Figure 2. Bar diagrams of average within-system functional connectivity of DMN subsystems in MDD and
HCs group. Figure 2. Bar diagrams of average within-system functional connectivity of DMN subsystems in MDD and
HCs group. Figure 3. Bar diagrams of average inter-system functional connectivity of DMN subsystems in MDD and
HCs group. Figure 2. Bar diagrams of average within-system functional connectivity of DMN subsystems in MDD and
HCs group. Figure 3. Bar diagrams of average inter-system functional connectivity of DMN subsystems in MDD and
HCs group. Figure 3. Bar diagrams of average inter-system functional connectivity of DMN subsystems in MDD and
HCs group Figure 3. Bar diagrams of average inter-system functional connectivity of DMN subsystems in MDD and
HCs group. Figure 4. Significantly positive correlations between RSQ-rumination score and DMN subsystem
connectivity in patients with MDD (p < 0.05). Figure 4. Significantly positive correlations between RSQ-rumination score and DMN subsystem
connectivity in patients with MDD (p < 0.05). subsystems was reported in dMPFC-MTL interaction in MDD patients (t = 2.039, p = 0.048), but not in the m
line core-dMPFC (t = 1.126, p = 0.267) or midline core-MTL interaction (t = 0.195, p = 0.846). Correlation analysis. After controlling for CES-D score, age, education and illness duration as covariates
RSQ-rumination scores were positively correlated with functional connectivity in patients with MDD, including
MPFC-TempP (r = 0.382, p = 0.038) and LTC-PHC (r = 0.416, p = 0.020) (Fig. 4). Correlation analysis. After controlling for CES-D score, age, education and illness duration as covariates,
RSQ-rumination scores were positively correlated with functional connectivity in patients with MDD, including
dMPFC-TempP (r = 0.382, p = 0.038) and LTC-PHC (r = 0.416, p = 0.020) (Fig. 4). Discussion To date, few studies have focused on the neural activity of DMN subsystems in MDD patients, although previous
work has investigated different aspects of the DMN. The present study aimed to investigate three DMN subsys-
tems resting-state functional connectivity and the association with rumination in patients with MDD. Four prin-
cipal findings emerged from our study. Firstly, we found significantly increased within-system connectivity in the Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 4 www.nature.com/scientificreports/ dMPFC subsystem in patients with MDD (dMPFC-TempP and TPJ-LTC). Secondly, the increased inter-system
connectivity was revealed between the dMPFC and MTL subsystems in MDD patients (TPJ-PHC, LTC-PHC,
TempP-vMPFC, TempP-pIPL, TempP-Rsp and TempP-PHC). Thirdly, the decreased connectivity was observed
between the midline core and dMPFC subsystem in individuals with MDD (aMPFC-dMPFC and PCC-dMPFC). Finally, correlation analysis revealed that the connectivity of dMPFC-TempP and LTC-PHC were positively cor-
related with depressive rumination in the MDD group.i p
g
p
In present study, we found significantly greater connectivity in the dMPFC subsystem in MDD group, includ-
ing dMPFC-TempP and TPJ-LTC. These observations might reflect resting-state hyperconnectivity in the dMPFC
subsystem in MDD patients. The dMPFC subsystem is thought to play more of a social-reflective role, allowing
individuals to infer the mental states of other people and reflect on their own mental states14,31,36. In particular,
as the key brain region in Mayberg’s model of MDD37,38, the dMPFC reportedly is involved in the process of
evaluating whether stimuli are self-referential39–41 and is linked to a variety of cognitive functions, appraisal and
the expression of negative emotion42. The neuropathological and neurochemical studies in major depression con-
sistently reported abnormalities in the dMPFC subsystem43–45. Our findings of positive correlations between the
rumination score and connectivity of dMPFC-TempP and LTC-PHC further support the view that the dMPFC
subsystem is involved in self-abnormalities in MDD. Our study provides novel evidence of the notion that the
dMPFC subsystem abnormalities in MDD might reflect the ruminative nature of MDD patients and further
suggests that the resting-state signal in the dMPFC subsystem may be a marker for a ruminative response style
in depression. p
Furthermore, increased inter-system connectivity was observed between the dMPFC and MTL subsystems
in MDD patients relative to HCs. It has been suggested that these two subsystems interact when individuals are
left to think alone and undisturbed46. Discussion In such cases, people tend to engage in self-relevant internal cognitive pro-
cesses, predominantly about significant past and future events31. The increased inter-system connectivity between
the dMPFC and MTL subsystems in our study showed stronger self-relevant neural activity in MDD patients,
which might be related to self-abnormalities in MDD. Consistent with this hypothesis, the following correlation
results indicated that a positive correlation existed between LTC-PHC connectivity with rumination score. As
far as we know, no study has investigated the relationship between rumination score and interaction within the
dMPFC and MTL subsystems in subjects with MDD. Our findings provide new pathophysiological evidence
that the interaction between the dMPFC and MTL subsystems may contribute to the ruminative response style
in MDD, which is consistent with the previous studies mentioned above46. Our results support the notion that
abnormal interaction between DMN subsystems might be involved in the pathophysiology of self-relevant symp-
toms in MDD. In MDD patients, we detected decreased connectivity between the midline core and dMPFC subsystem,
including aMPFC-dMPFC and PCC-dMPFC. Evidence from anatomical and imaging studies has revealed that
both aMPFC and PCC have extensive connectivity with the dMPFC subsystem47,48. It has been suggested that
midline core is critical to making self-relevant, affective decisions, which is strongly correlated with the dMPFC
subsystem, especially in affective self-referential cognition31. The current finding of decreased inter-system con-
nectivity might indicate impaired interaction between the midline core and dMPFC subsystem. It is worthy of
mention that the result relating to the diminished connectivity of PCC-dMPFC is consistent with our previous
study on first-episode treatment-naïve young adults with MDD49. The results were corroborated by other studies
on resting-state connectivity in MDD50, suggesting decreased interaction between the anterior and posterior
DMN in young people with depression. Nevertheless, increased PCC-dMPFC connectivity has been reported in
late-life depression51–53, which implies that there is a relationship between PCC-dMPFC connectivity and patient
age and medication status. Taken together, all the findings suggest that the pathophysiology of MDD could also
be related to reduced central role of the midline core subsystem in the DMN.hi y
The results of this study must be interpreted in light of its limitation. Discussion Firstly, as we selected first-episode,
medication-naïve young patients with MDD to minimize the confounding influences of chronicity, treatment
or comorbidity, the relatively small sample size may limit the generalizability of our results as well as our ability
to detect more relationships between clinical variables and neuroimaging findings. Secondly, consistent with
previous work31, seed-based analysis with the predefined 11 ROIs was used to calculate DMN subsystems con-
nectivity. However, defining DMN based on data-driven approaches is also worth studying54. Finally, the acqui-
sition parameters with large slices (4 + 1 mm), relatively short scanning time and high TR may have effect on the
consistency of resting-state findings. More optimized acquisition parameters deserve investigation in future work. Conclusions
h In summary, the present study examined the relationship between DMN subsystems connectivity and rumi-
nation in patients with MDD. Relative to HCs MDD patients demonstrated disrupted communication within
DMN subsystems, which supported and extended the widely acknowledged large-scale network dysfunction in
MDD55,56. The MDD patients showed increased within-system connectivity in the dMPFC subsystem, decreased
inter-system connectivity between the midline core and dMPFC subsystem, and increased inter-system connec-
tivity between the dMPFC and MTL subsystems. Importantly, abnormal connectivity in DMN subsystems had
a significant positive correlation with rumination scores in MDD patients. Our findings suggest that disrupted
integration of DMN subsystems may be closely associated with the pathophysiology of MDD patients, which
highlights the importance of DMN subsystems connectivity. Methods Participants. Patients with MDD were recruited from the psychiatric clinic at Second Xiangya Hospital of
Central South University in Changsha, China. Patients with MDD were diagnosed according to the Structured
Clinical Interview for DSM-IV57 by independent assessments of two psychiatrists. All of the patients were Participants.
l
h The following parameters were applied:
repetition time = 2000 ms, echo time = 30 ms, flip angle = 80°, field of view (FOV) = 240 mm × 240 mm,
matrix = 64 × 64, slice thickness = 4 mm, slice gap = 1 mm, number of slices = 32. For each participant, 216 vol-
umes were obtained, and the scan lasted 432 s. High-resolution 3-dimensional (3D) structural images were acquired using a T1-weighted, magnetization-
prepared rapid gradient-echo sequence. The following parameters were applied: repetition time = 1900 ms, echo
time = 2.01 ms, flip angle = 9°, FOV = 256 mm × 256 mm, matrix = 256 × 256, slice thickness = 1 mm, slice
gap = 0 mm, and number of slices = 176. Data preprocessing. Image preprocessing was performed using the Data Processing Assistant for
Resting-State fMRI (DPARSF) professional software62 (http://www.restfmri.net). For each individual participant,
the first 10 functional images were excluded from analysis. Subsequent images were corrected by slice timing and
realigned for head motion. Two MDD patients and one healthy subject were excluded because their translation or
rotation exceeded ±1.5 mm ±1.5°. The individual T1-weighted structural images were coregistered to functional
images. The transformed structural images were then segmented into gray matter, white matter, and cerebro-
spinal fluid and normalized to Montreal Neurologic Institute (MNI) space. These transformation parameters
were also applied to the functional images. The normalized functional images were resampled at a resolution of
3 × 3 × 3 mm3 and spatially smoothed with a 6-mm full width at half maximum Gaussian kernel. The sources of
spurious variance were regressed out including 6 parameters from head-motion correction (Friston 24-parameter
model)63, white matter and cerebrospinal fluid signal. Finally, functional images with linear trend were removed
by temporal bandpass filtering (0.01–0.08 Hz).l y
p
pi
g
In view of the influence of head motion on functional connectivity results64–66, the data was further performed
with the scrubbing method to remove time points affected by head motions. As recommended by Yan et al.66,67,
the framewise displacement (FD) was calculated by translation and rotation parameters of head motion based on
the formula from previous study64. The image frames (FD > 0.5 mm) were identified as bad time points. Along
with the bad time points, 1 preceding and 2 following points were deleted to assure exclusion of motion-related
confounds. Participants.
l
h Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 5 www.nature.com/scientificreports/ experiencing their first episode of depression and had never received medication. Closely matched healthy sub-
jects were recruited through advertisements from several colleges in Changsha. All subjects were right-handed. The shared exclusion criteria for patients and control subjects included any major medical illnesses; clinical diag-
nosis of neurologic trauma; any history of psychiatric disorder in the control subjects or any history of psychiatric
disorder, except major depression, in the MDD patients; any history of substance abuse or alcohol in the past 6
months; and any contraindications to imaging scanning. Finally, 33 patients with MDD and 33 matched HCs
were recruited. Research ethics. The study protocol was approved by Second Xiangya Hospital of Central South University. Written informed consent was obtained from all participants prior to the study, which was approved by the
Institutional Review Board of Second Xiangya Hospital of Central South University for Brain Research. The
methods were conducted in accordance with relevant approved guidelines and regulations. Measures. Depression. Depressive severity was measured using the CES-D scale58, a 20-item self-report
instrument to assess depressive symptoms in the general population. The Chinese version of the CES-D has been
found to have high degrees of reliability and validity59. In this study, the internal consistency of the CES-D was
good (Cronbach’s alpha = 0.93). Rumination. Level of rumination was assessed using the rumination subscale of RSQ in Chinese version60,61,
which focuses on the feelings and thoughts that are associated with negative events. The rumination subscale of
the RSQ exhibited high degrees of reliability and validity in both English and Chinese versions. Data acquisition. Images were obtained using a Siemens Skyra 3T scanner with a standard head coil. Participants wore a standard head coil fitted with foam padding to minimize head movement and diminish scan-
ner noise. During scanning, all participants were required to remain motionless, keep their eyes closed and sys-
tematically try not to think of anything. After scanning, the participants were asked about their statement during
scanning. Resting-state fMRI images were acquired with a single-shot, gradient-recalled echo-planar imaging sequence
oriented parallel to the line of the anterior-posterior commissure. Participants.
l
h No difference was observed for the mean FD between MDD and HCs groups (Mann Whitney U-test,
P = 0.27). For all subjects, the remaining imaging data contained more than 128 volumes. Definition of 11 ROIs. According to previous work31–34, 11 ROIs of the DMN were defined using 8-mm
radius spheres (see Fig. 5 and Table 2), including the anterior MPFC (aMPFC), PCC, dorsal MPFC (dMPFC),
temporo-parietal junction (TPJ), lateral temporal cortex (LTC), temporal pole (TempP), ventral MPFC (vMPFC),
retrosplenial cortex (Rsp), posterior inferior parietal lobule (pIPL), parahippocampal cortex (PHC) and hip-
pocampal formation (HF). The aMPFC and PCC constitute the midline core. The dMPFC subsystem includes the
dMPFC, TPJ, LTC and TempP. The MTL subsystem comprises the vMPFC, pIPL, Rsp, PHC and HF. Functional connectivity analysis. Functional connectivity analysis was performed on the left-lateralized
11 ROIs mentioned above (see Fig. 5 and Table 2) with the Resting-State fMRI Data Analysis Toolkit63. The mean
time series from the voxels within each ROI were extracted. Pearson’s correlations were calculated between any
two nodes of 11 ROIs. The correlation coefficients referred to the functional connectivity strength correspond-
ing to the paired ROIs. The 11 × 11 functional connectivity matrix was obtained for each subject68,69. Fisher’s
r-to-z transformation was applied to normalize the correlation coefficients. Apart from the individual edge Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 6 www.nature.com/scientificreports/ Figure 5. 11 regions of interest (ROIs) in the default mode network (DMN) are depicted with BrainNet
Viewer70, including the midline core denoted by red color, dMPFC subsystem denoted by green color,
and MTL subsystem denoted by blue color. The midline core includes posterior cingulate cortex (PCC) and
anterior medial prefrontal cortex (aMPFC). The dMPFC subsystem includes dorsal medial prefrontal cortex
(dMPFC), temporo-parietal junction (TPJ), lateral temporal cortex (LTC) and temporal pole (TempP). The
MTL subsystem includes ventral medial prefrontal cortex (vMPFC), posterior inferior parietal lobule (pIPL),
retrosplenial cortex (Rsp), parahippocampal cortex (PHC) and hippocampal formation (HF). Figure 5. 11 regions of interest (ROIs) in the default mode network (DMN) are depicted with BrainNet
Viewer70, including the midline core denoted by red color, dMPFC subsystem denoted by green color,
and MTL subsystem denoted by blue color. The midline core includes posterior cingulate cortex (PCC) and
anterior medial prefrontal cortex (aMPFC). The dMPFC subsystem includes dorsal medial prefrontal cortex
(dMPFC), temporo-parietal junction (TPJ), lateral temporal cortex (LTC) and temporal pole (TempP). Participants.
l
h group differences in average connectivity of three DMN subsystems, the two-sample t-tests were applied to
explore the average within-system and inter-system connectivity between patients with MDD and HCs. Correlation analysis. To examine the association of DMN subsystems connectivity with the level of
depressive rumination, linear correlations were calculated in MDD patients between the z-value of all edges and
RSQ-rumination scores with SPSS 18.0 software (IBM SPSS Inc., USA). Given that these analyses were explora-
tory, we used an uncorrected statistical significance level of p < 0.05. Correlation analysis. To examine the association of DMN subsystems connectivity with the level of
depressive rumination, linear correlations were calculated in MDD patients between the z-value of all edges and
RSQ-rumination scores with SPSS 18.0 software (IBM SPSS Inc., USA). Given that these analyses were explora-
tory, we used an uncorrected statistical significance level of p < 0.05. Participants.
l
h The
MTL subsystem includes ventral medial prefrontal cortex (vMPFC), posterior inferior parietal lobule (pIPL),
retrosplenial cortex (Rsp), parahippocampal cortex (PHC) and hippocampal formation (HF). DMN Subsystem Regions
Abbreviation
BA
MNIxyz
midline core
Anterior medial prefrontal cortex
aMPFC
10, 32
−6, 52, −2
Posterior cingulate cortex
PCC
23,31
−8, −56, 26
dMPFC self system
Dorsal medial prefrontal cortex
dMPFC
9, 32
0, 52, 26
Temporal parietal junction
TPJ
40, 39
−54, −54, 28
Lateral temporal cortex
LTC
21, 22
−60, −24, −18
Temporal pole
TempP
21,38
−50, 14, −40
MTL memory system
Ventral medial prefrontal cortex
vMPFC
11, 24, 25, 32
0, 26, −18
Posterior inferior parietal lobule
pIPL
39
−44, −74, 32
Retrosplenial cortex
Rsp
29, 30, 19
−14, −52, 8
Parahippocampal cortex
PHC
20, 36, 19
−28, −40, −12
Hippocampal formation
HF
20, 36
−22, −20, −26
Table 2. Coordinates of 11 regions of interest (ROIs) in the default mode network (DMN). BA: Brodmann
area; MNI: Montreal Neurological Institute; MTL: medial temporal lobe Table 2. Coordinates of 11 regions of interest (ROIs) in the default mode network (DMN). BA: Brodmann
area; MNI: Montreal Neurological Institute; MTL: medial temporal lobe. corresponding to one paired ROIs, the average connectivity of all edges is another indicator to represent the
global functional connectivity strength. Then, the average within-system and inter-system connectivity of three
DMN subsystems for each subject were calculated48. The average within-system connectivity is defined as the
mean of the sum of all edges in one subsystem, while the average inter-system functional connectivity is referred
to the mean of the sum of functional connectivity strength of all edges between two subsystems. Statistical analysis. To examine group differences in functional connectivity of 11 ROIs, the two-sample
t-tests were performed on half of 11 × 11 correlation coefficients matrices of MDD and HCs groups due to sym-
metry. The statistical significance level was set at p < 0.05 (FDR correction). The results with the undirected edges
and 11 nodes were laid out in the BrainNet Viewer software70 (http://www.nitrc.org/projects/bnv/). To investigate Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 7 www.nature.com/scientificreports/ group differences in average connectivity of three DMN subsystems, the two-sample t-tests were applied to
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collected data. Q.Z. and H.S. analyzed the results. X.Z. and Q.Z. wrote the main manuscript text. All authors
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50. Van, T.o.l. et al. Local cortical thinning links to resting-state disconnectivity in major depressive disorder. Psychol. Med. 44,
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52. Wu, M. et al. Default-mode network connectivity and white matter burden in late-life depression. Psychiatry Res. 194, 39–46 (2011). References Modulating dysfunctional limbic-cortical circuits in depression: towards development of brain-based algorithms for 37. Mayberg, H. S. Limbic-cortical dysregulation: a proposed model of depression. J Neuropsychiatry Clin Neurosci. 9, 471–481 (1997). 38. Mayberg, H. S. Modulating dysfunctional limbic-cortical circuits in depression: towards development of brain-based algorithms for
diagnosis and optimized treatment. Br. Med. Bull. 65, 193–207 (2003). 37. Mayberg, H. S. Limbic-cortical dysregulation: a proposed mo Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 8 Scientific RePorTS | 7:43105 | DOI: 10.1038/srep43105 Additional Informationi Competing financial interests: The authors declare no competing financial interests. Competing financial interests: The authors declare no competing financial interests. How to cite this article: Zhu, X. et al. Rumination and Default Mode Network Subsystems Connectivity in
First-episode, Drug-Naive Young Patients with Major Depressive Disorder. Sci. Rep. 7, 43105; doi: 10.1038/
srep43105 (2017). How to cite this article: Zhu, X. et al. Rumination and Default Mode Network Subsystems Connectivity in
First-episode, Drug-Naive Young Patients with Major Depressive Disorder. Sci. Rep. 7, 43105; doi: 10.1038/
srep43105 (2017). Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and
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English
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Myocardial Infarction-induced N-terminal Fragment of Cardiac Myosin-binding Protein C (cMyBP-C) Impairs Myofilament Function in Human Myocardium
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Journal of biological chemistry/The Journal of biological chemistry
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Myocardial Infarction-induced N-terminal Fragment of
Cardiac Myosin-binding Protein C (cMyBP-C) Impairs
Myofilament Function in Human Myocardium* y
y
Received for publication,December 9, 2013, and in revised form, January 24, 2014 Published, JBC Papers in Press,February 7, 2014 Namthip Witayavanitkul‡, Younss Ait Mou‡, Diederik W. D. Kuster‡, Ramzi J. Khairallah‡, Jason Sarkey‡,
Suresh Govindan‡, Xin Chen§, Ying Ge§, Sudarsan Rajan¶, David F. Wieczorek¶, Thomas Irving,
Margaret V. Westfall**, Pieter P. de Tombe‡1, and Sakthivel Sadayappan‡1,2
From the ‡Department of Cell and Molecular Physiology, Health Sciences Division, Loyola University Chicago, Maywood, Illinois
60153, the §Human Proteomics Program, School of Medicine and Public Health, University of Wisconsin-Madison, Madison,
Wisconsin 53706, the ¶Department of Molecular Genetics, Biochemistry, and Microbiology, University of Cincinnati College of
Medicine, Cincinnati, Ohio 45267, the Department of Biological and Chemical Sciences, Illinois Institute of Technology, Chicago,
Illinois 60616, and the **Department of Cardiac Surgery, University of Michigan, Ann Arbor, Michigan 48109 From the ‡Department of Cell and Molecular Physiology, Health Sciences Division, Loyola University Chicago, Maywood, Illinois
60153, the §Human Proteomics Program, School of Medicine and Public Health, University of Wisconsin-Madison, Madison,
Wisconsin 53706, the ¶Department of Molecular Genetics, Biochemistry, and Microbiology, University of Cincinnati College of
Medicine, Cincinnati, Ohio 45267, the Department of Biological and Chemical Sciences, Illinois Institute of Technology, Chicago,
Illinois 60616, and the **Department of Cardiac Surgery, University of Michigan, Ann Arbor, Michigan 48109 Background: Myocardial infarction (MI) leads to proteolytic cleavage of cMyBP-C (hC0C1f) and decreased contractility. Results: hC0C1f can incorporate into the human cardiac sarcomere, depressing force generation and increasing tension cost. Conclusion: Interaction between hC0C1f and both actin and -tropomyosin causes disruption of intact cMyBP-C function. Significance: Proteolytic cleavage of cMyBP-C is sufficient to cause contractile dysfunction following MI. Myocardial infarction (MI) is associated with depressed car-
diac contractile function and progression to heart failure. Car-
diac myosin-binding protein C, a cardiac-specific myofilament
protein, is proteolyzed post-MI in humans, which results in an
N-terminal fragment, C0-C1f. The presence of C0-C1f in cul-
tured cardiomyocytes results in decreased Ca2 transients and
cell shortening, abnormalities sufficient for the induction of
heart failure in a mouse model. However, the underlying mech-
anisms remain unclear. Here, we investigate the association
between C0-C1f and altered contractility in human cardiac
myofilaments in vitro. To accomplish this, we generated recom-
binant human C0-C1f (hC0C1f) and incorporated it into per-
meabilized human left ventricular myocardium. Mechanical
properties were studied at short (2 m) and long (2.3 m) sar-
comere length (SL). * Thisworkwassupported,inwholeorinpart,byNationalInstitutesofHealth
Grants R01HL105826 and K02HL114749 (to S. S.); HL007692 (to J. S.);
HL101297, HL75494, and HL62426 (to P. P. d. T.); 2P41RR008630-17 and
9P41GM103622–17 (to T. I.); and HL096971 (to Y. G.). This work was also
supported
by
American
Heart
Association
Midwest
Fellowships
11POST5260038
(to
Y.
A.),
13POST14720024
(to
S.
G.),
and
13POST17220009 (to D. W. D. K.). Use of the Advanced Photon Source, an
Office of Science User Facility operated for the U.S. Department of Energy
Office of Science by Argonne National Laboratory, was supported by the
U.S. Department of Energy under Contract DE-AC02-06CH11357. The Bio-
CAT facility is supported by grants from the National Institute of General
Medical Sciences of the National Institutes of Health (9 P41 GM103622-18).
Author’s Choice—Final version full access.
1 1 The abbreviations used are: MI, myocardial infarction; cMyBP-C, cardiac
myosin-binding protein-C; I/R, ischemia/reperfusion; -TM, -tropomyo-
sin; SL, sarcomere length; IAF, 5-iodoacetamidofluorescein. THE JOURNAL OF BIOLOGICAL CHEMISTRY VOL. 289, NO. 13, pp. 8818–8827, March 28, 2014
© 2014 by The American Society for Biochemistry and Molecular Biology, Inc.
Published in the U.S.A. THE JOURNAL OF BIOLOGICAL CHEMISTRY VOL. 289, NO. 13, pp. 8818–8827, March 28, 2014
© 2014 by The American Society for Biochemistry and Molecular Biology, Inc. Published in the U.S.A. THE JOURNAL OF BIOLOGICAL CHEMISTRY VOL. 289, NO. 13, pp. 8818–8827, March 28, 2014
© 2014 by The American Society for Biochemistry and Molecular Biology, Inc. Published in the U.S.A. Author’s Choice 2 To whom correspondence should be addressed: Dept. of Cell and Molecular
Physiology, Health Sciences Division, Loyola University of Chicago, 2160 S.
First Ave., Maywood, IL 60153. Tel.: 708-216-7994; Fax: 708-216-6308;
E-mail: ssadayappan@lumc.edu. EXPERIMENTAL PROCEDURES Human
Samples—Non-age/sex-matched,
deidentified
hearts were procured via the National Disease Research Inter-
change. Tissue procurement and processing was approved by
the University of Michigan Institutional Review Board, and the
Institutional Review Board at Loyola University Chicago
approved the protocol for the use of deidentified human donor
hearts. Prior to explant, hearts were flushed with ice-cold car-
dioplegia solution and arrived on ice 12 h postexplant. Sec-
tions of left ventricular tissue were then immediately frozen in
liquid N2 and stored at 80 °C until use. Top-Down Analysis by Mass Spectrometry—His-tagged
recombinant hC0C1f proteins were expressed in Escherichia
coli and purified as described previously (4, 15). Recombinant
hC0C1f (20–100 g) was desalted by offline reverse phase pro-
tein trap (Michrom Bioresources). The samples were intro-
duced to the mass spectrometer via an automated chip-based
nano-electrospray ionization source (Triversa NanoMate;
Advion BioSciences, Ithaca, NY) with a spray voltage of 1.2–1.6
kV versus the inlet of the mass spectrometer, resulting in a flow
of 50–200 nl/min. Intact protein molecular ions were analyzed
using 7T linear ion trap/FTICR (LTQ FT Ultra) hybrid MS
(Thermo Scientific Inc., Bremen, Germany) as described previ-
ously (17). Up to 1,500 transients were averaged per spectrum
to ensure high quality spectra. All FTICR spectra were pro-
cessed with in-house developed MASH Suite software (version
1.0), using the THRASH algorithm with a signal to noise
threshold of 3 and fit factor of 60% and validated manually. Single Permeabilized Cardiac Myocyte Preparation—Single
skinned cells were prepared as previously described (18, 19) with
adaptations to allow the use of human tissue. Briefly, frozen
human left ventricle samples were thawed in ice-cold relaxing
solution containing 10 mM EGTA, 10 mM creatine phosphate, 10
units/mlcreatinekinase,100mMN,N-bis(2-hydroxyethyl)taurine;
N,N-Bis-(2-hydroxyethyl)-2-aminoethanesulfonic acid, 6.3 mM
ATP, 6.48 mM magnesium chloride, 49.76 mM potassium propio-
nate, and protease inhibitors mixture (Sigma-Aldrich) at pH 7.0. Single cells were obtained by mechanically homogenizing left ven-
tricular tissues for 3 s at 10,000 rpm using a homogenizer (Power-
Gen 700D; Fisher Scientific). The homogenate was centrifuged at
120 g for 1 min, and the resulting pellet was filtered to remove
any unhomogenized tissue. The run-through was washed with
fresh ice-cold relaxing solution and resuspended in relaxing solu-
tion containing 1% Triton X-100 for 30 min at room temperature
under agitation to permeabilize the cells. N-terminal Fragment of cMyBP-C Impairs Myofilament Function myosin function by the presence of C0-C1f fragments in the
sarcomere, including its interaction with thick and thin fila-
ment proteins and its effects on ATPase activity, myofilament
Ca2 sensitivity, and contractility. myosin function by the presence of C0-C1f fragments in the
sarcomere, including its interaction with thick and thin fila-
ment proteins and its effects on ATPase activity, myofilament
Ca2 sensitivity, and contractility. cross-bridge cycling kinetics and tension cost, effectively break-
ing the interaction between cMyBP-C and actomyosin. Further,
we establish that the effects of hC0C1f occur through direct
interaction with the thin filament proteins actin and -TM. y
y
The C0-C1f fragment contains the cardiac-specific C0
domain, the proline-alanine (Pro-Ala)-rich region, the C1
domain, and the first 17 residues of the M domain (1–271 res-
idues; C0-C1f) of cMyBP-C (2). The N-terminal region of
cMyBP-C interacts with both actin and myosin and is believed
to function as a critical regulator of contraction (5–7). The car-
diac isoform of cMyBP-C differs from the two skeletal isoforms
by having an extra C0 domain at the N-end (8). Thus, the C0
domain is exclusively unique and specific to cMyBP-C. The C0
domain is able to directly bind with both actin (9, 10) and reg-
ulatory light chain of myosin (11), thereby placing the N-termi-
nal region of cMyBP-C in close proximity to the motor domain
of myosin (11). However, the relative binding affinity of the C0
domain to actin was found to be less than that of C1-C2 (9, 10). The C1-M-C2 region binds to the S2 fragment of myosin and
directly influences myofilament Ca2 sensitivity, myofibrillar
tension development, cross-bridge cycling kinetics, and sar-
comere length (SL)-tension relationships (5–7). The C1-M-C2
region also binds to actin, supporting the idea that cMyBP-C
may play a role in the regulation of cardiac contraction by mod-
ulating actin-myosin interaction (9, 10, 12, 13). Furthermore,
the interaction of the C1-M-C2 region of cMyBP-C with actin
and myosin is dynamically regulated in an on-off fashion by
phosphorylation within the M domain (6, 9, 13, 14). Phosphor-
ylation by PKA prevents the cMyBP-C N-region from binding
to actin (9) and myosin S2 (15) or altering myofilament Ca2
sensitivity (16). Interestingly, the cleaved C0-C1f fragment gen-
erated by the proteolytic degradation of cMyBP-C following
ischemia/reperfusion (I/R) injury does not contain these regu-
latory phosphorylation sites (2). N-terminal Fragment of cMyBP-C Impairs Myofilament Function Because the C0-C1f region
retains strong interaction with actin but lacks the phosphory-
lation sites necessary for phosphorylation-dependent on-off
interaction with myosin and actin, we hypothesized that the
presence of the C0-C1f fragments in the sarcomere would alter
actin-myosin interaction by constitutively interacting with thin
filament proteins, such as actin and -tropomyosin (-TM), in
turn having direct consequences on force generation of sar-
comere function. Myocardial Infarction-induced N-terminal Fragment of
Cardiac Myosin-binding Protein C (cMyBP-C) Impairs
Myofilament Function in Human Myocardium* Our data demonstrate that the presence of
hC0C1f in the sarcomere had the greatest effect at short, but not
long, SL, decreasing maximal force and myofilament Ca2 sen-
sitivity. Moreover, hC0C1f led to increased cooperative activa-
tion, cross-bridge cycling kinetics, and tension cost, with greater
effects at short SL. We further established that the effects of hC0C1f occur through direct interaction with actin and -tro-
pomyosin. Our data demonstrate that the presence of hC0C1f in
the sarcomere is sufficient to induce depressed myofilament
function and Ca2 sensitivity in otherwise healthy human donor
myocardium. Decreased cardiac function post-MI may result, in
part, from the ability of hC0C1f to bind actin and -tropomyo-
sin, suggesting that cleaved C0-C1f could act as a poison poly-
peptide and disrupt the interaction of native cardiac myosin-
binding protein C with the thin filament. Myocardial infarction (MI)3 leads to depressed cardiac con-
tractile function and often progresses to heart failure. This pro-
gression is preceded by impaired Ca2 handling, altered myo-
filament phosphorylation, and reduced cross-bridge cycling
rates. Surprisingly, cardiomyocytes in remote noninfarcted
regions, unaffected by ischemia, also display decreased contrac-
tility, despite restoration of blood flow. The mechanisms lead-
ing to this dysfunction are still incompletely understood (1). We have previously shown that MI is associated with catalytic
cleavage of cardiac myosin binding protein C (cMyBP-C), an
important regulator of cardiac contractility (2). Following MI, this proteolytic fragment can be detected in
the ischemic region but is also present in the noninfarcted adja-
cent and remote regions, as well as in nonischemic heart failure
(2–4), presumably after in situ cleavage. Furthermore, cleaved
cMyBP-C is released into the blood following MI, such that its
N-terminal fragment (C0-C1f) can be used as a marker of MI
(2). The C0-C1f fragment has been shown to disrupt myosin-
regulated contraction (3) and potentially lead to global dysfunc-
tion in the heart, not just in the ischemic area. Therefore, the
study aimed to identify the mechanisms underlying altered 8818 8818
JOURNAL OF BIOLOGICAL CHEMISTRY VOLUME 289•NUMBER 13•MARCH 28, 2014 N-terminal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function The aluminum clips holding the fiber were mounted
on hooks, slightly larger than the holes in the clips, so that they
were held securely between a force transducer (KG7; World
Precision Instruments, Sarasota, FL) and a servomotor (Cam-
bridge Technologies model 308B; Aurora Scientific, Aurora,
Canada) mounted on micromanipulators. During the experi-
ment, relaxing solution was continuously perfused through the
chamber using a peristaltic pump. All experiments were done at
room temperature. A highly sensitive CCD detector (Aviex
PCCD 16080) with a 160-mm 80-mm active area and 39-m
pixels was used to collect x-ray patterns. Prior to analysis, x-ray
diffraction patterns were corrected for dark current flat field
and spatial distortions. Spacings on detector images were meas-
ured using the FIT2D program on a UNIX work station. The
accuracy of these measurements reached to 1 pixel of 200–
600 (i.e., 0.5%) or better. Measured spacings of the 1,0 and 1,1
equatorial reflections from the diffraction pattern were con-
verted to d1,0 lattice spacing (thin filament spacing) using
Bragg’s Law, which can then be converted to the interfilament
spacing. Localization of C0-C1f after Exchange in Permeabilized
Cardiomyocytes—Permeabilized
human
cardiac
myocytes
were attached to two needles controlled by a custom microme-
chanics system using optical glue (NOA 63; Norland Products
Inc.) and mounted on a laser scanning confocal microscope
(LSM 410; LSM Tech Inc.). After attachment, nonspecific bind-
ing was blocked by incubating in blocking solution containing
1% (w/v) BSA, 0.1% (w/v) gelatin, and 0.1% (v/v) Tween 20 for
30 min at room temperature. Mouse monoclonal cardiac -ac-
tinin (Sigma; #A7811) was used to stain the Z-disc at a 1:200
dilution in blocking buffer overnight at room temperature. Unbound antibody was removed by washing three times with
PBS. The secondary antibody, goat anti-mouse IgG (Invitrogen;
A11001; 1:100 dilution in blocking buffer) conjugated to a flu-
orescein 488, was then added in blocking solution for 1 h at
room temperature. Unbound antibody was washed away with
PBS. The cell was then attached and imaged at 488/525 excita-
tion/emission. Following the first image, the cell was then incu-
bated with excess of hC0C1f-IAF (30 M) for 20 min at room
temperature. The unbound hC0C1f-IAF was washed three
times with relaxing solution, and the cell was imaged again at
488/525. Both -actinin and hC0C1f-IAF fluorescence signals
were measured simultaneously. N-terminal Fragment of cMyBP-C Impairs Myofilament Function col provided by Molecular Probes. Briefly, labeled hC0C1f-IAF
was prepared by dissolving lyophilized hC0C1f in dissolving
buffer containing 6 M urea, 0.02 M Tris-HCl, and 0.001 M DTT at
pH 8.0 to an initial concentration of 2 mg/ml and incubated at
room temperature until completely dissolved (3 h). The DTT
was removed by dialyzing the solution against a DTT-free
buffer containing 6 M urea, 0.1 M NaCl, 0.02 M Tris-HCl, and
0.001 M EDTA, at pH 7.5, overnight at 4 °C. To ensure high
labeling efficiency, hC0C1f was incubated with a 4 molar
excess of IAF for 8 h at room temperature. IAF covalently binds
via disulfide bonds with cysteine residues. After 8 h, the reac-
tion was stopped by adding 1 mM DTT. To remove unbound
IAF, the solution was dialyzed against a postlabeling buffer con-
taining 6 M urea, 1 M NaCl, 0.02 M Tris-HCl, 0.005 M MgCl2, and
0.001 M DTT at pH 8.0, overnight at 4 °C. Finally, the solution
was dialyzed against relaxing solution overnight at 4 °C. The
stock solution was aliquoted (10 l) and stored at 80 °C. Labeling efficiency was based on the relative molar concentra-
tions of dye/protein, where the dye concentration was deter-
mined spectroscopically by absorbance at 492 nm using a molar
extinction coefficient of 80,000–85,000 M1 cm1, and the
protein concentration was determined by the Bradford
method. Å wavelength). All flight paths were evacuated, except for a
small gap around the sample chamber itself (1 mm down-
stream, 2 mm upstream of the sample). The beam size at the
sample position was collimated to about 0.4 by 0.8 mm and
about 0.065 by 0.15 mm at the detector and contained a maxi-
mum incident flux of 3 1012 photons/s. Permeabilized
human cardiac tissue was clipped using aluminum T-clips and
mounted in a small trough with dimensions of 0.8 mm wide
40 mm long 5 mm deep. The bottom of the trough was a glass
coverslip allowing the muscle to be viewed by a long working
distance (4 mm) 4 objective and also allowed for sarcomere
length measurement via laser diffraction (21). One end of the
trough expanded into a larger reservoir for mounting the fiber. The slides of the trough were hollowed out, allowing the x-ray
beam to pass through the fiber via KaptonTM windows 0.0005
inch thick. N-terminal Fragment of cMyBP-C Impairs Myofilament Function The resulting image was then
subtracted from the -actinin staining to reveal the localization
of the hC0C1f-IAF. The experiment was also repeated, starting
with hC0C1f-IAF incubation and followed by -actinin
staining. p
g
Simultaneous
Measurement
of
Isometric
Tension
and
ATPase Activity—Permeabilized human cardiac tissue was
obtained by adapting previously described methods (19). Our
novel approach consisted of cutting the frozen tissue into
2–4-mm pieces (0.04 g) in ice-cold relaxing solution, fol-
lowed by homogenizing at low speed (1,000 rpm, 3 s) three
times (Power Gen 700D; Fisher Scientific) in relaxing solution. The preparation was allowed to settle, and the supernatant was
discarded, after which the tissue was resuspended in relaxing
solution. The tissue was permeabilized overnight in 1% Triton
X-100 (Amresco) at 4 °C, which served to remove cell mem-
branes and intracellular membrane-bound structures. The tis-
sues were then extensively washed in fresh ice-cold relaxing
solution, stored on ice, and used within 8 h. The strips of fibers
were selected for appearance under a dissecting microscope. They were then attached to a force transducer (KG4A; World
Precision Instruments, Sarasota, FL) and high speed length
controller (model 315C; Aurora Scientific) using aluminum
T-Clips. Muscle dimensions were determined using an ocular
micrometer mounted in the dissection microscope (resolution,
10 m). Sarcomere length was measured in the passive
relaxed condition by laser diffraction as previously described
(21), adjusted to 2 m and then 2.3 m. The skinned fibers were Simultaneous
Measurement
of
Isometric
Tension
and
ATPase Activity—Permeabilized human cardiac tissue was
obtained by adapting previously described methods (19). Our
novel approach consisted of cutting the frozen tissue into
2–4-mm pieces (0.04 g) in ice-cold relaxing solution, fol-
lowed by homogenizing at low speed (1,000 rpm, 3 s) three
times (Power Gen 700D; Fisher Scientific) in relaxing solution. The preparation was allowed to settle, and the supernatant was
discarded, after which the tissue was resuspended in relaxing
solution. The tissue was permeabilized overnight in 1% Triton
X-100 (Amresco) at 4 °C, which served to remove cell mem-
branes and intracellular membrane-bound structures. The tis-
sues were then extensively washed in fresh ice-cold relaxing
solution, stored on ice, and used within 8 h. The strips of fibers
were selected for appearance under a dissecting microscope. They were then attached to a force transducer (KG4A; World
Precision Instruments, Sarasota, FL) and high speed length
controller (model 315C; Aurora Scientific) using aluminum
T-Clips. EXPERIMENTAL PROCEDURES The resulting skinned
cells were washed three times with ice-cold relaxing solution and
suspended in 1.5 ml of fresh ice-cold relaxing solution. The cells
were kept on ice and used the same day. We have previously shown that expression of C0-C1f protein
in neonatal and adult cardiomyocytes induces contractile dys-
function (4). Furthermore, transgenic mice expressing this
fragment display sarcomere dysgenesis, increased fibrosis, and
impaired contractility, leading to the development of heart fail-
ure (3), abnormalities suggesting that cleaved C0-C1f could act
as a poison polypeptide. We hypothesize that C0-C1f interferes
with the binding of native cMyBP-C to thick and thin filaments,
thereby altering its regulation of actomyosin interaction. This
would then result in the depressed myocardial function
observed in post-MI and in heart failure. Here, to determine the
impact of C0-C1f on sarcomere function, we applied human
C0-C1f (hC0C1f) recombinant protein fragment to permeabi-
lized donor human myocardium and assayed force-Ca2/
force-ATPase relationships, length-dependent activation, and
cross-bridge cycling kinetics. We show that hC0C1f increases 5-Iodoacetamidofluorescein Labeling of hC0C1f—To deter-
mine whether the hC0C1f correctly localized to the sarcomere
after incubation, hC0C1f was labeled with a thiol-sensitive
maleimide fluorophore (5-iodoacetamidofluorescein (IAF)). The labeling procedure was conducted according to the proto- MARCH 28, 2014•VOLUME 289•NUMBER 13 8819 JOURNAL OF BIOLOGICAL CHEMISTRY MARCH 28, 2014•VOLUME 289•NUMBER 13 N-terminal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function Isometric tension and ATPase activity
were measured at various levels of Ca2 activation as previously
described (22, 23). Briefly, the isolated muscle was exposed to a
range of calcium solutions obtained by proportional mixing of
activating and relaxing solutions, and the force generated and
ATP consumed were measured simultaneously during the con-
traction. Activating solution contained 20 mM Ca2-EGTA,
1.55 mM potassium propionate, 6.59 mM MgCl2, 100 mM N,N-
bis(2-hydroxyethyl)taurine; N,N-Bis-(2-hydroxyethyl)-2-ami-
noethanesulfonic acid, 5 mM sodium azide, 1 mM DTT, 10 mM
phosphoenolpyruvate, 0.01 mM oligomycin, 0.1 mM PMSF, and
0.02 mM A2P5, as well as protease inhibitor mixture. Relaxing
solution was identical, except it contained 20 mM EGTA, 21.2
mM potassium propionate, and 7.11 mM MgCl2. The preactivat-
ing solution contained 0.5 mM EGTA, 19.5 mM 1,6-diamino-
hexane-N,N,N,N-tetraacetic acid, and 21.8 mM potassium pro-
pionate. All solutions contained 0.5 mg/ml pyruvate kinase and
0.05 mg/ml lactate dehydrogenase (Sigma) and had an ionic
strength of 180 mM, 5 mM free ATP, and 1 mM free magnesium,
as determined by the Fabiato program (24). Statistical Analysis—Tension- and ATPase-pCa relation-
ships were fit with a modified Hill equation (23, 26, 27), and
stiffness and tension costs were determined by linear fit to the
tension-stiffness and tension-ATPase data, respectively. The
data were analyzed using 1) one-way repeated measures analy-
sis of variance for the dose response of control and hC0C1f on
maximum force development, 2) an unpaired t test for compar-
ing control and hC0C1f, 3) two-way analysis of variance, and 4)
a Holm-Sidak post hoc test with a level of statistical significance
set at p 0.05. The data are presented as means S.E. y
g
ATPase activity was measured by a UV-coupled optical
absorbance enzyme assay (25). Briefly, ATP hydrolysis into
ADP and inorganic phosphate (Pi) inside the fiber was coupled
to the oxidation of NADH to NAD catalyzed by pyruvate
kinase and lactate dehydrogenase. Because NADH absorbs
light at 340 nm and NAD does not, the oxidation of NADH to
NAD, and thus ATP consumption, was determined by meas-
uring the absorbance of UV light at 340 nm at steady state
within the measurement chamber. This absorbance signal was
calibrated by repeated injections of 50 nl of 10 mM ADP into the
measuring chamber. N-terminal Fragment of cMyBP-C Impairs Myofilament Function The ADP injection induced a rapid step
reduction in fluorescence, and the rate of ATP consumption
was calculated by the magnitude of this step from the fluores-
cent decay rate at 340 nm. In addition, the ADP injection served
to confirm that the chemical response time and the bath stir-
ring were adequate. Furthermore, the rate of force redevelop-
ment following a release-restretch maneuver, ktr, was measured
during a final contraction at maximum Ca2 as previously
described (23). Only muscles that maintained greater than 80%
maximal tension were included for analysis. N-terminal Fragment of cMyBP-C Impairs Myofilament Function Briefly, the isolated muscle was exposed to a
range of calcium solutions obtained by proportional mixing of
activating and relaxing solutions, and the force generated and
ATP consumed were measured simultaneously during the con-
traction. Activating solution contained 20 mM Ca2-EGTA,
1.55 mM potassium propionate, 6.59 mM MgCl2, 100 mM N,N-
bis(2-hydroxyethyl)taurine; N,N-Bis-(2-hydroxyethyl)-2-ami-
noethanesulfonic acid, 5 mM sodium azide, 1 mM DTT, 10 mM
phosphoenolpyruvate, 0.01 mM oligomycin, 0.1 mM PMSF, and
0.02 mM A2P5, as well as protease inhibitor mixture. Relaxing
solution was identical, except it contained 20 mM EGTA, 21.2
mM potassium propionate, and 7.11 mM MgCl2. The preactivat-
ing solution contained 0.5 mM EGTA, 19.5 mM 1,6-diamino-
hexane-N,N,N,N-tetraacetic acid, and 21.8 mM potassium pro-
pionate. All solutions contained 0.5 mg/ml pyruvate kinase and
0.05 mg/ml lactate dehydrogenase (Sigma) and had an ionic
strength of 180 mM, 5 mM free ATP, and 1 mM free magnesium,
as determined by the Fabiato program (24). Pulldown Assay—His-tagged recombinant proteins, such as
mouse C0-linker, C0-C1f, C0-C2, C0-C3, and C0mC1f, were
expressed in E. coli and purified using nickel-nitrilotriacetic acid
column chromatography at room temperature as described previ-
ously (4, 15). Total tissue lysates from normal mouse left ventric-
ular myocardium were prepared in radioimmune precipitation
assay buffer (20 mM Tris-HCl, pH 7.5, 150 mM NaCl, 1 mM
Na2EDTA, 1 mM EGTA, 1% Nonidet P-40, 1% sodium deoxy-
cholate, 2.5 mM sodium pyrophosphate, 1 mM -glycerophos-
phate, 1 mM Na3VO4, and 1 g/ml leupeptin) at 1 mg protein/ml. 200 l of lysate was then incubated with 20 l of nickel-nitrilotri-
acetic acid-agarose beads (Qiagen) at 4 °C for 2 h and spun down,
and the pellet was discarded to remove proteins with nonspecific
binding to the beads. The supernatant was then incubated with
either His-tagged cMyBP-C protein fragments or SUMO-His-
tagged -TM and 20 l of beads for 2 h at 4 °C, followed by wash-
ing with Tris buffer (50 mM) containing 0.5% Triton X-100, 100
mMNaCl,10mMMgCl2,0.1mMPMSF,and1proteaseinhibitor. Proteins bound to the beads were eluted with Laemmli sample
buffer (Bio-Rad) and subjected to Western blot analysis. SDS-
PAGE and Western blot analyses were carried out as described
previously (2). incubated with peptide for 5 min in relaxing solution and
another 5 min in preactivating solution before measuring the
force development. The peptide was also present in all working
solutions thereafter. N-terminal Fragment of cMyBP-C Impairs Myofilament Function Muscle dimensions were determined using an ocular
micrometer mounted in the dissection microscope (resolution,
10 m). Sarcomere length was measured in the passive
relaxed condition by laser diffraction as previously described
(21), adjusted to 2 m and then 2.3 m. The skinned fibers were X-ray Diffraction—Experiments were performed using the
BioCAT undulator-based beamline at the Advanced Photon
Source (Argonne National Labs, Lemont, IL) (20). The high
x-ray flux density and low beam divergence delivered by this
instrument are highly advantageous for small angle x-ray stud-
ies of small specimens, such as those described here. Experi-
ments were done using a 3-m distance between the sample and
the detector and with the x-ray beam energy set to 12 keV (1.03 VOLUME 289•NUMBER 13•MARCH 28, 2014 8820 JOURNAL OF BIOLOGICAL CHEMISTRY VOLUME 289•NUMBER 13•MARCH 28, 2014 N-terminal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function FIGURE 1. hC0C1f incorporates into the human cardiac sarcomere. A, recombinant protein was run on SDS-PAGE and SYPRO-RUBY staining to show the
purity of recombinant hC0C1f. B, electrospray ionization-Fourier transform mass spectrometry spectrum of recombinant hC0C1f. C, a skinned stretched
cardiomyocyte stained with -actinin (red), a Z-line-specific protein, and incubated with IAF-labeled hC0C1f (green). The merged image highlights localization
of hC0C1f in the A-band of the sarcomere, and the imaging was performed at SL of 2.1 m. The scale bars represent 5 m. g
y
y FIGURE 1. hC0C1f incorporates into the human cardiac sarcomere. A, recombinant protein was run on SDS-PAGE and SYPRO-RUBY staining to show the
purity of recombinant hC0C1f. B, electrospray ionization-Fourier transform mass spectrometry spectrum of recombinant hC0C1f. C, a skinned stretched
cardiomyocyte stained with -actinin (red), a Z-line-specific protein, and incubated with IAF-labeled hC0C1f (green). The merged image highlights localization
of hC0C1f in the A-band of the sarcomere, and the imaging was performed at SL of 2.1 m. The scale bars represent 5 m. incubated with peptide for 5 min in relaxing solution and
another 5 min in preactivating solution before measuring the
force development. The peptide was also present in all working
solutions thereafter. Isometric tension and ATPase activity
were measured at various levels of Ca2 activation as previously
described (22, 23). RESULTS This concentration approximates the EC50 of inhibition in
force induced by hC0C1f. At 2-m SL, the fragment induced a
30% reduction in maximal force (Fmax) (Fig. 3, B and C) and a
50% increase in myofilament activation cooperativity, as
reported by the Hill coefficient (nH) (Fig. 3D). Remarkably,
hC0C1f did not affect Fmax or nH at long SL. Additionally,
hC0C1f induced a rightward shift of the force-pCa relationship
at both SLs, indicating decreased myofilament Ca2 sensitivity
(20% reduction of EC50 at both SL) (Fig. 3E). Despite
decreased myofilament Ca2 sensitivity, length-dependent
activation was enhanced with an increased Fmax (difference
between maximum force at long and short sarcomere lengths;
13.20 0.41 versus 20.56 1.70, control versus hC0C1f, p
0.05). Although the EC50 (difference between myofilament
Ca2 sensitivity at long and short sarcomere lengths) increased,
this change was not significant (0.28 0.06 versus 0.41 0.07,
control versus hC0C1f, p
0.07). rescently
labeled
anti--actinin
antibodies. IAF-labeled
hC0C1f was present specifically between Z-discs, likely on the
thin filaments (Fig. 1C). This is similar to the localization of
endogenous cMyBP-C. To determine whether hC0C1f incor-
poration disrupted sarcomere structure, x-ray diffraction
experiments were performed on permeabilized human donor
myocardium at short (2 m) and long (2.3 m) sarcomere
length (SL) (Fig. 2, A and B). As expected, spacing between each
thin and thick filament was reduced upon stretching. Further-
more, incubation with hC0C1f did not significantly change
interfilament spacing at either SL (Fig. 2B), suggesting that the
presence of hC0C1f in the sarcomere does not adversely affect
sarcomere lattice spacing. hC0C1f Disrupts Steady-state Force Development—To eval-
uate the effect of hC0C1f on myocardial contractile properties,
we developed a novel method to allow for the use of frozen
donor heart tissue. The tissue was cut into thin strips to resem-
ble trabeculae and then homogenized, permeabilized, and
attached to a length controller and force transducer. The sam-
ples were illuminated with a laser to reveal a diffraction pattern
allowing SL determination (Fig. 2C). To establish a dose-re-
sponse curve, permeabilized myocardium was incubated at pCa
5.85 (pCa50) with increasing concentrations of hC0C1f. As a
control, the muscle was also incubated with buffer alone or a
control protein, egg albumin, which has a molecular mass sim-
ilar to that of hC0C1f (44 kDa). RESULTS Purification and Characterization of hC0C1f—Recombinant
hC0C1f was generated, purified, and analyzed with SDS-PAGE
showing a pure protein that migrates at 40 kDa (Fig. 1A). To
further characterize the recombinant hC0C1f, high resolution
mass spectrometry analysis was performed, confirming the
amino acid sequences and a molecular mass of 31,989.20 dal-
tons (Fig. 1B). To determine hC0C1f incorporation into the
A-band of the sarcomere, permeabilized human ventricular
myocytes were incubated with IAF-labeled hC0C1f and fluo- MARCH 28, 2014•VOLUME 289•NUMBER 13 MARCH 28, 2014•VOLUME 289•NUMBER 13 8821 MARCH 28, 2014•VOLUME 289•NUMBER 13 JOURNAL OF BIOLOGICAL CHEMISTRY FIGURE 2. hC0C1f does not disrupt sarcomere structure. A, representative x-ray diffraction of skinned ventricular fibers with or without hC0C1f. B, thin and
thick filament spacings indicated by d1,0 (thin filament spacing) and d1,1 (thick filament spacing) reflections, respectively. CTRL, control. C, representative laser
diffraction pattern and intensity profile of skinned human ventricular muscle tissue at 2- and 2.3-m SL. Representative images are shown at 5 magnification. N-terminal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function FIGURE 2. hC0C1f does not disrupt sarcomere structure. A, representative x-ray diffraction of skinned ventricular fibers with or without hC0C1f. B, thin and
thick filament spacings indicated by d1,0 (thin filament spacing) and d1,1 (thick filament spacing) reflections, respectively. CTRL, control. C, representative laser
diffraction pattern and intensity profile of skinned human ventricular muscle tissue at 2- and 2.3-m SL. Representative images are shown at 5 magnification. This concentration approximates the EC50 of inhibition in
force induced by hC0C1f. At 2-m SL, the fragment induced a
30% reduction in maximal force (Fmax) (Fig. 3, B and C) and a
50% increase in myofilament activation cooperativity, as
reported by the Hill coefficient (nH) (Fig. 3D). Remarkably,
hC0C1f did not affect Fmax or nH at long SL. Additionally,
hC0C1f induced a rightward shift of the force-pCa relationship
at both SLs, indicating decreased myofilament Ca2 sensitivity
(20% reduction of EC50 at both SL) (Fig. 3E). Despite
decreased myofilament Ca2 sensitivity, length-dependent
activation was enhanced with an increased Fmax (difference
between maximum force at long and short sarcomere lengths;
13.20 0.41 versus 20.56 1.70, control versus hC0C1f, p
0.05). Although the EC50 (difference between myofilament
Ca2 sensitivity at long and short sarcomere lengths) increased,
this change was not significant (0.28 0.06 versus 0.41 0.07,
control versus hC0C1f, p
0.07). RESULTS A dose-dependent reduction
in force at pCa50 was observed with an EC50 (myofilament cal-
cium sensitivity; Ca2 concentration at which half of Fmax is
reached) of inhibition at 4.41 M and maximum inhibition
occurring at 30 M of hC0C1f (Fig. 3A), with no effect on
force development of vehicle or egg albumin. The effects of
hC0C1f on tension development and cross-bridge cycling were
determined by measuring force-pCa relationships in permeabi-
lized human ventricular myocardium in the absence and pres-
ence of 5 M hC0C1f. Given that length-dependent activation is
characterized by an increase in both maximum Ca2-activated
force and Ca2 sensitivity upon an increase in sarcomere
length, all experiments were done at both short and long SL. hC0C1f Alters Cross-bridge Cycling Properties—We also eval-
uated myocardium energy consumption by measuring the ATP
hydrolysis rate in the absence and presence of 5 M hC0C1f. Although no significant difference in maximum ATP hydroly-
sis was observed (Fig. 4A), the fragment significantly altered
stretch-induced reduction of ATP hydrolysis, as suggested by
reduced Max ATPase (Fig. 4B). We also observed a significant
increase in the tension cost with hC0C1f at both SL (Fig. 4, C
and D). The two-way analysis of variance determined that ten-
sion cost decreased with sarcomere length (p
0.0005) and
increased with hC0C1f treatment (p
0.0482; main effect). However, hC0C1f did not alter stretch-induced reduction in
the tension cost (1.06 0.22 versus 1.07 0.26, control versus
hC0C1f, p
0.9936). The rate of tension redevelopment (ktr)
was also significantly increased in the presence of hC0C1f at
both SLs (Fig. 4E). These data are consistent with increased
tension cost and indicate a significant increase in cross-bridge
cycling kinetics induced by hC0C1f. hC0C1f Interacts with the Thin Filament Proteins -TM and
Actin—Previously, we demonstrated that cMyBP-C colocalized
with -TM in the sarcomere (28), and many reports have VOLUME 289•NUMBER 13•MARCH 28, 2014 8822 8822
JOURNAL OF BIOLOGICAL CHEMISTRY URE 3. Altered force-Ca2 relationship and length-dependent activation in the presence of hC0C1f. A, dose-response curve of the inhibitory effect of
0C1f, compared with vehicle and egg albumin, on force development at pCa50 (pCa 5.85) for 2-m SL. CTRL, control. B, maximum force (Fmax) at pCa 4.5 in
nned ventricular fibers treated with hC0C1f at 2- and 2.3-m SL. C, myofilament force-Ca2 relationship (panel i) and force-Ca2 relationship normalized to
ximum force (panel ii). RESULTS D and E, normalized force-Ca2 relationships were fit to a modified Hill equation, and the Hill coefficient (D) and EC50 (E) were
termined. *, p 0.05 control versus hC0C1f; †, p 0.05 2 versus 2.3-m SL within groups. N-terminal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function g
y
p
y FIGURE 3. Altered force-Ca2 relationship and length-dependent activation in the presence of hC0C1f. A, dose-response curve of the inhibitory effect of
hC0C1f, compared with vehicle and egg albumin, on force development at pCa50 (pCa 5.85) for 2-m SL. CTRL, control. B, maximum force (Fmax) at pCa 4.5 in
skinned ventricular fibers treated with hC0C1f at 2- and 2.3-m SL. C, myofilament force-Ca2 relationship (panel i) and force-Ca2 relationship normalized to
maximum force (panel ii). D and E, normalized force-Ca2 relationships were fit to a modified Hill equation, and the Hill coefficient (D) and EC50 (E) were
determined. *, p 0.05 control versus hC0C1f; †, p 0.05 2 versus 2.3-m SL within groups. actomyosin interactions, thereby increasing the probability of
strong cross-bridge formation (Fig. 6, A and B). described its interaction with actin (9, 12). To demonstrate the
direct interaction with -TM and actin, we generated several
cMyBP-C N-terminal recombinant fragments (Fig. 5A) and
used heart protein lysate to pull down interacting partners. Consistent with previous reports (9), our results show a direct
interaction of hC0C1f with actin, but not with myosin (Fig. 5C). More interestingly and for the first time, we show a direct inter-
action of cMyBP-C with -TM via the Pro-Ala-rich linker
region (Fig. 5, C–E). This interaction, but not the interaction
with actin, is aborted following mutation of five proline resi-
dues to alanine (Fig. 5E), confirming that the interaction is
mediated through the Pro-Ala-rich linker region. These results
suggest a direct interaction of cMyBP-C N-region with both
-TM and actin and that disruption of that interaction by
C0-C1f leads to altered cross-bridge cycling. In the present
study, we show that incubation with hC0C1f increased cooper-
ativity, cross-bridge cycling, and ATP consumption. These
results suggest that hC0C1f fragments constantly interact with
thin filament proteins, such as actin and -TM, thereby altering DISCUSSION Catalytic cleavage of cMyBP-C, in human heart failure and
post-MI, has been associated with abnormal contractility; how-
ever, the underlying mechanisms have been unclear (2, 3, 29). Previously, we have shown that C0-C1f is released by catalytic
cleavage of cMyBP-C at a -calpain site and that this release
correlated with cardiac insults, such as I/R and MI (2, 30, 31). Further, overexpression of mouse C0C1f resulted in cell death,
impaired Ca2 handling, altered contractile function of the sar-
comere (4), and, finally, heart failure (3). Using in vitro motility
and laser trap assays, we also previously determined that
C0-C1f is sufficient to reduce actin sliding velocity in a manner
similar to that of either C0-C2 or full-length cMyBP-C (13). Furthermore, we determined that the 17 residues of M domain
in C0-C1f mediate interaction with actin (4, 13). In this study,
we extend these findings to show, for the first time, that the MARCH 28, 2014•VOLUME 289•NUMBER 13 8823 JOURNAL OF BIOLOGICAL CHEMISTRY MARCH 28, 2014•VOLUME 289•NUMBER 13 FIGURE 4. hC0C1f increases cross-bridge cycling kinetics. A, maximum ATPase hydrolysis rate. B, difference in ATPasemax at short and long SL. C, force-
ATPaserelationshipsinvarioustensionranges(binneddata)at2-m(paneli)and2.3-m(panelii)SL,andtheforce-ATPaserelationshipofeachindividualfiber
was fit by linear regression. D, the slope was then calculated to determine the tension cost. E, the rate of force redevelopment (ktr) was determined following
a rapid stretch-release-restretch maneuver. *, p 0.05 control versus hC0C1f. #, p 0.05, 2-m versus 2.3-m SL main effect. †, p 0.05 2-m versus 2.3-m
SL within groups. N-terminal Fragment of cMyBP-C Impairs Myofilament Function minal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function ent Function FIGURE 4. hC0C1f increases cross-bridge cycling kinetics. A, maximum ATPase hydrolysis rate. B, difference in ATPasemax at short and long SL. C, force-
ATPaserelationshipsinvarioustensionranges(binneddata)at2-m(paneli)and2.3-m(panelii)SL,andtheforce-ATPaserelationshipofeachindividualfiber
was fit by linear regression. D, the slope was then calculated to determine the tension cost. E, the rate of force redevelopment (ktr) was determined following
a rapid stretch-release-restretch maneuver. *, p 0.05 control versus hC0C1f. #, p 0.05, 2-m versus 2.3-m SL main effect. †, p 0.05 2-m versus 2.3-m
SL within groups. sue used in these experiments had normal levels of native
cMyBP-C, and simple addition of hC0C1f was enough to dis-
rupt the myofilament properties. DISCUSSION Of course, proteolysis of
cMyBP-C would in itself cause additional dysfunction in the
whole organ, and most likely, following I/R or MI, it is the addi-
tive effect of less cMyBP-C and the presence of hC0C1f that will
lead to contractile dysfunction in the heart. Furthermore, it is
still unclear whether the fragment can penetrate neighboring
cells. At this time, it is not possible to determine whether the
contractile dysfunction seen in the whole heart following I/R or
MI is solely due to hC0C1f acting within the cardiomyocyte or
due to “diffusion” to remote areas. Along the same lines, it is
unclear how long the fragment remains in the cells or attached
to the myofilaments. hC0C1f can be detected in tissue several
hours to several days following I/R and MI, and we do not know
whether there are any mechanisms to actively remove the frag-
ment from the cell. Additionally, it is very difficult to remove
hC0C1f from the myofilaments in vitro because it does not
wash off following removal. proteolytic fragment hC0C1f can incorporate into the human
cardiac sarcomere and that this incorporation is sufficient to 1)
decrease maximum force, 2) increase cooperativity, and 3)
increase tension cost and cross-bridge cycling. We also show
that these effects could be the result of a direct interaction of
hC0C1f with both -TM and actin, inhibiting, in turn, the role
of full-length cMyBP-C. This study provides a direct mecha-
nism that explains, at least in part, the depressed contractile
function observed post-MI in the noninfarcted myocardium. Although no attempt has been made to accurately measure
the concentration of hC0C1f in the myofilaments following MI
or I/R or in heart failure, Western blots done by us and others
shown that hC0C1f is easily detectable in myofilaments and is
of comparable intensity to the remaining cMyBP-C (2–4). Assuming that cMyBP-C is present at 15 M in cardiomyo-
cytes, we estimate that hC0C1f is present at concentrations
ranging from 3 to 15 M, depending on the conditions. In
the present study, we used a concentration of 5 M hC0C1f to
assay the effects of the fragment on myofilament properties. This concentration is well within the physiological range
observed following MI or in heart failure. To provide direct support that N-specific fragments of
cMyBP-C are sufficient to incorporate into the sarcomere, Her-
ron et al. DISCUSSION hC0C1f interacts with the thick and thin filaments. A, SDS PAGE,
followed by Coomassie staining of His tag recombinant proteins containing
the N-terminal domains of cMyBP-C and SUMO-tagged -TM. B, pulldown
assay with -TM (top panel) and C0C2 (bottom panel), followed by Western
blot analysis for cMyBP-C and -TM, respectively. C, pulldown assay with dif-
ferent N proteins of cMyBP-C, followed by Western blot analysis for myosin,
actin, and -TM. D and E, pulldown assay with C0-C1f (D) and C0mC1f (with
mutatedresiduesinthePro-Ala-richregion,E),followedbyWesternblotanal-
ysis for -TM and actin. IB, immunoblotting; IP, immunoprecipitation. ,
,
followed by Coomassie staining of His tag recombinant proteins containing
the N-terminal domains of cMyBP-C and SUMO-tagged -TM. B, pulldown
assay with -TM (top panel) and C0C2 (bottom panel), followed by Western
blot analysis for cMyBP-C and -TM, respectively. C, pulldown assay with dif-
ferent N proteins of cMyBP-C, followed by Western blot analysis for myosin,
actin, and -TM. D and E, pulldown assay with C0-C1f (D) and C0mC1f (with
mutatedresiduesinthePro-Ala-richregion,E),followedbyWesternblotanal-
ysis for -TM and actin. IB, immunoblotting; IP, immunoprecipitation. interaction is lost, and constant interaction occurs. Indeed, we
show that the C0-C1f region interacts with actin, but not myo-
sin, because of the absence of M and C2 domains. Furthermore,
part of the negative effects of hC0C1f could be due to disruption
of the link between the thick and thin filament that native
cMyBP-C provides. sarcomere. However, it remains unclear whether these frag-
ments directly interact with myosin or actin, or both, in a man-
ner that confers stability in the sarcomere. Recent studies dem-
onstrate that cMyBP-C N-regions structurally reach thin
filaments (32, 33) and dynamically interact with actin (12). In
the present study, we confirmed that recombinant C0-C1f pro-
teins were able to diffuse into the sarcomere and specifically
localize on the A-band and I-band in the presence of endoge-
nous cMyBP-C without changing the interfilament space at
both short and longer SL. Interestingly, we did not see any
staining of hC0C1f fragments on the Z-line, suggesting that
interaction of C0-C1f is very specific to thin and thick filament
proteins. However, further studies are needed to define the
exact region of interaction that is necessary in the N-region on
the thick and thin filaments. The N-region of cMyBP-C binds
transiently to actin and the myosin S2 region in a phosphory-
lation-dependent manner to regulate actomyosin interaction
and force generation. DISCUSSION (7) overexpressed either C0-C1 or C0-C2 regions into
neonatal cardiomyocytes and confirmed that these fragments
of cMyBP-C could specifically incorporate in the A-band of the Based on the data presented in this study, it is clear that
cleavage of cMyBP-C is not the most important/only mecha-
nism leading to contractile dysfunction. Indeed, the human tis- 8824
JOURNAL OF BIOLOGICAL CHEMISTRY VOLUME 289•NUMBER 13•MARCH 28, 2014 8824 FIGURE6.A,schematicdiagramofthesarcomerewiththick(titin,myosin,and
cMyBP-C) and thin (-TM, cTn complex, and actin) filament proteins. cMyBP-C
interacts with titin and myosin at its C-terminal, with actin at the C1 and M
domains, myosin at the M domain, and -TM at the Pro-Ala-linker region. B,
C0-C1f disrupts native cMyBP-C interaction with the thin filament by binding
actin and -TM at multiple locations. Our data indicate that the presence of
C0-C1f fragments in the normal sarcomere disrupts thin filament and thick
filament protein interactions, thereby interfering with the regulatory role of
native full-length cMyBP-C and, in turn, resulting in diminished force
generation. l Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function N-terminal Fragment of cMyBP-C Impairs Myofilament Function FIGURE 5. hC0C1f interacts with the thick and thin filaments. A, SDS-PAGE,
followed by Coomassie staining of His tag recombinant proteins containing
the N-terminal domains of cMyBP-C and SUMO-tagged -TM. B, pulldown
assay with -TM (top panel) and C0C2 (bottom panel), followed by Western
blot analysis for cMyBP-C and -TM, respectively. C, pulldown assay with dif-
ferent N proteins of cMyBP-C, followed by Western blot analysis for myosin,
actin, and -TM. D and E, pulldown assay with C0-C1f (D) and C0mC1f (with
mutatedresiduesinthePro-Ala-richregion,E),followedbyWesternblotanal-
ysis for -TM and actin. IB, immunoblotting; IP, immunoprecipitation. N-terminal FIGURE6.A,schematicdiagramofthesarcomerewiththick(titin,myosin,and
cMyBP-C) and thin (-TM, cTn complex, and actin) filament proteins. cMyBP-C
interacts with titin and myosin at its C-terminal, with actin at the C1 and M
domains, myosin at the M domain, and -TM at the Pro-Ala-linker region. B,
C0-C1f disrupts native cMyBP-C interaction with the thin filament by binding
actin and -TM at multiple locations. Our data indicate that the presence of
C0-C1f fragments in the normal sarcomere disrupts thin filament and thick
filament protein interactions, thereby interfering with the regulatory role of
native full-length cMyBP-C and, in turn, resulting in diminished force
generation. FIGURE 5. hC0C1f interacts with the thick and thin filam FIGURE 5. hC0C1f interacts with the thick and thin filaments. A, SDS-PAGE, FIGURE 5. REFERENCES Consistent with previous studies using various N fragments,
such as mouse C0-C1 (5, 7), C1-C2 (6), and C0-C2 (7) of
cMyBP-C, our results with the physiologically relevant hC0C1f
show decreased Fmax and accelerated cross-bridge cycling. Although these studies previously showed increased myofila-
ment Ca2 sensitivity (6, 7), we show decreased EC50 at both
short and long SLs. This difference might be partly explained by
the decreased phosphorylation status of endogenous cMyBP-C
seen in failing hearts, compared with the healthy donor tissue
used in this study. We and others have previously shown that
cMyBP-C phosphorylation is greatly depressed in HF and fol-
lowing MI (2, 34). Moreover, cMyBP-C binding to myosin (31)
and actin is modulated by phosphorylation status, and N-ter-
minal fragments do not bind actin when the cMyBP-C phos-
phorylation level is high (10, 31). Although Herron et al. (7)
determined an increase in Ca2 sensitivity following addition of
C0-C2 fragment, our report shows that the C0-C1f fragment
also contains the first 17 residues of the M-domain, which were
sufficient to interact with actin (4, 13). As opposed to the hC0-
Clf fragment, C0-C2 preserves the link between thick and thin
filaments. Also, C0-C2 has been shown to bind to myosin (10),
but hC0C1f does not, suggesting that cMyBP-C interaction
with myosin S2 is mediated through M and C2 domains of
cMyBP-C. 1. van der Velden, J., Merkus, D., Klarenbeek, B. R., James, A. T., Boontje,
N. M., Dekkers, D. H., Stienen, G. J., Lamers, J. M., and Duncker, D. J. (2004) Alterations in myofilament function contribute to left ventricular
dysfunction in pigs early after myocardial infarction. Circ. Res. 95, e85–95 2. Govindan, S., McElligott, A., Muthusamy, S., Nair, N., Barefield, D., Mar-
tin, J. L., Gongora, E., Greis, K. D., Luther, P. K., Winegrad, S., Henderson,
K. K., and Sadayappan, S. (2012) Cardiac myosin binding protein-C is a
potential diagnostic biomarker for myocardial infarction. J. Mol Cell Car-
diol. 52, 154–164 3. Razzaque, M. A., Gupta, M., Osinska, H., Gulick, J., Blaxall, B. C., and
Robbins, J. (2013) An endogenously produced fragment of cardiac myo-
sin-binding protein C is pathogenic and can lead to heart failure. Circ. Res. 113, 553–561 4. Govindan, S., Sarkey, J., Ji, X., Sundaresan, N. R., Gupta, M. P., de Tombe,
P. P., and Sadayappan, S. (2012) Pathogenic properties of the N-terminal
region of cardiac myosin binding protein-C in vitro. J. Muscle Res. REFERENCES Cell
Motil. 33, 17–30 5. Razumova, M. V., Shaffer, J. F., Tu, A. Y., Flint, G. V., Regnier, M., and
Harris, S. P. (2006) Effects of the N-terminal domains of myosin binding
protein-C in an in vitro motility assay. Evidence for long-lived cross-
bridges. J. Biol. Chem. 281, 35846–35854 6. Harris, S. P., Rostkova, E., Gautel, M., and Moss, R. L. (2004) Binding of
myosin binding protein-C to myosin subfragment S2 affects contractility
independent of a tether mechanism. Circ. Res. 95, 930–936 7. Herron, T. J., Rostkova, E., Kunst, G., Chaturvedi, R., Gautel, M., and
Kentish, J. C. (2006) Activation of myocardial contraction by the N-termi-
nal domains of myosin binding protein-C. Circ. Res. 98, 1290–1298 8. Yasuda, M., Koshida, S., Sato, N., and Obinata, T. (1995) Complete pri-
mary structure of chicken cardiac C-protein (MyBP-C) and its expression
in developing striated muscles. J. Mol Cell Cardiol. 27, 2275–2286 y
Recent studies in the literature report that the interaction of
cMyBP-C N terminus with thin filaments requires the Pro-Ala-
rich region (28, 32, 33, 35, 36). Herron et al. (7) proposed that
the Pro-Ala-rich region may promote binding of cMyBP-C to
thin filament proteins and thus enhance cross-bridge cycling. Here, we show, for the first time, that the hC0C1f region inter-
acts with -TM via the Pro-Ala-rich region. As suggested by the
increased Hill coefficient, hC0C1f interaction with -TM may
alter -TM movement and indirectly modulate troponin posi-
tion/structure leading to increased cooperativity. However,
more experiments are required to investigate the impact of
hC0C1f on -TM structure and regulation. We postulate that
the effects of hC0C1f are primarily mediated by interactions
with the thin filament through actin and -TM. Given the pro-
posed role of cMyBP-C as a brake on actomyosin dynamics (31),
we suggest that hC0C1f disables this brake by increasing ten-
sion cost and ktr. This, in turn, mimics the absence of cMyBP-C,
increases contraction velocity, and, in the working heart,
reduces the systolic phase, similar to what is observed in
cMyBP-C knock-out mice (37). This reduced efficiency of con-
traction then further exacerbates dysfunction in energy-starved
hearts. 9. Shaffer, J. F., Kensler, R. W., and Harris, S. P. (2009) The myosin-binding
protein C motif binds to F-actin in a phosphorylation-sensitive manner. J. Biol. Chem. 284, 12318–12327 10. Kulikovskaya, I., McClellan, G., Flavigny, J., Carrier, L., and Winegrad, S. N-terminal Fragment of cMyBP-C Impairs Myofilament Function enhance length-dependent activation with increased Fmax
and EC50. However, this effect is mostly caused by changes in
Fmax and EC50 at short, but not long, SL, as seen with C0-C2 (7). Therefore, the hC0C1f fragment seems to interfere with thin
filament function, as observed by changes in cooperativity and
Ca2 sensitivity, at short SL. Conversely, hC0C1f impacts
cross-bridge cycling at both SLs, suggesting that it acts on
strong cross-bridge kinetics by increasing the off rate. This is
supported by the colocalized immunofluorescence images,
highlighting the presence of the fragment in both the A- and
I-bands. In conclusion, we show, for the first time, that the cMyBP-C
cleavage product hC0C1f is sufficient to cause the contractile
dysfunction seen post-MI. We propose that its mechanism of
action results from constant interaction with thin filament pro-
teins, such as actin and -TM. Also, such an interaction dis-
rupts the regulation of actomyosin interaction by affecting
endogenous cMyBP-C regulation. Therefore, therapeutic
interventions aimed at preventing cleavage of cMyBP-C and
attenuating the effects of C0-C1f might confer protection fol-
lowing MI. DISCUSSION When the region is cleaved off and
C0-C1f is released from the full-length cMyBP-C, transient sarcomere. However, it remains unclear whether these frag-
ments directly interact with myosin or actin, or both, in a man-
ner that confers stability in the sarcomere. Recent studies dem-
onstrate that cMyBP-C N-regions structurally reach thin
filaments (32, 33) and dynamically interact with actin (12). In
the present study, we confirmed that recombinant C0-C1f pro-
teins were able to diffuse into the sarcomere and specifically
localize on the A-band and I-band in the presence of endoge-
nous cMyBP-C without changing the interfilament space at
both short and longer SL. Interestingly, we did not see any
staining of hC0C1f fragments on the Z-line, suggesting that
interaction of C0-C1f is very specific to thin and thick filament
proteins. However, further studies are needed to define the
exact region of interaction that is necessary in the N-region on
the thick and thin filaments. The N-region of cMyBP-C binds
transiently to actin and the myosin S2 region in a phosphory-
lation-dependent manner to regulate actomyosin interaction
and force generation. When the region is cleaved off and
C0-C1f is released from the full-length cMyBP-C, transient In the presence of C0-C2, studies show that maximal force is
significantly reduced but that no changes occur in ktr, tension
redevelopment, compared with the untreated control mouse
myocytes (7). In contrast, when human myocytes were permea-
bilized, myocardial preparations were treated with C0-C2,
maximal force, and ktr values were decreased and increased,
respectively, indicating that the difference in ktr largely resulted
from the presence of -myosin heavy chain in adult human,
compared with adult mouse, ventricle (7). Similarly, in our
study, the presence of hC0C1f was able to reduce maximal force
and increase ktr in human myocytes. It is unlikely that the
increase in ktr was due to accelerated thin filament activation
because this parameter was assessed at maximum saturating
levels of activating Ca2, a condition where thin filament acti-
vation kinetics is no longer rate-limiting for force development. Interestingly, our study determined that permeabilization of
hC0C1f causes a decrease in myofilament Ca2 sensitivity in
contrast to the C0-C2 effects. Furthermore, hC0C1f appears to MARCH 28, 2014•VOLUME 289•NUMBER 13 8825 JOURNAL OF BIOLOGICAL CHEMISTRY MARCH 28, 2014•VOLUME 289•NUMBER 13 N-terminal Fragment of cMyBP-C Impairs Myofilament Function 16. Chen, P. P., Patel, J. R., Rybakova, I. N., Walker, J. W., and Moss, R. L. (2010) Protein kinase A-induced myofilament desensitization to Ca2 as a
result of phosphorylation of cardiac myosin-binding protein C. J. Gen. Physiol. 136, 615–627 27. Chandra, M., Rundell, V. L., Tardiff, J. C., Leinwand, L. A., De Tombe, P. P.,
and Solaro, R. J. (2001) Ca2 activation of myofilaments from transgenic
mouse hearts expressing R92Q mutant cardiac troponin T. Am. J. Physiol. Heart Circ. Physiol. 280, H705–H713 17. Ge, Y., Rybakova, I. N., Xu, Q., and Moss, R. L. (2009) Top-down high-
resolution mass spectrometry of cardiac myosin binding protein C re-
vealed that truncation alters protein phosphorylation state. Proc. Natl. Acad. Sci. U.S.A. 106, 12658–12663 28. Barefield, D., and Sadayappan, S. (2010) Phosphorylation and function of
cardiac myosin binding protein-C in health and disease. J. Mol. Cell. Car-
diol. 48, 866–875 29. Sadayappan, S., and Robbins, J. (2008) The death of transcriptional chau-
vinism in the control and regulation of cardiac contractility. Ann. N.Y. Acad. Sci. 1123, 1–9 18. Aït Mou, Y., Reboul, C., Andre, L., Lacampagne, A., and Cazorla, O. (2009)
Late exercise training improves non-uniformity of transmural myocardial
function in rats with ischaemic heart failure. Cardiovasc. Res. 81, 555–564 30. Decker, R. S., Decker, M. L., Kulikovskaya, I., Nakamura, S., Lee, D. C.,
Harris, K., Klocke, F. J., and Winegrad, S. (2005) Myosin-binding protein C
phosphorylation, myofibril structure, and contractile function during
low-flow ischemia. Circulation 111, 906–912 19. Fentzke, R. C., Buck, S. H., Patel, J. R., Lin, H., Wolska, B. M., Stojanovic,
M. O., Martin, A. F., Solaro, R. J., Moss, R. L., and Leiden, J. M. (1999)
Impaired cardiomyocyte relaxation and diastolic function in transgenic
mice expressing slow skeletal troponin I in the heart. J. Physiol. 517,
143–157 31. Previs, M. J., Beck Previs, S., Gulick, J., Robbins, J., and Warshaw, D. M. (2012) Molecular mechanics of cardiac myosin-binding protein C in na-
tive thick filaments. Science 337, 1215–1218 20. Irving, T. C., Konhilas, J., Perry, D., Fischetti, R., and de Tombe, P. P. (2000)
Myofilament lattice spacing as a function of sarcomere length in isolated
rat myocardium. Am. J. Physiol. Heart Circ. Physiol. 279, H2568–H2573 32. Luther, P. K., and Craig, R. (2011) Modulation of striated muscle contrac-
tion by binding of myosin binding protein C to actin. Bioarchitecture 1,
277–283 y
y
y
21. de Tombe, P. REFERENCES (2003) Effect of MyBP-C binding to actin on contractility in heart muscle. J. Gen. Physiol. 122, 761–774 11. Ratti, J., Rostkova, E., Gautel, M., and Pfuhl, M. (2011) Structure and
interactions of myosin-binding protein C domain C0. Cardiac-specific
regulation of myosin at its neck? J. Biol. Chem. 286, 12650–12658 12. Kensler, R. W., Shaffer, J. F., and Harris, S. P. (2011) Binding of the N-ter-
minal fragment C0-C2 of cardiac MyBP-C to cardiac F-actin. J Struct Biol
174, 44–51 13. Weith, A., Sadayappan, S., Gulick, J., Previs, M. J., Vanburen, P., Robbins,
J., and Warshaw, D. M. (2012) Unique single molecule binding of cardiac
myosin binding protein-C to actin and phosphorylation-dependent inhi-
bition of actomyosin motility requires 17 amino acids of the motif domain. J. Mol Cell Cardiol. 52, 219–227 14. Gruen, M., Prinz, H., and Gautel, M. (1999) cAPK-phosphorylation con-
trols the interaction of the regulatory domain of cardiac myosin binding
protein C with myosin-S2 in an on-off fashion. FEBS Lett. 453, 254–259 15. Sadayappan, S., Osinska, H., Klevitsky, R., Lorenz, J. N., Sargent, M., Molk-
entin, J. D., Seidman, C. E., Seidman, J. G., and Robbins, J. (2006) Cardiac
myosin binding protein C phosphorylation is cardioprotective. Proc. Natl. Acad. Sci. U.S.A. 103, 16918–16923 8826
JOURNAL OF BIOLOGICAL CHEMISTRY 8826 VOLUME 289•NUMBER 13•MARCH 28, 2014 VOLUME 289•NUMBER 13•MARCH 28, 2014 N-terminal Fragment of cMyBP-C Impairs Myofilament Function P., and ter Keurs, H. E. (1990) Force and velocity of sarcom-
ere shortening in trabeculae from rat heart. Effects of temperature. Circ. Res. 66, 1239–1254 33. Luther, P. K., Winkler, H., Taylor, K., Zoghbi, M. E., Craig, R., Padrón, R.,
Squire, J. M., and Liu, J. (2011) Direct visualization of myosin-binding
protein C bridging myosin and actin filaments in intact muscle. Proc. Natl. Acad. Sci. U.S.A. 108, 11423–11428 22. de Tombe, P. P., and Stienen, G. J. (1995) Protein kinase A does not alter
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G. W., 2nd, Klevitsky, R., Seidman, C. E., Seidman, J. G., and Robbins, J. (2005) Cardiac myosin-binding protein-C phosphorylation and cardiac
function. Circ. Res. 97, 1156–1163 23. Rundell, V. L., Manaves, V., Martin, A. F., and de Tombe, P. P. (2005)
Impact of -myosin heavy chain isoform expression on cross-bridge cy-
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dent changes in myosin composition correlate with enhanced economy of
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https://openalex.org/W4280492487
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https://www.nature.com/articles/s41598-022-12216-2.pdf
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English
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One-year outcomes of intravitreal brolucizumab injections in patients with polypoidal choroidal vasculopathy
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Scientific reports
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cc-by
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Arisa Ito1, Maiko Maruyama‑Inoue1*, Yoko Kitajima2, Shoko Ikeda1, Tatsuya Inoue1 &
Kazuaki Kadonosono1 Arisa Ito1, Maiko Maruyama‑Inoue1*, Yoko Kitajima2, Shoko Ikeda1, Tatsuya Inoue1 &
Kazuaki Kadonosono1 To evaluate the 1-year visual outcomes and anatomic responses of Japanese patients who received
intravitreal brolucizumab (IVBr) injections for polypoidal choroidal vasculopathy (PCV). This was a
retrospective study of 17 treatment-naïve eyes with PCV that were treated with IVBr. We evaluated
the best-corrected visual acuity (BCVA), central macular thickness (CMT), central choroidal thickness
(CCT) and number of injections for 1 year. The eradication of polypoidal lesions was also evaluated
using by indocyanine green angiography during the 1-year follow-up. Non-infectious intraocular
inflammation developed in two (11.8%) eyes; 15 eyes were assessed at the 1-year follow-up
examination. The mean BCVA improved significantly from 0.28 at baseline to 0.13 (P < 0.05) at 1 year. The CMT and CCT decreased significantly after 1 year. The mean number of injections was 6.4 ± 0.13. The rate of complete resolution of polypoidal lesions at 1 year was 93.3%. A dry macula was achieved
in 13 eyes (86.6%) after the loading phase and in 11 eyes (73.3%) at 1 year. The IVBr injections
appeared to be effective for improving both functional and anatomic outcomes in Japanese patients
with PCV, with a high regression rate of polypoidal lesions. Polypoidal choroidal vasculopathy (PCV) was originally described by Yannuzzi et al.1,2 as a distinct subtype of
wet age-related macular degeneration (AMD). Recently, Spaide et al.3 described that PCV is a variant of type 1
macular neovascularization that is more prevalent in Asian individuals. Indocyanine green angiography (ICGA)
visualized a branching vascular network and various numbers of aneurysmal dilations at the outer edge of the
expanding lesion. In Japan, it has been reported that approximately half of the patients with wet AMD have PCV4.it In eyes with PCV, massive hemorrhages and significant loss of vision are evident after a long-term follow-up
period5. Previous studies have reported that anti-vascular endothelial growth factor (VEGF) agents, such as
ranibizumab (Lucentis, Genentech, Inc., South San Francisco, CA) and aflibercept (Eylea, Bayer Health Care,
Berlin, Germany) have had been effective in the treatment of PCV, although numerous injections are needed
to stabilize patients’ visual acuity (VA)6–13. Moreover, aflibercept is more effective than ranibizumab for achiev-
ing eradication of polypoidal lesions. www.nature.com/scientificreports www.nature.com/scientificreports www.nature.com/scientificreports Arisa Ito1, Maiko Maruyama‑Inoue1*, Yoko Kitajima2, Shoko Ikeda1, Tatsuya Inoue1 &
Kazuaki Kadonosono1 However, the eradication rate of polypoidal lesions at 1 year in Japanese
patients was not as high as the 13% to 39% reported with ranibizumab6–9 and the 39% to 55% reported with
aflibercept10–12. l
Recently, brolucizumab was sanctioned as a new anti-VEGF agent for the treatment of AMD. Brolucizumab
is a roughly 26-kDa single-chain antibody fragment14. The HAWK and HARRIER studies15,16, worldwide phase
3 clinical trials, showed that intravitreal brolucizumab (IVBr) injections administered at every-12-week/every-
8-week intervals were effective for improving and stabilizing VA for 96 weeks and were not inferior to the
every-8-week dosing interval for intravitreal aflibercept. Moreover, IVBr injections provided better intraretinal,
subretinal, and sub-retinal pigment epithelial fluid control than intravitreal aflibercept. p g
pll
p
However, the results of IVBr injections for treating PCV in a real-world clinical setting have not been reported. The purpose of this study was to evaluate the 1-year visual outcomes and anatomic responses of Japanese patients
with PCV treated with IVBr injections. 1Department of Ophthalmology, Yokohama City University Medical Center, 4‑57 Urafune‑cho, Minami‑ku,
Yokohama, Kanagawa 232‑0024, Japan. 2Department of Ophthalmology, Sakae Kyosai Hospital, Kanagawa,
Japan. *email: maicoo@urahp.yokohama-cu.ac.jp Scientific Reports | (2022) 12:7987 | https://doi.org/10.1038/s41598-022-12216-2 www.nature.com/scientificreports/ ntificreports/
Figure 1. Changes in the best-corrected visual acuity (BCVA) during the 12-month follow-up period. The
mean BCVAs at 4, 6, and 12 months improved significantly compared with the preoperative VA (*P < 0.05). Figure 1. Changes in the best-corrected visual acuity (BCVA) during the 12-month follow-up period. The i Figure 1. Changes in the best-corrected visual acuity (BCVA) during the 12-month follow-up period. The
mean BCVAs at 4, 6, and 12 months improved significantly compared with the preoperative VA (*P < 0.05). Figure 2. Changes in the central macular thickness (CMT) during the 12-month follow-up period. The mean
CMTs at 4, 6, and 12 months decreased significantly compared with baseline (**P < 0.01). Figure 2. Changes in the central macular thickness (CMT) during the 12-month follow-up period. The mean
CMTs at 4, 6, and 12 months decreased significantly compared with baseline (**P < 0.01). Resultsh This study included 17 consecutive eyes of 17 Japanese patients with treatment-naïve PCV. Of the 17 patients
enrolled in this study, two men developed intraocular inflammation (IOI) and were excluded. Therefore, the
1-year follow-up was completed in 15 eyes of 15 patients (13 men, 2 women; mean age, 77.8 ± 2.7 years; range,
53–90 years).hi y
The best-corrected visual acuity (BCVA) improved significantly from 0.28 ± 0.05 (range; -0.079 to 0.69) before
treatment to 0.17 ± 0.05 after the loading phase (4 months after starting treatment; P < 0.05). The BCVA was
maintained at 0.12 ± 0.04 (P < 0.05) at 6 months and 0.13 ± 0.06 (P < 0.05) at 1 year (Fig. 1).hi y
g
The central macular thickness (CMT) decreased significantly from 421 ± 53 µm before the initial treatment
to 219 ± 26 µm at 4 months (P < 0.01) and was maintained at 230 ± 22 µm (P < 0.01) at 6 months and 206 ± 18 µm
(P < 0.01) at 1 year (Fig. 2).hi y
g
The central choroidal thickness (CCT) decreased significantly from 226 ± 35 µm before the initial treatment
to 198 ± 32 µm at 4 months (P < 0.01) and was maintained at 197 ± 31 µm (P < 0.05) at 6 months and 181 ± 30 µm
(P < 0.01) at 1 year (Fig. 3). Scientific Reports | (2022) 12:7987 | https://doi.org/10.1038/s41598-022-12216-2 www.nature.com/scientificreports/ ntificreports/
Figure 3. Changes in the central choroidal thickness (CCT) during the 12-month follow-up period. The mean
CCTs at 4, 6, and 12 months decreased significantly compared with baseline (*P < 0.05, **P < 0.01). ntificreports/
Figure 3. Changes in the central choroidal thickness (CCT) during the 12-month follow-up period. The mean
CCTs at 4, 6, and 12 months decreased significantly compared with baseline (*P < 0.05, **P < 0.01). Figure 3. Changes in the central choroidal thickness (CCT) during the 12-month follow-up period. The mean
CCTs at 4, 6, and 12 months decreased significantly compared with baseline (*P < 0.05, **P < 0.01). During the 1-year follow-up, the mean number of injections administered was 6.38 ± 0.14. After the loading
phase, nine patients (60.0%) continued maintenance treatment every 3 months. The treatment interval for four
patients (26.6%) was shortened from every 3 months to every 2 months due to the appearance of fluid or new
hemorrhage. Resultsh Two patients (13.3%) continued maintenance treatment every 2 months due to the residual fluid
after the loading phase.ht t
g p
The proportion of patients with complete regression of polypoidal lesions after 1 year seen on ICGA was
93.3% (14 eyes). A dry macula was achieved in 13 eyes (86.6%) after the loading phase and 11 eyes (73.3%) at
1 year. One eye of four eyes without dry macular at 1 year was the eye without regression of the polypoidal lesion. Figure 4 shows a representative case.t g
p
Brolucizumab-related IOI developed in two (11.8%) of 17 eyes. A 72-year-old man had a vitreous opacity after
the third injection. His visual acuity reduced from 0.22 logMAR units at baseline to 0.30 logMAR units. 0.1%
betamethasone eye drops were started four times a day. A vitreous opacity resolved after two months, therefore
he stopped using eye drops. 10 months after IOI was diagnosed, the patient’s visual acuity improved to 0.22 log-
MAR units, and the fundus examination showed no recurrence of vitreous haze. OCT showed no recurrence of
exudation. Another man, 63-year-old man, had vascular sheathing without retinal vascular occlusion after the
fourth injection. His visual acuity did not reduce from 0.15 logMAR units at baseline to -0.079 logMAR units. The patients visited the hospital three months after the fourth injection without any symptoms. Therefore, we
could not treat the prompt treatment. Four months after vascular sheathing was diagnosed, the patient’s visual
acuity was maintained. The fundus examination showed vascular sheathing in the peripheral retina remained,
however there was no enlargement of the lesions or the appearance of vascular occlusion. OCT showed recur-
rence of exudation. Thereafter, the patient received additional intravitreal aflibercept injections. In both cases,
the BCVA after 12 months did not deteriorate compared to the baseline BCVA. No other severe adverse events,
such as infectious endophthalmitis, rhegmatogenous retinal detachment, cerebral infarction, or myocardial
infarction, occurred. Discussion Koizumi et al.19
also reported changes in subfoveal choroidal thickness after intravitreal aflibercept injections in 86 patients
with PCV at 1 year and reported a mean CCT decrease from 270 to 234 µm (13.3% decrease from baseline). Therefore, the reduction of the choroidal thickness in patients with PCV treated with IVBr injections tended to
be greater than that achieved with aflibercept. Kim et al.20 reported decreased choroidal thickness was observed
after anti-VEGF therapy for PCV. Choi et al.21 studied 88 patients with PCV who received anti-VEGF injections
and reported that faster chorioretinal atrophy (CRA) growth was related significantly to a decreased choroidal
thickness. A larger decrease in choroidal thickness might be a risk factor for long-term development of CRA in
patients with PCV.Choroidal thinning might affect the CRA in the longer term and CRA might be related to the
BCVA. Therefore, it is necessary to consider whether IVBr injections affect the VA for a longer follow-up period. h
y
jf
g
p p
In the current study, 60.0% of the patients with PCV completed the 12-week interval injections for 1 year. A dry macula was achieved in 73.3% after 1 year. Ogura et al.17 reported the efficacy of brolucizumab in Japa-
nese patients with PCV in a sub-analysis of the HAWK study. The authors reported that the probability of
12-week-interval injections after a loading phase through 48 weeks was 76%, which was similar to our result. The 12-week-interval injections were the longest intervals in the TAE regimen, which is widely used worldwide. The persistence of brolucizumab might derive from its high affinity for VEGF. Its low molecular weight allows
more delivery of drug per injection compared with other available anti-VEGFs and offers the potential for more
effective tissue penetration and increased duration of action14. Therefore, brolucizumab might reduce the treat-
ment burden for patients with PCV. p
In the current study, complete regression of polypoidal lesions at 12 months was achieved in 93.3%. Matsu-
moto et al.22 and Fukuda et al.23 reported the short-term outcomes of IVBr for PCV. In these two reports, the
regression rates of the polypoidal lesions after 3monthly IVBr injections were 78.9% (15eyes of 19eyes) and 78.6%
(11 eyes of 14 eyes). There were no previous studies which reports the regression rate of polypoidal lesions regard-
ing the 1-year outcomes. Discussion Twelve months after the first injection, the VA improved to 0.045 logMAR. (F) A color
fundus photograph shows no reddish-orange lesion or hemorrhage at the macula. (G) FA shows staining with
no leakage. (H) ICGA shows complete polyp regression, although an abnormal vascular network (arrow heads)
remained. (I) OCT shows no polypoidal lesion or SRF. Irregular retinal pigment epithelium elevation was
observed where the abnormal vascular network was located. Figure 4. The case of an 86-year-old man who presented with reduced visual acuity in his right eye. (A) A color
fundus photograph shows reddish-orange polypoidal lesions, including his fovea, submacular hemorrhage,
and a large area of subretinal fluid (SRF). (B) Fluorescein angiography (FA) demonstrates occult leakage. (C)
Indocyanine green angiography (ICGA) shows two polypoidal lesions (white arrows) and an abnormal vascular
network (arrow heads). (D, E) An optical coherence tomography (OCT) images obtained at baseline show SRF
with polypoidal lesions. (D through the superior polypoidal lesion, E through the inferior polypoidal lesion
involving the fovea) The visual acuity (VA) was 0.39 logarithm of the minimum angle of resolution (logMAR) in
the right eye, and the patient was diagnosed with PCV. The patient received IVBr injections during the loading
phase and during the maintenance phase, he was treated every 3 months IVBr injections. The exudative changes
did not recur for 1 year. Twelve months after the first injection, the VA improved to 0.045 logMAR. (F) A color
fundus photograph shows no reddish-orange lesion or hemorrhage at the macula. (G) FA shows staining with
no leakage. (H) ICGA shows complete polyp regression, although an abnormal vascular network (arrow heads)
remained. (I) OCT shows no polypoidal lesion or SRF. Irregular retinal pigment epithelium elevation was
observed where the abnormal vascular network was located. our study and previous results is difficult, our visual outcomes using brolucizumab seem favorable, with fewer
injections compared with aflibercept in Japanese patients with PCV.i j
pl
p
p
p
In the current study, the CCT decreased significantly from 226 to 181 µm (19.9% decrease from baseline)
at 1 year. We previously reported the 1-year results in treatment-naïve PCV treated with bimonthly intravitreal
aflibercept that had a mean CCT decrease from 265 to 231 µm (12.8% decrease from baseline)11. Discussion We investigated the 1-year outcomes of IVBr injections in Japanese patients with treatment-naïve PCV and
found that the BCVA improved significantly during the follow-up period. The CMT and CCT also decreased
significantly after 1 year. The polypoidal lesions regressed completely after 1 year in 93.3% of eyes. To the best of
our knowledge, no previous studies have evaluated the regression rate of polypoidal lesions regarding the 1-year
outcomes following IVBr injections for patients with PCV.fi g
j
p
In a sub-analysis of the HAWK study, Ogura et al.17 reported the efficacy of brolucizumab in Japanese patients
with PCV. They reported brolucizumab (n = 39) was as good as aflibercept (n = 30) in improving vision. Yama-
moto et al.12 conducted a multicenter study and reported the 1-year results in 90 treatment-naïve eyes with
PCV treated with bimonthly intravitreal aflibercept and reported a mean BCVA improvement from 0.31 to 0.17
logMAR. The mean number of injections during the first year was 7.1. Morimoto et al.18 reported the 2-year
results in 58 treatment-naïve eyes with PCV with good initial BCVA treated with intravitreal aflibercept using a
treat-and-extend (TAE) regimen and found a mean BCVA improvement from 0.27 to 0.12 logMAR during the
first year. The mean number of injections during the first year was 7.72. In the current study, the mean BCVA
improved from 0.28 to 0.13 logMAR. The number of injections was 6.4. Although a direct comparison between Scientific Reports | (2022) 12:7987 | https://doi.org/10.1038/s41598-022-12216-2 www.nature.com/scientificreports/ www.nature.com/scientificreports/ Figure 4. The case of an 86-year-old man who presented with reduced visual acuity in his right eye. (A) A color
fundus photograph shows reddish-orange polypoidal lesions, including his fovea, submacular hemorrhage,
and a large area of subretinal fluid (SRF). (B) Fluorescein angiography (FA) demonstrates occult leakage. (C)
Indocyanine green angiography (ICGA) shows two polypoidal lesions (white arrows) and an abnormal vascular
network (arrow heads). (D, E) An optical coherence tomography (OCT) images obtained at baseline show SRF
with polypoidal lesions. (D through the superior polypoidal lesion, E through the inferior polypoidal lesion
involving the fovea) The visual acuity (VA) was 0.39 logarithm of the minimum angle of resolution (logMAR) in
the right eye, and the patient was diagnosed with PCV. The patient received IVBr injections during the loading
phase and during the maintenance phase, he was treated every 3 months IVBr injections. The exudative changes
did not recur for 1 year. Scientific Reports | (2022) 12:7987 | Methods Received: 7 February 2022; Accepted: 6 May 2022 Received: 7 February 2022; Accepted: 6 May 2022 www.nature.com/scientificreports/ ranibizumab monotherapy. In another of our previous studies, the regression rates of polypoidal lesions in
patients treated with combination therapy with either intravitreal ranibizumab or intravitreal aflibercept were
higher (78.2% and 78.9%) than either monotherapy with those drugs24. In the current study, the regression rate
of polypoidal lesions with IVBr injections was much higher than with either ranibizumab or aflibercept mono-
therapy or PDT combination therapy, which might be why 73.3% of the patients had a dry macula at 1 year. However, brolucizumab-related IOI is a major adverse event associated with intravitreal brolucizumab. In
the current study, two (11.8%) of 17 eyes had IOI. The frequency of IOI was roughly the same compared with
a sub-analysis of HAWK study which Ogura et al. reported (15.4%)15–17. Although IVBr injections for PCV are
effective, we should be cautious about IOI and treat promptly if brolucizumab-related IOI develops.hh f
p
p y
p
This study had some limitations, a major one being the retrospective design. The second limitation was the
small sample size. A large-scale randomized study should confirm the current results. Furthermore, the long-term
outcomes are unknown. The results of this study must be evaluated with a longer follow-up period.f h
In conclusion, IVBr injections effectively improved the vision and exudative changes in patients with PCV
over a 1-year follow-up, along with higher resolution rate of polypoidal lesions as seen on ICGA. The results of
this analysis suggested that brolucizumab could be useful to treat PCV with a lower treatment burden. Discussion In our previous study, when intraocular aflibercept monotherapy was used to treat PCV,
complete regression of polypoidal lesions was achieved in 48.0% with a bimonthly fixed regimen and in 52.9%
with an as-needed regimen11. In the EVEREST II study8, which compared the effectiveness between ranibizumab
monotherapy with photodynamic therapy (PDT) combination therapy, the rates at 12 months were 69.3% in
the patients treated with PDT/intravitreal ranibizumab therapy and 34.7% in patients treated with intravitreal Scientific Reports | (2022) 12:7987 | https://doi.org/10.1038/s41598-022-12216-2 www.nature.com/scientificreports/ References e e e ces
1. Yannuzzi, L. A., Sorenson, J., Spaide, R. F. & Lipson, B. Idiopathic polypoidal choroidal vasculopathy (IPCV). Retina 10, 1–8. https://doi.org/10.1097/00006982-199001010-00001 (1990). 1. Yannuzzi, L. A., Sorenson, J., Spaide, R. F. & Lipson, B. Idiopathic polypoidal choroidal vasculopathy (IPCV). Retina 10, 1–8. https://doi.org/10.1097/00006982-199001010-00001 (1990).h p
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2. Yannuzzi, L. A. et al. The expanding clinical spectrum of idiopathic polypoidal choroidal vasculopathy. Arch Ophthalmol. 115
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ophtha.2019.11.004 (2020). p
4. Maruko, I. et al. Clinical characteristics of exudative age-related macular degeneration in Japanese patients. Am. J. Ophthalmol
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5. Uyama, M. et al. Polypoidal choroidal vasculopathy: Natural history. Am. J. Ophthalmol. 133, 639–648. https://doi.org/10.1016/
S0002-9394(02)01404-6 (2002). 6. Koh, A. et al. Polypoidal choroidal vasculopathy: Evidence-based guidelines for clinical diagnosis and treatment. Retina 33
686–716. https://doi.org/10.1097/IAE.0b013e3182852446 (2013). 7. Hikichi, T. et al. One-year results of three monthly ranibizumab injections and as-needed reinjections for polypoidal choroidal
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8. Koh, A. et al. Efficacy and safety of ranibizumab with or without verteporfin photodynamic therapy for polypoidal choroida
vasculopathy: A randomized clinical trial. JAMA Ophthalmol. 135, 1206–1213. https://doi.org/10.1001/jamaophthalmol.2017
4030 (2017). 8. Koh, A. et al. Efficacy and safety of ranibizumab with or without verteporfin photodynamic therapy for polypoidal choroidal
vasculopathy: A randomized clinical trial. JAMA Ophthalmol. 135, 1206–1213. https://doi.org/10.1001/jamaophthalmol.2017.
4030 (2017). (
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9. Mori, R., Yuzawa, M., Akaza, E. & Haruyama, M. Treatment results at 1 year of ranibizumab therapy for polypoidal choroidal
vasculopathy in eyes with good visual acuity. Jpn. J. Ophthalmol. 57, 365–371. https://doi.org/10.1007/s10384-013-0245-9 (2013).fil (
)
9. Mori, R., Yuzawa, M., Akaza, E. & Haruyama, M. Treatment results at 1 year of ranibizumab therapy for polypoidal choroida
vasculopathy in eyes with good visual acuity. Jpn. J. Ophthalmol. 57, 365–371. https://doi.org/10.1007/s10384-013-0245-9 (2013) (
)
9. Mori, R., Yuzawa, M., Akaza, E. & Haruyama, M. Treatment results at 1 year of ranibizumab therapy for polypoidal choroidal
vasculopathy in eyes with good visual acuity. Jpn. J. Ophthalmol. 57, 365–371. https://doi.org/10.1007/s10384-013-0245-9 (2013). 10. Lee, W. K. et al. Received: 7 February 2022; Accepted: 6 May 2022 Methods We retrospectively studied 17 consecutive eyes of 17 Japanese patients aged 50 years or older with previously
untreated PCV. All patients received initial treatment using IVBr injections at Yokohama City University Medical
Center between June 2020 and February 2021. The institutional review board of the Yokohama City University
Medical Center approved this study, which followed the tenets of the Declaration of Helsinki. All patients pro-
vided written informed consent before their medical record data were used in research. All patients received comprehensive ophthalmic examinations, including BCVA, slit-lamp biomicroscopy,
optical coherence tomography (OCT) (Spectralis Product Family Version 5.3; Heidelberg Engineering, Heidel-
berg, Germany), fluorescein angiography (FA) and ICGA (Spectralis Product Family Version 5.3; Heidelberg
Engineering, Heidelberg, Germany). The diagnosis of PCV was based on previous reports4,6. Inclusion criteria
included newly diagnosed PCV with the presence of polypoidal choroidal vascular lesion with branching vascular
networks on ICGA. Exclusion criteria were previous history of AMD, history of laser photocoagulation, PDT,
intravitreal injection of other anti-VEGF agents or intravitreal steroids. Patients with history of eye diseases such
as uveitis, retinal vein occlusion, rhegmatogenous retinal detachment, diabetic retinopathy were also excluded. g
g
p
y
During the loading phase, the patients received three monthly IVBr injections. In the maintenance phase,
patients received an IVBr injection every 12 weeks unless any fluid or a new hemorrhage was identified, in which
case the interval between IVBr injections was shortened to every 8 weeks. The endpoints were the BCVA, CMT,
CCT, number of intravitreal injections, regression of polypoidal lesions evaluated by ICGA after 1 year, and
complications. These endpoints were evaluated retrospectively 1 year after the initial treatment. CMT and CCT
were determined by using a caliper function of B-scan OCT. The CMT was defined as the thickness from the
internal limiting membrane to the retinal pigment epithelium at the center of the fovea; the CCT was defined
as the thickness between Bruch’s membrane and the inner surface of the choroidal-scleral junction at the fovea. The statistical analysis was performed with Ekuseru-Toukei (Social Survey Research Information, Tokyo, Japan). The BCVA measured on a Landolt chart was converted into the logarithm of the minimum angle resolution
(logMAR) for statistical analyses. Wilcoxon signed-rank test was used to compare the changes of BCVA, CMT,
and CCT between baseline and other time point. A value of P < 0.05 was considered statistically significant. p
g
The authors declare no competing interests. p
g
The authors declare no competing interests. www.nature.com/scientificreports/ www.nature.com/scientificreports/ 2. Yamamoto, A. et al. One-year results of intravitreal aflibercept for polypoidal choroidal vasculopathy. Ophthalmology 122, 1866–
1872. https://doi.org/10.1016/j.ophtha.2015.05.024 (2015).l p
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13. Maruyama-Inoue, M., Sato, S., Yamane, S. & Kadonosono, K. Intravitreal injection of aflibercept in patients with polyp
choroidal vasculopathy a 3-year follow-up. Retina 38, 2001–2009. https://doi.org/10.1097/IAE.0000000000001818 (2018). p
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14. Nguyen, Q. D. et al. Brolucizumab: Evolution through preclinical and clinical studies and the implications for the m
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127 963 976 htt
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hth 2019 1 14. Nguyen, Q. D. et al. Brolucizumab: Evolution through preclinical and clinical studies and the implications for the management of
neovascular age-related macular degeneration. Ophthalmology 127, 963–976. https://doi.org/10.1016/j.ophtha.2019.12.031 (2020). 14. Nguyen, Q. D. et al. Brolucizumab: Evolution through preclinical and clinical studies and the implications for the management of
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hth 2019 12 031 (2020) 14. Nguyen, Q. D. et al. Brolucizumab: Evolution through preclinical and clinical studies and the implications for the m
neovascular age-related macular degeneration. Ophthalmology 127, 963–976. https://doi.org/10.1016/j.ophtha.2019.1 g y
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neovascular age-related macular degeneration. Ophthalmology 127, 963–976. https://doi.org/10.1016/j.ophtha.2019.12.031 (2 15. Dugel, P. U. et al. HAWK and HARRIER: Phase 3, multicenter, randomized, double-masked trials of brolucizum
age-related macular degeneration. Ophthalmology 127, 72–84. https://doi.org/10.1016/j.ophtha.2019.04.017 (2 U. et al. HAWK and HARRIER: Phase 3, multicenter, randomized, double masked trials of brolucizumab for neovascular
d macular degeneration. Ophthalmology 127, 72–84. https://doi.org/10.1016/j.ophtha.2019.04.017 (2020). g
age-related macular degeneration. Ophthalmology 127, 72–84. https://doi.org/10.1016/j.ophtha.2019.04.017 (2020). g
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16. Dugel, P. U. et al. HAWK and HARRIER: Ninety-six-week outcomes from the phase 3 trials of brolucizumab for neovascula
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17. Ogura, Y. et al. Efficacy and safety of brolucizumab versus aflibercept in eyes with polypoidal choroidal vasculopathy in
participants of HAWK. Br. J. Ophthalmol. https://doi.org/10.1136/bjophthalmol-2021-319090 (2021). k
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18. Author contributionsh The authors were involved in the following aspects of the study: design and conduct (M.M-I.); collection of the
data (A.I., Y.K., S.I.); management (M.M-I., A.I.); analysis (A.I., T.I.); preparation of article (A.I.); review and
approval of the manuscript (M.M-I., K.K.). References Efficacy and safety of intravitreal aflibercept for polypoidal choroidal vasculopathy in the PLANET study a rand- p
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0. Lee, W. K. et al. Efficacy and safety of intravitreal aflibercept for polypoidal choroidal vasculopathy in the PLANET study a rand-
omized clinical trial. JAMA Ophthalmol. 136, 786–793. https://doi.org/10.1001/jamaophthalmol.2018.1804 (2018).l J
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11. Inoue, M. et al. Aflibercept for polypoidal choroidal vasculopathy. Retina 36, 1527–1534. https://doi.org/10.1097/IAE.0000000000
000933 (2016). 11. Inoue, M. et al. Aflibercept for polypoidal choroidal vasculopathy. Retina 36, 1527–1534. https://doi.org/10.1097/IAE.0000000000
000933 (2016). https://doi.org/10.1038/s41598-022-12216-2 Scientific Reports | (2022) 12:7987 | www.nature.com/scientificreports/ Morimoto, M., Matsumoto, H., Mimura, K. & Akiyama, H. Two-year results of a treat-and-extend regimen with aflibercept for
polypoidal choroidal vasculopathy. Graefe’s Arch. Clin. Exp. Ophthalmol. 255, 1891–1897. https://doi.org/10.1007/s00417-017-
3718-6 (2017).l (
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19. Koizumi, H. et al. Subfoveal choroidal thickness during aflibercept therapy for neovascular age-related macular degenera
twelve-month results. Ophthalmology 123, 617–624. https://doi.org/10.1016/j.ophtha.2015.10.039 (2016).l p
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20. Kim, J. H. et al. Short-term choroidal thickness changes in patients treated with either ranibizumab or aflibercept: A comparative
study. Br. J. Ophthalmol. 100, 1634–1639. https://doi.org/10.1136/bjophthalmol-2015-308074 (2016). 20. Kim, J. H. et al. Short-term choroidal thickness changes in patients treated with either ranibizumab or aflibercept: A comparative
study. Br. J. Ophthalmol. 100, 1634–1639. https://doi.org/10.1136/bjophthalmol-2015-308074 (2016). y
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21. Choi, E. Y. et al. Long-term incidence and growth of chorioretinal atrophy in patients with polypoidal choroidal vasculopathy. Ophthalmologica 243, 136–144. https://doi.org/10.1159/000501724 (2020). 21. Choi, E. Y. et al. Long-term incidence and growth of chorioretinal atrophy in patients with polypoidal choroidal vasculop
Ophthalmologica 243, 136–144. https://doi.org/10.1159/000501724 (2020). 22. Matsumoto, H. et al. Short-term outcomes of intravitreal brolucizumab for treatment-naïve neovascular age-related macular
degeneration with type 1 choroidal neovascularization including polypoidal choroidal vasculopathy. Sci. Rep. 11, 1–8. https://doi.
org/10.1038/s41598-021-86014-7 (2021).l 22. Matsumoto, H. et al. Short-term outcomes of intravitreal brolucizumab for treatment-naïve neovascular age-related macular
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23. Fukuda, Y. et al. Comparison of outcomes between 3 monthly brolucizumab and aflibercept injections for polypoidal chor
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24. Ito, A. et al. Comparison of one-year results of photodynamic therapy combined with ranibizumab or aflibercept for treating
polypoidal choroidal vasculopathy. PLoS ONE 15, 1–11. https://doi.org/10.1371/journal.pone.0235213 (2020). 24. Ito, A. et al. Comparison of one-year results of photodynamic therapy combined with ranibizumab or aflibercept for treating
polypoidal choroidal vasculopathy. PLoS ONE 15, 1–11. https://doi.org/10.1371/journal.pone.0235213 (2020). 4. Ito, A. et al. Comparison of one-year results of photodynamic therapy combined with ranibizumab or aflibercept for treating
polypoidal choroidal vasculopathy. PLoS ONE 15, 1–11. https://doi.org/10.1371/journal.pone.0235213 (2020). Additional information Correspondence and requests for materials should be addressed to M.M.-I. Correspondence and requests for materials should be addressed to M.M.-I. © The Author(s) 2022 Reprints and permissions information is available at www.nature.com/reprints. Reprints and permissions information is available at www.nature.com/reprints. Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and
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A consistent discrete version of a non-autonomous SIRVS model
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1. Introduction to some particular properties of discrete epidemiological
models obtained from continuous models by some NSFD
scheme [16]. We note that while the papers cited above
consider autonomous models, in the present work we discuss
dynamical consistency for a nonautonomous model. To the
best of our knowledge, this is the first work where the
consistency of a discretized epidemiological model with
the original continuous model is discussed in the nonau-
tonomous context. Most of the epidemiological models in the literature are
continuous models. In spite of this, recently, there has been a
growing interest in discrete-time models [1–7]. In this work,
we will use Mickens nonstandard difference (NSFD) scheme
to achieve a discretization of a family of continuous epi-
demiological models with vaccination and general incidence
function considered in [8]. We have multiple objectives:
firstly, we want to obtain conditions for extinction and per-
manence of the disease for the discrete family; next, having
a continuous and a corresponding discrete family of models,
we wish to discuss the problem of consistency of the discrete
models with the corresponding continuous ones; finally,
we intend to present some simulation results.h We remark that, even in the very particular case of
autonomous models with mass action incidence and assum-
ing that there is no disease related death, it follows from
results in [17] that, when R0 > 1, the continuous autonomous
model has one or two endemic equilibrium points coexisting
with the disease-free equilibrium. The existence and stability
of the equilibriums are governed by R0 and three additional
thresholds: R𝑞, RV, and R𝑞V. These thresholds determine
the qualitative behaviour of the system. This very particular
situation shows that, even in the autonomous context, the
qualitative behaviour can be difficult to determine. The dynamical consistency of a numerical scheme with
the associated continuous system is not a precise definition. By the expression “dynamical consistency,” it is meant that
the numerical solutions replicate some of the properties of the
continuous systems solutions. y
We will consider the dynamical consistency regarding the
permanence and extinction of the disease: whenever there is
extinction (permanence) of the disease for the continuous-
time model, the same holds for the discrete-time one. Several
papers [9–15] discuss the dynamical consistency with respect One of the motivations for our work was that the
difficulties increase considerably when we deal with a general
nonautonomous situation. Hindawi
Discrete Dynamics in Nature and Society
Volume 2018, Article ID 8152032, 18 pages
https://doi.org/10.1155/2018/8152032 Joaquim Mateus,1 César M. Silva
,2 and S. Vaz2 Joaquim Mateus, César M. Silva
, and S. Vaz
1Unidade de Investigac¸˜ao para o Desenvolvimento do Interior (UDI), Instituto Polit´ecnico da Guarda, 6300-559 Guarda, Portugal
2Centro de Matem´atica e Aplicac¸˜oes e Departamento de Matem´atica, Universidade da Beira Interior, 6201-001 Covilh˜a, Portugal 1Unidade de Investigac¸˜ao para o Desenvolvimento do Interior (UDI), Instituto Polit´ecnico da Guarda, 6300-559 Guarda, Portugal
2Centro de Matem´atica e Aplicac¸˜oes e Departamento de Matem´atica, Universidade da Beira Interior, 6201-001 Covilh˜a, Portugal Correspondence should be addressed to C´esar M. Silva; csilva@ubi.pt Received 24 February 2018; Revised 27 May 2018; Accepted 29 May 2018; Published 24 June 2018 Academic Editor: Pilar R. Gordoa Copyright © 2018 Joaquim Mateus et al. This is an open access article distributed under the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. A family of discrete nonautonomous SIRVS models with general incidence is obtained from a continuous family of models by
applying Mickens nonstandard discretization method. Conditions for the permanence and extinction of the disease and the stability
of disease-free solutions are determined. Concerning extinction and persistence, the consistency of those discrete models with the
corresponding continuous model is discussed: if the time step is sufficiently small, when we have extinction (permanence) for the
continuous model, we also have extinction (permanence) for the corresponding discrete model. Some numerical simulations are
carried out to compare the different possible discretizations of our continuous model using real data. 1. Introduction Thus, we decided to discuss an
aspect of consistency that, nevertheless, is very important Discrete Dynamics in Nature and Society 2 infective class) from the susceptible individuals; the function
𝜎𝑛𝜓is the incidence (into the infective class) from the vac-
cinated individuals; 𝜇𝑛are the natural deaths; 𝑝𝑛represents
the susceptibles vaccination; 𝜂𝑛represents the immunity loss
and consequence influx in the susceptible class; 𝛼𝑛are the
deaths occurring in the infective class; 𝛾𝑛is the recovery. We
will assume that (Λ 𝑛), (𝜇𝑛), (𝑝𝑛), (𝜂𝑛), (𝛼𝑛), (𝛽𝑛), (𝜎𝑛), and (𝛾𝑛)
are bounded and nonnegative sequences and that there are
positive constants 𝑤𝜇, 𝑤Λ, 𝑤𝑝, 𝑘𝜑, and 𝑘𝜓such that from the point of view of biomathematics: can we obtain
consistency between continuous and discrete-time models
from the point of view of persistence and extinction of the
disease? Regarding our simulation results, we considered two
different types of computational experiments. Our first set of
simulation results are designed to compare different possible
discretizations of our continuous models. After this discus-
sion, we apply our model to a real situation, considering data
from the incidence of measles in France in the period 2012-
2016. To the possible extent, this data is used to estimate our
model parameters and the computational results obtained are
compared with real data.h (H1) the functions 𝜑: R2 →R and 𝜓: R2 →R are
nonnegative and differentiable in (R+
0)2 and the func-
tions R+
0 ∋𝑥→𝜕2𝜑(𝑥, 0) and R+
0 ∋𝑥→𝜕2𝜓(𝑥, 0)
are nondecreasing and Lipschitz, with Lipschitz con-
stants 𝑘𝜑and 𝑘𝜓, (H1) the functions 𝜑: R2 →R and 𝜓: R2 →R are
nonnegative and differentiable in (R+
0)2 and the func-
tions R+
0 ∋𝑥→𝜕2𝜑(𝑥, 0) and R+
0 ∋𝑥→𝜕2𝜓(𝑥, 0)
are nondecreasing and Lipschitz, with Lipschitz con-
stants 𝑘𝜑and 𝑘𝜓, The law of mass action states that the rate of change in
the disease incidence is directly proportional to the product
of the number of susceptible and infective individuals and
was the paradigm in the classic models in epidemiology. This is why classical models usually consider a bilinear
incidence rate 𝛽𝑆𝐼, where 𝑆and 𝐼denote, respectively, the
number of susceptible and infective individuals, to model
the disease transmission. In spite of this, it is sometimes
important to consider other forms of incidence functions. 1. Introduction Another usual assumption is the time independence of the
parameters model parameters: in fact, the majority of the
epidemiological models in the literature are given by a system
of autonomous differential or difference equations. Neverthe-
less, the assumption that the parameters are independent of
time is not very realistic in many situations and it is useful
to consider nonautonomous models that, for instance, allow
the discussion of environmental and demographic effects that
change with time [18, 19]. In this work the family of models
considered is nonautonomous and the incidence rates are
taken from a large class of functions. (H2) we have 𝜑(𝑥, 0) = 𝜓(𝑥, 0) = 𝜑(0, 𝑦) = 𝜓(0, 𝑦) = 0 for
all 𝑥, 𝑦∈R+
0, (H2) we have 𝜑(𝑥, 0) = 𝜓(𝑥, 0) = 𝜑(0, 𝑦) = 𝜓(0, 𝑦) = 0 for
all 𝑥, 𝑦∈R+
0, (H2) we have 𝜑(𝑥, 0) = 𝜓(𝑥, 0) = 𝜑(0, 𝑦) = 𝜓(0, 𝑦) = 0 for
all 𝑥, 𝑦∈R+
0, (H3) lim sup𝑛→+∞∏
𝑛+𝜔𝜇
𝑘=𝑛(1/(1 + 𝜇𝑘)) < 1, (H3) lim sup𝑛→+∞∏
𝑛+𝜔𝜇
𝑘=𝑛(1/(1 + 𝜇𝑘)) < 1, (H4) lim inf𝑛→+∞∑𝑛+𝜔Λ
𝑘=𝑛+1Λ 𝑘>0andlim inf𝑛→+∞∑
𝑛+𝜔𝑝
𝑘=𝑛+1𝑝𝑘>
0, (H4) lim inf𝑛→+∞∑𝑛+𝜔Λ
𝑘=𝑛+1Λ 𝑘>0andlim inf𝑛→+∞∑
𝑛+𝜔𝑝
𝑘=𝑛+1𝑝𝑘>
0, In this work, we prove, when our conditions prescribe
extinction (permanence) for the continuous model we also
have extinction (permanence) for the corresponding discrete
model as long as the time step is smaller than some constant
(that depends on some model parameters and on the thresh-
old condition). We also consider a family of examples of the
periodic system of period 1 such that the continuous and the
discrete-time system with time step ℎ= 1/𝐿is not consistent,
highlighting the importance of knowing that for time steps
smaller than some explicit value we have consistency. Our model generalizes one obtained by Mickens nonstan-
dard finite difference method from the continuous model [8]
(see Section 2). In [20], a discrete nonautonomous epidemic
model with vaccination and mass action incidence was
obtained by Mickens method. We emphasize that, in the
particular mass action case, our model is not exactly similar
to the model in [20], although Mickens rules were considered
in both. We briefly compare computationally these two
slightly different models in Section 5.h Now, we need to make some definitions. We say that (i) the infectives (𝐼𝑛) are permanent if for any solution
(𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with initial conditions 𝑆0, 𝐼0,
𝑅0, 𝑉0 > 0 there are constants 0 < 𝑚< 𝑀such that (i) the infectives (𝐼𝑛) are permanent if for any solution
(𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with initial conditions 𝑆0, 𝐼0,
𝑅0, 𝑉0 > 0 there are constants 0 < 𝑚< 𝑀such that (i) the infectives (𝐼𝑛) are permanent if for any solution
(𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with initial conditions 𝑆0, 𝐼0,
𝑅0, 𝑉0 > 0 there are constants 0 < 𝑚< 𝑀such that Theorem 2 (Theorem 2 of [8]). Assume that conditions (C1),
(C2), and (C3) hold. Then if there is a constant 𝜆> 0 such
that 𝑅𝑢
𝐶(𝜆) < 0, then the infectives 𝐼go to extinction in system
(2). 𝑚< lim inf
𝑛→∞𝐼𝑛⩽lim sup
𝑛→∞
𝐼𝑛< 𝑀;
(10) (10) (ii) the infectives (𝐼𝑛) go to extinction if for any solution
(𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with initial conditions 𝑆0, 𝐼0,
𝑅0, 𝑉0 ⩾0 we have lim𝑛→∞𝐼𝑛= 0. (ii) the infectives (𝐼𝑛) go to extinction if for any solution
(𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with initial conditions 𝑆0, 𝐼0,
𝑅0, 𝑉0 ⩾0 we have lim𝑛→∞𝐼𝑛= 0. In the literature, several models were discretized using
Mickens NSFD schemes [22–35]. Next, we will apply Micken’s
nonstandard method to obtain a discrete version of system
(2). Let 𝜙: R+
0 →R be a positive continuous function such
that In the literature, several models were discretized using
Mickens NSFD schemes [22–35]. Next, we will apply Micken’s
nonstandard method to obtain a discrete version of system
(2). Similar definitions can be made for the other compartments. For instance, if there exist constants 0 < 𝑚< 𝑀such that for
any solution (𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with initial conditions 𝑆0, 𝐼0,
𝑅0, 𝑉0 > 0 we have Let 𝜙: R+
0 →R be a positive continuous function such
that lim
ℎ→0 𝜙(ℎ) = 0. (5) (5) 𝑚< lim inf
𝑛→∞𝑆𝑛⩽lim sup
𝑛→∞
𝑆𝑛< 𝑀
(11) (11) Given ℎ∈R+, we let 𝑡= 𝑛ℎ, with 𝑛∈N, and identify 𝑆(𝑡)
with we say that the susceptibles are permanent. 𝑆(𝑛ℎ+ ℎ) −𝑆(𝑛ℎ)
𝜙(ℎ)
. (6) (6) 2. Discretization of the Continuous Model We start with a nonautonomous SIRVS model that is slightly
less general than the one considered in [8] and generalizes the
one in [21]. Namely, we consider the model: 𝑆= Λ (𝑡) −𝛽(𝑡) 𝜑(𝑆) 𝐼−(𝜇(𝑡) + 𝑝(𝑡)) 𝑆+ 𝜂(𝑡) 𝑉
𝐼= [𝛽(𝑡) 𝜑(𝑆) + 𝜎(𝑡) 𝜓(𝑉) −𝜇(𝑡) −𝛼(𝑡) −𝛾(𝑡)] 𝐼
𝑅= 𝛾(𝑡) 𝐼−𝜇(𝑡) 𝑅
(2) g
yf
The model we will consider is the following: 𝑆𝑛+1 −𝑆𝑛= Λ 𝑛−𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛) −(𝜇𝑛+ 𝑝𝑛) 𝑆𝑛+1
+ 𝜂𝑛𝑉𝑛+1 (2) 𝐼𝑛+1 −𝐼𝑛= 𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛) + 𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛)
−(𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛) 𝐼𝑛+1 𝑉= 𝑝(𝑡) 𝑆−(𝜇(𝑡) + 𝜂(𝑡)) 𝑉−𝜎(𝑡) 𝜓(𝑉) 𝐼. (1) We assume that the functions Λ, 𝜇, 𝑝, 𝜂, 𝛼, 𝛽, 𝜎, and 𝛾
belong to the class 𝐶1(R+
0) and are nonnegative and bounded. We also require that 𝑅𝑛+1 −𝑅𝑛= 𝛾𝑛𝐼𝑛+1 −𝜇𝑛𝑅𝑛+1 𝑉𝑛+1 −𝑉𝑛= 𝑝𝑛𝑆𝑛+1 −(𝜇𝑛+ 𝜂𝑛) 𝑉𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) , 𝑉𝑛+1 −𝑉𝑛= 𝑝𝑛𝑆𝑛+1 −(𝜇𝑛+ 𝜂𝑛) 𝑉𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) , (C1) the functions 𝜑: R →R and 𝜓: R →R are non-
negative, nondecreasing, differentiable, and Lipschitz
with Lipschitz constants 𝑘𝜑and 𝑘𝜓, respectively; 𝑛∈N, where the classes 𝑆, 𝐼, 𝑅, and 𝑉correspond, respec-
tively, to susceptible, infective, recovered, and vaccinated
individuals and the parameter functions have the following
meanings: Λ 𝑛denotes the inflow of newborns in the sus-
ceptible class; the function 𝛽𝑛𝜑is the incidence (into the 𝑛∈N, where the classes 𝑆, 𝐼, 𝑅, and 𝑉correspond, respec-
tively, to susceptible, infective, recovered, and vaccinated
individuals and the parameter functions have the following
meanings: Λ 𝑛denotes the inflow of newborns in the sus-
ceptible class; the function 𝛽𝑛𝜑is the incidence (into the (C2) 𝜑(0) = 𝜓(0) = 0; (C3) there is 𝜔> 0 such that lim inf𝑡→+∞∫
𝑡+𝜔
𝑡
𝜇(𝑠)𝑑𝑠> 0. Discrete Dynamics in Nature and Society 3 Writing 𝑆𝑛= 𝑆(𝑛ℎ), 𝐼𝑛= 𝐼(𝑛ℎ), 𝑉𝑛= 𝑉(𝑛ℎ), Λ 𝑛= 𝜙(ℎ)Λ(𝑛ℎ),
𝛽𝑛= 𝜙(ℎ)𝛽(𝑛ℎ), 𝜇𝑛= 𝜙(ℎ)𝜇(𝑛ℎ), 𝑝𝑛= 𝜙(ℎ)𝑝(𝑛ℎ), and 𝜂𝑛=
𝜙(ℎ)𝜂(𝑛ℎ), we have Writing 𝑆𝑛= 𝑆(𝑛ℎ), 𝐼𝑛= 𝐼(𝑛ℎ), 𝑉𝑛= 𝑉(𝑛ℎ), Λ 𝑛= 𝜙(ℎ)Λ(𝑛ℎ),
𝛽𝑛= 𝜙(ℎ)𝛽(𝑛ℎ), 𝜇𝑛= 𝜙(ℎ)𝜇(𝑛ℎ), 𝑝𝑛= 𝜙(ℎ)𝑝(𝑛ℎ), and 𝜂𝑛=
𝜙(ℎ)𝜂(𝑛ℎ), we have In order to obtain threshold conditions for model (2), it was
considered in [8] the following auxiliary system: 𝑥= Λ (𝑡) −[𝜇(𝑡) + 𝑝(𝑡)] 𝑥+ 𝜂(𝑡) 𝑦
𝑦= 𝑝(𝑡) 𝑥−[𝜇(𝑡) + 𝜂(𝑡)] 𝑦. (3) 𝑆𝑛+1 −𝑆𝑛= Λ 𝑛−𝛽𝑛𝜑(𝑆𝑛+1) 𝐼𝑛−(𝜇𝑛+ 𝑝𝑛) 𝑆𝑛+1
+ 𝜂𝑛𝑉𝑛+1. 2. Discretization of the Continuous Model (8) (3) (8) And for each solution (𝑥∗(𝑡), 𝑦∗(𝑡)) of (3) with positive initial
conditions, it was shown that the numbers Proceeding similarly for the other equations, we obtain the
following discrete model: Proceeding similarly for the other equations, we obtain the
following discrete model: 𝑅ℓ
𝐶(𝜆) = lim inf
𝑡→∞∫
𝑡+𝜆
𝑡
𝛽(𝑠) 𝜑(𝑥∗(𝑠)) + 𝜎(𝑠) 𝜓(𝑦∗(𝑠)) 𝑆𝑛+1 −𝑆𝑛= Λ 𝑛−𝛽𝑛𝜑(𝑆𝑛+1) 𝐼𝑛−(𝜇𝑛+ 𝑝𝑛) 𝑆𝑛+1
+ 𝜂𝑛𝑉𝑛+1
𝐼𝑛+1 −𝐼𝑛= 𝛽𝑛𝜑(𝑆𝑛+1) 𝐼𝑛+ 𝜎𝑛𝜓(𝑉𝑛+1) 𝐼𝑛
−(𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛) 𝐼𝑛+1
𝑅𝑛+1 −𝑅𝑛= 𝛾𝑛𝐼𝑛+1 −𝜇𝑛𝑅𝑛+1
𝑉𝑛+1 −𝑉𝑛= 𝑝𝑛𝑆𝑛+1 −(𝜇𝑛+ 𝜂𝑛) 𝑉𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1) 𝐼𝑛,
(9) 𝑆𝑛+1 −𝑆𝑛= Λ 𝑛−𝛽𝑛𝜑(𝑆𝑛+1) 𝐼𝑛−(𝜇𝑛+ 𝑝𝑛) 𝑆𝑛+1 𝑆𝑛+1 −𝑆𝑛= Λ 𝑛−𝛽𝑛𝜑(𝑆𝑛+1) 𝐼𝑛−(𝜇𝑛+ 𝑝𝑛) 𝑆𝑛+1
+ 𝜂𝑛𝑉𝑛+1 −𝜇(𝑠) −𝛼(𝑠) −𝛾(𝑠) 𝑑𝑠,
𝑅𝑢
𝐶(𝜆) = lim sup
𝑡→∞
∫
𝑡+𝜆
𝑡
𝛽(𝑠) 𝜑(𝑥∗(𝑠)) + 𝜎(𝑠) 𝜓(𝑦∗(𝑠))
−𝜇(𝑠) −𝛼(𝑠) −𝛾(𝑠) 𝑑𝑠
(4) −𝜇(𝑠) −𝛼(𝑠) −𝛾(𝑠) 𝑑𝑠, (4) 𝐼𝑛+1 −𝐼𝑛= 𝛽𝑛𝜑(𝑆𝑛+1) 𝐼𝑛+ 𝜎𝑛𝜓(𝑉𝑛+1) 𝐼𝑛
−(𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛) 𝐼𝑛+1
𝑅𝑛+1 −𝑅𝑛= 𝛾𝑛𝐼𝑛+1 −𝜇𝑛𝑅𝑛+1
𝑉𝑛+1 −𝑉𝑛= 𝑝𝑛𝑆𝑛+1 −(𝜇𝑛+ 𝜂𝑛) 𝑉𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1) 𝐼𝑛,
(9) 𝑅𝑢
𝐶(𝜆) = lim sup
𝑡→∞
∫
𝑡+𝜆
𝑡
𝛽(𝑠) 𝜑(𝑥∗(𝑠)) + 𝜎(𝑠) 𝜓(𝑦∗(𝑠)) (9) 𝑅𝑛+1 −𝑅𝑛= 𝛾𝑛𝐼𝑛+1 −𝜇𝑛𝑅𝑛+1 are independent of the particular solution. are independent of the particular solution. Using the above numbers, the following results are con-
tained in results obtained in [8]. 𝑛∈N0. We will consider a model that contains this one
to obtain some of our results. Namely, based on model (9),
in Sections 3 and 4 we will study model (1) that has a more
general form for the incidence function.i Theorem 1 (Theorem 1 of [8]). Assume that conditions (C1),
(C2), and (C3) hold. Then, if there is a constant 𝜆> 0 such
that 𝑅ℓ
𝐶(𝜆) > 0, then the infectives 𝐼are permanent in system
(2). Now, we need to make some definitions. We say that 3.1. Auxiliary Results. Consider the auxiliary system, 3.1. Auxiliary Results. Consider the auxiliary system, 3. Permanence and Extinction in
the Discrete Model After deciding a nonlocal representation for the incidence
function and that terms that do not correspond to an
interaction will be considered in the 𝑛+ 1 time, the first
equation in (2) becomes In this section, we will extend the results obtained for the
model with the usual mass action incidence in [20] to our
generalized family of models. Namely, suitable thresholds are
defined and conditions for persistence and extinction of the
disease are obtained. As a corollary of our results, we consider
the periodic case where we have a unique number that
establishes the boundary between the regions of permanence
and extinction. Although the proofs of our results are inspired
in [20], some difficulties must be dealt with. In particular, it
was necessary to understand the right conditions to impose 𝑆((𝑛+ 1) ℎ) −𝑆(𝑛ℎ) = 𝜙(ℎ) [Λ (𝑛ℎ)
−𝛽(𝑛ℎ) 𝜑(𝑆((𝑛+ 1) ℎ)) 𝐼(𝑛ℎ) 𝑆((𝑛+ 1) ℎ) −𝑆(𝑛ℎ) = 𝜙(ℎ) [Λ (𝑛ℎ)
−𝛽(𝑛ℎ) 𝜑(𝑆((𝑛+ 1) ℎ)) 𝐼(𝑛ℎ)
−(𝜇(𝑛ℎ) + 𝑝(𝑛ℎ)) 𝑆((𝑛+ 1) ℎ) −𝛽(𝑛ℎ) 𝜑(𝑆((𝑛+ 1) ℎ)) 𝐼(𝑛ℎ) (7) −(𝜇(𝑛ℎ) + 𝑝(𝑛ℎ)) 𝑆((𝑛+ 1) ℎ) + 𝜂(𝑛ℎ) 𝑉(𝑛ℎ+ ℎ)] . 4 Discrete Dynamics in Nature and Society to the incidence functions in order to overcome the technical
difficulties. (v) when system (12) is 𝜔–periodic, it has a unique positive
𝜔–periodic solution which is globally uniformly attrac-
tive. fi
To lighten the reading, the proofs of our results are
presented in the appendix. Lemma 4. Assume that condition (H5) holds. Then we have
the following: 𝑥𝑛+1 = Λ 𝑛+ 𝜂𝑛𝑦𝑛+1 + 𝑥𝑛
1 + 𝜇𝑛+ 𝑝𝑛
𝑦𝑛+1 = 𝑝𝑛𝑥𝑛+1 + 𝑦𝑛
1 + 𝜇𝑛+ 𝜂𝑛
. (12) (i) all solutions (𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with nonnegative
initial conditions are nonnegative for all 𝑛∈N0; (12) (ii) all solutions (𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with positive initial
conditions are positive for all 𝑛∈N0; Note that the auxiliary system describes the behaviour of the
system in the absence of infection. If (Λ 𝑛), (𝜇𝑛), (𝑝𝑛), (𝜂𝑛),
(𝛼𝑛), (𝜇𝑛), (𝜎𝑛), and (𝛽𝑛) are constant sequences, then the
linear system (12) becomes autonomous and corresponds to
the linearization of the equations for (𝑆𝑛) and (𝑉𝑛) in the
classical (autonomous) SIRVS model. Note that the auxiliary system describes the behaviour of the
system in the absence of infection. If (Λ 𝑛), (𝜇𝑛), (𝑝𝑛), (𝜂𝑛),
(𝛼𝑛), (𝜇𝑛), (𝜎𝑛), and (𝛽𝑛) are constant sequences, then the
linear system (12) becomes autonomous and corresponds to
the linearization of the equations for (𝑆𝑛) and (𝑉𝑛) in the
classical (autonomous) SIRVS model. (iii) there is a constant 𝑀> 0 such that, if (𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛)
is a solution of (1) with nonnegative initial conditions,
then lim sup
𝑛→+∞𝑆𝑛+ 𝐼𝑛+ 𝑅𝑛+ 𝑉𝑛< 𝑀. (16) (16) In order to proceed we need to recall some notions. A
solution (𝑢𝑛) of some system of difference equations 𝑢𝑛+1 =
𝑓𝑛(𝑢𝑛) is said to be attractive if for all 𝑛0 ∈N and all 𝜀> 0
there is 𝜎(𝑛0) > 0 and 𝑇(𝜀, 𝑛0, 𝑢0) ∈N such that if (𝑢𝑛) is
a solution with ‖𝑢0 −𝑢0‖ < 𝜎(𝑛0) then ‖𝑢𝑛−𝑢𝑛‖ < 𝜀, for all
𝑛⩾𝑛0+𝑇(𝜀, 𝑛0, 𝑢0). Additionally, if some solution is attractive
and we can take 𝑇to be only dependent on 𝜀, we say that it is
uniformly attractive.h Proof. See the appendix. For each 𝜆and each particular solution 𝜉∗
𝑛= (𝑥∗
𝑛, 𝑦∗
𝑛) of
(12) with 𝑥∗
0 > 0 and 𝑦∗
0 > 0 we define the numbers Rℓ
𝐷(𝜉∗, 𝜆)
= lim inf
𝑛→∞
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0)
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
,
(17)
R𝑢
𝐷(𝜉∗, 𝜆)
= lim sup
𝑛→∞
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0)
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
,
(18) (17) The following theorem furnishes some simple properties
of system (12). Lemma 3 (lemma 2.2 of [20]). Assume that conditions (H3)
and (H4) hold. Then Lemma 3 (lemma 2.2 of [20]). Assume that conditions (H3)
and (H4) hold. Then (i) all solutions (𝑥𝑛, 𝑦𝑛) of system (12) with initial condi-
tion 𝑥0 ⩾0 and 𝑦0 ⩾0 are nonnegative for all 𝑛∈N0; where 𝜕𝑖𝑓denotes the partial derivative of 𝑓with respect to
the 𝑖-th variable. Contrarily to what one could expect, the next
lemma shows that the numbers above do not depend on the
particular solution 𝜉𝑛= (𝑥𝑛, 𝑦𝑛) of (12) with 𝑥𝑛(0) > 0 and
𝑦𝑛(0) > 0. (ii) each fixed solution (𝑥𝑛, 𝑦𝑛) of (12) is bounded and
globally uniformly attractive for all 𝑛∈N0; (iii) if (𝑥𝑛, 𝑦𝑛) is a solution of (12) and (̃𝑥𝑛, ̃𝑦𝑛) is a solution
of the system Lemma 5. Assume that (H1), (H3), and (H4) hold. If (𝜉∗
1 )𝑛=
((𝑥1)∗
𝑛, (𝑦1)∗
𝑛) and (𝜉∗
2 )𝑛= ((𝑥2)∗
𝑛, (𝑦2)∗
𝑛) are two solutions of
(12) with 𝑥∗
𝑖(0) > 0 and 𝑦∗
𝑖(0) > 0, 𝑖= 1, 2, then Lemma 5. Assume that (H1), (H3), and (H4) hold. If (𝜉∗
1 )𝑛=
((𝑥1)∗
𝑛, (𝑦1)∗
𝑛) and (𝜉∗
2 )𝑛= ((𝑥2)∗
𝑛, (𝑦2)∗
𝑛) are two solutions of
(12) with 𝑥∗
𝑖(0) > 0 and 𝑦∗
𝑖(0) > 0, 𝑖= 1, 2, then 𝑥𝑛+1 = Λ 𝑛+ 𝜂𝑛𝑦𝑛+1 + 𝑥𝑛+ 𝑓𝑛
1 + 𝜇𝑛+ 𝑝𝑛
𝑦𝑛+1 = 𝑝𝑛𝑥𝑛+1 + 𝑦𝑛+ 𝑔𝑛
1 + 𝜇𝑛+ 𝜂𝑛
,
(13) (13) Rℓ
𝐷(𝜉∗
1 , 𝜆) = Rℓ
𝐷(𝜉∗
2 , 𝜆) ,
R𝑢
𝐷(𝜉∗
1 , 𝜆) = R𝑢
𝐷(𝜉∗
2 , 𝜆) . (19) (19) with (̃𝑥0, ̃𝑦0) = (𝑥0, 𝑦0) then there is a constant 𝐿> 0,
only depending on 𝜇𝑛, satisfying with (̃𝑥0, ̃𝑦0) = (𝑥0, 𝑦0) then there is a constant 𝐿> 0,
only depending on 𝜇𝑛, satisfying Proof. We have the following lemma. 3.1. Auxiliary Results. Consider the auxiliary system, Therefore, we obtain the corollary. Therefore, we obtain the corollary. (23) Corollary 8 (periodic case). Assume that all coefficients are
𝜔-periodic in (1) and that conditions (H1) to (H5) hold. Then (a) if R𝑝𝑒𝑟
𝐷(𝜉∗) < 1 then the infectives (𝐼𝑛) go to extinction; (b) the disease-free solution (𝑥∗
𝑛, 0, 0, 𝑦∗
𝑛), where (𝑥∗
𝑛,
𝑦∗
𝑛)𝑛∈N is an disease-free 𝜔-periodic solution of (12), is
globally attractive; Then, (a) if R𝑢
𝐶(𝜆) < 0, then R𝑢
𝐷(⌊𝜆/ℎ⌋, ℎ) < 1 for all ℎ∈]0,
ℎ𝑢
𝑚𝑎𝑥[;
(b) if Rℓ
𝐶(𝜆) > 0, then Rℓ
𝐷(⌊𝜆/ℎ⌋, ℎ) > 1 for all for all ℎ∈
]0, ℎℓ
𝑚𝑎𝑥[. (c) if R𝑝𝑒𝑟
𝐷(𝜉∗) > 1, then the infectives (𝐼𝑛) are permanent. Proof. See the Appendix. Proof. Observe that (𝑥𝑛, 𝑦𝑛) = (𝑎, 𝑏), 𝑛∈N and (𝑥(𝑡), 𝑦(𝑡)) =
(𝑎, 𝑏), 𝑡∈R, where Proof. Observe that (𝑥𝑛, 𝑦𝑛) = (𝑎, 𝑏), 𝑛∈N and (𝑥(𝑡), 𝑦(𝑡)) =
(𝑎, 𝑏), 𝑡∈R, where (a) if there is a constant 𝜆> 0 such that R𝑢
𝐷(𝜆) < 1, then
the infectives (𝐼𝑛) go to extinction; (a) if there is a constant 𝜆> 0 such that R𝑢
𝐷(𝜆) < 1, then
the infectives (𝐼𝑛) go to extinction; (b) any solution (𝑥∗
𝑛, 0, 0, 𝑦∗
𝑛), where (𝑥∗
𝑛, 𝑦∗
𝑛) is a par-
ticular solution of system (12), is globally uniformly
attractive. (b) any solution (𝑥∗
𝑛, 0, 0, 𝑦∗
𝑛), where (𝑥∗
𝑛, 𝑦∗
𝑛) is a par-
ticular solution of system (12), is globally uniformly
attractive. Rℓ
𝐷(𝜆, ℎ) = lim inf
𝑛→∞
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽ℎ
𝑘𝜑(𝑥∗
𝑘+1) + 𝜎ℎ
𝑘𝜓(𝑦∗
𝑘+1)
1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘
,
R𝑢
𝐷(𝜆, ℎ) = lim sup
𝑛→∞
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽ℎ
𝑘𝜑(𝑥∗
𝑘+1) + 𝜎ℎ
𝑘𝜓(𝑦∗
𝑘+1)
1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘
,
(21) (21) Proof. See the Appendix. Proof. See the appendix. See the Appendix. Proof. See the Appendix. sup
𝑛∈N0
{̃𝑥𝑛−𝑥𝑛+ ̃𝑦𝑛−𝑦𝑛} ⩽𝐿sup
𝑛∈N0
(𝑓𝑛+ 𝑔𝑛) ;
(14) By Lemma 5 we can drop the dependence of the partic-
ular solution and simply write Rℓ
𝐷(𝜆) and R𝑢
𝐷(𝜆) instead of
Rℓ
𝐷(𝜉∗, 𝜆) an R𝑢
𝐷(𝜉∗, 𝜆), respectively. By Lemma 5 we can drop the dependence of the partic-
ular solution and simply write Rℓ
𝐷(𝜆) and R𝑢
𝐷(𝜆) instead of
Rℓ
𝐷(𝜉∗, 𝜆) an R𝑢
𝐷(𝜉∗, 𝜆), respectively. (14) (iv) there exist constants 𝑚, 𝑀> 0 such that, for each
solution (𝑥𝑛, 𝑦𝑛) of (12), we have (iv) there exist constants 𝑚, 𝑀> 0 such that, for each
solution (𝑥𝑛, 𝑦𝑛) of (12), we have 3.2. Extinction and Permanence. We have the following result
about the extinction of the disease. 𝑚⩽lim inf
𝑛→∞𝑥𝑛⩽lim sup
𝑛→∞
𝑥𝑛⩽𝑀, (15) Theorem 6 (extinction of the disease). Assume that condi-
tions (H1) to (H5) hold. Then 5 Discrete Dynamics in Nature and Society For a given time step ℎ> 0, the expressions Rℓ
𝐷(𝜆) and
R𝑢
𝐷(𝜆) in (17) and (18) become, in our context, Proof. See the Appendix. We have the following result about the permanence of the
disease. where (𝑥∗
𝑘, 𝑦∗
𝑘) is the solution of the (in our context
autonomous) system (12). where (𝑥∗
𝑘, 𝑦∗
𝑘) is the solution of the (in our context
autonomous) system (12). Theorem 7 (permanence of the disease). Assume that condi-
tions (H1) to (H5) hold. If there is a constant 𝜆> 0 such that
Rℓ
𝐷(𝜆) > 1 then the infectives (𝐼𝑛) are permanent in system (1). We have the following result. Theorem 9. For system (2), assume that Λ(𝑡) = Λ, 𝜇(𝑡) = 𝜇,
𝜂(𝑡) = 𝜂, and 𝑝(𝑡) = 𝑝for all 𝑡⩾0 and that the functions
𝛼(𝑡), 𝛾(𝑡), 𝛽(𝑡), and 𝜎(𝑡) are differentiable, nonnegative, and
bounded and have bounded derivative. Assume also that
conditions (C1) to (C3) hold and let Proof. See the Appendix. Proof. See the Appendix. We consider now the particular periodic case: assume
that all parameters of system (1) are periodic with period
𝜔∈N. By (v) in Lemma 3, there is an 𝜔-periodic disease-
free solution of (12), 𝜉∗= (𝑥∗
𝑛, 𝑦∗
𝑛)𝑛∈N. Thus, in the periodic
setting, (17) and (18) become both equal to ℎ𝑢
𝑚𝑎𝑥= −
𝑅𝑢
𝐶(𝜆)
sup𝑡⩾0 𝑓(𝑡)(𝜆+ 1),
ℎℓ
𝑚𝑎𝑥=
𝑅ℓ
𝐶(𝜆)
sup𝑡⩾0 𝑓(𝑡)(𝜆+ 1),
(22) ℎ𝑢
𝑚𝑎𝑥= −
𝑅𝑢
𝐶(𝜆)
sup𝑡⩾0 𝑓(𝑡)(𝜆+ 1), (22) R𝑝𝑒𝑟
𝐷(𝜉∗)
=
𝜔−1
∏
𝑘=0
1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0)
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
. (20)
ℎℓ
𝑚𝑎𝑥=
𝑅ℓ
𝐶(𝜆)
sup𝑡⩾0 𝑓(𝑡)(𝜆+ 1),
where
(
Λ (𝜇+ 𝜂)
) R𝑝𝑒𝑟
𝐷(𝜉∗)
=
𝜔−1
∏
𝑘=0
1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0)
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
. (20) (20) where where 𝑓(𝑡) = 𝛽(𝑡) 𝜑(
Λ (𝜇+ 𝜂)
𝜇(𝜇+ 𝜂+ 𝑝))
+ 𝜎(𝑡) 𝜓(
𝑝Λ
𝜇(𝜇+ 𝜂+ 𝑝)) −𝜇−𝛼(𝑡)
−𝛾(𝑡) . (23)h 𝑓(𝑡) = 𝛽(𝑡) 𝜑(
Λ (𝜇+ 𝜂)
𝜇(𝜇+ 𝜂+ 𝑝)) Therefore, we obtain the corollary. 4. Consistency Discrete Dynamics in Nature and Society 6 y contradiction, assume that
We also have By contradiction, assume that By contradiction, assume that We also have By contradiction, assume that
R𝑢
𝐶(𝜆) < 0,
(26)
and that there is a sequence (ℎ𝑚)𝑚∈N such that ℎ𝑚→0 as
𝑚→+∞and
R𝑢
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚)
(
)
We also have
𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑
𝑘=2
(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏))]
𝑘[𝜙(ℎ𝑚)]
𝑘
⩽ℎ
⌊𝜆/ℎ𝑚⌋
∑(⌊𝜆
ℎ⌋) [(𝛽𝑢( ) + 𝜎𝑢𝜓(𝑏))]𝑘[𝜙(ℎ)]
𝑘
(34) By contradiction, assume that
R𝑢
𝐶(𝜆) < 0,
(26)
and that there is a sequence (ℎ𝑚)𝑚∈N such that ℎ𝑚→0 as
𝑚→+∞and
We also have
𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑
𝑘2
(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏))]
𝑘[𝜙(ℎ𝑚)]
𝑘 R𝑢
𝐶(𝜆) < 0,
(26)
𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
R𝑢
𝐶(𝜆) < 0,
(26)
𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
R𝑢
𝐶(𝜆) < 0,
(26)
𝐴𝑛𝑚 R𝑢
𝐶(𝜆) < 0, (26) and that there is a sequence (ℎ𝑚)𝑚∈N such that ℎ𝑚→0 as
𝑚→+∞and
⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏))]
𝑘[𝜙(ℎ𝑚)]
𝑘 and that there is a sequence (ℎ𝑚)𝑚∈N such that ℎ𝑚→0 as
𝑚→+∞and
⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑(⌊𝜆
ℎ𝑚
⌋) [(𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏))]
𝑘[𝜙(ℎ𝑚)]
𝑘 and that there is a sequence (ℎ𝑚)𝑚∈N such that ℎ𝑚→0 as
𝑚→+∞and
⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝛽𝑢𝜑(𝑎) R𝑢
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚) R𝑢
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚)
= lim sup
𝑛→∞
𝑛+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛
1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏)
1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘
⩾1,
(27)
𝑘=2
(
𝑘
)
⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑
𝑘=0
(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏))]𝑘[𝜙(ℎ𝑚)]
𝑘
= ℎ[1 + (𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)) 𝜙(ℎ)]
⌊𝜆/ℎ𝑚⌋ (34) ⩽ℎ𝑚
⌊𝜆/ℎ𝑚⌋
∑
𝑘=0
(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏))]𝑘[𝜙(ℎ𝑚)]
𝑘 (27) (27) = lim sup
𝑛→∞
𝑛+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛
1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏)
1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘
⩾1, = ℎ𝑚[1 + (𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)) 𝜙(ℎ𝑚)]
⌊𝜆/ℎ𝑚⌋. = ℎ𝑚[1 + (𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)) 𝜙(ℎ𝑚)]
⌊𝜆/ℎ𝑚⌋. for all 𝑚∈N. By (27), we conclude that, for each 𝑚∈N,
there are sequences (ℎ𝑚)𝑚∈N and (𝑛𝑚,𝑟)𝑟∈N such that ℎ𝑚→0
as 𝑚→+∞, 𝑛𝑚,𝑟→+∞as 𝑟→+∞and
Noting that, by (5), we have
lim
𝑚→+∞[1 + (𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)) 𝜙 for all 𝑚∈N. 4. Consistency By (27), we conclude that, for each 𝑚∈N,
there are sequences (ℎ𝑚)𝑚∈N and (𝑛𝑚,𝑟)𝑟∈N such that ℎ𝑚→0
as 𝑚→+∞𝑛
→+∞as 𝑟→+∞and Noting that, by (5), we have Noting that, by (5), we have lim
𝑚→+∞[1 + (𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)) 𝜙(ℎ𝑚)]⌊𝜆/ℎ𝑚⌋ ,
as 𝑚→+∞, 𝑛𝑚,𝑟→+∞as 𝑟→+∞and 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛𝑚,𝑟
(1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏)) 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛𝑚,𝑟
(1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏))
=
lim
𝑚→+∞[(1 =
lim
𝑚→+∞[(1 (28) (28) (35) [(
+ 𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)
1/𝜙(ℎ𝑚)
)
1/𝜙(ℎ𝑚)
]
𝜙(ℎ𝑚)⌊𝜆/ℎ𝑚⌋
= e(𝛽𝑢𝜑(𝑎)+𝜎𝑢𝜓(𝑏))𝜆
(35) > (1 −ℎ𝑚)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛𝑚,𝑟
(1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘) . > (1 −ℎ𝑚)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛𝑚,𝑟
(1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘) . (28)
28), we have
+ 𝛽𝑢𝜑(𝑎) + 𝜎𝑢𝜓(𝑏)
1/𝜙(ℎ𝑚)
)
1/𝜙(ℎ𝑚)
]
𝜙(ℎ𝑚)⌊𝜆/ℎ𝑚⌋
= e(𝛽𝑢𝜑(𝑎)+𝜎𝑢𝜓(𝑏))𝜆
(35) By (28), we have By (28), we have = e(𝛽𝑢𝜑(𝑎)+𝜎𝑢𝜓(𝑏))𝜆 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
(𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏) −𝜇ℎ𝑚
𝑘−𝛼ℎ𝑚
𝑘−𝛾ℎ𝑚
𝑘)
>
(𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚)
ℎ𝑚
,
(29)
and that a convergent sequence is bounded; by (34) there is
𝐶1 > 0 such that
𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶1ℎ𝑚. (36)
Similarly, we have and that a convergent sequence is bounded; by (34) there is
𝐶1 > 0 such that 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
(𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏) −𝜇ℎ𝑚
𝑘−𝛼ℎ𝑚
𝑘−𝛾ℎ𝑚
𝑘)
and that a convergent sequence is bounded; by (34) there is
𝐶1 > 0 such that g
𝐶1 > 0 such that )
(29)
𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶1ℎ𝑚. (36) 𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶1ℎ𝑚. (36) (29) (36) Similarly, we have 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽
⌊𝜆/ℎ𝑚⌋
∑
𝑘=2
(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢)]
𝑘[𝜙(ℎ𝑚)]
𝑘
⩽
⌊𝜆/ℎ𝑚⌋
∑
𝑘=0
(⌊𝜆
ℎ𝑚
⌋
𝑘
) [(𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢)]𝑘[𝜙(ℎ𝑚)]𝑘
(37) where where 𝐴𝑛,𝐿,ℎfl−ℎ+ ℎ
𝑛+⌊𝐿/ℎ⌋
∏
𝑘=𝑛
(1 + 𝛽ℎ
𝑘𝜑(𝑎) + 𝜎ℎ
𝑘𝜓(𝑏))
−ℎ
𝑛+⌊𝐿/ℎ⌋
∑
𝑘=𝑛
(𝛽ℎ
𝑘𝜑(𝑎) + 𝜎ℎ
𝑘𝜓(𝑏)) ,
𝐵𝑛,𝐿,ℎfl−ℎ+ ℎ
𝑛+⌊𝐿/ℎ⌋
∏
𝑘=𝑛
(1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘)
−ℎ
𝑛+⌊𝐿/ℎ⌋
∑
𝑘=𝑛
(𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘) ,
𝐶𝑛,𝐿,ℎ= ℎ
𝑛+⌊𝐿/ℎ⌋
∏
𝑘=𝑛
(1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘) . 𝐴𝑛,𝐿,ℎfl−ℎ+ ℎ
𝑛+⌊𝐿/ℎ⌋
∏
𝑘=𝑛
(1 + 𝛽ℎ
𝑘𝜑(𝑎) + 𝜎ℎ
𝑘𝜓(𝑏)) (30) (37) −ℎ
𝑛+⌊𝐿/ℎ⌋
∑
𝑘=𝑛
(𝛽ℎ
𝑘𝜑(𝑎) + 𝜎ℎ
𝑘𝜓(𝑏)) , 𝐵𝑛,𝐿,ℎfl−ℎ+ ℎ
𝑛+⌊𝐿/ℎ⌋
∏
𝑘=𝑛
(1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘) = ℎ𝑚[1 + (𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢) 𝜙(ℎ𝑚)]
⌊𝜆/ℎ𝑚⌋. = ℎ𝑚[1 + (𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢) 𝜙(ℎ𝑚)]
⌊𝜆/ℎ𝑚⌋. 4. Consistency In this section, under the additional assumption that the
parameter functions Λ, 𝜇, 𝜂, and 𝑝are constant, we will get
a result stating that when our integral conditions prescribe
extinction (persistence) for the continuous-time model, then
the discrete-time conditions prescribe extinction (persis-
tence) for the corresponding discrete-time models, as long
as the time step is less than some constant. Throughout this
section, we assume that the parameter functions Λ, 𝜇, 𝜂, and
𝑝are constant functions and 𝜙(ℎ) will be the function used
in the discretization of the derivative. (𝑎, 𝑏) = (
Λ (𝜇+ 𝜂)
[𝜇(𝜇+ 𝜂+ 𝑝)],
𝑝Λ
[𝜇(𝜇+ 𝜂+ 𝑝)]) ,
(24) (24) are, respectively, solutions of system (12) and system (3). Thus, are, respectively, solutions of system (12) and system (3). Thus, R𝑢
𝐶(𝜆) = lim sup
𝑡→∞
∫
𝑡+𝜆
𝑡
𝛽(𝑠) 𝜑(𝑎) + 𝜎(𝑠) 𝜓(𝑏) −[𝜇(𝑠) + 𝛼(𝑠) + 𝛾(𝑠)] 𝑑𝑠, R𝑢
𝐷(⌊𝜆
ℎ⌋, ℎ)
(25) (25) We consider the continuous-time model (2) and, for a
given time step ℎ, the corresponding discrete-time model,
that is, the discrete-time model with parameters 𝛽ℎ
𝑘
=
𝜙(ℎ)𝛽(𝑘ℎ), 𝜎ℎ
𝑘= 𝜙(ℎ)𝜎(𝑘ℎ), Λℎ
𝑘= 𝜙(ℎ)Λ, 𝜇ℎ
𝑘= 𝜙(ℎ)𝜇, 𝑝ℎ
𝑘=
𝜙(ℎ)𝑝, 𝜂ℎ
𝑘= 𝜙(ℎ)𝜂, 𝛼ℎ
𝑘= 𝜙(ℎ)𝛼(𝑘ℎ), and 𝛾ℎ
𝑘= 𝜙(ℎ)𝛾(𝑘ℎ). = lim sup
𝑛→∞
𝑛+⌊𝜆/ℎ⌋
∏
𝑘=𝑛
1 + 𝛽ℎ
𝑘𝜑(𝑎) + 𝜎ℎ
𝑘𝜓(𝑏)
1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘
. 4. Consistency (42)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) + 𝑠𝑚,𝑟−∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠
2 ⌊𝜆⌋
2
(5 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶3ℎ𝑚. (42) (51) Thus, Thus, 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽(𝐶1 + 𝐶2 + 𝐶3) ℎ𝑚,
(43)
< 𝐶ℎ2
𝑚⌊𝜆
ℎ𝑚
⌋+ 𝐶ℎ2
𝑚< 𝐶(𝜆+ 1) ℎ𝑚,
thus
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
ℎ𝑓(𝑘ℎ) < ∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑓(𝑠) 𝑑𝑠
𝑠 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽(𝐶1 + 𝐶2 + 𝐶3) ℎ𝑚,
(43)
< 𝐶ℎ2
𝑚⌊𝜆
ℎ𝑚
⌋+ 𝐶
thus
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
ℎ𝑓(𝑘ℎ 𝑛𝑚,𝑟,𝜆,ℎ𝑚
𝑛𝑚,𝑟,𝜆,ℎ𝑚
𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽(𝐶1 + 𝐶2 + 𝐶3) ℎ𝑚,
(43)
thus
𝑛𝑚,𝑟+⌊𝜆/ℎ
∑ ⩽𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
(
)
(43)
thus 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) < ∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠−𝑠𝑚,𝑟
+ 𝐶(𝜆+ 1) ℎ𝑚,
(52) ⩽(𝐶1 + 𝐶2 + 𝐶3) ℎ𝑚,
for all 𝑚⩾𝑀. Since the right hand side of (43) is independent
of 𝑛𝑚,𝑟, we conclude that
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) < ∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠−𝑠𝑚,𝑟
+ 𝐶(𝜆+ 1) ℎ𝑚,
(52) for all 𝑚⩾𝑀. Since the right hand side of (43) is independent
of 𝑛𝑚,𝑟, we conclude that
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) < ∫
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠−𝑠𝑚,𝑟
𝐶(𝜆
1) ℎ
(52) for all 𝑚⩾𝑀. Since the right hand side of (43) is independent
of 𝑛𝑚,𝑟, we conclude that
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) < ∫
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠
𝑠𝑚,𝑟
+ 𝐶(𝜆+ 1) ℎ
(52) for all 𝑚⩾𝑀. Since the right hand side of (43) is independent
of 𝑛𝑚,𝑟, we conclude that (52) + 𝐶(𝜆+ 1) ℎ𝑚, 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚→0,
(44) 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚→0,
(44)
and therefore (44) and therefore as 𝑚→+∞, uniformly in 𝑟. as 𝑚→+∞, uniformly in 𝑟. 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
𝜙(ℎ𝑚) ℎ𝑚𝑓(𝑘ℎ𝑚)
< 𝜙(ℎ𝑚) [∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠+ 𝐶(𝜆+ 1) ℎ𝑚] . (53) y
On the other hand, we note that the 𝐶1 function 𝑓: R+
0 →
R given by On the other hand, we note that the 𝐶1 function 𝑓: R+
0 →
R given by On the other hand, we note that the 𝐶1 function 𝑓: R+
0 →
R given by (53) 𝑓(𝑡) = 𝛽(𝑡) 𝜑(𝑎) + 𝜎(𝑡) 𝜓(𝑏) −𝜇(𝑡) −𝛼(𝑡) −𝛾(𝑡)
(45) is Riemann-integrable on any bounded interval 𝐼⊂R+
0. h
h is Riemann-integrable on any bounded interval 𝐼⊂R+
0. 4. Consistency (31) (31) (31)
Using (5) again, we get lim
𝑚→+∞[1 + (𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢) 𝜙(ℎ𝑚)]⌊𝜆/ℎ𝑚⌋
⩽e(𝜇𝑢+𝛼𝑢+𝛾𝑢)𝜆. (38) (38) 𝐶𝑛,𝐿,ℎ= ℎ
𝑛+⌊𝐿/ℎ⌋
∏
𝑘=𝑛
(1 + 𝜇ℎ
𝑘+ 𝛼ℎ
𝑘+ 𝛾ℎ
𝑘) . (32)
⩽e(𝜇𝑢+𝛼𝑢+𝛾𝑢)𝜆. There is 𝐶2 > 0 such that (32)
⩽e(𝜇𝑢+𝛼𝑢+𝛾𝑢)𝜆
There is 𝐶2 > 0 such that (32) 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶2ℎ𝑚. (39) and, multiplying both sides by ℎ𝑚, we get
𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶2ℎ𝑚. (39) and, multiplying both sides by ℎ𝑚, we get and, multiplying both sides by ℎ𝑚, we get (39) 𝜙(ℎ𝑚)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚[𝛽(𝑘ℎ𝑚) 𝜑(𝑎) + 𝜎(𝑘ℎ𝑚) 𝜓(𝑏)
−𝜇(𝑘ℎ𝑚) −𝛼(𝑘ℎ𝑚) −𝛾(𝑘ℎ𝑚)] > 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
(33)
Finally, we have
𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
= ℎ𝑚
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛𝑚,𝑟
(1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘)
(40 𝜙(ℎ𝑚)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚[𝛽(𝑘ℎ𝑚) 𝜑(𝑎) + 𝜎(𝑘ℎ𝑚) 𝜓(𝑏) Finally, we have (33) (40) = ℎ𝑚(1 + 3𝜙(ℎ𝑚) max {𝜇𝑢, 𝛼𝑢, 𝛾𝑢})⌊𝜆/ℎ𝑚⌋+1 . −𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚. = ℎ𝑚(1 + 3𝜙(ℎ𝑚) max {𝜇𝑢, 𝛼𝑢, 𝛾𝑢})⌊𝜆/ℎ𝑚⌋+1 . −𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚. Discrete Dynamics in Nature and Society
7 Discrete Dynamics in Nature and Society Discrete Dynamics in Nature and Society 7 we obtain
of size ℎ𝑚of the interval [𝑛𝑚,𝑟, 𝑛𝑚,𝑟+ 𝜆]. Note that According to (5), we obtain of size ℎ𝑚of the interval [𝑛𝑚,𝑟, 𝑛𝑚,𝑟+ 𝜆]. Note that of size ℎ𝑚of the interval [𝑛𝑚,𝑟, 𝑛𝑚,𝑟+ 𝜆]. Note that 𝑠𝑚,𝑟fl(𝜆−⌊𝜆
ℎ𝑚
⌋ℎ𝑚) 𝑓(𝑛𝑚,𝑟ℎ𝑚+ ⌊𝜆
ℎ𝑚
⌋ℎ𝑚)
(49) (49) =
lim
𝑚→+∞[(1
(41)
𝑚
𝑚
⩽ℎ𝑚𝑓𝑢fl𝑠𝑚
(49)
d
0
if
l i =
lim
𝑚→+∞[(1 ⩽ℎ𝑚𝑓𝑢fl𝑠𝑚 [(
+ 3 max {𝜇𝑢, 𝛼𝑢, 𝛾𝑢}
1/𝜙(ℎ𝑚)
)
1/𝜙(ℎ𝑚)
]
𝜙(ℎ𝑚)(⌊𝜆/ℎ𝑚⌋+1)
(41)
and 𝑠𝑚→0 as 𝑚→+∞, uniformly in 𝑟. Since 𝑓is 𝐶1 with bounded derivative, for any ℎ> 0 we
have (41) + 3 max {𝜇𝑢, 𝛼𝑢, 𝛾𝑢}
1/𝜙(ℎ𝑚)
)
1/𝜙(ℎ𝑚)
]
𝜙(ℎ𝑚)(⌊𝜆/ℎ𝑚⌋+1)
(41)
and 𝑠𝑚→0 as 𝑚→+∞, uniformly in 𝑟. Since 𝑓is 𝐶1 with bounded derivative, for any ℎ> 0 we
have
𝑓
𝑓
ℎ
ℎ 𝜙( 𝑚)
)
]
= e3 max{𝜇𝑢,𝛼𝑢,𝛾𝑢}𝜆. 𝑓(𝑥) −𝑓(𝑥+ ℎ)⩽𝐶ℎ,
(50) 𝑓(𝑥) −𝑓(𝑥+ ℎ)⩽𝐶ℎ,
(50) (50) = e3 max{𝜇𝑢,𝛼𝑢,𝛾𝑢}𝜆. where 𝐶= sup𝑡⩾0|𝑓(𝑡)|. We conclude that There is 𝐶3 > 0 such that 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶3ℎ𝑚. (42)
,
𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚−𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
(43)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) + 𝑠𝑚,𝑟−∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠
< 𝐶ℎ2
𝑚⌊𝜆
ℎ𝑚
⌋+ 𝐶ℎ2
𝑚< 𝐶(𝜆+ 1) ℎ𝑚,
(51)
thus
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚) + 𝑠𝑚,𝑟−∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠
< 𝐶ℎ2
𝑚⌊𝜆
ℎ𝑚
⌋+ 𝐶ℎ2
𝑚< 𝐶(𝜆+ 1) ℎ𝑚,
(51) 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚
⩽𝐶3ℎ𝑚. 4. Consistency In fact, noting
that 𝑥∗(𝑡) + 𝑦∗(𝑡) = 1, we get Rℓ
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚)
= lim inf
𝑛→∞
𝑛+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛
1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏)
1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘
⩽1,
(57)
that 𝑥∗(𝑡) + 𝑦∗(𝑡) = 1, we get
Rℓ
C (1) = ∫
1
0
𝛽(𝑠) 𝑥∗(𝑠) + 𝜎(𝑠) 𝑦∗(𝑠) −𝜇−𝛼−
= ∫
1
0
𝑑[1 + 𝑐sin2 (2𝜋𝐿𝑡) (1 + cos (2𝜋𝑡)) Rℓ
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚)
= lim inf
𝑛→∞
𝑛+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛
1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏)
1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘
⩽1,
(57) Rℓ
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚)
(
) Rℓ
𝐷(⌊𝜆
ℎ𝑚
⌋, ℎ𝑚) Rℓ
C (1) = ∫
1
0
𝛽(𝑠) 𝑥∗(𝑠) + 𝜎(𝑠) 𝑦∗(𝑠) −𝜇−𝛼−𝛾𝑑𝑠
∫
1 Rℓ
C (1) = ∫
1
0
𝛽(𝑠) 𝑥∗(𝑠) + 𝜎(𝑠) 𝑦∗(𝑠) −𝜇−𝛼−𝛾𝑑𝑠
= ∫
1
0
𝑑[1 + 𝑐sin2 (2𝜋𝐿𝑡) (1 + cos (2𝜋𝑡))] 𝑑𝑠
(63) (57) = lim inf
𝑛→∞
𝑛+⌊𝜆/ℎ𝑚⌋
∏
𝑘=𝑛
1 + 𝛽ℎ𝑚
𝑘𝜑(𝑎) + 𝜎ℎ𝑚
𝑘𝜓(𝑏)
1 + 𝜇ℎ𝑚
𝑘+ 𝛼ℎ𝑚
𝑘+ 𝛾ℎ𝑚
𝑘
⩽1, (63) −𝜇−𝛼−𝛾= 𝑑(1 + 𝑐
2) −𝜇−𝛼−𝛾. it is possible to conclude that it is possible to conclude that 𝜙(ℎ𝑚)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚(𝛽(𝑘ℎ𝑚) 𝜑(𝑎) + 𝜎(𝑘ℎ𝑚) 𝜓(𝑏)
−𝜇(𝑘ℎ𝑚) −𝛼(𝑘ℎ𝑚) −𝛾(𝑘ℎ𝑚)) < 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚
−𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚, We can also compute Rℓ
𝐷(1, 1/𝐿). Namely, we have )
(58)
Rℓ
𝐷(1, 1
𝐿) =
1 + 𝑑/𝐿
1 + 𝜇+ 𝛼+ 𝛾. (64) Rℓ
𝐷(1, 1
𝐿) =
1 + 𝑑/𝐿
1 + 𝜇+ 𝛼+ 𝛾. (64) (64) (58) If we let 𝑑be sufficiently small so that 𝑑< (𝜇+ 𝛼+ 𝛾)𝐿,
or in other words, 𝑑< (𝜇+ 𝛼+ 𝛾) and 𝑐be sufficiently large
so that 𝑐> (2/𝑑)(𝜇+ 𝛾+ 𝛼−𝑑), we obtain −𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚+ 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚, where 𝐴𝑛𝑚,𝑟,𝜆,ℎ𝑚, 𝐵𝑛𝑚,𝑟,𝜆,ℎ𝑚, and 𝐶𝑛𝑚,𝑟,𝜆,ℎ𝑚are given, respec-
tively, by (30), (31), and (42) and still satisfy (36), (39), and
(42). Consequently, given 𝛿> 0, there is 𝑟𝑚∈N such that, for
all 𝑟⩾𝑟𝑚, Rℓ
𝐶(1) > 1 ⇐⇒
1 + 𝑑(1 + 𝑐/2)
1 + 𝜇+ 𝛼+ 𝛾> 1,
Rℓ
𝐷(1, 1
𝐿) < 1 ⇐⇒
(65) Rℓ
𝐶(1) > 1 ⇐⇒
1 + 𝑑(1 + 𝑐/2)
1 + 𝜇+ 𝛼+ 𝛾> 1,
Rℓ
𝐷(1, 1
𝐿) < 1 ⇐⇒
1 + 𝑑/𝐿
1 + 𝜇+ 𝛼+ 𝛾< 1. (65) (65) 𝜙(ℎ𝑚)
𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚)
> 𝜙(ℎ𝑚) [∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝑓(𝑠) 𝑑𝑠−𝛿+ 𝐶(𝜆+ 1) ℎ𝑚] . 4. Consistency In this context, (𝑥𝑛, 𝑦𝑛) = (𝑎, 𝑏), 𝑛∈N, and (𝑥(𝑡), 𝑦(𝑡)) =
(𝑎, 𝑏), 𝑡∈R, where 4. Consistency (59) (59) So we conclude that we do not have consistency for time step
1/𝐿. Recalling that Rℓ
𝐶(𝜆) > 0, by assumption and since 𝛿> 0 is
arbitrary, we obtain for sufficiently large 𝑚∈N, Let 𝐿= 6 and consider the continuous model with the
following parameters 𝜇= Λ = 0.25, 𝛾= 0.3, 𝛼= 0.05,
𝜂= 0.05, 𝑝= 2/3, 𝑑= 0.6, and 𝑐= 1.5. In Figure 1, we
plot function 𝛽(or similarly 𝜎) and the component 𝐼(𝑡) of
the solution of system (2) given by the solver of Mathematica
(that we take to represent the solution of the continuous-time
model) and the solution of the discrete-time model (9) with
time step 1/6. As can be seen, the infectives are persistent
in the continuous-time model but go to extinction in the
discrete-time model. We have inconsistency in this case. 0 ⩾𝜙(ℎ𝑚)
𝑛𝑚,𝑟𝑚+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟𝑚
𝑓(𝑘ℎ𝑚) > 0,
(60) (60) which is a contradiction. We obtain (b) and the theorem
follows. Next, for each 𝐿∈N, we give an example of a periodic
system of period 1 such that the continuous and the discrete-
time system with time step ℎ= 1/𝐿are not consistent;
namely, we will have persistence for the continuous-time
model and extinction for the discrete-time model with time
step ℎ= 1/𝐿. Note that, changing 𝛽(𝑡) and 𝜎(𝑡) slightly, we can con-
struct an example of a periodic system with period 1 where
the infectives in the continuous-time model go to extinction
but, in the discrete-time model with time step ℎ= 1/𝐿, the
infectives are persistent. Example 10. Let 𝐿∈N. Consider in system (2) that 𝜙(𝑥) =
𝜓(𝑥) = 𝑥, that, with the exception of 𝜎and 𝛽, all parameters
are constant, that Λ = 𝜇, and that Furthermore, we emphasize that this lack of consistency
is not a result of the discretization method used but simply a
result of the fact that the time steps lead to a situation where
the points 𝑛/𝐿where the functions 𝛽and 𝜎are evaluated (in
order to obtain the discrete-time parameters) correspond to
minimums of 𝛽and 𝜎. 𝜎(𝑡) = 𝛽(𝑡) = 𝑑[1 + 𝑐sin2 (2𝜋𝐿𝑡) (1 + cos (2𝜋𝑡))] . (61) We obtain a periodic system of period 1. In this context, (𝑥𝑛, 𝑦𝑛) = (𝑎, 𝑏), 𝑛∈N, and (𝑥(𝑡), 𝑦(𝑡)) =
(𝑎, 𝑏), 𝑡∈R, where We obtain a periodic system of period 1. 4. Consistency We have that g
We have that By (53) we conclude that, given 𝛿> 0, there is 𝑟𝑚∈N such
that, for all 𝑟⩾𝑟𝑚, 𝑛𝑚,𝑟+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟
ℎ𝑚𝑓(𝑘ℎ𝑚)
+ (𝜆−⌊𝜆
ℎ𝑚
⌋ℎ𝑚) 𝑓(𝑛𝑚,𝑟ℎ𝑚+ ⌊𝜆
ℎ𝑚
⌋ℎ𝑚)
(46)
that, for all 𝑟⩾𝑟𝑚,
𝜙(ℎ𝑚)
𝑛𝑚,𝑟𝑚+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟𝑚
ℎ𝑚𝑓(𝑘ℎ𝑚)
< 𝜙(ℎ𝑚) [R𝑢
𝐶(𝜆) + 𝛿+ 𝐶(𝜆+ 1) ℎ𝑚] . (54) (54) < 𝜙(ℎ𝑚) [R𝑢
𝐶(𝜆) + 𝛿+ 𝐶(𝜆+ 1) ℎ𝑚] . is a Riemann sum of Finally, recalling that R𝑢
𝐶(𝜆) < 0, by assumption, by the
arbitrariness of 𝛿> 0 and the fact that ℎ𝑚→0 as 𝑚→+∞,
we obtain for sufficiently large 𝑚∈N, ∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝛽(𝑠) 𝜑(𝑎) + 𝜎(𝑠) 𝜓(𝑏)
−[𝜇(𝑠) + 𝛼(𝑠) + 𝛾(𝑠)] 𝑑𝑠
(47)
Finally, recalling that R𝐶(𝜆) < 0, by assumption, by the
arbitrariness of 𝛿> 0 and the fact that ℎ𝑚→0 as 𝑚→+∞,
we obtain for sufficiently large 𝑚∈N,
0 ⩽𝜙(ℎ𝑚)
𝑛𝑚,𝑟𝑚+⌊𝜆/ℎ𝑚⌋
∑
ℎ𝑚𝑓(𝑘ℎ𝑚) < 0,
(55) ∫
𝑛𝑚,𝑟ℎ𝑚+𝜆
𝑛𝑚,𝑟ℎ𝑚
𝛽(𝑠) 𝜑(𝑎) + 𝜎(𝑠) 𝜓(𝑏)
−[𝜇(𝑠) + 𝛼(𝑠) + 𝛾(𝑠)] 𝑑𝑠 (47) 0 ⩽𝜙(ℎ𝑚)
𝑛𝑚,𝑟𝑚+⌊𝜆/ℎ𝑚⌋
∑
𝑘=𝑛𝑚,𝑟𝑚
ℎ𝑚𝑓(𝑘ℎ𝑚) < 0,
(55) (55) with respect to the partition which is a contradiction. We obtain (a). {𝑛𝑚,𝑟ℎ𝑚, 𝑛𝑚,𝑟ℎ𝑚+ ℎ𝑚, . . . , 𝑛𝑚,𝑟ℎ𝑚
⌊𝜆⌋
}
(48)
which is a contradiction. We obtain (a). A similar argument allows us to prove (b). In fact,
assuming by contradiction that {𝑛𝑚,𝑟ℎ𝑚, 𝑛𝑚,𝑟ℎ𝑚+ ℎ𝑚, . . . , 𝑛𝑚,𝑟ℎ𝑚 (48)
( )
A similar argument allows us to prove (b). In fact,
assuming by contradiction that (48)
( )
A similar argument allows us to prove (b). In fact,
assuming by contradiction that {
,
,
,
+ ⌊𝜆
ℎ𝑚
⌋ℎ𝑚, 𝑛𝑚,𝑟ℎ𝑚+ 𝜆}
(48)
A similar argument allows us to prove (b). In fact,
assuming by contradiction that
Rℓ
𝐶(𝜆) > 0,
(56) (48) + ⌊𝜆
ℎ𝑚
⌋ℎ𝑚, 𝑛𝑚,𝑟ℎ𝑚+ 𝜆} + ⌊𝜆
ℎ𝑚
⌋ℎ𝑚, 𝑛𝑚,𝑟ℎ𝑚+ 𝜆}
(
)
Rℓ
𝐶(𝜆) > 0,
(56) + ⌊𝜆
ℎ𝑚
⌋ℎ𝑚, 𝑛𝑚,𝑟ℎ𝑚+ 𝜆}
Rℓ
𝐶(𝜆) > 0, Rℓ
𝐶(𝜆) > 0,
(56) (56) Discrete Dynamics in Nature and Society 8 and that there is a sequence (ℎ𝑚)𝑚∈N such that ℎ𝑚→0 as
𝑚→+∞and are, respectively, solutions of system (12) and system (3). It is
now possible to compute the number Rℓ
𝐶(1). 5. Simulation Our objective in this section is twofold. On the one hand, we
want to consider different incidence functions 𝜑, correspond-
ing to different discretizations of our continuous model, and (𝑎, 𝑏) = (
𝜇+ 𝜂
𝜇+ 𝜂+ 𝑝,
𝑝
𝜇+ 𝜂+ 𝑝) ,
(62) (62) Discrete Dynamics in Nature and Society
9
0.5
1.0
1.5
2.0
2.5
3.0
0.5
1.0
1.5
2.0
2.5
R–K
Mick
20
40
60
80
100
0.2
0.4
0.6
0.8
Figure 1: Left: function 𝛽; right: inconsistency (time step=1/6). Mick–Zhang
Mickens
10
20
30
40
50
0.2
0.4
0.6
0.8
10
20
30
40
50
0.2
0.4
0.6
0.8
10
20
30
40
50
0.2
0.4
0.6
0.8
R–K
Mick–Zhang
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R–K Figure 2: 𝑆𝐼; 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 4, 1, 0.5. than 0.05. Clearly, there is numerical evidence that there is
consistency even for higher time steps. Figure 2 illustrates this
situation. ℓ compare the several discrete models obtained. We do this in
the first subsection. On the other hand, we want to use our
model to describe a real situation. We do this in the second
subsection where we consider data from the incidence of
measles in France in the period 2012-2016. Changing 𝑏to 0.9 we obtain Rℓ
𝐶(4) = 3.4 > 0 and thus we
conclude that we have persistence. 5. Simulation Taking time steps equal to
2, 1, and 0.5, we get Rℓ
𝐷(1, 2) = 3.201 > 1, Rℓ
𝐷(3, 1) = 5.9 >
1, and Rℓ
𝐷(7, 0.5) = 10.2 > 1 and we conclude that we have
persistence for all these time steps. Figure 3 illustrates this
situation. Figures 2 and 3 suggest that numerically our model
is slightly better than Zhang’s model, at least for large time
steps. 5.1. Simulation with Several NSFD Schemes. In this subsec-
tion we do some simulation to illustrate our results. To
begin, we compare our model (1) with mass action incidence
(𝜑(𝑆, 𝐼) = 𝑆𝐼and 𝜓(𝑉, 𝐼) = 𝑉𝐼) with Zhang’s model [20]. We
use the following set of parameters: 𝜙(ℎ) = ℎ+0.2ℎ2, Λ = 0.5,
𝜇(𝑡) = 𝛾(𝑡) = 𝛿(𝑡) = 0.3, 𝛼(𝑡) = 0.05, 𝜂= 0.05, 𝑝= 2/3 and Next, we compare our model with the discretized model
obtained by Euler method and the output of the Mathematica
solver ODE (that uses a Runge-Kutta method). Considering
𝑏= 0.3, we get extinction for the continuous-time model, as
we already saw. Taking time steps equal to 2, 1, and 0.5, we
can see in Figure 4 that for all methods considered and all
time steps we have extinction, although the behaviour of our
model shadows better the behaviour given by Mathematica’s
solver, at least for these time steps. 𝛽(𝑡) = 𝜎(𝑡) = 𝑏(1 + 0.3 cos (𝑡𝜋
2 )) . (66) (66) Setting 𝑏= 0.3 we obtain R𝑢
𝐶(4) = −0.6 < 0 and thus we
conclude that we have extinction for the continuous model. Taking time steps equal to 4, 1, and 0.5, we get R𝑢
𝐷(0, 4) =
Rℓ
𝐷(0, 4) = 1, R𝑢
𝐷(3, 1) = 0.644 < 1, and R𝐷(7, 0.5) =
0.601 < 1 and we conclude that we have extinction for time
steps 1 and 0.5. For these parameters, we have consistency
in the sense of Theorem 9 as long as the time step is less Changing 𝑏to 0.9 we already saw that we get persistence
for the continuous model. Figure 5 illustrates this situation. 5. Simulation 10
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Figure 3: 𝑆𝐼; 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. 10
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Figure 4: 𝑆𝐼; 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. Euler
Mickens
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Figure 5: 𝑆𝐼; 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. Next, we change our incidence function and consider
𝜙(𝑆𝐼) = 𝑆𝐼/(1 + 0 7𝐼) maintaining the set of parameters
extinction/persistence, relation to Zhang’s model and the
model obtained by Euler method 10
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F
8 M
l
(2012 2016)
i
l ti our parameters estimation, we gathered information from
several websites. We considered standard incidence functions
𝜓(𝑉𝑛+1, 𝐼𝑛) = 𝑉𝑛+1𝐼𝑛/𝑃𝑛and 𝜑(𝑆𝑛+1, 𝐼𝑛) = 𝑆𝑛+1𝐼𝑛/𝑃𝑛, where
𝑃𝑛is the total population. Inspired in the time series for the
infectives (https://ecdc.europa.eu), we considered 𝜎𝑛= 0.03
and 𝛽𝑛given by our parameters estimation, we gathered information from
several websites. We considered standard incidence functions
𝜓(𝑉𝑛+1, 𝐼𝑛) = 𝑉𝑛+1𝐼𝑛/𝑃𝑛and 𝜑(𝑆𝑛+1, 𝐼𝑛) = 𝑆𝑛+1𝐼𝑛/𝑃𝑛, where
𝑃𝑛is the total population. Inspired in the time series for the
infectives (https://ecdc.europa.eu), we considered 𝜎𝑛= 0.03
and 𝛽𝑛given by 𝛽𝑛=
{
{
{
{
{
3.8 + 10 sin ((𝑛+ 1) 𝜋
6
) ,
if ⌊𝑛
12⌋⩽5
2.7,
otherwise. (67) (67) The remaining parameters were considered time indepen-
dent and were inspired in data contained in the web-
sites http://www.worldbank.org, https://data.oecd.org, and
http://www.geoba.se. Namely, we took the mortality rate 𝜇𝑛=
0.0007, the newborns Λ 𝑛
= 50000, the disease induced
mortality 𝛼𝑛= 0.000375, the immunity loss 𝜂𝑛= 0.001, the
vaccination rate 𝑝𝑛= 0.001, and the recovery rate 𝛾𝑛= 0.957. We used the initial conditions 𝑆0 = 7.20428 × 106, 𝐼0 = 106,
𝑉0 = 5.84372 × 107, and 𝑅0 = 1.81918 × 104. In Figure 8, we
plot the real data for the infectives and the output given by
our model. Figure 8: Measles (2012-2016), simulation. 5. Simulation 10
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Figure 5: 𝑆𝐼; 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. 10
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0.8 Figure 5: 𝑆𝐼; 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. extinction/persistence, relation to Zhang’s model and the
model obtained by Euler method. extinction/persistence, relation to Zhang’s model and the
model obtained by Euler method. Next, we change our incidence function and consider
𝜙(𝑆, 𝐼) = 𝑆𝐼/(1 + 0.7𝐼), maintaining the set of parameters. Letting 𝑏= 0.3 we have extinction for the continuous model
and letting 𝑏= 0.9 we have persistence for the continuous-
time model. Note that the thresholds R𝑢
𝐶, Rℓ
𝐶, R𝑢
𝐷, and Rℓ
𝐷
are similar to the mass action case. Figures 6 and 7 illustrate
this situation. 5.2. Simulation with Real Data. In this subsection, we present
some simulation regarding measles. This disease is endemic
in some countries such as France. In that country, with the
measles outbreak in 2011, a vaccination policy that lowered
the number of reported cases was introduced. We will
focus on measles in France, between 2012 and 2016. For
a study concerning the period before 2012, see [36]. For Doing corresponding simulations and comparisons for
our model with 𝜙(ℎ) = (1−e−0.002ℎ)/(0.002) instead of 𝜙(ℎ) =
ℎ+ 0.2ℎ2, we can draw the same conclusions regarding Dynamics in Nature and Society
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Figure 7: 𝑆𝐼/(1 + 0.7𝐼); 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. meters estimation, we gathered information from
ebsites. We considered standard incidence functions
𝑛) = 𝑉𝑛+1𝐼𝑛/𝑃𝑛and 𝜑(𝑆𝑛+1, 𝐼𝑛) = 𝑆𝑛+1𝐼𝑛/𝑃𝑛, where
total population. Inspired in the time series for the
s (https://ecdc.europa.eu), we considered 𝜎𝑛= 0.03
ven by
{
{
{
{
{
3.8 + 10 sin ((𝑛+ 1) 𝜋
6
) ,
if ⌊𝑛
12⌋⩽5
2.7,
otherwise. 5. Simulation (67)
aining parameters were considered time indepen-
d were inspired in data contained in the web-
p://www.worldbank.org, https://data.oecd.org, and
ww.geoba.se. Namely, we took the mortality rate 𝜇𝑛=
he newborns Λ 𝑛
= 50000, the disease induced
Real Data
Simulation
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Figure 8: Measles (2012-2016), simulation. Discrete Dynamics in Nature and Society
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Figure 7: 𝑆𝐼/(1 + 0.7𝐼); 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. 11
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1.0 Discrete Dynamics in Nature and Society 11 11 Discrete Dynamics in Nature and Society
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ons
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03
67)
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𝑛=
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1.0 Figure 6: 𝑆𝐼/(1 + 0.7𝐼); 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. Figure 6: 𝑆𝐼/(1 + 0.7𝐼); 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. 10
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Figure 7: 𝑆𝐼/(1 + 0.7𝐼); 𝜙(ℎ) = ℎ+ 0.2ℎ2; step-size: 2, 1, 0.5. 6. Conclusions We considered a discretization procedure, based on Mick-
ens NSFD scheme, to get a discrete-time model from a
continuous time with vaccination and incidence given by
a general function. For a family of models containing the
previous discrete-time model, we achieved results on the
persistence and the extinction of the disease (Theorems 6 and
7). They contain the results of Zhang [20] as a particular case. Can be seen, in a general way, that our model behaves
in the same manner as the real data. It seems that if the
vaccination policy in France continues to be very strict, it may
decrease the number of cases. Discrete Dynamics in Nature and Society 12 and to carry out some numerical analysis in our general
nonautonomous setting. and to carry out some numerical analysis in our general
nonautonomous setting. Our threshold conditions depend on the parameters of the
model and of the incidence function derivative, with respect
to the infectives, computed on some disease-free solution. This agrees with the continuous counterparts of these results
[8]. Appendix We also considered the problem of establishing the
consistency of the continuous-time model and the discrete-
time model for small time steps, in the sense that if the time
step is small enough when we have persistence (extinction)
for the continuous-time model we also have persistence
(extinction) for the discrete-time model (at least for situations
where Theorems 1 and 2 allow us to conclude that we
have persistence or extinction). Assuming the differentiabil-
ity of parameters, our result on this direction, Theorem 9,
furnishes an interval [0, 𝑎], where 𝑎depends only on the
parameters of the model and their derivatives, where there is
consistency. Proof of the Results in Section 3 We begin this section by noting a simple consequence of our
assumptions that will be used several times throughout the
proofs: it follows from (H3) that there are constants 𝐾> 0
and 𝜃∈]0, 1[ such that 𝑛−1
∏
𝑘=𝑚
1
1 + 𝜇𝑘
< 𝐾𝜃𝑛−𝑚,
(A.1)fi (A.1) for 𝑚, 𝑛∈N sufficiently large. Additionally, using (H1), (H2),
and (H5), we have for 𝑚, 𝑛∈N sufficiently large. Additionally, using (H1), (H2),
and (H5), we have 𝜑(𝑥, 𝑦)
𝑦
= 𝜑(𝑥, 𝑦) −𝜑(𝑥, 0)
𝑦−0
⩽𝜑(𝑥, 𝑦) −𝜑(𝑥, 0)
𝑦−0
⩽𝜕2𝜑(𝑥, 0) = 𝜕2𝜑(𝑥, 0) −𝜕2𝜑(0, 0)
⩽𝑘𝜑𝑥
(A.2) We present an example of a periodic system of period 1
where the continuous and the discrete-time system with time
step ℎ= 1/𝐿are not consistent. Namely, for that time step,
we will have persistence for the continuous-time model and
extinction for the discrete-time model. These examples show
the importance of knowing that for time steps smaller than
some explicit value we have consistency, a type of result like
the one in Theorem 9. (A.2) and thus and thus 𝜑(𝑥, 𝑦) ⩽𝑘𝜑𝑥𝑦. (A.3) Finally, we carried out some simulations to illustrate our
results. As one might expect our simulations furnish evidence
that we may have consistency in intervals whose lengths are
several times bigger than the length of the given interval in
Theorem 9. Additionally, we used our model to describe a real
situation, namely, the case of measles incidence in France in
the period 2012-2016, and compared our results with the real-
time series for the infectives. We found in general that the
predictive behaviour of our model is very similar to the real
data. (A.3) Similarly Similarly 𝜓(𝑥, 𝑦) ⩽𝑘𝜓𝑥𝑦. (A.4) (A.4) We will now proceed with the proof of the results in
Section 3. Proof of Lemma 4. Let 𝑆𝑛> 0, 𝐼𝑛> 0, 𝑅𝑛> 0, and 𝑉𝑛> 0. By
(1), (A.3), and (A.4), we obtain We remark that this work is far from giving a complete
answer to the problem of consistency between our discrete
and continuous-time models. Proof of the Results in Section 3 To show that Rℓ
𝐷(𝜉∗, 𝜆) is independent of
the selection of 𝜉∗= (𝑥∗
𝑛, 𝑦∗
𝑛), a fixed solution of (12), it is
important to note that according to (ii) in Lemma 3, for any
𝜀> 0 and any solution 𝜉= (𝑥𝑛, 𝑦𝑛) of system (12) with initial
value 𝑥0 > 0, 𝑦0 > 0, there exists an 𝑁∈N+ such that, for
𝑘⩾𝑁, we have |𝑥𝑘−𝑥∗
𝑘| ⩽𝜀and |𝑦𝑘−𝑦∗
𝑘| ⩽𝜀. Hence, where where Θ𝜀= (
𝜆+ 1
𝜆) 𝑟𝜆𝐶𝜀+ . . . + (
𝜆+ 1
1 ) 𝑟𝐶𝜆𝜀𝜆
+ 𝐶𝜆+1𝜀𝜆+1. (A.17) Θ𝜀= (
𝜆+ 1
𝜆) 𝑟𝜆𝐶𝜀+ . . . + (
𝜆+ 1
1 ) 𝑟𝐶𝜆𝜀𝜆
(A.17) (A.17) + 𝐶𝜆+1𝜀𝜆+1. 𝑥∗
𝑘−𝜀⩽𝑥𝑘⩽𝑥∗
𝑘+ 𝜀
𝑦∗
𝑘−𝜀⩽𝑦𝑘⩽𝑦∗
𝑘+ 𝜀. (A.9) Analogously Analogously (A.9) 𝑛+𝜆
∏
𝑘=𝑛
(𝑟𝑘−
𝐿𝑘𝜀
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
) ⩾
𝑛+𝜆
∏
𝑘=𝑛
𝑟𝑘−Θ𝜀. (A.18) (A.18) By (H1), we have 𝜕2𝜑(𝑥𝑘, 0) −𝜕2𝜑(𝑥∗
𝑘, 0)⩽𝑘𝜑𝑥𝑘−𝑥∗
𝑘
⩽𝑘𝜑𝜀,
𝜕2𝜓(𝑦𝑘, 0) −𝜕2𝜓(𝑦∗
𝑘, 0)⩽𝑘𝜓𝑦𝑘−𝑦∗
𝑘
⩽𝑘𝜓𝜀. (A.10)
So,
By (A.12), we obtain
−Θ𝜀+ lim inf
𝑛→+∞
𝑛+𝜆
∏
𝑘=𝑛
𝑟𝑘⩽Rℓ
𝐷(𝜉∗, 𝜆)
(A 19) 𝜕2𝜑(𝑥𝑘, 0) −𝜕2𝜑(𝑥∗
𝑘, 0)⩽𝑘𝜑𝑥𝑘−𝑥∗
𝑘
⩽𝑘𝜑𝜀,
𝜕2𝜓(𝑦𝑘, 0) −𝜕2𝜓(𝑦∗
𝑘, 0)⩽𝑘𝜓𝑦𝑘−𝑦∗
𝑘
⩽𝑘𝜓𝜀. (A.10 By (A.12), we obtain By (A.12), we obtain (A.10) −Θ𝜀+ lim inf
𝑛→+∞
𝑛+𝜆
∏
𝑘=𝑛
𝑟𝑘⩽Rℓ
𝐷(𝜉∗, 𝜆) So, So, (A.19) 𝜕2𝜑(𝑥∗
𝑘, 0) −𝑘𝜑𝜀⩽𝜕2𝜑(𝑥𝑘, 0) ⩽𝜕2𝜑(𝑥∗
𝑘, 0) + 𝑘𝜑𝜀,
𝜕2𝜓(𝑦∗
𝑘, 0) −𝑘𝜓𝜀⩽𝜕2𝜓(𝑦𝑘, 0)
⩽𝜕2𝜓(𝑦∗
𝑘, 0) + 𝑘𝜓𝜀. (A.11)
⩽Θ𝜀+ lim inf
𝑛→+∞
𝑛+𝜆
∏
𝑘=𝑛
𝑟𝑘. (A.19)
Thus, 𝜕2𝜑(𝑥∗
𝑘, 0) −𝑘𝜑𝜀⩽𝜕2𝜑(𝑥𝑘, 0) ⩽𝜕2𝜑(𝑥∗
𝑘, 0) + 𝑘𝜑𝜀,
𝜕2𝜓(𝑦∗
𝑘, 0) −𝑘𝜓𝜀⩽𝜕2𝜓(𝑦𝑘, 0)
⩽𝜕2𝜓(𝑦∗
𝑘, 0) + 𝑘𝜓𝜀. (A.1 (A.11) Thus, Rℓ
𝐷(𝜉∗, 𝜆) −Rℓ
𝐷(𝜉, 𝜆)< Θ𝜀
(A.20) (A.20) Combining the previous computations, we get Combining the previous computations, we get 1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝐿𝑘𝜀
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘 and, by the arbitrariness of 𝜀, we obtain Rℓ
𝐷(𝜉, 𝜆) = Rℓ
𝐷(𝜉∗,
𝜆). Replacing lim inf by lim sup in the preceding argument,
we reach a similar conclusion for R𝑢
𝐷(𝜉∗, 𝜆). The result
follows. and, by the arbitrariness of 𝜀, we obtain Rℓ
𝐷(𝜉, 𝜆) = Rℓ
𝐷(𝜉∗,
𝜆). Replacing lim inf by lim sup in the preceding argument,
we reach a similar conclusion for R𝑢
𝐷(𝜉∗, 𝜆). The result
follows. ⩽1 + 𝛽𝑘𝜕2𝜑(𝑥𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦𝑘+1, 0)
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
(A.12) (A.12) Proof of Theorem 6. Proof of the Results in Section 3 Moreover, in addition to a
further discussion of the qualitative behaviour of the con-
tinuous and discrete systems, in future work it would be
very interesting to discuss the convergence of the methods 𝑉𝑛+1 =
𝑉𝑛+ 𝑝𝑛𝑆𝑛+1
1 + 𝜇𝑛+ 𝜂𝑛+ 𝜎𝑛𝑘𝜓𝐼𝑛
(A.5) (A.5) and thus and thus 𝑆𝑛+1 ⩾
𝜂𝑛(𝑉𝑛+ Λ 𝑛+ 𝑆𝑛) + (Λ 𝑛+ 𝑆𝑛) (1 + 𝜇𝑛+ 𝜎𝑛𝑘𝜓𝐼𝑛)
(1 + 𝜇𝑛+ 𝛽𝑛𝑘𝜑𝐼𝑛) (1 + 𝜇𝑛+ 𝜂𝑛+ 𝜎𝑛𝑘𝜑𝐼𝑛) + 𝑝𝑛(1 + 𝜇𝑛+ 𝜎𝑛𝑘𝜓𝐼𝑛)
. (A.6) (A.6) 𝐼𝑛> 0, 𝑅𝑛> 0, and 𝑉𝑛> 0 for all 𝑛∈N. In the same way, we
can conclude that, if 𝑆0 ⩾0, 𝐼0 ⩾0, 𝑅0 ⩾0 and 𝑉0 ⩾0, then
𝑆𝑛⩾0. 𝐼𝑛⩾0, 𝑅𝑛⩾0 and 𝑉𝑛⩾0 for all 𝑛∈N. This proves (i)
and (ii) in Lemma 4. By (1) we have Therefore, we conclude that 𝑆𝑛+1 > 0 and 𝑉𝑛+1 > 0. By the
second and third equations in (1) we obtain 𝐼𝑛+1 ⩾
𝐼𝑛
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
,
𝑅𝑛+1 = 𝛾𝑛𝐼𝑛+1 + 𝑅𝑛
1 + 𝜇𝑛
(A.7) ( )
By (1), we have (A.7) 𝑁𝑛+1 ⩽
Λ 𝑛
1 + 𝜇𝑛
+
𝑁𝑛
1 + 𝜇𝑛
,
(A.8) (A.8) and we conclude that 𝐼𝑛+1 > 0 and 𝑅𝑛+1 > 0. The previous
inequalities allow us to conclude by induction that 𝑆𝑛> 0, where 𝑁𝑛= 𝑆𝑛+𝐼𝑛+𝑅𝑛+𝑉𝑛is the total population. By Lemma
2 in [21] we obtain the result. where 𝑁𝑛= 𝑆𝑛+𝐼𝑛+𝑅𝑛+𝑉𝑛is the total population. By Lemma
2 in [21] we obtain the result. Discrete Dynamics in Nature and Society 13 Proof of Lemma 5. To show that Rℓ
𝐷(𝜉∗, 𝜆) is independent of
the selection of 𝜉∗= (𝑥∗
𝑛, 𝑦∗
𝑛), a fixed solution of (12), it is
important to note that according to (ii) in Lemma 3, for any
𝜀> 0 and any solution 𝜉= (𝑥𝑛, 𝑦𝑛) of system (12) with initial
value 𝑥0 > 0, 𝑦0 > 0, there exists an 𝑁∈N+ such that, for
𝑘⩾𝑁, we have |𝑥𝑘−𝑥∗
𝑘| ⩽𝜀and |𝑦𝑘−𝑦∗
𝑘| ⩽𝜀. Hence, Proof of Lemma 5. Proof of the Results in Section 3 First note that the original system (1) can
be rewritten as follows: Proof of Theorem 6. First note that the original system (1) can
be rewritten as follows: ⩽1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0) + 𝐿𝑘𝜀
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
, 𝑆𝑛+1 =
1
1 + 𝜇𝑛+ 𝑝𝑛
(Λ 𝑛+ 𝑆𝑛−𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛)
+ 𝜂𝑛𝑉𝑛+1)
𝐼𝑛+1 =
1
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛)
+ 𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) + 𝐼𝑛)
𝑅𝑛+1 =
1
1 + 𝜇𝑛
(𝛾𝑛𝐼𝑛+1 + 𝑅𝑛)
𝑉𝑛+1 =
1
1 + 𝜇𝑛+ 𝜂𝑛
(𝑝𝑛𝑆𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) + 𝑉𝑛) ,
(A.21) where 𝐿𝑘= 𝛽𝑘𝑘𝜑+ 𝜎𝑘𝑘𝜓. Let where 𝐿𝑘= 𝛽𝑘𝑘𝜑+ 𝜎𝑘𝑘𝜓. Let 𝑘
Let 𝑟𝑘= 1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0)
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
. (A.13) 𝐼𝑛+1 =
1
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛)
+ 𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) + 𝐼𝑛)
𝑅𝑛+1 =
1
1 + 𝜇𝑛
(𝛾𝑛𝐼𝑛+1 + 𝑅𝑛)
𝑉𝑛+1 =
1
1 + 𝜇𝑛+ 𝜂𝑛
(𝑝𝑛𝑆𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) + 𝑉𝑛) ,
(A.21) (A.21) Using (H1) and (iv) in Lemma 3, it is easy to see that Using (H1) and (iv) in Lemma 3, it is easy to see that 𝑘⩽
1 + 2𝛽𝑢𝑘𝜑𝑀+ 2𝜎𝑢𝑘𝜓𝑀
1 + 𝜇𝑙+ 𝛼𝑙+ 𝛾𝑙
š 𝑟,
(A.14)
large 𝑘∈N, and that
𝑅𝑛+1 =
1
1 + 𝜇𝑛
(𝛾𝑛𝐼𝑛+1 + 𝑅𝑛)
𝑉
=
1
(𝑝𝑆
−𝜎 𝑟𝑘⩽
1 + 2𝛽𝑢𝑘𝜑𝑀+ 2𝜎𝑢𝑘𝜓𝑀
1 + 𝜇𝑙+ 𝛼𝑙+ 𝛾𝑙
š 𝑟,
(A.14) 𝑟𝑘⩽
1 + 2𝛽𝑢𝑘𝜑𝑀+ 2𝜎𝑢𝑘𝜓𝑀
1 + 𝜇𝑙+ 𝛼𝑙+ 𝛾𝑙
š 𝑟,
(A.14)
ntly large 𝑘∈N, and that
𝑅𝑛+1 =
1
1 + 𝜇𝑛
(𝛾𝑛𝐼𝑛+1 + 𝑅𝑛)
𝑉𝑛+1 =
1
1 + 𝜇𝑛+ 𝜂𝑛
(𝑝𝑛𝑆𝑛+1 −𝜎𝑛𝜓(𝑉𝑛+1, 𝐼 for sufficiently large 𝑘∈N, and that for sufficiently large 𝑘∈N, and that 𝐿=
𝐿𝑘
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
⩽
𝛽𝑢𝑘𝜑+ 𝜎𝑢𝑘𝜓
1 + 𝜇𝑙+ 𝛼𝑙+ 𝛾𝑙š 𝐶. (A.15)
o, for sufficiently large 𝑛,
𝑛= 0, 1, . . .. Firstly, we will establish (a). Since R𝑢
𝐷(𝜆) < 1, we can
choose 𝜀0 > 0, 𝜀∈]0, 1[ and a sufficiently large integer 𝑁1 ∈N
such that 𝐿=
𝐿𝑘
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
⩽
𝛽𝑢𝑘𝜑+ 𝜎𝑢𝑘𝜓
1 + 𝜇𝑙+ 𝛼𝑙+ 𝛾𝑙š 𝐶. (A.15)
o, for sufficiently large 𝑛,
𝑛= 0, 1, . . .. Firstly, we will establish (a). Proof of the Results in Section 3 Since R𝑢
𝐷(𝜆) < 1, we can
choose 𝜀0 > 0, 𝜀∈]0, 1[ and a sufficiently large integer 𝑁1 ∈N
such that So, for sufficiently large 𝑛, So, for sufficiently large 𝑛, 𝑛+𝜆
∏
𝑘=𝑛
(𝑟𝑘+
𝐿𝑘𝜀
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
) ⩽
𝑛+𝜆
∏
𝑘=𝑛
(𝑟𝑘+ 𝐶𝜀) 6)
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽𝑘𝜕2𝜑(𝑥𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦𝑘+1, 0) + (𝛽𝑢𝑘𝜑+ 𝜎𝑢𝑘𝜓) 𝜀0
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
< 𝜀
(A.22)
for all 𝑛⩾𝑁1. (A.22) (A.16) (A.16) =
𝑛+𝜆
∏
𝑘=𝑛
𝑟𝑘+ Θ𝜀 for all 𝑛⩾𝑁1. 14 14 Discrete Dynamics in Nature and Society For any solution (𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (A.21) with initial con-
ditions 𝑆0 > 0, 𝐼0 > 0, 𝑅0 > 0 and 𝑉0 > 0, we have 𝜕2𝜓(𝑉𝑛+1, 0) ⩽𝜕2𝜓(𝑦∗
𝑛+1, 0) + 𝑘𝜓𝜀0. (A.30)h
erefore by (A 22) (A 26) (A 27) (A 29) and (A 30) we 𝜕2𝜓(𝑉𝑛+1, 0) ⩽𝜕2𝜓(𝑦∗
𝑛+1, 0) + 𝑘𝜓𝜀0. (A.30) (A.30) 𝑆𝑛+1 ⩽Λ 𝑛+ 𝜂𝑛𝑉𝑛+1 + 𝑆𝑛
1 + 𝜇𝑛+ 𝑝𝑛
𝑉𝑛+1 ⩽𝑝𝑛𝑆𝑛+1 + 𝑉𝑛
1 + 𝜇𝑛+ 𝜂𝑛
. (A.23) (A.23) (A.23)h
have 𝐼𝑛+1 By the comparison principle, we obtain 𝑆𝑛⩽𝑥𝑛and 𝑉𝑛⩽
𝑦𝑛for all 𝑛∈N, where (𝑥𝑛, 𝑦𝑛) is the solution of (12) with
initial condition (𝑥0, 𝑦0) = (𝑆0, 𝑉0). According to Lemma 3,
the solution (𝑥∗
𝑛, 𝑦∗
𝑛) is globally uniformly attractive and thus,
for the aforementioned 𝜀0 > 0, there exists an 𝑁2 ∈N such
that ⩽𝜀𝐼𝑛, for all 𝑛⩾𝑁2. We conclude that 𝐼𝑚⩽𝜀𝑚−𝑁2𝐼𝑁2 →0 as 𝑚→
∞. This completes the proof of (a). for all 𝑛⩾𝑁2. We conclude that 𝐼𝑚⩽𝜀𝑚−𝑁2𝐼𝑁2 →0 as 𝑚→
∞. This completes the proof of (a). h
p
p
Next, to establish (b), let us consider two arbitrary solu-
tions of the original system (𝑆(1)
𝑛, 𝐼(1)
𝑛, 𝑉(1)
𝑛, 𝑅(1)
𝑛) and (𝑆(2)
𝑛, 𝐼(2)
𝑛,
𝑉(2)
𝑛, 𝑅(2)
𝑛) and 𝜆a constant such that 𝑅𝑢
0(𝜆) < 1. Let 𝜄𝑛= 𝐼(1)
𝑛−
𝐼(2)
𝑛
and 𝜌𝑛= 𝑅(1)
𝑛−𝑅(2)
𝑛. By (9), we have 𝑥𝑛−𝑥∗
𝑛
⩽𝜀0,
𝑦𝑛−𝑦∗
𝑛
⩽𝜀0 𝑥𝑛−𝑥∗
𝑛
⩽𝜀0,
𝑦𝑛−𝑦∗
𝑛
⩽𝜀0
(A.24) (A.24) ∀𝑛⩾𝑁2 𝜌𝑛+1 −𝜌𝑛= (𝑅(1)
𝑛+1 −𝑅(1)
𝑛) −(𝑅(2)
𝑛+1 −𝑅(2)
𝑛)
= 𝛾𝑛(𝐼(1)
𝑛+1 −𝐼(2)
𝑛+1) −𝜇𝑛(𝑅(1)
𝑛+1 −𝑅(2)
𝑛+1)
= 𝛾𝑛𝜄𝑛+1 −𝜇𝑛𝜌𝑛+1. (A.32) 𝜌𝑛+1 −𝜌𝑛= (𝑅(1)
𝑛+1 −𝑅(1)
𝑛) −(𝑅(2)
𝑛+1 −𝑅(2)
𝑛) From this, it may be concluded that From this, it may be concluded that 𝑆𝑛⩽𝑥∗
𝑛+ 𝜀0,
𝑉𝑛⩽𝑦∗
𝑛+ 𝜀0
∀𝑛⩾𝑁2. Proof of the Results in Section 3 (A.25) 𝑆𝑛⩽𝑥∗
𝑛+ 𝜀0,
𝑉𝑛⩽𝑦∗
𝑛+ 𝜀0 𝑉𝑛⩽𝑦∗
𝑛+ 𝜀0 (A.25) = 𝛾𝑛𝜄𝑛+1 −𝜇𝑛𝜌𝑛+1. ∀𝑛⩾𝑁2. Because 𝑅𝑢
0(𝜆) < 1, we conclude that 𝜄𝑛→0 as 𝑛→+∞and
therefore, given 𝜀> 0, there is 𝑁∈N sufficiently large such
that, for 𝑛⩾𝑁, Because 𝑅𝑢
0(𝜆) < 1, we conclude that 𝜄𝑛→0 as 𝑛→+∞and
therefore, given 𝜀> 0, there is 𝑁∈N sufficiently large such
that, for 𝑛⩾𝑁, By the second equation of (1) we get By the second equation of (1) we get 𝐼𝑛+1 =
1
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛)
+ 𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) + 𝐼𝑛)
=
1
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(𝛽𝑛
𝜑(𝑆𝑛+1, 𝐼𝑛)
𝐼𝑛
+ 𝜎𝑛
𝜓(𝑉𝑛+1, 𝐼𝑛)
𝐼𝑛
+ 1) 𝐼𝑛. (A.26)
𝜌𝑛+1 + 𝜇𝑛𝜌𝑛+1 = 𝛾𝑛𝜄𝑛+1 + 𝜌𝑛< 𝜀+ 𝜌𝑛
(A.33)
Thus, since 𝜇𝑛> 0, we get
𝜌𝑛+1 <
𝜀
1 + 𝜇𝑛
+
𝜌𝑛
1 + 𝜇𝑛
. (A.34)
and proceeding by induction
(
𝑛−1
∏
1
)
𝑛−1
∑(
𝑛−1
∏
1
)
(A
) 𝜌𝑛+1 + 𝜇𝑛𝜌𝑛+1 = 𝛾𝑛𝜄𝑛+1 + 𝜌𝑛< 𝜀+ 𝜌𝑛
(A.33)
Thus, since 𝜇𝑛> 0, we get 𝜌𝑛+1 + 𝜇𝑛𝜌𝑛+1 = 𝛾𝑛𝜄𝑛+1 + 𝜌𝑛< 𝜀+ 𝜌𝑛
(A.33) (A.33) Thus, since 𝜇𝑛> 0, we get 𝜌𝑛+1 <
𝜀
1 + 𝜇𝑛
+
𝜌𝑛
1 + 𝜇𝑛
. (A.34) and proceeding by induction and proceeding by induction + 𝜎𝑛
𝜓(𝑉𝑛+1, 𝐼𝑛)
𝐼𝑛
+ 1) 𝐼𝑛. 𝜌𝑛+1 < (
𝑛−1
∏
𝑚=0
1
1 + 𝜇𝑚
) 𝜌0 + 𝜀
𝑛−1
∑
𝑚=0
(
𝑛−1
∏
𝑘=𝑚
1
1 + 𝜇𝑘
) . (A.35) (A.35) By (H5), we have 𝜑(𝑆𝑛+1, 𝐼𝑛)
𝐼𝑛
⩽𝜕2𝜑(𝑆𝑛+1, 0) ,
𝜓(𝑉𝑛+1, 𝐼𝑛)
𝐼𝑛
⩽𝜕2𝜓(𝑉𝑛+1, 0) . (A.27) By (H3) and (A.1), we conclude that By (H3) and (A.1), we conclude that lim sup
𝑛→+∞𝜌𝑛= 0. (A.36) (A.27)
lim sup
𝑛→+∞𝜌𝑛= 0. (A.36) (A.27) (A.36) Thus, 𝜌𝑛= 𝑅(1)
𝑛−𝑅(2)
𝑛
→0 as 𝑛→+∞. Similar computations
show that 𝑆(1)
𝑛−𝑆(2)
𝑛
→0 and 𝑉(1)
𝑛
−𝑉(2)
𝑛
→0. This proves (b)
and the result follows. By (H1), 𝑥→𝜕2𝜑(𝑥, 0) and 𝑥→𝜕2𝜓(𝑥, 0) are non-
decreasing and also Lipschitz, so, using (A.25) we obtain Proof of Theorem 7. Proof of the Results in Section 3 (A.40)
𝑆𝑛⩾𝑥𝑛⩾𝑥𝑛 (A.48) By (1), (A.39), and (A.40), we have By (1), (A.39), and (A.40), we have By (1), (A.39), and (A.40), we have for all 𝑛⩾𝑁2. Thus, since 𝑆𝑛⩽𝑥∗
𝑛and 𝑉𝑛⩽𝑦∗
𝑛for all 𝑛∈N,
we have 𝑆𝑛+1 ⩾
Λ 𝑛+ 𝜂𝑛𝑉𝑛+1 + 𝑆𝑛−𝛽𝑢𝑘𝜑𝑀𝜀1
1 + 𝜇𝑛+ 𝑝𝑛 𝑆𝑛+1 ⩾
Λ 𝑛+ 𝜂𝑛𝑉𝑛+1 + 𝑆𝑛−𝛽𝑢𝑘𝜑𝑀𝜀1
1 + 𝜇𝑛+ 𝑝𝑛
𝑉𝑛+1 ⩾
𝑝𝑛𝑆𝑛+1 + 𝑉𝑛−𝜎𝑢𝑘𝜓𝑀𝜀1
1 + 𝜇𝑛+ 𝜂𝑛
. 𝑆𝑛+1 ⩾
Λ 𝑛+ 𝜂𝑛𝑉𝑛+1 + 𝑆𝑛−𝛽𝑢𝑘𝜑𝑀𝜀1
1 + 𝜇𝑛+ 𝑝𝑛
𝑉𝑛+1 ⩾
𝑝𝑛𝑆𝑛+1 + 𝑉𝑛−𝜎𝑢𝑘𝜓𝑀𝜀1
1 + 𝜇𝑛+ 𝜂𝑛
. (A.41)
2h
𝑛
𝑛
𝑛
𝑦𝑛
we have
𝑆𝑛−𝑥∗
𝑛
⩽𝜀0,
𝑉𝑛−𝑦∗
𝑛
⩽𝜀0,
(A.49) (A.41)
𝑆𝑛−𝑥∗
𝑛
⩽𝜀0,
𝑉𝑛−𝑦∗
𝑛
⩽𝜀0,
(A.49) 1 + 𝜇𝑛+ 𝑝𝑛
𝑉𝑛+1 ⩾
𝑝𝑛𝑆𝑛+1 + 𝑉𝑛−𝜎𝑢𝑘𝜓𝑀𝜀1
1 + 𝜇𝑛+ 𝜂𝑛
. (A.41)
𝑆𝑛−𝑥∗
𝑛
⩽𝜀0,
𝑉𝑛−𝑦∗
𝑛
⩽𝜀0,
(A.49) (A.41) (A.49) l 𝑛⩾𝑁1. Given 𝜀1 > 0, consider the auxiliary system
𝑥𝑛+1 =
Λ 𝑛+ 𝜂𝑛𝑦𝑛+1 + 𝑥𝑛−𝛽𝑢𝑘𝜑𝑀𝜀1
1 + 𝜇+ 𝑝
for all 𝑛⩾𝑁2. By (H1), we have, for all 𝑛⩾𝑁2,
𝜕2𝜑(𝑆𝑘+1, 0) −𝜕2𝜑(𝑥∗
𝑘+1, 0)⩽𝑘𝜑𝑆𝑘+1 −𝑥∗
𝑘+1
⩽𝑘𝜑𝜀0
(A.50) ll 𝑛⩾𝑁1. Given 𝜀1 > 0, consider the auxiliary system
𝑥𝑛+1 =
Λ 𝑛+ 𝜂𝑛𝑦𝑛+1 + 𝑥𝑛−𝛽𝑢𝑘𝜑𝑀𝜀1
1 + 𝜇𝑛+ 𝑝𝑛
𝑦𝑛+1 =
𝑝𝑛𝑥𝑛+1 + 𝑦𝑛−𝜎𝑢𝑘𝜓𝑀𝜀1
1 + 𝜇𝑛+ 𝜂𝑛
. (A.42)
for all 𝑛⩾𝑁2. By (H1), we have, for all 𝑛⩾𝑁2,
𝜕2𝜑(𝑆𝑘+1, 0) −𝜕2𝜑(𝑥∗
𝑘+1, 0)⩽𝑘𝜑𝑆𝑘+1 −𝑥∗
𝑘+1
⩽𝑘𝜑𝜀0
(A.50)
and thus
𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 ⩽𝜕2𝜑(𝑆𝑘+1, 0)
𝜕
( ∗
)
𝑘
(A.51) for all 𝑛⩾𝑁1. 1
Given 𝜀1 > 0, consider the auxiliary system 1
Given 𝜀1 > 0, consider the auxiliary system (A.50) 𝑥𝑛+1 =
Λ 𝑛+ 𝜂𝑛𝑦𝑛+1 + 𝑥𝑛−𝛽𝑢𝑘𝜑𝑀𝜀1
1 + 𝜇𝑛+ 𝑝𝑛
𝑦𝑛+1 =
𝑝𝑛𝑥𝑛+1 + 𝑦𝑛−𝜎𝑢𝑘𝜓𝑀𝜀1
1 + 𝜇𝑛+ 𝜂𝑛
. (A.42)
and thus (A.42)
and thus (A.42) 𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 ⩽𝜕2𝜑(𝑆𝑘+1, 0)
⩽𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝑘𝜑𝜀0. (A.51) 𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 ⩽𝜕2𝜑(𝑆𝑘+1, 0)
⩽𝜕𝜑(𝑥∗
0) + 𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 ⩽𝜕2𝜑(𝑆𝑘+1, 0) −𝑘𝜑𝜀0 ⩽𝜕2𝜑(𝑆𝑘+1, 0)
⩽𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝑘𝜑𝜀0. (A.51) (A.51) (A.51) ⩽𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝑘𝜑𝜀0. (A.51 For any 𝑛0 ∈N and 𝑥0, 𝑦0 ∈R+, let (𝑥𝑛, 𝑦𝑛) be the solution
of (12) with initial condition (𝑥𝑛0, 𝑦𝑛0) = (𝑥0, 𝑦0) and let
(𝑥𝑛0, 𝑦𝑛0) = (𝑥0, 𝑦0) be the solution of (A.42) with the same
initial condition. Proof of the Results in Section 3 Since Rℓ
0(𝜆) > 1, there are 𝜀, 𝜀0 > 0 such
that 𝜕2𝜑(𝑆𝑛+1, 0) −𝜕2𝜑(𝑥∗
𝑛+1, 0)
⩽𝜕2𝜑(𝑥∗
𝑛+1 + 𝜀0, 0) −𝜕2𝜑(𝑥∗
𝑛+1, 0)
= 𝜕2𝜑(𝑥∗
𝑛+1 + 𝜀0, 0) −𝜕2𝜑(𝑥∗
𝑛+1, 0)⩽𝑘𝜑𝜀0
(A.28) 𝜕2𝜑(𝑆𝑛+1, 0) −𝜕2𝜑(𝑥∗
𝑛+1, 0) (A.28) lim inf
𝑛→+∞
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑢
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
> 1 + 𝜀,
(A.37) lim inf
𝑛→+∞
𝑛+𝜆
∏
𝑘=𝑛
1 + 𝛽𝑘𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑘𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑢
1 + 𝜇𝑘+ 𝛼𝑘+ 𝛾𝑘
(A.37) (A.37) and thus and thus > 1 + 𝜀, 𝜕2𝜑(𝑆𝑛+1, 0) ⩽𝜕2𝜑(𝑥∗
𝑛+1, 0) + 𝑘𝜑𝜀0. (A.29)
for all 𝑢∈[0, 𝜀0]. Discrete Dynamics in Nature and Society 15 Noting that (A.41) can be written as We claim that there is 𝜀1 > 0 such that Noting that (A.41) can be written as 𝑆𝑛+1
⩾
𝜂𝑛𝑝𝑛(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛) (Λ 𝑛+ 𝑆𝑛−𝛽𝑢𝑀𝑘𝜑𝜀1)
(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛)
+
(𝑉𝑛−𝜎𝑢𝑀𝑘𝜓𝜀1) (1 + 𝜇𝑛+ 𝑝𝑛)
(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛)
(A.47) lim sup
𝑛→∞
𝐼𝑛> 𝜀1
(A.38) 𝑆𝑛+1 (A.38) ⩾
𝜂𝑛𝑝𝑛(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛) (Λ 𝑛+ 𝑆𝑛−𝛽𝑢𝑀𝑘𝜑𝜀1)
(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛) for every solution with positive initial conditions of system
(1). +
(𝑉𝑛−𝜎𝑢𝑀𝑘𝜓𝜀1) (1 + 𝜇𝑛+ 𝑝𝑛)
(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛)
(A.47) ( )
We proceed by contradiction. Assume that (A.38) does
not hold. Then, for each 𝜀1 there is 𝑁1 ∈N and a solution
(𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) with positive initial conditions such that 𝐼𝑛⩽
𝜀1 for all 𝑛⩾𝑁1. By (iii) in Lemma 4, we can assume that 𝑆𝑛,
𝑉𝑛< 𝑀for all 𝑛⩾𝑁1. By (A.3) we have (A.47) 𝑉𝑛+1 ⩾
𝑝𝑛(Λ 𝑛+ 𝑆𝑛−𝛽𝑢𝑀𝑘𝜑𝜀1) + (𝑉𝑛−𝜎𝑢𝑀𝑘𝜓𝜀1) (1 + 𝜇𝑛+ 𝑝𝑛)
(1 + 𝜇𝑛) (1 + 𝜇𝑛+ 𝑝𝑛+ 𝜂𝑛) 𝜑(𝑆𝑛+1, 𝐼𝑛) = 𝑘𝜑𝑆𝑛+1𝐼𝑛⩽𝑘𝜑𝑀𝜀1
(A.39) (A.39) and, using (A.42) and (A.46), we conclude that and likewise, by (A.4), we get 𝜓𝑀𝜀1. (A.40)
𝑆𝑛⩾𝑥𝑛⩾𝑥∗
𝑛−𝜀0,
𝑉𝑛⩾𝑦𝑛⩾𝑦∗
𝑛−𝜀0,
(A.48) 𝜓(𝑉𝑛+1, 𝐼𝑛) = 𝑘𝜓𝑉𝑛+1𝐼𝑛⩽𝑘𝜓𝑀𝜀1. (A.40)
𝑆𝑛⩾𝑥𝑛⩾𝑥∗
𝑛−𝜀0,
𝑉⩾
⩾
∗ 𝜓(𝑉𝑛+1, 𝐼𝑛) = 𝑘𝜓𝑉𝑛+1𝐼𝑛⩽𝑘𝜓𝑀𝜀1. (A.40)
(A.39), and (A.40), we have
𝑆𝑛⩾𝑥𝑛⩾𝑥∗
𝑛−𝜀0,
𝑉𝑛⩾𝑦𝑛⩾𝑦∗
𝑛−𝜀0,
(A.48) 𝜓(𝑉𝑛+1, 𝐼𝑛) = 𝑘𝜓𝑉𝑛+1𝐼𝑛⩽𝑘𝜓𝑀𝜀1. Proof of the Results in Section 3 Letting 𝜀2 > 0 in (A.69) be sufficiently
small and (𝑥𝑛, 𝑦𝑛) be the solution of (12) with 𝑥𝑠𝑛,𝑘+1 = 𝑆𝑠𝑛,𝑘+1
and 𝑦𝑠𝑛,𝑘= 𝑉𝑠𝑛,𝑘, we have by (iii) in Lemma 3 Recall that, since Rℓ
0(𝜆) > 1, there are 𝜀, 𝜀0 > 0 such that
(A.37) holds for all 𝑢∈[0, 𝜀0]. Recall that, since Rℓ
0(𝜆) > 1, there are 𝜀, 𝜀0 > 0 such that
(A.37) holds for all 𝑢∈[0, 𝜀0]. If (A.59) does not hold, then, given 𝜀0 > 0, there must be a
sequence of solutions of (1), ((𝑆𝑛,𝑘, 𝐼𝑛,𝑘, 𝑅𝑛,𝑘, 𝑉𝑛,𝑘)𝑛∈N)𝑘∈N, with
initial conditions (𝑆0,𝑘, 𝐼0,𝑘, 𝑅0,𝑘, 𝑉0,𝑘) such that 𝑥𝑛−𝑥𝑛+ 𝑦𝑛−𝑦𝑛
⩽𝜀0
2
(A.70) (A.70) lim inf
𝑛→+∞𝐼𝑛,𝑘< 𝜀0
𝑘2 . (A.60) (A.60) (A.60) for all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N. We conclude that for all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N. We conclude that From (A.38), for each 𝑘∈N, there must be two sequences
(𝑠𝑚,𝑘)𝑚∈N and (𝑡𝑚,𝑘)𝑚∈N such that 𝑠𝑚,𝑘→+∞as 𝑚→+∞, 𝑆𝑛⩾𝑥𝑛⩾𝑥𝑛−𝜀0
2 > 𝑥∗
𝑛−𝜀0,
𝑉𝑛⩾𝑦𝑛⩾𝑦𝑛−𝜀0
2 > 𝑦∗
𝑛−𝜀0
(A.71) s
1)
𝑆𝑛⩾𝑥𝑛⩾𝑥𝑛−𝜀0
2 > 𝑥∗
𝑛−𝜀0,
𝑉𝑛⩾𝑦𝑛⩾𝑦𝑛−𝜀0
2 > 𝑦∗
𝑛−𝜀0
(A.71) (A.71) 0 < 𝑠1,𝑘< 𝑡1,𝑘< 𝑠2,𝑘< 𝑡2,𝑘< ⋅⋅⋅< 𝑠𝑚,𝑘< 𝑡𝑚,𝑘
<
(A.61) 0 < 𝑠1,𝑘< 𝑡1,𝑘< 𝑠2,𝑘< 𝑡2,𝑘< ⋅⋅⋅< 𝑠𝑚,𝑘< 𝑡𝑚,𝑘 (A.61) (A.61) < . . . ,
𝐼𝑠𝑚,𝑘,𝑘> 𝜀0
𝑘,
𝐼𝑡𝑚,𝑘,𝑘< 𝜀0
𝑘2 ,
(A.62)
𝜀0
𝑘2 ⩽𝐼𝑛,𝑘⩽𝜀0
𝑘,
∀𝑛∈[𝑠𝑚,𝑘+ 1, 𝑡𝑛,𝑘−1] ∪N. (A.63)
Given 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘−1] ∪N, we have
for all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N. Proceeding like before, we obtain (A.51) and (A.52) with
𝑆𝑘and 𝑉𝑘replaced by 𝑆𝑛+1,𝑘and 𝑉𝑛+1,𝑘, respectively, and, for
sufficiently large 𝑛∈N,
𝜑(𝑆𝑛+1,𝑘, 𝐼𝑛,𝑘)
𝐼𝑛,𝑘
⩾𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 −𝜃1 (𝜀1) ,
𝜓(𝑉𝑛+1,𝑘, 𝐼𝑛,𝑘)
𝐼𝑘
⩾𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑘𝜓𝜀0 −𝜃2 (𝜀1) ,
(A.72) for all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N. for all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N. 𝑚,𝑘
𝑚,𝑘
Proceeding like before, we obtain (A.51) and (A.52) with
𝑆𝑘and 𝑉𝑘replaced by 𝑆𝑛+1,𝑘and 𝑉𝑛+1,𝑘, respectively, and, for
sufficiently large 𝑛∈N, 𝑚,𝑘
𝑚,𝑘
Proceeding like before, we obtain (A.51) and (A.52) with
𝑆𝑘and 𝑉𝑘replaced by 𝑆𝑛+1,𝑘and 𝑉𝑛+1,𝑘, respectively, and, for
sufficiently large 𝑛∈N, (A.62) 𝐼𝑡𝑚,𝑘,𝑘< 𝑘2 ,
𝜀0
𝑘2 ⩽𝐼𝑛,𝑘⩽𝜀0
𝑘,
∀𝑛∈[𝑠𝑚,𝑘+ 1, 𝑡𝑛,𝑘−1] ∪N. (A.63)
Given 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘−1] ∪N, we have
𝜑(𝑆𝑛+1,𝑘, 𝐼𝑛,𝑘)
𝐼𝑛,𝑘
⩾𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 −𝜃1 (𝜀1) ,
𝜓(𝑉𝑛+1,𝑘, 𝐼𝑛,𝑘)
𝐼𝑛,𝑘
⩾𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑘𝜓𝜀0 −𝜃2 (𝜀1) ,
(A.72) (A.72) 𝐼𝑛+1,𝑘
𝐼𝑛,𝑘
where 𝜃1(𝜀1) →0 as 𝜀1 →0. Therefore, 𝐼𝑛+1,𝑘 where 𝜃1(𝜀1) →0 as 𝜀1 →0. Therefore, where 𝜃1(𝜀1) →0 as 𝜀1 →0. Proof of the Results in Section 3 By (iii) in Lemma 3 we obtain For any 𝑛0 ∈N and 𝑥0, 𝑦0 ∈R+, let (𝑥𝑛, 𝑦𝑛) be the solution
of (12) with initial condition (𝑥𝑛0, 𝑦𝑛0) = (𝑥0, 𝑦0) and let
(𝑥𝑛0, 𝑦𝑛0) = (𝑥0, 𝑦0) be the solution of (A.42) with the same
initial condition. By (iii) in Lemma 3 we obtain Reasoning similarly we obtain, for all 𝑛⩾𝑁2, Reasoning similarly we obtain, for all 𝑛⩾𝑁2, 𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑘𝜓𝜀0 ⩽𝜕2𝜓(𝑉𝑘+1, 0)
⩽𝜕2𝜓(𝑦∗
𝑘+1, 0) + 𝑘𝜓𝜀0. (A.52) (A.52) sup
𝑛∈N0
{𝑥𝑛−𝑥𝑛+ 𝑦𝑛−𝑦𝑛}
⩽𝐿𝑀(𝛽𝑢𝑘𝜑+ 𝜎𝑢𝑘𝜓) 𝜀1
(A.43) (A.43) By (H1) and (H2), we conclude that By (H1) and (H2), we conclude that ⩽𝐿𝑀(𝛽𝑢𝑘𝜑+ 𝜎𝑢𝑘𝜓) 𝜀1 𝜑(𝑆𝑛+1, 𝐼𝑛) = 𝜑(𝑆𝑛+1, 0) + 𝜕2𝜑(𝑆𝑛+1, 𝜀𝑛) (𝐼𝑛−0)
= 𝜕2𝜑(𝑆𝑛+1, 𝜀𝑛) 𝐼𝑛,
(A.53) and thus we can take 𝜀1 > 0 small enough such that (A.53) sup
𝑛∈N0
{𝑥𝑛−𝑥𝑛+ 𝑦𝑛−𝑦𝑛} ⩽𝜀0
2 . (A.44) for some 𝜀𝑛∈[0, 𝜀1], and all 𝑛⩾𝑁2. Thus, by continuity of
𝜕2𝜑, On the other hand, by (ii) in Lemma 3, there is 𝑁2 ⩾𝑁1
sufficiently large such that 𝜑(𝑆𝑛+1, 𝐼𝑛)
𝐼𝑛
= 𝜕2𝜑(𝑆𝑛+1, 𝜀𝑛)
(A.54) 𝜑(𝑆𝑛+1, 𝐼𝑛)
𝐼𝑛
= 𝜕2𝜑(𝑆𝑛+1, 𝜀𝑛)
⩾𝜕2𝜑(𝑆𝑛+1, 0) −𝜃1 (𝜀1) ,
(A.54) 𝑥∗
𝑛−𝑥𝑛+ 𝑦∗
𝑛−𝑦𝑛⩽𝜀0
2 ,
(A.45)h
𝜑(𝑆𝑛+1, 𝑛)
𝐼𝑛
= 𝜕2𝜑(𝑆𝑛+1, 𝜀𝑛)
⩾𝜕2𝜑(𝑆𝑛+1, 0) −𝜃1 (𝜀1) ,
(A 𝑥∗
𝑛−𝑥𝑛+ 𝑦∗
𝑛−𝑦𝑛⩽𝜀0
2 ,
(A.45)
𝐼𝑛 𝑥∗
𝑛−𝑥𝑛+ 𝑦∗
𝑛−𝑦𝑛⩽𝜀0
2 ,
(A.45)
𝐼𝑛
= 𝜕2𝜑(𝑆𝑛+1, 𝜀𝑛)
⩾𝜕2𝜑(𝑆𝑛+1, 0) −𝜃1 (𝜀1) , 𝑥∗
𝑛−𝑥𝑛+ 𝑦∗
𝑛−𝑦𝑛⩽𝜀0
2 ,
(A.45) (A.54) (A.45) ⩾𝜕2𝜑(𝑆𝑛+1, 0) −𝜃1 (𝜀1) , for all 𝑛⩾𝑁2. Therefore, for all 𝑛⩾𝑁2. Therefore, Therefore,
𝑥𝑛−𝑥∗
𝑛
+ 𝑦𝑛−𝑦∗
𝑛
⩽𝜀0,
(A.46)
with 𝜃1(𝜀1) →0 as 𝜀1 →0, for all 𝑛⩾𝑁2. Thus, by (A.51)
𝜑(𝑆𝑛+1, 𝐼𝑛)
𝐼𝑛
⩾𝜕2𝜑(𝑥∗
𝑘+1, 0) −𝑘𝜑𝜀0 −𝜃1 (𝜀1) ,
(A.55) (A.55) for all 𝑛⩾𝑁2. Discrete Dynamics in Nature and Society 16 where 𝜃1(𝜀1) →0 as 𝜀1 →0. Similarly, for all 𝑛⩾𝑁2, we
h
b (A 52)
and therefore where 𝜃1(𝜀1) →0 as 𝜀1 →0. Similarly, for all 𝑛⩾𝑁2, we
have, by (A.52)
and therefore
𝜀0
𝑡
𝜎𝑡𝑛,𝑘−𝑠𝑛,𝑘𝜀0
(A
) where 𝜃1(𝜀1) →0 as 𝜀1 →0. Similarly, for all 𝑛⩾𝑁2, we
have, by (A.52)
and therefore
𝜀 where 𝜃1(𝜀1) →0 as 𝜀1 →0. Similarly, for all 𝑛⩾𝑁2, we
have, by (A.52)
(𝑉
𝐼)
and therefore
𝜀0
𝑘2 > 𝐼𝑡𝑛𝑘,𝑘⩾𝜎𝑡𝑛,𝑘−𝑠𝑛,𝑘𝐼𝑠𝑛𝑘,𝑘> 𝜎𝑡𝑛,𝑘−𝑠𝑛,𝑘𝜀0
𝑘
,
(A.65) y
𝜓(𝑉𝑛+1, 𝐼𝑛)
𝐼𝑛
⩾𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑘𝜓𝜀0 −𝜃2 (𝜀1) ,
(A.56)
where 𝜃2(𝜀1) →0 as 𝜀1 →0. Proof of the Results in Section 3 From the second equation in (1),
we have
𝜀0
𝑘2 > 𝐼𝑡𝑛,𝑘,𝑘⩾𝜎𝑡𝑛,𝑘−𝑠𝑛,𝑘𝐼𝑠𝑛,𝑘,𝑘> 𝜎𝑛,𝑘
𝑛,𝑘𝜀0
𝑘
,
(A.65)
where
𝜎=
1
1 + 𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢. (A.66) 𝜓(
𝑛+1
𝑛)
𝐼𝑛
⩾𝜕2𝜓(𝑦𝑘+1, 0) −𝑘𝜓𝜀0 −𝜃2 (𝜀1) ,
(A.56)
where 𝜃2(𝜀1) →0 as 𝜀1 →0. From the second equation in (1),
we have
where
𝜎=
1
1 + 𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢. (A.66) →0. From the second equation in (1),
𝜎=
1
1 + 𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢. (A.66) where 𝜃2(𝜀1) →0 as 𝜀1 →0. From the second equation in (1),
we have
𝜎=
1
1 + 𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢. (A.66) (A.66) Thus, by (A.62) and (A.65), 𝐼𝑛+1 = 1 + 𝛽𝑛𝜑(𝑆𝑛+1, 𝐼𝑛) /𝐼𝑛+ 𝜎𝑛𝜓(𝑉𝑛+1, 𝐼𝑛) /𝐼𝑛
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
𝐼𝑛
⩾1 + 𝛽𝑛𝜕2𝜑(𝑥∗
𝑘+1, 0) + 𝜎𝑛𝜕2𝜓(𝑦∗
𝑘+1, 0) −𝑢
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
𝐼𝑛,
(A.57)
Thus, by (A.62) and (A.65),
𝑡𝑛,𝑘−𝑠𝑛,𝑘⩾
ln 𝑘
ln (1/𝜎) →+∞
as 𝑘→+∞. (A.67)
In view of (A.67), we can choose 𝑘0 ∈N such that (A.67) In view of (A.67), we can choose 𝑘0 ∈N such that In view of (A.67), we can choose 𝑘0 ∈N such that 𝑡𝑛,𝑘−𝑠𝑛,𝑘> 𝑁+ 𝜆+ 1,
(A.68) where (A.68) 𝑢= 𝛽𝑢(𝑘𝜑𝜀0 + 𝜃1 (𝜀1)) + 𝜎𝑢(𝑘𝜓𝜀0 + 𝜃2 (𝜀1)) ,
(A.58) for all 𝑘⩾𝑘0. Letting 𝑚and 𝑘⩾𝑘0 be sufficiently large and 𝜀2 > 0 be
sufficiently small, we may assume that for all 𝑛⩾𝑁2. Letting 𝜀1 be sufficiently small, we conclude,
according to (A.37) and (A.57), that 𝐼𝑛→+∞as 𝑛→+∞, a
contradiction. Thus, we conclude that (A.38) holds. 𝑆𝑛,𝑘⩾
Λ 𝑛+ 𝜂𝑛𝑉𝑛,𝑘+ 𝑆𝑛,𝑘−𝛽𝑢𝑘𝜑𝑀𝜀2
1 + 𝜇𝑛+ 𝑝𝑛
𝑉𝑛,𝑘⩾
𝑝𝑛𝑆𝑛,𝑘+ 𝑉𝑛,𝑘−𝜎𝑢𝑘𝜓𝑀𝜀2
1 + 𝜇𝑛+ 𝜂𝑛
(A.69) h
Next we will prove the permanence of the infectives. By
(iii) in Lemma 4, it is only necessary to prove that there is an
𝜀2 > 0 such that, for any solution (𝑆𝑛, 𝐼𝑛, 𝑅𝑛, 𝑉𝑛) of (1) with
positive initial conditions, we have (A.69) lim inf
𝑛→+∞𝐼𝑛> 𝜀2. (A.59) holds for all 𝑛∈[𝑠𝑚,𝑘+ 1, 𝑡𝑚,𝑘−1] ∩N. holds for all 𝑛∈[𝑠𝑚,𝑘+ 1, 𝑡𝑚,𝑘−1] ∩N. (A.59) [ 𝑚,𝑘
, 𝑚,𝑘
]
Let (𝑥𝑛, 𝑦𝑛) be a solution of (A.42) with initial value
(𝑥𝑠𝑚,𝑘, 𝑦𝑠𝑚,𝑘) = (𝑆𝑠𝑚,𝑘, 𝑉𝑠𝑚,𝑘). We have 𝑆𝑛⩾𝑥𝑛and 𝑉𝑛⩾𝑦𝑛for
all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N. References [19] P. E. Kloeden and C. P¨otzsche, “Nonautonomous dynamical
systems in the life sciences,” in Nonautonomous dynamical
systems in the life sciences, vol. 2102 of Lecture Notes in Math.,
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demiology of Infectious Diseases, Model Building, Analysis and
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class of discrete SIRS epidemic models with nonlinear incidence
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non-autonomous epidemic model with vaccination,” Applicable
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solutions of a discrete epidemic model,” Discrete Dynamics in
Nature and Society, Art. ID 434537, 10 pages, 2015. [22] L. J. Allen, M. A. Jones, and C. F. Martin, “A discrete-time
model with vaccination for a measles epidemic,” Mathematical
Biosciences, vol. 105, no. 1, pp. 111–131, 1991. [5] J. Liu, B. Peng, and T. Zhang, “Effect of discretization on
dynamical behavior of SEIR and SIR models with nonlinear
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infectives and susceptibles,” Theoretical Population Biology. An
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the article is available in the websites: http://www.worldbank
.org, https://data.oecd.org, and http://www.geoba.se. [14] R. E. Mickens, “Discretizations of nonlinear differential equa-
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sion,” Journal of Difference Equations and Applications, vol. 18,
no. 3, pp. 525–529, 2012. Proof of the Results in Section 3 Therefore, = 1 + 𝛽𝑛𝜑(𝑆𝑛+1,𝑘, 𝐼𝑛,𝑘) /𝐼𝑛,𝑘+ 𝜎𝑛𝜓(𝑉𝑛+1,𝑘, 𝐼𝑛,𝑘) /𝐼𝑛,𝑘
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(A.64)
1(
𝐼𝑛+1,𝑘 = 1 + 𝛽𝑛𝜑(𝑆𝑛+1,𝑘, 𝐼𝑛,𝑘) /𝐼𝑛,𝑘+ 𝜎𝑛𝜓(𝑉𝑛+1,𝑘, 𝐼𝑛,𝑘) /𝐼𝑛,𝑘
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(A.64)
1
1
1h
𝐼𝑛+1,𝑘
1
𝛽𝜕
( ∗
0)
𝜕
(
∗
0)
(A (A.73) ⩾1 + 𝛽𝑛𝜕2𝜑(𝑥∗
𝑛+1, 0) + 𝜎𝑛𝜕2𝜓(𝑦∗
𝑛+1, 0) −𝑢
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
𝐼𝑛,𝑘,
(A.73) ⋅𝐼𝑛,𝑘⩾
1
1 + 𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢𝐼𝑛,𝑘
⩾1 + 𝛽𝑛𝜕2𝜑(𝑥∗
𝑛+1, 0) + 𝜎𝑛𝜕2𝜓(𝑦∗
𝑛+1, 0) −𝑢
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
𝐼𝑛,𝑘,
(A.73) ⋅𝐼𝑛,𝑘⩾
1
1 + 𝜇𝑢+ 𝛼𝑢+ 𝛾𝑢𝐼𝑛,𝑘 Discrete Dynamics in Nature and Society 17 for all 𝑛∈[𝑠𝑚,𝑘, 𝑡𝑚,𝑘] ∩N, where 𝑢is given by (A.58). Thus, [10] Q. Cui and Q. Zhang, “Global stability of a discrete SIR epidemic
model with vaccination and treatment,” Journal of Difference
Equations and Applications, vol. 21, no. 2, pp. 111–117, 2015. 𝜀0
𝑘2 > 𝐼𝑡𝑚,𝑘,𝑘
⩾𝐼𝑡𝑚,𝑘−𝜆,𝑘
𝑡𝑚,𝑘
∏
𝑛=𝑡𝑚,𝑘−𝜆
1 + 𝛽𝑛𝜕2𝜑(𝑥∗
𝑛+1, 0) + 𝜎𝑛𝜕𝑛𝜓(𝑦∗
𝑛+1, 0) −𝑢
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
> 𝜀0
𝑘2
(A.74) [11] D. Ding, Q. Ma, and X. Ding, “A non-standard finite difference
scheme for an epidemic model with vaccination,” Journal of
Difference Equations and Applications, vol. 19, no. 2, pp. 179–190,
2013. 𝑘2
𝑚,𝑘,
⩾𝐼𝑡𝑚,𝑘−𝜆,𝑘
𝑡𝑚,𝑘
∏
𝑛=𝑡𝑚,𝑘−𝜆
1 + 𝛽𝑛𝜕2𝜑(𝑥∗
𝑛+1, 0) + 𝜎𝑛𝜕𝑛𝜓(𝑦∗
𝑛+1, 0) −𝑢
1 + 𝜇𝑛+ 𝛼𝑛+ 𝛾𝑛
(A.74) [12] D. Ding and X. Ding, “Dynamic consistent non-standard
numerical scheme for a dengue disease transmission model,”
Journal of Difference Equations and Applications, vol. 20, no. 3,
pp. 492–505, 2014. which is a contradiction. The theorem follows. [13] S. Jang and S. Elaydi, “Difference equations from discretization
of a continuous epidemic model with immigration of infec-
tives,” Canadian Applied Mathematics Quarterly, vol. 11, no. 1,
pp. 93–105, 2003. Conflicts of Interest The authors declare that they have no conflicts of interest. [16] R. E. Mickens, “Nonstandard finite difference schemes for
differential equations,” Journal of Difference Equations and
Applications, vol. 8, no. 9, pp. 823–847, 2002. Acknowledgments [17] C. Sun and W. Yang, “Global results for an SIRS model
with vaccination and isolation,” Nonlinear Analysis: Real World
Applications, vol. 11, no. 5, pp. 4223–4237, 2010. Joaquim
Mateus,
C´esar
M. Silva,
and
S. Vaz
were
partially supported by FCT through CMA-UBI (Project
UID/MAT/00212/2013). [18] A. A. Khasnis and M. D. Nettleman, “Global warming and
infectious disease,” Archives of Medical Research, vol. 36, no. 6,
pp. 689–696, 2005. References 119, no. 1, pp. 97–114, 1994. [8] E. Pereira, C. M. Silva, and J. A. da Silva, “A generalized
nonautonomous SIRVS model,” Mathematical Methods in the
Applied Sciences, vol. 36, no. 3, pp. 275–289, 2013. [26] K. Hattaf, A. A. Lashari, B. El Boukari, and N. Yousfi, “Effect
of discretization on dynamical behavior in an epidemiological
model,” Differential Equations and Dynamical Systems, vol. 23,
no. 4, pp. 403–413, 2015. [9] Q. Cui, X. Yang, and Q. Zhang, “An NSFD scheme for a class of
SIR epidemic models with vaccination and treatment,” Journal
of Difference Equations and Applications, vol. 20, no. 3, pp. 416–
422, 2014. [27] V. Kaitala, M. Heino, and W. M. Getz, “Host-parasite dynamics
and the evolution of host immunity and parasite fecundity Discrete Dynamics in Nature and Society 18 strategies,” Bulletin of Mathematical Biology, vol. 59, no. 3, pp. 427–450, 1997. [28] G. Di Lena and G. Serio, “A discrete method for the iden-
tification of parameters of a deterministic epidemic model,”
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Infectious disease with a random number of contacts between
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immunity: a synthesis,” Mathematical Biosciences, vol. 82, no. 1,
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|
https://openalex.org/W2988193221
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http://addi.ehu.es/bitstream/10810/40406/1/ijerph-16-04381.pdf
|
English
| null |
Causal Effects of Prenatal Exposure to PM2.5 on Child Development and the Role of Unobserved Confounding
|
International journal of environmental research and public health/International journal of environmental research and public health
| 2,019
|
cc-by
| 8,579
|
Causal Effects of Prenatal Exposure to PM2.5 on Child
Development and the Role of
Unobserved Confounding Viola Tozzi 1, Aitana Lertxundi 2,3,4, Jesus M. Ibarluzea 2,3,5,6
and Michela Baccini 1,* 1
Department of Statistics, Computer Science, Applications, University of Florence, 59 50134 Florence, Viale
Morgagni, Italy, viola.t@hotmail.it Faculty of Medicine, University of the Basque Country (UPV/EHU), 48940 Leioa, Spain
5
Sub-Directorate for Public Health of Guipúzcoa, Department of Health, Government of the Basque Country,
20013 San Sebastian, Spain 6
Faculty of Psychology, University of the Basque Country (UPV/EHU), 20018 San Sebastian, Spain
*
Correspondence: michela.baccini@unifi.it; Tel.: +39 055 2751563 y
y
gy
y
q
y
p
*
Correspondence: michela.baccini@unifi.it; Tel.: +39 055 2751563 Received: 12 September 2019; Accepted: 6 November 2019; Published: 9 November 2019 ceived: 12 September 2019; Accepted: 6 November 2019; Published: 9 November 2019
Abstract: Prenatal exposure to airborne particles is a potential risk factor for infant neuropsychological
development. This issue is usually explored by regression analysis under the implicit assumption
that all relevant confounders are accounted for. Our aim is to estimate the causal effect of prenatal
exposure to high concentrations of airborne particles with a diameter < 2.5 µm (PM2.5) on children’s
psychomotor and mental scores in a birth cohort from Gipuzkoa (Spain), and investigate the
robustness of the results to possible unobserved confounding. We adopted the propensity score
matching approach and performed sensitivity analyses comparing the actual effect estimates with
those obtained after adjusting for unobserved confounders simulated to have different strengths. On average, mental and psychomotor scores decreased of −2.47 (90% CI: −7.22; 2.28) and −3.18
(90% CI: −7.61; 1.25) points when the prenatal exposure was ≥17 µg/m3 (median). These estimates
were robust to the presence of unmeasured confounders having strength similar to that of the observed
ones. The plausibility of having omitted a confounder strong enough to drive the estimates to zero
was poor. The sensitivity analyses conferred solidity to our findings, despite the large sampling
variability. This kind of sensitivity analysis should be routinely implemented in observational studies,
especially in exploring new relationships. Keywords: child development; airborne particles; propensity score matching; sensitivity analysis;
bias analysis; Monte Carlo simulations International Journal of
Environmental Research
and Public Health International Journal of
Environmental Research
and Public Health 1. Introduction Many studies around the world documented long term and short term effects of air pollution on
population mortality and morbidity [1,2]. Recently, several researches focused on the effects of prenatal
exposures to air pollution on children’s neuropsychological development [3]. Harris et al. [4] analyzed
1109 mother–child pairs from Eastern Massachusetts (USA) concluding that parental proximity to
major roadways may negatively influence performance across a range of mental domains in childhood. Guxens et al. [5] found significant associations between high levels of nitrogen dioxide (NO2) and
benzene and infant neurodevelopment in the Spanish cohorts participating in the INMA (Infancia
y Medio Ambiente, the Spanish for Childhood and Environment) Project. High levels of NO2 were Int. J. Environ. Res. Public Health 2019, 16, 4381; doi:10.3390/ijerph16224381 www.mdpi.com/journal/ijerph Int. J. Environ. Res. Public Health 2019, 16, 4381 2 of 12 found to be associated also with delayed psychomotor development in children between one and six
years of age [6]. Lertxundi et al. [7,8] investigated the effect of prenatal exposure to airborne particles
with a diameter < 2.5 µm (PM2.5) and NO2 on infant cognitive and psychomotor development and on
the cognitive functions in children of 4–6 years of age, reporting the existence of negative associations. All these studies accounted for the confounding effect of several mother and child characteristics
through the specification of regression models, under the implicit assumption that all relevant
confounders were known and observed (assumption of unconfoundedness [9]). However, given the
complex nature of the problem, the presence of residual confounding due to unknown or unmeasured
factors related to the familiar and environmental context where children live should not be excluded. In the literature, the problem related to the presence of unobserved confounders and the evaluation of
the associated bias has been addressed through sensitivity analyses that check whether and to what
extent the estimated effect or association is robust to possible deviations from unconfoundedness [10–18]. Usually, the aim is to assess how strong an unobserved confounder would have to be to substantially
change the study results. However, despite the importance of this issue, quantitative evaluations
of the potential bias associated with residual confounding have rarely received consideration in
epidemiological applications (see chapter 19 in [10]). 1. Introduction In the present paper, with reference to a birth
cohort of the province of Gipuzkoa [7], we estimated the causal effect of high PM2.5 exposures during
maternal pregnancy on child mental and psychomotor outcomes at 15 months of age, within a potential
outcomes approach to causal inference [19]. Then, we checked the robustness of the results to the
possible presence of unobserved confounders through a sensitivity analysis based on simulations. In particular, we compared the actual causal effect estimates obtained by using propensity score (PS)
matching [9], to those obtained after controlling also for simulated unobserved confounders of different
strengths, according to the method proposed by Ichino et al. [11] and implemented in the sensatt
function of STATA (StataCorp LP., College Station, TX, USA) [20,21]. 2.1. Data As part of the Environment and Childhood Research Network (INMA network) [22], the birth
cohort of Gipuzkoa was recruited between May 2006 and January 2008 among the residents of a
study area of 519 km2 spanning three narrow valleys: high Urola-Goierri, mid Urola and high Oria. The total population of the area was approximately 88,000 inhabitants, spread across 25 small localities. Data on exposures, outcomes and potential confounders were available for 438 mother–child pairs. Children’s mental and psychomotor developments were assessed at around 15 months of age (range
13–18 months), according to the Bayley Scales of Infant Development [23], by one of two trained
neuropsychologists, blinded to the child’s exposure status. The raw scores were standardized to a mean
of 100 points with a standard deviation of 15 points, with higher scores indicating better developments. Data on the mother’s characteristics and habits were obtained through questionnaires administered
during the first and third trimesters of pregnancy. The daily levels of PM2.5 in the area were measured
from the beginning of the study until the last birth by three fixed-site samplers belonging to the Air
Quality Network of the Basque Government and three Digitel DHA-80 high-volume aerosol samplers,
two of which were monthly rotated in different locations (Figure 1). Due to the presence of moving monitors, daily measurements were characterized by the presence
of design-missing values. With the aim to assess the average exposure of the mother–child couples
during pregnancy, we used a multiple imputation (MI) procedure starting from the weekly data. First,
we derived weekly means from the daily levels of PM2.5 measured in each location. Considering a
weekly mean as missing when daily measurements were missing for more than two days during the
week, the percentage of missing weekly averages during the study period was around 30%. Then,
under the Missing At Random (MAR) assumption, we applied an MI procedure [24] on the incomplete
data set including weekly averages concentrations of the air pollutants measured by the monitors, 3 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 meteorological variables and seasonality indicators. In this way, we generated five imputed data
sets, thus five imputed values for each missing weekly mean of PM2.5. Finally, we averaged over
weekly means, obtaining five imputed values of the average level of PM2.5 during the pregnancy for
each mother–child couple. Figure 1
2.2. Notation 2.2. Notation
Unless otherwise specified the notation and the methods described hereinafter refer to a single
Unless otherwise specified, the notation and the methods described hereinafter refer to a single
imputed data set. 2.2. Notation
U l
th
i
ifi d th
t ti
d th
th d d
ib d h
i
ft
f
t
i
l
Unless otherwise specified, the notation and the methods described hereinafter refer to a single
imputed data set. Unless otherwise specified, the notation and the methods described hereinafter refer to a single
imputed data set. Being i = 1, N the indicator of the child, let Ei be a binary exposure indicator, which was equal to
0 if the average level of PM2.5 during the pregnancy was <17 µg/m3 (control) and equal to 1 if it was
≥17 µg/m3 (treatment). The threshold of 17 µg/m3, which approximated the median exposure of the
it
th i
t d d t
t
fi
d
i
i i
d
t
id
t h
d t d
d ti
[25]
Being i = 1, N the indicator of the child, let Ei be a binary exposure indicator, which was equal to 0
if the average level of PM2.5 during the pregnancy was <17 µg/m3 (control) and equal to 1 if it was
≥17 µg/m3 (treatment). The threshold of 17 µg/m3, which approximated the median exposure of the
units across the imputed data sets, was fixed a priori in order to avoid post-hoc data dredation [25]. Let Yi be the observed mental or psychomotor score for child i and Xi a vector of observed covariates. units across the imputed data sets, was fixed a priori in order to avoid post-hoc data dredation [25]. Let Yi be the observed mental or psychomotor score for child i and Xi a vector of observed covariates. According to the potential outcome approach to causal inference, under the Stable Unit
Treatment Value Assumption, we associated with each child two potential outcomes Yi(1) and Yi(0),
representing the score under treatment and under control [9]. 2.1. Data We assumed that the mother–child couples were exposed to PM2.5 levels
measured at the monitoring site located in their town or in the closest one. For more detail on the MI
procedure see [7]. generated five imputed data sets, thus five imputed values for each missing weekly mean of PM2.5. Finally, we averaged over weekly means, obtaining five imputed values of the average level of PM2.5
during the pregnancy for each mother–child couple. We assumed that the mother–child couples were
exposed to PM2.5 levels measured at the monitoring site located in their town or in the closest one. For more detail on the MI procedure see [7]. The study was approved by the hospital ethics committee and all participating mothers provided
informed consent. The study was approved by the hospital ethics committee and all participating mothers
provided informed consent. Figure 1. Study area and location of the particles with a diameter < 2.5 µm (PM2.5) air samplers. Figure 1. Study area and location of the particles with a diameter < 2.5 µm (PM2.5) air samplers. Figure 1
2.2. Notation For each child it was possible to observe
only one of the two potential outcomes: for children belonging to the treated group we observed only
According to the potential outcome approach to causal inference, under the Stable Unit Treatment
Value Assumption, we associated with each child two potential outcomes Yi(1) and Yi(0), representing
the score under treatment and under control [9]. For each child it was possible to observe only one of
the two potential outcomes: for children belonging to the treated group we observed only Y(1) (Yi(1) =
Yi if Ei = 1); for children belonging to the control group we observed only Y(0) (Yi(0) = Yi if Ei = 0). 2.4. Assumption and ATT estimation We included in the PS model not only pre-treatment variables, as usually
recommended [27], but also some post-treatment variables (smoking and diet during pregnancy, child
care mode, breastfeeding) which were not affected by the exposure and could be thought of as proxies
for unmeasured pre-treatment covariates, mainly related to the socio-economic status of the family. We expect that adjusting for them could reduce the bias without inflating the variance [28,29]. (1) For each child, we estimated the PS as predicted value from a logistic model defined on the
exposure Ei, given Xi [26]. The model accounted for the following variables: child’s sex, year of
birth, nursery attendance (yes, no), main caregiver (only mother, both parents, other relative, other
caregiver), maternal education (secondary school or less, university), maternal work (non-manual
worker, manual worker), mother’s age (<25, 25–34, 35+), mother’s body mass index (underweight,
normal, overweight, obese), mother’s parity (0, 1+), mother’s vegetable and fruit intake during
pregnancy (≤405 g/day, >405 g/day), smoking during pregnancy (yes, no), smoking at 32 weeks of
pregnancy (yes, no), breastfeeding (yes, no), indicator of the neuropsychologist who evaluated the
child (first, second evaluator). We included in the PS model not only pre-treatment variables, as usually
recommended [27], but also some post-treatment variables (smoking and diet during pregnancy, child
care mode, breastfeeding) which were not affected by the exposure and could be thought of as proxies
for unmeasured pre-treatment covariates, mainly related to the socio-economic status of the family. We expect that adjusting for them could reduce the bias without inflating the variance [28,29]. Regarding the need of adjusting for the neuropsychologist indicator, it is worth noting that the
probability of exposures ≥17 µg/m3 was very low (from 1% to 5%) for the children examined by the
second neuropsychologist, who substituted the first one at the end of the study period. The observed
decrease in the exposures over time was mainly due to the substantial reduction of the atmospheric level
of PM2.5 in 2007, likely attributable to the high level of precipitations (58.8 mm in 2006 versus 126.5 mm
in 2007) and to the closure of a polluting industry in the area in July of the same year (Figure S1) [30]. This, coupled with the fact that the scores assigned by the second psychologist were systematically
higher than those assigned by the first one, made crucial to adjust for the neuropsychologist indicator. y
p
Y(1) (Yi(1) = Yi if Ei = 1); for children be
if E
0)
2.3. Research Question and Estimands if Ei = 0). 2.3. Research Question and Estimands
The research question we were interested in was the following: what would have been the
mental and psychomotor development of the children with prenatal exposure ≥17 µg/m3, if their
exposure were been <17 µg/m3? In order to reply to this questions, we focused, separately for
h
d
l
h
l ff
f
h
d (ATT)
hi h
The research question we were interested in was the following: what would have been the mental
and psychomotor development of the children with prenatal exposure ≥17 µg/m3, if their exposure
were been <17 µg/m3? In order to reply to this questions, we focused, separately for psychomotor
and mental score, on the average causal effect of treatment on the treated (ATT), which compares the
average score observed among the “treated” children, with the average score that we would have
observed on them if they were not “treated”: Int. J. Environ. Res. Public Health 2019, 16, 4381 4 of 12 ATT =
PN
i=1 Yi(1)Ei
PN
i=1 Ei
−
PN
i=1 Yi(0)Ei
PN
i=1 Ei
=
PN
i=1 YiEi
PN
i=1 Ei
−
PN
i=1 Yi(0)Ei
PN
i=1 Ei
. (1) (1) 2.4. Assumption and ATT estimation In order to estimate the ATT, we imputed the missing potential outcome Yi(0) in (1) through a
matching procedure based on the PS [9]. This required two assumptions [26]: (i) Unconfoundedness for controls: Ei⊥Yi(0)|Xi. This assumption states that treatment assignment
is independent of the potential outcome under control, conditional on the observed covariates. In other
words, this assumption requires that there are no unobserved confounders. (i) Unconfoundedness for controls: Ei⊥Yi(0)|Xi. This assumption states that treatment assignment
is independent of the potential outcome under control, conditional on the observed covariates. In other
words, this assumption requires that there are no unobserved confounders. p
q
(ii) Weak positivity assumption: P(Ei = 1|Xi = x) < 1 for each i, for each x. This assumption implies
that for each treated unit there is a similar untreated unit that can be used as a matched control. p
q
(ii) Weak positivity assumption: P(Ei = 1|Xi = x) < 1 for each i, for each x. This assumption implies
that for each treated unit there is a similar untreated unit that can be used as a matched control. (ii) Weak positivity assumption: P(Ei = 1|Xi = x) < 1 for each i, for each x. This assumption implies
for each treated unit there is a similar untreated unit that can be used as a matched control. Being the PS a balancing score, the conditions (i) and (ii) are valid also conditionally to the PS [26]. Under (i) and (ii), we applied a PS matching procedure for AT estimation which consisted of
two steps: p
(1) For each child, we estimated the PS as predicted value from a logistic model defined on the
exposure Ei, given Xi [26]. The model accounted for the following variables: child’s sex, year of
birth, nursery attendance (yes, no), main caregiver (only mother, both parents, other relative, other
caregiver), maternal education (secondary school or less, university), maternal work (non-manual
worker, manual worker), mother’s age (<25, 25–34, 35+), mother’s body mass index (underweight,
normal, overweight, obese), mother’s parity (0, 1+), mother’s vegetable and fruit intake during
pregnancy (≤405 g/day, >405 g/day), smoking during pregnancy (yes, no), smoking at 32 weeks of
pregnancy (yes, no), breastfeeding (yes, no), indicator of the neuropsychologist who evaluated the
child (first, second evaluator). 2.5. Sensitivity Analysis The sensatt function generates U starting from four tuning parameters: pij = Pr(U = 1
E = i, Y∗= j),
i = 0, 1, j = 0, 1, and calculates the odds ratios Γ and Λ from the simulations. In our study, we focused on unobserved confounders with a distribution similar to the empirical
distribution of relevant observed confounders, by deriving the pij from the observed data. Successively,
we explored the whole space of the hypothetical Us by setting pij = 0.1, 0.25, 0.5, 0.75, 0.9, i = 0, 1, j = 0, 1,
in order to search for the so-called “killer” confounder, i.e., the confounder U able to explain
away the estimated effect, driving the estimated ATT to zero. In this second case, it is crucial to
assess the plausibility of the configurations of Γ and Λ which lead to the killer confounder: if the
killer configuration is unlikely, the ATT estimate can be considered robust to the violation of the
unconfoundedness assumption. All the sensitivity analyses were separately performed on each of the five multiple-imputed data
sets and the results combined according to Rubin [32]. 2.4. Assumption and ATT estimation Once the relevant covariates for the PS estimation have been defined, the key criterion for driving
the specification of the PS model was to obtain PS estimates that balanced the covariate distribution
between treated and controls [9]. According to the procedure implemented in the STATA command
pscore, we checked the balancing property of the PS by comparing the covariate distribution between
the two groups of exposure within strata of units with similar estimated PS [31]. We also checked for
the weak positivity condition (ii), by the inspection of the PS distributions among treated and controls. (2) On the basis of the estimated PS, we matched each treated child with the child in the control
group having closest PS, Yc
i . The same matched control could be used for different treated units,
in order to reduce the bias due to inappropriate matching. Finally, the ATT was estimated as: ˆ
ATT =
PN
i=1 YiEi
PN
i=1 Ei
−
PN
i=1 Yc
i Ei
PN
i=1 Ei
. (2) (2) The standard error of
ˆ
ATT was estimated without accounting for the uncertainty around the
estimated PS [31]. 5 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 The estimation procedure was separately performed on each of the five multiple-imputed data
sets, and the five results combined according to Rubin [32]. It is worth noticing that, for each
multiple-imputed data set, the ATT was calculated on different treated populations. This required
a cautionary interpretation of the ATT, which referred to different sets of children depending on
the imputation. 2.5. Sensitivity Analysis The Ichino’s sensitivity analysis, which is mainly employed in social science and economics,
is based on Monte Carlo simulations [11]. Let us suppose that the unconfoundedness assumption (i) is
not valid because of the presence of a binary unobserved confounder U. A value of U is attributed to
each subject by sampling from a hypothetical distribution, then U is included in the set of variables
used to estimate the ATT. This procedure can be repeated many times (e.g., 1000) and, at the end,
an average ATT is estimated. Comparing this average ATT with the causal estimate obtained without
adjusting for U, the robustness of the result to violation of the unconfoundedness assumption can be
checked: if the ATT estimates before and after adjusting for U are close, the result can be considered
robust to the presence of an unobserved confounder similar to U. In the present paper, we explored the space of the possible unobserved confounders by generating
different versions of U, which covered different hypotheses on confounding, as expressed by the
associations of U with the outcome and with the exposure. According to Ichino et al. [11], indicating
with Y∗a binary version of the outcome (for example Y∗= 1 if Y > 75th percentile, Y∗= 0 otherwise),
the association of U with the outcome (outcome effect) is measured by the odds ratio Γ comparing the
odds of Y∗= 1 when U is equal to 1 with the odds of Y∗= 1 when U is equal to 0, among the controls: Γ =
P(Y∗= 1|E = 0,U = 1,X)
P(Y∗= 0|E = 0,U = 1,X)
P(Y∗= 1|E = 0,U = 0,X)
P(Y∗= 0|E = 0,U = 0,X)
(3) (3) The association of U with the exposure (selection effect) is measured by the odds ratio Λ comparing
the odds of being exposed when U is equal to 1 with the odds of being exposed when U is equal to 0: Λ =
P(E = 1|U = 1,X)
P(E = 0|U = 1,X)
P(E = 1|U = 0,X)
P(E = 0|U = 0,X)
(4) (4) The sensatt function generates U starting from four tuning parameters: pij = Pr(U = 1
E = i, Y∗= j),
i = 0, 1, j = 0, 1, and calculates the odds ratios Γ and Λ from the simulations. 3. Results The analyses were conducted on the subset of the 391 children without missing covariates. The
covariates distribution in this subset did not substantially differ from that observed on the entire data
set (Table 1). 6 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 Table 1. Distribution of the characteristics of the child–mother couples in the entire data set and in the
subset of the complete observations by exposure. N. of Missing
Values
Entire Data Set
(n = 438)
Complete Cases
(n = 391)
Percentage/Mean
Percentage/Mean
Treated
Controls
Treated
Controls
Gender
0
Male
45.7
44.7
47.6
45.4
Female
54.3
55.3
52.4
54.6
Neuropsychologist
0
First
98.4
47.8
97.8
50.6
Second
1.6
52.2
2.2
49.4
Maternal fruit and
veg. intake
5
>405 g/day
76.2
70.6
77.7
70.4
≤405 g/day
23.8
29.4
22.3
29.6
Maternal smoke
13
Yes
18.2
28.3
18.4
27.4
No
81.8
71.7
81.6
72.6
Smoke at 32 weeks
of pregnancy
34
Yes
11.3
11.0
11.0
11.1
No
88.7
89.0
89.0
88.9
Nursery attendance
26
Yes
48.6
44.6
48.7
44.9
No
51.4
55.4
51.3
55.1
Caregiver
28
Mother
55.6
48.2
55.2
48.0
Other
44.4
51.8
44.8
52.0
Maternal education
2
University
47.6
53.9
44.8
53.0
Secondary
school or less
52.4
46.1
55.2
47.0
Maternal work
0
Manual
45.6
38.3
46.0
36.6
Non manual
54.4
61.7
54.0
63.4
Mother’s age
0
(mean)
31.2
31.7
31.3
31.7
Mother’s Body
Mass Index
1
(mean)
23.1
22.7
23.2
22.8
Parity
0
1+
46.4
44.4
45.6
42.8
0
53.6
55.6
54.4
57.2
Breastfeeding
24
No
46.6
47.6
47.1
45.5
Yes
53.4
52.4
52.9
54.5
The reported proportions (for the categorical variables) and means (for the continuous variables) come from the
combination of the same quantities arising from the 5 imputed data sets [32]. The reported proportions (for the categorical variables) and means (for the continuous variables) come from the
combination of the same quantities arising from the 5 imputed data sets [32]. The reported proportions (for the categorical variables) and means (for the co
combination of the same quantities arising from the 5 imputed data sets [32]. The procedure for the balancing check implemented in the pscore function did not find a relevant
difference between the covariate distribution of treated and controls within strata of children with
similar estimated PS (results not reported). 3. Results Moreover, the distributions of the estimated PS among
treated and controls substantially overlapped (Table S1), indicating that it was possible to find, for each
treated, a matched control with similar PS, as stated by the positivity condition (ii). We would like to
stress that the fact that the probability of being treated was close to zero for the children evaluated by
the second neuropsychologist (Table 1) did not affect the assumption (ii). The number of subjects in the high exposure group varied from 183 to 213 depending on the
imputed data set. Similarly, the number of matched controls varied from 62 to 69. The population
of the treated units, which the ATT referred to, was quite stable across imputed data sets: for more
than 90% of the 391 analyzed children the exposure status did not change across the imputed data
sets: 172 children were “always exposed” and 181 were “always unexposed” across the five imputed
data sets (Figure S2). Likely, the exposure status was uncertain only for children with actual average
prenatal exposure close to the threshold of 17 µg/m3. Combining the estimates of the ATT from the five imputations, we found that the expected scores
under PM2.5 ≥17µg/m3 were lower than the average score under PM2.5< 17 µg/m3, with final ATT
estimates equal to −2.47 (90% CI: −7.22, 2.28) for the mental score and −3.18 (90% CI: −7.61, 1.25) for
the psychomotor score. In all the analyses, the statistical variability was due to both the within and
between-imputation variances, with the second being less than 10% of the first one (Table S2). 7 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 Tables 2 and 3 report the results of the sensitivity analyses conducted by simulating several
versions of U mimicking the behavior of the observed confounders. In doing this we selected some
relevant categories of the confounders and simulated U according to the corresponding indicators
(this was necessary because the pscore function permits only for the definition of binary unobserved
confounders). The odds ratios Γ and Λ were calculated defining Y* = 1 if Y > 75th percentile and 0
otherwise. Even if in principle the selection effect Λ should be the same in the mental and psychomotor
scores analyses, the values reported in Tables 1 and 2 are different because they arise from different
simulations. 3. Results The odds ratio Γ ranged from 0.20 to 2.25 for mental score and from 0.31 to 4.30 for
psychomotor score; Λ ranged from 0.57 to very large values (>103). This indicates that we were
exploring a fairly wide class of possible unobserved confounders. Table 2. Results of the simulation-based sensitivity analysis for mental score. Observed Confounder Mimicked by U
ATT
90% CI
Outcome
Effect (Γ)
Selection
Effect (Λ)
Gender: male, female (ref)
−1.88
−7.12; 3.35
0.57
1.07
Neuropsychologist: first, second (ref)
0.78
−7.02; 8.57
0.20
>103
Maternal fruit and vegetable intake:
≤405 g/day (ref), >405 g/day
−1.89
−7.24; 3.47
1.04
1.55
Smoke: yes, no (ref)
−1.73
−7.11; 3.65
1.18
0.62
Smoke at 32 weeks of pregnancy:
yes, no (ref)
−2.07
−7.35; 3.22
1.76
1.19
Nursery attendance: yes (ref), no
−1.88
−7.13; 3.37
1.05
0.89
Caregiver: both parents, other (ref)
−1.81
−7.16; 3.55
1.75
0.90
Caregiver: mother, other (ref)
−1.85
−7.14; 3.44
0.85
1.42
Caregiver: relative, other (ref)
−2.11
−7.28; 3.07
0.39
0.74
Maternal education: secondary school or less,
university (ref)
−1.74
−7.13; 3.65
0.74
1.43
Maternal work: non-manual worker (ref),
manual worker
−1.75
−7.12; 3.61
0.63
1.52
Mother’s age: <25, 25+ (ref)
−2.12
−7.29; 3.06
0.31
3.00
Mother’s age: <35 (ref), 35+
−1.92
−7.25; 3.41
0.70
0.58
Body Mass Index: Normal weight, other (ref)
−1.89
−7.22; 3.43
2.25
1.00
Parity: 0 (ref), 1+
−1.92
−7.16; 3.32
0.97
1.15
Breastfeeding: no, yes (ref)
−1.88
−7.18; 3.42
1.67
1.16
Results of the simulation-based sensitivity analysis on the average causal effect of treatment on the treated (ATT) for
the mental score. Unobserved confounders U were simulated to mimic the observed covariates. The estimated
ATTs (90% CI) after adjusting for the different versions of U are reported, as well as the odds ratios for outcome and
selection effects (Γ and Λ) calculated from the simulations. All the reported results arose from the combination of
the results from 5 multiple imputed data sets, according to Rubin [32]. Table 2. Results of the simulation-based sensitivity analysis for mental score. Results of the simulation-based sensitivity analysis on the average causal effect of treatment on the treated (ATT) for
the mental score. Unobserved confounders U were simulated to mimic the observed covariates. The estimated
ATTs (90% CI) after adjusting for the different versions of U are reported, as well as the odds ratios for outcome and
selection effects (Γ and Λ) calculated from the simulations. 3. Results All the reported results arose from the combination of
the results from 5 multiple imputed data sets, according to Rubin [32]. Overall, including among the confounders in the PS matching, the simulated Us also did not
bring to qualitatively different ATT estimates. Only when U mimicked the neuropsychologist indicator,
generating low values of Γ and very large values of Λ, the estimated effects were fully explained away. It is worth noticing that extremely high values of Λ when U mimicked the neuropsychologist indicator
originated from the fact that the probability of being treated for the children evaluated by the second
neuropsychologist ranged, in the imputed data sets, from 1% to 5%. The results obtained in searching for the killer confounder are reported in Figure 2. Unobserved
confounders positively associated with the outcome and negatively associated with the exposure, or
vice versa, could kill the effect if at least one of the odds ratios Λ and Γ was approximately lower than
1:10 or larger than 10:1. However, the plausibility of having omitted a confounder of such strength was
poor in this context, especially considering that many relevant covariates had already been included in
the PS model. 8 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 Table 3. Results of the simulation-based sensitivity analysis for psychomotor score. 3. Results Public Health 2019, 16, x
9 Results of the simulation-based sensitivity analysis on the average causal effect of treatment on the treated (ATT)
for the psychomotor score. Unobserved confounders U were simulated to mimic the observed covariates. The
estimated ATTs (90% CI) after adjusting for the different version of U are reported, as well as the odds ratios for
outcome and selection effects (Γ and Λ), calculated from the simulations. All the reported results arose from the
combination of the results from 5 multiple imputed data sets, according to Rubin [32]. J. Environ. Res. Public Health 2019, 16, x
9 Figure 2. Configurations of the outcome and selection effects (Γ and Λ) driving the causal effect
estimate to zero, as obtained from the simulation-based sensitivity analysis a. Bold points correspond
to unobserved confounders that could kill the effect. a Simulations were performed setting the tuning
parameters 𝑝ଵ, 𝑝ଵଵ, 𝑝ଵ, 𝑝 to 0.1, 0.25, 0.5, 0.75, 0.9. 4. Discussion
Our results suggest that prenatal exposure to average levels of PM2 5 ≥17 µg/m3 had a negative
Figure 2. Configurations of the outcome and selection effects (Γ and Λ) driving the causal effect
estimate to zero, as obtained from the simulation-based sensitivity analysis a. Bold points correspond
to unobserved confounders that could kill the effect. a Simulations were performed setting the tuning
parameters p10, p11, p01, p00 to 0.1, 0.25, 0.5, 0.75, 0.9. Selecting the 50th percentile of Y for the definition of Y* did not lead to substantially
ifferent conclusions. Figure 2. Configurations of the outcome and selection effects (Γ and Λ) driving the causal effect
estimate to zero, as obtained from the simulation-based sensitivity analysis a. Bold points correspond
to unobserved confounders that could kill the effect. a Simulations were performed setting the tuning
parameters 𝑝ଵ, 𝑝ଵଵ, 𝑝ଵ, 𝑝 to 0.1, 0.25, 0.5, 0.75, 0.9. Figure 2. Configurations of the outcome and selection effects (Γ and Λ) driving the causal effect
estimate to zero, as obtained from the simulation-based sensitivity analysis a. Bold points correspond
to unobserved confounders that could kill the effect. a Simulations were performed setting the tuning
parameters p10, p11, p01, p00 to 0.1, 0.25, 0.5, 0.75, 0.9. 4. Discussion
Our results suggest that prenatal exposure to average levels of PM
≥17 µg/m3 had a negative
Selecting the 50th percentile of Y for the definition of Y* did not lead to substantially
different conclusions. 4. 3. Results Observed Confounder Mimicked by U
ATT
90% CI
Outcome
Effect (Γ)
Selection
Effect (Λ)
Gender: male, female (ref)
−2.59
−8.13; 2.95
0.58
1.12
Neuropsychologist: first, second (ref)
−0.88
−9.38; 7.61
0.31
>103
Maternal fruit and vegetable intake:
≤405 g/day (ref), >405 g/day
−2.70
−8.32; 2.91
1.60
1.65
Smoke: yes, no (ref)
−2.33
−7.97; 3.31
2.42
0.63
Smoke at 32 weeks of pregnancy: yes, no (ref)
−2.89
−8.35; 2.57
3.77
1.27
Nursery attendance: yes (ref), no
−2.69
−8.28; 2.89
1.07
0.90
Caregiver: both parents, other (ref)
−2.67
−8.19; 2.84
1.62
0.90
Caregiver: mother, other (ref)
−2.60
−8.22; 3.02
1.03
1.44
Caregiver: relative, other (ref)
−2.88
−8.31; 2.54
0.36
0.83
Maternal education: secondary school or less,
university (ref)
−2.47
−8.19; 3.25
0.93
1.48
Maternal work: non-manual worker (ref),
manual worker
−2.44
−8.14; 3.26
0.71
1.53
Mother’s age: <25, 25+ (ref)
−2.91
−8.28; 2.45
4.30
3.18
Mother’s age: <35 (ref), 35+
−2.70
−8.39; 2.99
0.71
0.57
Body Mass Index: Normal weight, other (ref)
−2.77
−8.2; 2.67
0.84
0.95
Parity: 0 (ref), 1+
−2.69
−8.26; 2.89
1.80
1.23
Breastfeeding: no, yes (ref)
−2.66
−8.16; 2.85
0.76
1.07
Results of the simulation-based sensitivity analysis on the average causal effect of treatment on the treated (ATT)
for the psychomotor score. Unobserved confounders U were simulated to mimic the observed covariates. The
estimated ATTs (90% CI) after adjusting for the different version of U are reported, as well as the odds ratios for
outcome and selection effects (Γ and Λ), calculated from the simulations. All the reported results arose from the
combination of the results from 5 multiple imputed data sets, according to Rubin [32]. nt. J. Environ. Res. Public Health 2019, 16, x
9 o Table 3. Results of the simulation-based sensitivity analysis for psychomotor score. Results of the simulation-based sensitivity analysis on the average causal effect of treatment on the treated (ATT)
for the psychomotor score. Unobserved confounders U were simulated to mimic the observed covariates. The
estimated ATTs (90% CI) after adjusting for the different version of U are reported, as well as the odds ratios for
outcome and selection effects (Γ and Λ), calculated from the simulations. All the reported results arose from the
combination of the results from 5 multiple imputed data sets, according to Rubin [32]. J. Environ. Res. 4. Discussion Our results suggest that prenatal exposure to average levels of PM2.5 ≥17 µg/m3 had a negative
effect on a child’s mental and psychomotor development at around 15 months of age, even if the
confidence intervals of the effects estimates were wide, reflecting large sampling variability. Unobserved confounders moderately associated with both outcome and exposure could kill the
effects in the sense of driving the point estimates of the causal effects to zero. However, the required
associations were stronger than those calculated for most of the observed confounders, making the
omission of such relevant factors quite implausible, in particular if the sensitivity analysis results are
interpreted “in context”, i.e., accounting for number and relevance of the observed covariates which
we already adjusted for [13]. In fact, the sensitivity analysis explores the robustness of the results to
the existence of unmeasured confounders that act through pathways independent of the observed
covariates. Therefore, the same strength of association required to an unobserved confounder to
kill the effect can be indicative of large robustness if the original model accounts for many relevant
covariates, as in the present paper, or of poor robustness if the original model accounts just for few
confounding factors. g
In interpreting the results of the sensitivity analysis, we focused on the point estimates of the
effects and not on the confidence intervals limits as sometimes suggested (searching for possible
unobserved confounders able to explain away the upper or lower limit of the interval, depending on
the sign of effect estimate) [11,12]. This choice was motivated by our will to distinguish between the
evaluation of bias and evaluation of uncertainty. In particular, we think that the sensitivity analysis
should be performed regardless the wideness of the confidence interval and its position in respect to
the null hypothesis, because its aim is to evaluate the possible presence of major bias due to omission
of relevant confounders, which is different from evaluating the “significance” of the result. When the
confidence interval of the estimate is wide as in our analysis, the robustness of the point estimate, if
demonstrated, can confer a certain strength to the conclusions, despite the large sampling variability. As
other kinds of selective reporting, limiting the sensitivity analysis to the subset of “significant” results
may produce bias in the literature, with only studies with narrow confidence intervals evaluated for
their robustness in respect to unobserved confounding. 3. Results Discussion
Our results suggest that prenatal exposure to average levels of PM2 5 ≥17 µg/m3 had a negativ
Selecting the 50th percentile of Y for the definition of Y* did not lead to substantially
different conclusions. Int. J. Environ. Res. Public Health 2019, 16, 4381 9 of 12 4. Discussion This issue is crucial especially in a perspective
where, as desirable, this kind of procedure can become standard in observational studies. Finally,
even if it refers to a specific case study, the sensitivity analysis may provide useful insights about
the confounders to be accounted for or collected in future studies on the same research question,
contributing to enhance the knowledge about the phenomenon of interest. Ichino’s sensitivity analyses rely on the definition of a single binary unobserved confounder
U, and on the assumption of no interaction between the effects of the exposure and U on the
outcome. A sensitivity analysis that relaxes these assumptions has been recently proposed by Ding
and VanderWeele [12,13], but it was not considered in this paper. A second drawback of the simulation
method is its high computational demand. In fact, it requires that the statistical procedure used to
estimate the effect is repeated n times in order to generate a distribution of the effect estimate adjusted
for the hypothetical unobserved confounder. This is particularly expensive when complex analyses
are required, as in our application, where an MI procedure is used. The computational burden of the
simulation approach is particularly evident when the aim is to search for the killer confounder over
the whole space of the hypothetical Us. Our findings qualitatively agree with those obtained for the same cohort by Lertxundi et al. [7]
using a regression approach. In Lertxundi et al. [7], the linearity assumption on the exposure-outcome
relationship, together with the parametric specification of the exposure-confounders-outcome
relationship, contributed to increase the precision of the effect estimates, but, at the same time,
could have generated a certain amount of bias due to problems of inappropriate model specification. On the contrary, the PS matching protected us from possible bias due to nonlinearity or inappropriate
modeling of the exposure-confounders-outcome relationship, but at the prize of a lower precision of
the effect estimates. Anyway, the fact that different statistical methods brought to consistent results
strengthens our conclusions regarding the negative effect of the exposure. Int. J. Environ. Res. Public Health 2019, 16, 4381 10 of 12 10 of 12 Study Limitations Our study has several limitations. First, we limited the analyses to the subset of children for which
all covariates included in the PS model were observed. This affected the power of the analysis and
could have brought a certain amount of bias, although it has been suggested that in several situations
estimating the PS on the complete cases can yield a valid estimate of the causal effect in the subjects
without missing values [33]. Second, exposure during pregnancy was not based on personal exposure, that, if available, could
provide more accurate results. Always in relation with the exposure definition, because of to the
small dimension of the area and the small number of available monitors, we preferred to associate to
each mother–child couple the level of PM2.5 measured by the monitor located in the municipality of
residence instead of using more complex exposure assessment procedure, e.g., kriging interpolation,
to predict the level of the exposure at the residence address. Finally, we treated continuous exposure as a binary one, after having defined an arbitrary
threshold. Focusing on a binary version of the exposure allowed us to apply the sensitivity analysis as
implemented in the sensatt function without complex adaptations, but at the same time led to a certain
degree of arbitrariness related to the choice of the threshold for the exposure, and to loss of information
which resulted in large sampling variability. A future promising extension of the analysis could use
methods based on the generalized propensity score, which, treating the exposure as continuous, allows
for estimating the whole exposure-response relationship [34,35]. References 1. WHO. Health Risks of Air Pollution in Europe—HRAPIE Project: Recommendations for Concentration–Response
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Regional Office for Europe: Copenhagen, Denmark, 2013. 3. Suades-González, E.; Gascon, M.; Guxens, M.; Sunyer, J. Air Pollution and Neuropsychological Development:
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psychomotor development: Six European birth cohorts. Epidemiology 2014, 25, 636–647. [CrossRef] 7. Lertxundi, A.; Baccini, M.; Lertxundi, N.; Fano, E.; Aranbarri, A.; Martínez, M.D.; Ayerdi, M.; Alvarez, J.;
Santa-Marina, L.; Dorronsoro, M.; et al. Exposure to fine particle matter, nitrogen dioxide and benzene during
pregnancy and cognitive and psychomotor developments in children at 15 months of age. Environ. Int. 2015,
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Ibarluzea, J. Prenatal exposure to PM2.5 and NO2 and sex-dependent infant cognitive and motor development. Environ. Res. 2019, 174, 114–121. [CrossRef] 9. Imbens, G.W.; Rubin, D.B. Causal Inference: For Statistics, Social, and Biomedical Sciences: An Introduction,
1st ed.; Cambridge University Press: New York, NY, USA, 2015. [CrossRef] 10. Rothman, K.J.; Greenland, S.; Lash, T.L. Modern Epidemiology, 3rd ed.; Lippincott Williams & Wilkins:
Philadelphia, PA, USA, 2008. 11. Ichino, A.; Mealli, F.; Nannicini, T. From temporary help jobs to permanent employment: What can we learn
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12. Ding, P.; VanderWeele, T.J. Sensitivity Analysis Without Assumptions. Epidemiology 2016, 27, 368–377. 5. Conclusions Although the variability of the estimates was large, our findings indicated that prenatal exposure
to high levels of PM2.5 had a negative effect on a child’s mental and psychomotor development in the
analyzed cohort. The causal effect estimates appeared substantially robust to the possible presence of
unmeasured confounders, even considering that the causal estimates were already adjusted for many
relevant factors. Checking the robustness of the estimates to the presence of unobserved confounders is very
important in observational studies and should not be considered less relevant than quantifying the
sampling uncertainty around the estimates. Especially when new relationships are investigated,
sensitivity analyses aimed to evaluate the robustness of the results to the omission of relevant factors
in the analysis should become a standard. In this paper, we proposed a simulation approach to be
employed after PS matching that, even if computationally expensive, is implemented in standard
statistical software and provides results easy to be interpreted. The use of other methods for sensitivity
analysis, which eventually require less computational effort and rely on less stringent assumptions,
should be explored as well. Supplementary Materials: The following are available online at http://www.mdpi.com/1660-4601/16/22/4381/s1,
Figure S1: Time series of monthly average PM2.5 exposures and monthly average mental and psychomotor scores. The red line indicates the time when the second neuropsychologist substituted the first one; Figure S2: Distribution
of the average prenatal exposure to PM2.5, by the imputed data set; Table S1: Range of the estimated propensity
score among treated and controls, by imputation; Table S2: Between imputation variance, within imputation
variance and their ratio, calculated for the average causal effect of treatment on the treated (ATT) for mental and
psychomotor scores. Author Contributions: M.B. and A.L. designed the study. M.B. designed and supervised the statistical analyses. M.B. and V.T. performed the statistical analyses and wrote the first version of the manuscript. A.L. acquired
data and controlled their quality. M.B., A.L., J.M.I. discussed the results. All the authors critically reviewed and
approved the final version of the manuscript. Funding: This study was partially supported by local research funds (ex 60% funding program University
of Florence 2018), by Basque Government funds (reference: 2009111069) and Spanish Health Ministry funds
(reference: PI06/0867). Conflicts of Interest: The authors declare no conflict of interest. 11 of 12 11 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 References The cost of dichotomising continuous variables. BMJ 2006, 332, 1080. [CrossRef 25. Altman, D.G.; Royston, P. The cost of dichotomising continuous variables. BMJ 2006, 332, 1080. [CrossRef]
26. Rosenbaum, P.R.; Rubin, D.B. The central role of the propensity score in observational studies for causal
effects. Biometrika 1983, 70, 41–55. [CrossRef] 26. Rosenbaum, P.R.; Rubin, D.B. The central role of the propensity score in observational studies for causal
effects. Biometrika 1983, 70, 41–55. [CrossRef] 27. Brookhart, M.A.; Schneeweiss, S.; Rothman, K.J.; Glynn, R.J.; Avorn, J.; Stürmer, T. Variable selection for
propensity score models. Am. J. Epidemiol. 2006, 163, 1149–1156. [CrossRef] 28. D’Agostino, R.B.; Rubin, D.B. Estimating and Using Propensity Scores with Partially Missing Data. J. Am. Stat. Assoc. 2000, 95, 749–759. [CrossRef] 29. Rosenbaum, P.R. The consequences of adjustment for a concomitant variable that has been affected by the
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online: http://prtr.ec.europa.eu/#/home (accessed on 30 January 2018). 31. Becker, S.O.; Ichino, A. Estimation of average treatment effects based on propensity scores. Stata J. 2002, 2,
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of propensity score analysis. Eur. J. Epidemiol. 2019, 34, 23–36. [CrossRef] 34. Hirano, K.; Imbens, G.W. The Propensity Score with Continuous Treatments; Gelman, A., Meng, X.L., Eds.;
Applied Bayesian modeling and causal inference from incomplete-data perspectives; Wiley InterScience:
West Sussex, UK, 2004; pp. 73–84. 34. Hirano, K.; Imbens, G.W. The Propensity Score with Continuous Treatments; Gelman, A., Meng, X.L., Eds.;
Applied Bayesian modeling and causal inference from incomplete-data perspectives; Wiley InterScience:
West Sussex, UK, 2004; pp. 73–84. pp
35. Imai, K.; van Dyk, D.A. Causal inference with general treatment regimens: Generalizing the propensity score. J. Am. Stat. Assoc. 2004, 99, 854–866. [CrossRef] pp
35. Imai, K.; van Dyk, D.A. Causal inference with general treatment regimens: Generalizing the propensity score. J. Am. Stat. Assoc. 2004, 99, 854–866. [CrossRef] © 2019 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access
article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/). © 2019 by the authors. Licensee MDPI, Basel, Switzerland. References f 11. Ichino, A.; Mealli, F.; Nannicini, T. From temporary help jobs to permanent employment: What can we learn
from matching estimators and their sensitivity? J. Appl. Econom. 2008, 23, 305–327. [CrossRef] 12. Ding, P.; VanderWeele, T.J. Sensitivity Analysis Without Assumptions. Epidemiology 2016, 27, 368–377. [CrossRef] 13. VanderWeele, T.J.; Ding, P. Sensitivity Analysis in Observational Research: Introducing the E-Value. Ann. Intern. Med. 2017, 167, 268–274. [CrossRef] 14. Cornfield, J.; Haenszel, W.; Hammond, E.C.; Lilienfeld, A.M.; Shimkin, M.B.; Wynder, E.L. Smoking and
lung cancer: Recent evidence and a discussion of some questions. J. Natl. Cancer Inst. 1959, 22, 173–203. [CrossRef] 15. Rosenbaum, P.R.; Rubin, D.B. Assessing sensitivity to an unobserved binary covariate in an observational
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Everitt, B.S., Howell, D.C., Eds.; John Wiley & Sons, Ltd.: Chichester, UK, 2005; Volume 4, pp. 1809–1814. [CrossRef] 17. VanderWeele, T.J.; Arah, O.A. Bias formulas for sensitivity analysis of unmeasured confounding for general
outcomes, treatments, and confounders. Epidemiology 2011, 22, 42–52. [CrossRef] [PubMed] 17. VanderWeele, T.J.; Arah, O.A. Bias formulas for sensitivity analysis of unmeasured confounding for general
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18. Ding, P.; VanderWeele, T.J. Generalized Cornfield conditions for the risk difference. Biometrika 2014, 101,
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f] 8. Ding, P.; VanderWeele, T.J. Generalized Cornfield conditions for the risk difference. Biometrika 2014,
1–13. [CrossRef] 19. Hernán, M.A. Invited commentary: Hypothetical interventions to define causal effects—Afterthought or
prerequisite? Am. J. Epidemiol. 2005, 162, 618–620. [CrossRef] 20. Nannicini, T. A Simulation-Based Sensitivity Analysis for Matching Estimators. Stata J. 2007, 7, 334–350. [CrossRef] 12 of 12 Int. J. Environ. Res. Public Health 2019, 16, 4381 21. StataCorp. Stata Statistical Software: Release 13; StataCorp LP.: College Station, TX, USA, 2013. 22. Ramón, R.; Ballester, F.; Rebagliato, M.; Ribas, N.; Torrent, M.; Fernández, M.; Sala, M.; Tardón, A.; Marco, A.;
Posada, M.; et al. The Environment and Childhood Research Network (“INMA” network): Study protocol. Rev. Española Salud Publica 2005, 79, 203–220. [CrossRef] 3. Bayley. Bayley Scales of Infant Development; Psychological Corporation: San Antonio, TX, USA, 1977. 24. Li, F.; Baccini, M.; Mealli, F.; Zell, E.R.; Frangakis, C.E.; Rubin, D.B. Multiple imputation by ordered monotone
blocks with application to the anthrax vaccine research program. J. Comput. Graph. Stat. 2014, 23, 877–892. [CrossRef] Altman, D.G.; Royston, P. References This article is an open access
article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/).
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Aspirin has little additional anti‐platelet effect in healthy volunteers receiving prasugrel
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Introduction The use of dual anti-platelet therapy comprising aspirin and a
P2Y12 inhibitor, such as clopidogrel or prasugrel, has become a
standard therapy in patients at risk of thrombotic vascular
events [1–4]. The rationale of this approach is to target the
primary drivers of platelet aggregation: thromboxane A2
(TXA2) and ADP. Aspirin reduces platelet TXA2 production
and P2Y12 receptor blockers prevent binding of the ligand
ADP. The observation that strong P2Y12 inhibition alone can
reduce TXA2-dependent platelet aggregation in both in vitro
and in vivo models [5–8] has raised the question of whether
aspirin provides an additional anti-platelet effect in the
presence of strong P2Y12 receptor blockade [9,10]. In addition
to its anti-platelet effects, aspirin also reduces the formation of
other intravascular prostanoids, such as the vasoprotective
hormone prostacyclin (PGI2) [11]. PGI2 may be important in
controlling platelet responses to pro-aggregatory stimuli and
we have proposed that the administration of aspirin in the
setting of strong P2Y12 receptor blockade removes this
protective factor, while the reduction of TXA2 adds little to
the anti-platelet effect [9]. The production of each prostanoid
within the body can be assessed by measurement of the stable
urinary metabolites: 2,3-dinor-6-keto PGF1a (PGI-M) for
PGI2 and 11-dehydro-TXB2 (TX-M), for TXA2. In the present study, we aimed to investigate, in healthy male
volunteers, whether co-administration of either low or high
doses of aspirin with prasugrel provide any additional bene-
ficial effects on platelet reactivity and intravascular prostanoid
levels, over those seen with prasugrel alone. Correspondence: Tim D. Warner, The William Harvey Research
Institute, Barts & the London School of Medicine & Dentistry, Queen
Mary University of London, Charterhouse Square, London EC1M
6BQ, UK. Tel.: +44 20 7882 2100; fax: +44 20 7882 8251. Tel.: +44 20 7882 2100; fax: +44 20 7882 8251. E-mail: t.d.warner@qmul.ac.uk E-mail: t.d.warner@qmul.ac.uk Aspirin has little additional anti-platelet effect in healthy
volunteers receiving prasugrel P. D. M. LEADBEATER*, N. S. KIRKBY*, S. THOMAS, A.-R. DHANJI, A. T. TUCKER§, G. L. MILNE,
J. A. MITCHELL* and T. D. WARNER *Department of Cardiothoracic Pharmacology, Imperial College London, National Heart and Lung Institute; The William Harvey Research
Institute, Barts and The London School of Medicine & Dentistry, Queen Mary University of London, London, UK; Departments of Pharmacology
and Medicine, Vanderbilt University, Nashville, TN, USA; and §The Ernest D Cooke Clinical Microvascular Unit, St Bartholomews Hospital,
London, UK London, UK To cite this article: Leadbeater PDM, Kirkby NS, Thomas S, Dhanji A-R, Tucker AT, Milne GL, Mitchell JA, Warner TD. Aspirin has little additional
anti-platelet effect in healthy volunteers receiving prasugrel. J Thromb Haemost 2011; 9: 2050–6. Keywords: aspirin, PGI-M, platelets, prasugrel, TX-M. Summary. Background: Strong P2Y12 blockade, as can be
achieved with novel anti-platelet agents such as prasugrel, has
been shown in vitro to inhibit both ADP and thromboxane A2-
mediated pathways of platelet aggregation, calling into question
the need for the concomitant use of aspirin. Objective: The
present study investigated the hypothesis that aspirin provides
little additional anti-aggregatory effect in a group of healthy
volunteers taking prasugrel. Study participants/methods: In all,
9 males, aged 18 to 40 years, enrolled into the 21-day study. Prasugrel was loaded at 60 mg on day 1 and maintained at
10 mg until day 21. At day 8, aspirin 75 mg was introduced and
the dose increased to 300 mg on day 15. On days 0, 7, 14 and 21,
platelet function was assessed by aggregometry, response to
treatments was determined by VerifyNowTM and urine samples
were collected for quantification of prostanoid metabolites. Results: At day 7, aggregation responses to a range of platelet
agonists were reduced and there was only a small further
inhibition of aggregation to TRAP-6, collagen and epinephrine
at days 14 and 21, when aspirin was included with prasugrel. Urinary prostanoid metabolites were unaffected by prasugrel,
and were reduced by the addition of aspirin, independent of
dose. Conclusions: In healthy volunteers, prasugrel produces a
strong anti-aggregatory effect, which is little enhanced by the
addition of aspirin. The addition of aspirin as a dual-therapy
with potent P2Y12 receptor inhibitors warrants further inves-
tigation. 2011 International Society on Thrombosis and Haemostasis DOI: 10.1111/j.1538-7836.2011.04450.x DOI: 10.1111/j.1538-7836.2011.04450.x Journal of Thrombosis and Haemostasis, 9: 2050–2056 Methods Re-use of this article is permitted in accordance with the Terms and
Conditions
set
out
at
http://wileyonlinelibrary.com/onlineopen#
OnlineOpen_Terms 2011 International Society on Thrombosis and Haemostasis Blood collection for platelet aggregation studies Blood was collected by venepuncture into tri-sodium citrate
(3.2%, 1:9 v/v; Sigma, Poole, Dorset, UK). Platelet-rich plasma
(PRP) was obtained by centrifugation at 175 · g for 15 min at
25 C. Platelet-poor plasma (PPP) was obtained by centrifu-
gation of PRP at 15 000 · g for 2 min. All experiments were
completed within 2 h of blood collection. 96-well plate light transmission aggregometry PGI-M (2,3-dinor-6-keto PGF1a) and TXB-M (11-dehydro-
TXB2) were quantified using stable isotope dilution assays with
gas
chromatography/mass
spectrometry,
as
previously
described [13]. To assess the aggregation of platelets in 96-well plates a
modified light transmission method was used. Briefly, 100 lL
samples of PRP were placed into the individual wells of a 96-
well microtiter plate (Nunc, Lutterworth, Leicestershire, UK)
containing 10 lL of vehicle or agonist: ADP (0.1–30 lM;
LabMedics, Salford, Manchester, UK), arachidonic acid (AA;
0.03–1 mM; Sigma), Horm collagen (0.1–30 lg mL)1; Ny-
comed, Linz, Austria), epinephrine (0.001–100 lM; LabMed-
ics), TRAP-6 amide specific for PAR1 (SFLLRN; 0.1–30 lM; VerifyNowTM P2Y12 and aspirin cartridge assays Blood was collected by venepuncture into 2-mL partial fill
vacuum tubes containing tri-sodium citrate (3.2% final
concentration; Griener Bio-One, Stonehouse, UK). Verify-
NowTM assays for effects of P2Y12 receptor blockade and
aspirin activity were performed in accordance with the
manufacturers instructions (VerifyNowTM; Accumetrics, Eli-
tech, UK). A positive response to aspirin was taken as an
aspirin response unit (ARU) score < 550, as described in the
cartridge package insert. Study participants Study participants In all, 9 healthy male volunteers, aged 18–40 years, were
recruited and participated in the present study. The volunteers Received 28 February 2011; accepted 7 July 2011 Leadbeater: prasugrel and aspirin 2051 Bachem, Bubendorf, Switzerland) and the stable TXA2-
mimetic U46619 (0.1–30 lM; Cayman Chemical Company,
Ann Arbor, MI, USA). The plate was then placed into a 96-
well plate reader (Tecan Sunrise, Tecan, Reading, UK) at
37 C, and absorbance was measured at 595 nm every 15 s for
16 min with vigorous shaking between readings. Percentage
aggregation was calculated with reference to the absorbance of
PPP as a surrogate for 100% aggregation. Graphs shown are
for aggregation responses at 16 min. health statuses were determined through their medical histories
and physical examination including blood pressure, pulse rate,
blood chemistry and urinalysis. Volunteers with normal clinical
profiles were included in the study. The study was approved by
the St Thomass Hospital Research Ethics Committee (Ref. 07/
Q0702/24), conducted according to the Declaration of Helsinki
and all volunteers gave written informed consent before
entering the study. Urine collection and storage A fresh mid-stream urine sample was collected into sterile
containers and the preservative chlorhexidine-propyl gallate
(1:1, 20% (wt/v) aqueous chlorhexidine digluconate and 20%
(wt/v) n-propyl gallate in methanol) added at 1:500 parts. Samples were then stored at )40 C for later assay of urinary
prostanoid metabolites. All samples were obtained 3–6 h after
dosing on each study day. Thromboxane B2 assay Before starting this 21-day study, all volunteers had abstained
from aspirin, non-steroid anti-inflammatory drugs (NSAIDs),
paracetamol or any other anti-platelet therapy for 14 days. The
volunteers received a 60-mg loading dose of prasugrel on day 1
of the study and a maintenance dose of 10 mg prasugrel per day
on days 2–21 (Efient, Eli Lilly and Company, Basingstoke,
UK). On day 8 of the study, volunteers began taking 75 mg
aspirin and on day 15 the aspirin dose was increased to 300 mg
per day until day 21 (Angettes 75; Bristol-Myers Squibb,
Uxbridge, UK). Compliance was assessed by interview. Blood
and urine samples were collected on day 0, before commencing
drug treatment, and on days 7, 14 and 21 of treatment. At the end of the PRP aggregation monitoring (i.e. 16 min),
cyclo-oxygenase activity was halted by the addition of 1 mM
diclofenac (Sigma), the samples were centrifuged at 1300 · g
for 10 min at 5 C, and the supernatants removed and frozen. In samples from PRP stimulated with either arachidonic acid
or collagen, plasma thromboxane B2 (TXB2) levels, as a
surrogate for TXA2 production, were determined using a
selective, competitive EIA (Cayman Chemicals) in accordance
with the package insert. In our previous studies we have found
readily detectable levels of TXB2 after platelet activation by
either collagen or AA, but assessments to be less robust for
other agonists [7,8,10,12]. Samples were diluted between 1:10
and 1:1000 in diluent and assayed parallel to known TXB2
standards and a maximum binding control. The percentage
binding of known standards was calculated in reference to the
maximum binding (zero TXB2) control wells, plotted against
the logarithm of concentration and analyzed by non-linear
regression
using
a
four-parameter
logistical
fit
model. Unknown samples were expressed in a similar fashion,
interpolated from this standard curve and corrected for
dilution. All statistical analyzes were performed in Prism v5.0
(GraphPad software, La Jolla, CA, USA). 2011 International Society on Thrombosis and Haemostasis Stastical analysis Although the present study included only nine subjects we are
confident that this is sufficient to detect differences between the 2011 International Society on Thrombosis and Haemostasis 2052 P. D. M. Leadbeater et al 100
A
B
800
600
400
200
80
60
40
P2Y12 reactivity (% inhibition)
Asprin reactivity units
20
0
0
-
-
-
-
-
-
7
14
21
0
7
14
21
Day
Prasugrel
Aspirin
Day
Prasugrel
Aspirin
10 mg o.d. 10 mg o.d. 10 mg o.d. 10 mg o.d. 75 mg o.d. ↑ Aspirin non-responder
↓ Aspirin responder
300 mg o.d. 75 mg o.d. 300 mg o.d. 10 mg o.d. 10 mg o.d. 100
A
B
800
600
400
200
80
60
40
P2Y12 reactivity (% inhibition)
Asprin reactivity units
20
0
0
-
-
-
-
-
-
7
14
21
0
7
14
21
Day
Prasugrel
Aspirin
Day
Prasugrel
Aspirin
10 mg o.d. 10 mg o.d. 10 mg o.d. 10 mg o.d. 75 mg o.d. ↑ Aspirin non-responder
↓ Aspirin responder
300 mg o.d. 75 mg o.d. 300 mg o.d. 10 mg o.d. 10 mg o.d. Fig. 1. VerifyNowTM aspirin and P2Y12 cartridge reactivity. Upper panel,
percent inhibition of P2Y12 cartridge reactivity, normalized to baseline for
values for each of nine study participants. Lower panel, Aspirin Reactivity
Units (ARUs) for each of nine study participants at baseline, and study
days 7, 14 and 21. ARU values < 550 indicate a response to aspirin
(abnormal platelet response to arachidonic acid). In each panel, group
means are indicated by a horizontal line. o.d., once daily. conditions that we have investigated. The variability of platelet
aggregation responses between healthy volunteers in this study
was 12% for AA, 8% for ADP, 5% for collagen, 8% for
TRAP-6 and 6% for U46619 and we propose this low level of
variability allowed changes in platelet reactivity to be detected
readily, as we have found in previous studies [8,12]. Indeed,
power calculations demonstrate that with our sample size
(n = 9) in healthy volunteers and standard deviation (SD) of
response (< 0.45) we had a > 98% power to detect after
analysis of data by two-way ANOVA and Bonferronis post tests
[14]. Similarly, others have detected effects in volunteer groups
of 8–12 subjects [15,16]. A All statistical analyzes were conducted using GraphPad
Prism
v5
(GraphPad). Results Upper panel,
percent inhibition of P2Y12 cartridge reactivity, normalized to baseline for
values for each of nine study participants. Lower panel, Aspirin Reactivity
Units (ARUs) for each of nine study participants at baseline, and study
days 7, 14 and 21. ARU values < 550 indicate a response to aspirin
(abnormal platelet response to arachidonic acid). In each panel, group
means are indicated by a horizontal line. o.d., once daily. were also moderately inhibited by prasugrel alone, and at the
highest concentrations of these agonists there was a further
effect seen with aspirin, although there was no variation
between the effects of the two aspirin doses (Fig. 2, panels C, D
and E). Stastical analysis Agonist
concentration
response
curves were plotted and analyzed according to the four
parameter logistic equation: Y = Bottom + (Top ) Bot-
tom)/[1 + 10^((LogEC50-X) · HillSlope)]. B Concentration-response curves were compared using two-
way ANOVA and Bonferronis post-tests whereas urinary metab-
olite data were compared by one-way ANOVA with Bonferronis
post-tests. In each case statistical significance was taken as
P < 0.05. Other analyzes are stated in text as appropriate. Asprin reactivity units Effects of prasugrel alone and in combination with aspirin on
ex vivo platelet reactivity, in 96-well plate aggregometry Effects of prasugrel alone and in combination with aspirin on
ex vivo platelet reactivity, in 96-well plate aggregometry Platelet aggregation to AA, ADP and U46619 was strongly
inhibited at all concentrations by prasugrel alone, and prasu-
grel in combination with aspirin, although there was no further
reduction with aspirin at either dose (Fig. 2, panels A, B and
F). Responses to collagen, epinephrine and TRAP-6 amide Thromboxane production in response to AA and collagen by
96-well plate aggregometry Prasugrel administration reduced the production of TXA2 in
response to both AA and collagen by platelets in 96-well
aggregometry (Fig. 3A and B, respectively). Co-administration
of either dose of aspirin with prasugrel resulted in the complete
abolition of any remaining TXA2 production. 2011 International Society on Thrombosis and Haemostasis Results Responses to prasugrel alone and in combination with aspirin
in the VerifyNowTM P2Y12 and aspirin cartridge assays Responses to prasugrel alone and in combination with aspirin
in the VerifyNowTM P2Y12 and aspirin cartridge assays To confirm the response of the study subjects to both prasugrel
and aspirin we used the VerifyNowTM cartridge assay system. None of the subjects had positive tests for either P2Y12
inhibitors or aspirin at baseline (Fig. 1). With prasugrel alone,
eight of the nine study participants had > 95% inhibition of
baseline results on the P2Y12 assay, whereas the ninth
participant had only 50% inhibition on this assay, and these
results were maintained with the addition of both low- and
high-dose aspirin (Fig. 1A). Although this subject was an
outlier from the group, based on this assay, all data relating to
this subject were included in further analyzes. It would have
been interesting to test the efficacy of the prasugrel active
metabolite in a PRP sample of this individual to explore
reasons for this difference, but we could not arrange further
blood sampling. After the addition of low-dose aspirin (75 mg),
all subjects were identified as aspirin responders, and these
responses were maintained at the higher dose of aspirin
(300 mg) (Fig. 1B). Aspirin responsiveness was determined as
an aspirin response units (ARU) score of < 550. Fig. 1. VerifyNowTM aspirin and P2Y12 cartridge reactivity. Upper panel,
percent inhibition of P2Y12 cartridge reactivity, normalized to baseline for
values for each of nine study participants. Lower panel, Aspirin Reactivity
Units (ARUs) for each of nine study participants at baseline, and study
days 7, 14 and 21. ARU values < 550 indicate a response to aspirin
(abnormal platelet response to arachidonic acid). In each panel, group
means are indicated by a horizontal line. o.d., once daily. Fig. 1. VerifyNowTM aspirin and P2Y12 cartridge reactivity. Upper panel,
percent inhibition of P2Y12 cartridge reactivity, normalized to baseline for
values for each of nine study participants. Lower panel, Aspirin Reactivity
Units (ARUs) for each of nine study participants at baseline, and study
days 7, 14 and 21. ARU values < 550 indicate a response to aspirin
(abnormal platelet response to arachidonic acid). In each panel, group
means are indicated by a horizontal line. o.d., once daily. Fig. 1. VerifyNowTM aspirin and P2Y12 cartridge reactivity. 2011 International Society on Thrombosis and Haemostasis Production of urinary prostanoid metabolites 100
75
50
25
0
**
*
Day 0
Day 7
Day 21
Day 14
% aggregation
–5
–4
–3
AA (log M)
A 100
75
50
25
0
***
Day 0
Day 7
Day 21
Day 14
% aggregation
–8
–7
–6
–5
–4
ADP (log M)
B B A 100
75
50
25
0
100
75
50
25
0
*
*
**
***
**
1
4
1
4
1
4
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
% aggregation
% aggregation
100
75
50
25
0
% aggregation
U46619 (log M)
–8
–7
–6
–5
–4
ADP (log M)
Epinephrine (log M)
–10
–8
–6
–4
–8
–7
–6
–5
–4
B
D
F *
**
Day 0
Day 7
Day 21
Day 14
100
75
50
25
0
% aggregation
–5
–4
–3
AA (log M)
Collagen (log g mL–1)
–8
–7
–6
–5
–4
C 100
75
50
25
0
*
**
Day 0
Day 7
Day 21
Day 14
% aggregation
ADP (log M)
Epinephrine (log M)
–10
–8
–6
–4
D D C *
**
D
D
D
D
100
75
50
25
0
% aggregation
Collagen (log g mL–1)
TRAP-6 amide (log M)
–8
–7
–6
–5
–4
E E Fig. 2. Platelet aggregation induced by arachidonic acid (0.03–1 mM; panel A), ADP (0.1–30 lM; panel B), collagen (0.1–30 lg mL)1; panel C), epi-
nephrine (0.001–100 lM; panel D), TRAP6 (0.1–30 lM; panel E) and U46619 (0.1–30 lM; panel F) at baseline and on days 7, 14 and 21 (prasugrel,
prasugrel + low aspirin and prasugrel + high aspirin, respectively). Data shown are mean ± standard error of the mean (SEM) of responses measured
by 96-well plate aggregometry in citrated platelet-rich plasma (PRP) prepared from nine different individuals. All data points were analyzed by two-way
ANOVA with Bonferronis post test. *Represents a significant difference, P-value < 0.05, compared with baseline; represents a significant difference, P-
value < 0.05, between prasugrel and prasugrel + low aspirin. Symbols at the end of rows indicate a difference between all points; symbols at individual
points signify particular differences. production of vascular PGI2. Production of urinary prostanoid metabolites To assess the changes in urinary
prostanoid levels we studied the stable urinary metabolites of
thromboxane, 11-dehydro-TXB2 (TX-M), and PGI2, 2,3
dinor-6-keto-PGF1a (PGI-M), at each time point of the study. Neither urinary PGI-M nor TX-M were reduced from
baseline values with prasugrel alone (Fig. 4, panel A and B,
respectively, P > 0.05). When prasugrel and low-dose aspirin
were co-administered, the levels of both urinary prostanoid
metabolites decreased (Day 0 or Day 7 vs. Day 14, PGI-M,
P < 0.01; TX-M, P < 0.001), although there was no further
decrease seen with prasugrel in combination with high-dose
aspirin (Day 14 vs. Day 21, PGI-M, P > 0.05; TX-M,
P > 0.05). production of vascular PGI2. To assess the changes in urinary
prostanoid levels we studied the stable urinary metabolites of
thromboxane, 11-dehydro-TXB2 (TX-M), and PGI2, 2,3
dinor-6-keto-PGF1a (PGI-M), at each time point of the study. Neither urinary PGI-M nor TX-M were reduced from
baseline values with prasugrel alone (Fig. 4, panel A and B,
respectively, P > 0.05). When prasugrel and low-dose aspirin
were co-administered, the levels of both urinary prostanoid
metabolites decreased (Day 0 or Day 7 vs. Day 14, PGI-M,
P < 0.01; TX-M, P < 0.001), although there was no further
decrease seen with prasugrel in combination with high-dose
aspirin (Day 14 vs. Day 21, PGI-M, P > 0.05; TX-M,
P > 0.05). dosing protocol, prasugrel inhibited platelet aggregation to a
range of agonists, with additional effects of aspirin seen only
against the highest concentrations of collagen, epinephrine and
TRAP-6. The urinary levels of TXA2 and PGI2 metabolites
were unaffected by prasugrel alone, but were both reduced by
aspirin. Higher doses of aspirin in combination with prasugrel
did not have any additional effects on either platelet reactivity
or urinary prostanoid metabolites, compared with those seen
with prasugrel and low-dose aspirin. The effects of prasugrel alone, and in combination with
aspirin, were similar to those seen in a recent in vitro study
reported by our group [10], although we observed a slightly
larger effect of aspirin ex vivo. The observations presented here
confirm that P2Y12 receptor blockade by prasugrel strongly
inhibits platelet aggregation to ADP as measured by 96-well
plate aggregometry, although a small residual response was
seen to the highest concentrations of this agonist. In addition,
platelet production of TXA2 and TXA2 (TP) receptor-depen-
dent platelet aggregation was reduced. Production of urinary prostanoid metabolites In contrast to our recent
in vitro findings [10], TP receptor-dependent aggregation was 2011 International Society on Thrombosis and Haemostasis Production of urinary prostanoid metabolites In addition to reducing platelet-derived TXA2 production,
aspirin can, by inhibition of cyclooxygenases, reduce the 2011 International Society on Thrombosis and Haemostasis Leadbeater: prasugrel and aspirin 2053 100
75
50
25
0
100
75
50
25
0
100
75
50
25
0
**
*
*
*
*
**
*
*
**
**
**
**
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
Day 0
Day 7
Day 21
Day 14
% aggregation
100
75
50
25
0
% aggregation
100
75
50
25
0
% aggregation
% aggregation
% aggregation
100
75
50
25
0
% aggregation
–5
–4
–3
AA (log M)
Collagen (log g mL–1)
TRAP-6 amide (log M)
U46619 (log M)
–8
–7
–6
–5
–4
ADP (log M)
Epinephrine (log M)
–8
–10
–8
–6
–4
–7
–6
–5
–4
–8
–7
–6
–5
–4
–8
–7
–6
–5
–4
A
B
C
D
E
F
Fig. 2. Platelet aggregation induced by arachidonic acid (0.03–1 mM; panel A), ADP (0.1–30 lM; panel B), collagen (0.1–30 lg mL)1; panel C), epi-
nephrine (0.001–100 lM; panel D), TRAP6 (0.1–30 lM; panel E) and U46619 (0.1–30 lM; panel F) at baseline and on days 7, 14 and 21 (prasugrel,
prasugrel + low aspirin and prasugrel + high aspirin, respectively). Data shown are mean ± standard error of the mean (SEM) of responses measured
by 96-well plate aggregometry in citrated platelet-rich plasma (PRP) prepared from nine different individuals. All data points were analyzed by two-way
ANOVA with Bonferronis post test. *Represents a significant difference, P-value < 0.05, compared with baseline; represents a significant difference, P-
value < 0.05, between prasugrel and prasugrel + low aspirin. Symbols at the end of rows indicate a difference between all points; symbols at individual
points signify particular differences. Discussion *Shows P < 0.01 difference from Day 0
by one-way ANOVA plus Bonferronis post test. grel + high aspirin, respectively). Data shown are mean ± standard
error of the mean (SEM) of responses collected from 96-well plate
aggregometry in citrated platelet-rich plasma (PRP) prepared from nine
study particpants. *Shows P < 0.05 difference from baseline by two-way
ANOVA plus Bonferronis post test; Shows P < 0.05 difference between
prasugrel and prasugrel + low aspirin. Symbols at the end of lines signify
a difference in set; symbols at individual points signify particular differ-
ences. by the residual potential for TP receptor-induced platelet
aggregation, as described above. not entirely abolished by prasugrel. As TP-mediated platelet
aggregation relies upon the ADP-P2Y12 axis [5,6,10] this may
be explained by the residual responses to ADP observed in the
present study. Alternatively, this may reflect a P2Y12-indepen-
dent response to TP receptor stimulation, although it is unclear
why this may be apparent ex vivo after prasugrel treatment,
although not after in vitro treatment with prasugrel active
metabolite. not entirely abolished by prasugrel. As TP-mediated platelet
aggregation relies upon the ADP-P2Y12 axis [5,6,10] this may
be explained by the residual responses to ADP observed in the
present study. Alternatively, this may reflect a P2Y12-indepen-
dent response to TP receptor stimulation, although it is unclear
why this may be apparent ex vivo after prasugrel treatment,
although not after in vitro treatment with prasugrel active
metabolite. Although the combination of prasugrel and aspirin severely
limits the platelet aggregation response to soluble mediators
(ADP and TXA2) there is still a reasonable aggregation
response to high concentrations of platelet agonists that act
through different signaling pathways. For instance, PAR1
activation by TRAP-6 amide is able to induce approximately
50% aggregation, even when ADP is rendered inactive and
TXA2 formation is inhibited. Additionally, high concentrations
of collagen induced a moderate level of aggregation in the
absence of soluble mediator activities. This is consistent with
strong signaling from the G-protein coupled receptors, a topic
that has been addressed in two informative reviews [17,18], and
from platelet integrins [19]. Measurable residual platelet reac-
tivity is unsurprising given the wealth of clinical data that
reportstheoccurrenceofocclusivethromboticeventsinpatients
taking both aspirin and P2Y12 receptor blockers [20–23]. When aspirin is used in combination with prasugrel there is a
complete inhibition of platelet TXA2 production, although the
further reduction of platelet aggregation, compared with
prasugrel alone, is limited. Discussion In this healthy volunteer study we looked at the effects of a
strong P2Y12 receptor blocker, prasugrel, alone and in
combination with low- and high-dose aspirin, on platelet
reactivity and urinary prostanoid metabolites. After a standard 2054 P. D. M. Leadbeater et al by the residual potential for TP receptor-induced platelet
800
Day 0
Day 7
*
*
*
*
Day 14
Day 21
Day 0
Day 7
Day 14
Day 21
600
400
200
300
200
100
0
0
–5
–8
–7
–6
–5
–4
Arachidonic acid (log M)
Collagen (log g mL–1)
Thromboxane B2
(ng mL–1)
Thromboxane B2
(ng mL–1)
–4
–3
A
B
Fig. 3. Platelet production of TXA2 induced by arachidonic acid (0.03–
1 mM; panel A) and collagen (0.1–30 lg mL)1; panel B) at baseline and
days 7, 14 and 21 (prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Data shown are mean ± standard
error of the mean (SEM) of responses collected from 96-well plate
aggregometry in citrated platelet-rich plasma (PRP) prepared from nine
study particpants. *Shows P < 0.05 difference from baseline by two-way
ANOVA plus Bonferronis post test; Shows P < 0.05 difference between
prasugrel and prasugrel + low aspirin. Symbols at the end of lines signify
a difference in set; symbols at individual points signify particular differ-
ences. 0.20
0.4
0.3
0.2
0.1
0.0
0.15
0.10
0.05
2,3 dinor-6-keto PGF 1α
(ng mg–1 creatinine)
11-dehydro-TXB2
(ng mg–1 creatinine)
0.00
0
7
14
21
0
7
14
21
Day
Prasugrel
Aspirin
Day
Prasugrel
Aspirin
10 mg o.d. 10 mg o.d. 10 mg o.d. 300 mg o.d. 75 mg o.d. -
-
-
10 mg o.d. 10 mg o.d. 10 mg o.d. 300 mg o.d. 75 mg o.d. -
-
-
A
B
*
*
*
*
Fig. 4. Urinary PGI-M (panel A) and TX-M (panel B) production at days
0, 7, 14 and 21 (baseline, prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Values for urinary prostanoid metab-
olites (PGI-M or TX-M) are corrected against urinary creatinine
concentration. Data shown are mean ± standard error of the mean
(SEM) of nine study participants. *Shows P < 0.01 difference from Day 0
by one-way ANOVA plus Bonferronis post test. Discussion -
-
-
A
B
*
*
*
*
Fig. 4. Urinary PGI-M (panel A) and TX-M (panel B) production at days
0, 7, 14 and 21 (baseline, prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Values for urinary prostanoid metab-
olites (PGI-M or TX-M) are corrected against urinary creatinine
concentration. Data shown are mean ± standard error of the mean
(SEM) of nine study participants. *Shows P < 0.01 difference from Day 0
by one-way ANOVA plus Bonferronis post test. 800
Day 0
Day 7
*
*
Day 14
Day 21
600
400
200
0
–5
Arachidonic acid (log M)
Thromboxane B2
(ng mL–1)
–4
–3
A A 5
Arachidonic acid (log M)
4
3 B *
*
Day 0
Day 7
Day 14
Day 21
300
200
100
0
–8
–7
–6
–5
–4
( g
)
Collagen (log g mL–1)
Thromboxane B2
(ng mL–1)
B B Collagen (log g mL–1) Fig. 3. Platelet production of TXA2 induced by arachidonic acid (0.03–
1 mM; panel A) and collagen (0.1–30 lg mL)1; panel B) at baseline and
days 7, 14 and 21 (prasugrel, prasugrel + low aspirin and prasu-
l + hi h
i i
ti
l ) D t
h
± t
d
d Fig. 3. Platelet production of TXA2 induced by arachidonic acid (0.03–
1 mM; panel A) and collagen (0.1–30 lg mL)1; panel B) at baseline and
days 7, 14 and 21 (prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Data shown are mean ± standard
error of the mean (SEM) of responses collected from 96-well plate
aggregometry in citrated platelet-rich plasma (PRP) prepared from nine
study particpants. *Shows P < 0.05 difference from baseline by two-way
ANOVA plus Bonferronis post test; Shows P < 0.05 difference between
prasugrel and prasugrel + low aspirin. Symbols at the end of lines signify
a difference in set; symbols at individual points signify particular differ-
ences. Fig. 4. Urinary PGI-M (panel A) and TX-M (panel B) production at days
0, 7, 14 and 21 (baseline, prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Values for urinary prostanoid metab-
olites (PGI-M or TX-M) are corrected against urinary creatinine
concentration. Data shown are mean ± standard error of the mean
(SEM) of nine study participants. 2011 International Society on Thrombosis and Haemostasis Discussion 0.20
0.4
0.3
0.2
0.1
0.0
0.15
0.10
0.05
2,3 dinor-6-keto PGF 1α
(ng mg–1 creatinine)
11-dehydro-TXB2
(ng mg–1 creatinine)
0.00
0
7
14
21
0
7
14
21
Day
Prasugrel
Aspirin
Day
Prasugrel
Aspirin
10 mg o.d. 10 mg o.d. 10 mg o.d. 300 mg o.d. 75 mg o.d. -
-
-
10 mg o.d. 10 mg o.d. 10 mg o.d. 300 mg o.d. 75 mg o.d. -
-
-
A
B
*
*
*
*
Fig. 4. Urinary PGI-M (panel A) and TX-M (panel B) production at days
0, 7, 14 and 21 (baseline, prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Values for urinary prostanoid metab-
olites (PGI-M or TX-M) are corrected against urinary creatinine
concentration. Data shown are mean ± standard error of the mean
(SEM) of nine study participants. *Shows P < 0.01 difference from Day 0
by one-way ANOVA plus Bonferronis post test. 800
Day 0
Day 7
*
*
*
*
Day 14
Day 21
Day 0
Day 7
Day 14
Day 21
600
400
200
300
200
100
0
0
–5
–8
–7
–6
–5
–4
Arachidonic acid (log M)
Collagen (log g mL–1)
Thromboxane B2
(ng mL–1)
Thromboxane B2
(ng mL–1)
–4
–3
A
B
Fig. 3. Platelet production of TXA2 induced by arachidonic acid (0.03–
1 mM; panel A) and collagen (0.1–30 lg mL)1; panel B) at baseline and
days 7, 14 and 21 (prasugrel, prasugrel + low aspirin and prasu-
grel + high aspirin, respectively). Data shown are mean ± standard
error of the mean (SEM) of responses collected from 96-well plate
aggregometry in citrated platelet-rich plasma (PRP) prepared from nine
study particpants. *Shows P < 0.05 difference from baseline by two-way
ANOVA plus Bonferronis post test; Shows P < 0.05 difference between
prasugrel and prasugrel + low aspirin. Symbols at the end of lines signify
a difference in set; symbols at individual points signify particular differ-
ences. 0.20
0.4
0.3
0.2
0.1
0.0
0.15
0.10
0.05
2,3 dinor-6-keto PGF 1α
(ng mg–1 creatinine)
11-dehydro-TXB2
(ng mg–1 creatinine)
0.00
0
7
14
21
0
7
14
21
Day
Prasugrel
Aspirin
Day
Prasugrel
Aspirin
10 mg o.d. 10 mg o.d. 10 mg o.d. 300 mg o.d. 75 mg o.d. -
-
-
10 mg o.d. 10 mg o.d. 10 mg o.d. 300 mg o.d. 75 mg o.d. Discussion For instance, the abolition of
TXA2 in collagen-induced aggregation results in a further
reduction of platelet aggregation only at high concentrations of
this agonist. Aspirin also has a small additional effect on the
platelet aggregation induced by strong PAR1 activation (by
TRAP-6 amide; SFLLRN) and the highest concentrations of
epinephrine. In these circumstances, it is likely that the effect of
TXA2 (that is lost by the addition of aspirin) is partly mediated The predominant choice of which aspirin dose to use in dual
anti-platelet therapy varies with geographic location [24] and so Leadbeater: prasugrel and aspirin 2 2055 it was useful to include both a low- and a high dose of aspirin in
combination with prasugrel in the present study. Consistent
with meta-analysis of aspirin as a monotherapy, which show
that the efficacy of aspirin is not enhanced at higher doses [1],
there was no evidence that the higher dose of aspirin produced
any further anti-platelet effects than a strong P2Y12 inhibitor
together with low-dose aspirin. The present study did not
determine the effects of aspirin alone on platelet aggregation
using the 96-well plate assay method, although previous work
by our group has demonstrated that this assay readily detects
the effects of aspirin in healthy volunteers [12]. contribute to the efficacy of drugs targeting the P2Y12 receptor
pathway. Clearly, this can only occur if the production of
vascular prostacyclin is preserved, whereas the co-administra-
tion of aspirin with P2Y12 receptor blockers results in reduced
PGI2 formation and a potential limitation of the anti-throm-
botic effect. Furthermore, loss of vascular PGI2 has been
proposed as the underlying mechanism of cardiovascular risk
that is associated with NSAIDs based on data derived from
urinary PGI-M measurements [29]. A limitation of the present study was the continuous
administration of treatments without any wash-out periods
to differentiate the effects of each drug combination from a
potential effect of time. An alternative study design could have
also included a treatment period of aspirin alone. Clearly, such
a study would have been a much larger undertaking and would
have provided further levels of information. However, it is our
belief that for the mechanisms of drug action that we wished to
explore the study design chosen was appropriate. In addition to platelet reactivity, we measured the urinary
metabolites of both TXA2 and PGI2, which are considered a
marker of in vivo prostanoid levels [11]. Discussion Urinary metabolites
were measured within 6 h of dosing on the relevant study day,
which is within the time period in which the effects of aspirin
are maintained in healthy subjects taking consecutive daily
doses of this drug [11,25]. In healthy volunteers PGI-M and
TX-M were unaffected by prasugrel alone, although these
results contrast with the effects of clopidogrel, which we have
previously reported in a similar study [8]. Urinary TX-M
formation was reduced after 7 days of clopidogrel dosing in
healthy volunteers, although in this case the samples were
measured by enzyme immunoassay rather than mass spectros-
copy [8] and we have greater confidence in the data we present
here; although the small number of individuals included in each
study makes a definitive analysis difficult, especially because of
the large inter-individual variability in urinary PGI-M and TX-
M levels. The failure of prasugrel to reduce TX-M excretion
seems contradictory to the reduction of ex vivo TXA2
formation that is associated with clopidogrel [8,26] and now
prasugrel; however, TX-M is used as a marker of physiological
TXA2 levels whereas ex vivo TXA2 formation describes the
production of this lipid mediator in response to strong
thrombotic stimuli, which are absent in young healthy volun-
teers. In patients with vascular inflammation [13] and athero-
sclerosis [27], urinary TXA2 formation is increased and the
effects of prasugrel on urinary TX-M may be different in these
pathologic states where in vivo platelet activation may contrib-
ute more to urinary TX-M levels. In conclusion, the present study indicates that strong P2Y12
receptor blockade produces an inhibition of platelets that is
only slightly enhanced by aspirin, as assessed ex vivo. Aspirin
also reduces the levels of PGI2 in vivo, an effect that is suggested
to increase the risk of thrombosis. As no clinical trial has tested
the effects of aspirin in the presence of strong P2Y12 receptor
blockade it is unclear whether or not aspirin enhances or limits
the anti-thrombotic effect that can be achieved with drugs such
as prasugrel. We would therefore suggest that dedicated clinical
trials should be considered to ascertain the true benefits, and
rates of adverse events, when using aspirin alongside an
effective, strong P2Y12 receptor blocker. Acknowledgements This research was supported by the William Harvey Research
Foundation, the National Heart and Lung Institute Founda-
tion and the Wellcome Trust (085255/Z/08/Z). This work
forms part of the research themes contributing to the
translational research portfolio of Barts and the London
Cardiovascular Biomedical Research Unit, which is supported
and funded by the National Institute of Health Research. In the present study, both prostanoids were reduced,
independently of dose, when aspirin was co-administered. It
has been previously suggested in a similarly sized study that the
reduction of PGI-M by aspirin is dose dependent, although
when scrutinizing these earlier data we found a large overlap
between the values for groups taking 75 and 325 mg aspirin
similar to the results we report here [11]. It is unclear from our
data but it is possible that in larger studies we may have seen a
dose-dependent relationship between aspirin and urinary
PGI-M. Disclosure of Conflict of Interest The authors state that they have no conflict of interest. 2011 International Society on Thrombosis and Haemostasis 2 Saw J, Madsen EH, Chan S, Maurer-Spurej E. The ELAPSE (Eval-
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JA, Warner TD. Utility of 96-well plate aggregometry and measure-
ment of thrombi adhesion to determine aspirin and clopidogrel effec-
tiveness. Thromb Haemost 2009; 102: 772–8. 27 Belton O, Byrne D, Kearney D, Leahy A, Fitzgerald DJ. Cyclooxy-
genase-1 and -2-dependent prostacyclin formation in patients with
atherosclerosis. Circulation 2000; 102: 840–5. 13 McAdam BF, Byrne D, Morrow JD, Oates JA. Contribution of
cyclooxygenase-2 to elevated biosynthesis of thromboxane A2 and
prostacyclin in cigarette smokers. Circulation 2005; 112: 1024–9. 28 Cattaneo M, Lecchi A. Inhibition of the platelet P2Y12 receptor for
adenosine diphosphate potentiates the antiplatelet effect of prostacy-
clin. J Thromb Haemost 2007; 5: 577–82. 14 Lenth RV. Statistical power calculations. J Anim Sci 2007; 85: E24–9. 15 Madsen EH, Schmidt EB, Gehr N, Johannesen NL, Kristensen SR. Testing aspirin resistance using the Platelet Function Analyzer-100:
some methodological caveats and considerations. J Thromb Haemost
2008; 6: 386–8. 29 Grosser T, Fries S, FitzGerald GA. Biological basis for the cardio-
vascular consequences of COX-2 inhibition: therapeutic challenges and
opportunities. J Clin Invest 2006; 116: 4–15. 16 Madsen EH, Schmidt EB, Maurer-Spurej E, Kristensen SR. Effects of
aspirin and clopidogrel in healthy men measured by platelet aggrega-
tion and PFA-100. Platelets 2008; 19: 335–41. 2011 International Society on Thrombosis and Haemostasis
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The Future and Journey of Social Media Towards Virtual World
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The Future and Journey of Social Media Towards
Virtual World Raju Sitaram Nandkar, BSc Agri., MBA(BII) , MA(RD), ISTD
Indira Gandi National Open University ,New Delhi Raju Sitaram Nandkar, BSc Agri., MBA(BII) , MA(RD), ISTD
Indira Gandi National Open University ,New Delhi Abstract:- Social media has covered and permeated the
entire human life and also existence of human beings. Sociability is a basic human characteristic and gossiping
is an integral part of social dialogue. Social Media is a
highly developed form of social gossiping. In one click we
can interacts with any person from any country in the
world through social media. Friendship and interaction
with many people at a time is an important feature of
social media. Social media allows sharing of feelings. Ideas and information is exchanged through social media. Social contact continues to growing day by day. There are
lots of examples people help each other through Social
contact. Despite these many benefits, social media cannot
demonstrate spatial presence. Although social media
enhances interaction, it represents a virtual existence. Obviously, virtual world issues also come into focus on
social media. Hence, an anonymous fear seems to be
forming that social media is not moving towards the
virtual world. Many problems arise in the virtual world,
whether it is family or social, its consequences can be felt
by the entire human race. The virtual world will remain
a superficial and imperfect world. The virtual world is not
perfect and how to deal with it is a huge challenge. Good
and bad effects of social media also affect your emotions. Social media creates problems with properly controlling
and channeling emotions because it creates emotional
hanging. Mental pressure builds up as emotions go up and
down. Mentally, the person also becomes depressed. The
use of social media reduces efficiency. As the time that
should be spent on productive work is not available, it has
a far-reaching impact on overall growth. Due to social
media there is a huge increase in cyber crime. Overall,
people are still experimenting in the virtual world of social
media and facing lot of issues in adjustment. In this article
we try to find out in which direction this journey of the
virtual world going? What are its possible side effects? And information about how all mankind will find a way
out of it has been made available. revolutionized human communication. Volume 8, Issue 1, January – 2023 Volume 8, Issue 1, January – 2023 International Journal of Innovative Science and Research Technology
ISSN No:-2456-2165 II. Social media is now spread everywhere in the world. One who said ‘no one can survive without social media. It is
the fastest medium of communication. Social media
encompasses every sphere and corner of mankind. The first
recognizable social media site Six Degrees was created in
1997. It helps users to upload a profile and make friends with
other users. The first blogging sites became popular in 1999,
creating a social media sensation. After the invention of
blogging, social media began in popularity. Sites like The Future and Journey of Social Media Towards
Virtual World Script is a distinctive
writing system, based on a repertoire of specific elements or
symbols, or that repertoire. Language is the system of
communication in speech and writing that is used by people
of a particular country. There is two types of communication
one is verbal and another is non verbal. In ancient period we
can found some of the oldest forms of human communication
include making sounds, drawing or painting, dancing, acting,
and using symbols. The journey of communication includes
Cave Paintings, Smoke Signals, Carrier Pigeons, Postal
System,
Newspapers,
Radios,
Telegraph, Telephone,
Television, Internet, Email, Text Message and Social media. The Internet has made evolution of communication more
effective. We can send messages with just one click. Computers, mobile phones, laptops, radios, etc, all help us to
communicate. The latest mode of communication in the
digital world is the social media. People share their entire life
events on social media. Social media platforms help people
share pictures, videos, and almost everything on the internet. Social media has reduced the geographical boundaries and
distances between two people and the time gap has been
reduced to a fraction of a second. All in all, social media has
captured the entire human life. Just as the force of gravity
works, the social media worked to connect the bonds of
contact and communication between two or more people. Social media is emerging as a new innovation in human
communication. However question marks have been always
raised about this. Fake accounts, cyber crime, cheating,
pornography, blackmailing, defamation etc. now became
common type fraud on social media. Therefore all users are
in pressure however the cant able to keep away from social
media because there are lot of benefits they are enjoying from
social media platform. However there is urgent need to
balance the use of social media in our daily life and need to
avoid virtual world in life. In this article, we will try to explain
the future of social media and the journey towards the virtual
world. We also try to shed light on the emergence of the social
media and its use and its future. II.
LITERATURE REVIEW Keywords:-
Social,
Virtual,
Media,
Communication,
Interaction. Keywords:-
Social,
Virtual,
Media,
Communication,
Interaction. I.
INTRODUCTION Food, clothing, and shelter are the basic human needs. Now, social media has become the fourth basic need. Some
day before, some important applications in social media were
shut down and life seemed to come to a standstill. Communication has been a natural gift from immemorial
times. The discovery of scripts and languages has www.ijisrt.com IJISRT23JAN498 604 Volume 8, Issue 1, January – 2023 International Journal of Innovative Science and Research Technology
ISSN No:-2456-2165 ISSN No:-2456-2165 MySpace and LinkedIn gained prominence in the early
2000s, and sites like Photobucket and Flickr facilitated online
photo sharing. YouTube came out in 2005, creating an
entirely new way for people to communicate and share with
each other across great distances. In 2006, Facebook and
Twitter both became available to users throughout the world. These sites remain some of the most popular social networks
on the Internet. Other sites like Tumblr, Spotify, Foursquare
and Pinterest began popping up to fill specific social
networking niches. Today, there is a tremendous variety of
social networking sites, and many of them can be linked to
allow cross-posting. This creates an environment where users
can reach the maximum number of people without sacrificing
the intimacy of person-to-person communication. and a considerable space for the virtual interactions of
intellectual and popular society categories; through the
worldwide known media and interactive gates. Gogolin, et al.(2014) critically discussed about the use
of virtual worlds and social media has grown to the point that
more than one-quarter of the world's population utilize it in
some manner. Security and privacy concerns regarding the
use and capabilities of current and emerging technologies
such as gaming, blogging, podcasting, virtual meetings,
virtual worlds and Web 3.0 is examined. Security and privacy
concerns are investigated in the context of exploits,
vulnerabilities, and related security risks; confidential access
control; communication trends and patterns in the use of
massive communication strategies; intellectual property and
product risk management; resource management; financial
considerations and accountability; and safety. Several
technologies and personal practices are reviewed, as well as
ways to mitigate or eliminate their associated risks. The core
principles of information security-confidentiality, integrity,
and availability-provide an overall framework. Mohammed et al., 2021; observed in research that the
social media is becoming a critical part of everyone’s life. Social media has numerous platforms including Fecebook,
Twitter, Instagram, and LinkedIn. I.
INTRODUCTION Impersonation is a
common phenomenon found nearly on all social media
platforms; it is the act of attempting to deceive someone by
pretending that he is another person. Impersonators always
try to hide a real account by making another similar profile to
spread the fake contents on social media platforms making it
very difficult to know the real accounts from the fake ones. The article explains on the social media impersonation,
impersonation types, how to identify the social media
impersonation, cases of social media impersonation, how to
prevent impersonation, and how to protect the security of a
social media user. Social media impersonation is the act of
pretending that a person is another person which usually
occurs on all social media platforms. Eraslan, et al., (2019), illustrated about the living in the
age of constant technology developments shifted social
communication patterns and shifted social relations to virtual
environments. The socialization process that takes place in
digital platforms also transferred many negative elements
experienced in social life to the virtual environment. That is,
the aggression behaviours concerning these negative
processes have also been transferred to the virtual
communication. The current study examines the effects of
social media aggression (SMA) regarding digital platforms
on the social relations in human life in the context of various
variables. Results of the study revealed that the counter-
comments towards participants’ values have a significant
effect
on
participants’ demonstration
of
aggressive
tendencies. Messinger,et al.,(2009), discussed about the virtual
worlds,
where
thousands
of
people
can
interact
simultaneously within the same environment, represent a
frontier in social computing with critical implications for
business, education, social sciences, and our society at large. In this paper, they first trace the history of virtual worlds back
to its antecedents in electronic gaming and on-line social
networking. They provide an overview of extant virtual
worlds,
including
education-focused,
theme-based,
community-specific, children-focused, and self-determined
worlds and they analyze the relationship among these worlds
according to an initial taxonomy for the area. Papp, Raymond;(2010), discussed thoroughly about
virtual
worlds
and
social
networking. As
online
communication
and
collaboration
becomes
more
commonplace, universities are exploring the educational
possibilities of online virtual environments for reaching the
Millennial. III.
DISCUSSION, ANALYSIS AND
INTERPRETATION Three million years ago, a truly wise man started
roaming the earth. For many years, everything went
smoothly, such as living in the same territory, roaming for
food, and reproducing. Gradually, as man began to
comprehend, analyze, and reflect on the information he
received, he became involved in the process of evolution. Such an evolution had begun not only on a physical but also
on a mental level. As a result, he had to adapt to the
environment in which he lived, as his body adapted. However, these changes were taking place at a very subtle
and very slow pace. On the other hand, mental changes were
advancing at a slower pace, and so human evolution was
advancing faster than other animals. In the past, we used to express these feelings on
occasion, but now they are completely broken. Therefore,
emotions and thoughts are harmonized. Everyone is
dependent on this social media. The world is constantly
changing and science and technology are accelerating this
change. As the changes took place, we became more and
more aware of that change. But the big question for
researchers is whether the current change has brought about
mental and physical harmony. Among them, it will be
necessary to see whether the massive use of social media and
the mental and physical changes it requires are taking place. Mobile and its electromagnetic waves cover the whole world. The mobile revolution happened, but with it came the social
media revolution. Social media has shocked many
superpowers globally and social media has done a disservice
to the family. This pace of evolution began with the discovery of fire,
increased with the discovery of the wheel, speeded up due
with discovery of agriculture, and the discovery of script and
language gained real momentum for human evolution and
development. Printing, Radio, Internet, and Mobile Phones
have brought the world closer together. Social media brought
man closer and the geographical distance of man-to-man
became zero. Evolutionary changes are very slow, while revolutions
happen in a few years and suddenly. In simple terms,
evolution means change and getting used to that change,
while revolution is sudden change. Whether social media is
evolution or revolution? When changes happen and those
changes are not accustomed, it has physical and mental
trauma. As the man sat up and started working, he started
getting back and neck ailments. I.
INTRODUCTION Both virtual worlds and social networking
constitute a large part of the Millennial' time and
incorporating these technologies into the classroom can foster
a more collaborative and diverse learning atmosphere.Virtual
campus tours, recruiting, advising, simulations and classes
are all part of the growing virtual environment. Corporate and
educational institutions are exploring their use in cutting
costs, delivering higher customer satisfaction and catering to
a more tech-savvy clientele. Madouni, et al., (2020) elaborated about the virtual
media emerged and disseminated immensely ; specifically in
the last ten years of the twenty first Century, through
innumerable channels and virtually broadcasting pages, as
strong equivalently as the traditional mass media in almost
life critical domains and areas ; as a result and feature of the
technological progress. The technology of the twenty-first
Century gave to hands a wide reach and availability of
information, it allows people and communities to participate
even in producing and making influential public opinions
towards local and international issues and topical; as ways of
social interaction behind devices screens. Technology and
changes create a sort of circumstantial adaptation which did
not exist before. Furthermore and notably, the traditional
mass media amid this advance; they specify regular corners Pradeep,et al.,(2016), shade light on the Social
Networking Sites (SNSs) such as Fecebook have made
inroads in the life of users. The current study used a
quantitative approach to explore adolescents usage of SNS in
Mumbai and their perceptions and experiences of the same. Easily accessible Internet and availability of devices such as
smart phones influenced access to SNS. Computer-mediated-
communication was the preferred means of communication. www.ijisrt.com IJISRT23JAN498 605 Volume 8, Issue 1, January – 2023 International Journal of Innovative Science and Research Technology
ISSN No:-2456-2165 ISSN No:-2456-2165 Selective self-representation and online social comparison
were pertinent themes seen through the study. Gender and age
differences were found in the overall usage and experiences
of SNS. Although social media is not as physically and mentally
accustomed as it is, the attraction of the information available
and the interaction and communication that takes place is so
great that man is drawn to it like a force of gravity. This power
does not allow him to sit still and think. As a result,
information, data, language, scripts, videos, audio and
pictures are simply exploding. I.
INTRODUCTION How to store so much
information is a challenge in front of the brain, so it
immediately began to put the information in the dustbin. There is so much information that it takes a lot of time and
energy for the brain to grasp and analyze it, so the brain may
be stopped grasping and analyzing. It is certainly not
desirable for a wise person to accept that the brain is moving
in the right direction. Naturally, social media will have far-
reaching effects on human comprehension and analysis. Naturally, the connection between the brain and the mind is
disrupted. The confusion between what the mind tells the
brain and what the brain tells the mind has created confusion
and roughness in the emotions that arise from the mind. Therefore, there is an upheaval in happiness, sorrow, anger,
fear, hatred, surprise, love. III.
DISCUSSION, ANALYSIS AND
INTERPRETATION As the man gained weight, he
began to develop diabetes and high blood pressure. Of course,
social media is a part of human evolution or revolution is a
subject of research, but the radical changes brought about by
social media did not take many years to get used to physically
and mentally, and suddenly we started riding on social media. Eventually, many of its good and bad consequences are
beginning to emerge. The above mentioned changes, which
do not get used to the physical and mental, certainly cause
physical and mental disorders. No one should be upset that
social media has increased the physical complaints of man. Eye strain, strain on the cervical spine, the effect of mobile
light on the skin of the face is some of the complaints that
need to be mentioned deliberately. No one should be offended
by the fact that mental problems are exacerbated by the
slowness of thought processes, constant distractions due to
incoming messages, persistence of thought processes, grief
and frustration, fear and irritability by mobile and social
media as a whole. Everyone has a situation where they
understand but do not turn around. We are all connected to the virtual relationship and the
virtual world, rather than personal and interpersonal
relationships. In a virtual relationship, even though the
connection of contacts and communication is fast, they
cannot create intimacy. As we aware eyes contact and touches
and actual emotional and mental support play an important
role in creating intimacy. Virtual relationships seem to lack
this. Although social media has accelerated communication
and interaction, the disadvantages outweigh the benefits. The
benefits of social media are obvious at the business level and
at the administrative level; however social media has not been
able to work beyond virtual communication at the individual
level. Social media has also contributed to the decline in
human efficiency and effectiveness. There is a fear that we
will lose creativity and innovation. Although social media is
attractive, it is not conducive to long-term human
development. The time that you have is so precious that it is
spent in the virtual world and just listening to, viewing and
reading information so no one else has time left to be for
constructive work. III.
DISCUSSION, ANALYSIS AND
INTERPRETATION There are also some gentlemen who are IJISRT23JAN498 IJISRT23JAN498 www.ijisrt.com 606 Volume 8, Issue 1, January – 2023 International Journal of Innovative Science and Research Technology
ISSN No:-2456-2165 spending more than ten hours out of twenty four hours on
mobile and social media. The power of human capital as a
whole is weakening and diminishing. As a result, its problems
are becoming more prevalent in the business, social spheres
and at the family level. Going even further, some organized
cybercriminals are on the rise, increasing the number of
people being deceived in various ways. Fake accounts, cyber
crime, cheating, pornography, blackmailing, defamation etc. now became common type fraud on social media. Instead of
that the Fake friends, Free app downloads, Quizzes ,Hidden
URLS, Lottery and Free Gift Card Scam, Gossip Scam,
Healthcare Scam, Cat fishing, Photo of You Scam, Account
Cancelled Scam, Nigerian Scam, Stuck Abroad Scam, IQ
Scam, See Who Viewed Your Profile Scam etc type of frauds
now became very common on social media. permanent. Emotions cannot be controlled and flowed
properly in virtual world because the reality is different. Taking all these factors and things for granted, there is a
global need for a comprehensive effort to stop the society
polarization towards a virtual world. We need to realize that
social media is a necessity not a must. Anything in this world
is difficult but not impossible. REFERENCES [1]. Eraslan, Levent & Kukuoğlu, Ahmet. (2019). Social
relations in virtual world and social media aggression. World Journal on Educational Technology: Current
Issues. 11. 140-149. 10.18844/wjet.v11i2.4145. [2]. Gogolin, Greg & Gogolin, Erin & Kam, Hwee-Joo. (2014). Virtual Worlds and Social Media. International
Journal of Artificial Life Research. 4. 30-42. 10.4018/ijalr.2014010103. Man is such an intelligent creature on earth that he is
always looking for reuse from waste. Naturally, even though
the world is occupied by social media, a lot of changes are
expected in the future. Also, the use of social media needs to
be as efficient and effective as it needs to be. Otherwise the
possibility of human beings running away from this social
media cannot be ruled out. There is nothing that can defeat a
human being, deprive him, because at the same time the
human race has many options in front of it and it is capable
of choosing the right option at the right time. Although it is
not possible to say how long social media will control and
cover to human being, many more good and bad effects of
social media are yet to come. Naturally, the four generations
that exist on the surface of the earth need to use social media
in a timely, limitedly, efficiently, effectively and to make
human life simpler, easier, straightforward, convenient and
enjoyable. [3]. Messinger, Paul & Stroulia, Eleni & Lyons, Kelly &
Bone, Michael & Niu, Run & Smirnov, Kristen &
Perelgut, Stephen. (2009). Virtual worlds past, present,
and future: New directions in social computing. Decision
Support
Systems. 47. 10.1016/j.dss.2009.02.014. [4]. Mohammed, Gharawi & Badawy, Ahmed & Ramadan,
Doaa & Elsayed, Shaymaa. (2021). SOCIAL MEDIA
IMPERSONATION IN THE VIRTUAL WORLD. Al
Hikmah International Journal of Islamic Studies and
Human Sciences. [5]. Papp, Raymond. (2010). Virtual worlds and social
networking: Reaching the millennials. Journal of
Technology Research. [6]. Pradeep, Pooja & Sriram, Sujata. (2016). The Virtual
World of Social Networking Sites: Adolescents Use and
Experiences. Psychology & Developing Societies. 28. 10.1177/0971333615622911. IV. The changes are inevitable we can’t deny it. The social
media is one of the major changes in our daily life. It is now
became a part and parcel of our routine life. No one can avoid
it and no one can deny it. Social media has touched every
aspect of human life and made it easy and comfortable. Smart
phones and social media are the inventions of science and
technology. Social media has definitely revolutionized for
human life. Social media works on the three goals that are
communication, relationship and interaction. Social media
brings two or more people together so the exchange of
feelings and thoughts takes place at a faster pace. Useful
information is also available quickly through social media. Social media has permeated and covered the entire life
of human being although it has also been created some
serious problems for mankind. This will include unnecessary
information bombardment, spreading false and misleading
information, inflammatory emotions, wastage of time, and
absenteeism from work and cyber crime, family conflict. These all problems are the due to virtual feature of social
media. The entry and spread of this virtual world is a
challenge created by social media for the entire world. Since
the virtual world is mostly superficial, nothing substantial
happens in it. The virtual world is shallow in nature. Relationships in this virtual world are not that much 607 www.ijisrt.com www.ijisrt.com IJISRT23JAN498
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W4239069418.txt
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de
|
Geistige Stromungen der Gegenwart.
|
The Philosophical review
| 1,909
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public-domain
| 148,483
|
%
BOOK 190.EU22G 1913 c.
EUCKEN # GEISTIGE STROEMUNGEh
1
DERGEGENWART
3
T153 000t,ME73
fl
(B
Geistige Strömungen
Geistige Strömungen ^^-
DER Gegenwart
VON
RUDOLF EUCKEN
DER GRUNDBEGRIFFE DER GEGENWART
VIERTE UMGEARBEITETE AUFLAGE
Neue Ausgabe
VERLAG VON VEIT&COMP. IN LEIPZIG
1913
Druck von Fr. Richter, O. m. b. H., Leipzig.
Vorwort zur dritten Auflage
-----
Die dritte Auflage ist gegen die zweite noch mehr verändert
S
als diese
C:
legung den Grundstock, den die sachliche Erörterung nur um-
gegen die erste; bildete bei ihr die geschichtliche Dar-
säumte, so
diese in der zweiten Auflage weit selbständiger
ist
geworden und hat in der dritten die volle Herrschaft erlangt;
das Buch
nun vor allem ein Ausdruck einer eigentümlichen
ist
ß
philosophischen Gesamtüberzeugung und will
4
würdigt sein.
j
als solcher
ge-
Das mußte auch die Darstellung wesentlich ver-
ändern, das verlangte namentlich eine präzisere Anordnung und
Einteilung
des Stoffes
bis
in
die einzelnen Abschnitte hinein.
j
Den Grundgedanken der früheren Behandlungen: die Ver-
<5^
knüpfung von Historischem und Sachlichem einerseits, die Zer-
-V legung in einzelne Abschnitte andererseits glaubte ich dabei
festhalten zu können.
Daß das Geschichtliche mir mehr ist
i
als
ein
Gegenstand gelehrter Beschäftigung, daß
es,
freilich
f^ unter bestimmten Voraussetzungen, zur Erhöhung der eignen
-J
Arbeit kräftig beizutragen vermag, dafür kämpft das Buch ebenso
als Ganzes
wie
in
besonderen Erörterungen, die sich hier nicht
vorwegnehmen lassen.
Das Ausgehen von einzelnen Problemen
aber gewährte den Vorteil greifbarer Angriffspunkte, von denen
l/l
sich
rasch
Allerdings
y
-^
zu
verblieb
Überzeugung
sich
in
irgendwelcher Entscheidung
dabei
der Mißstand,
vordringen
ließ.
daß das Ganze der
nicht als solches volle Rechenschaft
geben und
einem fortlaufenden Zusammenhange darlegen kann.
Vorwort
VI
Dieser Mangel
sei
bereitwillig zugestanden,
dieser Behandlungsweise verbunden, als
helfen
ist
zu eng mit
daß sich ihm hier ab-
Gewisse Ergänzungen bieten
ließe.
er
dieser Hinsicht
in
meine früheren Bücher, die größte Lücke liegt in dem Mangel
genügenden
einer
erkenntnistheoretischen
Fundamentierung,
mein nächstes Buch wird einer prinzipiellen Erörterung dieses
Problems gewidmet sein.
Mehr noch
die Art
als
der Behandlung aber ist es eine
durchgehende Grundüberzeugung,
welche
verschiedenen
die
Auflagen zusammenhält, die Überzeugung von der Unsicherheit
des Bodens,
dem
auf
unser ganzes Kulturleben und mit ihm
auch unsere wissenschaftliche Arbeit
steht,
die Überzeugung,
daß dieses Leben nicht nur einzelne Probleme
in
Hülle und
Fülle enthalte, sondern daß es auch als Ganzes einer energischen
und
Revision
einer
gründlichen
Erneuerung
Streben danach aber schien mir auch
teiligen
zu
müssen,
Mitarbeit berufen.
ja
sie
bedürfe.
Am
die Philosophie sich be-
besonders schien
hier zu eifriger
Das brachte mich in Gegensatz zum Haupt-
zuge der heutigen deutschen Philosophie, der seine wissenschaftliche Arbeit unbeirrt
können meint.
fortführen zu
namentlich
durch jene Fragen und Zweifel ruhig
Wie viel Wertvolles diese Arbeit,
der genaueren Durchbildung der einzelnen Er-
in
kenntnisgebiete, geleistet hat
und dankbar anerkannt.
und weiter leistet, das sei freudig
Aber zugleich sei auch auf dem Rechte
und der Notwendigkeit jenes allgemeineren Problems mit aller
Entschiedenheit bestanden; wir werden uns in der Arbeit dafür
in keiner Weise
durch die Sorge um die Stellung anderer dazu
beirren lassen, sondern lediglich
und
allein
der inneren Not-
wendigkeit der Sache vertrauen.
Aber
Fülle
es sprechen neuerdings
dafür,
eintreten, ihre
unserer
auch Zeichen in Hülle und
daß weitere Kreise den Problemen,
für die wir
Teilnahme zuwenden. Die inneren Verwicklungen
Kultur,
ja
unserer gesamten
geistigen
Lage werden
Vorwort
VII
immer augenscheinlicher, mehr und mehr empfinden wir darin
schwere Unwahrheiten, Phrasen,
wo
wir Brot suchten.
Sinn
unseres
Nun
wo wir Wirklichkeiten, Steine,
das Glück und der
dabei
steht
dem Spiele; so erhebt sich
eignen Daseins auf
immer dringender das Verlangen nach Klärung wie nach Befestigung,
so
wird auch die Philosophie immer zwingender
zur Arbeit an diesen Lebensfragen aufgerufen.
Neue Wogen
des Lebens steigen auf, neue Stimmungen ergreifen die Gemüter
und heißen sie neue Ziele suchen.
Diese inneren Wandlungen haben auch meinen Büchern
mehr und mehr Freunde zugeführt und mir das Bewußtsein
engen geistigen Kontaktes mit der Zeit gegeben, das ich
eines
früher
nicht
haben konnte.
Mit besonderer Freude begrüße
ich die unerwartet rasch wachsende Teilnahme des aufsteigenden
jüngeren Geschlechts; möchte solche Teilnahme auch diesem
Buche zugute kommen, und möchte
Weiterführung der hier bloß
sehr
entworfenen
die Idee
eines
und
sicherlich oft
ist
schließlich
nichts
Denn was
geringeres
neuen Menschen und einer neuen Kultur;
nur ein Zusammenschluß der
alles
namentlich zu einer
unvollkommen behandelten Probleme wirken.
uns gemeinsam vorschwebt,
als
sie
Kräfte,
nur eine Überwindung
bloß Individuellen, nur das Entstehen einer durchgehenden
Bewegung kann uns
bei
einer so
gewaltigen Frage weiter-
bringen.
Jena, im Februar 1Q04.
Rudolf Eucken
Vorwort zur vierten Auflage
Die
vierte
Auflage
ist
gegen die
dritte
nicht
so
umge-
wandelt worden wie diese gegen die zweite, immerhin bringt
sie
Mehrere Abschnitte sind völlig um-
manche Veränderung.
einer
gearbeitet,
(über den
Wert des Lebens) neu hinzugefügt
worden; durchgängig aber war das Bemühen, die Darstellung
flüssiger,
stalten,
dem
den Inhalt gesättigter, die Hauptthesen präziser zu ge-
die
allen
Probleme der Zeit unmittelbarer zu
das
Ganze
anschaulicher
und
erfassen,
mit
eindringlicher
zu
machen; auch wurden diesmal weit mehr die Bewegungen des
Auslandes
in die Betrachtung hineingezogen.
So hoffe ich; daß
die neue Auflage als Ganzes einen merklichen Fortschritt bringt.
Jena, Ende August 1Q08.
Rudolf Eucken
Inhaltsübersicht
Einleitung.
Seite
'Die Zeitlage
A.
und unsere Aufgabe ihr gegenüber
1
Zum Grundbegriff des Geisteslebens.
1.
Subjektiv -Objektiv.
a)
Geschichtliches
b)
Das
c)
Die positive Behauptung.
a.
•
.
19. Jahrhundert
18
Einführung
27
Der Grundbegriff des Geisteslebens
Y. Das Verhältnis des Menschen zum Geistesleben
30
ß.
o.
2.
10
....
Ergebnisse für den Wahrheitsbegriff
33
35
Theoretisch - praktisch (Intellektualismus -Voluntarismus).
a)
37
Geschichtliches
Behauptung des Voluntarismus
c) Der Pragmatismus
d) Die eigene Behauptung: der Aktivismus
e) Intellekt und Intellektualismus
a. Die Überflutung des modernen Lebens durch den Intellekb) Die
43
.
y.
o.
£.
Die Begründung des Erkennens im Lebensprozesse
Die bewegende Kraft im Wahrheitsstreben
Konsequenzen
Konsequenzen
Philosophie
51
52
53
tualismus
ß.
47
...
für die Erkenntnisarbeit
für
die
56
59
63
Behandlung der Geschichte der
65
Inhaltsübersicht
3.
B.
Seite
Idealismus- Realismus.
a)
Die Ausdrücke
b)
Zum Kampf der Lebensgestaltungen
68
70
a.
Der Realismus des
ß.
Die Schranken des neuen Realismus
y.
Kritik der
o.
Erörterung des Wirklichkeitsproblemes
79
£.
Die Forderungen eines neuen Idealismus
81
19. Jahrhunderts
überkommenen Formen des Idealismus
72
74
...
76
Zum Erkenntnisproblem.
1.
Denken und Erfahrung (Metaphysik).
a)
Geschichtliches
b)
Das Recht einer selbständigen Philosophie
84
Die Wendung zur Metaphysik
d) Der Gesamtanblick der menschlichen Erkenntnisarbeit
93
104
c)
e)
2.
Würdigung des Rationalismus und des Empirismus.
.
111
...
117
.
.
Mechanisch - organisch (Teleologie).
Zur Geschichte der Ausdrücke und Begriffe
Problems
c) Erwägungen zum Kampf der Gegenwart
a. Das Problem im Gebiet der Philosophie
ß. Das Problem in der Naturwissenschaft
y. Das Problem auf gesellschaftlichem Gebiet
3.
a)
126
b) Zur Geschichte des
130
142
144
149
Gesetz.
a)
Zur Geschichte
b) Der
154
Kampf um das Gesetz in der Neuzeit
159
C. Zum Weltproblem.
1.
2.
Monismus und Dualismus.
a) Zur Geschichte und Kritik der Begriffe
b) Der Monismus der Gegenwart
170
183
Entwicklung.
Zur Geschichte des Ausdrucks
Zur Geschichte des Begriffs und Problems
c) Die Verwicklungen und Schranken der bloßen Entwicklungs-
a)
192
b)
193
d)
lehre
206
Forderungen für einen neuen Lebenstypus
221
D. Zu den Problemen des Menschenlebens.
1.
Kultur.
a)
Zur Geschichte des Ausdrucks und
Begriffs
228
Inhaltsübersicht
XI
Erwägung.
Das Problem des Wesens und Wertes der Kultur
Das Problem des Inhalts der Kultur
Das unsichere Verhältnis des Menschen zur Kultur
Seite
b) Kritische
a.
.
.
.
.
.
240
a.
Die Notwendigkeit einer tieferen Begründung
243
ß.
Die Notwendigkeit einer inneren Weiterbildung der Kultur
247
ß.
y.
c)
2.
237
Forderungen für ein wahrhaftiges Kulturleben.
Geschichte.
Zur Entwicklung des Problems
Forderungen und Ausblicke
Anhang: zum Begriff des Modernen
3.
235
.
a)
253
b)
262
273
und Individuum (Sozialismus).
Das Verhältnis von Gesellschaft und Individuum.
Gesellschaft
a)
a.
Geschichtliches
ß.
Die Probleme der Gegenwart.
,
283
aa.
Die Unzulänglichkeit einer bloßgesellschaftlichen Kultur
292
ßß.
Die Unzulänglichkeit einer bloßen Individualkultur
302
ff.
Die
Notwendigkeit
einer
inneren
.
Überwindung des
Gegensatzes
312
»
b) Die sozialdemokratische
4.
Bewegung
313
Probleme der Moral.
Die unsichere Stellung der Moral
in der Gegenwart
und Metaphysik
c) Moral und Kunst (ethische und ästhetische Weltanschauung)
a)
.
b) Moral
5.
a.
Die Geschichte des Problems
ß.
Die Probleme der Gegenwart.
.
322
324
329
aa.
Der moderne Ästhetizismus
ßß.
Die Stellung der Kunst im modernen Leben
Persönlichkeit
.
336
.... 339
und Charakter.
a) Persönlichkeit.
Zur Geschichte des Ausdrucks
Zur Geschichte des Begriffs
343
ß.
y.
Untersuchung des Problems
348
a.
345
b) Charakter.
6.
a.
Geschichtliches zum Ausdruck
ß.
Die Lage der Zeit
und Begriff
355
358
Freiheit des Willens.
a)
Einleitung
b)
Erwägungen zur Behauptung des Determinismus
363
.... 366
Inhaltsübersicht
XII
E. Letzte
1.
2.
Seite
Probleme.
Der Wert des Lebens.
a)
Einleitendes
376
b)
Die Verwicklung der Gegenwart
378
Das Problem der Religion (Immanenz und Transzendenz).
Zur Geschichte der Ausdrücke
b) Die Bewegung der Neuzeit zur Immanenz
c) Die Verwicklungen im Begriffe der Immanenz
d) Das Wiedererwachen des religiösen Problems
390
392
a)
e)
Forderungen für die gegenwärtige Lage der Religion
.
394
396
.
.
398
Schlußwort
404
Sachregister
405
Einleitung.
Die Zeitiage und unsere Aufgabe ihr gegeniiber.
VV /er
• '^
die
Lage der Zeit überblickt und
geistige
der wird
prüft,
vor allem eine starke Verworrenheit und eine peinliche Un-
sicherheit
über das Hauptziel des Strebens empfinden; überall ein
Auseinandergehen
spaltensein
Menschheit
der
des Menschen bei sich
unsichere Stand
Überlieferung
in
Parteien,
selbst.
oft
auch
ein
Ge-
Dieser verworrene und
mag zunächst als eine Wirkung der geschichtlichen
Denn von der Vergangenheit her um-
erscheinen.
fangen uns verschiedenartige,
ja
einander feindliche Strömungen, ein
Erbe und eine Last aus tausendjähriger Arbeit; nichts unterscheidet
moderne Kultur mehr von der einfacheren des Altertums als
solches Durchtränktsein von Gegensätzen. Das Mittelalter überlieferte
die
ein
Lebensganzes, das die grundverschiedene antike und
liche,
künstlerische
und
religiöse,
weltfreudige
und
altchrist-
weltfeindliche
Denkweise weniger gegenseitig ausglich als geschickt zusammenfügte.
Diesem Gefüge
setzte die Neuzeit einen
neuen Lebensdrang, ein Ver-
langen nach unbegrenzter Entfaltung der Kraft und voller Beherrschung
der Dinge entgegen,
aber die nähere Ausführung dessen entzweite
das Neue alsbald bei sich selbst: einerseits verlangte das Seelenleben
mit seinem Denken, andererseits die Natur mit ihrem Mechanismus
die Herrschaft über Leben
mus).
Alle
historischen
und Welt (Intellektualismus und Naturalis-
solche Gegensätze
läßt
das 19. Jahrhundert mit seiner
Bildung und seiner grüblerischen Reflexion
in
greller
Deutlichkeit sehen und mit voller Stärke empfinden, es drängt zwingend
zu einer schärferen Scheidung der verschlungenen Gedankenmassen, es
verbietet ihnen
immer strenger ein friedliches Zusammengehen. Und
wie viel hat das 1 9. Jahrhundert bei sich selbst erlebt, wie eingreifende
Eucken, Grundbegriffe.
4. Aufl.
1
Einleitung.
2
hat es durchgemacht,
Wandlungen
äußerer Zurückdrängung uns
deren einzelne Phasen auch bei
innerlich nahe bleiben und nach wider-
streitenden Richtungen ziehen: die künstlerische Geisteskultur unserer
klassischen Literaturepoche, ein kraftvoller
und selbstbewußter Realis-
mus, ein Rückschlag gegen diesen Realismus in einem Subjektivismus
seelischen Fürsichseins,
schwebender Stimmung!
Wie viele Gegen-
wir aus altem und neuem Bestände in uns,
sätze
tragen
haben
wir zu tun,
wie
viel
um ihrer innerlich Herr zu werden!
Um diese verschiedenartigen Anregungen zu verarbeiten und
mit einander auszugleichen, bedürfte es überlegener geistiger Kraft.
Da
diese
so
fehlt,
erscheinen
alle
Mißstände,
welche das
Be-
Menschen von seinen eignen Erlebnissen, das
des
wältigtwerden
Unterliegen unter die Zerstreutheit des Daseins herbeiführen muß.
Keine festen Ziele beherrschen das Streben, keine einfachen Ideen
dem Chaos und
entwinden sich
befreien
von seinen Wirren und
Zweifeln.
Vielmehr bewältigen uns die unmittelbaren Eindrücke und
zerreiben
das Leben
unsicher auf den
unter
ihren
Widersprüchen.
Wogen der Zeit einher,
So treiben
wehrlos gegen
alles,
mr
was
uns mit starkem Bewußtsein und kecker Behauptung naht, wehrlos
auch gegen die eigenen Einfälle und Leidenschaften, ein Spiel von
wechselnden Lagen und Launen.
Diese Lage wird namentlich dadurch gespannt, daß die Wandlungen,
die wir erfahren, sich schließlich zu Einer Frage verbinden
und uns vor ein einziges Entweder-Oder stellen, das keine Verschleierung
duldet
und
eine
Entscheidung
des ganzen Menschen
Das stille, aber unablässige und unwiderstehliche Wirken
der modernen Arbeit hat an der überkommenen Lebensführung nicht
verlangt
nur
alle
einzelnen Punkte verändert, es hat sie als Ganzes unter-
graben und unhaltbar gemacht
offen
oder
versteckt,
Die
ältere
Denkweise behandelte,
gröber oder feiner, sinnlicher oder
geistiger,
den Menschen als das Maß und den Mittelpunkt des Alls, verwandelte
die
Wirklichkeit
in
ein
Reich
von
menschenartigen Größen
machte das Wohlergehen des Menschen zum Ziele
und
alles Geschehens.
Diesen Anthropismus hat das Ganze der modernen Arbeit gründlich
zerstört;
nicht
nur die unermeßliche Erweiterung der äußeren Welt,
auch die Aufdeckung innerer Notwendigkeiten und sachlicher Zusammenhänge im eigenen Bereich der Menschheit, ein weites Hinauswachsen des geistigen Schaffens über das bloße Subjekt machen jenes
Sicheinspinnen in das Menschliche zu einer unerträglichen Enge, sie
Einleitung.
erwecken
zugleich
glühendes Verlangen nach einem weiteren,
ein
freieren, gehaltvolleren
3
Durst nach einem Leben
Sein, einen starken
mit der Unendlichkeit und der Wahrheit des
lungen treten immer mehr auch
und verlangen stürmisch
Alls.
Diese
Wand-
das Bewußtsein der Menschheit
in
ihr Recht.
Aber aus dem Nein entspringt dabei keineswegs rasch
und der Erschütterung entspricht nicht eine Befestigung.
neue Lage eröffnet zwei Möglichkeiten,
ein Ja,
Denn die
die, als schroffe Gegensätze,
Hat jene weltgeschichtliche Bewegung
keinerlei Ausgleichung dulden.
gegeii das Beharren beim Bloßmenschlichen den Sinn, daß der Mensch
sich als ein bloßes Naturwesen
Tun dem Rahmen
zu verstehen und all sein Sinnen und
Dann wäre
habe?
der Natur einzufügen
alles
unterscheidend und auszeichnend Menschliche als ein verderblicher
Wahn zu entfernen,
Größen und Güter unseres Lebens hätten
empfangen. Oder besagt
jene Bewegung, daß innerhalb des Menschen selbst eine neue Weib
eine geistige Welt emporsteigt und ihn wie über die Natur auch
über sich selbst hinaushebt? Beginnt mit ihm eine neue Stufe der
Wirklichkeit, und kann sein Seelenleben sich von innen her zu einer
Welt erweitern? Dann würde zur Hauptaufgabe die Ergreifung,
Aneignung und Ausbildung dieser Welt, dann müßte der Mensch
vor allem hier sich befestigen und all sein Sinnen und Streben
nicht sowohl rückwärts als vorwärts richten.
So ist der Mensch
alle
ihr Gesetz und ihre Gestalt von der Natur zu
entweder weniger oder mehr,
je
als er sich
heute einzuschätzen pflegt;
nach der Entscheidung für dieses oder jenes muß aber das Leben
sich
vom Größten
bis
zum
Kleinsten
verschieden gestalten.
Aber
so unerläßlich die hier gebotene Entscheidung ist, jene Schwäche des
Einheitsstrebens läßt die Zeit unsicher zögern
wechselnden Eindrücken neigt
sie
sie
im Gesamturteil das eine billigt,
bald
und schwanken, nach
hieher
bald
dahin;
indem
will sie zugleich von dem anderen
nicht lassen; so bejaht sie hier,
was sie dort verneint, so setzt sie an
keiner Stelle ihr ganzes Wollen
und Wesen
ein.
Oft genug ist diese
Lage geschildert worden, ihr rascher Wechsel der Strömungen und
Stimmungen,
lichkeit
ihr
Mangel an Logik, wie ihn sowohl die Unempfind-
auch für die härtesten Widersprüche und das Ineinander-
schieben verschiedenartigster Gedankenmassen, als die Schwäche des
Ausdenkens, des Verfolgens der Behauptungen in ihre Voraussetzungen
wie ihre Konsequenzen
zeigt.
In allem
Sinken des inneren Lebensstandes,
ja
diesem erscheint ein starkes
eine
innere
Verarmung des
Einleitung.
Lebens, das aber inmitten staunenswerter Fortschritte an seiner Peripherie, inmitten nie gesehener, nie geahnter Virtuosität von technischen
Leistungen, inmitten überströmenden Reichtums an äußeren Erfolgen
Augenscheinlich befinden wir uns
uns zu übermannen droht.
heit
Aber es
in
ist
einer geistigen Krise, die
diese Krise nicht der Bos-
oder der Zweifelsucht Einzelner, sondern
der weltgeschichtlichen Lage entsprungen.
dem Ganzen
ist
Sollten wir nicht hoffen
daß die Notwendigkeit, welche eine solche Krise erzeugte,
dürfen,
uns
sie
irgendwelche Mittel
auch
und Hülfen gewährt,
die
über sie
Widerständen
und
Gegen-
hinausführen könnten?
Wahrheit
In
fehlt
es
nicht
an
wirkungen gegen jene chaotische Lage, an Versuchen,
ihr eine ein-
heitliche Gestaltung des Lebens, ein einheitliches Bild
der Wirklich-
keit
entgegenzusetzen; leider bleiben aber diese Versuche meist unter
dem Einfluß dessen, das sie überwinden möchten.
Die Zeit des selbst-
bewußten Spezialismus, den die Arbeit an der endlosen Breite der
Dinge
alle
hinter
sich.
meist so,
Sorge
um das Ganze vergessen ließ,
Aber das Streben zur Einheit
hat ihren Gipfel
gestaltet
sich
zunächst
daß die einzelnen Lebens- und Wissensgebiete die Sache
an sich reißen und das Bild
Eindrücken,
vom Ganzen
nach ihren besonderen
Zwecken entwerfen.
Erfahrungen,
Mehr
als
es sonst
geschah, erzeugen sie innerhalb ihres besonderen Kreises geschlossene
Gedanken massen, dringen damit kühnlich über die Grenzen jenes
Ihre
Kreises hinaus und möchten die ganze Wirklichkeit meistern.
besonderen Aufgaben werden vor alle übrigen gestellt, ihre Begriffe,
Maßstäbe, Methoden sollen schlechtweg gelten, ihr Gebiet wird ihnen
zum beherrschenden Mittelpunkt der gesamten Wirklichkeit. So bildet
sich die Religion, so oft auch die Kunst ihre eigene Welt, so erzeugt
die soziale
Bewegung ihre besondere Weltanschauung, so erweitern
sich auf intellektuellem Gebiet namentlich oft die Naturwissenschaften
Das tat zunächst die Zoologie
zu einer allumfassenden Philosophie.
unter
dem Einfluß des Darwinismus,
Physikern,
Physiologen
Weltgedankens
ist
jetzt
gewandert, und es
u.
s.
das sehen wir
w. unternommen.
jetzt
auch von
Die Kühnheit des
von den Philosophen zu den Naturforschern
fehlt hier nicht
an kecken Husarenritten in das
Land der Wahrheit; die Verquickung der philosophischen Behauptung
mit tüchtiger Forschungsarbeit läßt dabei manche das Ungeheuerliche
des Wagnisses
kaum empfinden.
So entstehen eigentümliche Durchblicke,
Partialweltbilder, deren
Einleitung.
sinnliche
Nähe und
leichte
Faßlichkeit die Geister gewinnt
gutes Stück mit sich fortreißt.
schließlich
wird
5
Wahrheit der Dinge Widerstand
die
und
Jedoch immer nur ein Stück.
zu knappe Maß zersprengen;
ein
Denn
und
leisten
wird es um
das aufgedrängte,
viel
so eher, als jene
Bewegung die verschiedenen Ansprüche bald zu-
sammenstoßen und
und daß
samtwahrheit
Nun
vom Teil zum Ganzen bauen
sich ihr Recht gegenseitig bestreiten läßt.
wird offenbar, daß sich nicht wohl
läßt,
Überspannung zur Ge-
die Teilwahrheiten mit ihrer
sich
sie
in
Soweit aber jene Partial-
Irrung verkehren.
bewegungen Macht behaupten, einander hemmen und durchkreuzen,
müssen
sie
die Verwirrung,
die sie
bekämpfen, vielmehr steigern;
kaum etwas so sehr zur Entzweiung als jenes
vielleicht wirkt heute
unzulängliche Streben nach Einheit.
Nie war so
viel
die
Rede von
Monismus wie heute, und nie ging die Menschheit so weit auseinander wie heute.
Aber so unzulänglich jene Versuche sind,
sie bleiben
wertvoll
durch ihre Lehren.
Namentlich zeigt ihr Scheitern mit voller Klar-
daß sich
von den einzelnen Punkten her ausrichten läßt,
heit,
nichts
sondern es eine der Zerstreuung überlegene Einheit zu suchen
ohne eine Erhebung über das Ganze der
Zeitlage,
ohne
gilt;
ein Er-
greifen neuer Anfänge gibt es keine Hoffnung, der Krise gewachsen
zu
Aber warum
werden.
schichte
ist
für
sollte
jenes
unmöglich sein?
das Innere des Lebens kein
Die Ge-
fortlaufender Anstieg
zur Höhe; da wesenhaftes Geistesleben sich in der Geschichte nicht
nur entwickelt, sondern sich auch
in
ihr auslebt, so kommen immer
wieder Zeiten, wo es aus der Wirkung im menschlichen Dasein zu sich
selbst
zurückkehren und die Wurzeln seiner Kraft neu beleben muß.
So allein kann es der Zeit überlegen werden und dahin wirken, das
Wahre in ihr von dem Problematischen zu befreien, das uns beirrt
und
gilt
entzweit.
Eine
solche Zeit
eine Selbstbesinnung auf die
unser Grundverhältnis
zur Welt,
bloßen Zeit an das Ewige
die überlegenen Gewalten
in
ist
einmal wieder gekommen,
es
Grundlagen unseres Daseins, auf
es
eine
gilt
der Zeit,
Berufung von
der
vom bloßen Menschen an
die aus dem Menschen
und Ordnungen,
mehr als ein bloßes Naturwesen machen.
Bei solcher Lage hat jeder, der den Notstand durchschaut, nach
dem Maße seines Vermögens für jenes Ziel der Vertiefung des Lebens
und der Erneuerung der Kultur zu wirken. Der Weg, den unser Werk
dabei einschlagen soll, wird namentlich durch drei Merkmale bestimmt.
Einleitung.
Unsern nächsten Vorwurf sollen die der Zeit charakteristischen
1.
Hauptbewegungen, die geistigen Strömungen,
Wir sprechen von
heißen mag.
um
oder Ideen,
bei
nicht
als
ob
um bloß intellektuelle Vorgänge handle und
Mag sich
diesen die Entscheidung liege.
auf
Kürze
von Begriffen
von Anfang an die Meinung fernzuhalten,
es sich an erster Stelle
vorwiegend
bilden, wie es in
und
solchen
Gebiet
intellektuellem
der Streit
äußerlich
dahinter
abspielen,
stehen
Lebensbewegungen aus dem Ganzen, dahinter steht eine eigentümAbsteckung der Wirklichkeit,
liche
des
Lebens;
inmitten
vielfachen
Probleme hindurch kann
Gestaltung
eigentümliche
eine
und durch verschiedene
Streites
bei diesen Voraussetzungen der Zeit eine
Gemeinschaft walten; so
Heraushebung besonders geeignet,
ihre
ist
zu einem Gesamtbilde der Zeit zu verhelfen und das Eigentümliche
der Zeit klar erkennen zu lassen.
Das Ausgehen von der Vielheit
aber hat den Vorteil, uns die Behauptungen und Probleme der Zeit
und anschaulicher zu machen;
greifbarer
die
Erörterung
dem
rasch
Notwendigkeiten
sachliche
Wege
Denken
zeigen,
zu
es hat den weiteren Vorteil,
bestimmten Punkt zu führen,
einen
auf
hervorzubrechen
daß wir überall auf dieselben Fragen kommen,
ja
an
unserem
Untersuchung
Die
vermögen.
weisen
und
wird
daß ein
und dasselbe Hauptproblem durch alle Mannigfaltigkeit wirkt, auch
wird
sie
wird,
so
zeigen,
die
daß, wie an jeder Stelle
um das Ganze gekämpft
Entscheidung über das Ganze
in
alle
Verzweigung
Der eignen Behauptung darüber dürfen wir uns aber
hineinreicht.
um so sicherer fühlen, je mehr die Erfahrungen und Forderungen
der einzelnen Punkte zu ihr drängen und sie als die einzige Möglichkeit einer glücklichen
2.
Lösung zeigen.
Was wir aber näher bei den einzelnen Strömungen ermitteln
und wonach wir sie prüfen wollen, das ist der von ihnen behauptete
oder doch
in
die Frage
beschäftigen,
ihnen enthaltene Lebensprozeß;
namentlich
ob dieser Lebensprozeß
Geistesleben
möglich macht.
lebens pflegt,
wenn auch
werden; wie
viel
aber in
ein
soll
uns
selbständiges
Ein gewisser Tatbestand des Geistes-
oft widerwillig,
ihm
liegt,
von jedem anerkannt zu
und was er über die nächste
Erscheinung hinaus verlangt,
an welche Voraussetzungen und Be-
dingungen er geknüpft
das
ist,
meist in völligem Dunkel.
bleibt
Wie sich die Bewegungen der Zeit zu diesem Problem, zum Problem
der Möglichkeit des Geisteslebens,
darauf
sei
stellen,
vornehmlich das Augenmerk
und was sie dafür leisten,
gerichtet.
So soll uns
nicht
Einleitung^
die Breite der Leistungen
sondern wir streben rasch zu
festhalten,
dem sie durchwaltenden Leben, als dem letzten Punkt, der erreichbar ist und von dem aus sich unsere Gedankenwelt aufzubauen
hat; es bringt uns solche Wendung zum Lebensprozesse wohl am
sichersten an den Punkt, wo die Probleme dem Einzelnen zum eignen
wo
Erlebnis werden,
er
am
eigene Erfahrungen einsetzen
ehesten
und am wenigsten eine eigne Entscheidung ablehnen kann.
3. Wo der Gehalt der Zeit den Ausgangs- wie den Endpunkt
da empfiehlt sich ein Heranziehen der geschichtlichen Be-
bildet,
trachtung zur
Jene Be-
Unterstützung der philosophischen Arbeit.
trachtung helfe zunächst dazu, die geistige Art der Gegenwart durch
Aufdeckung
ihres
beleuchten
und
Werdens und
deutlicher
Zusammenhänge
ihrer
Für
abzugrenzen.
die
heller
Fassung
zu
und
Schätzung dessen, was die Zeit beherrscht, kann es nicht gleichgiltig
ihm eine Woge des bloßen Augenblickes oder
einen bleibenden Lebensstrom erkennen, ob das heutige Erlebnis
schon öfter erlebt wurde und einem wiederkehrenden Rhythmus anob wir
sein,
gehört,
Neues und Eigenartiges aufeine Gegenwirkung,
oder ob
in
ihm etwas
völlig
ob
es
mehr
Wirkung oder
auch
steigt,
in
eine
mehr einen Vorstoß oder einen Rückschlag
schichtliche
Betrachtung hat
schieden weit in
Bewegung
Oft wird die
Hauptphasen durch das Ganze der europä-
Kulturentwicklung
ischen
den verschiedenen Punkten ver-
bei
die Vergangenheit zurückzugreifen.
ihren
in
Diese ge-
bedeutet.
zu
verfolgen
sein,
an
anderen
Stellen
wird die nächstvorangehende Epoche zur Aufklärung der Gegenwart
genügen.
Hand der
mag eine selbständige Untersuchung vorbereiten, wenn
Solche hellere Beleuchtung des Tatbestandes an der
Geschichte
anders ein Ding in seiner Eigentümlichkeit schärfer sehen, zugleich
seine
Grenzen deutlicher sehen und
ein
Problem
in
ihm erkennen
Aber nicht nur die Behauptung der Gegenwart, auch die
heißt
geschichtlichen
Zusammenhänge,
wandeln
bei
sich
ja
Aufdeckung des
die Geschichte als
in
Ganzes ver-
ihnen wirksamen Lebenspro-
zesses in ein Problem; der Lebensprozeß mit seiner Bewegung läßt
sich
aus
dem Chaos
der Erscheinungen
nicht
wohl heraussehen
ohne eine Versetzung aus der geschichtlichen Betrachtung
zeitlose
Seiner Wahrheit
in
eine
Aufnehmen der Frage nach
und seinem Rechte; das Ganze läßt sich nicht
und unmittelbare, ohne
ein
durchleuchten, ohne daß sich ursprüngliche, eigene, letzte Tatsachen
Einleitung.
von den übermittelten scheiden.
Wendung zu
eine
und Erörterung
Umkehrung mit ihrer Verwandlung
der Sache vollziehen; erst diese
der Geschichte
mag sich so eine Umkehrung,
Es
einer unmittelbaren Betrachtung
Lebens macht
Entfaltung eines zeitlosen
die
in
es
den Befund des Daseins von innen her zu durchschauen,
möglich,
von der Erscheinung zur Tatsache, vom bloßen Faktum zur be-
gründenden Wahrheit vorzudringen,
erkennen,
ja
ihrem
Ganzen
den Bewegungen der Ge-
in
und durchgehende Richtungen zu
innere Notwendigkeiten
schichte
irgendwelchen
Sinn
zu
entringen.
Auf Grund solcher Betrachtung aus bleibender Wahrheit
erst
läßt sich
Bedeutung der einzelnen Epochen ermessen, sowie an der
die
der
Leistung
Gegenwart eine immanente
werde an dem
hauptung der Zeit
Die
Be-
weltgeschichtlichen Stande
der
üben.
Kritik
mehr Gehalt
und Tiefe erschlossen, als jene Behauptung in sich aufnehmen kann,
hat die Geschichte schon
geistigen Evolution geprüft;
so wird das Streben notwendig über sie hinausgetrieben, auch
es zugleich
es weiter
eine
Anweisung über
zu suchen
mit
Bei solcher Verflechtung
hat.
ischen
Arbeit
Kritik
nicht zurückschauend
der
die Richtung empfangen,
weltgeschichttichen
und
die sie fordert,
von
der philosoph-
Erfahrung
braucht
reflektierend zu bleiben,
produktiv und vordringend werden,
sie
mag
in der
sie
die
kann
kann die Weiterbewegung,
sich aus fördern.
Eine solche Untersuchung hat darauf vor allem ihr Augenmerk
zu
richten,
die Selbstverständlichkeit zu zerstören,
mit der sich die
Bestrebungen einer Zeit zu geben gewohnt sind, und zugleich von dem
Dogmatismus zu
befreien,
Bedingung dafür
bildet
unternimmt und
erreicht,
der jenen anzuhaften
präziseres
ein
präzise
Umfang der Leistung sehen, und
Die
pflegt.
erste
Sehen dessen, was die Zeit
sehen
heißt
hier
zugleich
den
das allein macht es möglich, zu
einem Urteil zu gelangen, das selbständiger und
kräftiger
Art
ist,
ohne ungerecht zu werden und Paradoxie für Unabhängigkeit zu
geben.
Darauf vornehmlich
der Mannigfaltigkeit und
sei
dem
also
das
scheinbaren
Augenmerk
gerichtet,
in
Durcheinander der Be-
wegungen durchgehende Züge, einfache Grundlinien aufzudecken;
von hier aus am ehesten werden wir hoffen dürfen, den Wahrheitsgehalt der Zeit, ihre inneren Notwendigkeiten von der entstellenden
Zutat
menschlicher Irrung
und Leidenschaft zu
befreien,
aber Anhaltspunkte für das eigne Streben zu gewinnen.
zutreffend
beurteilen
zugleich
Die Zeit
kann nur, wer sie innerlich mitzuerleben ver-
Einleitung.
mag; wer sich
ihr
von vornherein
9
krittelnd
und nörgelnd entgegen-
dessen Urteil hat keinen Wert.
stellt,
Endlich
sei
noch hinzugefügt, daß, wie
in
den früheren Auf-
lagen des Buches, so auch in dieser die Bezeichnungen der Hauptbegriffe
sorgfältig
Gebrauch verschuldet zum
Indem einmal
Gegenwart.
strengere
Fassungen
Ihr
schwankender
nicht geringen Teil die
Verwirrung der
werden
beachtet
oft
bei
sollen.
demselben Ausdruck laxere und
durcheinanderlaufen,
erschleichen
hauptungen mehr Sicherheit und mehr Gehalt,
heit
indem
gebührt;
ferner
verschiedene Bedeutungen
Sache
und
versteckt
sich
nicht
hat,
selten
verwirrt
als
dasselbe
sich
leicht
leicht
Be-
ihnen in Wahr-
Wort
wesentlich
der Anblick der
der Punkt der Entscheidung.
Zu
allen
Zeiten decken die Ausdrücke und die Begriffe sich nur in annähernder
Weise,
heute
Bekämpfung
aber
gehen
sie
besonders
weit
auseinander.
Zur
eines solchen Mißstandes bedarf es auch eines Blickes
auf die Geschichte der Ausdrücke; so sei auch dieser ein bescheidner
Platz
gewährt.
Zum Grundbegriff des Geisteslebens.
A.
Subjektiv
1.
a)
- Objektiv.
Geschichtliches.
|as Verhältnis von Subjekt und Objekt
I
*—^
punkte der Arbeit und des Kampfes;
des einen oder des anderen
die Begriffe
gestalten
Hauptbewegung des Lebens entweder vom Men-
zum Menschen.
schen zur Welt, oder von der Welt
Stand
müssen
In
alle
die
übrigen Probleme einmünden,
Behandlung
Phasen werden
wir
Lösung erkennen und zugleich
Wie
so trägt sein
Wirkungen der gesamten Geschichte
bei seiner
diesen
Bilder
die
von der Wirklichkeit, die Fassungen der Wahrheit ver-
schieden, geht die
Problem
dem Vorwiegen
vom Leben,
nach
je
sich
heute im Mittel-
steht
in
dies
in
heutiger
und
sich,
es
ihre
Hauptphasen gegenwärtig sein.
die
wichtigsten
eine
Möglichkeiten
der
durchgehende Bewegung
ge-
wahren, welche, die Arbeit in eine gewisse Richtung drängt.
Eigentümliche Verwicklungen der Sache verrät schon die merk-
würdige Geschichte der Ausdrücke subjektiv und objektiv;
sie
im
Lauf
vertauscht.
Bei
Duns Scotus (f 1308), der zuerst
der Jahrhunderte
ander gegenüberstellte,
das
Subjekt
der
ihre
geradezu
sie als
also
auf
die
haben
Kunstausdrücke ein-
subjectivum dasjenige, was
»/hieß
Urteile,
Bedeutung
sich
auf
konkreten Gegenstände des
Denkens bezieht; hingegen objectivum jenes, was im bloßen obicere,
d. h.
im Vorstelligmachen,
liegt
stellenden
fällt"
IIL 208).
In diesem Sinne
sophen
stück
(s.
Prantl,
und hiemit auf Rechnung des Vor-
gehen die Ausdrücke
bis in das 17. und 18. Jahrhundert,
des
gebräuchlicheren
Logik
Geschichte der
objektive
im Abendlande,
bei
den
Philo-
doch bildet das Gegen-
öfter
formaliter
oder
auch
Subjektiv — Objektiv.
schon
U
Auch zeigen die Ausläufer der Scholastik
realiter. 1
Schwanken,
ein
gebrauch vorbereitet.
das
die
Wendung zum
objectivus
bei
neueren
Sprach-
2
Die völlige Umkehrung der Bedeutung erfolgte aber
Übergang
schen
in die deutsche Sprache,
so
Schule,
z.
erst beim
und zwar innerhalb der Woffi-
bei A. F. Müller (Einleitung
B.
sophische Wissenschaft,
1733),
die
philo-
Baumgarten und Gottsched.
Doch
in
bleiben die Ausdrücke (man sagt übrigens «subjektivisch"
an
zunächst
jektivisch")
gebunden,
Schule
die
und » ob-
so erscheinen
sie
dem Streit zwischen Lessing und Goetze als gelehrte
Termini; erst die Kantische Philosophie hat sie dem allgemeinen
Sprachgebrauch zugeführt, in dem sie zu Beginn des 19. Jahrhunderts einen breiten Raum erlangen.
Von Deutschland aus ist die
z.
noch
B.
in
neue Bedeutung zu den anderen Völkern gewandert und von ihnen
zunächst oft als fremdartig empfunden worden.
Der
heutige
Sprachgebrauch
so
ist,
deutlich
er
vom
sich
Schwankung und Un»Subjektiv bezeichnet zunächst das, was der bloßen
mittelalterlichen abhebt, bei sich selbst
sicHerheit.
voll
Vorstellung des Individuums angehört, nicht selten aber auch, namentich
Naturforschem,
bei
Wesen
fühlende
bei
und
alles
sich
selbst
jedes,
erlebt,
was
das
denkende
auch wird wohl
alle
und
Über-
zeugung, die den unmittelbaren Tatbestand überschreitet, subjektiv
und damit
genannt
und
für eine
erscheinen
Flachstes
nachträgliche
damit
als
Zutat erklärt.
gleichen
Tiefstes
»Objektiv
Rechtes.
leidet
namentlich an der Zweideutigkeit, daß es das Gegenständliche
bald
als
ihrer
befindlich
aller
seelischen Betätigung
innerhalb
moderne Naturalismus sein.
In
den Erörterungen
subjective
= formaliter in se
^
als
Gegenständlich wollte Goethe, gegen-
bezeichnet.
ständlich will auch der
gegenüber, bald
unterscheidet (oeuv. div. 1727,
zwischen Descartes und Qassendi
ipsis,
III.
findet sich
objective =: idealiter in intellectu.
Bayle
334a) objectivement dans notre esprit und
und noch bei Berkeley heißt es (Ausgabe von
phaenomena are only natural appearances. They
such as we see and perceive them. They real and objective
reellement hors de notre esprit,
Fräser
are,
II.
477): „Natural
therefore,
nature
'
are,
therefore, the
same."
So heißt es z. B. in Chauvins lexicon rationale (1692) unter certitudo:
objectiva nonnullis est ipsa necessitas objecti, seu propositio necessaria objectiva.
AHis autem
intellectus
per
nihil
aliud
est
quem objectum
nimmt ratio objectiva
quam denominatio quae sumitur ab actu
repraesentatur.
Goclen
(lex.
philos.
= res ipsa quatenus definitioni respondet.
1613)
Zum Grundbegriff des Geisteslebens.
12
In der Sache handelt es sich augenscheinHch
um das Verhältnis
des Menschen und seines Gedankenkreises zur Welt, der er angehört.
Soweit das Denken selbständig wird,
mit ihr unablässig beschäftigen
sich
Scheidung
zu
zugleich
selbst
Welt gegen-
es sich der
aber zugleich kann es nie vergessen,
über,
und
stellt
daß
muß;
zu ihr gehört
es
so entsteht mit der
unabweisbares Verlangen,
ein
sie
irgend
überwinden, das Geschiedene wieder zusammenzubringen
zu verwickeln,
mehr wir uns mit ihr befassen.
je
und
Diese Aufgabe aber scheint sich immer mehr
fest zusammenzuhalten.
Die Verwick-
lungen hat schon das griechische Altertum stark empfunden, aber
es
Die Lösung,
ist.
versuchte,
noch
mit ihnen
hat sich
möglich
ist
am
meisten
abgefunden,
leichter
als
uns Späteren
Höhe der klassischen Zeit
zu geschichtlicher Wirkung gelangt; was
welche
hier die leitenden Denker, ein Plato
die
und
ein Aristoteles, an Lehren
entwickeln, das schöpft vornehmlich daraus eine Überzeugungskraft,
daß es ein Ganzes der Lebensführung hinter sich
hat.
griechische Lebensführung hat ihre Eigentümlichkeit
und ihre Stärke
Diese
alt-
darin, das naive Verhältnis des Menschen zur Natur ohne einen schroffen
Bruch ins Geistige zu heben und zugleich zu veredeln, den Menschen
in
die Welt hineinzusehen,
zu
sich
selbst
Äußeres, haben
aber aus der Spiegelung ihn geläutert
zurückzuführen.
die
anfängliche
Mensch und Welt, Inneres und
Vermengung überwunden, aber sie
sind noch nicht so schroff geschieden,
um sich nicht durch geistige
Arbeit rasch wieder zusammenzufinden.
ander wesensverwandt
seiner eigenen
und
Denn
innerlich zugetan,
beides scheint ein-
jedwedes
bedarf zu
Vollendung der Ergänzung durch das Gegenstück:
von innerem Leben erfüllte Natur erreicht ihre Höhe in der
Aneignung durch den Menschen; was aber in diesem an Kräften
die
schlummert, das wird
erst
vollem Leben geweckt.
durch die Berührung mit der Welt zu
In der Einigung,
wie
sie
Anschauung und
Liebe bewirken, gewinnt das Leben die Höhe und Seligkeit geistigen
Schaffens.
Eine solche Überzeugung kann unbedenklich die Wahr-
heit als eine
Übereinstimmung unseres Denkens mit dem Gegenstande
(adaequatio intellectus et
rei)
fassen.
Diese Fassung genügt aber nur für einen Stand des Lebens,
wo
die
schien,
Natur
noch
seelischer
und der Mensch noch
wo weder jene eine volle Selbständigkeit in
natürlicher
eigentümlichen
und Gesetzen gewonnen, noch das Innenleben sich zu einer
eignen Welt vertieft hatte.
So gewiß jene größere Nähe und jene
Kräften
Subjektiv — Objektiv.
13
fruchtbare Wechselwirkung beider Seiten eine großgesinnte,
frohe,
künstlerische Kultur
hat bilden helfen,
lebens-
jener enge Anschluß
des Geisteslebens an das naive Weltbild war für die Dauer nicht
zu erhalten.
So hat denn schon das
Neuplatonismus neue
spätere Altertum in der Stoa
Wege versucht,
und im
aber stärker als diese hat auf
die Neuzeit jene ältere Art gewirkt, da sie eine nicht unbedeutende
Nachblüte
in
der mittelaltedichen Scholastik fand und durch diese
unmittelbar die Neuzeit berührte; namentlich in Auseinandersetzung
mit ihr hat diese ihre Eigentümlichkeit gefunden.
Es erscheint aber die neue Art zunächst
des Subjektes, einer trotzigen Losreißung von der
Entfaltung
und das Leben zu
her die Welt zu bilden
Welt zu empfangen und an
sie
sie
gestalten,
statt
Anschluß zu suchen.
aus der
Gewaltiger
hat die Wissenschaft den Anblick der Dinge verändert;
alles
ausscheidet,
was
Um-
vom Menschen und seinem Denken
gebung, einem kühnen Versuch,
als je
einer kräftigeren
in
Prüfung nicht besteht,
ihre
indem
das
Ver-
bleibende aber durchleuchtet und enger verbindet, wird das ganze
Dasein des Menschen
das Element des Gedankens getaucht und
in
Das Innere erkennt
seine Einheit und befestigt sich sicher im eigenen Kreise, die Außenwelt weicht davor zurück, sie verliert alles innere Leben, indem sie
Ideelle gehoben.
ins Gedankenhafte, Begriffliche,
zur räumlichen
Bewegung
sie verliert alle
bunte Farbe, indem die ganze Fülle der sinnlichen
keiner Seele
mehr zu bedürfen
scheint,
Eigenschaften aus einem eigenen Besitz der Dinge zu einem bloßen
Gewände wird, womit sie die
fremd;
innerlich
und mit
So wird nunmehr
von leblosen Massen und Bewegungen der
die Natur als ein Reich
Seele
Seele umkleidet.
als
auf sich selbst gestellt
die Unendlichkeit
bezwingend, fühlt sich
die Seele
ihrer Denkkraft
aber,
ihr weitaus überlegen.
Das
ein Hauptstück,
ist
der Neuzeit,
nicht
aber
ist
vielleicht
es
das Hauptstück des Lebens
das Ganze dieses Lebens.
Denn
unverkennbar hat die Neuzeit neben dem Drange zur Steigerung
des Subjektes
auch den entgegengesetzten Zug,
des Menschen heraus zur
gegenüber
dem
menschlichen
wirren
Kreises
ein
aus der Kleinheit
Größe der umgebenden Welt zu flüchten,
Getriebe und der dumpfen Enge des
weiteres,
gehaltvolleres,
reineres
Leben
dem unermeßlichen All zu schöpfen. So ein Trieb zum Objekt,
ein Streben, sich in sein Wesen zu versenken, seinen Gehalt ohne
aus
Zum Grundbegriff des Geisteslebens.
14
Trübung
aufzunehmen.
sich
in
Dinge, der Erfahrung,
Hier wird von der Mitteilung der
der Mensch darf seine
Heil erwartet;
alles
Art der Welt nicht irgendwie aufdrängen wollen, er
dienstwillig einfügen,
die Verstärkung
Denn
indem
selbst
alle
des Subjekts
das
Subjekt
Ende
bereitet,
heit
entfalten,
Ganzen
bei
alten
verketten; jetzt erst,
Reich
von
die Natur
auf
sich
Das
Gesetzen.
klammern,
Vermenschlichung des Weltbildes ein
leerer
wie
ein
dem Men-
Wahn, und
es wird
Von
zu unterwerfen suchen.
alles
Fürsichsein des Subjekts
gefordert,
daß das Leben sich
anschmiege und lediglich ihren Geboten
folge.
Zusammenschluß des Menschen mit der Umgebung,
So
ein engerer
ein
neuer Lebenstypus unter der Herrschaft des Objekts.
Demnach
sie
unverbrüchlichen
zunächst gegenüber
ganz und gar sich
willig
Schleier ge-
muß es sich zu ihm zurückwenden, ihn um-
aus erscheint mehr und mehr
den Dingen
zu einem
Autonomie und wird
volle
alles entwickelt sich
ihn
fester
nachdem jener trübende
eine
Zusammenhängen
lückenlosen
schen, aber schließlich
ein
Selbst
Wendung.
Konzentration
kräftigerer
enger verbinden und
seine Vielheit
erlangt
als
diese
kann das Objekt seine eigene Natur mit voller Rein-
fallen,
hier
indirekt
unterstützt
den Dingen geliehenen Eigenschaften von ihnen zurück-
und damit der
fordert
muß sich ihr
um seinem Leben Wahrheit zu sichern.
es
ist
nicht eine einzige,
sondern es sind zwei Be-
wegungen, die durch die Neuzeit gehen und auf ihrem Boden ein
gleiches Recht
zweit,
und
ihr
und Unruhe.
die
meisten
behaupten.
Leben
Diesen
der
Solcher Zwiespalt
So
enthält
ist
sie
innerlich bei sich selbst ent-
von Haus aus eine
starke
Spannung
Doppelcharakter der Neuzeit werden
von uns behandelten Probleme erkennen
stellt
auch
lassen.
der geistigen Arbeit eine schwere, aber un-
abweisbare Aufgabe; nicht aus der unmittelbaren Lage heraus, sondern
nur durch ihre Weiterbildung, nur durch Aufdeckung eines neuen
Grundgefüges der Wirklichkeit läßt sich eine überlegene Einheit und
zugleich eine sichere Wahrheit erhoffen.
Daher war
es kein bloßer Eigensinn der Spekulation,
es
war
eine innere Notwendigkeit, welche große Forscher auf neue Bahnen
trieb
und
sie
eine
vom Denken getragene Wirklichkeit dem ersten
Lebens- und Weltbilde entgegenstellen hieß.
sind
als
von Spinoza und Kant;
jener
Von diesen Versuchen
Ausdrücke eines neuen Lebenstypus besonders wichtig die
das Objekt,
um den Gegensatz zu überwinden, verstärkt
dieser das Subjekt,
jenem dringt das Objekt in
Subjektiv — Objektiv.
15
das Subjekt, diesem das Subjekt in das Objekt vor.
und
den Menschen
bindet
die
Spinoza ver-
Welt durch die Aufdeckung einer
Weltkraft im Menschen und ihre scharfe Abhebung von aller bloß-
menschlichen Art; diese Weltkraft
sinnliche
Umgebung befreite,
ist
das von aller Bindung an die
auf sich selbst gestellte und
lediglich
von seiner eigenen Notwendigkeit getriebene Denken, wie z. B. die
Mathematik es zeigt, das Kleinmenschliche dagegen besteht in dem
bloßsubjektiven Fürsichsein mit seinen Affekten und Zwecken.
Die
Wendung von
und
heit
der Dinge und
keit
Die
damit
So ,waren
welche
es
Bewegung zugleich die Wahrund Unendlich-
unmittelbar ihre Ewigkeit
und
Lebens
die Wissenschaft,
Vollendung zu
Klarheit
einem auch die Dinge tragenden Allleben
in
teilt
des
Seele
zur
Befangenheit
so erfaßt es in seiner
ist,
wird
und
den Menschen ein Weltleben gewinnen;
läßt
denn wie das Denken
gegründet
Enge
seiner
Weite des Denkens
von
Erlöserin
die
die Wissenschaft,
die
und künstlerischer Kontemplation
religiöser
Nöten
allen
sich
ihrer
in
gestaltet.
auch vor allem künstlerische und kontemplative Naturen,
Lebenstypus
dieser
mit
herben
seiner
und
Größe
stillen
anzog; aber weit über den Kreis der Anhänger hinaus wirkte diese
Denkweise durch das Scheiden von Kosmischem und Kleinmenschlichem
innerhalb
des Menschen
durch
selbst,
die
energische Be-
kämpfung des Anthropismus des Denkens nicht nur, sondern auch
der
Gesinnung,
der
sich
im
so
Mittelalter
eingenistet
fest
hatte.
Die Kleinheit des gewöhnlichen Glückverlangens und die Enge des
kommen
Vorstellungskreises
landläufigen
zu
jetzt
Emp-
deutlicher
findung; einmal empfunden und als unzulänglich befunden, können
sie die alte Selbstverständlichkeit
in
nun und nimmer wieder erlangen.
Das aber bleibt eine Frage, ob unser geistiges Leben gänzlich
Denken aufgeht, ob nicht die Wendung vom Schein der Sinne
zur Wahrheit
des Denkens
verlangt, die jenseit
selbst
eine Tat
des bloßen Denkens
des ganzen
Auch
liegt.
Menschen
die Voraus-
setzung dieser Lösung: das Zusammenstimmen unseres Denkens mit
der Welt
um uns,
Leben des Alls,
ist
das
Umfangensein beider von einem einzigen
keineswegs zweifellos;
unseres Denkens unsicher wird, da
ist
wo aber der Weltcharakter
sofort die
Wahrheit des ge-
botenen Lebens erschüttert.
Dieses Bedenken
entgegengesetzten
der Dinge
in
wirkte
auch
Weg einschlug.
eine
unzugängliche
Jbei
Denn
Kant,
bei
als
er einen
ihm weicht
Feme zurück, und
es
gerade
die
fällt
Welt
alle
Zum Grundbegriff des Geisteslebens.
16
Möglichkeit,
Übereinstimmung mit ihnen gewiß zu werden.
einer
irgendwelche Wahrheit für uns bestehen bleiben,
Soll
also
sie
innerhalb
des
Subjektes
zum Objekt zu suchen.
von diesem Nein
nicht
selbst,
in
so
ist
einem Verhältnis
Das bedeutet ein entschiedenes Nein, aber
findet sich für Kant ein
er innerhalb unseres Lebenskreises
Weg zu einem Ja, indem
große Gesamtleistungen
aufzeigt,
vornehmlich die Bildung einer wissenschaftlichen Erfahrung und die
Was immer an diesen Leistwird vom Subjekt geleistet; so
eines Reichs
des sittlichen Handelns.
ungen
geistiger Art
von
ist,
das
muß es auch bei sich selbst der herkömmlichen Fassung entwachsen.
Es
nun
ist
sowohl
Einzelpunkt,
geistiges
Gewebe; was
nicht
geistige
Struktur,
Tätigkeit
erfaßt,
individuelle
Existenz,
von
und
es
sich-
gewinnt damit eine Giltigkeit für
alle
als
seiner
Einzelnen;
so entsteht eine neue Art von Objektivität,^ ein neuer Begriff der
Wahrheit.
Sein
näherer Inhalt bestimmt sich nach der Beschaffen-
heit
und der Bedeutung der Tätigkeit,
bei
der
und
theoretischen
menschliche
Erkenntnis
bleibt
bei
er ist daher grundverschieden
der
Vernunft.
praktischen
Alle
nach Kant an eine undurchsichtige
Welt gebunden; die Gedankenwelt, die wir auf ihre Anregung hin
entwickeln,
gilt
nur für uns
selbst
und unser
Vorstellen,
unser
Weltbild reicht nicht über uns hinaus, nicht nur die Formen der
sinnlichen
bloßmenschlicher Art.
das
Denkens sind
und bleiben
Anders auf praktischem Gebiet.
Hier erlangt
Anschauung,
Tun des Menschen
auch
die
des
eine volle Ursprünglichkeit
und vermag es
aus sich selbst eine Welt zu erzeugen; hier, wo das Unterscheidende
in
der Unterordnung aller menschlichen Besonderheit unter allgemeine
Normen liegt, ist die Wahrheit nicht bloßmenschlicher, sondern abArt.
Der Mensch steht hier unmittelbar in den tiefsten
Gründen der Dinge, als moralisches Wesen wächst das Subjekt bei
sich selbst zum Träger einer Welt.
So wird hier die Moral zu
soluter
'
Dieser neue Begriff der Objektivität
ist
freilich
voller
Verwicklung
und wurde von den Gegnern Kants hart angegriffen. So sagt z. B. Plattner,
Philosophische Aphorismen I, § 6Q9 Anmerkung: „Wenn nun aber damit
bewiesen werden
soll,
daß unsere Erkenntnis objektive Giltigkeit hat: so übt
man doch fürwahr an dem Worte Objektiv eine Gewalttätigkeit aus, die
bisher in dem philosophischen Sprachgebrauche unerhört war; denn man
deutet
damit gerade den entgegengesetzen Begriff Subjektiv an. Wirklich
Schmid, der nie von seiner Liebe zur Wahrheit abweicht, in
hat sich Herr
der Notwendigkeit gesehen, die Kantische Objektivität subjektive Objektivität
zu nennen.
Wörterb., Art. Objektiv."
Subjektiv — Objektiv.
17
einem selbständigem Reich und zugleich zum Kern des Lebens, die
und bekommt zur höchsten
Erkenntnisarbeit rückt in die Peripherie
Welt vor Störungen
moralische
Aufgabe,
die
entsteht
damit
ein
zu
bewahren.
zu
dem Spinozas:
dort die
Ruhe der Kontemplation,
ruf
zur Aktivität;
dort
Vordringen
ein
vorhandenen Welt,
hier
Ausgleichung
Gegensätze
aller
das Schaffen
in
eine Spaltung der Wirklichkeit
sätze.
Es
neuer Typus des Lebens in vollem Gegensatz
hier der Auf-
zu den Grundlagen einer
einer
neuen Welt;
dort die
einer allumfassenden Einheit, hier
und
eine Verschärfung aller
Gegen-
Beiden ist aber das Streben gemeinsam, unserem Leben irgend-
wie einen Weltcharakter zu geben,
den Menschen aus sich
selbst
herauszureißen und ihn zu neuen Tiefen zu führen.
Eine Erörterung des Kantischen Denk- und Lebenstypus bleibe
der Betrachtung
der Gegenwart
Leben erweckt hat.
an
dem
Belassen
eines Dinges
menschlichen Vermögens,
sich
fiel
die
die Scheidung
neuem
zu
ihn
als Söhne
und freudigem Lebensgefühl, schweren An-
einer Zeit von kräftigem
stoß-
vorbehalten,
Die unmittelbaren Nachfolger nahmen,
an sich und der Begrenzung
die daraus hervorgeht
Mit dem Ding an
von theoretischer uud praktischer Vernunft,
mehr im Wege, das Leben in einen einzigen
Zusammenhang zu verwandeln. So wurde denn kühnen Mutes
unternommen, alle Wirklichkeit aus unserer geistigen Tätigkeit, im
besonderen aus dem mit innerer Bewegung ausgestatteten Denken
hervorzubringen.
Das Denken, so hatte schon Plotin gezeigt, vermag im eigenen Kreise den Gegensatz von Subjekt und Objekt zu
überwinden, indem es sich gegen sich selbst kehrt, das Denken selbst
und
es
stand
zum Vorwurf
nichts
Denkens macht
Das brauchte nur mit voller
Konsequenz entwickelt, vom bloßen Individuum abgelöst und auf
das Ganze der weltgeschichtlichen Arbeit übertragen zu werden, und
es
entstand
des
das
Hegeische System,
eine
Selbstentwicklung
ein
Selbstbewußtwerden
des Denkens
des
das
die
ganze Wirklichkeit
in
Wahrheit
als
verwandelt,
Geistes
versteht
die
und den Menschen
an dieser absoluten Wahrheit vollauf teilnehmen läßt
alle
Eigenwilligkeit
eines
subjektiven
Nur muß er
Meinens aufgeben und
allein
den Notwendigkeiten des Denkprozesses folgen.
Dies Unternehmen ergriff nicht nur wie ein brausender Sturm
seine eigne Zeit, es hat sich mit seinem Flüssigmachen aller Größen
und seinem Zusammenschweißen
aller
Bestand des Geisteslebens eingegraben.
Eucken Orundbegriffe. 4. Aufl.
,
Mannigfaltigkeit
tief
in
den
Aber auch ein Rückschlag
2
Zum Grundbegriff des Geisteslebens.
18
konnte nicht fehlen, sobald jener kühne Aufschwung nachließ, und
das Ausleben selbst die Grenzen des Unternehmens zur Empfindung
Unabweisbar wurde nun die Frage, ob der Prozeß nicht
brachte.
über
sich
selbst
will
und
es
hinausweist, da er als geistiger wieder erlebt sein
dazu eines überlegenen Punktes bedarf;
unabweisbar
die andere, ob die ausschließliche Verwandlung des Lebens in Denken
der Wirklichkeit
nicht
Gewebe von
ist
weniger durch
als
durch
worden.
und aus
ihr
— Wie immer dem
bloßes
Wendung
des Lebens
wir den Boden,
betreten
der
dem
9. Jahrhundert
hat das Problem
unmittelbarer zur Empfindung wirken
und den Gegensatz mehr
erfährt
es läßt sie
und sich breiter entfalten.
Überwindung aber
gebracht, wie schon das stete
Das Problem
19. Jahr-
Das 19. Jahrhundert.
zum Bewußtsein gebracht als irgendwelche frühere Zeit,
einer
überwunden
zurückgedrängt
ist.
b)
den Versuch
ein
sein mag, das Ganze
wissenschaftliche Gegenarbeit
tatsächliche
Damit aber
1
ist,
eine
hundert eigentümlich
Das
raubt
hier nicht viel zu rasch die menschliche Geistigkeit
zur absoluten erhoben
eine
Inhalt
Formen und Formeln macht; unabweisbar
logischen
ob
endlich die,
allen
hat
es
Für
kaum etwas neues
Zurückgehen auf Kant bekundet.
zunächst den Einfluß der
Wendung von
den Zielen der inneren Bildung zur Beherrschung der sichtbaren
Welt durch Naturwissenschaft, Technik und politisch-soziales Wirken.
Solche Richtung des Lebens heißt den Menschen
stets
den engsten
Anschluß an die Dinge suchen und nur von der Bindung seiner
Kräfte
an
sie
eine
Realität
und Wahrheit erwarten, während
ein
davon abgelöstes Leben zu einem bloßen Schattenreich, einer leeren
Einbildung
sinkt.
So
verlegt
das Leben
seinen
Schwerpunkt
ins
Objektive und findet seinen Kern in der mit den Gegenständen befaßten
und durch ihre Natur bedingten Arbeit; diese Arbeit vollzieht
eine Emanzipation
von den bloßen Individuen, entwickelt bei
sich
Zusammenhänge ausgedehntester Art und macht mit ihrem
unaufhörlichen Anschwellen den Menschen mehr und mehr zu einem
bloßen Diener und Werkzeug. So zunächst in der technischen
Arbeit mit ihren Fabriken, so mehr und mehr auch in den anderen
Lebensgebieten.
Je mehr überall das Sinnen und Streben sich auf
gemeinsame und sichtbare Leistungen richtet, desto nebensächlicher
selbst
Subjektiv — Objektiv.
was
wird,
der Seele des Einzelnen vorgeht, desto gleichgültiger
in
und Befinden, desto mehr wird das Subjekt
erscheint sein Ergehen
am Eimer, zu einer Größe, die sich ohne
zu einem bloßen Tropfen
Schaden zurückstellen und ausschalten läßt.
Ausdruck
findet
das
Einen wissenschaftlichen
der Theorie des Positivismus,
in
Grundsätze folgerichtig durchführt,
ihre
19
sie
soweit sie
nicht mit andersartigen
Gedankenmassen verquickt
Dieser
Zug
überwiegt noch immer im Leben der
Aber
Zeit.
stärker und stärker empfinden wir die Grenzen eines solchen Strebens,
und immer deutlicher steigt das Gefühl der Leere auf; bekundet nicht
schon das ein Wiedererwachen des Subjekts und die Unmöglich-
sich
nach dieser Richtung
damit nahe, das Subjekt beginnt
tritt
seiner Zuständlichkeit
in
das Erste und
als
zusehen, es entwickelt die Neigung,
zugeben,
Ein jäher Um-
Verzichtes auf alle innere Befriedigung?
keit eines
schlag
alles Geschehen
alle
Überlegene an-
Bindung nach außen auf-
nach der Wirkung auf das eigne Befinden
zu messen, schließlich das Leben möglichst in freischwebende Stimmung
zu verwandeln.
Literatur,
So geht es neuerdings in breiten Wogen durch die
das künstlerische Schaffen und auch durch das gesellschaft-
um die Seele zu befriedigen und
Aber
Zusammensein.
liche
Gegner zu besiegen, dazu entbehrt
Gehalts.
eines
ihr
All
Aufrufen
diese
der
Bewegung
individuellen
viel
den
zu sehr
ergibt
Kräfte
keineswegs eine zusammenhängende Innenwelt und eine gemeinsame
Wahrheit;
von der
schließlich
er
befreien
Subjektivismus darf
mittelbar
dieser
wollte.
Weg
in
dieselbe Leere zurück,
Als wissenschaftlicher Vertreter dieses
am ehesten der Psychologismus gelten, der un-
von der Seele des Einzelnen her zu einer Gedankenwelt
Zeitweise hat dies viele überwältigend fortgerissen, aber der
strebte.
Rückschlag
daß
führt
sich
ist
rasch
gekommen und immer
wird
deutlicher
uns,
von dieser schwankenden Grundlage aus zu einer Wissen-
schaft,
zu einem Reich der Wahrheiten nun und nimmer gelangen
läßt^
Auch
mehr
die
jenseit
Schranken
der Wissenschaften
des
Subjektivismus.
erkennen
mehr und
wir
Aber da wir zugleich
zum Objektivismus im geschilderten Sinne unmöglich zurückkehren
können, so verbleiben wir
*
in
peinlichem Zwiespalt,
und
es
drohen
Die durchschlagendste Widerlegung des Psychologismus enthalten die
Logischen Untersuchungen von Husserl, 1900 und 1901; im besonderen wird
hier
überzeugend erwiesen, wie
tief
der Psychologismus auch in die Ge-
dankenwelt solcher Forscher eingedrungen
ist,
die ihn prinzipiell verwerfen.
2*
Zum Grundbegriff des Geisteslebens.
20
und
Arbeit
sich
immer mehr zu
einander
Seele
des Lebens läßt sich unmöglich
solcher Zerfall
Ein
verfeinden.
endgültig hin-
als
muß die Kluft überwunden werden.
An Bemühungen darum fehlt es nicht, am meisten verbindet
nehmen, irgendwie
von innen her so zu verstärken
die Geister ein Streben, das Subjekt
und auszuweiten, daß damit ein neuer Weltdurchblick und ein neues
Leben gewonnen wird. Es geschieht das aber in engem, wenn auch
keineswegs völligem Anschluß an Kant.
sowohl die Theologie
und dort
— es handelt
um die von Ritschi ausgehende Bewegung —
hier namentlich
sich
es gestaltet sich aber hier
In der Theologie
verschiedener Weise.
in
Ein derartiges Streben zeigt
als die Philosophie,
wird unternommen, die religiöse Wahrheit von der Unsicherheit der
Spekulation und
der Metaphysik zu
Wesen der Seele
einen
und
befreien
Grund zu
festen
im innersten
ihr
Vornehmlich
geben.
in
der Moral, in der Ausbildung der sittlichen Persönlichkeit, scheint
das Geistesleben
über
sicher
und
zu erzeugen
eignes Reich
ein
alles
ihm
sich mit
Nach diesem Ge-
andere Dasein hinauszuheben.
dankengange bedarf, was zur geistigen Selbstbehauptung nötig
keiner Bestätigung von außen her,
Vermögen, durch
eignes
sein
Lebens.
es erweist seine Wahrheit
die Steigerung
ist,
durch
des ethisch-religiösen
Die nähere Entwicklung der Gedankenwelt wird hier vor-
nehmlich durch die „Werturteile" bestimmt, die jene Beziehung zum
Kern
des Lebens
vertreten
und
damit
werden;
das
moralischreligiöse
führung überlegen
nach
theoretischen
aller
ihm innewohnenden Notwendigkeiten
den
nur
in
steter
Ganzes
ein
Überzeugungen, das freilich keine Welterklärung sein
Geltung
Beweis-
Leben entwirft
will
von
und seine
Zurückbeziehung auf die Grundwahrheiten
jenes Lebens hat.
Dieses Streben,
verschiedene Färbungen
es
die
barer
begründen
schehen kann,
denken hervor.
handelt
Gefühl
will,
die nähere
Als
sich selbst
insofern gewiß in
ist".
III,
Kern
(Ritschi,
142).
als
des Menschen fester und unmittel-
manche Be-
Lösung dieser Aufgabe
ruft
des Lebens
das Gefühl be-
ein
pflegt
Christi.
Aber
hier
Aufbau versucht zu werden.
nun einmal die geistige Funktion,
Versöhnung
gutem Recht,
Leben in höherem Grade einen Tatcharakter
und von ihm aus
ist
ist
das durch theoretische Erwägungen ge-
als
als es dem
Aber
verleiht.
hat,
Überzeugungen
letzten
Durchführung übrigens recht
nähere
dessen
in
„Das
welcher das Ich bei
Lehre von der Rechtfertigung und
läßt
in
Wahrheit das Gefühl
schon
Subjektiv - Objektiv.
ein Beisichselbstsein des Lebens erreichen?
und
flach
leer
Es erzeugt doch
sein?
sondern es gewinnt ihn nur
Inhalt,
des Lebens.
artigsten
nicht
von
weiteren
nicht
sich aus einen
Zusammenhängen
Wie das Gefühl in stetem Fluß und der verschieden-
Deutung
dem Leben
in
21
Kann das Gefühl
ein
fähig
so läßt sich von ihm aus unmöglich
ist,
wie
Halt
ein
Inhalt
geben.
Der Versuch
eines
Aufbaues einer Gedankenwelt vom Gefühl und damit vom Subjekt
würde
sich
vom bloßen Subjektivismus kaum unterscheiden, wenn
sich nicht das Gefühl als
als
der
notwendig und der von ihm bejahte Inhalt
Besonderheit
natürlichen
des Menschen
dartun
überlegen
Wie aber sollte das von der bloßen Tatsächlichkeit des Seelen-
ließe?
Mag ein Gefühl noch so unabweismag
lebens her geschehen können?
bar scheinen, es
ist
das zunächst nur für das besondere Subjekt;
einem bestimmten Inhalt noch so verwachsen scheinen, diese
Verbindung besagt mehr als der unmittelbare Eindruck enthält, sie
ist das Ergebnis einer Deutung, und eine solche Deutung kann irre
es mit
So
gehen.
leistet
die Stärke eines Gefühls nicht die mindeste
Ge-
währ für die Wahrheit des aus ihm entwickelten Gedankengehaltes.
Das bekundet u. a. die Vielheit und der Streit der Religionen. Jede
von ihnen glaubt
ihres
sein als die anderen,
Grundgefühles mindestens ebenso sicher zu
und doch gelangen sie zu grundverschiedenen
notwendig einer höheren Instanz,
um
über das Recht der verschiedenen Ansprüche zu entscheiden,
und
Wahrheiten.
diese
So bedarf
es
kann nicht das Gefühl
Menschen aus zu
Überhaupt aber läßt sich vom
sein.
einer Wahrheit
nur gelangen, wenn
in
ihm
ein
seiner natürlichen Besonderheit überlegenes Leben durchbricht; eine
an jene Besonderheit geknüpfte Wahrheit
ist
keine Wahrheit.
Schon
das bringt es mit sich, daß der Mensch das Problem seines Grundverhältnisses zur Wirklichkeit
nur zurückschieben kann.
träglich
Art
aufgedrängt,
Das Leben
es
nun und nimmer aufgeben oder auch
Dies Problem wird ihm nicht
gehört von
erst
nach-
vornherein zu seiner geistigen
eines geistigen Wesens erschöpft sich einmal nicht
in die bloße Zuständlichkeit, es
umspannt auch das Gegenständliche
und muß sich mit ihm auseinandersetzen, es muß auf einer Überwindung jener Spaltung zwingend bestehen, während ihm die Beschränkung
auf
die
bloße Zuständlichkeit zu einer unerträglichen
Enge wird.
Was hier an Verwicklung vorliegt, das verdunkelt und vergißt
sich namentlich deshalb leicht, weil die
Regung des Gefühls durch
Zum Grundbegriff des Geisteslebens.
22
eine geschichtlich überlieferte Gedankenwelt ergänzt zu werden pflegt,
der Halt scheint damit fester und der Inhalt reicher zu werden.
ist
zu
und das kann
erweisen,
Gefühl
geschehen;
her
In
die Wahrheit der geschichtlichen Überlieferung erst
der Tat aber
diesem Zusammenhange nur
in
ebenso
könnte
auch nur
dieses
vom
darüber
entscheiden,
was vom Gehalt jener Überlieferung
gelten hätte.
So kommen wir hier auf Umwegen immer wieder auf
als
wertvoll
das Gefühl zurück und bleiben an seinen Kreis gebannt.
zu
mehr
Je
es aber auf sich selbst gestellt wird, desto weniger Inhalt gewährt es,
desto mehr droht es sich in einzelne zerstreute und sinnlose Vorgänge
aufzulösen.
So führt dieser Weg mehr in die Verwicklung hinein als aus
ihr heraus.
Was immer aber uns von der hier gebotenen Gedanken-
entwicklung trennt, es kann uns nicht verhindern, die entschiedene
Kräftigung des ethisch-religiösen Lebens anzuerkennen, die mit jenem
Streben
erreicht
ist;
nur können wir die wissenschaftliche Fassung
dessen nicht glücklich finden.
dem Gebiet der
wo der Begriff des Wertes^ den Mittelpunkt einer be-
Wesentlich anders gestaltet die Sache sich auf
Philosophie,
deutenden und fruchtbaren Bewegung bildet
^
Über den Begriff und
die
Diese Bewegung vertritt
Bedeutung des Wertes ist in den letzten
deren Betrachtung und
Würdigung hier nicht wohl möglich ist; nur das Werk Meinongs „Psychologisch-ethische Untersuchungen zur Wert-Theorie", 1894, möge hier angeführt
Wünschenswert wäre eine Gesamtgeschichte des Wertproblems und
sein.
Jahrzehnten eine ausgedehnte Literatur entstanden,
Wertbegriffes;
nur aus Höffdings Religionsphilosophie folgende
„Der Philosophie Kants verdanken wir die Selbständig-
hier seien
Stellen hervorgehoben
:
keit des Wertproblems
dem Erkenntnisproblem gegenüber.
Er lehrte uns die
Schätzung von der Erklärung unterscheiden." (S. IL) Femer „Kant redet häufiger
von Zwecken
von Werten. Es ist aber klar (obschon Kant dies weder
noch in seiner Ethik recht beachtete), daß der Begriff
des Zweckes den Begriff des Wertes voraussetzt, da ich zum Zwecke nur das
machen kann, dessen Wert ich erfahren habe. Wenn Kant von dem Reiche
der Zwecke' im Gegensatz zur kausalen Naturordnung redet, so meint er
hiermit dasselbe, was spätere Philosophen ,das Reich der Werte' nannten.
Der Kantianer Fries geht von dem Begriff des Wertes aus (System der Philoals
in seiner Psychologie,
,
sophie, Leipzig 1804; §§ 238, 255, 330.
berg 1807.
III.
S. 14).
— Neue Kritik der Vernunft. Heidel-
Vorzüglich sind es aber Herbart und Lotze, die dem
Begriffe des Wertes in weiteren Kreisen Eingang verschafften.
Nach Lotze
nahmen der Theolog Albrecht Ritschi und dessen Schüler denselben auf."
Über den Wertbegriff bei Fries ist neuerdings eine Schrift von Pöschmann
erschienen.
lich
modern
Es ist aber der Begriff mit seinem Problem keineswegs ausschließ;
er
tritt
hervor, sobald das Subjekt eine größere Selbständigkeit
Subjektiv - Objektiv.
23
Ganzes angesehen die moderne Denkweise gegenüber der antiken,
Wo den Hauptgegen-
als
namentlich sofern diese durch Plato bestimmt ist.
satz der Wirklichkeit der des beharrenden Seins und des flüchtigen Wer-
dens bildet, wie bei diesem, da liegt es nahe, das wesenhafte Sein zugleich
als das Gute und Wertvolle zu fassen, beide Begriffe möglichst zusammen-
zuschmelzen. Diesem Gedankengange kann das Gute als etwas von der
Tätigkeit Abgelöstes,
dem Menschen Gegenüberliegendes erscheinen.
Die moderne Denkweise verficht dagegen, daß von einem Guten nur
in
Beziehung auf ein lebendtätiges Wesen die Rede sein kann, und
daß nur nach der Bedeutung für dieses sich die Schätzung näher bestimmen läßt; damit empfahl es sich, statt von Gütern von Werten zu
Dieser Grundgedanke kann aber sehr verschiedene Fassungen
reden.
annehmen und
hat das in Wahrheit getan.
Gilt als der Träger des
Lebens lediglich das empfindende und fühlende Einzelsubjekt, und
demnach der Wert der Geschehnisse nach der Leistung
und Behagen bemessen, wird Lust und Unlust
zum entscheidenden Maßstab gemacht, so sieht man nicht, wie davon
wird
sich*
für dessen Befinden
Bewegungen oder gar Erhöhungen des Lebens ausgehen
könnten. Denn die Lust schmiedet den Menschen an die eigne dunkle
fruchtbare
Zuständlichkeit,
sie
macht das Leben innerlich eng bei
aller
Weite
Erhöhung des Wesens, alle
unmittelbare Freude an Menschen und Dingen, alles Aufnehmen des
nach außen,
sie
verwehrt
Gegenstandes
in
den Lebensprozeß.
lich
alle
derjenige schwer nehmen,
innere
Diesen Mißstand wird nament-
der in
dem
seelischen Stande des
Menschen große Aufgaben und Verwicklungen erkennt Denn eine
solche Lage bringt die Forderung eines kräftigen Emporklimmens,
ja einer inneren
Umwälzung mit sich
;
diese aber sind ausgeschlossen,
wenn das Leben starr an den bloßen Zustand gebannt bleibt.
Unvergleichlich
kritisch-idealistische
versetzt
und
bestrebt ist ^
gewinnt.
bildeten.
sie
viel
höher
steht
eine
andere Art, welche die
Denkweise Kants auf den Boden der Gegenwart
den Erfahrungen der Zeit gemäß weiterzuführen
Hier bildet die Tatsache den Ausgangspunkt, daß unser
So zuerst bei den Stoikern, die auch einen Terminus (a^ia) für ihn
Nikolaus von Kues, der erste moderne Deiiker, nennt Gott den
Wert der Werte (valor valorum).
^
Einen besonders klaren und bedeutenden Ausdruck findet diese Be-
wegung in den „Präludien" Windelbands, namentlich in den Abschnitten
„Was ist Philosophie?", „Normen und Naturgesetze", „Kritische oder genetische
Methode".
Zum Grundbegriff des Geisteslebens.
24
Leben und Tun nicht in eine bloße und blinde Tatsächlichkeit des
Geschehens aufgeht, sondern
daß unsere
uns zu
geistige Art
steter
Beurteilung zwingt; diese Beurteilung aber erfolgt nach bestimmten
Normen,
die
Willkür überlegen und auch von der Verwirk-
aller
lichung beim Menschen unabhängig sind.
Werte
ersichtlich,
Lust
und Unlust
und
vollziehen
die
jenseit
liegen,
für
sich
die
eine
In diesen Normen
werden
alles
bloßen
Nutzens,
eine
innere
Erhöhung des Lebens
jenseit
aller
Unbedingtheit in Anspruch nehmen
dürfen. ^ So erkennen wir hier ein bedeutendes Streben, dem Leben des
Menschen von innen her einen Halt und einen Inhalt zu geben, ihn
durch
machen,
zu
ohne ihn
in
der Mensch
die Fährnisse
zugleich
verwickeln,
Aufgabe abzustecken.
anderen Wege
dem
Selbstbesinnung
kritische
als
Naturgetriebe
überlegen
zu
einer spekulativen Metaphysik
auch der Philosophie
eine
eigentümliche
In der Tat ist nicht zu ersehen,
wie auf einem
dem der Selbstbesinnung und der Selbstvertiefung
durch die Verfeindung von Subjekt und Objekt
die
drohende Spaltung des Lebens überwinden könne.
Was uns Bedenken erregt,
gewiesenen
hier
Stelle
ist
selbst mit innerer Notwendigkeit
ob
ob man an der
nicht die
über sie hinaustreibe.
Fragen mögen sich dabei erheben.
Erlebnisse die
lediglich dieses,
abschließen könne,
Bewegung
Verschiedene
Können die Werte als einzelne
genügende Sicherheit erlangen, wird
nicht ihre Ur-
sprünglichkeit anfechtbar sein, so lange sie ein bloßes Nebeneinander
bilden, sich nicht zur Einheit eines Ganzen zusammenfassen ? 2
ferner die in den
Werten
eröffnete höhere Stufe des
Wird
Lebens gegen
die bestrickende
Macht der natürlichen und sozialen Selbsterhaltung
aufkommen und
sich
durchsetzen
können, wenn
so
faltet
behauptet?
sie
uns nicht ein
den Werten wie
ent-
Dieses aber wird schwerlich ohne eine
Um-
neues geistiges Selbst verschafft, das sich
kehrung der vorgefundenen Lage möglich
in
sein,
und das führt doch
der Werte vertritt namentlich Münsterberg in
Werte" (1908) mit großer Kraft und Wärme.
" Die Notwendigkeit einer solchen Verbindung hebt auch Münsterberg
nachdrücklich hervor; er sagt im Vorwort zu seiner „Philosophie der Werte":
„Die Gesamtheit der Werte muß grundsätzlich geprüft und aus einer Grund^
Diese
Überlegenheit
seiner „Philosophie der
tat
einheitlich
abgeleitet werden.
fehlt, ist ein in sich
Das, was unserem Philosophieren heute
geschlossenes System der reinen Werte; erst dann kann
die Philosophie auch wieder aufs neue zur wirklichen Lebensmacht werden,
wie es zu lange ausschließlich die Naturwissenschaft gewesen
ist"
(VI).
Subjektiv
— Objektiv.
25
mag sie von
wieder zu irgendwelcher Metaphysik,
der alten noch
so verschieden sein.
So erblicken wir in der Lehre von den Werten weniger einen
Abschluß
Bewegung.
Über die
Bewegung wenig hinaus, und
die Menschheit verbleibt in dem peinlichen Hin- und Herschwanken
zwischen Arbeit und Seele, zwischen der Absorbierung des Subjekts
fertigen
Philosophie
als
kommt
eine
aussichtsreiche
einstweilen diese
durch das übermächtige Objekt und der Verflüchtigung des Objekts
durch die Selbstherrlichkeit des Subjekts.
Diese Lage mit ihrer Verwicklung treibt die Frage hervor, ob
ganze Scheidung von Subjekt und Objekt, ob nicht
nicht die
Anerkennung
inneren
eines
neben
Bereiches
alle
Außenwelt von
der
Haus aus verfehlt sei, ob nicht bei solcher Fassung das Wahrheitsstreben den unlösbaren Widerspruch enthalte, zugleich scheiden und
Von
verbinden, auseinanderhalten und zusammenführen zu wollen.
entgegengesetzten Ausgangspunkten
her sind
neuerdings Avenarius
und Mach zu dem gleichen Ergebnis gekommen, jene Scheidung als
unnütze und irreleitende Verdoppelung aufzugeben.
Die Ver-
eine
setzung der Empfindungen in ein Inneres, die Introjektion, erschien
ebenso
verfehlt,
wie die Herausstellung von Bewußtseinsvorgängen
nach außen, die Projektion.
eine
einzige,
und
es verbot
Erfahrung zu
mittelbare
Statt
sich
zweier Welten ergab sich damit
jedes Hinausgehen
Dingen. ^
jenseitigen
Wendung des Problems, deren
Streben
über die un-
Diese
scharfsinnige
nach Vereinfachung einen
sichtlichen Eindruck auf die Zeit macht, nach ihrer technischen Seite
zu
prüfen,
^
liegt
außerhalb
Aufgabe;
unserer
daß
sie
2.
Aufl.
Siehe Mach, Die Analyse der Empfindungen.
gibt keine Kluft zwischen Psychischem
und Physischem,
keine Empfindung, der ein äußeres, von ihr
Draußen,
auf
ihrem
S. 206:
«Es
und
verschiedenes Ding
kein Drinnen
Es gibt nur einerlei Elemente, aus welchen sich das vermeintliche
Drinnen und Draußen zusammensetzt, die eben nur, je nach der temporären
entspräche.
Betrachtung, drinnen oder draußen sind.
— Die sinnliche Welt gehört dem
physischen und psychischen Gebiet zugleich an."
S. 33:
»Ich sehe keinen
Gegensatz von Psychischem und Physischem, sondern einfache Identität
bezug auf diese Elemente."
Siehe
auch Wlassak
Nr. 18, S. 202): »Kein naiver Mensch findet einen
findung
in
seinem Bewußtsein,
(in
in
der »Zukunft" 1902,
Baum irgendwie als Emp-
sondern immer nur
als
Bestandteil seiner
auch dann, wenn der Baum nicht gesehen, sondern
nur erinnert wird; auch das blasse Gedankenbild steht in keinem anderen
Umgebung.
Dies
gilt
Verhältnis zum Beschauer als der gesehene
Baum."
Zum Grundbegriff des Geisteslebens.
26
nächsten Gebiet
der
Wahrnehmungs-
physiologisch-psychologischen
lehre scheinbar abgeschlossene Fragen wieder in Fluß bringt und den
problematischen Charakter des landläufigen wissenschaftlichen Naturbildes aufdeckt,
ist
sicherlich ein Verdienst.
Der Grundbehauptung
aber beizutreten verbietet uns schon die Erwägung,
in Wirklichkeit
daß schon
unser
Erkennen
daß namentlich aber
jenseit
Innenleben
entwickelt,
durch
Wechsel
allen
erweist.^
daß unser Ich
mehr ist als ein Strom von sinnlichen Empfindungen,
ein
selbständiges
aller
intellektuellen
gegenüber
das
Verarbeiten
enthält,
Vorgänge
sich ein
Mannigfaltigkeit
aller
und Wandel hindurch
eine
und
beharrende Art
Für solche Selbständigkeit des Inneren spricht auch das
Ganze der
geschichtlichen
Bewegung; denn durch
alle
Arbeit
und
Verwicklung hindurch hat sich der Mensch immer weiter von der
ein
immer mehr das äußere Begegnis in
inneres Erlebnis verwandelt, immer mehr Gegenwirkung gegen
die
zuströmende Fülle geübt.
bloßen Sinnlichkeit entfernt,
Wenn Mach
^
Das alles ist kein bloß
das Ich als etwas Unbeständiges
intellektuelles
und Unselbständiges
behandelt, so liegt das zum guten Teil an einer Vermengung von Bewußtsein
des Ich
und lebendigem
Ich selbst.
So heißt
es
z.
B.
a.
a.
O. S. 3: »Die
scheinbare Beständigkeit des Ich besteht vorzüglich nur in der Kontinuität,
in der
langsamen Änderung.
Die vielen Gedanken und Pläne von gestern,
welche heute fortgesetzt werden, an welche die
Umgebung im Wachen fort-
während erinnert (daher das Ich im Traume sehr verschwommen, verdoppelt
sein oder ganz fehlen kann), die kleinen Gewohnheiten, die sich unbewußt
und unwillkürlich längere Zeit erhalten, machen den Grundstock des Ich aus.
Größere Verschiedenheiten im Ich verschiedener Menschen, als im Laufe der
Jahre in einem Menschen eintreten, kann es kaym geben. Wenn ich mich
heute meiner frühen Jugend erinnere, so müßte ich den Knaben (einzelne
wenige Punkte abgerechnet) für einen anderen halten, wenn nicht die Kette
S. 17: „Man wird dann auf das Ich, welches schon
der Erinnerung vorläge."
während des individuellen Lebens vielfach variiert, ja im Schlaf und bei Versunkenheit in eine Anschauung, in einen Gedanken, gerade in den glück-
ganz fehlen kann, nicht mehr den hohen
Aber besteht denn nicht mit lebendiger Kraft eine Einheit
geistiger Art gegenüber allen Wandlungen und Verdunklungen des Bewußtseins, und wirkt nicht aus solcher Einheit geistiger Individualität alles vordringende wissenschaftliche und künstlerische Schaffen, entspringt aus ihr nicht
alle durchgreifende Leistung auch auf praktischem und technischem Gebiete?
lichsten Augenblicken, teilweise oder
Wert
legen."
Diese
Erfahrungen
des
geistigen
Lebens bestätigen gegenüber jener Ver-
flüchtigung des Ich vielmehr die Überzeugung Goethes:
„Und keine Zeit und keine Macht zerstückelt
Geprägte Form, die lebend sich entwickelt."
— Objektiv.
Subjektiv
Phänomen,
kein bloßer Erklärungsversuch,
reicher Tatsächlichkeit, der nächsten
und
27
sondern eine Eröffnung
sichersten, die wir kennen,
die allein uns die sinnlichen Eindrücke denken und überdenken lehrt.
So wenig
erklären
sich
eine
solche Tatsächlichkeit für eine bloße Illusion
und das Rad der Weltgeschichte zurückdrehen läßt, so gewiß
behauptet jene Scheidung zwischen Subjekt und
zwischen
Objekt,
Innenwelt und Natur eine unabweisbare Notwendigkeit.
c)
Die positive Behauptung.
a.
In
Einführung.
welcher Richtung sollen wir nun weiter suchen?
Ist
die
Scheidung nicht zurücknehmbar und führt kein Weg von der einen
Seite zur anderen, so bleibt keine andere Möglichkeit, als den Gegensatz in
den Lebensprozeß
selbst
aufzunehmen, diesen von innen her
so zu erweitern, daß er sich nicht erst nachträglich auf eine neben
ihm befindliche Welt bezieht, sondern daß er selbst eine Welt enthält;
innerhalb
des
Menschen
selbst
muß
Ganze
das
Welt zur
einer
Wirkung kommen, die dem Gegensatz überlegen ist, und diese Welt
muß nicht durch
die Besonderheit des Punktes hindurch,
unmittelbar uns zugänglich
sein;
sondern
dann und nur dann kann es für
den Menschen irgendwelche Wahrheit geben.
Diese
in
Wendung mag beim
Wahrheit
die
fehlt
es
ihr
nicht
Anblick auffallend dünken,
ersten
an geschichtlichen Anknüpfungen,
um
nur einer Zusammenfassung bedürfen,
in
dem
scheinbar
Neuen Altes entdecken zu lassen. Wie kam denn die Menschheit
dazu, die Begriffe des Wahren und des Guten zu bilden und sie von
der bloßen Tatsächlichkeit und Nützlichkeit abzuheben, wie konnte
sie
überhaupt über die Meinungen und Neigungen des bloßen
Menschen irgend hinausstreben? Ein merkwürdiges Phänomen ist
Denn mag noch
hier
nicht zu
sein,
was als wahr urtd als gut zu gelten habe, vor aller Ungewißheit
verkennen.
so
der Antwort steht die Tatsache der Frage, und diese
etwas Großes
Denn
und Folgenreiches.
brechung des bloßpunktuellen Seins,
des Wesens, die in
sie
sie
Streit
viel
ist
enthält
darüber
unmittelbar
eine
Durch-
bezeugt eine innere Weite
dem scheinbar Fremden etwas Eigenes sieht und
sucht Sicherlich nämlich kann sich der Mensch
kümmern und bekümmern, was
nicht
um nichts ernstlich
irgendwie zu seinem Leben
Zum Grundbegriff des Geisteslebens.
28
und Wesen
Beziehung
in
vöUiges Jenseits kann
ihn
schHeßlich zu
ihm gehört;
steht,
ja
nicht
im mindesten erregen.
beim Wahren und Guten das Streben auf eine Welt,
des nächsten Lebenskreises
nicht die
einem
an
auch
aus
sich
Bewegung
viel
muß
Kreise teilhaben,
Welt umspannen, wenn
uns üben, so
die
jenseit
müssen wir dann nicht von Haus
liegt;
weiteren
ein
Nun geht
unser Leben
Anziehung auf
ihr Inhalt so viel
uns erzeugen soll?
in
Freilich
muß
dann der Begriff unseres Selbst verändern, aber die Begriffe haben
den Tatsachen und nicht die Tatsachen den Begriffen zu dienen, warum
sollten wir
uns also gegen eine solche Veränderung sträuben?
Eine schwere Frage und Aufgabe bleibt es
danken einer Weltnatur des Menschen
eine
jenem Ge-
freilich,
präzisere
Fassung zu
geben, aber auch dafür haben die letzten Jahrhunderte auf der Höhe
Leistungen
ihrer
Hauptstück der
deutlich genug einen Weg gezeigt.
Es ist ein
Größe Kants, von der bloß psychologischen Er-
klärung die Erforschung der Möglichkeit geistiger Inhalte abzuheben,
z.
von der Frage, wie der einzelne Mensch zum Erkennen, zur
B.
Moral
u. s.
w.
kommt,
die zu
unterscheiden,
welche inneren Be-
dingungen das Bestehen von Wissenschaft und Moral
sowohl die logische
als die ethische
über der psychologischen.
hat.
So wird
Betrachtung selbständig gegen-
Das mag zunächst bloß als eine neue
hinfällig ohne ein
Methode erscheinen, aber diese Methode wäre
Leben
neues Leben,
ein
Daseins,
Leben aus dem Ganzen,
charakter.
ein
jenseit
der Einzelvorgänge des seelischen
ein
Leben mit einem Welt-
Was sich aber von- solchem Leben nach einzelnen Rich-
tungen entwickelt, das hat keinen festen Grund und Halt, wenn es
sich nicht zu
einem Ganzen zusammenfaßt und als Erweisung einer
neuen Stufe anerkannt wird,
die
nicht unter,
Gegensatz von Subjekt und Objekt
sondern über
dem
steht.
In anderer Art und doch zu verwandtem Ziele wirkt die moderne
Kunst auf der Höhe ihres Schaffens.
Die Objektivität eines Goethe
wird gefeiert, er selbst begrüßte es mit freudigem Dank,
sein
Denken
ein gegenständliches nannte.
lichkeit bedeutet aber keineswegs eine
Aufsaugung des Subjekts durch
als
Heinroth
Eine solche Gegenständ-
Unterdrückung und möglichste
das
Objekt,
ein
bloßes
Wieder-
geben des äußeren Eindrucks der Dinge, sondern eine Begegnung
und gegenseitige Durchdringung von Objektivem und Subjektivem
auf dem gemeinsamen Boden des Innenlebens; die Dinge empfangen
dabei selbst eine Seele und vermögen ihre eigene Art getreulich mit-
Subjektiv
das
zuteilen;
menschliche
Leben
zu einem Inhalt geführt.
Leere
— Objektiv.
29
aber wird von der anfänglichen
Hier wird nicht
eine
subjektive
Stimmung den Dingen aufgedrängt, sondern ihnen ihr eigenes Leben
und Weben abgelauscht oder abgerungen; der Dichter „erscheint
damit wie ein Zauberer, der die sonst stummen Wesen zum Sprechen
bringt,
dem
öffnet,
der
ganze Unermeßlichkeit der Welt seelisch
die
sich
alle
ihrer eigenen
Mannigfaltigkeit
er-
Natur zuführt und
zugleich das Lebendige, Wesentliche, Wirksame aus den Dingen heraussieht"
Lebensanschauungen der großen Denker,
(s.
7. Aufl.,
446).
Goethe nennt das eine Synthese von Geist und Welt, „die von der
ewigen Harmonie
des
Wahrheit
diese Synthese
erfolgt
sondern
Außenwelt,
Daseins die seligste Versicherung gibt«;
innerhalb
nicht zwischen der Seele
der
So bestehen nicht
bloß zwei, sondern drei Arten künstlerischen Schaffens:
satz einer objektivistischen
eine
sich
stellt
und
überlegene
Art
und der
zu einer Innenwelt erweiterten
zwischen Seiten und Polen ihres Lebens.
Seele,
in
einer
subjektivistischen
entgegen,
die
dem GegenBehandlung
wir eine souveräne
Erst diese souveräne Behandlung dringt sowohl über die
nannten. ^
seelenlose Gegenständlichkeit, als über die formlose Zuständlichkeit
zu einer Eigenständlichkeit vor, bei welcher der Lebensprozeß eine
Welt nicht nachträglich sucht, sondern
damit
gewinnt er einen
erst
sie
aus sich selbst
Inhalt, nicht als
entfaltet;
Abbildung eines vor-
handenen Daseins, sondern durch schöpferische Synthese einer neuen
Welt.
hat,
Sollle,
was so
in der
Kunst eine unbestreitbare Wirklichkeit
nicht auch für das Ganze des Geisteslebens gelten, könnte über-
haupt die Kunst sich darum bemühen, wenn nicht hinter ihr ein Ganzes
So sollten wir getrost den hier gewiesenen
und ihn mutig zu Ende gehen, mag er uns noch
des Geisteslebens stünde?
Weg
verfolgen
so weit von
daran
ist
dem üblichen Welt- und Lebensbilde entfernen.
nicht zu zweifeln,
Denn
daß nur im Widerspruch damit sich eine
Welt von innen her aufbauen und das Leben und Wirken eigentümlich gestalten
des
begriff
läßt.
Erwägen wir also, wie
Geisteslebens,
2.
das
Verhältnis
der Grund-
sich
1.
des
Menschen zur
und zugleich der Anblick des geschichtlichen Lebens,
3. das Problem der Wahrheit ausnimmt, wenn wir jenen Weg be-
Geistigkeit
treten
und damit
die Voraussetzungen
dessen entwickeln,
Ergebnis jeder irgendwie festhalten muß.
^
S.
„Wahrheitsgehalt der Religion".
2. Aufl.,
S. 95.
was im
Zum Grundbegriff des Geisteslebens.
30
ß.
Der Grundbegriff des Geisteslebens.
Ein Leben geistiger Art
es
als das,
gilt
was den Menschen aus-
So muß
und über die Stufe des bloßen Tieres hinaushebt
zeichnet
wohl mehr
sein als das natürliche, uns mit den Tieren gemein-
same Seelenleben;
Wahrheit zeigt schon eine flüchtige Betrachtung
in
Das Seelenleben der niederen
mehr als eine Begleiterscheinung und eine Hülfe des
Naturprozesses; alle Entwicklung von Intelligenz und Geschicklichkeit bleibt hier ein bloßes Werkzeug zur Erhaltung des Individuums
oder der Gattung; als solches Werkzeug gelangt es nicht zu einem
inneren Zusammenhange, einem sicheren Beisichselbstsein, einem
alsbald einen wesentlichen Unterschied.
Stufe
nicht
ist
eigentümlichen
Dies
Inhalt.
aber
es,
ist
was die Wendung zur
Hier erscheint ein neuer Lebens-
geistigen Stufe des Lebens bringt.
prozeß: das Innere, bisher eine bescheidene Zutat, ein Anhang einer
fremden
Welt,
will
auf
jetzt
sich
und
selber stehen
eigne
eine
Zu einem Ganzen verbunden wäre demnach
Wirklichkeit bilden.
geistiges Leben selbständig
gewordene und zu einem Inhalt gelangte
Innerlichkeit; die sonst in eine unermeßliche Vielheit zerstreute
in
lauter
Abhängigkeitsverhältnisse
verstrickte
Wirklichkeit
und
erreicht
Zusammenhang und ein Leben, das allererst ein
hier einen inneren
Selbstleben heißen kann.
Wird
Aber aus solcher Behauptung erwächst sofort eine Frage.
mit jenem Selbstleben ein Sonderreich
lichkeit der
Dinge
liegt,
zufrieden
erstrebt,
und
das neben der Wirk-
oder
sicher bei sich selbst,
verbleibt auch in jener Wendung ein Zusammenhang mit der großen
Nur dies letztere entspricht dem Befunde des Lebens.
Welt?
im Streben zu
sich
selbst
bleibt
großen Welt befaßt,
es
sich zu ziehen, es
kann nicht ruhen und rasten,
und
überwunden
in
Denn
das Geistesleben zugleich mit der
kann sich selbst nicht finden, ohne diese an
sich
bis
es sie vollauf
Darum
aufgenommen hat
ist
all
sein
Gehalt zugleich eine Behauptung, die Behauptung, das Letzte, Ganze,
Allumfassende, der Kern der gesamten Wirklichkeit zu sein.
aber
kann
es
Drngen durch
nur
sein,
wenn
die Weiterbildung,
die es an
Dies
den
die Aneignung bewirkt, diese zur Höhe ihres eigenen
Wfesens führt, wenn der Gehalt des Geisteslebens die eigene Wahrheit der
Dinge bedeutet
unerträglicher
Das Geistesleben wird
Widerspruch,
wenn
es
neben
in sich selbst ein
und gegenüber der
Subjektiv
nicht innerhalb ihrer steht,
Welt,
ihm
— Objektiv.
wenn
31
Wendung zu
nicht in der
sich die Wirklichkeit selbst vollendet.
Die Anerkennung dessen
verwandelt
sie in
ein
in
Fluß und
Reich von aufsteigender Bewegung.
Die An-
unsere Welt
versetzt
fangsstufe bildet die Natur, aus ihr quillt das natürliche Seelenleben
Aber
auf.
Leben
dies
durchgängigen
einen
bildet
indem es eine gewisse Innerlichkeit entwickelt und
die
völlige
Bindung an
Selbstlebens wieder aufhebt
sie zugleich
Diesen Widerspruch
und
Stärke des Lebensaffektes gehalt-
durch
die Versagung alles
durch
Äußeres,
ein
Widerspruch,
das
stellt
aller
in
sinnlose Getriebe der Tierwelt
jedem denkenden Beobachter eindringlich,
ja erschütternd
vor Augen.
Erst
im Geistesleben beginnt eine Lösung des Widerspruches, indem
sich
nun das Leben
gegen sich selber
mehr bloß nach außen hin, sondern auch
nicht
kehrt.
Als eine solche Stufe des Alls kann das Geistesleben nicht eine
und
bloße Eigenschaft einzelner Punkte sein
erst
nachträglich aus
einzelnen Betätigungen zu einem Ganzen zusammenschießen, es muß
vielmehr von Haus aus ein Ganzes, ein selbständiges und sich selbst
angehöriges Leben sein.
Mannigfaltigkeit
aller
so
Zu einem solchen Ganzen gehört eine wie
auch dem Gegensatz von Subjekt und
Objekt überlegene Einheit.
Dies Ganze entwickelt sich mittels des
Gegensatzes von Subjekt und Objekt,
aber
es
bleibt
ihm überlegen und
von Kraft und Gegenstand,
hält
beide Seiten
auch
in
der
Scheidung zusammen, auf geistigem Boden kann jede einzelne sich
nur zusammen mit der anderen
entfalten
und
ihre
eigene
Höhe
So sind hier nicht sowohl die beiden Seiten einander ent-
finden.
gegengesetzt, als vielmehr der Stand ihrer Einigung, der Stand der
Volltätigkeit
dem der Spaltung, dem des halbseitigen und zugleich
Das bloße Subjekt ist vom Geistesleben aus ange-
leeren Lebens.
sehen ebenso etwas äußerliches wie das Objekt; nicht die Beziehung
der
einen
auf
Seite
die
andere,
sondern
nur die schöpferische
Synthese erzeugt eine Innerlichkeit und zugleich eine
selbst
befindliche Wirklichkeit;
eine solche
volle,
bei sich
kann nie von draußen
dargeboten werden.
Es hat aber solche Überwindung des Gegensatzes von Subjekt
Bedingung und Voraussetzung: die
Betrachtung darf ihren Ausgangspunkt nicht in einem gegebenen
Geschieht
Sein, sie muß ihn im Lebensprozeß selber nehmen.
und Objekt
nämlich
eine
jenes,
so
unerläßliche
wird
entweder
die
Welt oder das Subjekt
als
Zum Grundbegriff des Geisteslebens.
32
etwas für sich bestehendes und
dann aber
läßt sich
bei
sich abgeschlossenes festgelegt;
vom einen zum anderen
nie
gelangen,
immer unter der Macht des Gegensatzes.
verbleiben wir
dann
Im Lebens-
prozeß dagegen kann von vornherein das eine auf das andere angewiesen
und
sein,
die Beschaffenheit
jeder Seite sich
nach
dem
bemessen, was im Ganzen geschieht und erreicht wird; dann verschwindet die Starrheit des Gegensatzes, dann kann der Scheidung
ein
Zusammenhang überlegen bleiben.
Endlich diene zur Aufhellung und Abgrenzung dieser Fassung
des Geisteslebens noch eine Bemerkung historischer Art.
klärung
dem Mechanismus
neben
kannte
Die Auf-
der Natur keine andere
Wirklichkeit als das Nebeneinander einzelner Seelen, sie sprach nicht
von
einer
nur von
sondern
Geisteswelt,
Kant brachte die
einer
Geisterwelt.
Erst
Wendung in Fluß, welche die geistige Arbeit des
19. Jahrhunderts beherrscht,
Abhebung eines GeistesDenn bei ihm erscheint jenseit
die deutliche
lebens vom bloßseelischen Getriebe.
des Unterschiedes der Individuen eine gemeinsame geistige Struktur,
ein
Grundgewebe, das
Aber
staltet.
als
sich
die
alle
Sache
geistige Betätigung beherrscht
blieb
weder das Neue
fest
bei
ihm
zugleich
behandelten
sie
unbedenklich das menschliche
Geistesleben als absolutes Geistesleben
zeuger
Wirklichkeit.
aller
besonderes Tun,
Die speku-
erhoben das Geistesleben zur vollen Selbständig-
lativen Nachfolger
aber
ge-
genug bei sich selbst zusammenschloß,
noch eine deutliche Abgrenzung des Geistigen erfolgte.
keit,
und
insofern noch unvollendet,
und machten jenes zum Er-
Das konnten
nicht
sie
und mehr und mehr ward
dies
wohl ohne
ein
das Denken, für
das Ganze des Geisteslebens einzusetzen; daraus entstand nicht nur
ein viel zu knappes,
auch zu anthropomorphes Bild der Welt, es
drohte sich zugleich
alle
Wirklichkeit in einen
Dementgegen
verflüchtigen.
das menschliche Dasein
sei
hinausgehoben:
das Geistesleben, sondern er gewinnt
mit
an
einer
höheren Stufe
das Geistesleben nicht
rastlosen
teil
der Mensch erzeugt nicht
am Geistesleben und da-
der Wirklichkeit.
als eine
Prozeß zu
entschieden das Geistesleben über
Zugleich erscheint
besondere Betätigung, nicht als eine
besondere Seite des Lebens, sondern
als bei sich
selbst befindliches,
wirklichkeitbildendes Leben, das unser menschliches Tun keineswegs
ganz
erfüllt,
sondern
als ein
hohes Ziel zu ihm wirkt.
— Objektiv.
Subjektiv
y.
33
Das Verhältnis des Menschen zum Geistesleben.
Dieses Selbständigwerden des Geisteslebens mit seiner Abhebung
vom bloßen Menschen
verwandelt
dessen Verhältnis
zum Geistes-
leben aus einer scheinbar selbstverständlichen Tatsache in ein schweres
Problem.
Wie kann der Mensch,
verschwindender Punkt,
befindlichen Welt teilhaben,
Er kann es
leben darstellt?
für die nächste Betrachtung ein
am Ganzen einer Welt, einer bei sich selbst
welche sich nunmehr das Geistes-
als
wenn von vornherein
Wesen angelegt, er ihm
Das Geistesleben darf ihm
sicherlich nur,
das Geistesleben als Möglichkeit in seinem
irgendwie
verbunden
unmittelbar
ist.
nicht durch die Vermittlung seiner besonderen Natur zugeführt
dabei
all
sich
selbst
entfremdet werden,
es
muß ihm
seiner Unendlichkeit irgend gegenwärtig sein, es
als
und
Ganzes mit
muß ihn damit
von innen her, wenn auch zunächst nur der Möglichkeit nach, zu
Ohne ein solches Inne-
einem Weltleben und Weltwesen erweitern.
wohnen der Geistigkeit
gibt es für den
des Weiterkommens; würde er in
Menschen keine Hoffnung
dem Geistesleben nicht sein echtes
Selbst ergreifen, so könnte es nie eine Macht für ihn
werden; böte
und hielte es nicht mit richtender Kraft allem menschlichen Unternehmen Ziele und Maße vor, so
wären wir dem Wechsel und Wandel der Erscheinungen wehrlos
preisgegeben, so entfiele für uns alle Möglichkeit einer Wahrheit
jenes nicht einen unwandelbaren Pol
Nur das Geistesleben, nicht der bloße Mensch, kann
eine absolute
Festigkeit gewähren.
Solches Teilhaben des Menschen
Gesamtanblick
seines
Wesens.
am Geistesleben verändert den
Jenes
ist
nur möglich
jenseit des
unmittelbaren Daseins, so gewinnt sein Leben einen tieferen geistigen
Grund; zugleich
Betrachtung,
scheidet
sich
von der empirisch-psychologischen
welche mit den unmittelbaren Vorgängen des Seelen-
lebens zu tun hat, eine noologische, welche sich jene geistige Grundlage mit ihrer Selbsttätigkeit
Bei solchem
zum Vorwurf macht.
zwiefachen Anblick erscheint der Mensch in sich
Das Geistesleben ist
und eine Aufgabe, unerschütterliche
Ruhe und nie befriedigtes Streben, innerster Kern und fernes Ziem-
selbst
bei
als
ein
Gegensatz und ein Problem.
ihm zugleich
eine Tatsache
er selbst aber erscheint zugleich
klein
im Abstände, sein Leben
als
groß
wird
ein
der Verbindung,
als
unablässiges Suchen
des
in
eigenen Wesens und wird damit erst einer wahrhaftigen Geschichte
3
Eucken, Grundbegriffe. 4. Aufl.
Zum Grundbegriff des Geisteslebens.
34
Denn wie
fähig.
i<önnte
wo
entstehen,
das Streben
von außen abhängig wäre, nicht von innen her durch
lediglich
und
Ziel gelenkt
festes
solche
eine
gerichtet
ein
würde?
Auf dem Boden der menschlichen Geschichte überwindet das
Geistesleben
langsam die anfängliche Zerstreutheit und Ohnmacht;
es geschieht das, indem unter besonders günstigen
Umständen eine
Kernbildung innerhalb des Lebensprozesses
indem Tätigkeits-
komplexe
indem
suchen,
Art
geistiger
erfolgt,
zusammenschießen und
schließlich
ein
durchzusetzen
sich
Lebenssystem,
charakteristisches
Aufbau einer geistigen Wirklichkeit, unternommen wird.
licheres Beispiel gibt es
davon
als
das griechische Schaffen,
eine
wie es
Wie eine solche
mit ausgeprägter Art Leben und Welt umspannt.
Synthese
der
Kein deut-
Wahrheit des von ihr dargebotenen
ausschließliche
Lebensinhaltes behauptet, so zerlegt sie das Dasein in ein Für und ein
kann nichts dulden, was ihr fremd oder feindlich entgegen-
Wider,
sie
steht.
So erwächst Bewegung und Kampf, diese erschließen Erfahr-
ungen und treiben das Leben dadurch
weiter; so
mögen sich neue
mag
Konzentrationen vorbereiten, die ähnliche Schicksale haben, so
durch Werden und Vergehen der einzelnen Phasen der Wahrheitsgehalt des
Ganzen wachsen.
von
einem
wird,
während
begründenden
sich
Das freilich
nur,
wenn alle Bewegung
und richtenden Geistesleben umspannt
ohne das gegenüber den
starren
Widerständen
und den schweren Hemmungen der menschlichen Lage nun und
nimmer irgendwelche Wahrheit durchsetzen könnte.
der
geschichtliche
Prozeß
als
eine
fortschreitende
nicht subjektiver, sondern substantieller Art; es
Damit erscheint
Verinnerlichung
muß sich damit eine
immer größere Entfernung von der unmittelbaren Lage vollziehen,
die unter dem Gegensatz steht und daher einer vollen Innerlichkeit
wie einer echten Wirklichkeit entbehrt.
Innerhalb dieser
Bewegung findet sich auch ein
Platz für
den
Gegensatz, den die Ausdrücke „subjektiv" und „objektiv" recht unzulänglich bezeichnen.
Das Geistesleben
ist
Selbstleben
und Welt-
leben zugleich, ein Selbst entfaltet sich hier zur Welt und die Welt
gewinnt ein
Selbst, das eine gehört
sammengehörigkeit verhindert
wegung
das
nicht,
zum anderen. Aber solche Zudaß
in
der geschichtlichen Be-
Leben bald mehr zur Konzentration,
Expansion geht;
tiefung in sich
dort das Streben
selbst,
hier
bald
mehr zur
nach Verinnerlichung und Ver-
das nach Weite und Sachlichkeit, dort
die Gefahr eines Einströmens bloß menschlicher Elemente, hier die
Subjektiv
— Objektiv.
35
des Vordringens einer seelenlosen Welt; vielleicht besteht ein Rhyth-
mus
des Geschehens,
der
gibt.
Aber durch
Wandlungen hindurch
alle
bald
bald jenem
diesem,
die
Führung
beharrt das Streben
des Geisteslebens nach einer den Gegensätzen überlegenen Einheit.
Auch
bleibt
von jenem subjektiven oder objektiven Zuge innerhalb
des Geisteslebens grundverschieden ein Subjektivismus oder Objektivismus gegenüber
dem
Geistesleben,
Subjektivismus,
ein
der aus
den Zuständen des bloßen Subjekts eine Welt hervorspinnen möchte,
ein Objektivismus, der eine Wahrheit glaubt
von den bloßen Dingen
Das eine
her durch Austreibung des Geistes erreichen zu können.
muß rasch ins Leere sinken, wenn sie nicht versteckt aus eben dem überlegenen Geistesleben schöpfen, dessen Anwie das andere
erkennung sie ablehnen.
Ergebnisse für den Wahrheitsbegriff.
S.
Was an Wandlungen ersichtlich wurde, das muß sich auch auf
'
den Begriff der Wahrheit erstrecken und ihn eigentümlich
gestalten.
Wahrheit bedeutet jetzt nicht mehr die Übereinstimmung mit einem
draußen befindlichen Gegenstande, sondern ein Aufsteigen zu einem
Leben, das aller menschlichen Willkür überlegen
Gegensatz von Subjekt und Objekt durch
die
umspannt.
um eine Verwandlung des Daseins in Selbst-
Es handelt sich hier
tätigkeit,
und das den
ist,
tätiges Schaffen
mit ihrem umbildenden Wirken von aller bloßen Be-
tätigung innerhalb eines gegebenen Daseins wesentlich verschieden
ist.
Dies Wahrheitsstreben hat nichts mit einem ruhenden Sein zu
tun,
das unabhängig von allem Leben bestünde, vielmehr
und
sie
ist
Aber das Leben, das hier
in
Frage
Wahrheit innerhalb des Lebens,
das Leben.
Sache des bloßen Menschen, sondern
Wirklichkeit
Inhalten
zu
nur erreichbar durch
steht,
ist
nicht eine
ihm gelangt das Ganze der
in
und zugleich
einem Beisichselbstsein
allererst
und Werten, die Wahrheit aber ist nicht ein bloßes
zur Erhöhung dieses Lebens, sondern
Wahrheit
Alle
intellektuelle
einer
gesamtgeistigen
Wahrheitserkenntnis
der
liegt die
auf
einer
aller
gehört zu seinem Wesen.
ruht
schließlich
auf
wesentliche
Fortschritt
der
prinzipieller
Wahrheit,
Gewinn der Wahrheit
sie
Art
Weiterbildung
nicht Sache
zu
Mittel
eines
des
Lebens.
So
ist
einzigen Augenblicks,
sondern nur durch den Lauf der weltgeschichtlichen Arbeit mit ihren
Versuchen, Erfahrungen, Wandlungen dringt der Mensch allmählich
3*
36
Zum Grundbegriff des Geisteslebens.
zu ihr vor, es gibt kaum etwas so törichtes als der Anspruch philosophischer Systeme, in einem gegebenen Zeitpunkt die ganze
der Wahrheit
erschöpfen
und
alle
wir so im Suchen verbleiben und
Rätsel
lösen
zu wollen.
Fülle
Daß
zugleich unvermeidlich im Irren,
das kann uns in keiner Weise erschrecken,
wenn wir der Über-
zeugung sind, daß alles menschliche Streben eine Welt des Geisteslebens hinter sich hat,
die sich
nur durch Freiheit aneignen
die aber nicht auf unsere Willkür gestellt
ist.
läßt,
Theoretisch - praktisch.
(Intellektualismus
a)
I
— Voluntarismus.)
Geschichtliches.
|ie Frage des Intellektualismus
und Voluntarismus steht mit der
*-^ eben behandelten in engstef Verbindung, es wird bei ihr nur
direkter ins Seelische gewandt, was uns dort als Weltanblick beschäftigte.
Auch hier ein Gegensatz von Lebenstypen, auch hier eine Bewegung
der Jahrtausende. Nur der Unterschied, daß sich hier unsere Zeit
ihres eigenen Zieles weit sicherer fühlt.
Denn entschieden überwiegt
hier das Streben,
den Schwerpunkt des Lebens
in
das Wollen zu
verlegen als in dasjenige, was allein ihm Wärme, Kraft und Festigkeit
Wie mag es kommen, daß eine alte Streitfrage uns
geben könne.
plötzlich
Sehen
so einig findet?
wir,
ob die Geschichte zur Auf-
klärung dessen einiges beitragen kann.
Die Ausdrücke
Intellektualismus
und Voluntarismus sind mo-
dernster Prägung; jenes taucht in den philosophischen
Kämpfen zu
Beginn des
in
19. Jahrhunderts auf,
es findet sich z. B.
Schellings
Bruno (s. Werke IV, 309), als Gegensatz zu Materialismus; Voluntarismus
ist
die
erst
in
den
Ausdrücke
griechischen
letzten
Jahrzehnten aufgekommen.'^
theoretisch
Philosophie
— praktisch
zurück; der
Dagegen führen
der
in
die
Blütezeit
Gegensatz
der
theoretischen
bis
und der praktischen Vernunft erscheint zuerst bei Aristoteles (voG?
und 7rpaxTi>cd?). Jene hat die Aufgabe, die große Welt
^ewpTjTixd;
^
Gebildet ist das Wort von Tönnies, der darüber in der Wiener „Zeit"
vom 23. März 1901 folgendes berichtet: „Diese Termini (d. h. Voluntarismus
und voluntaristisch) sind zuerst vom Verfasser dieses Memoire gebraucht
worden in seiner Abhandlung »Zur Entwicklungsgeschichte Spinozas", Viertel-
Von Paulsen, der sie bald
angenommen und durch seine Autorität in Umlauf
Der Begriff der „voluntaristischen" Psychologie ist mehr und mehr
jahrsschrift für wissenschaftliche Philosophie 1883.
adoptierte, hat Wundt sie
gebracht.
zu allgemeiner Geltung gelangt".
Zum Grundbegriff des Geisteslebens.
38
mit ihren ewigen Ordnungen zu erkennen, die praktische Vernunft
dagegen befaßt
und Wandel
mit den menschlichen Dingen, die dem Wechsel
sich
unterliegen.
Sie
nicht bloß Einzelerkenntnis (An-
ist
wendung der allgemeinen Sätze auf den einzelnen Fall), sondern sie
hat
auch
eigentümliche Prinzipien,
aber ihre Gesamtleistung wird
der theoretischen Vernunft entschieden untergeordnet.
So auch in der
Gedankenwelt und dem Sprachgebrauch der Scholastik; wenn Thomas
von Aquino von cognitio practica redet, so versteht er darunter einfach
eine Erkenntnis, die sich auf das Handeln bezieht.
praktischer Philosophie
Für die Neuzeit
von theoretischer und
hat namentlich Ch. Wolff die Unterscheidung
Umlauf gebracht und dabei jener den
in
unbedingten Vorrang gegeben. ^ Wie in der Einteilung der Philosophie
so folgt
ihm auch im Sprachgebrauch Kant,
zieht er die bekannte Umwälzung dahin, daß
»was durch Fi-eiheit möglich
als das,
der Sache aber voll-
in
nunmehr das Praktische
das Übergewicht gewinnt
ist",
und zugleich einen selbständigen Gedankenkreis hervorbringt.
«Die
ihm in das Gebiet des Theoretischen
praktische Vernunft greift bei
zurück, indem sie Postulate erzeugt, also theoretische Voraussetzungen,
welche der Kritik der reinen Vernunft zweifelhaft waren" (s. TrendelenLogische Untersuchungen,
burg,
3.
Aufl.
457).
II,
Da nach ihm
nur auf diesem Gebiet die Vernunft eine volle Selbsttätigkeit erreicht,
so ergeben sich von hier die tiefsten Einblicke in die Wirklichkeit,
und
eröffnet
es
sich
absolute Wahrheit.^
nirgend anders als hier dem Menschen eine
Von Kant ist nur ein Schritt zu Fichtes Lehre:
praktische Vernunft
»die
einmal das Praktische
aber
die
ist
als
bloße
Wurzel
aller
als
Quell neuer Wahrheiten übergeordnet.
Es
spiegelt
sich
aber
So z. B. Logica § 92
'
versam
ex Metaphysica
:
der Geschichte
in
Palam
principia
Vernunft".
So wird
Anwendung untergeordnet, sodann
igitur est,
petere
der Ausdrücke
ein
philosophiam practicam uni-
debere.
§ 93: Metaphysica philo-
sophiam practicam praecedere debet.
Die Art, wie Kant aus der praktischen Vernunft Überzeugungen ab-
"
leitet,
z.
B.
nicht ohne Bedenken und fand manchen Widerspruch.
So sagt
Harms (Gesch. der Philosophie seit Kant S. 247): »Kant nennt die
ist
Ideen Postulate der praktischen Vernunft,
sie
sind aber gar keine Postulate
der praktischen, sondern sie sind Postulate der theoretischen Vernunft in der
Erkenntnis der praktischen, der handelnden Vernunft im sittlichen Leben des
Das Wort praktische Vernunft ist bei Kant selbst zweideutig; denn
wird darunter verstanden die handelnde Vernunft, dann aber auch
die Erkenntnis von der praktischen Vernunft."
Geistes.
einerseits
— praktisch.
Theoretisch
39
fundamentaler Gegensatz, die Frage, ob der Welterkenntnis oder dem
— denn vornehmlich wird unter
Führung unseres Lebens und
Vernunft verstanden —
moralischen Handeln,
dies
,
unserer
die
Überzeugungen gebühre.
Die Antwort darauf entscheidet
zugleich über unsere Stellung zur Wirklichkeit
und damit über das
Es entstehen zwei widerstreitende Lebens-
Bild der Wirklichkeit selbst.
typen, der eine vornehmlich auf Weite
Wärme und Kraft bedacht,
und Klarheit, der andere auf
der eine mehr
um
Gesetzlichkeit,
der
um Freiheit bemüht.
andere mehr
Die griechischen Denker
Intellekt,
praktischer
die Leitung
erteilen
einmütig
den Vorzug dem
oder maßvollere
es unterscheidet sie lediglich die schroffere
Ausführung des Grundgedankens.
Diese Schätzung des
war der naturgemäße Ausdruck der
hier
Intellekts
vorhandenen Überzeugung,
daß der Mensch einer unwandelbaren Weltordnung angehöre, die ihn
mit sicherem Dasein und herrlicher Pracht umfängt.. Nichts Größeres
gab es hier zu erstreben
Anschauung eines solchen Kosmos
als die
mit .ihrer Befreiung von aller Kleinheit des Alltages und allem Wirrwarr
So
verficht Aristoteles, dieser reinste
Ausdruck der griechischen Kultur,
die unbedingte Überlegenheit des
der menschlichen Verhältnisse.
über
Lebens
forschenden
veränderlichen Dingen
das
befasse
handelnde,
das
den Menschen
mit
und ihn von seiner Umgebung ab-
Nur soweit die Forschung reicht, soweit reicht nach
Glückseligkeit.
Auch die Wendung zur Moral, welche
hängig mache.
ihm
echte
die Stoa
brachte,
vom Denken,
eine
bedeutet
nicht
sowohl eine Ablösung des Lebens
Aufnehmen tätiger Energie
Steigerung des Denkens zur Denkhandlung.
ein
als
in
das Denken,
Und
das
letzte
Aufleuchten des griechischen Geistes in Plotin bringt eine Erhebung
des Denkens zu voller Souveränität und weltschaffender Größe; im
Untergange
selbst
Geisteskraft,
der
steigert
seine
das Griechentum das Bekenntnis zu der
Kulturarbeit
eine
unermeßliche Weite
und
eine wunderbare Klarheit verdankt.
Das Christentum mußte nach
Schätzung brechen.
in
Denn wo
das Verhältnis des
Erscheinen
sich
seiner innersten Natur mit jener
das Hauptproblem des
Lebens
Menschen zu Gott verlegt, und wo mit dem
neuer Tiefen
auch
schwere Verwicklungen,
ja
dunkle
Abgründe im Menschenwesen zur Empfindung kommen, wo es demnach
einen Aufstieg
und
eine
Erneuerung
gilt,
da
muß sich
das
Streben des Menschen von der Erkenntnis der Welt auf den Stand
seiner Seele
und weiterhin auf den Aufbau
eines neuen
Zusammen-
Zum Grundbegriff des Geisteslebens.
40
hanges der Menschheit
richten.
Der Intellektualismus wird damit in
Aber diese innere Wandlung ist in der Ge-
der Wurzel gebrochen.
staltung und den allgemeinen Verhältnissen wenig zum Ausdruck
gekommen was die Herzen erfüllte, fand nicht die Kraft zur Schöpfung
;
einer entsprechenden Gedankenwelt
Lediglich bei Augustin erscheinen
bemerkenswerte Ansätze dazu, wie er denn alle Wirklichkeit auf Wollen
(nihil aliud quam voluntates) und in seiner Psychologie
dem Willen als der die Seele einigenden Kraft die leitende Stellung
zurückführt
Aber ein durchgebildetes Lebenssystem und eine entsprechende
gibt.
Gedankenwelt hat auch Augustin daraus nicht entwickelt; so
ist
die
Gestaltung des Christentums unter einem starken Einfluß derselben
Denkweise verblieben, die es überwinden wollte, so leidet das Christen-
tum bis zum heutigen Tage an einem Zwiespalt innerster Gesinnung
und greifbarer Ordnung. Der griechische Intellektualismus beherrscht
auch das christliche Dogma; mögen an die Stelle der Lehren von
der Welt Lehren von Gott getreten sein,
es
daß die
bleibt dabei,
Erkenntnis über die Wahrheit und den Wert des Lebens
richtige
Auf der Höhe der Scholastik wächst noch der Einfluß
entscheidet.
des griechischen Intellektualismus, ein logisches Räsonnement dringt
bis
die
in
fehlt nicht
Scotus,2
Tiefen der christlichen
letzten
Gedankenwelt
vor.^
dem Nominalismus,
der praktisch
gerichteten
Mystik;
Reformation bringt diesen Zug zum siegreichen Durchbruch.
bemüht
des
Es
an Gegenwirkungen zu gunsten des Willens, so bei Duns
mit höchstem
sich
griechischen
platonischer
Eifer,
Intellektualismus
Prägung zu
befreien,
die
Luther
das Christentum von der Macht
sowohl
der,
aristotelischer
als
neu-
wie er meint, seinen echten
Tatbestand verdunkelt oder verflüchtigt hat; Melanchthon aber nennt
«das Herz mit seinen Affekten« «den wichtigsten
teil
Aber
des Menschen".
und den Haupt-
bei aller Verstärkung des Willens hat auch
der Protestantismus nicht die Kraft gefunden, den innersten Lebenstrieb
in
ein
Lebenssystem zu verwandeln, auch er hat schließlich
wieder der Macht des Intellektualismus gehuldigt.
^
Denn mochte die
Nur äußerlich angesehen dient im Mittelalter die Philosophie als Magd
der Theologie, innerlich hat damals weit mehr die Philosophie der Theologie
ihren Stempel aufgedrückt,
"
Er sagt
z.
B.
(s.
est habitus speculativus,
Stöckl,
Philos. d. Mittelalt.
credere est visio speculativa, sed practica.
fruitioni.
II,
S. 788):
Fides
non
nee credere est actus speculativus, nee visio sequens
Nata est enim ista visio conformis
Theoretisch
für
Spekulation
die
— praktisch.
Dauer abgelehnt werden,
41
ein
Wissen anderer
Wissen von geschichtlichen Daten, aber immerhin ein Wissen
Art, ein
erschien als unerläßlich zur Rettung der Seele; auch der Begriff des
Glaubens
erhielt
eine
Färbung,
intellektualistische
stark
die
neue
Kirche aber wurde vornehmlich eine Gemeinschaft der Lehre, eine
Schule des
Es entstand eine neue Orthodoxie, an
reinen Wortes.
Selbstgerechtigkeit und Unduldsamkeit der griechischen voll gewachsen.
Die Neuzeit beginnt mit voller und freudiger Ergreifung der
Denkarbeit; das Denken
ist
es,
von dem
sie
eine
Befreiung
vom
Druck der geschichtlichen Überlieferung hofft, das Denken, das eine
Klärung des unerträglich gewordenen Chaos verspricht, das Denken
endlich, welches das Gewebe kleinmenschlicher Interessen zerreißt und
die Unendlichkeit des Weltalls eröffnet.
Art
ja
ist
Gegenüber der griechischen
das Denken aus ruhigem Anschauen weit mehr rastlose Arbeit,
stürmisches Vordringen, aus Aneignung einer gegebenen Welt zum
Aufbau einer neuen geworden.
vom, Großen
Eine solche beherrscht die Aufklärung
bis ins Kleine, sie beherrscht nicht nur ihre spekulative
Richtung mit der Kühnheit ihrer Weltentwürfe,
sondern auch die
empiristische mit ihrer Richtung auf das Menschenleben.
hier
wird von def klaren und deutlichen Erkenntnis
Denn auch
alles
Heil er-
eine andere Art des Erkennens, aber es ist ein Erkennen,
wartet, es
ist
das den
Kern der Arbeit
trug freilich auch
Wie
bildet.^
die Aufklärung
in
jede große Bewegung,
selbst einen
sich
so
Rückschlag:
und Überspannung des Erkennens erzeugte mit Notwendigkeit einen Zweifel an seinem Vermögen gegenüber der Welt
die Steigerung
und
seiner Herrschaft über den Menschen.^
Aber ein bloßer Rück-
schlag hat noch nie die Herrschaft über die Geister gewonnen, es
bedurfte
positiver
Wandlungen,
um die Menschheit in eine andere
Bahn zu lenken.
Solche Wandlungen erfolgten auf philosophischem Boden in Kant;
sowohl sein Nein als sein Ja hatten bei diesem Problem eine unver^ S. z. B.
Locke zu Beginn seines Essay: Our business here is not to
know all things, but those which concem our conduct.
^ So z. B. bei Pascal, so mehr noch bei Bayle, dem bedeutendsten SkepEr sagt z. B. oeuv. div. 1727, III, 89 b: Ce ne sont
tiker der Aufklärung.
pas les opinions generales de l'esprit, qui nous determinent ä agir mais
passions presentes du coeur.
les
Wie bei Bayle die Moral allein dem Leben eine
Festigkeit verleiht, so sagt auch Friedrich II., sein getreuer Schüler: Les sciences
doivent etre considerees
cite
comme des moyens qui nous donnent plus de capa-
pour remplir nos devoirs (s. Zeller,
Fr. d.
Gr. als Philosoph
S. 183).
Zum Grundbegriff des Geisteslebens.
42
gleichlich größere
je
Wucht als eine andere wissenschaftliche Leistung
und gründlicher als je wird das Vermögen des
Schärfer
zuvor.
bloßen Erkennens geprüft und werden die Bedingungen seines Gelingens ermittelt; das Ergebnis
diese Erschütterung wird
ist
eine schwere Erschütterung, aber
mehr als ausgeglichen durch die Steigerung
des moralischen Tuns zu einer moralischen Welt und die Anerkennung
dieser Welt als des Kernes aller Wirklichkeit.
Erst mit solcher
Um-
wälzung erhielt der Intellektualismus eine ebenbürtige Gegenwirkung,
nun
gelangte
erst
ein
seit
Jahrtausenden
vorhandenes Streben zu
Wir wissen, daß trotzdem der Intellektualismus im Panlogismus Hegels noch einmal sein
wissenschaftlicher Klärung und Durchbildung.
Haupt erhob, so kühn erhob wie nur je, aber wir wissen auch, daß nur
ein
rasches Hinwegeilen
bald ein
über Kant dies möglich machte, und daß
um so stärkerer Rückschlag kam.
zug der Zeit gegen den Intellektualismus.
Einfluß Schopenhauers
der Neigung
praktische
und seiner Willenslehre, wir gewahren
der Religion
und Theologie,
die
ihren
es in
Schwerpunkt
in
Aufgaben und Forderungen zu verlegen, wir gewahren
es in dem Zuge des gesamten Lebens, das
hinter
Seitdem geht der Haupt-
Wir gewahren das in dem
praktisch -sozialen
Grübeln über Weltprobleme
Fragen zurückzustellen, die sich mit
immer größerer Dringlichkeit erheben.
Auf dem eigenen Gebiet der
Wissenschaft aber wirkt zu gunsten der neuen Überzeugung namentlich
die Psychologie,
und
Interessen
eigne
indem
sie die
beherrschende Macht der Triebe
auch über das Vorstellungsleben aufdeckt,
ja seine
Bewegung als vom Willen geleitet aufweisen möchte.
Solcher Schätzung
des Willens
entspricht
die
Neigung,
dem
Überwiegen des Verstandes alle möglichen Schäden der gegenwärtigen
Die Grundlage unserer geistigen Existenz und
Lage aufzubürden.
die Hauptrichtung unseres Strebens sind unsicher geworden: das verschuldet, so heißt
es,
der Intellekt, der alles beweisen,
d. h. auf
Um-
wegen ermitteln möchte und damit alles unmittelbare Leben mit seiner
Gewißheit verscheucht;
der Meinungen
uns umfängt eine unsägliche Zersplitterung
und Schätzungen: das kommt, so sagt man, vom
Vorwalten der Verstandesarbeit, welche die Individuen auf ihre eigene
Reflexion stellt und sie damit unvermeidlich auseinandertreibt; beklagt
wird ein Mangel an Ehrfurcht vor göttlichen und heiligen Dingen:
das liegt, so hören wir, an einer Überhebung des menschlichen Selbst-
bewußtseins, und die Haupttriebkraft dabei ist der Intellekt mit seinem
Kraftgefühl
und seinem Wissensdünkel.
Trägt demnach der Intellekt
Theoretisch
— praktisch.
43
die Hauptschuld aller Irrung, so läßt die Befreiung von seiner Herrschaft eine Gesundung des gesamten Lebens erwarten.
Hat der heutige
Voluntarismus die Kraft zu solcher Befreiung?
Der Voluntarismus.
b)
Der Voluntarismus
Erscheinung
keine
ist
einfacher
Art,
die
Hauptepochen der Geschichte haben ihn ihrer Hauptrichtung gemäß
verschieden
In
der
gestaltet,
religiösen
jede
besonderen Voluntarismus.
ihren
hat
Gedankenwelt ging die
—
daß
Behauptung dahin,
sowohl die Offenbarung Gottes als ihre Annahme durch den Menschen
Sache
eine
des Willens
sei,
der
nicht
sich
Darin lag eine Betonung der Selbständigkeit,
weiter
ableiten
lasse.
Ursprünglichkeit und
reinen Tatsächlichkeit des religiösen Geschehens, die Abweisung aller
Versuche, es aus weiteren Zusammenhängen verständlich zu machen.
Zum
Ausdruck
den
bringt
Gegensatz
bekannte Formel,
die
nach, Thomas von Aquino Gott das Gute gebietet,
nach Duns Scotus das Gute gut
Spezifische
der
gleichlichkeit
gleich
Religion,
ihre
ist,
weil
Gott es
weil
Unabhängigkeit
es gut
ihre
ist,
Das
gebietet.
und
daß
Unver-
kam bei dieser Denkweise voll zur Geltung, aber zu-
entstand
die
Gefahr
einer
Ablösung vom
übrigen
Leben,
Zusammenhängen; wie es hier nicht
2u einer vollen geistigen Durchdringung und Aneignung des Wahrheitsgehaltes kam, so konnte leicht die Tatsächlichkeit in starre und
eines
Herausfallens aus allen
äußerliche Positivität
und
daß,
die Ursprünglichkeit
Man möchte
Willkür umschlagen.
wer über die Vernunft
hier
hinausstrebt,
und Freiheit in blinde
an Plotins Wort denken,
leicht
in
Gefahr kommt,
aus ihr herauszufallen.
Auf dem Gebiete der Philosophie stellt der Voluntarismus
anders dar: hier
gilt
es
eine Verlegung des
sich
Zentrums des Lebens
dem Erkennen ins Wollen, namentlich in das moralische Wollen.
Antrieb dazu gab vornehmlich ein Irrewerden am Vermögen
des Erkennens; wie es unfähig dünkte, zum letzten Grunde der
Dinge durchzudringen, so schien es auch dem Leben keinen sicheren
aus
Den
Halt gewähren
zu können.
Sollte
also
nicht auf alle
Wahrheit in
vollem Sinne verzichtet werden, so war eine andere Quelle zu suchen,
und
eine solche schien nach Erschütterung des religiösen
nichts anderes
dem großen
zu bieten
Sinne,
als
des Menschen
wie Kant dies versteht,
sittliches
Glaubens
Handeln.
In
wird dies zur Offen-
Zum Grundbegriff des Geisteslebens.
44
barung einer neuen Welt, welche die
Tiefe
letzte
Diese Welt aber läßt sich ebenso wie das
bildet.
der Wirklichkeit
Handeln
sittliche
selbst nicht jedem theoretisch demonstrieren oder als ein Vorhandenes
sondern
aufweisen,
der das
sie
besitzt eine
Überzeugungskraft nur für den,
Grundgeschehen anerkennt und auf
sittliche
nimmt;
sich
so geht eine Tat dem Erkennen voran, und was aus ihr an leitenden
Überzeugungen hervorgeht, das hat nicht den Charakter theoretischer
Erkenntnisse, sondern den praktischer Postulate.
Die gewaltige Er-
regung und eingreifende Wandlung, die das bewirkt hat, stehen uns
vor Augen.
allen deutlich
richtige Würdigung dieser Wendung
Die
schwer,
und
notwendige
weil
ist
namentlich deshalb
Wahrheiten
fruchtbarste
hier
mit
problematischen Fassungen aufs engste verquickt sind.
Ein solche
Wahrheit
daß
liegt
vor
allem
letzten
Überzeugungen
Welt,
sondern
durch
der
in
deutlichen
Einsicht,
unsere
nicht durch den Befund der uns umgebenden
was im Bereich des
bestimmt werden,
das
Innenlebens vorgeht und an Wirklichkeit ersichtlich wird; das macht
Versuchen ein Ende, durch Spekulation
allen
in ein
inneres
Wesen
der Dinge einzudringen und von ihm her die Wirklichkeit zu verstehen.
In
andere,
daß
schlossen
enger Verbindung mit dieser Wahrheit steht aber die
der Bestand des
allen
geweckt und
geschieht,
vor Augen
entfaltet sein
dem
Einzelnen
Innenlebens nicht mühelos und ge-
liegt,
sondern
daß
dieses
Leben
erst
will,
und daß nach dem Maße, wie das
auch
das Bild
Die Entzweiung der Menschen im
Streit
der Welt
um
sich
gestaltet.
die Wahrheit,
sowie
der persönliche Faktor im Wahrheitsstreben können hier ihre volle
Würdigung finden.
Aber sobald wir vom Allgemeinen des Gedankens zur n;iheren
Ausführung
fortschreiten,
erwachen Zweifel über Zweifeln.
etwas anderes, die Grundtatsachen des Innenlebens in den
Es
ist
Mittel-
punkt der Erkenntnisarbeit zu stellen und in ihnen die entscheidenden
Daten zu finden, etwas anderes, jene direkt zu einer eignen Erkenntnisquelle zu machen.
ist
dieses.
Ebenso notwendig wie jenes, ebenso unmöglich
Denn der Befund des Innenlebens
läßt
sich
nicht un-
mittelbar, wie er vorliegt, als sicherer Ausgangspunkt ergreifen, sondern
er
ist
erst durch die Erkenntnisarbeit zu klären
und zu durchleuchten;
was dabei aber in ihm als Grundtatsache und als Wahrheit befunden
wird, das hat eine Allgemeingültigkeit und zugleich einen
Zwang, das
läßt
sich
inneren
unmöglich auf eine persönliche Zustimmung
Theoretisch
45
Es gibt mathematische Wahrheiten von solcher Schwierig-
stellen.
keit,
— praktisch.
daß nur ein kleiner Teil der Menschen sie klar zu durchschauen
vermag,
das ihrer Allgemeingültigkeit auch nur den mindesten
tut
Wenn daher auch die Lebenswahrheiten ihre volle Über-
Abbruch?
zeugungskraft erst bei Entwicklung des entsprechenden Lebens
langen können, ja
des Menschen in sich
so werden sie dadurch in keiner Weise
trägt,
zu bloßen Möglichkeiten,
sondern
werfen könnte,
er-
wenn die Wendung zu ihnen eine Entscheidung
der eine annehmen, der andere ver-
die
sie
behalten
und
die volle Notwendigkeit
Allgemeingültigkeit; die Subjektivität liegt nicht in der Sache, sondern
nur im Verhältnis des Menschen zu
besitzen,
das
mit
innerlich
einem
ihr,
nichts
kann volle Wahrheit
subjektiven Faktor behaftet
ist.
Von solchen Erwägungen aus müssen wir den Begriff der praktischen
Vernunft als schief und irreleitend grundsätzlich ablehnen, es gibt nicht
und eine praktische Vernunft nebeneinander, son-
eine theoretische
dern es gibt nur eine einzige Vernunft, die mit
zu
t)an
hat;
dem ganzen Leben
den Begriff dieser Vernunft aber ist als wesentliches
in
Und es schwebt die
Merkmal das der Selbsttätigkeit aufzunehmen.
Vernunft nicht
der Luft, sondern
in
frei
sie
ist
die Vertreterin eines
bei sich selbst befindlichen Lebens, eines Beisichselbstseins der Wirk-
Ohne ein solches gibt es keinerlei Wahrheit.
lichkeit
Dazu behauptet der Begriff der praktischen Vernunft meist nicht
die
Höhe,
die
er
Hier vollzieht er eine
Kant einnahm.
bei
wälzung des ganzen Lebens, bewirkt er eine Versetzung
Um-
in ein ur-
sprüngliches Schaffen, und gibt er dem Leben in besonderer Richtung
wenn irgend
wenn auch eine andere Metaphysik
als die der ontologischen Spekulation.
Je mehr sich aber dieser
metaphysische Charakter verwischt, desto mehr wird das Reich der
einen Weltcharakter
etwas,
so
ist
mit
strenger Allgemeingültigkeit;
dies Metaphysik,
praktischen Vernunft aus der Tiefe der ganzen Wirklichkeit zu etwas
besonderem neben anderem, desto weniger
allgemeingültigen
die daraus hervorgeht,
wird
läßt sich hier zu einer
Die Lebensgestaltung aber,
Wahrheit gelangen.
leicht ein
Gebiet des praktischen und
moralischen Lebens gegen die Kulturarbeit isolieren, damit aber jenes
und auf
zu subjektiv und empfindungsmäßig,
dieses
die bloße Leistung gerichtet gestalten,
den Gesamtstand des Lebens
zu
aber durch solche Spaltung stark herabdrücken.
die Kulturarbeit
und
auseinanderfallen;
je
äußerlich
In
Wahrheit dürfen
die letzten
Überzeugungen des Menschen
mehr
es tun,
sie
desto
nicht
mehr muß unserem
Zum Grundbegriff des Geisteslebens.
46
Leben
durchwaltende
eine
Größe
Seele
und
sich
der Voluntarismus im Leben der
zugleich
alle
ent-
weichen.
Wieder anders gestaltet
Hier erscheint er als wissenschaftliche Theorie zunächst
Gegenwart.
dem
auf psychologischem Gebiet in
von
Vorstellungslebens
den
auch
gesamten
den
Streben,
Trieben
die Abhängigkeit des
und Neigungen aufzuweisen,
des Vorstellungslebens
Verlauf
als
durch ein
Willensmoment bedingt zu zeigen, wie das vornehmlich in
ist
neue und fruchtbare Erkenntnis
Viel
Apperzeptionslehre geschieht.
Wundts
auf diesem Wege gewonnen, ja der Gesamtanblick der Sache ver-
Nur in der Richtung mögen
tieft.
entstehen,
und Wille als zwischen
zentraler
der durch den ganzen
liegt,
hier
noch Fragen und Zweifel
ob dabei nicht oft weniger ein Gegensatz zwischen Intellekt
und peripherer Seelentätigkeit vor-
Umfang des Lebens geht.
gestaltet sich auf modernem Boden
Für das Ganze des Lebens
der Voluntarismus
und das Überwiegen der praktischen Aufgaben
und zwar
dahin, daß in weitem Zuge das Befinden der Menschheit,
Daseins,
der Menschheit des unmittelbaren
für
das Ziel der Ziele
und auch das Erkennen zu einem bloßen
erklärt
macht wird;
es erscheint als eine
zur Förderung des Wohlseins der Menschen wirkt.
Richtung des modernen Lebens entspricht,
die
Geschichte ersehen.
Eine
Mittel dafür ge-
unnütze Grübelei, wenn es nicht
ließ
Daß
dies der
schon der Blick auf
Ermüdung an dem Kampf um
die
Weltprobleme ist eingetreten, und die Fragen der inneren Bildung, der
Bildung des ganzen Menschen zu einer weltumspannenden Persön-
werden
lichkeit,
durch
die
wirtschaftlichen, technischen
unablässig
anschwellenden politischen,
Probleme weit zurückgedrängt, im be-
nimmt der Kampf um die ökonomische Selbsterhaltung
immer mehr alle Kräfte in Anspruch und stellt immer mehr das
Leben und Handeln unter den Gesichtspunkt der Zweckmäßigkeit;
wie sollte in solchen Zusammenhängen noch irgendwelcher Platz
sondern
für einen Selbstwert des Wissens bleiben?
Ein Versuch, von jener
praktischen Haupttendenz aus eine eigentümliche Theorie des Wissens
zu
entwickeln,
ist
namentlich im Pragmatismus gemacht,
Amerika und England aus immer mehr die Kulturwelt
so
sei
er etwas genauer betrachtet.
der von
beschäftigt;
Theoretisch
47
Der Pragmatismus.
c)
Der Pragmatismus
daß
— praktisch.
ist
in
Deutschland noch so wenig bekannt,
es vor aller Erörterung einer kurzen Orientierung bedarf;
halten uns dabei namentlich an die Vorlesungen
wir
von William James,
Zweck einer Aufklärung über den Pragmatismus verDer Ausdruck und der Begriff Pragmatismus im hier ge-
die eben den
folgen.^
meinten Sinne stammt von Charles Pierce, der ihn zuerst
amerikanischen Zeitschrift „Populär Science"
wandte.
Zwanzig Jahre
und
glänzender Weise weitergebildet.
in
später
hat James
in
der
vom Jahre 1878
ver-
Sache aufgenommen
die
Als weitere Führer
sind
Dewey (New York) und Schiller (Oxford) zu nennen; von letzterem
stammt der Ausdruck «Humanismus."
Es
nicht
ist
daß hier wohl zuerst
geschichtliches Interesse,
in
ohne
ein kultur-
einer philosoph-
ischen Bewegung Amerika vorangeht, wie denn auch dort vornehmlich
der Pragmatismus eine weitverbreitete Strömung geworden
Was, Europa anbelangt, so hat er außer
England namentlich
in
ist.
in
Einfluß gewonnen.
Italien
Über das Verhältnis des Pragmatismus zu anderen. Denkrichtäußert James sich folgendermaßen:
»Der Pragmatismus
ungen
repräsentiert
sophie,
eine
uns
durchaus
vertraute
nämlich die empirische Richtung,
in einer radikaleren
Richtung
allein
der Philo-
in
er repräsentiert sie
und zugleich einwandfreieren Form als die war,
die sie bisher angenommen hatte" (S. 31
a. a.
O.), ferner,
»er stimmt
dem Nominalismus darin überein, daß er sich überall an das
Einzelne hält, mit dem Utilitarismus darin, daß er überall den
praktischen Standpunkt betont, mit dem Positivismus in der Vermit
achtung, die er den bloß sprachlichen Problemlösungen, überflüssigen
Fragestellungen
O. S. 33).
(a. a.
und metaphysischen Abstraktionen entgegenbringt"
Der Pragmatismus
sondern eine Methode zu
sein,
legt
diese
Wert
darauf,
kein System,
Methode aber besteht in der
Beziehung alles Erkenntnisstrebens auf das menschliche Dasein und
was hier
sich
bewährt,
Das Wahre wird damit
ein
Teil
Förderung;
seine
gelten.
nur
kann
des Guten,
als
Wahrheit
„wahr heißt
„Der Pragmatismus". Ein neuer Name für alte Denkmethoden. VolksAns dem Englischen übersetzt von
Wilhelm Jerusalem, 1908. — Beachtenswert ist auch der Aufsatz von Jerusalem
in der Deutschen Literaturzeitung vom 25. Januar 1908: „Der Pragmatismus.
Eine neue philosophische Methode."
*
tümliche philosophische Vorlesungen.
Zum Grundbegriff des Geisteslebens.
48
was
alles,
dem
auf
sich
Überzeugung
Gebiete der intellektuellen
aus bestimmt angebbaren Gründen
als
gut erweist"
«Alle
48).
(S.
Theorien sind nur Werkzeuge, nur Anpassungen der Gedanken an
und keineswegs Offenbarungen oder intellektualistische
die Tatsachen,
Lösungen eines göttlichen
Welträtsels"
122).
(S.
In
solchem Ge-
dankengange gelten dem «Humanismus" die Wahrheiten als Erzeug«Die Wahrheit macht keinen anders-
nisse des Menschengeschlechts.
gearteten
Anspruch und
heit und
Reichtum, Alle diese Ansprüche sind bedingter Art« (S. 146).
legt
Fassung gibt
Solche
keine andere Pflicht auf als Gesund-
Forschung insofern
der
durchaus
eine
eigentümliche Richtung, als es nunmehr nicht sowohl die Begründung
von
als
Prinzipien
nicht
jetzt
Menschen
Sache
in
sondern
betrachtet,
Konsequenzen
gilt,
Natur gegenüber
dem
auf den Menschen gerichtet
und
Entwicklung
die
als
die
alles
ihre
in
eignen
einer
nach der Leistung für ihn bemessen wird.
Was
ehesten
Der
das besagt und welche
deutlich
Streit
durch
die
Wandlung
Beispiele,
des Materialismus
wird
bewirkt,
es
welche James
und des Spiritualismus
tritt
am
bringt.
selbst
in
eine
neue Beleuchtung und wird zugleich einer Entscheidung zu-
völlig
geführt,
wenn
nicht das Recht der Prinzipien erörtert,
sondern die
Wohl der Menschheit erwogen wird. Materialismus bedeutet in diesem Zusammenhange die
Denkweise, welche* die höheren Phänomene durch die niederen erLeistung eines jeden der Gegner für das
klärt
und
blinden Kräfte bestimmen
Leitung
und ihre
die Geschicke der Welt durch ihre blinden Teile
läßt,
Spiritualismus diejenige,
welche die
dem höheren Elemente zuweist und damit den Geist nicht
zu einem bloßen Zeugen und Berichterstatter des Weltlaufs macht,
sondern ihn
Frage,
denselben eingreifen
tätig in
welche der beiden
läßt.
so kann die Antwort nicht zweifelhaft sein.
tischen
Ergebnisse
Spiritualismus
mit
des
Stellen wir
nun die
Beleuchtungen mehr das Leben fördert,
Denn die letzten prak-
Materialismus
sind
trostlos,
während der
Bejahung
einer
sittlichen
Weltordnung
gibt.
«Der Glaube an geistige
seiner
unseren Hoffnungen freien Spielraum
Wesen in all seinen Formen hat es immer mit einer Welt der Vertun, während die Sonne des Materialismus
Meer der Enttäuschung untergeht" (a. a. O. S. 67). Dem
heißung zu
die
Erörterung
des
religiösen
Prinzipien her wird es
Problems:
statt
von
in
einem
entspricht
spekulativen
vom Bedürfnis des Menschen aus behandelt.
»Nach pragmatischen Grundsätzen
ist
die Hypothese
von Gott wahr,
— praktisch.
Theoretisch
wenn
im weitesten Sinne des Wortes befriedigend
sie
nun immer noch
besteht
den
andern
O.
(a. a.
so
daß
sie
wirkt,
in
und das Problem
daß
auszubauen,
Wahrheiten
wirkenden
Einklang
mit
sich
sie
bringen
läßt"
S. 192).
Wer sich
weite
zeigt,
Hypothese
die
darin,
Was
wirkt.
dieser Hypothesen
Schwierigkeiten
restlichen
die
mögen, die Erfahrung
sein
49
in
Ganze einlebt, kann wohl
das
der Zeit
Kreise
gewinnen
zu
verstehen,
daß
Indem
etwas,
vermochte.
was sonst nur nebenbei und an einzelnen
Stellen
es
es
beachtet wurde,
zur Hauptsache macht, wird eine Beleuchtung der Dinge gewonnen,
den Vorzug der Einfachheit und der
die
hat.
Eine
große
Fragen, die keine
Vereinfachung
teien
diese
gestattet;
als
alle
unnütz
Beziehung aber scheint einen Maßstab
in
würdigen und damit die Sache dem
zu entziehen
indem
augenscheinlich,
ist
zu gewähren, der die verschiedenen Behauptungen
fangenheit zu
Verständlichkeit
Beziehung zur Lebenshaltung haben,
müssen;
ausscheiden
leichten
voller
Streit
Unbe-
der Par-
die Wahrheit erhält hier mehr Unmittel-
mehr Beweglichkeit und Flüssigkeit, indem
sie so in die Bewegung des Lebens hineingestellt wird und zu ihrer
barkeit und Fruchtbarkeit,
Förderung mitzuwirken
hat;
gerade unsere Zeit mit ihrer Zersplitter-
ung der Überzeugungen scheint eine derartige Lösung zu empfehlen.^
Das Positive der Leistung wird aber durch eine scharfe Kritik des
überkommenen Wahrheitsbegriffes wesentlich unterstützt.
Aber so viel Anregung das alles, vertreten von bedeutenden
und geistvollen Männern, bringen mag, als Letztes und Ganzes angesehen
druck
muß
des
es
uns doch
als
ein
Pragmatismus stammt
Irrtum gelten.
Der starke Ein-
namentlich daher,
daß hier die
gewöhnliche Betrachtungsweise umgekehrt wird; wie aber, wenn dabei der Begriff der Wahrheit selbst auf den
So aber geschieht
es
in
Wahrheit
des Wahrheitsbegriffes, und das
ist
Kopf zu stehen kommt?
Das eben
die
ist
das Wesentliche
bewegende Seele des Wahr-
daß der Mensch dabei etwas erreicht, was jenseits aller
Meinungen und Neigungen liegt, und was völlig unabhängig
von seiner, ja von aller Menschen Zustimmung gilt. Damit scheint
heitsstrebens,
seiner
'
James (a. a. O. S. 122) bemerkt in dieser Hinsicht: „Die tatsächliche
Unruhe, die jetzt in der theoretischen Philosophie herrscht, der Wert, den
jeder Denktypus für gewisse Zwecke besitzt, die Unfähigkeit eines jeden dieser
Denktypen, alle anderen zu verdrängen, alles das legt die pragmatische An-
schauung nahe."
Eucken, Grundbegriffe.
4.
Aufl.
4
Zum Grundbegriff des Geisteslebens.
50
dem Menschen
neues Leben zu winken,
wesentlich
ein
und Großwerden des Wesens,
eine Befreiung
Wirklichkeit,
von
ein
innere Gemeinschaft
eine
bloßmenschlichen
aller
Weit-
mit der
Art.
Wo
dagegen das Wohl des Menschen und der Menschheit zum höchsten
und zum richtenden Maß gemacht wird, da sinkt die Wahrheit
Meinung herab, und das ist eine innere
Ziel
zu einer bloßnützlichen
sie in
Was immer sie an Überzeugungskraft hätte, das müßte
dem Augenblick verlieren, wo sie als ein bloßes Mittel durch-
schaut
würde.
Zerstörung.
Wahrheit
»instrumentale" Wahrheit
nur
ist
ist
möglich,
Selbstzweck
als
eine
keine Wahrheit.
Das heißt nicht behaupten, daß die Wirkung der verschiedenen
Lehren auf den Zustand des Menschen ein unwichtiger Gegenstand
kann
sicherlich
sei;
Aufklärung und Anregung davon ausgehen,
viel
wenn jene Wirkung mehr beachtet und ihren Gründen nachgeforscht
wird. Aber zunächst haben wir es dabei mit einem bloßen Phänomen
zu tun; was in ihm Kern und Schale, Recht und Unrecht sei, das
ist
erst herauszustellen.
Dem
Pragmatismus
Wahrheiten
es
so
und
auf,
daß
sicher,
ander geraten?
lauter
in
er selbst sieht darin einen Vorteil.
diese Wahrheiten
gegenseitig vertragen,
Wahrheit
die
sich
löst
daß
nicht
sie
sich
friedlich
einzelne
Aber ist
und freundlich
schwere Konflikte mit ein-
in
Wer sollte wohl bei solchen Konflikten der Schieds-
richter sein?
Endlich
ist
das
betrachtet,
was
das,
Gelingen
allem Zweifel enthoben.
Kulturleben,
das
als
nimmt;
um
dieses
wie
der
Pragmatismus
als
höchstes
und Gedeihen des Lebens,
Denn
es
das
sich
Leben
in
ist
hier
Ziel
keineswegs
nichts anderes
der Breite des Daseins aus-
Leben aber als ein sicheres Gut zu betrachten,
muß man von dem
Kulturenthusiasmus
und
der
optimistischen
Stimmung beherrscht sein, die früheren Zeiten näher als der unsrigen
lagen.
Denn ist dies Leben, wenn es für sich selbst als das Letzte
genommen wird, all der Mühe und Arbeit, all der Sorgen und Aufregungen, all der Schmerzen und Entsagungen wert, die es kostet?
Ist
dies Leben, genauer betrachtet, mit
und
all
seinem prunkenden Schein
seiner inneren Leere, mit seiner durchgängigen Unlauterkeit und
Unwahrhaftigkeit nicht ein schwerer Widerspruch?
Und ein Mittel
zur Aufrechterhaltung dieses problematischen Lebens sollte das Streben
nach Wahrheit sein?
Uns will von allen Arten des Glaubens keiner
gewagter
als
erscheinen
ein
derartiger
unfundierter
Lebensglaube.
Theoretisch
— praktisch.
51
Die eigne Behauptung: der Aktivismus.
d)
Jerusalem hebt im Vorwort zur Übersetzung von James' »Pragma-
meine Annäherung an die neue Richtung hervor und be-
tismus"
»Euckens
merkt dabei:
Aktivismus
ruht
auf
bestimmten
meta-
physischen Voraussetzungen, während der Pragmatismus rein empirisch
Wahrheit begrüße
In
(S. VII).
ist"
ich
sympathisch
die Wahrheit in eine engere Beziehung mit
sie
nicht als eine Sache des bloßen Intellektes
minder
Wahrheit
Sein
da
ich
teile
als
die
das
Streben,
dem Leben zu bringen,
zu verstehen,
nicht
Ablehnung eines Wahrheitsbegriffes, dem
Übereinstimmung
mit
einem
die
neben uns befindlichen
Es fragt sich nur, was unter Leben verstanden wird, und
gilt.
zwischen
dürfte
was
dem,
in
jenen Worten „empirisch" und
Dort be-
„metaphysisch" genannt wird, eine weite Kluft bestehen.
sagt Leben die menschliche Zuständlichkeit, das menschliche Befinden
— ob des Einzelnen oder der Gattung, kommt
hinaus — wir aber denken beim Suchen
selbe
,
schließlich auf das-
einer engeren Ver-
bindung der Wahrheit mit dem Leben an das Geistesleben
Beisichselbstsein
des
Lebens,
aller menschlichen Zuständlichkeit
bildet,
ja
eine
völlige
Daher bedeutet auch
gegenüber etwas wesentlich Neues
Umkehrung
die
als ein
das mit seinen Inhalten und Werten
des
nächsten
Standes
besagt.
Verbindung der Wahrheit mit dem Leben
dem Pragmatismus und dem Aktivismus
etwas sehr verschiedenes:
dort wird die Wahrheit ein bloßes Mittel für ein höheres Ziel,
das
gilt
uns
als eine innere
licher Bestandteil des
Lebens
Zerstörung,
selbst
hier wird sie
und darf daher
und
ein
wesent-
nie ein
bloßes
Mittel werden.
So
wenn
hat es auch hier
die
verschiedenen
und dort einen wesentlich anderen Sinn,
Leistungen
im
Kampf um
die Wahrheit
nach ihrer Fruchtbarkeit für die Lebensentfaltung bemessen werden.
Dort bildet das Maß der Nutzen des Menschen mit all seiner Wandelbarkeit, hier
wird es das Bestehen und der lühalt des Geisteslebens,
und die verschiedenen Behauptungen werden daraufhin zu prüfen
sein, was sie dafür an sachlicher Vertiefung und Erweiterung bringen.
Eine Denk-
Beides kann bis zu vollem Gegensatz auseinandergehen.
weise
kann
dem Menschen
als
Menschen
harte Opfer
—
auferlegen
von allem
und ihm das Leben eher schwerer als leichter machen
wahrhaft Großen läßt sich dieses behaupten -, aber zugleich kann
es das geistige Leben gehaltvoller und größer machen, während um-
Zum Grundbegriff des Geisteslebens.
52
was dem Menschen
gekehrt,
arg herabdrücken
und Genuß
geistig
recht
behagliches Dasein gewährt,
ein
Geistesleben
leer
kann.
Daß
sein
können,
Zeiten
zeigt
voller
das
Leistung
Gegenwart
die
deutlich genug. ^
Mit
Lebens
dem Pragmatismus
in Tätigkeit,
wollen auch wir eine Versetzung des
aber diese Versetzung dünkt uns nicht möglich
vom gegebenen Dasein mit seinen festen Verkettungen aus, sondern
nur durch eine Umkehrung dieses Daseins, durch die Ergreifung
eines neuen Ausgangspunktes
und die Entfaltung eines neuen Lebens.
Das ist eine Art von Metaphysik: wir leugnen
es
nicht,
vielmehr
Um-
verlangen wir aufs entschiedenste eine Metaphysik, da nur in
kehrung der ersten Lage
möglich
und
wird,
irgendwelche
ein ursprüngliches
daher keine
es
Metaphysik
gibt
und
geistige
selbsttätiges
Leben
Selbsterhaltung
ohne
Wiederum kommen wir damit auf
die Notwendigkeit eines selbständigen Geisteslebens als einer
neuen
Stufe der Wirklichkeit, als der Entfaltung ihrer eignen Tiefe.
Nach dem allen scheint uns der Gegensatz des Intellektualismus
und des Voluntarismus
genügt
nicht,
nicht
den Kern der Sache zu
Es
treffen.
den Schwerpunkt des Lebens von der einen seelischen
Tätigkeit in eine andere zu verlegen, da dies keine wesentliche Ver-
änderung und Erhöhung bewirkt und den abgesteckten Kreis nicht
überschreiten
läßt.
Vielmehr
handelt es
sich
um
den Gegensatz
und eines bei aller Emsigkeit innerlich geDamit aber tritt die ganze Frage unter einen
eines selbsttätigen, freien
bundenen Lebens.
wesentlich neuen Anblick.
Intellekt
e)
und Intellektualismus.
Die Eigentümlichkeit der aktivistischen Behauptung dürfte
ehesten erhellen durch eine Betrachtung ihrer Stellung sowohl
Intellektualismus
herabzusetzen,
als
zur intellektuellen Arbeit.
dazu hat
sie
nicht
am
zum
Diese Arbeit irgend
den mindesten Anlaß,
ihr
kann
dem Hauptleben Befindliches und ganz
wohl Entbehrliches gelten. Denn die geforderte Umkehrung des
Lebens, die Wendung zur Selbsttätigkeit, wird sich ohne energische
sie
nicht als
etwas neben
intellektuelle Arbeit nun
*
Für eine weitere
und nimmer vollziehen und behaupten lassen.
Erörterung des
Wahrheitsbegriffs
sei
„Grundlinien einer neuen Lebensanschauung" (1907) verwiesen.
auf
meine
Theoretisch
Hierfür
daß
sie
— praktisch,
53
auch das Zeugnis der Geschichte beachtet, welche
sei
überall
zeigt,
wo von der Erkenntnisarbeit groß gedacht wurde,
da,
nicht als eine bloße Begleitung des Lebens oder als eine nach-
trägliche Deutung eines
sondern vielmehr
Entfaltung
Plato,
wesentliches Stück des Lebens, ohne dessen
Höhe
eigene
seine
es
„gegebenen" fertigen Tatbestandes erschien,
als ein
Spinoza und Leibniz; überall schien
gehalt des Lebens
bei
bei
den Geistes-
ganzen Tiefe nach zu eröffnen und ihn
seiner
dem Menschen zu
vollem Besitze
Unrecht das Wissen
als
Mochte dabei zu
machen.
zu
das Ganze des Lebens erscheinen, jedenfalls
war es mehr als ein bloßes Abbild,
sondern in dem Leben.
jedenfalls stand es nicht neben,
Aber so entschieden wir es demnach ab-
lehnen müssen, den Intellekt
was an der Gegenwart uns
liches Geistesleben verlangt
zum Sündenbock für alles zu machen,
wer ein bei sich selbst befindund nur von ihm einen wesenhaften Inmißfällt:
des menschlichen Lebens erwartet, der
halt
•
und Origenes, so
das Wissen
erst
So
könne.
erreichen
nicht
wie Clemens
so bei Kirchenvätern,
ist
gegen
Hinneig-
alle
ung zu einer intellektualistischen Lebensgestaltung gesichert, der wird
vielmehr die Überflutung namentlich der Neuzeit und Gegenwart
von
intellektualistischen
wird
aber
ihn
Überflutung
in
Strömungen besonders stark empfinden.
anderer Weise betrachten und beurteilen lassen
als
So sei zunächst auf diese Machtentwicklung des
den Voluntaristen.
Intellektualismus das
Auge gerichtet;
dann
erst
läßt
sich
ermessen,
ob die erstrebte Gegenwirkung ihr in Wahrheit gewachsen
a.
Es
Sorge für das Ganze des Geisteslebens diese
die
ist.
Die Überflutung des modernen Lebens durch den Intellektualismus.
Der
Intellektualismus
wirkt
auf
uns
zunächst
von
der
Ge-
schichte her in mannigfacher Gestalt: er wirkt aus dem Altertum, das
Geist
und
Intellekt
dem christlich- kirchlichen Leben, dem
wirkung
immer
zu behandeln
gleichbedeutend
als
wieder
eine
pflegte,
aus
der Glaube trotz aller Gegen-
überwiegend
intellektuelle
Tätigkeit
wurde, aus der Neuzeit, welche die von ihr erstrebte Lebenserhöh-
ung namentlich von der Tätigkeit des Intellektes erwartete. Sie veraber solche Schätzung bis in die Gegenwart hinein nicht nur
ficht
in
der Richtung,
welche
Innenleben ausging,
sie
als
Aufklärung,
tut es fast
erkennen beherrschten Denkweise.
Spekulation
noch stärker
in
der
u. s. w.
vom
vom Natur-
Denn noch immer pflegen die
Zum Grundbegriff des Geisteslebens.
54
Naturforscher Geist und Bewußtsein einander gleichzusetzen und das
Geistesleben als eine bloße Abspiegelung einer draußen befindlichen
Welt zu verstehen.
Dem entspricht es, alles Heil,
auch
Er-
alle
höhung des moralischen Standes, vornehmlich von einer Berichtigung
der Begriffe zu erwarten.^
aber
Nichts
zeigen
oft
Macht des Intellektualismus deutlicher
die
dürfte
daß
Tatsache,
die
als
die
selbst
wieder zu ihm zurückgekehrt
wollte einen neuen Inhalt, aber
ist
Gegenwirkung gegen ihn
So zieht es sich durch
unterlag damit alsbald der feindlichen Macht.
die
ganze Geschichte des Christentums,
Der
19. Jahrhundert hinein.
den
setzung
aller
Kraft
positive
und
irrationale
das
Man
und ihn nur verstärkt hat.
man gab ihn in der alten Form und
so
reicht
spätere Schelling
eingewurzelten
z.
auch
es
das
in
B. sucht mit Ein-
Rationalismus
Denkweise zu verdrängen.
durch
eine
Aber er bringt
Neue als eine bloße Lehre; die Annahme dieser Lehre, das Bezu ihr, scheint das ganze Leben ins Wahre zu stellen.
kenntnis
Was
ist
leicht
macht
das anders als Rationalismus
es
und Intellektualismus?
Viel-
mancher heutige Gegner des Intellektualismus ähnlich
wie Schelling.
Gefährlicher noch, weil tiefer eindringend und
Wirkungen des Intellektualismus durch
sind die
neuen Gewöhnungen
und
alten
Erkenntnisarbeit gaU von
alters
des
her
Denkens.
die
mehr versteckt,
sein Festwurzeln in
Als der Kern der
Heraushebung allgemeiner
Größen aus der grenzenlosen Vielheit der Erscheinungen, im Altertum in vollem Einklang mit der Gesamtauffassung der Wirklichkeit,
indem einfache und unwandelbare Formen ihr Grundgerüst zu bilden
schienen; seit dem Verlassen solcher Überzeugung mit entschiedenem
Unrecht, indem das Heraussehen der großen Züge aus der Erfahrung und die Verbindung der Mannigfaltigkeit zu einem Ganzen weit
^
So
zeigt es
System des
mit
besonderer Anschaulichkeit
19. Jahrhunderts,
das System Comtes.
das gi'ößte
realistische
Es seien nur einige be-
zeichnende Stellen aus dem cours de philosophie positive (4. Aufl. 1877) angeführt.
opinions.
I,
40/41
heißt es:
le
mecanisme
social
repose finalement sur des
Nach IV, 113 trägt die Hauptschuld an der unerquicklichen Lage
der Gegenwart die anarchie intellectuelle; so
eine philosophie convenable.
ist
das dringendste Bedürfnis
Als der tiefste Grund der politischen Korruption
erscheint l'impuissance et le discredit des idees generales.
Überhaupt
ent-
Comte die Epochen der Geschichte den Stufen des Erkennens.
Auch der heutige Monismus glaubt durch eine Berichtigung der Begriffe das
sprechen bei
ganze Leben heben zu können.
— praktisch.
55
verlangt als ein
bloßes Abstrahieren
Theoretisch
mehr besagt und
mehr
weit
gleicher Züge.^
Der
intellektualistischen
Überschätzung des Strebens
zum All-
gemeinen entspricht ein merkwürdiger Kultus abstrakter Begriffe, der
im
Jahrhundert
19.
besondere
eine
Höhe
erreicht
hat.
Wie
viel
Macht üben jetzt Begriffe wie Vernunft, Kultur, Gesetz, Wert, FortHumanität
schritt,
u. s. w.,
Unbestimmtheit scheint
sie
Gerade ihre
Begriffe höchst vager Art!
zu empfehlen,
liebsamen Entscheidung enthebt;
sie
indem
sie
uns einer un-
sind oft wie leere Anweisungen,
Dabei wird Hegel getadelt,
die jeder nach seinem Belieben ausfüllt.
dessen Begriffen doch eine zusammenhängende Gedankenwelt einen
bestimmten Inhalt gab!
Den Einfluß intellektualistischer Denkweise verrät auch die landläufige Neigung, unser Handeln nach Art eines logischen Schlusses als
eine Subsumtion eines einzelnen Falles unter ein allgemeines Gesetz
zu verstehen.
In
Wahrheit würde die wissenschaftliche Arbeit selbst
nicht viel vermögen,
logischen
Formen
erfüllenden
im besonderen nichts neues erreichen, wären die
nicht das bloße
Denkprozesses.
Jenseit
Gefäß eines sie belebenden und
der Wissenschaft aber wird die
Verkehrung noch augenscheinlicher, wenn z. B. das politische Leben,
ja alle menschliche Handlung nach jenem
Schema der Anwendung allgemeiner Sätze auf den besonderen Fall
die richterliche Tätigkeit,
^
Auch der Terminus Abstraktion bekundet jene Wandlung;
ihr ent-
sprechend hat er zwei Hauptphasen durchlaufen, eine logisch-metaphysische
und
eine psychologische,
Abstrakt
zurückgeht.
(s?
von denen jene auf
äipatpEaew? Xeyo'tAeva)
Aristoteles,
diese auf Locke
heißen bei Aristoteles die von
der Materie abgesonderten Formen, namentlich die mathematischen Größen;
so hält es auch das Mittelalter fest (abstrahere
formam a materia
intellectu).
moderne Auffassung sieht im Abstrahieren ein allmähliches Herausheben gemeinsamer Eigenschaften aus der Breite der Erscheinungen. Die
alte Bedeutung überdauert die Herrschaft der antiken Formenlehre; so heißt
Erst die
es
z.
def.
B. in Baumeisters definitiones philosophicae ex systemate Wolfii collectae,
DCCXXXV: absü-ahere ea dicimur, si ea, quae in percepüone distingu-
untur,
tanquam a
re
percepta sejuncta intuemur.
Kant versteht
in
seiner
Logik (s. VIII, 92, Hartenst.) unter Absü-aktion „die Absonderung alles Übrigen,
worin die gegebenen Vorstellungen sich unterscheiden." Er will daher nicht
sagen „etwas abstrahieren" (abstrahere aliquid), sondern „von etwas abstrahieren"
und meint „abstrakte
(conceptus
abstrahentes)
gebrauches erklärt sich
Bedeutungen.
Begriffe sollte
nennen."
man
eigentlich
abstrahierende
Das Schwanken des heutigen Sprach-
zum guten Teil aus dem Durcheinanderlaufen beider
Zum Grundbegriff des Geisteslebens.
56
verstanden
Denn das
wird.
heißt
eine
in
sie
starre
Schablone
und sowohl der Ursprünglichkeit als der Individualität berauben.
Das ist auch eine Wurzel des viel bestrittenen und inpressen
mitten aller Bestreitung unablässig anschwellenden Bureaukratismus.
Schließlich
sei
noch daran
daß der Intellektualismus
erinnert,
mit seiner Neigung, Denken und Geist einander gleichzusetzen und
die
Welt hauptsächlich
handeln,
eingesickert
die
in
tief
als
einen Vorwurf der Betrachtung zu be-
Sprache,
namentlich
in
die der Wissenschaft,
Die Ausdrücke aber, so unverbindlich
ist.
sie
scheinen,
führen leicht unter die Macht der Sache.
So umfängt uns der Intellektualismus von allen Seiten mit einem
dichten
und feingewobenen Netz;
subjektive
aller
Affekt
vermag
daraus nicht zu befreien, selbst die Behauptung des direkten Gegenteiles
wie wir sahen,
lenkt,
zurück.
Wendung
Eine
auf
Umwegen
leicht in die alte
Bahn
könnte nur aus der Anerkennung dessen
hervorgehen, daß die eigene Arbeit des Intellektes einen positiven
und produktiven Charakter erst erlangt, wenn sie sich einem Ganzen
des Geisteslebens empfangend und fördernd einfügt, wenn sie von
gesamtgeistigen Synthesen geleitet und von solchen Energien getrieben
Daß
wird.
als
aber
es
Wahrheit so
in
steht,
läßt sich
sowohl direkt
indirekt erweisen: alle echte Leistung intellektueller Art stand in
engem Zusammenhange mit Gesamtbewegungen des Geisteslebens; wo
immer hingegen die Arbeit solchen Zusammenhang fallen ließ, da ist sie
rasch zu leerem Formalismus oder schwankender Reflexion gesunken.
Ein solches Verfechten der Abhängigkeit des Intellektes
ist
mit der Anerkennung einer Größe
Ganzen
vom Ganzen
und Bedeutung innerhalb des
aufs beste vereinbar.
ß.
Die Begründung des Erkennens im Lebensprozesse.
Wer vom Erkennen
ein Registrieren
seine nähere Gestaltung
Geisteslebens
gering denkt, wer in ihm nicht mehr
bloßer Erscheinungen
keinerlei
sieht,
als
der braucht sich über
und über sein Verhältnis zum Ganzen des
zu machen. Wer aber in ihm eine
Sorge
Durchleuchtung und innere Aneignung der Wirklichkeit sucht, dem
wird jenes zu einem schweren Probleme.
Wirklichkeit
unterwerfen
und aneignen,
irgendwelches Vermögen an
die Widerstände
sie
Wie läßt sich eine fremde
wenn die Arbeit nicht
heranbringt, nicht eine Kraft gegen
der Dinge einzusetzen
hat,
wie kann eine Erfahr-
ung uns wertvoll werden, wenn sie nicht eine von innen kommende
Theoretisch
— praktisch.
57
Bewegung aufnimmt und weiterführt, wie kann sie eine Antwort erteilen, wenn ihr nicht zuvor eine Frage gestellt ist?
Woher aber
soll das Erkennen die Kraft zu jener Leistung finden, wenn nicht
der gesamte Lebensprozeß eine innere Konzentration vollzieht, seine
zu einem Ganzen verbindet und aus ihm
Tätigkeiten
den Kampf
Umgebung aufnimmt? Eine derartige Bewegung würde
aller
Betätigung,
wie
so auch dem Erkennen eine eigentümliche Art
gegen die
und Richtung
Ein solches Zusammenschießen des Lebens
erteilen.
zu einem charakterhaften Ganzen steckt einen eigentümlichen
seinsraum ab und
eigentümlich wie die Erfahrung,
gestaltet
Da-
so das
Grundverhältnis des Menschen zur Wirklichkeit und seine Arbeits-
Daraus hat auch
welt.
das Erkennen
Wie könnte
empfangen.
Großen und Unterscheidenden
die
die
seine
und Wege zu
Ziele
einer die griechische Philosophie in
dem
ihrer Art verstehen, der sie nicht als
Wendung derjenigen Lebenssynthese zur Wissenschaft verstünde,
dem Ganzen der griechischen Kultur zugrunde liegt? Diese
Synthese
erfolgte
unabhängig von der
nicht
intellektuellen Arbeit,
war nicht
sie
bedurfte vielmehr ihrer unablässigen Hülfe;
ein
Werk des bloß auf seine eigene Kraft gestellten Erkennens. So
überhaupt nur ein Erkennen,
besitzt
gegründet
ist
aber
sie
das in einer Lebenssynthese
und aus ihrer Tiefe schöpft,
sichere Richtungen
und
zwingende Notwendigkeiten, nur ein solches vermag den Gegenstand
zu erfassen und zu durchdringen, nur ein solches kann der Wirk-
Zusammenhang geben. Warum macht die
lichkeit einen
lebendigen
Scholastik
bei
allem
Eindruck,
warum hat sie, trotz breitester Wirkung im menschlichen
Fleiß
und allem Geschick einen so dürftigen
Kreise, geistig nichts Erhebliches- gefördert?
Weil ihr die gestaltende
Kraft eines charakteristischen Lebens fehlte, weil sie daher ihren Be-
keine
griffen
innere
zu geben, vermochte.
Nähe und keine zwingende Überzeugungskraft
Schon deshalb mußte sie der neueren Philo-
sophie erliegen, weil in dieser und durch sie ein neues Leben emporstieg.
Eben
dieses
ist
Leibniz
und
Kant,
von
es
auch,
tüchtigen
was schöpferische
Schulgelehrten,
Denker,
wie
wie Wolff und
Herbart, unterscheidet,
daß jene neue Lebenssynthesen zum Durch-
bruch bringen und
ihrer Arbeit eine
vollziehen.
So sind
in
siö
Erhöhung der Wirklichkeit
Mehrer im Reich des
Geistes,
nicht bloße
Bearbeiter und Zergliederer eines gegebenen Befundes, sie reflektieren
nicht
bloß über die Wirklichkeit,
sie
erweitern unsere Wirklichkeit.
Es bekräftigt aber solchen Zusammenhang des Erkennens mit
Zum Grundbegriff des Geisteslebens.
58
dem Ganzen des Geisteslebens nichts nachdrüci<licher als die eigenen
Erfahrungen
der
Logik,
die
enthoben glaubt.
leicht aller
Die Unverbrüchlichkeit jener Gesetze ist klar und
Aber
unbestritten.
und
Bindung und Beziehung
sich mit der Allgemeingültigkeit
Umwandelbarkeit ihrer Gesetze
alle
und Formen ergeben keineswegs
Gesetze
schon ein lebendiges Denken; das wirkliche Denken des Menschen
ist
keineswegs bloß eine gleichförmige Anwendung jener Denkgesetze,
es hat
darüber hinaus eine Eigentümlichkeit, die alle Mannigfaltigkeit
beherrscht
und durchdringt, und
die
Lebensprozesses hervorgehen kann.
nur aus dem Ganzen eines
Demgemäß
das Denken in
ist
seiner feineren Struktur verschieden nach der Art des Lebensganzen,
dem es angehört.
Daß das künstlerisch gerichtete Griechentum das
Denken enger mit der Anschauung zusammenschloß, daß es rasch
und unmittelbar zu einer Synthese strebte und alles Grenzenlose
mied, daß es die Elemente des Lebens als gegeben und unveränder-
hinnahm,
lich
das
auch seine Erkenntnisarbeit
gestaltet
einzelnen logischen Operationen
ferner die
Denkweise des
alters den Einfluß eines
eigentümlich.
Wie
ausgehenden Altertums
bis in
die
sehr bekundet
und des Mittel-
neuen, von der Religion beherrschten Lebens!
Indem dort alles sichtbare Dasein zum bloßen Gleichnis einer unsichtbaren
die
Ordnung
Behauptungen
ahnt
wird,
verlieren
die Begriffe
ihre Ausschließlichkeit.
und schaut durch den
ihre Sprödigkeit,
Die allegorische Deutung
sinnfälligen Tatbestand hindurch eine
höhere Welt, ohne dabei jenen zu einer gleichgültigen Erscheinung
So ist derselbe Gegenstand Bild und Sache, Sinnund Geistiges in Einem; daß darin zugleich eine Bindung
herabzusetzen.
liches
und eine Befreiung, eine Bejahung und eine Verneinung, damit aber
ein unhaltbarer Widerspruch liegt, das empfindet jene von Stimmung
und Ahnung beherrschte, beinahe traumhafte Denkweise
dieser
Denkweise aber ruht der
Sakramentenlehre u. s. w.
nicht.
mittelalterliche Kirchenbegriff,
Auf
die
Die Höhe der Scholastik wird klarer und
nüchterner, aber da es ihr bei aller Tüchtigkeit syllogistischen Ver-
fahrens an einer selbständigen Synthese
und an einer entsprechenden
Energie des Denkens fehlt, so fehlt ihr auch die Kraft eines disjunktiven
Verfahrens; grundverschiedene Welten, wie der weltfrohe Aristotelismus
und das weltfremde alte Christentum, ferner innerhalb des Christentums die kirchliche Ordnung und die alle Gestalt überfliegende
Mystik, vertragen hier sich miteinander friedlich und freundlich; sie
scheinen zu völliger Harmonie gebracht,
wenn eine geschickte An-
Theoretisch
— praktisch.
59
Ordnung und Abstufung einen direkten Zusammenstoß verhütet. Man
als auch, wo einem kräftigen Denken
denkt im Schema eines Sowohl
Entweder
sich alsbald ein
—
— oder ergeben
hätte.
Auch
die
logische
Methode der neuen Wissenschaft mit ihrer energischeren Disjunktion
und ihrer schärferen Analyse, ihrem Flüssigmachen auch der Elemente
und ihrem Streben ins Unbegrenzte, sie zeigt deutlich genug einen
Zusammenhang
engen
dem modernen
mit.
Lebensideal
der Kraft
und Bewegung.
wer
So bejaht eine besondere Art des Geisteslebens,
dieser Art der Forschung den Typus aller Forschung sieht.
in
Wie aber jede ausgeprägte Zeit
so
ihre besondere Art
der Logik hat,
auch jeder selbständige Denker; ohne eigene Logik kann
hat sie
keine eigene Denkweise, keine eigene Lebensgestaltung bestehen.
kräftiger diese Gestaltung,
desto tiefer
wird ihr Einfluß bis
Je
in die
einfachsten Elemente und Tätigkeiten des Denkens sich hineinerstrecken.
So
muß die Verkettung mit dem Lebensganzen die Denkarbeit
konkreter, individueller, reicher gestalten.
neue -Fragen und Aufgaben.
dem Lebensganzen
Es
gilt
Zugleich entstehen damit
zu zeigen, was das Erkennen
näher aufzuweisen,
wie es in ihm zur
vom
zur Verkettung der
leistet,
Wesentlichen,
Scheidung des
Zufälligen
Mannigfaltigkeit,
zur Herausbildung einer Allgemeingültigkeit wirkt.
Auf den
mag gerade die Allgemeingültigkeit als bedroht
ersten Blick
wenn das Erkennen in so nahe Beziehung zu besonderen
Lebenssystemen tritt. Wird sich nicht damit die Wahrheit in eine
Vielheit von Wahrheiten auflösen und ein zerstörender Relativismus
erscheinen,
gänzlich
Das würde doch nur der Fall
das Feld gewinnen?
sein,
wenn alle Lebenssynthesen gleichwertig nebeneinander stünden, nicht
ihrer aller Arbeit einer einzigen umfassenden Synthese diente, an der sich
messen hätte. Aber könnte nicht eine solche Synthese der Bewegung als Endziel vorschweben und zugleich von Anfang an zur
alles zu
und Richtung des Lebens und mit ihm des Denkens
Gestaltung
wirken?
Daß
Behauptung neue Probleme hervorruft,
eine
gegen
keineswegs
sie;
sind
Grundanschauung nicht sowohl
Y-
Daß
stätigt
es
jede
spricht
echte Probleme, so werden sie die
es
belasten als verstärken.
Die bewegende Kraft im Wahrheitsstreben.
sich
in
der Tat
Erwägung der
um echte Probleme handelt, das bewas im Kampf um die Wahrheit
Frage,
Macht hat und die Entscheidung bringt.
Daß es nicht die bloßen
Gründe und Beweise sind, das läßt uns jede Disputation zwischen
Zum Grundbegriff des Geisteslebens.
60
abweichenden Überzeugungen
Gedankenarbeit,
anders
sollte es
auf
dem
beim Zusammenstoß der Geister in
größeren Felde der Disputation,
der
ersehen;
deutlich
stehen?
Jeder überträgt die
Gründe
des anderen in die eigene Sprache und Denkweise und verändert sie
damit völlig; so steht gewöhnlich Monolog neben Monolog, zu einem
wirklichen Dialoge
kommt es selten.
Überzeugungskraft
ihre
nicht
ihr
In
Wahrheit gibt den Gründen
logisches
und
dialektisches Ver-
mögen, sondern der Gehalt und die Gewalt des Geisteslebens, der
geistigen Konzentrationen, der Lebensenergien, aus denen sie schöpfen.
So
verteidigt in aller Erörterung prinzipieller Fragen
im Grunde ein
jeder sich selbst und seine eigene Art; erst aus solcher geistigen Selbst-
erhaltung strömt Kraft,
Glut und Leidenschaft
Eine fruchtbare Aussprache und die Möglichkeit einer
Bewegung.
Verständigung entsteht nur
Art
in die intellektuelle
gemeinsamen
einen
da,
wo die Verwandtschaft der geistigen
Boden
würden
bereitet;
Aristoteles
und
Thomas und Voltaire, sie alle treffliche Logiker, sich wohl
gegenseitig überzeugt haben, wenn sie noch so lange miteinander
gestritten hätten? Mur einen flachen und haltlosen Menschen können
bloße Gründe aus seiner geistigen Art hinauswerfen; auf bloß
intellektuelle Erwägung gestellt, könnte der Mensch seines eigenen
Wesens nimmer sicher und froh werden; denn er müßte in steter
Furcht sein, daß nicht ein stärkerer Dialektiker komme, ihn überwinde und zum Gegenteil zwinge.
Augustin,
So sind es auch im geschichtlichen Leben
Gedankenbilder,
geistigen
die
Energien,
freischwebenden
die
nicht die abgelösten
sondern es sind die
Ideen,
Lebenskonzentrationen, welche die Geister
beherrschen und die Leidenschaften entzünden.
bewußte Anhänger der Hegeischen
die
mit
Ideen
hervortreiben,
weitertreibt
als
überwältigender
und daß
Notwendigkeit
nichts stärker aufrüttelt,
logischer Widerspruch,
ein
Bewußte und un-
Denkweise sagen uns oft, daß
ihre
Konsequenzen
nichts zwingender
Gewiß, Konsequenzen
und Widersprüche
können eine unwiderstehliche Gewalt über den
Menschen erlangen.
Aber sie tun das nicht von der bloßen Logik
aus.
Konsequenzen
nicht gezogen,
doch
nicht
können sehr
nahe
empfunden.
Es
kommt
und werden doch
hier alles darauf an,
daß die
aufgenommen werden, daß
Lebensenergien entfalten und ein geistiger
Probleme
in die geistige Selbsterhaltung
durch
hindurch sich
sie
liegen
Widersprüche mögen handgreiflich sein und werden
Existenzkreis gestaltet; nur eine solche Aneignung, eine solche Auf-
— praktisch.
Theoretisch
61
macht
die
Konsequenzen unabweisbar und die Widersprüche unerträglich;
es
nähme
ist
der
Arbeit
intellektuellen
Eigenleben
das
in
der Einigung, die Kraft der Zusammen-
im besonderen der Grad
welche der Logik die Macht
fassung des Lebens,
die
erst verleiht,
Vermögen zu üben wähnt. Das ruhige Ertragen
widersprechender Gedanken massen verrät immer eine geringere
Konzentration des Lebens; wie jenes für das kindliche Denken
charakteristisch ist, so ist es das auch für naivere Zeiten und für
sie oft aus eigenem
den Durchschnittsstand der Menschheit gegenüber den Forderungen
selbständiger
Geistigkeit.
Der Mangel an Logik verschuldet
nicht,
er bekundet nur jenen Mangel.
So
licher
und
der geistigen Lage der Gegenwart nichts verdrieß-
in
ist
hemmender,
nichts
die Unempfindlichkeit
als
Widerspruch von Gedanken massen;
an Lebensenergie
Selbsttätigkeit inmitten
sie
verrät einen starken
den
Mangel
wahrhaftigem Eigenleben und
an
zentraler Art,
für
rührigster Betätigung.
Der Lebensbestand
ist
Kompromisse an der Oberfläche sollen sie beschwichtigen, sie scheinen ausgeglichen, wenn die
Schroffheit des direkten Zusammenstoßes einigermaßen vermindert
heute voll fundamentaler Gegensätze;
ist.
Oder auch
Widerspruch
vermengt.
alten
Gedankenmassen werden
die
unbedenklich
ineinander
bei allem sachlichen
geschoben
und miteinander
Wie oft müssen sich die grundverschiedenen Welten des
und der modernen KulturentwickNicht minder laufen
solche Behandlung gefallen lassen.
ethisch-religiösen Idealismus
lung eine
bei allermodernsten
Schriftstellern
So
ungen durcheinander.
z.
grundverschiedene Lebensstimm-
B. bei Nietzsche antike
und moderne,
klassische und romantische, künstlerische und dynamische Denkweise;
wer für solche Dinge ein Ohr
der
muß darin
grelle
Disso-
Die Masse der sog. Gebildeten aber, wie
nanzen empfinden.
alles
hat,
kräftigen Eigenlebens entbehrt,
sie
empfindet nicht den mindesten
Schmerz an geistigen Dissonanzen, eher sieht sie darin
ein
buntes
und unterhaltendes Spiel; je mehr Widersprüche, desto «eigenartiger«,
desto «interessanter"!
Nichts
zeigt
die
Abhängigkeit des Denkens von der Energie
Lebens greifbarer
des
geistigen
Alle
eingreifenden
der
jeweilige
als
die
Bewegungen der Religion.
Wendungen kamen hier dadurch in
Befund
unerträgliche Widersprüche
Fluß,
daß
empfinden
ließ,
daß im besonderen, was der Lauf der Zeit und die Anpassung an
die menschliche Lage an äußeren Einrichtungen,
Übungen, Formeln
Zum Grundbegriff des Geisteslebens.
62
hervorgebracht
mit den Forderungen einer gesteigerten Inner-
hatte,
Aber wie wenig war die
zusammenstieß.
unversöhnlich
hchkeit
Empfindung, Durchlebung, Überwindung solcher Widersprüche das
Werk bloßlogischer Betrachtung!
lag der Kontrast zwischen
und dem Verlangen
vor
aller
Im Zeitalter der Reformation
z.
B.
der Veräußerlichung des Kirchenwesens
ernster
Seelen
nach Verinnerlichung deutlich
Augen; der größte Gelehrte jener Zeit, Erasmus, sah
ihn,
Warum ist
nach Ausweis seiner Schriften, ebenso klar wie Luther.
nun Luther der Reformator geworden und nicht Erasmus? Sicherlich
nicht, weil jener der größere Logiker war, denn das war entschieden
Erasmus.
Sondern weil ihm jene Lage mit ihren Widersprüchen
und geistreicher Reflexion blieb,
nicht eine Sache kühler Betrachtung
sondern weil
sie
ihm zu einer persönlichen Angelegenheit und zu-
gleich zu einem heftigen Schmerz, einer unerträglichen Notlage wurde.
Bei solcher völligen Aneignung wurde ihm eine Lösung des Konfliktes
zur zwingenden Notwendigkeit,
zur Seele seines Lebens,
geistigen Selbsterhaltung, die alle andere
zu einer
Erwägung mit elementarer
Die Gewalt einer solchen Selbsterhaltung gab
Wucht verdrängte.
dem schlichten Manne das Vermögen und auch das Recht, eine
überkommene, in der Überzeugung der Menschheit geheiligte Ordnung
anzugreifen
aus
und den Aufbau
ursprünglicher
Helden, an
geistiger
einer
das Wirken
neuen zu wagen;
Notwendigkeit
machte ihn zu
einem
dem gemessen Erasmus mit allem überlegenen Wissen,
Geschmack und Scharfsinn
als
klein erscheint
Entscheiden so in den geistigen Kämpfen nicht freischwebende
intellektuelle
Erwägungen, sondern die begründenden Lebensprozesse
und der Gehalt der von ihnen umspannten
geistigen Wirklichkeit,
so sind auf sie die Gedankenmasseh zurückzuführen, so hängt aller
wesentliche Fortschritt an einer Weitererschließung jener Wirklichkeit.
So vollzog und
vollzieht sich die Überwindung alternder Gedanken-
massen nicht durch
ein plötzliches Erscheinen
sondern dadurch, daß die Schranken des
in
überlegener Gründe,
jenen Gedankenmassen
verkörperten Lebens bemerklich werden, neue Konzentrationen oder
doch Bewegungen aufstreben, durch
das in scheinbar sicherem
wird;
mag
es äußerlich
Besitz
Befindliche veraltet
sicher zu herrschen glaubt.
ist
Leben
und entwertet
den früheren Glanz behaupten,
die Herrschaft über die Seelen; es
Ideen in die Energien,
ein frisch sich regendes
geschlagen, auch
es verliert
wo es noch
Daß sich so die Entscheidung von den
von den
intellektuellen
Erwägungen
in
die
Theoretisch
schöpferischen
— praktisch.
53
das
muß zur Vertiefung
Lebensentfaltungen
verlegt,
der Arbeit und zur Befestigung des Strebens wirken.
Unvergleichlich
größer wird das Bild des geschichtlichen Lebens,
wenn
sowohl Lehren gegen Lehren
wird das Problem
durchgängig
stehen;
hier nicht
Lebensmächte gegen Lebensmächte
als
zurückverlegt,
beträchtlich
wenn es die Wurzeln der Lehren erst zu ermitteln, die letzten Triebkräfte erst aufzudecken,
stoßes
den entscheidenden Punkt des Zusammen-
herauszustellen
erst
Aber
gilt.
alle
Mühe der Arbeit wird
dann von der Überzeugung getragen und beseelt, daß im Menschenleben ursprünglichere Kräfte, tiefer gegründete Notwendigkeiten walten,
Gedankenarbeit aus eigenem Vermögen
als die
8.
sie
Konsequenzen für die Erkenntnisarbeit.
Solche Verbindung der Erkenntnisarbeit mit
und dem Aufbau
Geisteslebens
aufbringen kann.
dem Ganzen des
einer geistigen Wirklichkeit
muß für
ihre
eigene Gestaltung eingreifende Folgen haben;
bei
'
der dringend gebotenen Beschränkung nur insofern betrachtet,
als
mit der Lösung wichtige Aufgaben lösbar oder doch angreifbar
diese seien hier
werden, die sich sonst einer erfolgreichen Behandlung entziehen.
Immer noch waltet darüber viel Unsicherheit, wie die Philosophie
eine
selbständige Aufgabe
finden
könne.
jener
zur
Einigung
Die
Einheit
oft
verbinden,
entweder
ist
gegenüber den einzelnen Wissenschaften
erteilte
eine
Antwort,
kann
bloße
sie
nicht
solle
die
genügen.
Zusammenstellung,
Ergebnisse
Denn
jene
dann
muß
man sehr freigebig mit der Bezeichnung Wissenschaft sein, um eine
solche Enzyklopädie als Wissenschaft anzuerkennen, oder sie bedeutet
eine Weiterbildung
Prinzipes,
Prinzip
bloß
aus
kann
und Umwandlung, dann bedarf es eines neuen
dem eine solche hervorgehen könnte. Das gesuchte
nun weder von außen gegeben werden, noch aus
intellektuellen
Punkt, der uns zugänglich
ungen
es muß im Ganzen
kommen wir auf den letzten
Bewegungen entspringen,
des Lebensprozesses liegen.
Hier
ist;
erst
nach seiner Art und seinen Erfahr-
muß sich das Grundverhältnis des Menschen zur Wirklichkeit,
sowie die Bedeutung seines Lebens und Seins bestimmen;
erst
von
hier aus kann eine Verbindung, eine Abschätzung, eine Weiterbildung
der Ergebnisse der einzelnen Wissenschaften erfolgen.
prozeß
erst
tritt
Dieser Grund-
uns nicht im unmittelbaren Eindruck entgegen,
herauszuarbeiten;
das aber
ist
die
er
ist
Aufgabe der Zentraldisziplin
der Philosophie, die von alters her Metaphysik heißt; die anderen
Zum Grundbegriff des Geisteslebens.
64
haben dann die neue Beleuchtung den einzelnen Ge-
Disziplinen
zuzuführen.
bieten
Solche
Fassung
engen Zu-
den
auch
erklärt
sammenhang der Philosophie mit der Persönlichkeit des Menschen,
ohne jene zu einer bloßen Ausstrahlung der individuellen Art herab-
Denn
zusetzen.
ohne Erweisung einer
erfolgen
nicht
Lebens;
Durchdringen zu jenem Grundprozesse kann
das
insofern
letzthin
ist
Weite und Tiefe des
Kraft,
Maß
das
Maß
des Lebens auch das
des Denkens.
Wendung dem
Eine wesentliche Förderung bringt ferner jene
Daß
Probleme der Wahrheit.
wenn
wäre,
sie
auf Wahrheit endgültig zu verzichten
Übereinstimmung unseres Denkens mit einer
eine
draußen befindlichen Welt bedeutete, darüber
Um so zweifelhafter
Mit
könnte.
Anknüpfung
der
heute kein Zweifel.
Nein entgegentreten
Lebensprozeß
das
erhält
es gibt keine intellektuelle
Wahr-
ohne eine gesamtgeistige Wahrheit, diese aber bedeutet
nichts
anderes
als eine
Verwandlung der Welt in Eigenleben, eine innere
Bewältigung der Wirklichkeit.
den
den
an
Problem eine neue Beleuchtung;
heit
ist
das solchem
das Ja,
ist
Menschen
hinaus
das
Das
hat zur Voraussetzung,
den
Geistesleben
den Menschen aber
Wirklichkeit bildet;
für
aufhörliches Streben
und Weiterstreben,
ist
daß über
Grund
der
die Sache ein
un-
letzten
ein Vordringen
und Empor-
klimmen, ein wachsendes Ringen mit den Widerständen ungeistiger
und
Innerhalb dieses Strebens gibt
halbgeistiger Art.
sahen,
fruchtbares
keinerlei
Erkennen
des
sächlichkeit
zunächst nicht mehr
als
Versuche,
der Welt drinnen und draußen können
es,
wie wir
Begründung
eine
Aber diese Synthesen sind
Synthesen
Lebens.
ohne
erst
bei
aller
in
Tat-
im Kampf mit
Vermögen erweisen;
sie ihr
das Erkennen aber hat bei solcher Auseinandersetzung eine leitende
Stellung, es ist unentbehrlich zur Klärung und Prüfung, unentbehrlich
zur Erringung der Allgemeingültigkeit, zur Austreibung
alles
Klein-
menschlichen und zur Herausbildung des Weltcharakters des GeistesEin
lebens.
solches
kritisches
Wirken
kann
es
nicht
üben ohne
ein gewisses Heraustreten aus der Besonderheit jener Synthese, aber
die Kritik kann
nicht
weiterführen,
wenn
sie
nicht einer neu
auf-
steigenden Synthese dient.
Als
festen
weiteres
Problem
gehört
Zusammenhang des Menschen
ging,
hierher
die
Ausgangspunkt der Erkenntnisarbeit.
ist
mit
Frage
Seit
der
der sinnlichen
jene Frage unabweisbar geworden.
nach
einem
unmittelbare
Welt verloren
Aber vergeblich hat
Theoretisch
Erkennen
das
einen
— praktisch.
Punkt
festen
bei
sich
55
und
Letzte
erreichen
Nicht anders
Sicherste auftrat.
als
gesucht;
selbst
wieder erschienen unerwiesene Voraussetzungen
ist
in
immer
dem, was als das
jener feste Punkt zu
durch eine Zusammenfassung des ganzen Lebens zur
Einheit
und durch eine gleichzeitige Verwandlung in eigene Tat; nur
damit
kann eine axiomatische Gewißheit entstehen und auch
Erkennen
zugehen.
bedeutet diese Einheit
im
dem
Für
den
stets
eine Aufgabe; eine volle Tatsache könnte
Streben
befindlichen
Menschen
Weges bringen, was in unabsehbarer Ferne liegt.
Aber das Streben selbst wäre unmöglich, wenn nicht das, was für
uns Menschen eine unermeßliche Aufgabe ist, im Geistesleben selbst
erst
das Ende des
die
begründende Tatsache wäre.
Es
ein alter Einwand
ist
gegen die Philosophie, daß sie lediglich
Meinungen neben Meinungen
stelle
und
diese
tausende bis zur Unübersehbarkeit anhäufe,
im Lauf der Jahr-
ohne
die spätere Leist-
ung der früheren sicher überlegen zu machen. Gewiß verbleibt in der
Philosophie ein Element der Freiheit und der Entscheidung, sie teilt
mit der Religion, der Moral, der Kunst und allen edlen Dingen die
Eigenschaft,
immerfort der eigenen Tat zu bedürfen und sich nie-
mandem aufzwingen zu lassen.
Aber deshalb wird
sie
noch kein
bloßes Neben- und Nacheinander menschlicher Meinungen: davor behütet sie sicher die Erkenntnis ihres engen Zusammenhanges mit dem
Das bringt
Bewegung in enge Verkettung mit der Evolution
Lebens in der Menschheit, und wie die Wendepunkte
Streben des Menschen nach einer geistigen Wirklichkeit.
ihre geschichtliche
des geistigen
dieser Evolution
fundamentale Tatsachen erschließen, so treiben
auch die philosophische Arbeit
in
neue Bahnen.
sie
Unser Verhältnis
zu den großen Problemen kann nicht mehr das der alten Griechen
sein,
seit
prozeß
das Christentum so durchgreifende Wandlungen im Lebens-
vollzogen,
so
schwere Konflikte,
Tiefen in ihm aufgedeckt hat; es
wachsen,
seit
hat.
solche
fruchtbare
dem Mittelalter entdem Menschen und der
und das Innenleben zu größerer
Zeigt
so
aber auch
die Neuzeit schärfer zwischen
Welt geschieden
geweckt
ist
aber auch
Selbständigkeit
Erfahrung nicht den Denker
in
enger
Verbindung mit der Geschichte und dem Ganzen der Menschheit?
Denn
immer nur Mög-
Seine Selbständigkeit braucht er darüber nicht einzubüßen.
was
die
lichkeiten
Umgebung dem Menschen zuführt,
sind
und Anregungen; eine Wirklichkeit und
greifbare
eine
Gestalt erwächst daraus nur durch eine vordringende Tat, die immer
Eucken, Grundbegriffe.
4. Aufl.
5
Zum Grundbegriff des Geisteslebens.
66
So wird das eine auf das andere
eine Sache des Einzelnen bleibt.
angewiesen, das Ganze aber, das beides umspannt, gewinnt unver-
kennbar an Größe.
£.
Konsequenzen für die Behandlung der Geschichte der Philosophie.
Die Anerkennung solches Zusammenhanges der Philosophie mit
dem Ganzen
muß auch auf die Behandlung der Ge-
des Lebens
schichte
der Philosophie einen starken Einfluß üben.
kann
genügen,
es
einzelnen
die
Systeme
in
mehr
Nicht
ihrem unmittelbaren
Bestände darzustellen und aneinanderzureihen, sondern zur Aufgabe
wird,
begründenden
die
Lebensinhalte
und
herauszuarbeiten
die
Leistung der Denker dadurch in größere Zusammenhänge zu stellen.
Nicht sowohl was jene sagten,
als
und
Aussprache
welche
Art
geistige
ihre
wie
dazu kamen, es zu sagen,
sie
zeigte,
wird
jetzt
Hauptproblem.
Das
geschichtlichen
und menschlichen Umgebung aufzuklären,
zum
das Verhältnis des Denkers zu seiner
zwingt,
freilich
nicht in der Art eines landläufigen kulturgeschichtlichen Verfahrens,
das
Dinge auf den
die
Kopf
indem
stellt,
es
das
Innere
vom
Äußeren, das Große aus der Summierung des Kleinen, das Ewige
vom Zeitlichen
herleitet.
aber wird sich
nunmehr danach bemessen, was sie für die Eröff-
nung neuer Tiefen,
In
taten.
diesem
für
Die Bedeutung der einzelnen Leistungen
die Erweiterung
Sinne
alles
ist
der geistigen Wirklichkeit
große Denken
ein
Vordringen,
Neubilden und Schaffen.
Macht solche Zurückbeziehung des philosophischen Strebens
die
Behandlung seiner Geschichte
in
gewisser Richtung verwickelt,
Denn nach jenem Maß-
so wirkt sie in anderer zur Vereinfachung.
stabe gemessen,
heit
als
dürfen nur einige wenige Erscheinungen in Wahr-
schöpferisch,
als
Weiterbildner
der Substanz des Lebens
gelten; auch heben sich dann aus der scheinbar chaotischen Fülle
einige wenige
Typen hervor,
wie der Begriffe
sich
schärfer
meiste
oder
ein
die bei aller
subtile
lebens.
Erörterung,
der Menschen,
Wir
So mag
Kern von der bloßen Umhüllung scheiden.
zeigt,
ist
bloße
Das>
Um-
Gelehrsamkeit verschiedener Art, mehr
minder geistreiches Räsonnement,
schäftigung
Veränderung der Lagen
der Hauptsache immer wiederkehren.
was der unmittelbare Anblick
aber,
gebung:
in
untauglich
alles
das tauglich zur Be-
zur Erhöhung des Geistes-
sind ärmer und reicher, als wir gewöhnlich meinen,
ärmer im Umfang, reicher im Inhalt unseres
Besitzes.
— praktisch.
Theoretisch
bloßen Form des Systems entgegenwirken, die
von der Hauptsei
keines-
Die Verkettung zum System drängt die einzelnen
unterschätzt.
Sätze enger
leicht
Die Bedeutung der systematischen Form
sache ablenkt.
wegs
57
mag jene Zurückverlegung auch der Überschätzung der
Endlich
zusammen,
sie
Widersprüche schärfer heraus,
treibt die
zur gleichmäßigen Durchbildung, zur Organisation der Ge-
sie wirkt
Aber das
dankenwelt.
alles
tut
nur unter Voraussetzung eines
sie
lebendigen und belebenden Inhaltes, den nur Synthesen und Energien
des gesamten Lebens erzeugen.
Fehlt solcher Inhalt,
so kann
alle
und Geschicklichkeit in Aufbau und Anordnung das
System nicht vor einem Sinken zu einem leeren Gehäuse bewahren.
logische Kraft
Wie
ist
viel
durchgearbeiteter
ist
Wolffs System
die schroffen
Ausbau
Gedanken
seiner
gewirkt
verhindert,
der Geisteswelt
wie
wenig andere.
und doch
auch
auf
als
gerichtet,
der bloßen
die
hat er durch seine
Gedankenarbeit so
Auf jenes Wesentliche, auf
schaffende Kraft und den treibenden Kern,
Augenmerk
das von Leibniz,
Widersprüche seines Wesens an einem systematischen
Weiterbildungen
stark
als
Augustin wurde schon durch
jener damit der größere Philosoph?
sei
Form aber
die
also vornehmlich das
nicht
mehr gegeben
ihr gebührt.
Doch genug der Erörterungen; auch
würden
sie
bloße Ausschnitte aus einem
bei weiterer
Ausführung
größeren Gedankenkreise
bleiben.
Wir verweilten hier et\('as länger, weil der Nachweis wichtig
schien,
daß das eigene Interesse der Erkenntnisarbeit über die bloße
Erkenntnisarbeit hinaustreibt.
Zugleich aber wurde ersichtlich, daß es
nach einer anderen Richtung
treibt als
Vielleicht
manche der Männer,
stehen
pflegen,
dem
könnten
wir nur freudig begrüßen.
nach der des Voluntarismus.
die Voluntaristen
nahe, was wir versuchen.
Persönlichkeiten
stehen
zu heißen
Solche Übereinstimmung
Aber wie immer es mit den
mag, der Unterschied einer Verschiebung
innerhalb des Seelenlebens und einer Erhebung über alles empirische
Seelenleben
sei
in
keiner Weise verdunkelt.
Idealismus
a)
Die Ausdrücke.
|ie Ausdrücke Idealismus
I
und Realismus sind durch übermäßigen
*-^ Gebrauch so verschliffen
Wissenschaft
treten
sie
— Realismus.
und
sind.
für
die
ver-
einen alten und bleibenden Gegensatz,
bewegende
Frage
Gegenwart
der
daß
Aber immerhin
abgegriffen,
unbrauchbar geworden
fast
ist.
Wegen
sie
der zugleich eine
solcher
Beziehung
mögen zunächst die Termini kurz erläutert sein.
Das Wort Idealist taucht in der Philosophie zuerst gegen das
Ende des 17. Jahrhunderts auf;^ wenn Leibniz das Wort im Gegensatz
zu Materialist
(s.
186a Erdm.) verwendet wie sonst Formalist,
so denkt er dabei an Philosophen, welche wie Plato
in
der
Form das Wesen der Dinge sehen.
und Aristoteles
Alsbald aber übte die
moderne Bedeutung des Wortes Idee eine Wirkung auch hierher.
Aus einer urbildlichen Form wurde Idee, zuerst in der französischen
Sprache, zur bloßen Vorstellung, zu einem subjektiven Gedankenbilde;
durch Descartes und Locke drang — nicht ohne Widerspruch
Neuerung auch
System, das
die
Realität
alle
in
Lehre
wöhnlich
in
Idealismus bedeutete dann ein
Wirklichkeit jenseit der Vorstellungen
einer
Berkeley's
die Philosophie;
Außenwelt
mit jenem
tadelndem Sinn
leugnet.
Im
und damit
besonderen
Ausdruck bezeichnet.
als
- die
wurde
Und zwar ge-
eine Verflüchtigung der Wirklich-
So rechnet Wolff die Idealisten samt den Materialisten und
zu den «drei schlimmen Sekten" (s. Wolff von seinen
keit
Skeptikern
Schriften S. 583); durchgängig war man bis gegen
^
Ende des 18. Jahr-
Näheres darüber s. Vaihinger in den „Straßburger Abhandlungen zur
In der Kunsttheorie scheint der Ausdruck noch weiter
Philosophie" S. 94 ff.
zurückzureichen.
Wenigstens ward mir von befreundeter Seite die mir hier
mitgeteilt, daß schon in Pacheco's arte de la
nicht kontrollierbare Notiz
Pintura (Sevilla 1649) Idealist zur Bezeichnung einer Kunstrichtung dient.
Idealismus
— Realismus.
hunderts ebenso eifrig darauf bedacht,
zu verwahren,
als
später,
sich
sich
69
gegen den Idealismus
zu ihm zu bekennen.^
stück zu Idealismus in diesem Sinne bedeutete
dem
Als Gegen-
18. Jahrhundert
Realismus die Behauptung einer außerhalb des Denkens befindlichen
Durch Herbart und
Welt. 2
seine Schule hat sich diese
Bedeutung
der Ausdrücke durch das 19. Jahrhundert hindurch bis zur Gegenwart erhalten.
Dann aber bewirkte, wie bei vielen Ausdrücken,
so auch hier
die Kantische Philosophie eine wesentliche Verschiebung. ^
noch der herkömmlichen
Terminologie
folgt
zunächst
daher
(z.
nunft)
den Idealismus mit dem Skeptizismus zusammen.
Kant selbst
und
stellt
B. in der Vorrede zur 2. Aufl. der Kritik der reinen Ver-
Die Präg-
ung des Ausdruckes transzendentaler (auch formaler oder kritischer)
Idealismus erfolgt im Hinblick nicht auf Plato, sondern auf Berkeley;
seinem
»empirischen",
er
einen
Dingen
jenseit
stellt
neuen
„psychologischen« Idealismus
„materiellen",
Idealismus
entgegen,
der
die
Existenz
von
der Vorstellung keineswegs leugnet oder bezweifelt,
der aber die Formen der Anschauung und des Denkens für bloßsubjektiv
erklärt;
damit werden
Gegenstände einer uns mög-
alle
lichen Erfahrung zu bloßen Erscheinungen,
danken keine an
„die außer unseren Ge-
Diese Verschieb-
sich gegründete Existenz haben."
ung enthält insofern einen fruchtbaren Keim zur Weiterbildung, als
den Träger der Formen,
das Subjekt der Erkenntnis,
dabei nicht
sowohl der Einzelmensch
in seiner Besonderheit als die
gemeinsame
Struktur unseres Wesens,
Indem
bildet.
sich
so
die
geistige
das Problem
Organisation des Menschen
aus
der Psychologie in die
Geisteslehre verlegte, konnten bald in weiterem Sinne Idealisten alle
welche die Überlegenheit der geistigen Tätigkeit gegen die
heißen,
Macht der Außenwelt verfechten.
So
schreibt
W. von Humboldt (Briefw. S. 485): „Am Ende
Idealisten
^
z.
Schiller
B.
sind
an
wir ja beide
und würden uns schämen, uns nachsagen zu
lassen,
daß
Wolff (de differentia nexus rerum sapientis et fatalis necessitatis S. 75)
will in keiner
Weise Plato zu den
Idealisten
gerechnet wissen;
wohl nenne
den
er die Körperwelt Erscheinung, aber er verstehe darunter keineswegs mit
Idealisten eine
bloße Vorstellung.
Im Mittelalter bildete bekanntlich Realismus den Gegensatz zu Nominalismus; seine Anhänger hießen gewöhnlich reales; realista erwähnt Prantl
(Geschichte der Logik IV, 221) zuerst bei Petrus Nigri (um 1475).
*
^
II,
Näheres darüber s. Trendelenburg, Logische Untersuchungen,
512 ff.
3. Aufl.
70
Zum Grundbegriff des Geisteslebens.
die
Dinge uns formten und nicht wir die Dinge".^
Zur Durch-
setzung dieser Bedeutung hat niemand kräftiger gewirkt als Fichte.
In deutlicher Verwandtschaft damit, aber zugleich in
eigentüm-
licher
Färbung, verwendet den Ausdruck der deutsche Neuhumanis-
mus,
diese
neueste
Phase
der
So
Renaissance.
mit der F. A. Wolf
dankenreiche Abhandlung,
erklärt
das
ge-
die
„Museum der
Altertumswissenschaft" (1807) eröffnet, als die „erste Bedingung aller
höheren Ausbildung"
Richtung des Geistes";
»ideale
die
ver-
er
gemäß seinem Lieblingsspruch, daß, „überall
das Nützliche zu suchen ganz und gar nicht für großgesinnte und
steht
aber
freie
Menschen paßt"
einer
harmonischen Entfaltung
nicht
irgendwelcher Folgen
darunter
Polit.
(Aristot.,
aller
wegen, die Richtung des Lebens
auf das Nützliche, sondern das Schöne.
mus
läßt
Realisten
die
nicht
Diese Fassung des Idealis-
durch Persönlichkeit und Lebenswerk niemand mehr ge-
hat
fördert als Goethe, mit so
einen
1338 b2), das Streben nach
Geisteskräfte allein ihrer selbst,
gutem Rechte er
in
anderer Hinsicht sich
Der Sprachgebrauch des
nannte.
philosophische und
die
19.
künstlerische Fassung
Jahrhunderts
in
eins zu-
sammenfließen; indem so der Idealismus ein Bekenntnis zur Selbsttätigkeit
und zum Selbstwert des Geisteslebens wird,
des erkenntnistheoretischen Schulproblems des
Stelle
1 8.
tritt
an die
Jahrhunderts
und bleibende Frage der Menschheit.
eine alte
b)
Zum Kampf der Lebensgestaltu,ngen.
Die Formulierung des Gegensatzes von Idealismus und Realismus
läßt verschiedene
Fassungen zu, die
hinauskommen.
selbe
Ist
in
der Sache aber alle auf das-
der Hauptstandort des Lebens die sichtbare
oder eine unsichtbare Welt, und sind die Hauptziele des Lebens hier
Besonders eingehend hat sich Schiller mit den Ausdrücken
in
der
Abhandhmg ,,Über naive und sentimentalische Dichtung" beschäftigt.
Als
^
ihm hier, wer sich durch die Notwendigkeit der Natur bestimmen
läßt, als Idealist, wer sich durch die Notwendigkeit der Vernunft bestimmt.
Die Schulgelehrten empfanden die Wandlung wohl und widerstrebten ihr.
So sagt Plattner (Phil. Aphorismen I, 412): „Man fängt itzt an, den Begriff
des Idealismus gar zu weit auszudehnen. Der zeither gewöhnlich gewesenen
Bestimmung nach ist es dasjenige System, welches das Dasein alles dessen
leugnet, was nicht Geist ist." — „So wie man itzt den Idealismus versteht,
Realist gilt
wären alle die, welche die Sinnenwelt als eine Erscheinung betrachten, mit
alle Philosophen ohne Ausnahme Idealister."
anderen Worten
— Realismus.
Idealismus
sich
nicht
ist
sondern nur
der Natur,
als eine
neue und höhere Stufe der Wirklichkeit verstehen
wesentlich
oder
Fortführung
eine
als
71
Entwickelt sich im Menschen ein Leben, das
oder dort zu suchen?
alle
läßt,
Betätigung nur eine Begleiterscheinung oder
geistige
Werkzeug eines seinem Kerne nach naturhaften Lebens? Hat
kein anderes Ziel als die Erhaltung und Pflege des
ein
der Mensch
menschlichen Kreises, wie er im unmittelbaren Dasein vorliegt, oder
gewinnt das Menschenleben einen Sinn und Wert nur durch das
Teilhaben an
Ordnung?
niedere
zu
bloßmenschlichen
allem
einer
Ist,
wo
gewöhnliche
die
und höhere Stufen
und
erklären
damit
Befinden
Ansicht
scheidet, das
die
Höhere vom Niedern her
ihm zurückzuziehen,
zu
Höhere den Schlüssel zum Verständnis des Niedern?
durch
der
alle
vom Größten
Gehalt
Fassungen
diese
bis
zum
und Wert
bis
hindurchscheint,
einzelnen Gebiete
alle
oder bildet das
Der Gegensatz,
muß das Leben
im Denken wie im Handeln,
Kleinsten,
in
überlegenen
Wirklichkeit in
in
hinein grundver-
Das wird auch für den Begriff
schieden, ja entgegengesetzt gestalten.
der Wirklichkeit selber gelten, und es wird sich der Idealist mit Fug
und
dagegen
Recht
dem
nach
sträuben,
Realismus gemessen zu werden.
des
Wirklichkeitsbegriff
Das aber geschieht, wenn die Welt
des Idealismus zu einer gegebenen und gesicherten Welt nur hinzu-
kommen scheint, wenn sie wie ein bloßer Zusatz und Schmuck behandelt wird; demgegenüber wird der Idealist darauf bestehen, daß
seine
Gedankenwelt
allererst
Wirklichkeit möglich mache,
den
Begriff
einer
Welt und
einer
und daß die sinnliche Welt einen Halt
wie einen Wert nur aus jener gewinne.
Dem Idealismus ergeht es hier oft ähnlich wie der Religion.
So
lange diese das Leben beherrschte, galt ihre Welt als die allernächste
und
unbestreitbare,
ein
Augustin
Idee eines höchsten Wesens aus,
überwand
alle
Zweifel von der
und ein Thomas von Aquino nannte
Erst nachdem
die überirdische Welt schlechtweg das Vaterland (patria).
die
Stellung
der Religion
erschüttert
und
ihr Inhalt verblaßt war,
konnte bei ihr der Gedanke der Jenseitigkeit und der Transzendenz
in
ist,
den Vordergrund
treten.
Wie sie in Wahrheit schon aufgegeben
wo sie in erster Linie als transzendent erscheint, so ist auch die
Sache des Idealismus schon verloren gegeben, wenn seine Welt
eine ferne
schließende
als
und fremde, durch mühsame Gedankengänge erst zu ergilt.
Den Gegensatz aber in solcher Schärfe denken, das heißt alle
Zum Grundbegriff des Geisteslebens.
72
Vermittlung zur Unmöglichkeit machen, auch den sog. Realidealismus,
falls
er eine solche bedeuten will.
Wohl mag, ja muß der Idealist
anzueignen streben, auf die sich der Realist stützt,
sich die Tatsachen
und dieser wird das in umgekehrter Richtung tun. Beide aber tun
es, indem sie den Tatbestand von ihrer Überzeugung aus neu beleuchten
ein
und umgestalten, das aber ist weniger ein Ausgleichen
als
Vertiefen des Gegensatzes.
Der Realismus des 19. Jahrhunderts.
a.
Zu einer Untersuchung des Problems drängt zwingend die Tatsache,
daß der Verlauf des
neue Phase
gebracht
hat.
Aufgaben vornehmlich
tat
es
die
neuere
in
überkommene
Kultur
hatte
19. Jahrhunderts
,
alten Streit in eine
der Richtung des Idealismus;
Lebensführung,
religiöse
dahin
bis
den
Bis dahin suchte das Kulturleben seine
die Lebensarbeit
vor allem
aber auch die
vornehmlich von
innen her geführt und den Forderungen des Denkens die äußeren
Verhältnisse zu unterwerfen gesucht. Es fehlte dabei nie an einer Gegen-
wirkung realistischer Art, aber sie bot weniger eine charakteristische
Gestaltung des Ganzen
als sie ein
zäher Widerstand der Individuen
war, die von den Freuden
und Leiden der sinnlichen Welt viel zu
wurden,
um sich zur geforderten Höhe des Lebens
stark festgehalten
aufschwingen und auf ihr halten zu können.
wirkung von
sie
Eine solche Gegen-
Mochte
bemessene Grenzen.
lauter Kleinkräften hatte
unablässig zur Abbröcklung und Herabdrückung wirken,
keines-
wegs war sie im stände, den Idealismus durch Entgegenhaltung eines
neuen Lebenssystems bis zum Grunde zu erschüttern.
es,
was der Realismus des
19. Jahrhunderts
Das aber ist
unternommen
hat;
die
seine Meinung, kann alle Ziele der Menschheit
nächste Welt, so
ist
aufnehmen und
alle
Wünsche
erfüllen,
ohne
sie
kömmliche Fassung des Idealismus herabzustimmen.
gegen
die
her-
Solches Unter-
nehmen ist mehr als eine andere Deutung, eine neue Zurechtlegung
des überkommenen Tatbestandes, es schöpft seine Kraft vornehmlich
aus der
Tatsache,
Menschheit
Realismus
er
das
daß
die
mehr geworden
Welt des unmittelbaren
ist
als je
zuvor.
Nur
Daseins der
weil
der neue
dem Idealismus eine neue Wirklichkeit entgegenhält, kann
Denken und Streben der Menschheit zu gewinnen
So stoßen
hier nicht
eine Bestätigung der
sowohl Lehren
als
Wirklichkeiten
hoffen.
zusammen,
Behauptung des vorigen Abschnittes, daß der
Idealismus
— Realismus.
73
Kampf der Geister nicht sowohl auf die Deutung
als
auf die
Ge-
staltung des Tatbestandes geht.
Zur Steigerung der nächsten Wirklichkeit verbinden sich im
mannigfachsten Bewegungen. Weit tiefer hat uns
19. Jahrhundert die
die Natur in ihr Gewebe einblicken lassen, weit
mehr beschäftigt sie
das Tun und beherrscht sie das Denken; den Gewinn des Wissens
aber verwandelt das technische Geschick alsbald in einen Gewinn
für das Leben und bringt diesem die erfreulichste Bereicherung, Beschleunigung und Kräftigung; ein staunenswertes Wachstum menschlichen Vermögens treibt mehr und mehr das starre Schicksal aus
der Welt und nimmt selbst den Widerständen ihre Bitterkeit, indem
Über-
es sie in einen Antrieb zur Tätigkeit, eine Aufforderung zur
windung
Zugleich eröffnet das menschliche Zusammen-
verwandelt.
sein größere
und größere Aufgaben.
uns, wie viel die Gestaltung des
Immer mehr überzeugen wir
gemeinsamen Lebens bedeutet, und
dem vorgefundenen Stande eine erhebliche
Steigerung möglich, eine Hebung der Wohlfahrt und ein allgemeineres
Glück erreichbar ist. Wie innerhalb der Staaten die einzelnen Kräfte
wie
gegenüber
hier
zu einer freieren und volleren Betätigung gelangen, so findet zugleich
und Unterscheidende der Völker eine bereitwillige
läßt Gesinnungen und
das Eigentümlich
Anerkennung, die Ausbildung nationaler Art
Auf wirtschaftlichem Gebiet trifft das Streben nach
Kräfte erstarken.
gleichmäßigerer Verteilung der Güter mit schweren Verwicklungen
der
aus
technischen Gestaltung der Arbeit
unermeßliche
Leidenschaften;
die
Macht
zusammen und erzeugt
der
materiellen
Lebens-
bedingungen gelangt nun zuerst zu deutlicher Anschauung und voller
Würdigung;
scheint
an
steigert sich
auch
die
innere
Lage
Probleme zu
diesem
wie
hängen.
das
Glück
Das
alles
des
Lebens
ergänzt
gegenseitig, die Erfolge wie die Probleme dieses
und
neuen
Lebens schmieden den Menschen immer fester an die unmittelbare Welt.
In solchen
Leistungen wächst auch der Träger der Arbeit, die
Menschheit, und zwar die Menschheit wie
wie eine Gedankenwelt
in
neuem
Bilde
verklärte.
sie
sie
leibt
und
lebt,
nicht
Geschichte und Gesellschaft
wirken dahin zusammen.
Im Nebeneinander wie
im Nacheinander rücken die Kräfte einander näher, verbinden sich
zu gemeinsamem Werk und gewinnen das Bewußtsein einer durch-
gängigen
heit,
wie
Solidarität.
So steht vor uns in großen Zügen die Mensch-
sie sonst zerstreute
Zusammenhängen
einfügt,
Kräfte verbindet, den Einzelnen festen
das
Vermögen des Ganzen unermeßlich
Zum Grundbegriff des Geisteslebens.
74
So kann
steigert.
zum Gegenstand
sie
der
Verehrung und
des
Glaubens werden, so kann sie alle praktische und ethische Betätigung
des Menschen an sich zu ziehen suchen.
muß alle einzelnen Gebiete,
Diese neue Denkweise
und Wissenschaft, eigentümlich
Kunst
aller Tätigkeit
gestalten,
Berührung mit
den Dingen
Ablösung von ihnen,
Gewebe,
alles
wie
scheint
die
und
idealistischen
Nur die
zu führen, während
ein Sicheinspinnen der Seele in ihr eigenes
Streben schattenhaft, matt und unwahr macht.
treibt;
solchen
in
B.
aber
menschlichen Kräfte von
Wandlungen scheinen
Lebensgestaltungen
So ist
Bewegung
es das Verlangen nach echter Wirklichkeit, das hier alle
trägt
z.
gebietet
den engsten Anschluß an die Welt um uns.
blasser Möglichkeit zu lebendiger Wirklichkeit
die
sie
älteren,
alle
alle
wie Nebelgebilde der siegreichen
Klarheit eines neuen Tages zu weichen.
ß.
Ist
Die Schranken des neuen Realismus.
das eigene Schicksal
nicht
alle Schatten und die Wendung
Daß die Sache nicht ganz einfach liegt, zeigt
der realistischen Lebenswelle. Gewiß hat sie
das Licht dieses Tages ohne
ohne allen Zweifel?
nur die menschliche Meinung überwältigend fortgerissen,
sie
hat auch die Arbeit ein gewaltiges Stück gefördert, sie hat in unser
Dasein
einen
rascheren
Fluß,
ein
mannhafteres Ringen
mit
den
Widerständen, mehr siegreiches Vordringen gegen die Unvernunft
gebracht.
erzeugt,
und
Aber zugleich hat das Anschwellen der Bewegung Probleme
die
den vom Realismus abgesteckten Kreis überschreiten
die Selbständigkeit
dieses
Kreises gefährden.
Zur
ausschließ-
lichen Wirklichkeit des
Menschen konnte das realistische System nur
werden, wenn sich
Verwicklungen durch den eignen Fortgang
alle
wenn alle selbständige Innerlichkeit mehr
mehr verschwunden, und der Mensch ganz und gar in ein
Werkzeug der Arbeit verwandelt wäre. Statt dessen hat jener Fort-
der Arbeit gelöst hätten,
und
gang deutlich
gezeigt,
Arbeit aufgeht.
daß der
Mensch keineswegs
in
die
Zunächst hat er die Arbeit immer mehr
in
bloße
einen
harten Kampf ums Dasein verwandelt, in einen Kampf der Individuen,
Stände und Völker; die Gegensätze sind immer schroffer, die Schlachtlinien
immer breiter geworden.
verraten
deutlich
genug,
Die Leidenschaften dieses Kampfes
daß hinter der Arbeit empfindende und
glücksdurstige Wesen stehen, die von der Arbeit etwas begehren und
verlangen, sei es selbst auf Kosten der Arbeit.
Läßt sich den daraus
Idealismus
— Realismus.
75
erwachsenden Gefahren begegnen, ohne die Gesinnung anzurufen,
d. h.
eine
Größe, für die
strenger Realismus keinen Platz hat?
ein
Es reichen aber die Verwicklungen über den Zusammenstoß der
dem
Wesen der
immer nur
einen Teil der menschlichen Kräfte, sie entwickelt einen immer
kleineren Teil, je feiner und verzweigter sie wird; immer geringer
wird der Bruchteil des Ganzen, den das Individuum zu umspannen
arbeitenden
vermag.
Leben
wirkliche
ist
folgerichtig
Menschen an
nicht
in
nimmt
aber
sie
jenen
Umgebung
zuerkennen
Verzicht
als einen Verlust
mehr in ihm,
als
darf.
da ihm
der
besteht;
nicht
gleichgültig
und Schmerz. Also
der Realismus ihm zuerkennt
Weiter
die Arbeit
bindet
den
die Leistung, ihr gilt alle Kraft als verloren, die sich
Leistung
nach außen,
kann
eigenen
entwickelt
gleichgültig sein,
Berührung mit der
der
in
Mensch
augenscheinlich
und
Die Arbeit
müßte dem Realismus
sondern er empfindet ihn
hin,
scheinen
sie
verbunden.
Solches Liegenlassen von Kräften, solcher Verzicht auf den
ganzen Menschen
alles
hinaus,
Kräfte
untrennbar
Arbeit
umsetzt.
macht
einen
solchen Stand
Streben nach Leistung,
Damit
aber
richtet
sie
Sinnen
alles
gleichgültig gegen den Stand der Seele,
sie
Erfolg
den Menschen aufsaugen und
überhaupt
nicht gelten lassen.
Das
und Anerkennung muß immer mehr
alles
selbständige Seelenleben unter-
Aber freuen
drücken, es hat es in Wirklichkeit sehr zurückgedrängt.
können wir uns solcher Zurückdrängung
wir eine peinliche Leere;
ja
nicht,
vielmehr empfinden
wo aber eine solche Empfindung erwacht,
da verfliegt alsbald die Befriedigung an der Arbeit, und
uns mit allen ihren Erfolgen
in eine seelische Ferne.
sie
rückt
Beim Ganzen
der Menschheit entspricht jener völligen Verwandlung des Daseins
in
Arbeit
ein
Verblassen eines geistigen Lebensgehaltes;
mehr gemeinsame Ideen und Überzeugungen
lich
wo nicht
die Menschheit inner-
zusammenhalten, da entschwindet immer mehr eine gemeinsame
Gedankenwelt.
Darauf aber scheinen wir nicht so
leicht
und nicht
ohne schweren Schaden verzichten zu können, denn damit hängt
alles
zusammen, was unserem Leben einen
und eine Seele
gibt.
Das sind
nicht
bloße Erwägungen
Selbstwert,
eine
Größe
grüblerischer Reflexion, es
sind unbestreitbare Erlebnisse und Erfahrungen der modernen Menschheit
Oder kann jemand leugnen, daß
alle
glänzenden Triumphe
der Arbeit ein Aufkommen und Vordringen einer tiefen Unzufriedenheit,
einer pessimistischen
Lebensstimmung
nicht verhindert
haben?
Zum Grundbegriff des Geisteslebens.
76
Das
19.
Jahrhundert
hat
den Anblick der Welt und die Lage des
Menschen so gefördert, wie kein anderes Jahrhundert;
so ließ sein
Schluß ein stolzes und freudiges Kraftgefühl erwarten.
Wenn der
wirkliche Anblick der Dinge ein völlig anderer ist, so enthält gewiß
die
Rechnung einen
die
realistische
Lebensbewegung
die
Seele
nicht eliminieren läßt;
sich
Fehler.
die
Seele eliminieren wollte,
und
der Versuch der Verneinung
wieder stärker hervorgetrieben.
selbst hat die Seele
überkommenen Formen des Idealismus.
Kritik der
y.
Dieser aber dürfte darin bestehen, daß
Solche Erfahrung zwingt, die ganze Frage zu
revidieren
und
Recht und Unrecht bei den streitenden Parteien möglichst zu scheiden.
Das Verlangen nach
Lebens
voller Wirklichkeit des
soviel
Macht
erlangt,
kundet,
wenn
nicht die
überkommenen
idealistischen
eine solche Wirklichkeit hätten vermissen lassen.
in
Wahrheit:
jene Arten
eine religiöse,
be-
sie
Lebensformen
So aber stand
des Idealismus wurzelten
im eignen Wesen des Menschen.
nehmlich zwei:
hätte schwerlich
wie das Vordringen des Realismus
nicht
es
mehr fest
Es waren aber dieser Arten vor-
die zu uns
vom Christentum durch
seine verschiedenen Gestaltungen wirkt,
und
eine künstlerische,
vom
oft
unterdrückten, so doch
Griechentum her einen wenn auch
die
nie ganz versiegten Lebensstrom bildet.
Die religiöse Lebensgestaltung mit ihrer Begründung des menschlichen Daseins auf eine überweltliche Ordnung, ihrer Erhebung
von
der Zeit zur Ewigkeit, von allem Außenleben zu einer reinen Innerlichkeit
behauptet
Macht; auch
trotz
aller
Schwächung immer noch eine große
wo sie abgelehnt wird, wirkt sie im Verborgenen fort.
Nähe aber und ihre sichere Überzeugungskraft hat
Menschen verloren.
Sie hat sie verloren
schon deshalb, weil zwischen der überkommenen Gestalt der Religion
und der modernen Gedankenwelt eine tiefe Kluft entstand; selbst
wer diese Kluft überbrücken zu können hofft, der hat nicht mehr
Ihre
sie
seelische
für
den
modernen
die Unmittelbarkeit
und
die
volle
Gewißheit des alten Glaubens.
Wenn
aber die Religion nicht das Allergewisseste
leicht
zum
ist,
so wird sie
Allerungewissesten.
Mehr noch
hat
die
Religion dadurch an Macht verloren, daß
dem Menschen der Neuzeit nicht mehr in derselben Weise aus
eigenen Erfahrungen hervorquillt wie dem Christen der alten Zeit.
Dieser Zeit entsprang die Wendung zur Religion aus stärkster
sie
Idealismus
— Realismus.
Empfindung menschlicher Ohnmacht, aus
und
windlicher Schranken
nur
Lebens;
ort des
eine Rettung
ließ
und sicherste Welt, der schlechthin feste Standals
ein
Abglanz oder ein Symbol jener Welt
Die Neuzeit hingegen ver-
behielt das nächste Dasein einen Wert.
Aufkommen wie ihre Eigentümlichkeit einem jugendlichen
dankt ihr
einem starken Lebenstriebe des Menschen; von da aus
Kraftgefühl,
verwandelt sich ihm die Welt in eine unermeßliche Aufgabe,
Bearbeitung
fallen
nur die
des geistigen Selbst er-
wurde daher tieferen Gemütern, wie einem
warten, diese Überwelt
Augustin, die nächste
So
Widersprüche.
starrer
Wendung zu einer Überwelt
77
einer Erfahrung unüber-
er
Schranken
eigene Entwicklung
scheint
deren
innerlich wächst;
hier
endgültigen Verzichte,
ihre
und auch
weitet
sich
selbst
starren
alle
in
und
alle
Welt zu höchster Vollendung
die
hier
Vielleicht liegt die Sache nicht
zu führen.
Anhänger moderner Denkart
meint,
es
ganz so einfach, wie der
vielleicht
entfaltung selbst schließlich unsere Grenzen,
zur ^Empfindung bringen.
der Stärke, und es
wird die Kraft-
unser Unvermögen
ja
Aber einstweilen herrscht das Bewußtsein
fehlt zugleich
ein eigener, ein unmittelbarer, ein
Damit aber verliert sie
überwältigender Antrieb zur Religion.
ihre
zwingende Kraft und sichere Wahrheit.
Größer noch ist die Gefahr eines Unwahrwerdens beim künstlerischen
Idealismus.
legenen
Standort
Er suchte die Welt nicht von einem übersondern
her,
durch
ein
ihr
selbst
angehöriges
Wirken zu vollenden: die Gestaltung, die in Begegnung von Innerem
und Äußerem, von
gebung
alle
Seele
und Welt erfolgt, schien mit ihrer Form-
Mannigfaltigkeit
des
Lebens zusammenzufügen,
einzelnen Glieder gegeneinander abzugrenzen
Ebenmaß zu verbinden.
das Geistige
aber
aus
Alle bloße Naturkraft wurde damit veredelt,
dem dunklen
Wirklichkeit
tagesheller
Schacht
Mit
gefördert.
künstlerische
Lebensform
erzeugt,
hat das menschliche Dasein
sie
der Seele
verfeinert,
ihre
und zu harmonischem
sie
ein
hat
ebenso
sich
Durchbildung des Lebens gezeigt.
solcher
tätiges
als
der Möglichkeit
Leistung
hat
zu
die
wie vornehmes Leben
gehoben und das Gewebe
unentbehrlich zur geistigen
Aber ist sie stark und gehaltvoll
genug,
um es ganz ausfüllen zu können?
Gehört nicht eine be-
sondere
Naturbegabung,
Vermögen dazu,
hier
schöpferisches
um
den Schwerpunkt des Lebens zu finden, erwächst daraus nicht
ein Aristokratismus,
sich
ein
auch
solcher
der nicht nur die anderen ausschließt, sondern
Ausschließung
freut?
Muß
ferner
nicht
ein
Zum Grundbegriff des Geisteslebens.
78
Mensch, ein Volk, eine Zeit schon
um
im Gestalten Großes zu
müssen
nicht
Gestaltung
fiihren
zu
jenes
künstlerische
leicht
zu
endlich
und Großes zu
Wo solche Tiefe fehlt, da bleibt
können?
Leben
erreichen,
um sie zur
eine Tiefe der Seele irgend besitzen,
sie
an die Oberfläche gebannt,
da sinkt es
zu
Und wenn
Tändelei,
einer
der Fülle des Lebens stehen,
in
erfahren
leerem
Scheine
herab.
schweren Verwicklungen und harten Widersprüche,
die
unheimlichen Abgründe
menschlichen Daseins
des
ja
anerkannt
voll
— und dahin drängt eben die Erfahrung des Jahrhunderts, — kann da wohl die Kunst den Anspruch behaupten,
werden,
19.
von
sich
Leid
in
sie
leicht
aus
Schwere zu heben,
alle
Kann
Freude zu verwandeln?
dahin neigen,
jene
alles
sie
des
Licht,
alles
aber nicht, so
mag
in
Unvernunft abzuschwächen und das
Das aber weckt bald den
Dasein möglichst ins Schöne zu malen.
Widerspruch
Trübe
es
und
Wahrheitssinnes,
dessen
als
Vertreter
darf
sich der Realismus fühlen.
Noch
augenscheinlicher
ist
sein
Recht gegen den landläufigen
inmitten aller Erschütterung und Zerreibung der
Formen des Idealismus das Allgemeine der Richtung
ohne es irgend näher zu bestimmen und zu begründen.
der
Idealismus,
besonderen
festhält,
Ein
solcher
wissen,
„Schöne",
geben.
Idealismus
ohne
über
ihren
Vollauf verständlich
idealistischen
Lebensformen
nicht genügen;
friedigt,
schwärmt
was dieses »Höhere«
ob
wie er ihn
freilich
für
„Höheres",
sei,^
irgendwelche
demnach, daß
dem
neu
das
die
erwachten
der Realismus
eifrig vertritt,
ohne zu
er preist das „Gute", „Wahre",
Inhalt
ist
etwas
ist
ihn
Rechenschaft
zu
überkommenen
Wahrheitsdrange
ebenso
völlig
be-
eine andere Frage.
*
„Höher" ist als Lieblingsausdruck für eine neue, vermeintlich vornehmere Denkweise wohl namentlich in der Sturm- und Drangzeit der
deutschen Literatur aufgekommen. Dann suchte mit Vorliebe die Romantik
dadurch die eigenen Ziele und Begriffe von denen des Durchschnitts zu
scheiden, so verwenden auch Schleiermachers Jugendschriften das Wort sehr
oft.
Man spricht von „höherem" Leben, „höheren" Gefühlen, „höherer"
Bildung, „höherer" Sittlichkeit u. s. w., bis der Ausdruck endlich dem Spott
verfiel („höherer Blödsinn").
Der soliden und klaren Denkweise Kants widerstrebte der Ausdruck gründlich.
Als Feder ihm einen „höheren" Idealismus
zugeschrieben hatte, bemerkt er dagegen (IV, 121 Hart.): „Bei Leibe nicht
der höhere.
Hohe Türme und die ihnen ähnlichen metaphysisch -großen
Männer, um welche beide gemeiniglich viel Wind ist, sind nicht für mich.
Mein Platz ist das fruchtbare Bathos der Erfahrung."
Idealismus
79
Erörterung des Wirklichkeitsproblemes.
5.
des Lebens scheint
Eine Wirklichkeit
durch
reichbar
— Realismus.
ob
er-
Tuns mit der
des
Verkettung
fortwährende
eine
sichtbaren Umgebung;
dem Realismus nur
Wahrheit durch eine solche Verkettung
in
Wirklichkeit für den Menschen entsteht, das läßt sich sehr bezweifeln.
Denn nur um
kann
keit
des Tuns
mit
in
keiner Weise
Umgebung
der
Hülle und Fülle, nicht aber ergibt
lebnis
doch erlebbare Wirklich-
erlebte oder
handeln, alle andere Wirklichkeit liegt außerhalb
und kann ihn
seines Kreises
kettung
ihm
eine von
sich
es
wird die Leistung
sie
indem
erst,
kümmern.
ergibt
Jene Ver-
nun Leistungen
in
zum
Er-
damit Erlebnisse;
auf eine Einheit zurück-
sie
des Seelenlebens umspannt wird.
bezogen und von einem Ganzen
Ein solches Seelenleben aber kann der Realismus mit seinen Mitteln
unmöglich erklären,
für
sich
und doch bedarf
Denn
selbst.
er seiner aufs
dringendste
ihm eigentümliche Welt
entwickelt die
er
eigenem Vermögen,
er
würde,
eigenen Mittel angewiesen, wie
alle
inneren Zusammenhänge, so alle
nicht
aus
und
Lebenssysteme
So
zugleich
sich
bildet seine stillschweigende
selbst
auf
ausschließlich
als
ein
Ganzes
seine
zerstören.
Voraussetzung ein Seelenleben, das
und auch den Gegensatz von Subjekt und Objekt
die Mannigfaltigkeit
Grund solcher Voraussetzung läßt sich dartun,
daß die Weltumgebung für den Menschen weit mehr bedeutet, daß
er aus ihr weit mehr zu gewinnen vermag, als der DurchschnittsDann aber wird in Wahrheit der Realismus
idealismus zugestand.
umfaßt.
von
auf
Erst
einer' Gedankenwelt des Idealismus
Lebenssystem
Wird aber
er
ist
zugleich
umsäumt,
ja
umspannt,
als
überhaupt nicht möglich ohne den Idealismus.
der Seele jede Selbständigkeit versagt und
sie
möglichst von draußen abgeleitet, so entsteht ein schreiender Wider-
spruch;
mag
er
sich
leidlich
verstecken
schleichungen in den Begriffen und
Betrachten
Realismus.
Es
lassen,
unablässige
Er-
Lehren verraten ihn bald.
wir das System Comtes, des größten Denkers des
ist
Grundlegung
bei
der
alles
zu entfernen, was irgend
aus den
Begriffen
stammt.
Aber sobald
es
eifrigst
darauf
vom
bedacht,
Idealismus
vom Entwurf zur Ausführung, von
der
Kritik zum Aufbau vordringt, verschiebt sich der Anblick der Sache.
Je
mehr nämlich
jenes
geschieht,
desto
mehr sehen wir
jene
an-
und einer idealistischen Fassung
nähern, namentlich wird nur mit Hülfe einer solchen Verwandlung
fänglichen
Größen
sich verändern
Zum Grundbegriff des Geisteslebens.
80
Überganges vom Erkennen zum Handeln
der kritische Punkt des
und der physische Zwang
überwunden
ein
in
moralisches Sollen
So ergibt sich schließlich ein Lebensganzes, aber es ergibt
verwandelt.
sich
nur mit ständiger Hülfe desselben Gegners, an dessen Vernichtung
die
Wahrheit des Lebens zu liegen schien.
nun wohl eine derartige zwiespältige Welt die Bedürfnisse
Sollte
im besonderen das ethische Verlangen befriedigen
des Geisteslebens,
Wiederum mag Comte herangezogen sein, wiederum voll-
können?
Umschlag, nur tut er es hier in umgekehrter Richtung.
zieht er einen
Der Ausgangspunkt nämlich
der Abschluß
idealistischer,
ist
realist-
ischer Art. Die tiefangelegte Natur des Mannes empfindet die Schäden
der Zeit durchaus im Sinne des Idealismus, er nimmt
sie
so schwer,
daß ohne ein Aufbieten ursprünglichen Schaffens, ohne eine MögErneuerung
durchgängiger
lichkeit
Gegenwirkung
alle
verloren
Was er aber vom Realismus her als Heilmittel bringt,
scheint.
genug,
dürftig
es
Aufklärungen über die Natur,
sind
ist
sowie Ver-
der gesellschaftlichen Organisation, von denen jene
änderungen
in
Umwälzung,
jener Sieg des
Das kann
Guten erwartet wird.
nicht
geschehen ohne einen krassen Optimismus gegenüber dem Menschen,
sonst
würde der Widerspruch zwischen der Größe der Aufgabe und
Das aber ist
der Kleinheit der Mittel allzu handgreiflich werden.
typisch
für
den Realismus: entweder
er
nimmt das Lebensproblem
sehr flach, oder er verwickelt sich in Widersprüche, die, konsequent
durchdacht,
ihn
zerstören.
selbst
Befriedigt
nun wohl ein Lebens-
system das Verlangen nach Wahrheit und den Durst nach Wirklichkeit,
das
um so widerspruchsvoller wird, je mehr es dem Gesamt-
befunde des menschlichen Lebens gerecht werden will?
Dieser
begrifflichen
war.
ihrer
Atmosphäre,
Denn
mögen
hat
in
vom
welche
gänzlich
dessen
Gestaltungen
Behauptung noch so
der Kultur
entspricht
die
Erfahrung der
Die Bewegung zum Realismus erfolgte innerhalb einer
Menschheit
geistigen
Erörterung
erschüttert
sein,
Idealismus
in
dem
gesättigt
Besonderen
die Gesamtentwicklung
jahrtausendlanger Arbeit ein Allgemeineres der
Denkweise, der Empfindung und Schätzung von jener Besonderheit
abgelöst
und
es
tief
in
das Ganze des Lebens, auch in das Innere
der Seele einsickern lassen; das umfängt auch die realistische Lebens-
ordnung
und
dient
ihr
zu
unablässiger
Ergänzung,
Milderung,
Berichtigung.
Je mehr sich aber der Realismus zur Selbständigkeit
erhebt und seiner Eigenart bewußt wird, desto gründlicher muß er
— Realismus.
Idealismus
jene
idealistischen
Elemente vertreiben.
81
Aber zugleich
wird eine innere Niederlage.
vollzieht
er
dem äußeren Sieg
eine Selbstverengung und Selbstzerstörung, aus
Bei solcher weltgeschichtlicher Dialektik
wäre der Verlauf der Sache mit voller Ruhe zu betrachten, und es
könnte das gewaltige Wechselspiel der Gedankenmassen reine Freude
um ein Schauspiel handelte. Aber es
handelt sich um das Geschick des Menschen, um Vernunft oder
Unvernunft seines Daseins, um den Gewinn oder Verlust einer Seele.
wenn
bereiten,
es
sich
nur
Und das läßt sich nicht so ruhig betrachten.
Die Notwendigkeit eines neuen Idealismus.
s.
So wenig der Realismus mit
digen kann,
viel
zu
viel
seiner Daseinskultur
daß eine einfache Rückkehr zum
als
uns
befrie-
im Bestände des Lebens verändert,
hat sich
alten Idealismus
möglich wäre.
draußen und drinnen weit mehr Unvernunft zutage
Nicht bloß
ist
getreten als
jenem Idealismus vor Augen
stand,
auch die weite Aus-
delyiung starrer Tatsächlichkeit und blinder Gleichgültigkeit des Weltlaufs
gegen die Zwecke des Geisteslebens hat für uns eine
viel
zu
Nähe und Gegenwart, als daß wir darüber so rasch
hinwegkommen könnten als jener. Sind aber die Verwicklungen
und die Widerstände so sehr gewachsen, so muß sich auch die
Gegenwirkung verstärken, so muß sich der Idealismus zu größerer
eindringliche
Tiefe
zurückverlegen und eine festere Grundlage suchen.
nur
das
aber
ihm
nicht die
faltung
des
können,
wenn zur Anerkennung
Er wird
gelangt,
daß bei
Aufbringung besonderer Leistungen, nicht eine Ent-
Lebens nach besonderen Richtungen,
sondern
die
Er-
reichung eines wesen- und wahrhaften Lebens schlechthin in Frage
und damit überhaupt
kein
echtes
Leben ohne eine das Dasein begründende Tiefe und
ihre
Aneig-
steht,
daß
kein
es
Selbstleben
nung gibt Zugleich aber muß eine schärfere Abhebung des Geisteslebens vom Stande des bloßen Menschen erfolgen.
Der Welt der
Natur und des sichtbaren Daseins, die uns mit überlegener Macht
umfängt, kann die geistige Betätigung gewachsen nur werden, wenn
sie
eine neue Stufe der Wirklichkeit,
Gesamtleben der Geistes-
ein
welt vertritt
und aus seinen Kräften schöpfen darf.
Sonst fehlt dem
Idealismus
ein
und
nur wenn eine
Halt
fester
ein
gutes
bei sich selbst befindliche Geisteswelt in
Recht;
uns wirkt und uns zu
er-
füllen vermag, wird die
dem Idealismus wesentliche Forderung ver-
und
daß die Größen und Güter der neuen Welt
ständlich
erfüllbar,
Eucken, Grundbegriffe.
4.
Aufl.
140095
6
Zum Grundbegriff des Geisteslebens.
82
Zwecken der Menschen unvergleichlich überlegen und
als allen
als
unabhängig von allem menschlichen Mögen und Meinen, von allem
menschlichen Tun und Treiben behandelt werden; nur dann wird es
möglich, daß nicht
daß
hat.
Die Menschheit
nach
sich
Wahrheit
ihre
sie
dern
vom Menschen erhalten, son-
bemißt, wie
ihnen
zum Maß
des
viel
das heißt diese innerlich zerstören; wie aber
heit
Wahrheit sein Leben
Wahren und Guten machen,
über die Mensch-
ist
hinauszukommen, wenn das unmittelbare Dasein
Wirklichkeit giltpi
Durch
Sache wird immer wieder die eine Frage
Entscheidung fordern, ob
Dasein zu dienen
hat,
oder ob
in
der
in
dem mensch-
einem gegebenen
Wendung zum Geistesleben
und zugleich ein Reich von echten Gütern
eine neue Art Wirklichkeit
Gewinnt das Geistesleben
hervorbricht.
die ganze
hervorbrechen und eine
unser Streben lediglich
all
Wohlsein, der menschlichen Wohlfahrt
lichen
als
Verdunklung und Abschwächung der
alle
nicht
eine
Überlegenheit
gegen das menschliche Getriebe, so muß aller Idealismus fallen, aber
mit ihm
fällt
läuft in
völlige
auch
menschlichen
gefährden,
sicher
und weit überlegen.
lebens
in
Ideen
gelangen,
heit
es ver-
so kann alle
Unzu-
jene Überlegenheit anerkannt,
des
Weise
die
und Wert unseres Lebens, und
Leere.
Wird aber
länglichkeit
aller Sinn
so
bleibt
Standes
die
das
Geistesleben
Grundtatsache
aller
in
keiner
Verwicklung
Mag sich an alle Entfaltung des Geistes-
unserem Bereich Kleinmenschliches anhaften und mögen
nur mit Hilfe der Interessen zur Wirkung
gewöhnlich
mag ferner das Geistesleben auf dem Boden der Mensch-
von verschwindenden Anfängen her geworden und
sich
sehr
langsam, auch unter mannigfachen Rückfällen, fortbewegt haben, das
alles gefährdet
nach jener Scheidurig nicht im mindesten die Grund-
des Geisteslebens;
tatsache
unwillige
Anerkennung der
ja
der Widerstand der Menschen,
die
auch
der
geistigen
Notwendigkeiten,
der Geistigkeit,
womit das menschliche Tun und Treiben
zu
umkleiden
das alles kann nur die Überzeugung ver-
stärken,
daß mehr
Schein
sich
liebt,
in
der Menschheit vorgeht, als aus dem Menschen
des unmittelbaren Daseins stammt.
Wenn daher heute auf der Höhe der geistigen Arbeit die Wage
Es sei dabei das Wort Kants gegenwärtig (III, 260 Hart.): »In Ansehung der sittlichen Gesetze ist Erfahrung (leider!) die Mutter des Scheines,
'-
und es ist höchst verwerflich, die Gesetze über das, was ich tun soll, von demjenigen herzunehmen oder dadurch einschränken zu wollen, was getan wird."
Idealismus
wieder mehr sich
— Realismus.
83
zum Idealismus neigt, so ist nur zu wünschen, daß
solche
Bewegung
bleibe,
sondern daß das Entweder - oder mit voller Schärfe hervor-
nicht
abschwächenden Vermittlungen stecken
in
gekehrt und die unerläßliche Umkehrung mit vollem Nachdruck gefordert
werde.
muß
auch
auch
positiver
Der Idealismus darf
vordringender,
Art
sein.
er
darf
nicht
nicht
Denn nur
so
bloß
abwehrender,
kann
es
er
er
muß
gelingen,
der
bloß kritischer,
wachsenden Veräußerlichung, Verflachung und Scheinhaftigkeit einer
bloßen
Menschenkultur
eine
echte
Geisteskultur
entgegenzusetzen
und der Überwältigung und Unterdrückung siegreichen Widerstand
zu leisten, womit jetzt Natur, Geschichte und Gesellschaft den Menschen
bedrohen.
Ohne einen Glauben an die Größe und den Wert der
Menschheit
feste
kommen wir nicht weiter, ein solcher Glaube aber muß
Grundlagen haben.
6*
Zum Erkenntnisproblem.
B.
Denken und Erfahrung.
1.
(Metaphysik.)
Geschichtliches.
a)
I—« inige Notizen zur Terminologie seien vorangeschickt. Der Aus*~^ druck Erfahrung ist im Lauf der Zeit immer vieldeutiger geworden und zeigt auch bei den einzelnen Denkern so viele Schwank-
daß
ungen,
er
kaum
einen
Terminus
festen
Abgrenzung der
eine sprachliche
vorwissenschaftlichen
Erfahrung von der wissenschaftlichen Erfahrung
hervorgegangen.
— Schon
Erfahrung
schaftlichen
Nicht einmal
bildet.
alltäglichen,
ist
aus
aller
Arbeit
die Stoiker bildeten den Begriff der wissen-
(ejATrsipia
Die
jjtsö-oSixTj).
Erfahrungsphilo-
sophen der Neuzeit waren geneigt, durch Herabdrückung der griechischen Ausdrücke Empirie,
empirisch, Empiriker zur Bezeichnung
der niederen Stufe eine Scheidung herbeizuführen, auch die deutsche
Schulphilosophie des 18. Jahrhunderts suchte eine »empirische oder
gemeine" und eine »gelehrte" Erfahrung auseinanderzuhalten.
Kant verwendet
fahrungsphilosoph
energisch
ist
empirisch in jenem Sinne,
oft
gegen
des
den
19.
Comte,
Jahrhunderts,
„empirisme".
Zu
Auch
und der größte Erverwahrt
diese Unterscheidung nicht gelangt; lediglich die Sonderung
»Empiriker«
für
niedere und
die
die aus der kantischen Philosophie
gedrungen
von
„Empirist« für die höhere Stufe,
stammen
dürfte,
kann
als durch-
gelten.
Bedeutender
a priori
sich
allgemeiner Geltung aber
ist
die Geschichte der hierher gehörigen Ausdrücke
und a posteriori;
ihre Wandlungen spiegeln die Hauptphasen
des Erkenntnisstrebens und erstrecken Wirkungen bis in die Gegenwart.
— Entsprungen
fahren,
sind die Ausdrücke
dem aristotelischen Ver-
das Allgemeine als das (begrifflich)
Frühere,
das Einzelne
Denken und Erfahrung.
als
einen festen Sprachgebrauch bildete
das Spätere zu bezeichnen;
Höhe des Mittelalters. Ex prioribus beweisen,
einem Albert dem Großen von den Gründen her, ex
daraus aber
das
85
heißt
erst die
posterioribus von den Folgen her beweisen; a priori und a posteriori
in
Bedeutung erwähnt Prantl
gleicher
Abendlande
IV,
78) zuerst bei Albert von Sachsen, einem Gelehrten
Diese
des 14. Jahrhunderts.
bis
die Neuzeit,^
in
dem Ende
im
(Geschichte der Logik
sie
ist
Bedeutung behielten die Ausdrücke
auch heute noch nicht erloschen.
Mit
gemäß der stärkeren
des 17. Jahrhunderts aber beginnt,
Hervorkehrung der Frage nach dem Ursprünge der Erkenntnis, eine
Verschiebung von der Methodenlehre zur Erkenntnislehre.
A priori
So vor-
heißt nun,
was aus der Vernunft,
a posteriori, was aus der Erfahrung stammt.
Diese Unterscheidung
nehmlich bei Leibniz.
aber
sich
ließ
relativ
und
flacher
absolut,
und
tiefer
verstehen.
A priori erkennen bedeutete zunächst nicht mehr als ein Erkennen
aus
gewonnenen Einsichten vor näherer Befassung mit der
schon
besonderen Sache, also durch bloße Schlußfolgerung; ^ woher das
Erkennen
letzthin
Aber schon
stamme, blieb dabei unerörtert.
bei
Leibniz selbst und dann bei seinen Nachfolgern bezeichnet a priori
auch, was von aller Erfahrung unabhängig ist und lediglich der VerBei
Kant
erreicht
Erfahrung
selbst
ihm
Bewegung
nunft
angehört.^
indem
die
griffen
und Sätzen a priori möglich zu werden
er gebraucht nicht selten
diese
Höhe,
ihre
durch ein Gefüge von Be-
erst
den Ausdruck
in
scheint.
Aber auch
dem laxeren Sinne.
Um
diese Zeit dringen die Wörter über die Schule hinaus in den allge-
meinen Sprachgebrauch, zugleich gewinnt a priori eine feste deutsche
^
penser)
So heißt es z. B.
:
seit
a priori,
in der
sogenannten Logik von Port Royal
en prouvant les effets par
seit
les causes, ce
en demontrant au contraire
les
(l'art
de
qui s'appelle demontrer
causes par les effets,
ce qui
s'appelle prouver a posteriori.
'
quod experiundo
quod vero ratiocinando nobis
§435: quicquid ex iis coUigimus,
So sagt z. B. Wolff (psychologia empirica § 434)
:
addiscimus, a posteriori cognoscere dicimur:
innotescit,
a priori cognoscere dicimur.
quae nobis jam innotuere, cum ante ignotum esset,
innotescit, adeoque idem a priori cognoscimus.
'
Leibniz
stellt
id
ratiocinando
nobis
der philosophie experimentale qui procede a posteriori
la pure raison ou a priori entgegen (s, 778 b. Erdm.).
Lambert sagt im „Neuen Organon" §639: „Wir wollen es demnach gelten
lassen, daß man absolute und im strengsten Verstände nur das a priori
die Erkenntnis durch
heißen könne, wobei wir der Erfahrung nichts zu danken haben."
Zum Erkenntnisproblem.
86
Der
Bezeichnung.!
Sprachgebrauch
laxere
moderne Empirismus, namentlich mit
wenn der
vor,
liegt
der Entwicklungslehre,
Hilfe
auch das a priori aus der Erfahrung abzuleiten sucht.
A priori ist
was nicht das einzelne Individuum
sondern was
dann
das,
Niederschlag der Erfahrung des gesamten Menschengeschlechts
als ein
kommt und seinem Denken bestimmte
durch Vererbung an jenes
Bahnen
So schöpft
vorschreibt.
nicht der Einzelne,
anderes Problem
gröbliches
erst
Kants,
ein völlig
und es ist
wenn man glaubt, Kant
und Spencer
zu
denken
schärfer
das des absoluten a priori
Mißverständnis der Sache,
Darwin
durch
als
wohl aber die
Das ist jedoch
Menschheit lediglich aus der Erfahrung.
ein
erwirbt,
lernen,
widerlegen
ehe
man
Man
können.
sich m.it solchen
sollte
Problemen
befaßt.
Wenn schon solcher Wandel und solche Unsicherheit im Ausdruck verwickelte Probleme der Sache vermuten
so
stellt
die
Geschichte der Philosophie diese klar und deutlich vor Augen,
sie
jahrtausendlangen
einen
zeigt
^
Kampf,
Dieser Kampf war aber bei
schwillt.
der
läßt,
immer
gewaltiger
an-
aller seiner Leidenschaft wenig
Als Übersetzung von a priori diente in früheren Jahrhunderten «von
vornen her", das schon
Luthers Tischreden vorkommt
in
(s.
Ausgabe von
Förstemann IV, 399) und sich bis ins 18. Jahrhundert erhält. Als
Quelle für „von vornherein" bezeichnet Campe Lessings Ernst und
erste
Falk,
auch ich kann den Ausdruck nicht weiter zurückverfolgen. Er bezeichnet
aber den Begriff nur in dem laxeren, bloß relativen Sinne, Absolut verstanden,
begegnet sich a priori mit „rein", das auch eine lange Geschichte
Anaxagoras voüc xaO^apd;
hat.
Seit
yoCiv
cpif)aiv
«utov
(d. h.
(s.
Aristoteles
de anima 405 a 16:
(xo'vov
tov vouv) tüJv ovtwv octcXoGv Eivat y.ai «[Aty^ T£ xa"i xaO-apov)
dem älteren Sprachgebrauch das Einfache, Lautere, Ungemischte
zum Gemisch der sinnlichen Welt. Die Neu-
bedeutet es
des Geistigen im Gegensatz
platoniker
und
ihrem Gefolge das Mittelalter übertragen den Begriff auf
in
das Erkennen und nennen rein eine von allen sinnlichen Bildern freie Erkenntnis
(s.
z.
B. Scotus Erigena,
pura
erklärt die intellectio
reas versatur.
stand",
ihr
si
in
de div. nat. 657 D, 658 B).
Auch Descartes
eine solche quae circa nullas imagines corpo-
In diesem Sinne
während
nimmt
die Wolffsche Schule
„reinen Ver-
den Gegensatz zu Erfahrung bildet
ratio pura
entspricht (s. Wolff psych, emp. § 495
„reine Vernunft"
und damit dem a priori
est,
als
:
non admittimus nisi definitiones ac propositiones
Auch Gottsched folgt diesem Sprachgebrauch, s. z. B.
ratiocinando
a priori cognitas).
=
Erste Gründe der gesamten Weltweisheit (1739), S. 485 reiner Verstand
ohne sinnliche Vorstellungen, S. 486 reine Vernunft, == wenn sich in unsere
Vernunftschlüsse keine Erfahrungssätze mit einmengen.
So entspricht Kants
Verwendung von reiner Vernunft dem gelehrten Sprachgebrauch.
—
Denken und Erfahrung.
weil
fruchtbar,
er
das Problem
Stelle
wo
aufnahm,
Man stritt darüber, woher das Erkennen
Entscheidung liegt
seine
87
an der
nicht
stamme, ob aus der Mitteilung der Dinge oder der Selbsttätigkeit
Das wäre aber
des Denkens.
direkt zu entscheiden nur,
wenn das
Was des Erkennens, unser Erkenntnisbesitz, allem Zweifel enthoben
wäre, nicht die Frage nach dem Woher immer wieder in die nach
dem Was zurückgriffe. Dieses aber geschieht in Wahrheit. Wir
sind keineswegs einig über den Tatbestand des Erkennens, sondern
die
grundverschiedene Bilder ein und beweisen
setzen
Streitenden
von ihnen aus, sie beweisen damit immer nur für sich selbst, nicht
für die anderen;
die geschichtliche
Bewegung wird
Monologen, welche die Gegner nicht
einen fruchtbaren Austausch
in
die eigene
Behauptung
nicht
zu ermitteln
Das Was des Erkennens aber
hineintreibt.
ohne
in
sondern jeden nur immer weiter
bringt,
ein Zurückgreifen
eine Folge von
ist
auf die letzten Fragen,
im besonderen auf
dem unsere Untersuchung überall begegnet,
das eine Grundproblem,
das Problem, ob das Leben und Streben des Menschen lediglich die
Bewegung der Natur fortsetzt, oder ob es eine neue Stufe der Wirklichkeit einführt.
Im eigenen Gebiet des Erkennens aber erzeugt der
über seinen Ursprung immer wieder die Frage,
Streit
ob neben
den Einzelwissenschaften auch eine selbständige Philosophie möglich
und
nötig
ist,
so daß
den
in
Streit
auch dieses Problem mit hin-
einspielt.
So gewiß die Frage des Ursprunges der Erkenntnis die Arbeit
der Philosophie
seit
Plato
begleitet,
der Neuzeit erlangt.
erst in
zuerst das Seelenleben
eine
leitende
hat
sie
Das aber aus dem Grunde, weil nun
und die Weltumgebung deutlich auseinander-
zugleich aber die einzelnen Seiten ihr
traten,
zulegen
Stellung
und gegeneinander abzugrenzen
Vermögen näher dar-
hatten.
Sie
traten
aber
weiter auseinander, nicht weil ein gesteigerter Scharfsinn auf diesen
Gedanken kam, sondern
weil
das Leben in seinem Grundbestande
nach entgegengesetzten Richtungen auseinanderging.
wann
die durch lange Arbeit
Einerseits ge-
und mannigfache Erfahrungen
selbst vertiefte Innerlichkeit ein so starkes Selbstgefühl,
in
sich
um sich für
den Kern der Welt erklären und den Versuch eines Aufbaues der
ganzen
Wirklichkeit
aus
selbständiger
Gedankenarbeit
wagen
zu
können; andererseits erhob sich die sinnliche Welt, unter Abwerf ung
der
mittelalterlichen
Verschleierung,
so
mächtig
gegenüber
dem
Zum Erkenntnisproblem.
88
Menschen und
eine
zeigte
ermeßlichkeit des Lebens,
daß
Baues,
Festigkeit des
solche
Un-
wie
aus sicherer Überlegenheit auch
sie
dem menschlichen Dasein und damit dem Erkennen
seinen
Inhalt
zuzuführen schien.
Ein so schroffer Gegensatz verbot
alle friedliche
auf einer der Seiten mußte der Schwerpunkt
Vereinbarung,
und
liegen,
je
nach
der Entscheidung das Erkenntnisbild sich grundverschieden gestalten.
So entstanden die Systeme des Rationalismus und des Empirismus
mit
entgegengesetzten
ihren
Durchblicken
der
Wirklichkeit.
Der
Empirismus nimmt seinen Standort beim Bewußtsein des Einzelnen,
mit einleuchtender Klarheit vermag er zu zeigen, wie dies Bewußt-
sondern ihn langsam, von
und unter Leitung der Umgebung ge-
seinen Inhalt nicht fertig mitbringt,
sein
Eindrücken
einzelnen
her
Die Philosophie
winnt.
hatte
hier
Bewußtsein zurückzubeziehen, nur
sie
nur die Erkenntnisse auf das
als empirische
sich in diesen Zusammenhängen halten.
schließlich
'
eine bloße Assoziation
Psychologie konnte
Das Erkennen wird hier
von Empfindungen und
Vorstell-
ungen ohne allen inneren Zusammenhang, auf eine Durchleuchtung
der Wirklichkeit wird ganz und gar verzichtet.
heißen
ob
kann,
ja
hinauskommen und
ein
gemeinsamer
bleibt
bestreitbar
schaft
winnen
das
läßt,
Gründen
worden.
bestritten
dabei
sich
Ob das noch Wissen-
über die bloßen Individuen
Besitz
und
ist
der
Menschheit
mit
alsbald
ge-
triftigen
Völlig anders der Rationalismus.
Sein
Ausgangspunkt ist die Tatsache der Wissenschaft; ihre Eigentümlichkeit präzis
daß
erfassen, das scheint
nicht
sie
ihm die Überzeugung zu begründen,
von außen her dem Menschen
sondern nur
zufallen,
aus der Werkstätte selbsttätigen Denkens hervorgehen kann.
lich
sind
nisse,
anders
es
welche
Nament-
formale Eigenschaften der wissenschaftlichen Erkenntalle
Ableitung von außen zu verbieten scheinen.
Was
kann die Quelle der ewigen und allgemeingültigen Wahr-
heiten sein, die das Gebäude der Wissenschaft tragen, als die eigene
Natur des Geistes?
Es wird aber damit das Erkennen vornehmlich
ein volles Herausarbeiten dessen,
was dem Vernunftwesen von Haus
aus innewohnt; das analytische Verfahren bildet den Kern der wissenschaftlichen,
namentlich der philosophischen
gestaltet sich die
Arbeit;
einem Leibniz
Philosophie zu einer universalen Mathematik, welche
die Voraussetzungen des Erkennens
immer weiter zurückschiebt und
Gleichungen
mehr und mehr
die
umsetzen möchte.
Aber wenn damit ein systematisches Gefüge der
ganze Wirklichkeit
in
rationale
Denken und Erfahrung.
89
Wissenschaft erreicht wird, so verwandelt sich zugleich die Welt mehr
und mehr in
von bloßen Formen und Beziehungen, die
immer blasser und ärmer zu werden. So mußte
ein Reich
Wirklichkeit droht
der
Empirismus
dem unermeßlichen
beherrschende
keine
Stoffe
Form, der Rationalismus aber den Formen keinen genügenden Inhalt
zu geben.
Kant strebte
Kraft nach
ganzer
mit
Überwindung des
einer
Gegensatzes, dessen beide Seiten in seiner eigenen Natur zusammen-
Er gehört insofern auf die
trafen.
Erkennen
energisch
hinaushebt
und
über
ihm
aus
einen
macht.
Aber dieser Rationalismus
schlag
dadurch,
Selbsttätigkeit
daß
das Denken
erzeugt,
sondern
gebunden
Stoffes
ist;
rationalistische Seite, als er das
bloße Assoziation der Vorstellungen
die
bei
einen empiristischen Ein-
die Erkenntnis
ihr
an
nicht aus reiner
Darbietung
die
eines
so kann es nicht eine Welt der Dinge, sondern
nur ein Reich der Erscheinungen erreichen.
wandt ist auch
Zusammenhang
systematischen
erhält
Dem Empirismus ver-
ein starker Tatsachensinn, der überall auf eine präzise
Erfassung des Eigentümlichen und Unterscheidenden dringt, während
der Rationalismus dies zu Gunsten einfacher Gedankenreihen abzu-
Ebenso entschieden wie Leibniz vorwiegend
schleifen geneigt war.
denkt Kant
quantitativ,
denkt dieser
in
vorwiegend
qualitativ,
wie jener
in Stufen,
Das hier unternommene schiedsrichter-
Gegensätzen.
liche Verfahren hat nicht nur den Vorzug, das Problem systematischer
zu
behandeln,
als
je
zuvor geschehen war, es hat auch das
dem
Menschen eigentümliche Erkenntnisvermögen mit besonderer Schärfe
zu begrenzen versucht.
eine neue
Aber bei aller Größe der Behandlung, die
Epoche für das gesamte Problem beginnt,
ruft
die neue
Kann das Denken
zugleich an eine fremde Welt gebunden werden und eine Selbstständigkeit bewahren?
Verrät nicht schon der Umstand die große
Antwort sofort neue Fragen und Zweifel hervor.
Verwicklung der Sache, daß Kants Untersuchung nirgends umständ-
wo es gilt,
die Denkfunktionen
und die Sinneseindrücke zusammenzubringen?
Auch kann das Er-
licher
und
gebnis
des
künstlicher
ist
als
Schiedsspruches
Rationalisten nicht,
da,
keine der
Parteien
befriedigen.
Den
weil Kants gewaltige Steigerung der Denkarbeit
unvermeidlich über die Bindung an ein Ding an sich und die Be-
schränkung auf ein Reich von Erscheinungen hinaustreibt, den Empiristen
nicht, weil er die
Frage aufwerfen kann,
ja
aufwerfen muß,
ob jenes Flechtwerk von Formen, das nach Kant die Erfahrung erst
Zum Erkenntnisproblem.
90
möglich macht, nicht umgekehrt aus ihr selbst allmählich erwachsen
sei;
damit aber würde auch sein Sinn ein wesentlich anderer werden.
Die
unsichere
Lage,
die
in
das Erkennen
solcher Streit
würde weit mehr empfunden werden, brächte
versetzt,
nicht die praktische
Philosophie der Gedankenwelt eine Ergänzung wie eine Befestigung.
Aber auch
ihre
Begründung ist
nicht allem Zweifel enthoben.
So kann es nicht befremden, daß die Bewegung der Philosophie
über die kantische Lösung des Erkenntnisproblemes hinaustrieb,
daß
der
Auch
nun
Gegensatz
erst
Wandlung,
hierher wirkte die geistige
Charakter des
19. Jahrhunderts
schichtlich-gesellschaftlichen
noch
fernlag. ^
Ansicht
war.
Spannung
die
am
ja
erreichte.
den
meisten
das Aufsteigen einer ge-
bestimmt,
der Wirklichkeit,
wie
Kant
sie
Dieser neuen Denkweise konnten sich beide Parteien
bemächtigen und mit ihrer Hilfe zu
gelungen
höchste
seine
Die
Geschichte
leisten
erhielt
suchen, was bisher nicht
dabei
hier
und dort
ein
grundverschiedenes Ansehen: dort ward sie eine einzige, von innerer
Notwendigkeit getriebene Bewegung, hier ein Sichaufschichten einer
Wendung
endlosen Mannigfaltigkeit.
Mit der
wuchs der
zur spekulativen
Rationalismus
zu jener Geschichte
Konstruktion,
den
die
Denkprozeß -durch seinen eignen Fortgang die ganze Wirklichkeit
erzeugen
ließ
und
Tatbestand
allen
ein
Die Erfahrung
bloße
der Vernunft zu verwandeln suchte.
fahrung
sollte
hier ganz
Werk
mehr und mehr in
verschwinden.
Das
als
'Er-
analytische Verfahren
des älteren Rationalismus wich damit einem synthetischen, die Philo-
sophie gestaltete sich zu
einer
weltumspannenden,
Geschichte durchwaltenden Logik,
sie
zog
alles
namentlich
die
echte Erkennen an
sich und ließ den Einzelwissenschaften keinerlei Selbständigkeit.
Völlig
Ihm bot namentlich die Entwicklungslehre naturwissenschaftlicher Art Mittel und Handhaben zu
umgekehrt verfuhr der
Empirismus.
dem Unternehmen, allen vermeintlichen Eigenbesitz des Geistes aus
der Erfahrung abzuleiten.
Das Erkennen wurde hier eine wachsende
Wie Kant, in Einklang mit dem älteren Rationalismus, der PrinzipienBewegung versagt, zeigt u. a. folgende Stelle aus
der Vorrede zur Kritik der reinen Vernunft (III, 11 Hart.): »Nun ist Meta*
lehre
eine geschichtliche
physik nach den Begriffen,
aller Wissenschaften,
die wir hier davon geben werden,
die sich eine solche Vollendung,
die einzige
und zwar
in
kurzer
und mit nur weniger, aber vereinigter Bemühung versprechen darf, so
daß nichts für die Nachkommenschaft übrig bleibt, als in der didaktischen
Manier alles nach ihren Absichten einzurichten, ohne darum den Inhalt im
Zeit,
mindesten vermehren zu können."
Denken und Erfahrung.
91
die Umgebung, der Kampf ums Dasein sollte diese
immer zweckdienlicher, immer ökonomischer gestalten; daraus sollte
alles entstanden
sein, was unser Denken an durchgehenden Rich-
Anpassung an
tungen und Formen aufweist, und was sich vom bloßen Individuum
her als ein a priori ausnimmt.
und
Alles innere
logische
Gefüge
der Erkenntnis weicht dabei der bloßen Tatsächlichkeit, alles Erklären
dem bloßen
Schildern.
Philosophie,
alles
Hier
ist
kein Platz
eine selbständige
für
Erkennen echter Art wird zur Naturwissenschaft.
Nur als ein Herausheben und Zusammenstellen ihrer Hauptergebnisse
kann die Philosophie sich behaupten.
Die wirkliche Arbeit des
1 9.
Jahrhunderts ging zwischen diesen
Gegensätzen ihren Weg; begünstigte in seinen ersten Jahrzehnten das
hochgespannte Selbstgefühl des Menschen und die vorwaltende Beschäftigung mit den Fragen der inneren Kultur den Rationalismus,
so unterstützte die Wendung des Lebens
zum sinnlichen Dasein und
der unermeßliche Zustrom von Tatsächlichkeit aus Natur, Geschichte,
politisch-praktischem Leben den Empirismus.
Fühlte sich vorher der
Mensch im Mittelpunkt der Wirklichkeit und
durfte er glauben, bei
sich ihre
Fäden zusammenfassen und aus seiner geistigen Tätigkeit
ihr anfängliches
Dunkel gänzlich klären zu können, so übermannt
ihn jetzt das Bewußtsein seiner verschwindenden Kleinheit; aus
Zentrum
in
die
die Wirklichkeit
Peripherie verwiesen,
von
darf
nicht
er
sich aus hervorzubringen,
dem
mehr hoffen,
muß er vielmehr be-
scheiden und unterwürfig ihre Eröffnung erwarten.
Aber nicht nur
solche Notwendigkeit, auch ein inneres Verlangen treibt den Menschen
zur Hingebung an die Erfahrung.
Unmittelbarkeit,
als
sie
die
der Welt
in
mehr
Es
Tatsächlichkeit,
ist
die Sehnsucht nach mehr
mehr Reichtum des Lebens,
Gedankenwelt des Rationalismus mit ihrem Einfangen
ein Netz
freischwebender Begriffe
und Formen
bot.
Dies Verfahren beginnt als eine Verarmung und Verflüchtigung des
Lebens empfunden zu werden; dem gegenüber wird »ein unersättliches
Verlangen nach Realität die gewaltige Seele der gegenwärtigen
Wissenschaft" (Dilthey).i
^
Mit Recht sagt James (Pragmat.
S. 9):
„Seit hundertundfünfzig Jahren
scheint der Fortschritt der Wissenschaft nichts anderes zu bedeuten als eine
Vergrößerung der materiellen Welt und eine stete Verminderung der
Bedeutung des Menschen. Das Resultat ist die Zunahme der naturalistischen
stete
und der positivistischen Fühlweise." S. auch S. 8: «Niemals hat es so viele
Menschen von entschieden empirischer Geistesrichtung gegeben als heutzutage."
Zum Erkenntnisproblem.
92
Natürlich fehlte es nicht an Versuchen, zu vermitteln
und
aus-
Eine Wiederaufnahme kantischer Denkweise lieferte den
zugleichen.
Nachweis, daß die Erfahrung, so wertvoll
mag, aus eignem
sie sein
Vermögen nun und nimmer ein wissenschaftliches Erkennen erzeugt,
daß es dazu einer fortwährenden Hilfe des Denkens bedarf. In
ähnlicher Gesinnung
ließ
schaften
Voraussetzungen
fertigen
vermögen,
solche
sich
daß die einzelnen Wissen-
dartun,
enthalten,
die sie selbst
nicht zu
die vielmehr über sie hinausweisen.
Gegenbewegung war mehr
recht-
Aber eine
negativer als positiver Natur, sie
mochte ungelöste Rätsel jenseit der Erfahrungswelt zeigen, nicht aber
eröffnete sie ihr
gegenüber einen neuen Lebens- und Gedankenkreis,
nicht trieb sie zu einer eigentümlichen philosophischen Betrachtungs-
weise und
in diesen
die Philosophie
einer selbständigen Philosophie;
wurde
Zusammenhängen lediglich eine kritische und reflektierende
Umsäumung
der Einzelwissenschaften,
dem
die
Fachgelehrten die
anziehendste Beschäftigung bieten mag, die aber zur Erhöhung des
Geisteslebens
kaum etwas beiträgt, die zugleich, beim Mangel eines
beherrschenden Prinzips,
zu
die Subjektivität
überwinden vermag.
punkte
nicht
Ideen-
und Überzeugungswelt,
tausenden besessen
hatte.
wie
die
bloß individueller Stand-
So entfiel eine gemeinsame
Menschheit
sie
seit
Jahr-
Und das Ungeheure dieses Verlustes, die
damit drohende Zersplitterung und innere Verarmung der Mensch-
kam gegenüber der Freude an der massenhaft zuströmenden
kaum zur Empfindung. Das kann jedoch nicht lange
Denn das Verlangen nach einer zusammenhängenden
so bleiben.
Gedankenwelt und einer inneren Einheit des Lebens wurzelt zu tief,
heit
Tatsächlichkeit
um sich auf die Dauer unterdrücken zu lassen; schon heute ist der
Beginn einer Gegenbewegung deutlich genug.
die
Einzelwissenschaften
selbst,
indem
ihr
Mehr Einheit fordern
eigner
Ausbau
sie
zu
einer gründlicheren Beschäftigung mit ihren Prinzipien und Voraussetzungen führt, diese aber zur Aufdeckung der Zusammenhänge
mit anderen Gebieten
treibt.
Von den
und zu einem Streben nach einem Ganzen
Seiten erschallt wieder der Ruf
verschiedensten
nach einer Synthese.
Die Synthese
ist
aber nicht echt, so lange die
Verbindung eine bloße Zusammenstellung bleibt; zur Wurzel durchgreifen kann sie nur bei kräftiger Herausarbeitung gemeinsamer
Ideen
und Überzeugungen, das aber fordert einen dem Nebenein-
ander überlegenen Standort, es fordert eine selbständige Philosophie.
Stärker noch drängt dahin das
gemeinsame Leben.
Die Kehr-
Denken und Erfahrung.
Seite
93
der völligen Hingebung an die unmittelbare Welt, der völligen
Verwandlung des Lebens
in Arbeit
immer weniger über-
läßt sich
sehen: der Mangel einer Zurückbeziehung auf eine überlegene Eindie
heit,
das
Leben
verwandelt,
Besitz
allererst
die
und damit
Selbstleben
in
in
äußeren Eindrücke, die Ungewißheit über das Ganze bei
festigung
im
Einzelnen.
Alles
geistige
eignen
überströmender Fülle der
geistige Leere bei
aller
und zugleich
Leben
Bealler
Sinn und Wert unseres Daseins gerät damit in Zweifel, der Boden
entschwindet
dem Menschen unter den Füßen; so treibt es zwingend,
wieder
die
auf
letzten
und einen Kampf
Grundlagen seiner Existenz zurückzugehen
um die Erhaltung einer Seele zu führen.
Solche
Probleme verbieten einen Abschluß bei der bloßen Erfahrung,
treiben uns, neue Möglichkeiten zu suchen
sie
und unser Verhältnis zur
Wirklichkeit gründlich zu revidieren. Zugleich aber tritt die Philosophie
wieder auf den Plan, die Philosophie nicht
als
zur Bearbeitung der Erfahrungswelt, sondern
eine bloße Gehilfin
als die
Trägerin eines
eigenen Gedankenreiches, als eine Kraft des Schaffens und Weiterbildens.
b)
Das Recht einer selbständigen Philosophie.
Die Frage nach der Selbständigkeit der Philosophie bilde den
Beginn der Untersuchung, da ihre Beantwortung über den Gesamtanblick des Erkennens entscheidet.
Aber wie ist hier zu einer Antwort
Daß sich beim Erkenntnisproblem nicht unmittelbar an
zu gelangen ?
die Vergangenheit anknüpfen, sich nicht
aufnehmen und fortführen
läßt,
von dort ein Faden einfach
das zeigte der Überblick der welt-
geschichtlichen Bewegung, und das fand sich bestätigt durch die eigen-
tümliche Lage der Gegenwart; wir empfinden
heute
mehr die Ab-
weichung als die Übereinstimmung mit früheren Leistungen, wir sehen
uns von ihnen eher Wege versperrt
des
geistigen
Lebens
trieb
als
gewiesen.
Natur und Seele
weiter auseinander, sie verwehrt damit auch
Die Entfaltung
innerlich
weiter
und
dem Erkennen ein un-
mittelbares Zusammenfassen beider, sie zwingt es zu einer Entscheidung
für
das
eine
oder das andere.
So geschah
es,
grundverschiedene
Weltbilder erschienen und rissen die Wahrheit an sich.
war stark genug,
trieb es die
Aber keins
um das Feld gänzlich einzunehmen, immer wieder
Forschung vom einen zum andern zurück.
Solche Er-
fahrung empfahl eine friedliche Verständigung, eine schiedsrichterliche
Abgrenzung; eine solche schien am ehesten erreichbar durch
Zum Erkenntnisproblem.
94
ein
Anerkennen verschiedener Faktoren im Erkennen und die Zu-
Das geschah durch
weisung des einen hierher, des anderen dahin.
von
Unterscheidung
kantische
die
Form und
Lösung scheitert an der Schwierigkeit,
durchaus
sondern
nur verschiedenen,
zusammenzubringen,
das
von
eine
beiden
die nicht
Faktoren
logische
Tätigkeit
So scheinen wir uns weder
entscheiden,
noch
beides
miteinander
Zu solcher Erfahrung aus dem Ganzen der
zu können.
festhalten
Aber diese
verschiedenartigen
Empfindung und
sinnliche
zu gemeinsamer Wirkung zu verknüpfen.
für
Stoff.
UnmögUchkeit,
ja
Geschichte gesellen sich widerstreitende Eindrücke und Antriebe der
Gegenwart.
bloß
mit
zugleich
Macht.
Wir empfinden mehr und n^ehr die innere Leere eines
befaßten Lebens und Denkens,
der Erfahrungswelt
aber
umklammert uns die Erfahrung mit immer wachsender
Wir wollen mehr Selbständigkeit des Denkens, aber die Ab-
neigung gegen die spekulativen Systeme
läßt
uns bei jedem Schritte
vorwärts zaudern und macht uns mißtrauisch gegen alle Metaphysik.
Eine so verworrene Lage zwingt, das Problem direkt ins Auge
zu
fassen
und
in
eigner Weise
zu
behandeln.
Was ist es, von
was den Menschen über die Erfahrungsund solchem Streben eine Macht verleiht?
Ist es das Denken selbst, dessen Natur auf diesen Weg führt und
auch die Mittel zu seiner Verfolgung gewährt? So hieß es von
dieser Frage sei begonnen,
welt hinausstreben
alters her,
es,
läßt
so hören wir vielfach auch heute.
Das Denken, so scheint
enthält Forderungen, welche die Erfahrungswelt nicht befriedigt,
auf deren Befriedigung aber eine innere Notwendigkeit seines eignen
Wesens
ihr
es
bestehen
heißt.
So muß
es jene
Welt verwandeln,
ja
gegenüber eine neue entwerfen, da doch die eigne, innere Not-
wendigkeit ihm mehr als
alle
Eindrucke der Umgebung gelten muß.
Das wäre einfach und überzeugend, wenn nur nicht die vom Denken
behauptete Notwendigkeit über das Denken hinaus gelten wollte und
die
von ihm entworfene Welt den Anspruch erhöbe, die eigne Wahr-
heit
der Dinge zu sein.
Das aber geschieht, und indem es geschieht,
überschreitet das Denken den eignen Bereich; das Recht dazu kann
nur durch künstliche Annahmen stützen, deren Verfolgung in
immer weitere Schwierigkeiten verwickelt. Wie ließe sich hier dem
es
Vorwurf entrinnen, daß das Denken nur menschliche Bilder
in
die
weite Welt hineinsieht?
Auch ist es ein eignes Ding mit jener vermeintlichen Denknotwendigkeit. Da sie selbst jede weitere Begründung
ablehnt und ablehnen muß, so
käme die Entscheidung an das Ge-
Denken und Erfahrung.
das Gefühl
an
fühl,
einer
unwiderstehlichen Zwanges.
und
fühlen,
alle
in
schlechterdings
unvermeidlich
nicht
Subjektive
ins
als
Hegel
denknotwendig:
läßt
Wirklichkeit in Bewegung umsetzen, Herbart möchte
Bewegung aus der
alle
Wahrheit ein solcher
Forderungen
widerstreitende
das Denken
eines
läßt sich
In Wahrheit verfechten hervorragende Philosophen
und Individuelle?
direkt
Unabweisbarkeit,
Aber
das Gefühl
führt
Q5
entfernen;
Wirklichkeit
jener
feiert
den
Widerspruch als die treibende und erhöhende Kraft des Weltprozesses,
Wessen Denknotwendigkeit
dieser will ihn in keiner Weise dulden.
ist
die echte, welche soll uns anderen binden?
nun
Eine Weltaufhellung vom bloßen Denken her wäre nur unter Einer
Bedingung erreichbar: das Denken müßte die ganze Wirklichkeit in
sich tragen oder durch seine Bewegung erzeugen; dann wäre das
Selbsterkennen
des Denkens
zugleich
und der
Welterkennen,
ein
Lebensprozeß hätte seine Wahrheit bei sich selbst, er brauchte sie nicht
So trieb die Konsequenz der Sache
bestätigt zu haben.
von außen
von Plotin
die 'Philosophie
und
ihn
ließ
immer wieder auf diesen Weg
bis Hegel
mögliche Überwindung des Spaltes
einzig
die
als
Denken und Sein begrüßen.
zwischen
Aber wie
der Absor-
viel
bierung der ganzen Wirklichkeit durch das Denken widersteht, und
wie sich
dabei
die
Welt
in
ein
zu verwandeln droht, das
griffe
ungen des Hegeischen Systems
bloßes Schattenreich formaler Be-
der Neuzeit durch die Erfahr-
ist
viel
zu deutlich geworden,
um sich
bald vergessen zu lassen.
Wenn
aber
weder das Denken mit dem Sein zusammenfällt,
vDn ihm ein draußen befindliches Sein erreichen läßt, so ist
Denken aus überhaupt kein Erkennen möglich,
freischwebenden
vom
im besondern kein Aufbau eines selbständigen Gedankenreiches neben
noch
sich
der
Erfahrungswelt.
darauf,
Alle
daß das Denken
Aussicht
eines
Gelingens
in weitere Zusammenhänge
beruht
tritt
also
und damit
ein anderes Verhältnis zur Wirklichkeit gewinnt; das aber tut es in
von vornherein den ganzen
intellektuellen Kreis des Menschen, sondern dieser wird zunächst von
den Assoziationen der einzelnen Vorstellungen mit ihrem mecha-
Wahrheit.
nischen
auf
Das Denken
Getriebe
nicht
eingenommen; das Denken mit
das Gegenständliche,
tischen
bildet
seinen
inneren Gesetzen,
seiner
Richtung
seinem synop-
Umspannen der Mannigfaltigkeit gegenüber dem sukzessiven
Verlaufen der Vorstellungsketten hat sich dagegen erst aufzuringen
und durchzusetzen.
Das aber kann
es
nur,
und
es
ist
überhaupt
Zum Erkenntnisproblem.
96
lebendige
eine
neuen
kommen
nur
Kraft
Lebensstufe,
die
vom bloßen
Damit aber
wie wir ihn im
des Geisteslebens,
lernten.
Im
geistigen
Wendung der gesamten Wirklichkeit,
eine Tiefe entwickelt
sie
Ausdruck einer
ein
aufsteigt
Seelenleben fassen
Leben erkannten wir eine
der
Menschen
im
wir auf den Begriff
Unterschied
Stück und
ein
als
erst
und
sich zugleich zu
bei
einem Weltleben
am Geistesleben teilgewinnen heißt daher zugleich an
zusammenfaßt;
einem Weltleben teilgewinnen; es sind nicht Erfahrungen des bloßen
Punktes,
es sind
Erfahrungen
vom Ganzen,
ungen und Wandlungen hervorgehen.
die aus seinen
Dies neue Leben zeigte sich
auch dem Gegensatz von Subjekt und Objekt überlegen, es
halbes Sein, das zu seiner Ergänzung eines
dern es
ist
trägt
in
es
kein
ist
Fremden bedürfte, son-
Leben über jenen Gegensatz hinausgehoben,
als volltätiges
sich
Beweg-
den Umriß einer selbständigen Wirklichkeit und
Bewegung zur vollen Durchbildung dieser Wirklichnun ist der Träger des Denkens und
alles Erkenntnisstrebens, nicht der bloße Mensch und das einzelne
Individuum; das Erkennen aber erscheint in neuem Lichte, wenn es
strebt in seiner
—
keit.
Dies
Geistesleben
weder auf sich
selbst
noch auf
ein
draußen gelegenes Sein, sondern
erstwesentlich auf das Geistesleben geht,
von dem es selbst umspannt
wird; Welterkennen kann es nur werden,
wenn das Geistesleben den
Kern der Wirklichkeit
bildet.
Eine solche Begründung im Geistesleben und zugleich ein Weltcharakter
kommt allem Wissen zu, aber es ist leicht zu ersehen, wie
hier die Philosophie eine besondere Aufgabe findet.
Alles Erkenntnis-
streben beruht auf einem Verhältnis von Ganzem zu Ganzem.
Verhältnis
dies
kann
als
eine
stillschweigende
Aber
Voraussetzung
im
Hintergrunde bleiben, und die Arbeit kann sich auf einzelne Gebiete
oder einzelne Beziehungen richten; es bedarf einer eignen Wissenwelche die Sache
schaft,
gründende Tatsache zu
wie
halt
ihre
Stellung
als
Ganzes behandelt,
vor allem die be-
voller Klarheit herausarbeitet
und ihren Ge-
zur umgebenden Welt zu ermitteln sucht;
ist die Philosophie.
So gewiß das GeistesZusammensetzung aus einzelnen Punkten, sondern ein
diese Wissenschaft aber
leben
keine
inneres Ganzes
ist,
so zuversichtlich
läßt
sich
erwarten,
daß
sich
mit der Philosophie ein neuer Weltanblick eröffnet, daß sie bei allem,
was
sie
von den einzelnen Wissenschaften empfangen muß,
ihnen
auch eine selbständige Leistung entgegenzusetzen vermag und von hier
aus
alle
dargebrachten Tatsachen von neuem in Probleme verwandelt.
Denken und Erfahrung.
So
97
der Angelpunkt aller philosophischen
ist
Betrachtung und
das Axiom der Axiome die Tatsache eines weltumspannenden Geistes-
Schon daß überhaupt eine neue Stufe der Wirklichkeit über
lebens.
die Natur
hinaus anerkannt wird, verändert den Anblick der Welt
und
auch
zeigt
Geistesleben
es
ist
es
den Anspruch erheben,
an
sich
zu
Beleuchtung.
Aber
das
der Wirklichkeit zu ihrer eignen
es das Beisichselbstsein des Lebens bedeutet,
gewiß
Stufe sein wollen; als solche aber muß
und abschließende
die letzte
anderer
in
es die Wendung
muß, so gewiß
Innerlichkeit, so
Natur
die
nicht nur irgendwelches Mehr gegenüber der Natur,
alles
von
sich aus zu verstehen
Dieser Anspruch
messen.
aber
führt
und alles
notwendig zur
Frage, wie weit das im
Menschen gegenwärtige Geistesleben solcher
Aufgabe gewachsen
die Widerstände wollen erwogen, die
einer
lichkeit
zusammen
sei,
Überwindung geprüft
ergibt sich eine
Mög-
aus Größe und Schranke
sein,
dem Menschen eigentümliche Art.
zusammen stellt der Philosophie eine besondere Aufgabe
Alles
und
eröffnet
wird
damit
ihr
einen
selbständigen
charakteristische
Züge
Weltdurchblick,
empfangen,
von
ihre
Arbeit
denen
hier
namentlich drei herausgehoben sein mögen.
Wenn die Philosophie vom Ganzen des Geisteslebens zum
1.
Ganzen der Wirklichkeit strebt,
eines gegebenen
stellen,
sie
so liegt ihre Arbeit nicht innerhalb
Raumes, sondern
findet nicht ihre Welt,
das Ganze, das sie sucht,
sie
hat diesen
sondern
sie
Raum erst herzu-
hat sie erst zu bilden;
tritt
ihr nie von außen her entgegen, es will
von innen her entworfen sein,
es verlangt eine Synthese schöpferischer
Art.
Zur Selbständigkeit
wird
dieses
Weltbild
der
Philosophie
namentlich dadurch getrieben, daß das von ihrer Synthese umspannte
Dasein ohne
Umwandlung
was
ist
es bietet,
sammenzufügen.
viel
nicht in sie einzugehen vermag.
Denn
um sich ohne weiteres zu-
zu verschiedenartig,
Namentlich das Zusammentreffen von Natur und
Innenwelt in Einer Wirklichkeit treibt zwingend zur Umwandlung des
ersten Anblicks.
Schon dadurch
ist
namentlich der modernen Ge-
dankenarbeit ein Trieb zur Metaphysik unzerstörbar eingepflanzt, daß
die Neuzeit den Gegensatz
gebracht
hat,
und daß
von Natur und Seele zur vollen Klarheit
dieser Gegensatz
sich
beim Versuch einer
unmittelbaren Zusammenfassung notwendig zu einem unerträglichen
Widerspruch
sophischen
steigert.
Tätigkeit
Zugleich
selbst,
wie
ist
auch das eine Sache der philo-
viel
von
dem Umfange
unseres
Lebens- und Gedankenkreises in jene Synthese eingeht und zu ihrer
Eucken, Grundbegriffe.
4. Aufl.
7
Zum Erkenntnisproblem.
98
Denn
Gestaltung mitwirkt.
nicht alles uns irgend Bekannte
ist
bei
Auch der beherrschende
Mittelpunkt, der alles um sich gruppiert und das Ganze eigentümlich
gestaltet, will immer erst gewonnen sein und läßt sich an verschie-
jener philosophischen Synthese gegenwärtig.
denen
suchen.
Stellen
Nachdem
die
Die
mittelalterliche
Gestaltung
Umkreis der Religion
ganzen
gehen
Zeiten
weit
hier
Geisteslebens
des
unterworfen
auseinander.
seinen
erhob die Auf-
hatte,
klärung das Verlangen nach einer größeren Weite der Wirklichkeit;
sie
fand
aber
eine
solche
in
dem Nebeneinander von Natur und
einzelnen Seelen, einem Nebeneinander, das sich nicht ohne Gewalt-
beherrschenden
samkeit einer
Einheit
unterwerfen
Die
ließ.
von
Kant anhebende Bewegung erzeugt den Begriff eines selbständigen
Geisteslebens und macht es mit seiner geschichtlich-gesellschaftlichen
zum Kern des Ganzen.
Entfaltung
Aber da
sie
das Geistesleben
mehr und mehr in das bloße Denken setzt, so wird
die davon
um-
spannte Wirklichkeit zu eng, ein Rückschlag wird unvermeidlich, und
dem Gesichts-
dieser Rückschlag droht das Geistesleben wieder aus
kreis der Philosophie zu rücken
und damit in das Wirklichkeitsbild
der Aufklärung zurückzufallen; zugleich wird der Mangel eines be-
herrschenden Mittelpunktes peinlich empfunden, den in Wahrheit nur
ein selbständiges Geistesleben zu
Daß
so
immer von
Wirklichkeit in
springende
die
gewähren vermag.
zum Probleme
neuem
Punkt der Synthese
liegt,
Freiheit der philosophischen Arbeit;
den einzelnen Wissenschaften wird
deutlich
zeigt
bei allem
sie
wie
wird,
philosophische Synthese eingeht,
viel
und wo der
größere
die
Zusammenhange mit
durch die Forderung kühnen
Vordringens und mutigen Vorausentwerfens zur Spekulation getrieben.
Die Hilfe
intellektueller Phantasie
dabei unentbehrlich; was aber
an Gestalten entwirft, das wird
diese Phantasie
sie
dem Menschen
machen können, ohne eben der Erfahrungswelt
eindringlich
nicht
ist
Bilder zu entlehnen, über welche die Philosophie hinausführt.
Das
alles
ist
Gefahren,
voller
großes Unternehmen, und
sein
in
Freiheit
Welt
in
eine
Gefahren
der
aber ohne solche gibt es kein
wenn die Philosophie unser ganzes Da-
verwandeln möchte und uns aus einer gegebenen
eigne,
selbstgebildete
Freiheit
tragen.
versetzt,
Immerhin
so
muß sie auch die
erscheint
das
Wagnis
der Philosophie bei unserer Fassung in ganz anderer Beleuchtung
als in
den Systemen freischwebender Begriffskonstruktion.
Denn bei
uns geht das Streben vor allem auf eine Tatsache, die das Denken
Denken und Erfahrung.
begründet, auf die Tatsache
selbst
lebens;
ist
was
ihm
in
das
liegt,
gg
weltumfassenden
eines
als eine Tatsache
ist
Geistes-
zu ermitteln, das
aufzuweisen, nicht abzuleiten; wie es zur umgebenden Welt steht,
welchen Widerstand es
windung
ihr
in
der Tatsächlichkeit, aber
und wie
findet,
weiterzubilden
selber
sich
das
hat,
es zu seiner
Über-
eine
Frage
alles
ist
einer Tatsächlichkeit, die nicht
freilich
von
Zusammenfassung des Lebens,
außen her zufallen kann, sondern
in
im Aufklimmen zu
und Messen vom Ganzen zum
einem Sehen
Ganzen immer neu zu erringen ist.
Darin steckt eine freie Tat,
die sich keiner Zeit und keinem Individuum aufzwingen läßt, die
damit aber keineswegs eine Sache
und Ge-
individuellen Beliebens
schmackes wird.
2.
Erst die Philosophie rechtfertigt ein Streben über ein bloßes
Kennen der Dinge hinaus zu einem Erkennen.
nichts anderes als ein
selbstfinden,
ein
Hineinziehen
Selbsterkennen.
Erfahrungswelt mit ihrem
nahme der Dinge
Denn Erkennen ist
das eigene Leben,
in
Das
bietet
nun und nimmer die
Nebeneinander, aber auch
das subjektive Seelenleben,
in
liche
Fürsichsein, wird es nicht erreicht.
sein
trägt
ein Sich-
mit der Auf-
das bloßmensch-
Denn ein solches Fürsich-
bloß die eigne Zuständlichkeit in die Welt hinein, es ver-
menschlicht
sie
und
daher auch
ist
in
Vollendung
seiner höchsten
dem Grade nach von der kindlichen Personifizierung der Um-
nur
gebung verschieden, welche die Anfänge kennzeichnet. Eine Innerlichkeit, die den Dingen nicht von draußen aufgedrängt wird, sondern
eignes Sein
ihr
ihnen
erschließt,
sucht
selbst
in
Kraft
die Widerstände
Kampf
mit
phie aber
zum
erst
im Geistesleben, das
findet,
das
mit seiner
in
innere
ist es,
sie
in
sich
umspannenden
verwandelt und den
Die Philo-
gestaltet.
welche diese Bewegung zur inneren Durchleuchtung,
der Wirklichkeit
auf
sich
nimmt.
dem Menschen
Vergeistigung der Welt
wie weit
Hemmungen
ihnen zu einem inneren Erlebnis
Verstehen
solche
erscheint
und
einer gegebenen Lage gelingt,
Wie
gelingen
das
weit eine
kann,
und
eine andere
ist
Frage, aber schon die Aufwerfung des Erkenntnisproblems bekundet
eine
völlige
bloßen
Wendung und
Kennen der Dinge.
zerstört
Alle
lassen die Tatsache unangetastet,
alle
Befriedigung
Hemmungen und
bei
alle
einem
Zweifel
daß beim Menschen eine Aufhellung
der Wirklichkeit beginnt; wie aber könnte er über das Ganze der
Welt irgend denken, dächte er nicht aus dem Ganzen der Welt?
So
treibt
die
Bewegung zwingend über alles bloße Anordnen und
Zum Erkenntnisproblem.
100
Aufschichten
der
Erscheinungen hinaus
wäre das
Erringen einer Seele;
ihnen
nicht
irgendwie
Die berufene Vertreterin dieses Verlangens nach Seele
Philosophie;
die
ist
Leben und Denken
menschliche
überlegen.
aber
zum
könnten nicht als Schranken empfunden werden,
selbst die Schranken
sie
kann
Aufgabe
jene
der
Verinner-
lichung der Wirklichkeit mit besonderem Nachdruck angreifen,
wo
das Geistesleben mit voller Klarheit als der Träger jenes Strebens
anerkannt und die ganze Weite des Daseins zu ihm in Beziehung
gesetzt wird.
Endlich
3.
ist
es die
des Erkenntnisstrebens mit
Philosophie, welche den Zusammenhang
dem Ganzen des Geisteslebens deutlicher
herausstellt, als es irgend sonst geschieht,
und damit jenem Streben
mehr
verleiht.
Sicherheit,
Kraft
und
Bedeutung
Die Philosophie
bedarf jenes Lebens, weil nur sein Gehalt und seine Kraft
den Stand
Suchen
einer
fruchtlosen
Reflexion
in ein sicheres Schaffen
erhebt
bringt; das
sie
über
und aus tastendem
Leben bedarf der Philo-
sophie, weil es nur durch sie seine volle Durchleuchtung, Einigung
und Ursprünglichkeit erreicht.
Wie die Philosophie aus dem Gesamtleben hervorgeht und sich
nach seiner eigentümlichen Lage verschieden
jede Vergleichung verschiedener Zeiten
gestaltet,
das zeigt uns
und verschiedener Kulturen.
Wie grundverschieden ist z. B. die Art und das Streben der Philoim indischen und im vorderasiatisch -europäischen Kulturgemäß dem Typus des Lebens hier und dort! Dort weniger
eine Durchdringung und Überwindung der Welt als eine Ablösung
und Befreiung von ihr, nicht eine Steigerung des Lebens, um es
sophie
kreise
auch gegen die härtesten Widerstände durchzusetzen, sondern eine
Herabstimmung,
Verschwinden,
dagegen ein
ein
eine
Auflösen
tiefsinnige,
aller
Härte,
ein
aber tatenlose
kräftiger Lebenstrieb,
Verschwimmen und
Kontemplation;
hier
ein zähes Festhalten des Daseins
gegenüber allen Widerständen, ein Zurückkehren zur Selbstbejahung
aus
aller
durch
Erschütterung und scheinbaren Vernichtung, ein Vordringen
alle
stellung
Hemmungen zur Entwerfung neuer Welten und zur Her-
neuer Lebensformen.
Zugleich wird auch die Philosophie
mehr Eindringen in die Welt, mehr Ringen mit ihren Widerständen,
mehr Fortschreiten durch das Überwinden dieser Widerstände.
Doch wir brauchen nicht in die Ferne zu schweifen, um den engen
Zusammenhang des philosophischen Strebens mit dem Gesamtstande
—
des Geisteslebens zu
gewahren; die eigne Erfahrung des
19. Jahr-
Denken und Erfahrung.
mit
ihn
zeigt
Systeme
freischwebender Spekulation
während
fortreißen,
die
einer
zeugungskraft gewinnen?
zugleich
ändert
hat.
so
schien
Verlangen.
Heute
winzigen
der
statt
heute
die
seitdem
sich
Über-
rechte
Gesamtlage und
die
Tiefe der Welt zu
letzte
der
Vollbesitz
dagegen
des
Kleinheit
und
Dinge,
Wiederbelebung keine
Weil
alle
Wirk-
Vernunft zu verwandeln schien, so durften seine Begriffe
mutigem Vordringen
Welt,
unwiderstehlich
im Mittelpunkte der Welt; wie dieses Schaffen
lichkeit in
hoffen,
die
Grundstimmung des Lebens aufs wesentlichste verDamals fühlte sich der Mensch mit seinem geistigen
die
Schaffen
bei
unsere Väter
konnten
uns ganz fremdartig anmuten und auch durch
sie
Versuche
eifrigsten
Warum
Klarheit.
völliger
Hunderts
101
beherrscht
Menschen
erschließen
Wahrheit
kein
uns
Empfindung der
die
gegenüber
der
kühnes
zu
unermeßlichen
im Zentrum fühlen wir uns an der Peripherie der
faßt
sich
das geistige
Leben nicht zu einheitlichem
Schaffen zusammen, heute bedrängen zugleich uns schwere Verwick-
lungen im eignen Kreise des Menschen.
überhaupt ein
Lage
wieder Kraft dafür sammeln
sichtig
vom Saume
Behauptet sich bei solcher
philosophisches Streben,
Dinge
der
so müssen
wir
erst
und scheinen nur langsam und vorher
einigermaßen
zu
vordringen
können.
Wie aber
wirkt sie auch
trägt
sich
in
Philosophie aus
ein Aufstreben
bloß
Erzeugnis
die
dahin zurück.
Jede
dem Gesamtleben
des ganzen Geisteslebens,
intellektueller
Geschicklichkeit,
und eine Bekräftigung der gesamten
schöpft,
so
große philosophische Leistung
sondern
sie
ist
ein
kein
Werk
geistigen Art, auch eine Selbst-
erhaltung weltumspannender Persönlichkeit.
Das eben kennzeichnet
wahrhaft große philosophische Leistungen, daß sich in ihnen nicht
bloß eine Klärung der Begriffe, eine Erweiterung des intellektuellen
Horizontes vollzieht,
sondern durch ihre Arbeit eine Weiterbildung
des Lebensprozesses
selbst,
Die Philosophie
erfolgt.
Dinge,
sie
sondern
nach
ihrer
sie
ein Wachstum der geistigen Wirklichkeit
liefert
keineswegs bloße Abdrücke fertiger
nimmt selbst am Bilden und Bauen
innersten Natur
teil;
so
ist
keineswegs eine kühle Betrachtung,
Nur ein solcher Zusammenhang mit dem Gesamtleben erklärt die Stellung und Bedeutung
sondern eine Sache kräftigen Lebensaffektes.
der Philosophie im menschlichen Dasein, die sonst einen rätselhaften
Widerspruch enthält
sophie
als
ein
Denn äußerlich angesehen, erscheint die Philo-
Durcheinander von Systemen, die sich gegenseitig
Zum Erkenntnisproblem.
102
und
widersprechen
sich
ihrer
in
Ablehnung
als
Leben
geistige
Wirkung
und ganzen
scheinen, die dazu im großen
aufzuheben
einander
mehr
bei der Menschheit
Zustimmung fanden.
verarmen und verkümmern, wo es alle Beziehung
Aber zugleich sehen wir das
So
zur Philosophie aufgibt.
es
ist
am
wie dürftig wird
Religion;
wie eng,
entsagt!
Der Widerspruch
löst
sich
augenscheinlichsten
der
bei
sie,
wo sie aller Philosophie
bei
Anerkennung jener Ver-
kettung der Philosophie mit dem Ganzen des Lebens.
Denn damit
wird zu ihrer Hauptleistung nicht eine Ablieferung fertiger Lehren,
sondern die innere Erhöhung des Lebensprozesses, der Gewinn an
und Ursprünglichkeit, die Befähigung, mehr Ganzes,
mehr Wesenhaftes in den Dingen zu sehen.
Diese Verbindung der Philosophie mit dem Leben ist auch
Selbständigkeit
mehr
Inneres,
geeignet,
eine
Auseinandergehen
ihr
Richtungen
verschiedene
in
ver-
machen, ohne diese einander gleichzusetzen und damit
ständlich zu
allgemeingültige
Wahrheit aufzugeben.
Das
nämlich
die Entscheidung auch über die Art der Philosophie,
bringt
was als Mittel-
punkt des Lebens erklärt wird, und wie die Gestaltung, die daraus
hervorgeht,
in
Frage,
zum Ganzen
der Wirklichkeit steht
kommt
Zunächst
ob überhaupt eine Zusammenfassung zum Ganzen
voll-
zogen wird, oder das Leben bei einem bloßen Nebeneinander verbleibt;
in
diesem Falle entsteht überhaupt keine Philosophie.
Versuch einer Zusammenfassung aber wird die Hauptfrage
oder
gehört,
sichselbstseins
ob
wozu auch das Durchschnittsleben der Ge-
das natürliche Dasein,
sellschaft
Beim
die,
ob
ein
ihm
überlegenes
Reich
des
Bei-
Lebens mit geistigen Inhalten und Werten den
des
Hauptstandort des Denkens
dort
bilde;
entsteht
der Naturalismus
mit seinem Empirismus, hier der Idealismus mit seiner Verfechtung
eines
a priori.
Eine
Entscheidung
weitere
entsteht
innerhalb
des
Idealismus bei Beantwortung der Frage, wie das aufstrebende geistige
Leben
sich zu
entgegensetzt.
eignen
Kraft
den Widerständen verhält, welche die Weltlage ihm
Glaubt
allen
der
Idealismus
Widerstand
durch
unmittelbar
volle
Entfaltung
überwinden
der
und auch
das scheinbar Feindliche sich ganz und gar assimilieren zu können,
so entsteht ein reiner Idealismus; dieser aber wird
zu spekulativer
Konstruktion und zur Geringschätzung der Erfahrung neigen.
hingegen
der Widerstand
geistige Kraft
entfalten
und
als
überwinden zu
in
zu
groß
lassen,
erscheint,
um
sich
Wo
durch
da wird sich der Pessimismus
der Erkenntnislehre einen Skeptizismus erzeugen;
Denken und Erfahrung.
Denkweise
diese
so
erklärt,
im Sinne des Idealismus und gehört
steckt das Ziel
daher auf seine
Seite,
aber da
sie
das Leben
beläßt
103
es für schlechthin
sie
unerreichbar
unter der peinlichen Macht eines
Wo aber der Widerstand wohl in
fundamentalen Widerspruches.
seiner Schwere anerkannt wird, aber eine Weiterbildung des Lebens
trotzdem möglich scheint, die wenigstens seinen innersten Kern von
Lähmung befreit, da wird sich ein Idealismus entwickeln, den
man einen positiven nennen könnte; er drängt zu einer Metaphysik,
jener
die
von
schieden
einer
bloßen
Begriffskonstruktion
bleibt.
Von
aus würde
der Wirklichkeit
eindringt
darf
der
als
friedlich
sich
ma^cht solche
unterals
den Tatbestand
in
schwer genug mit ihren
gleichwertige Möglichkeiten
als
miteinander
Ausdruck
volle
genug
tief
nicht
es
können nicht
damit entstehen,
und
und
deutlichste
reiner Idealismus
Die Haupttypen des philosophischen Denkens,
Widerständen nimmt.
gelten
der nicht
erscheinen,
ein abstrakter
die
hier
aufs
jener
vertragen,
der Wahrheit
sondern nur eine
gelten;
aber zugleich
Verbindung mit dem Leben augenscheinlich, daß
die
Entscheidung des Menschen wesentlich durch seine eigne Lage und
Erfahrung wie
auch
durch die Arbeit und
weiligen Zeit bestimmt sein wird,
die
Stimmung der je-
und daß wir daher, so gewiß nur
eine einzige Wahrheit besteht, uns schwerlich je einträchtig bei ihr
zusammenfinden werden.
Wir brauchen dabei nicht zu fürchten, daß solche engere Verknüpfung der Philosophie mit dem Ganzen des Lebens sie dem
Wechsel und Wandel geschichtlicher Lagen preisgebe und einem
zerstörenden Relativismus überliefere. Denn das würde nur geschehen,
wenn das
Leben
geistige
gesellschaftlichen
Menschen wäre.
lediglich
Erzeugnis des geschichtlich-
ein
Daseins und zugleich eine Erscheinung
In
Wahrheit
ist
am bloßen
geschichtlich-gesellschaftliche
alle
Geistigkeit nur die Entfaltung eines zeitlosen, allem bloßmenschlichen
Dasein
überlegenen
Geisteslebens;
und
nur echter
Art,
ist
teilhat.
die Kultur
soweit sie an
nur eine Seele
Es wirkt etwas Überzeitliches in jeder großen geschichtlichen
Erscheinung, etwas Übermenschliches
des Menschen.
in
jedem geistigen Aufschwung
Dies Überzeitliche und Übermenschliche, mit Einem
Worte das Absolute, herauszuarbeiten, dazu
sonders
hat
einem solchen Geistesleben
berufen.
Gesichtskreises, sie
Denn
nicht
kann
kraft
nur hat
sie
ist
die Philosophie be-
die
größte Weite des
der Freiheit des Denkens
zu den ursprünglichen Tatsachen und
am ehesten
zugleich zu einer Betrachtung
Zum Erkenntnisproblem.
104
sub specie aeterni vordringen;
sie
kann
einer durchgreifenden
in
Umkehrung unser Leben dem bloßen Strom der Dinge entreißen
und
es bei sich selbst befestigen, sie
Leistungen
liegenden
indem
üben,
kann eine Kritik an
sie
dieselben
allen vor-
auf den
Grund-
prozeß und die inneren Notwendigkeiten zurückführt und daran mißt,
sowie ihnen von da aus neue Aufgaben vorhält.
Mit solcher
Um-
kehrung der ersten Lage bringt die Philosophie nur eine durchgängige Notwendigkeit des Geisteslebens zum Ausdruck, und dient sie
seiner
Erhebung zur
Schon
in
vollen
Selbständigkeit
und Ursprünglichkeit.
dem Streben danach liegt eine Wendung des Lebens und
eine Befreiung, es verändert den Anblick des Lebens wie der ganzen
Wendung bedeutend
Wirklichkeit.
Das
allein
machen, daß
ihre
Vorhaltung absoluter Forderungen die Gering-
genügt,
zum Bewußtsein
fügigkeit unseres Besitzes
Tiefen ahnen
läßt.
c)
diese
bringt
zu
und uns weitere
^
Die Wendung zur Metaphysik.
Die Philosophie, so sahen wir, gewinnt eine eigentümliche Auf-
Erhebung über die Erfahrungswelt,
gabe nur
in
nicht erst
nachträglich,
ihre Arbeit
sich aber
^
sie
enthält sie
sie
von vornherein.
kommt dazu
Daher hat
von Haus aus eine beträchtliche Spannung; diese steigert
zu einem schroffen Widerspruch durch die besonderen Er-
Es sei hiezü eine Stelle des tiefsinnigen Steffensen angeführt, obschon
Gedankengang sich nicht völlig mit dem unsrigen deckt. Er sagt (siehe
Gesammelte Vorträge und Aufsätze S. 6): »Nicht aus sich selber, aus ihren
Werken oder der besonderen Macht oder Reinheit ihrer Leidenschaft schöpft
sie (d. h. die Philosophie) ihren Ruhm, sondern aus der lichten Höhe, in
welcher der Gegenstand schwebt, dem sie sich ergeben hat und dessen Mitteilungen sie empfängt. Darum mag sie ohne Gefahr ihre eigene Ohnmacht
bekennen und dann und wann auf eine Weile verstummen und sehr unscheinbar einhergehen; ihr altehrwürdiges Dasein bezeugt doch den Menschen
das Hereinleuchten einer vollkommenen Erkenntnis in den veränderlichen
Schein dieser Welt und unserer alltäglichen Denkart.
Wie die irdischen
sein
Entfernungen zu der Tiefe des Fixsternhimmels, so verhalten sich die Begriffe und Maße der empirischen
Wissenschaft zu dem Erkennen, nach welchem
die gewaltigsten Überzeugungen der gemeinen Ansicht, wenn sie dieselben mit der Gewißheit vergleicht, von deren
Ahnung sie ausgeht, erscheinen ihr nur als schwankende Meinungen des
die Philosophie emporstrebt,
Augenblicks.
auftut,
und
Ein Standpunkt, vor dem ein so unermeßlicher Horizont sich
wird seine Selbständigkeit und Unabhängigkeit schon zu behaupten
wissen."
Denken und Erfahrung.
Die Art, wie hier das Geistes-
fahrungen des menschlichen Kreises.
leben
durchaus
widerspricht
vorliegt,
Widerspruch
deutlich
erkennt,
ausweichen,
entweder
das
Welt zu
eignen
der
Entscheidung
aufzugeben
Geistesleben
oder
im
es
und zum Träger
behaupten
Das Geistesleben kann
machen.
nicht
die
Wirk-
beherrschen und an sich ziehen ohne eine volle Selb-
lichkeit nicht
ständigkeit,
Wesen; wer diesen
seinem
kann
der
Gegensatz zur vorliegenden Welt zu
einer
105
in
unserem Kreise aber
bloße Begleiterscheinung,
bildet es
von der Natur aus eine
vom geschichtlichen Dasein aus ein nachZusammenseins; das Geistes-
trägliches Erzeugnis des menschlichen
vom Ganzen zum Einzelnen, im nächsten Dasein ist alle
Verbindung eine Zusammensetzung aus einzelnen Elementen; zum
leben geht
und Ursprünglichkeit, das Dasein
Geistesleben gehört Selbsttätigkeit
Verkettung und damit eine Gebundenheit
durchgängige
zeigt
eine
alles
Wirkens; das Geistesleben gibt seine Wahrheit als zeitüberlegen,
das menschliche Leben verläuft in der Zeit und
Nun
folgen.
kann das Geistesleben
Welt-
als
ohne auch einen eigentümlichen Weltanblick zu
wirken,
kraft
muß ihrem Wandel
uns unmöglich
bei
er-
zeugen; müssen wir also auf einem solchen bestehen und begegnen
wir dabei einem durchgängigen Widerstände der nächsten Welt, so
ist
Sache im
die
Gegensatz
Steigerung
der
Spekulation
zur Metaphysik,
istischen
stellt,
zu
durchzuführen.
ihr
zur
Weltüberlegenheit
Wie
wird
Gegensätzlichkeit
überhaupt
diese
die
charakter-
die
Züge der Philosophie kräftiger ausprägt und klarer heraus-
so wird sie namentlich die Umkehrung des Weltbildes verstärken,
welche
in
jener
liegt;
sie
wird zugleich erkennen lassen,
gegebene Welt sich nicht völlig
Seins verwandeln
läßt,
sie
dagegen Widerstand
leistet;
und
harte
aber
muß
Kämpfe erzeugen.
Ja in
das Gesamtbild unserer Welt
schwere
ein geschichtliches Element; nichts
physik
als ein
zu lassen.
In
solches,
wenn auch
ist
nicht
die
kommt damit
charakteristischer für die Meta-
so doch ahnen
dem allen wachsen die Probleme, es wächst der Ab-
das Unternehmen müßte
des Lebens.
Verwicklungen
nicht erkennen,
stand zwischen den Zielen der Sache
stünde
daß die
eine Entfaltung eines geistigen
in
sondern daß
Widerstand
dieser
das
Mit solcher
hinter
In
als
ein
und den Mitteln des Menschen,
vermessenes Wagnis erscheinen,
der Metaphysik des Denkens eine
Wahrheit
trägt
alles
Leben
das
Metaphysik zur deutlichen Aussprache bringt.
echte Geistesleben
entwickelt
sich
bei der
Metaphysik
Problem
in
sich,
Denn alles
Menschheit nicht nur
in
Zum Erkenntnisproblem.
106
sondern
einer Überlegenheit,
Welt; die Moral
z.
Selbsterhaltung,
sondern
ernem
B.
Weltgetriebe
in
Reich
Das
sie
möglich
das
uns
läßt
muß
sie
im direkten Gegensatz zu
sich
und
Interessen
bereitet,
sie
diesen
sich
kleinlicher
eigner
ein
Zu
ohne Zweifel,
Weltanblick.
aber nicht wir
Un-
Verwicklungen durch einen Rückzug auf
Phänomen
und etwa die
entfliehen
Denn nicht
moralische Persönlichkeit als ein sicherer Halt ergreifen.
nur
Zwecke
sind uns zugefallen und auferlegt.
moralische
unmittelbare
auch
gehört
aber
Verwicklungen,
schwere
ergibt
zur nächsten
hartem Kampf dagegen ihr Reich erbauen.
behaupten und
haben
einem Widerspruch
in
nicht nur etwas anderes als die natürliche
selbstischer
einem
solchen
ist
der ihr notwendigen Einheit des
steht solche Persönlichkeit mit
Lebens und Ursprünglichkeit des Handelns in schroffem Widerspruch
zu
dem Nebeneinander und
sie
enthält
unmittelbar
der Gebundenheit der nächsten Welt,
Weltbehauptung,
eine
sie hat damit selbst
Dieser Weltcharakter aber wird
einen Weltcharakter.
dem Menschen
nur mit Hülfe eines Bildes der Wirk-
zur eindringlichen Gegenwart
lichkeit;
Behauptung von
die
der Gegenwart einer neuen Ordnung der Dinge,
so treibt eben die Selbsterhaltung der Persönlichkeit zu einer
Demnach wird
Metaphysik.
um die Aufrecht-
der Metaphysik
in
erhaltung einer selbständigen Philosophie gekämpft, die Philosophie
zusammen ohne den Fortgang zu jener. Die Leugnung der
Metaphysik bekundet entweder, daß die Bewegung zur Philosophie
bricht
nicht Kraft
um gegenüber den Widerständen der nächsten
Weg zu verfolgen, oder daß ein flacher Optimismus
genug hat,
Welt ihren
die Widerstände unterschätzen läßt.
Mit der Aufgabe wachsen
immer der Aufbau
zu
überwinden
Aufschwung
Charakter,
die
das
wird
sich
hier
bekommt
Festhalten
lassen,
überschreitet,
Aber wenn damit
auf
sondern
die
in allem,
noch
jetzt
sich
Hemmungen
einen
heroischen
hier
nie rein
das
in
was den andeutenden
sein.
noch weiteren Spielraum gewinnt,
Ganze ein bloßes Bild; durch alle Unzuläng-
der Darstellung können Notwendigkeiten wirken,
angesehen,
Der
steigern.
Hülfe von Bildern angewiesen
die Phantasie
nie wird deshalb das
lichkeit
das
Gedankenforderungen werden
Begriffe umsetzen
Umriss
auch die Widerstände; was
einer selbständigen Philosophie an
hatte,
wie
freilich
Ursprünglichste
und Gewisseste
die,
unseres
geistig
ganzen
Lebens sind; gerade jene Unzulänglichkeit der Darstellung kann die
Gewißheit der Grundtatsache
nur
noch
stärker
empfinden
lassen.
Denken und Erfahrung.
des
Alle Tiefe
was
in
Lebens hat einen symbolischen Charakter;
geistigen
und von
ihr ursprünglich aufsteigt
hier aus die ganze Wirk-
Formen nur
geht in die menschlichen und seelischen
lichkeit trägt,
unvollkommen
die Leistung dieser hat nur in der Zurückbezieh-
ein;
ung auf jenen Grund und
Wahrheit;
107
sie
der Durchleuchtung von daher eine
in
der Unwahrheit, sobald
verfällt
sie
davon ablöst
sich
So geschieht es vor allem
und mehr als ein bloßes Mittel sein will.
der Religion, die zur bloßen Mythologie zu sinken droht,
in
ihre
Formen
und
Begriffe
den
auf
unablässig
nicht
wenn
geistigen
Orundprozeß zurückbezogen und von da aus beseelt werden. Auch
auf der Höhe der Kunst waltete oft die Empfindung, daß das
Schaffen
durch
alles
das
wohl
anregen und
sich
drücken
lasse.
;,lch
hindurch
Darstellen
habe
etwas
erweise,
Tieferes
angemessen aus-
nicht aber
beleben,
mein Wirken und Leisten immer nur
all
symbolisch angesehen, und es ist mir im Grunde ziemlich gleichgiltig
gewesen, ob ich Töpfe machte oder Schüsseln", so bekennt es von
Überall der Wider-
sich, ein
Goethe (Gespräche mit Eckermann).
spruch,
daß der Lebensprozeß im innersten Kern über das Bloß-
menschliche hinaus zu selbständiger Geistigkeit und absoluter Wahrheit
gehoben
wird,
menschlichen Art
Aufforderung, durch
alle
führung zu behaupten.
für allen Zweifel
klug
wie
der
überall damit die
Unzulänglichkeit hindurch das Notwendige
Aber
gegen
hier
Verwicklung der Aus-
alle
bequemer Angriffspunkt
ein
ist
und Kleinglauben, nirgends mehr als
hier werden
lange die Sache von draußen her kühl
sich die Geister scheiden: so
und
Entfaltung die Schranken
seine
überwinden vermag,
zu
Grundtatsache
die
festzuhalten,
und daß
nicht
etwas fremdes
hat der Zweifel ge-
betrachtet wird,
Überwindung wird nur möglich, wo die Aufgabe als der Kern des eignen Lebens ergriffen und als eine Sache
wonnenes Spiel;
seine
Eine Vermittlung gibt
der geistigen Selbsterhaltung betrieben wird.
es zwischen
solchem Entweder
— Oder
nicht.
Wenn demnach die Metaphysik nur das Geschick alles Geisteslebens
teilt,
Aufgabe
das selbständig sein
darin,
hervorzukehren,
gültigkeit
will,
den Widerspruch
damit
aufzurütteln
trieb einzupflanzen.
hat
sie
eine
Klarheit
aller
aber
das
Leben
und
ihm
einen
Denn indem
so
mit voller
aus
zwingenden
besondere
und Schärfe
trägen
ihm gegenüber
und
eine
dann
aber
Fort-
das zur geistigen Selbsterhaltung
Notwendige dem Durchschnitt entwunden,
durchgebildet,
Gleich-
inneren
ihm
als
nicht
befestigt
abzulehnende
Zum Erkenntnisproblem.
108
Aufgabe entgegengehalten wird, kommt eine Unzufriedenheit, eine
Bewegung
Unruhe,
eine
treibt es
zu einer Emporarbeitung und weist zugleich durch
innere
dem Streben bestimmte Bahnen.
Ziele
sammenhange unserer Betrachtung
was
gelten,
in
Leben;
das
in
die
dem Zu-
Metaphysik nicht
als
sie
den Bewegungen dieser Arbeit aufs engste verflochten
bedeutende Kultur hat ihre eigne Metaphysik,
und Wesen
aus,
etwas
den Bestrebungen und Erfahr-
farbloser Gestalt über
Wollen
jene
Dabei kann nach
ungen der weltgeschichtlichen Arbeit schwebt, sondern
innerstes
Zerlegung
jene
jede
sie spricht darin
sich
will
sie
wird mit
sein,
selbst
hier
ihr
einen
wesenhaften Charakter und zugleich eine lebendige Seele erringen,
wird
in
greifen,
seits
ihr sich selbst zum Ideal.
Die Metaphysik muß einerseits er-
was als vorwaltende Kraft eine Kultur durchdringt, anderer-
hebt sie das Ergriffene über
Standes hinaus zu
öffnet
sie
von
sie
hier
Bloßmenschliche,
alle
Schranken des vorgefundenen
und absoluter Giltigkeit,
vollendeter Gestalt
aus
Kampf gegen
einen
alles
Unzulängliche,
Niedere der üblichen Lebensführung,
eine energische Scheidung in ein Für oder Wider.
er-
bewirkt sie
So vollzog die
platonische Ideenlehre eine Erhebung der künstlerischen Weltanschau-
ung des Griechentums ins Metaphysische, wobei
wandelbaren Ewigkeit voransteht, so
Aufklärung
eine
erhielt
und
die Gedankenwelt der
im System Leibnizens mit
metaphysische Gestalt
seiner Unendlichkeit des Kleinen
die Idee einer un-
seiner Verwandlung der Philo-
sophie in eine universale Mathematik.
Überall ein Bestreben,
vom
Gipfelpunkt menschlicher Leistung ins Absolute vorzudringen und in
Umkehrung der
nächsten
Lage
solche
sie
ist
ein
Herausarbeiten
dessen,
was von den be-
sonderen Zeitlagen aus an zeitloser Wahrheit erreichbar war;
sich dabei ergab, das versank nicht mit der Zeit,
wenigstens
Aus
als
Möglichkeit und Aufforderung,
solcher Fassung
Schon der
Name
konnte
was
sondern es bleibt,
stets
gegenwärtig.
der Metaphysik läßt sich ganz wohl den
Angriffen begegnen, mit denen
*
eine
Die Metaphysik wird durch
Beziehung zur Geschichte keine Hingebung an die bloße
sondern
Zeit,
Denken und Sein
unserem
Selbständigkeit geistiger Art zu erringen.
sie
von
alters
leicht Vorurteile
her zu kämpfen hatte.
erwecken,^ aber auch in
Der Ausdruck Metaphysik entstand dadurch, daß Andronikus Rhodius,
in seiner Anordnung der aristotelischen Schriften
ein Zeitgenosse Ciceros,
die Untersuchungen über die „erste Philosophie" (rptÖTT) cp(Xoao9(a) hinter die
Physik stellte:
[jL£Ta
xa tpuatzä (näheres s. Bonitz,
Kommentar zur aristotelischen
Denken und Erfahrung.
der Sache wird
die Metaphysik einen anderen
jetzt
müssen, als ihn frühere Zeiten versuchten.
Bruch
mit
Weg einschlagen
Es gilt einen entschiedenen
freischwebenden Spekulation,
jener
109
welche aus bloßem
Denken glaubte, eine neue Welt erzeugen zu können.
welche im Wissen
jener älteren Denkweise,
gehalt
des
Lebens glaubte
und
ermitteln
Das entspricht
den gesamten Geistesihn
von
hier aus
den
übrigen Gebieten zuführen zu können, während wir jetzt das Wissen
ein
in
tiefer
gegründetes Geistesleben
hineinstellen
und
es vereint
um die Entfaltung dieses Lebens und zu-
mit den übrigen Gebieten
um Wahrheit kämpfen lassen.
Besonders aber bildet die neue
Metaphysik den schärfsten Gegensatz
zum ontologischen und damit
gleich
und dogmatischen Charakter der
Meta-
zugleich
abstrakten
physik.
Die Art, wie Aristoteles die Aufgabe der »ersten Philosophie"
dahin bestimmt
6v),
hat,
älteren
das Seiende als Seiendes zu betrachten (to ov
Eigenschaften
die allgemeinsten
des Seins zu ermitteln,
hat
•jl
von vornherein
die Sache
in
eine
erschienen
damit
der Dinge
und wurden zum Hauptgerüst,
als
eine
gewisse
Bahn
schiefe
formale
gelenkt.
Beschaffenheiten
als
Denn es
das Wesen
in das alle Besonderheit
bloße Ausführung einzutragen war; die Metaphysik wurde
Metaphysik,
S. 3 ff.).
Daraus wurde schon
eine Bezeichnung der Disziplin selbst (xä
-paYp.aT£ta).
im
[ista
ersten Jahrhundert n. Chr.
Ta
tpnaixä,
tj
puta xa «puaixi
Die Singularform metaphysica gehört der Scholastik an, sie dürfte
aus der Übersetzung des Averroes stammen.
Der Name war insofern nicht
von Anfang an den Begriff mit der Vorstellung behaftete,
als habe die Metaphysik mit etwas Femliegendem, Jenseitigem, zur nächsten
Wirklichkeit erst Hinzugedachtem zu tun.
So heißt es schon bei dem Neuplatoniker Herennius (s. Brandis, Abh. der Berl. Akad. 1831, p. 80): {j-et« xa
glücklich,
cpuatxa
als er
XEyovTat,
ä~tp
cpuasoj;
Ü7:£pf,pTai
xai jjTup
atriav
za\ Xdyov slaiv.
Auch
den Scholastikern, z. B. Thomas, gilt die metaphysica soviel als transphysica.
Kant aber sagt (VIII, 576 Hart.): „Der alte Name dieser Wissenschaft [xera
Ta »uitxa gibt schon eine Anzeige auf die Gattung von Erkenntnis, worauf
die Absicht mit derselben gerichtet war.
Man will vermittelst ihrer über alle
Gegenstände möglicher Erfahrung (trans physicam) hinausgehen, um womöglich das zu erkennen, was schlechterdings kein Gegenstand derselben sein
kann." Die Freunde der Metaphysik strebten dem gegenüber nach neuen
Bezeichnungen.
Clauberg, der bedeutendste Cartesianer Deutschlands,
emp-
Ontosophie oder Ontologie, aber bald übertrug sich die Ungunst auf
das neue Wort; schon Wolff klagte (s. Philos. prima sive ontologia 1): vix
aliud hodie contemtius est nomen quam Ontologiae. Auch würde Ontologie
fahl
nur die ältere Art der Metaphysik bezeichnet haben, die uns heute unmöglich
dünkt.
Ist
es übrigens nicht bemerkenswert, daß nie ein Denker ersten
Ranges eine „Metaphysik" unter diesem Namen geschrieben hat?
Zum Erkenntnisproblem.
110
Das ergab eine Verschiebung der Ge-
damit zur bloßen Ontologie.
dankenwelt ins Abstrakte und Formale, eine Zurückstellung des eigentümlichen Inhalts des Menschenlebens.
Zugleich aber ergab es einen
Dogmatismus, indem jene formalen Eigenschaften vor aller näheren
Erfahrung und unabhängig von
erkennbar dünkten und daher von der Metaphysik den
allemal
für
Bewegung ein-
aller geschichtlichen
übrigen
Wissensgebieten
wurden.
Ein
solches
als
Wahrheiten
unantastbare
dogmatisches Verfahren
nahm
zugeführt
zugleich
der
Metaphysik eine innere Bewegung und den anderen Wissenschaften
ihre Selbständigkeit.
Kein Wunder, daß jene ontologische und dog-
matische Metaphysik von den verschiedensten Seiten her Widerstand
Entwicklung der modernen Forschung
die
fand;
ist
möglich
nur
geworden unter Abschüttelung der alten Metaphysik.
Aber
ist
die
Abweisung
ein Verzicht
nicht
einer
auf alle
besonderen
Metaphysik
der
Art
und jede Metaphysik, vielmehr wird
Kant wohl Recht behalten mit der Überzeugung »irgend eme Metaphysik
immer in der Welt gewesen und wird auch wohl ferner -
ist
anzutreffen
darin
sein"
(Hart.
111,
25).
wird
Jedenfalls
Meta-
die
physik,
welche sich aus den Zusammenhängen unserer Betrachtung
ergibt,
nicht
von den Vorwürfen
getroffen,
welche die Zerstörung
wo das Erkennen selbst
eine Entfaltung des Lebens in sich trägt, und wo es vor allem auf
der alten Metaphysik bewirkten.
die
Denn
hier,
Durchleuchtung und Vertiefung dieses Lebens geht,
Metaphysii^
das
Denken
und
Leben
nicht
ins
wird die
Abstrakte
locken,
sondern es seiner eignen Tatsächlichkeit und Bestimmtheit zuführen,
sie
wird mit ihrem Zusammenschluß aller Mannigfaltigkeit die einzig-
artige Individualität
Klarheit
unseres Seins und unserer Welt erst mit voller
herausstellen.
Alle einzelnen
gesamte Gebiete wie Religion,
Kunst,
Größen und Aufgaben, auch
Moral, werden die klägliche
Farblosigkeit der üblichen Fassung erst überwinden,
wenn sie inner-
halb eines umfassenden
Lebenszusammenhanges einen
und
erhalten,
ein
bestimmtes Ziel
auch wird nur der
die Wirklichkeit in jener Verdichtung
festen
Platz
Inhalt,
den
zum Ganzen erschließt, über
das Recht und die Bedeutung der Formen des Seins entscheiden.
So
ist
es nicht eine Lust an allgemeinen Formeln, sondern das Ver-
langen nach mehr Charakter, nach ursprünglicher Tatsächlichkeit, nach
energischer Durchbildung unseres Lebenskreises, was die Forschung
zur Metaphysik
treibt.
Ebenso sicher ist eine Metaphysik, welche den Zusammenhang
Denken und Erfahrung.
Hl
des Erkenntnisstrebens mit einem begründenden und umfassenden
dem Vorwurf eines erstarrenden Dogmatis-
vor
Geistesleben wahrt,
Eine solche Metaphysik wird mit der weltgeschichtlichen Be-
mus.
wegung
nehmen und zugleich selbst eine Ge-
engste Fühlung
die
ohne damit zur bloßen Zeitgeschichte zu sinken.
schichte gewinnen,
Wir haben heute keine Metaphysik, und es gibt nicht wenige,
die das für einen Gewinn erachten.
nur,
wenn
stande
sich
befände,
Aber ein Recht darauf hätten sie
unsere Gedankenwelt
einem vortrefflichen Zu-
in
wenn auch ohne Metaphysik
unser Leben und Streben beherrschten,
hielten
und vom kleinmenschlichen Getriebe
heit
eine grenzenlose Zersplitterung,
in
ist
allem
Überzeugungen
In
befreiten.
Wahr-
eine klägliche Unsicherheit
Überzeugungen,
der
Prinzipiellen
feste
hohe Ziele uns zusammen-
Ohnmacht gegen
eine
das Kleinmenschliche, eine Seelenlosigkeit in überströmender äußerer
Wer das ruhig zu ertragen vermag, den
Fülle nicht zu verkennen.
werden
theoretischen Erörterungen nicht zur Metaphysik führen;
alle
wer 'aber eine zwingende Aufgabe darin
Kulturleben
und
sich
zu
erkennt,
daß auch unser
einem charaktervollen Ganzen zusammenfasse
gewinne zu engerer Verbindung wie
eine innere Selbständigkeit
zu schärferer Scheidung der Geister, der wird mit uns an der Metaphysik festhalten und für die
d)
alte
Aufgabe neue Bahnen suchen.
Der Gesamtanblick der menschlichen Erkenntnisarbeit.
Die bisherigen Untersuchungen enthalten prinzipielle Überzeug-
um
ungen vom Erkennen, die nur entwickelt zu werden brauchen,
ein eigentümliches Gesamtbild zu
die
Fassung des
verfochtene
hier
schichtlich
Geistesleben
gilt
uns zugleich
unmittelbaren
Stufen
welche
ist
den
es
ge-
befreien
und
strebens
vollauf
steigern,
was
neue Stufe der Wirklichkeit
als eine
Von
ursprüngliche Tiefe gegenüber
eine
als
Seelenleben,
zusammentreffen.
dem
in
hier
Entfaltungen
aller
die Bedingtheit
unseres
anzuerkennen;
ja
es
beiden
der
aus wird es möglich,
zugleich
Abhängigkeit nach außen
menschlichen
mag nun
sich sonst feindselig gegenüberstand
wendig Abbruch
Das
besonderen
Geisteslebens,
den Kern der Erkenntnisarbeit von
zu
Im
überkommenen Gegensatz zu überwinden verspricht. Das
gegenüber der Natur, und
dem
ergeben.
sich
Erkenntnisgegenseitig
und einander not-
tat.
Geistesleben
galt
uns
als
volltätiges
Leben,
das
nicht
Zum Erkenntnisproblem.
112
zwischen Subjekt und Objekt verläuft, sondern den Gegensatz von
Haus aus umspannt; die Aufgabe kann
Seins,
sondern nur
in
muß dann
in sich selbst
verschiedene Stufen der Aus-
eines jenseitigen
Es
liegen.
hier nicht in der Abbildung
der eigenen Durchbildung
prägung enthalten, die Bewegung zur höheren Stufe aber durch eine
Notwendigkeit
Ganzen gefordert werden; was
des
das wird
irgend schon angehört,
der Verwandlung
So
steht
es
erst
mit
beim Er-
auch
Bewegung liegt innerhalb des gesamten Lebens.
kennen, auch seine
Denn auch
Tätigkeit
zu seinem vollen Besitz
Selbsttätigkeit.
in
seiner
der Vorwurf,
dem es zu tun hat, muß sich inner-
mit
halb, nicht jenseit des Geisteslebens befinden; ein gänzlich
draußen
Gelegenes könnte nichts erregen und nichts bewegen, es würde das
Denken gar nicht berühren und könnte ihm nun und nimmer auch
nur zum Probleme werden.
wenn
Dies geschieht nur dann,
ein
Gegenstand der Gedankenwelt irgend schon gegenwärtig ist, die Art
wie er
aber,
entspricht,
ja
zwingender
gegenwärtig
Antrieb
zur
der Natur
ist,
widerspricht;
dieser
Weiterbildung.
behauptung, die das Geistesleben
Vordringen zur höheren Stufe
in
des
Geisteslebens
Widerspruch
So
ist
eine
es
nicht
dann
wird
ein
Selbst-
der Erkenntnisarbeit mit ihrem
vollzieht.
kann zum Erkenntnisproblem nur
dem Lebensprozeß irgend schon einverleibt ist, so muß
eine innere Erweiterung des Lebens vorangehen, wo das Erkennen
in Fluß kommen soll.
Die weltgeschichtliche wie die alltägliche
Steht die Sache derart,
so
werden, was
Erfahrung
bestätigt
diese
Behauptung mit überzeugender
Klarheit.
Denn sie zeigt, daß auch solches, was den Menschen mit eindringlicher Nähe umfängt und mit den stärksten sinnlichen Wirkungen
berührt,
ihm
innerlich
völlig
fremd bleiben kann und seinem Er-
kennen nicht zum Probleme wird.
der
an
keiten,
sie
Fragen
stellt;
Nur dem antworten die Dinge,
nur dem erschließen sich Wirklich-
der ihnen Möglichkeiten entgegenbringt;
auch
der härteste
Widerstand übt eine seelische Wirkung nur nach Verwandlung
innere
eine
Hemmung;
Völker und Zeiten
treffen,
welcher Gegenwehr
mitgebracht
fertig
verfechten
^
zu
schwerste
können
Mißstände
in
Individuen,
ohne sie stark zu erregen und zu irgendtreiben.
werden,
übereinstimmend
Daß
sondern
große
die geistigen
erst
zu
Künstler
Organe
bilden
und große
sind,
nicht
das
Erzieher. ^
Bekannt ist Herbarts Auslassung über den neunzigjährigen Dorfschul-
meister (Werke
X,
8):
„Wollten wir nur sämtlich
bedenken,
daß jeder
Denken und Erfahrung.
Auch
ja
113
die weltgeschichtliche Betrachtung zeigt, wie spät ganz Nahes,
äußerlich schon Angehöriges zu eignem Leben geworden
das eigne Streben bewegt hat;
und
Wie langsam
dünkt.
deckung der Natur,
des Sehens zu
erfolgte
erschloß
spät
die Kunst
verfeinern,
künstlerische
die
sich
z.
der Gegenwart,
um dann
Ent-
der
die Gestalt
B.
zuerst
den Dingen mehr
in
und neue Seiten an ihnen zu eröffnen! Auch sich
entdecken
das Menschsein und was daraus an Gemeinschaft, des Lebens
selbst,
und Empfindens
fand
wie
wie müht sich auch
Landschaft,
die Art
zu
erkennen, was später leicht selbst-
die Vorbereitungen dessen
verständlich
und
ist
läßt zugleich die Voraussetzungen
sie
es
nicht
fließt,
vor,
fertig
der Mensch erst entdecken müssen,
hat
innerer Art
Fortbildungen
errang
er
Die
er
Bewegungen und
durch
es
Erziehungslehre
spricht
von der
stellungskreis
Aufnahme neuer Eindrücke in den Vordes Individuums; nun wohl, es gibt auch eine welt-
geschichtliche
Apperzeption, auch das Ganze der Menschheit
Apperzeption
als
einer
mag, nichts aufzunehmen, dem
innere
es nicht eine
ver-
Bewegung ent-
gegenbringt.
Was so im Einzelnen bereitwillig anerkannt wird, das muß, ins
Ganze gehoben, das Erkenntnisproblem
rücken.
Denn damit wird
klar,
Menschen
liegt,
daß
und
in
alle
eine
neue Beleuchtung
Erkenntnis innerhalb der
daß
Arbeitswelt
des
Fortschritte
des Erkennens gibt ohne ein V/achstum dieser Arbeits-
Wahrhaft große
welt.
solche,
die
nicht
den Lebenskreis
Leistungen
auch
des
es
keine
wesentlichen
Erkennens sind
einem vorgefundenen Kreise angehören,
selbst verändern.
nur
sondern
Die moderne Wissenschaft wäre
unmöglich gewesen ohne den modernen Menschen mit seiner kühnen
Erhebung über die Welt und
seiner Befestigung in der eignen Seele.
Solche Zurückverlegung gestattet
erst,
den Erkenntnisprozeß
als ein
immanentes Vorgehen zu fassen und jenes Dilemma zu vermeiden,
daß das Denken entweder mit einem fremden Sein zu tun habe
oder aus sich selbst
Aber eben
alles Sein
diese
hervorspinnen müsse.
Anerkennung der Selbständigkeit des
Geistes-
nur erfährt, was er versucht! Ein neunzigjähriger Dorfschulmeister hat
die Erfahrung seines neunzigjährigen Schlendrians; er hat das Gefühl seiner
langen Mühe; aber hat er auch die Kritik seiner Leistungen und seiner
Methode?" Fröbel aber meint, daß der Mensch, ,,um die Natur zu verstehen, sie durch eine ihm eigentümliche Kunstweise gleichsam von Neuem
in und aus sich schaffen muß."
Eucken, Grundbegriffe.
4.
Aufl.
8
Zum Erkenntnisproblem.
114
und der Immanenz ^ des Erkenntnisprozesses ist geeignet, das
Eigentümliche der menschlichen Lage und zugleich die Bedeutung
lebens
der Erfahrung zur vollen Geltung zu bringen.
und
Je selbständiger
und mit ihm das Erkennen
überlegener nämlich das Geistesleben
gefaßt wird, desto größer wird der Abstand der nächsten Lage, desto
deutlicher erhellt,
daß nur unter gewissen Bedingungen und durch
harte Arbeit der
Mensch daran teilzunehmen vermag, daß auch das
Geistesleben ihm nur durch irgendwelche Erfahrung zugänglich wird.
Der Mensch wird zunächst von der untergeistigen
eingenommen,
lichkeit
mit
mechanischen
ihren
könnte diese Stufe
höhere
die
Höhere
es
hat
erfolgt
Stufe der Wirk-
die intellektuell in der sinnlichen Vorstellung
Verkettungen
keiner Weise
in
zum Ausdruck kommt;
Aber dieses
wirksam.
irgendwie
er
wäre nicht auch
überschreiten,
in
seinem
hat
für
den Lebensprozeß keine volle Gegenwart, sondern
eine
solche
Bereiche
zu
erst
gewinnen;
selbst
die
Anregung dazu
gewöhnlich nur unter besonderen Bedingungen, bei Verwick-
lungen und Widersprüchen der niederen Stufe; die Geschichte zeigt
wie mühevoll und langsam das Erkenntnisstreben
deutlich,
in
Fluß
Und eben im Vordringen mußte es auch eine eigentümliche
kam.
Lage beim Menschen anerkennen, die
sondern nur
als eine
sich
Tatsache hinnehmen
nicht begrifflich ableiten,
läßt.
Insofern trägt
das
menschliche Erkennen einen Erfahrungscharakter. Aber das anerkennen
heißt keineswegs
dem Empirismus huldigen. Denn jener Erfahrungs-
charakter
wäre nicht zu erkennen
selbst
gegen die bloße Erfahrung;
ohne eine Überlegenheit
die Schranken
des Menschen gelangen zur Einsicht nur,
ständigen
und überlegenen Geistesleben
seine Lage zu
und
die
Gebundenheit
sofern er an einem selbteil
hat
und von
hier aus
würdigen vermag.
Es hat aber die Erfahrung für das Erkennen eine zwiefache
Bedeutung:
sie
im Innern.
Jenes
ist
Begrenzung nach außen und sie ist Determination
ist
Bedingungen gebunden
tätigkeit
sie,
wenn
bleibt
die
geistige Tätigkeit
an jenseitige
und sich daher nicht zur vollen Selbst-
zu erheben vermag; dieses, wenn
sie die volle
Bestimmtheit
ihrer
eignen Art erst im Zusammenstoß mit Widerständen erreicht,
erst
durch Versuchen und Erfahren hindurch ihrer selber inne wird
und zu
reiner Selbsttätigkeit gelangt.
In beiderlei
Bedeutung
ist
das
Wir nehmen hier Immanenz in dem alten und ursprünglichen Sinne,
wonach es ein Geschehen bezeichnet, das innerhalb des Lebensprozesses ver*
bleibt, nicht
über ihn hinausgeht; siehe den Artikel Immanenz-Transzendenz.
Denken und Erfahrung.
115
Erkennen des Menschen auf Erfahrung angewiesen;
entbehrHch
sowohl
gebung
als
für
fang
für seinen Inhalt.
als
für
eigne Beschaffenheit, sowohl für seinen
seine
un-
sie ist hier
das Verhältnis des Geisteslebens zur
UmUm-
Was sich beim Menschen an Erkennen entwickelt, das befindet
zunächst gegenüber der Unermeßlichkeit einer fremden Welt,
sich
und das kommt nur
weiter in
aus der Welt lediglich
des Empfangenen
arbeitung
Berührung mit dieser Welt, das kann
empfangen scheinen.
zu
kann
namentlich die sinnliche Natur,
sich
es
nie
Auch
für weite
in
der Be-
wie
Gebiete,
davon ablösen; was daraus
die Gedankenwelt des Menschen eingeht,
ist
nicht rein in
in
Gedanken-
größen umzusetzen, es behält eine Bindung an etwas Jenseitiges und
zugleich
eine
Undurchsichtigkeit.
mit den Dingen
Aber mag
und eine Beziehung auf
die
hier
eine
Berührung
Dinge noch so not-
wendig sein, diese Berührung und Beziehung erzeugt
nicht das Er-
kennen; es entwickelt sich unter Bedingungen und Beschränkungen,
aber es bleibt auch dabei vornehmlich ein
Werk des Geisteslebens;
es entwickelt sich nicht aus der Erfahrung,
sondern nur an der Er-
fahrung, wie Eindrücke nicht in die Gedankenwelt eingehen können,
ohne eine wesentliche Umwandlung zu
erleiden.
Wie grundver-
nimmt sich dasselbe Naturphänomen in der unmittelbaren
Empfindung des naiven Menschen und in der Gedankenwelt des
schieden
Mit Recht sagt Hegel.
Naturforschers aus!
»Die Natur des Geistes
— nicht ein anderes Ursprüngliches in sich aufzunehmen, oder
es
ist
eine Ursache sich in ihn kontinuieren zu lassen,
nicht
sondern
sie
abzubrechen und zu verwandeln" (Wke. IV, 229).
Aber
nicht
Geisteslebens
ist
Ausdehnung,
nur
die
für
uns Menschen eine Frage und eine Aufgabe.
auch
die
innere
Art
des
Weder erfüllt das Geistesleben
in fester
und klarer Gestah unmittel-
noch
zieht es
uns
bar unser eignes Leben,
schritt
hinein,
in einen sicheren
Fort-
wie der intellektuelle Optimismus der konstruktiven
Philosophie es annahm, sondern wir müssen von kleinen, nicht ein-
mal unbestreitbaren Anfängen allmählich vordringen, und wir finden
in
solchem Streben die mannigfachsten
Hemmungen und Gefahren,
wir unternehmen vieles in froher Zuversicht, was sich im Verlauf als
unmöglich
erweist,
hergeworfen.
wir
nicht
wir werden scheinbar oft im Zickzack hin- und
Was wir aber mit aller Mühsal erringen, das gewinnen
durch
kluge
Überlegung,
sondern durch ein zu Ende
Gehen eingeschlagener Richtungen; unseres Vermögens wie unserer
Zum Erkenntnisproblem.
116
Schranken werden wir inne erst durch Lebensentwicklung und Lebens-
Im besonderen
erfahrung.
erreicht unser
Leben nur durch Kampf
seine volle Tiefe, erst der Widerstand treibt es zur Aufbietung seiner
ganzen Kraft und
Wachstum der
zu voller Ursprünglichkeit.
Dabei bedeutet das
reinen Sieg
über das Feindliche
keinen
Geistigkeit
und bringt keine volle Aufhellung. Vielmehr mag die innere Steigerung
Probleme und Widerstände hervorrufen, und
neue Ansprüche,
sich
damit der Anblick der Wirklichkeit immer positiver, immer irrationaler
muß aus dem Erkennen etwas
Eine solche Tatsächlichkeit
gestalten.
wesentlich anderes machen, als es der Rationalismus wollte; Punkt für
Punkt wird
es hier auf die
gewiesen.
Einem
glatten
Erfahrungen des gesamten Lebens hin-
Abschluß nahe glaubte man
den ersten Anfängen, aus wachsender Einsicht
sondern
klarer,
rätselhafter
geworden.
in
immer mehr An-
ist
erkennung ungelöster Probleme hervorgegangen, die Welt
nicht
nur
sich
So
ist
ist
uns
eben auf der
Höhe des modernen Lebens der Gesamtanblick des Erkennens alles
Die Wirklichkeit erhebt sich vor unseren Augen
eher als einfach.
als ein Stufenreich mit einem Fortgang vom Unorganischen zum
Organischen, vom Unlebendigen zum Lebendigen und zur Seele,
von der naturgebundenen zur geisterfüllten Seele. Von jeder Stufe
aus ergibt sich ein eigentümlicher Durchblick der Wirklichkeit; der
Kampf wird
nicht enden,
ob die unterste oder die oberste Stufe den
Ausgangspunkt der Erklärung zu bilden habe.
Die Philosophie kann
nicht umhin, die Wirklichkeiten, welche sich auf der höchsten Lebens-
Offenbarungen zu behandeln und von
stufe eröffnen, als die tiefsten
ihnen
sie,
aus
das
Aber nun
des Ganzen zu entwerfen.
erfährt
daß die von dort entwickelten Größen für die Welt unter uns
daß diese ihnen eine starre Eigenart entgegensetzt, auch
nicht passen,
erfährt sie,
daß diese Welt durch
höhere Stufe
mit
Bild
seinen
als
Begriffen
ihr ganzes
Wirken und Sein jene
gleichgültige Nebensache
behandelt.
Weder
noch
scheint
unserer
eine
mit
seinen
Kräften
in
Welt sich durchsetzen zu können, was wir für den Kern aller Wirklichkeit
zu erachten nicht lassen können.
daß an
den Menschen
seine
geistige
Überall der Widerspruch,
Natur Anforderungen
stellt,
denen seine bloßmenschliche Art nicht zu entsprechen vermag; die
geistige
sein
Selbsterhaltung zwingt ihn,
intellektuelles Vermögen
Wahrheiten zu bejahen, denen
nicht voll gewachsen
ist,
ihren Grund-
gedanken energisch zu behaupten, ohne ihm eine angemessene Ausführung geben zu können.
So bewirkt
es
notwendig eine geistige
Denken und Erfahrung.
117
Verarmung, wenn das intellektuelle Vermögen über den Gesamtinhalt
des Lebens entscheiden
e)
soll.
Würdigung des Rationalismus und des Empirismus.
Nach diesen Erörterungen läßt sich eine schiedsrichterliche
Würdigung der streitenden Gegner unternehmen; sie wird zeigen,
daß jeder bedeutende Wahrheitsmomente
gegen
daß
den anderen kehrt,
vertritt
er aber ins
und
sie siegreich
Unrecht gerät und die
eigne Stellung nicht behaupten kann, sobald er bei sich selbst einen
Abschluß
letzten
sucht.
Der Rationalismus
des
Selbständigkeit
alle
Umgebung,
in
hat
seine
Geisteslebens
Stärke
und
der Verfechtung
in
der
Überlegenheit gegen
seiner
der Verfechtung der Überzeugung, daß das Leben
daß
erstwesentlich nicht von außen nach innen geht,
sich,
um mit
Plato zu reden, nicht einem Blinden von außen her Augen einsetzen'
Ohne diese Überzeugung gibt es überhaupt keine Wahrheit.
lassen.
Denn
der
die
Auslieferung unserer Erkenntnis an die Eindrücke
volle
Umgebung würde ihr alle Festigkeit, allen Zusammenhang, alle
innere Durchleuchtung rauben,
Individuen preisgeben.
würde sie der Zufälligkeit der bloßen
Es ist eine axiomatische Notwendigkeit, wenn
der Rationalismus dagegen ein a priori
Nur sei dabei das
verficht.
a priori nicht als eine in der Seele jedes Einzelnen fertig vorhandene
Größe, sondern es
sei
als
der Mensch
standen,
das
a priori
enthält
ein
Grundgesetz des Geisteslebens ver-
sich
erst
die Behauptung,
aneignen
muß.
Ein
solches
daß das Geistesleben Normen
in
sich trägt, die das Streben aus aller Irrung immer wieder zur Wahrheit
lenken; es enthält auch die Behauptung, daß das Geistesleben seinem
Wesen nach
übergeschichtlicher
Geschichte bildet.
Art
ist,
kein
bloßes Produkt der
Ohne solche Übergeschichtlichkeit könnte es nie die
geschichtlichen Bildungen einer überlegenen Kritik unterwerfen, sondern
würde
es
ganz und gar ihrem Wechsel
und Wandel
ausgeliefert.
Mit der Verfechtung so unerläßlicher Wahrheit hat der Rationalismus ein überlegenes Recht gegenüber
gerät ins Unrecht,
reichen
zu
oder
doch
indem
er
indem
können,
leicht
dem Empirismus.
er jene Wahrheiten
indem er das hohe
Ziel als eine gegenwärtige
zugängliche Tatsache behandelt.
Dies
ohne weiteres das menschliche Geistesleben
leben an sich,
als
Aber er
unmittelbar glaubt er-
absolutes Geistesleben behandelt
geschieht,
als
Geistes-
und damit den
Zum Erkenntnisproblem.
118
auch für die Schranken
Sinn für das Charakteristischmenschliche,
des Menschen abstumpft; es geschieht, indem er Leistungen, die das
Denken nur im Zusammenhange mit dem Ganzen eines selbständigen
ihm
vermag,
Geisteslebens aufzubringen
den Gedankengrößen
selber zuweist
nimmt;
ihre belebende Tiefe
und damit
es geschieht,
in-
dem er unser Geistesleben ohne weiteres auf sicherem Wege glaubt
und
innere Verwicklungen anerkennt.
keinerlei
Alles zusammen gibt dem Rationalismus die Neigung, das Dunkle
und Feindliche unserer Weltlage abzuschwächen und wegzuerklären, das
dem
Individuelle
— So
der
Inhalt
Form zu
opfern.
zu glatte, dünne, blutlose Wirklichkeit, das
viel
Denken wie das Leben
hafte.
den
Allgemeinen,
Daraus entsteht eine
Formale und Schatten-
gerät ins Abstrakte,
zeigt es besonders anschaulich das Bild der Geschichte,
Wendung zur spekulativen Begriffs-
das der Rationalismus
in
konstruktion
Die Bewegung scheint hier von vornherein
erzeugt.
der
im Element der Vernunft befindlich, da
hat;
dünken
hier
Gegensätze und Kämpfe nur ein Mittel zur
alle
Steigerung der Vernunft, und
in
eine große
nicht bloß
und
Mehr der Vernunft
jedes
auch die Unvernunft zu steigern
hältnissen
eine
alles Irrationale
scheint sich schließlich
Harmonie aufzulösen, während in Wahrheit der Kampf
innerhalb der Vernunft, sondern
wird,
führt
Wahrheit den Ver-
in
sie
und immer von neuem zu bestätigen
nunftcharakter erst zu erringen
mehr noch um sie gemenschlichen Ver-
in
pflegt;
hier
scheint die
Epoche auf der anderen sicher fortzubauen und, was an
welt-
geschichtlicher Erfahrung erwächst, dauernd gesichert zu sein, da in
Wahrheit der Kampf immer wieder
greift,
eine
feste
in
Grundlage immer neu zu erringen
der Mensch
als
ein
reines
ist,
zum Probleme wird;
Erfahrung geistiger Art immer neu
scheint
die letzten Elemente zurück-
Werkzeug der
und
alle
hier er-
geistigen Arbeit,
da vielmehr seine Neigung überwiegend dahin geht, das Geistesleben
der natürlichen und sozialen Selbsterhaltung unterzuordnen, es damit arg zu
verkehren und seinen eignen Zwecken zu entfremden.
Bei
Verkennung des Dunklen und Feindlichen
solcher
Geschichte
listische
ihre
Kraft
und
Tiefe;
je
ausschließlicher
geschichtliche
Leben
daß immerfort
des Ganzen
um
nicht
in
desto
mehr
entleert
und
klar,
daß
das
sicherem Fortgang aufbaut,
das Ganze gekämpft,
vollzogen
werden
die
rationa-
Wird dagegen
Behandlung durchgeführt wird,
verflüchtigt sie die Wirklichkeit.
verliert
jene
muß, so
sich
sondern
immerfort eine Bejahung
tritt
vor den Prozeß die
Denken und Erfahrung.
und
Tat,
freie
es
entfällt
alle
Möglichkeit
\\g
einer
Kon-
rationalen
struktion.
muß das Sichausleben des Rationalismus
So
in
der Richtung des Empirismus mit seinem
lichkeit
einen Rückschlag
Durst nach Tatsäch-
und seiner willigen Anerkennung der Schranken des Menschen
erzeugen, wie denn auch auf geschichtlichem Boden der Empirismus
namentlich da zu Macht und Ansehen gelangt
wo die Mängel
ist,
überkommenen Rationalismus augenscheinlich wurden.
eines
den Hintergrund
des
neuesten
Empirismus
Auch
der Widerwille
bildet
gegen die spekulative Begriffskonstruktion.
Aber der Empirismus bringt den Erfahrungscharakter unserer
Gedankenwelt keineswegs zu angemessenem Ausdruck, er
Prozeß der Erfahrung
keit,
er
faßt
ohne die es doch kein wissenschaftliches Erkennen gibt.
alles
Geistesleben
selbständige
Menschen aus eine
den
einem schroffen Gegensatz zur Selbsttätig-
in
Geistigkeit
leugnet,
muß
und zugleich
eine
Indem
vom bloßen
er
Erkenntnis zu
Dies aber, was in Wahrheit unmöglich ist,^ kann
entwickeln suchen.
einen leidlichen Schein des Gelingens nur erreichen, indem es versteckterweise eine geistige Welt voraussetzt und ihr entlehnte Größen
verwendet.
Damit aber ergibt
sich
schiefer Anblick der Wirklichkeit.
bis
ins
Einzelne
hinein
ein
Der Empirismus richtet beim Er-
kenntnisprozeß das Augenmerk gänzlich auf die Leistung und übersieht die geistige Tätigkeit,
und
vergißt,
daß
er für
die darin wirkt; er haftet am
Gegenstand
uns etwas nur durch unser Aneignen wird;
er sieht die Determination des Erkennens durch die Erfahrung, aber
er sieht nicht,
daß diese Determination innerhalb eines umfassenden
Gedankenraumes und durch die eigne Bewegung des Geistes,
durch
^
eine
Mitteilung
außen
von
erfolgt; -
er
ist
so
nicht
ausschließ-
Die Unmöglichkeit mit den Mitteln des Empirismus zu einer Wissen-
schaft zu gelangen,
ist
eben neuerdings von hervorragenden Forschern nach-
drücklich hervorgehoben.
es einen
Windelband (Präludien,
2. Aufl.,
S.
303)
nennt
„hoffnungslosen Versuch, durch eine empirische Theorie dasjenige
zu begründen, was selbst die Voraussetzung jeder Theorie bildet", und Husserl
L Bd S. HO) bemerkt zum gleichen Gegenstande:
„Der schwerste Vorwurf, den man gegen eine Theorie der Logik erheben
kann, besteht darin, daß sie gegen die evidenten Bedingungen der Möglich-
(Logische Untersuchungen
keit einer
,
Theorie überhaupt verstoße."
ist der Sprachgebrauch, indem er Erfahrung
und Denken einander entgegensetzt, als vermöge die Erfahrung etwas ohne
das Denken zu leisten. Mit Recht bemerkt dagegen schon Robert Boyle (the
-
Nicht ohne Schuld daran
Zum Erkenntnisproblem.
120
von der Fülle des Einzelnen eingenommen, daß
lieh
Zu-
er ihren
sammenhang wie selbstverständlich erachtet und über den Bäumen
den Wald nicht sieht. Dem Empirismus scheint aus den Dingen
entgegenzukommen,
gelegt
wie
hat,
was
Wahrheit die Tätigkeit
in
in
sie
hinein-
der Begriff der Erfahrungswelt alles eher
B.
z.
als
Daß es ein Gesamtproblem
der Erfahrung gibt, d. h. daß der Boden, auf dem Erfahrung entein Erzeugnis der bloßen
steht,
zu gewinnen
erst
um einzelne Daten,
nur
Erfahrung ist.^
ist,
auch daß wir bei der Wahrheit nicht
sondern
um Gesamtgestaltungen und Ge-
samtüberzeugungen
kämpfen,
verdunkelt werden.
Es kann sich aber
das
sollte
nach
Kant nicht so
leicht
dem Empirismus nur ver-
dunkeln, weil ihm nur besondere Seiten der Wirklichkeit gegenwärtig
sind, die ihre Tiefe
Es
und ihren Umfang in keiner Weise erschöpfen.
sowohl
das
geschieht
nach Seite
des
des Objekts, wie es in Kürze heißen mag.
Leben
sich zunächst in
Subjekts als
nach Seite
Da unser Denken und
Bewußtseinsvorgängen
abspielt,
so bleibt der
Empirismus dabei stehen und verkennt, daß der Inhalt des Bewußtseins selbst nicht
verständlich
sie
schon
ist,
in
namentlich
alle
es
einander
ein tiefer gegründetes Selbstleben des Geistes
ohne eine Umkehrung der ersten Ansicht, wie
der Bildung eines einheitlichen Ich
aber
gibt
ohne
nicht
erfolgt,
innere Synthese erst möglich macht;
keine
einzelner
Wissenschaft.
Das Seelenleben
Bewußtseinsvorgänge
auflösen,
wie sie aber
ohne eine solche
in
das
ein
Neben-
heißt
allen
Zusammenhang preisgeben und damit auch die Möglichkeit
einer Wissenschaft von Grund aus zerstören.
Innern
Christian virtuose gegen Schluß):
'tis
When we say, experience corrects reason,
an improper way of speaking, since
tis
reason
formation of experience corrects the judgment
it
itself,
that
upon the In-
had made before.
Merkwürdig ist es überhaupt, wie oft man sich heute auf eine Erfahrung
ohne zuvor ihre Bedingungen irgend zu untersuchen und ihre Möglichkeit zu sichern.
Nirgends dürfte dies heute mehr geschehen als auf
pädagogischem Gebiet. Man richtet neue Schulgattungen ein und erklärt
bald, die Erfahrung habe sie als vortrefflich erwiesen; man neigt dahin, Einrichtungen fremder Völker einzuführen und beruft sich dafür auf die Bewährung durch die Erfahrung bei jenen. Aber ist gesagt, daß was dem einen
Volke frommt, für das andere, vielleicht unter wesentlich anderen Lebensbedingungen befindliche, ebenfalls passe? Und wenn eine Einrichtung hie
und da vielleicht unter besonders günstigen Bedingungen gute Erfolge hat,
ist damit
ein durchgängiger Vorzug erwiesen?
Die Erfahrung hefert ein
Zeugnis nur, wo wesentlich gleiche Umstände vorliegen; ob aber dies der
Fall, darum pflegt man sich wenig zu kümmern.
*
beruft,
Denken und Erfahrung.
Nach
121
Seite des Objekts aber haftet der Empirismus viel zu aus-
schheßlich an der äußeren Natur
und verkennt die EigentümHchkeit
Was von seinen Lehren ein gewisses
Recht gegenüber der Natur hat, das wird zum Unrecht in der Ausdehnung über die ganze Welt. Was uns an sinnlicher Wirkung
der anderen Daseinsgebiete.
zugeht,
das
nie
sich
läßt
voll
in
umsetzen und
geistige Tätigkeit
von innen aus entwickeln; so verbleibt hier immer eine Fremdheit
und Gebundenheit, und
es
nicht hinauszukommen.
Aber schon beim ersten Anblick des mensch-
ist
über ein Registrieren und Beschreiben
und Strebens stellt sich
lichen Lebens
die Sache anders.
Auch hier
werden zunächst uns einzelne Vorgänge zugeführt, aber es verbleibt
beim bloßen Eindruck, jene
nicht
der Betrachtende sich
diesen
in
lassen
sich auf
den erzeugenden
und mit einander verbinden; indem
Lebensprozeß zurückverfolgen
kann
Prozeß zu versetzen vermag,
das Fremde in eignes Leben verwandeln.
Kann aber so der
Mensch mit dem Menschen leben und fühlen, ihn nicht bloß wie
ein fremdes Ding von außen betrachten, so ergibt sich über das bloße
er
Noch weiter aber gelangen wir,
wenn innerhalb des menschlichen Kreises ein Geistesleben anerkannt,
Beschreiben hinaus ein Erkennen.
der Standpunkt der Erkenntnisarbeit gewählt, die Gesamtheit
hier
des geschichtlich-gesellschaftlichen Lebens sowohl als der Erfahrungen
am
einzelnen Punkt
darauf
bezogen,
von dort durchleuchtet und
Nun und nimmer kann
zusammengefaßt wird.
sich
die Be-
hier
handlung eine bloße Feststellung der dargebotenen Erscheinungen
muß
genügen
lassen,
Messen
und Umwandeln
sie
eine
innere Aneignung, mit ihr aber ein
vollziehen.
Denn was im menschlichen
Kreise an Entfaltung des Geisteslebens vorliegt, das ist seinem ersten
Befunde
nach
mit
behaftet,
daß
keine Klärung ohne
vielem
so
Menschlichen, ZufäUigen
Zeitlichen,
eine
energische Scheidung und
eine
Zurückführung zur eignen Art erfolgen kann, zugleich
hier,
aus
jenes
Leben
besonderen Beziehungen und Richtungen,
den
vorliegt
herauszuheben,
sie
und
aufstrebt,
in
gilt
umfassendes Ganzes
ein
es
denen
erst
von ihm her jene Mannigfaltigkeit aufzuhellen und
zu einem Zusammenhange zu verbinden.
Höhe dessen, was sich dem Menschen an
In
Wahrheit
liegt die
Erkenntnis erschließt, an
dieser Stelle: in den charakteristischen Entfaltungen des Geisteslebens
und dem Aufbau
einer
geistigen
Welt;
hier
liegt
daher auch die
Entscheidung über das Ganze der Weltanschauung, von hier aus ist
der Gesamttypus
unserer Weltanschauung
zu entwerfen,
von hier
Zum Erkenntnisproblem.
122
muß irgendwelche Würdigung auch das finden, was das mensch-
aus
ist
Das alles
Dasein an Schranken und Widersprüchen enthält.
liche
Erfahrungen,
voller
voller
Bewegungen und Vertiefungen,
nun und nimmer aus bloßen Begriffen hervorgehen können,
damit
durchaus jenseit
liegt aber ebenso gewiß
Beiden
bloßen
Sphäre des
der
die
es liegt
Rationalismus,
es
über allem Vermögen des bloßen Empirismus.
deutliche Abhebung des Geisteslebens vom mensch-
fehlt die
das treibt den Rationalismus zur Überspannung des
lichen Dasein:
Menschen, den Empirismus zur Verleugnung des Geisteslebens; jener
vermag dem Erkennen keinen lebendigen
Inhalt,
keinen wissenschaftlichen Charakter zu geben.
dieser vermag
Beiden
ihm
auch der
ist
Fehler gemeinsam, das Erkennen nicht einem größeren Ganzen des
Geisteslebens
einzufügen
kenntnisproblem
und
Zusammenhang damit
im
behandeln;
zu
solcher
bei
Isolierung
Erkennen entweder zu niedrig oder zu hoch bewertet.
treten
dabei
dem Erkennen
Ursprünglichkeit,
Er-
das
Beide ver-
unentbehrliche Momente: der eine die
andere
der
das
wird
die
Tatsächlichkeit.
Aber es bedarf
eines neuen Standortes, um diese Wahrheitsmomente zu einem Ganzen
zu verbinden und weder einseitig an der Größe noch an der Schranke
des menschlichen Erkennens zu haften, sondern Schranke und Größe
mit einander anzuerkennen.
Wenn der Empirismus trotz aller augen-
scheinlichen Schwächen immer von neuem aufsteigt und die Menschen
überwältigend fortreißt, so liegt das weniger an seiner eignen Leistung
als
an
den Mängeln des Wahrheitsbegriffes, der im Rationalismus
zu überwiegen
Daß er die Wahrheit über die Meinung und
pflegt.
Spaltung der Menschen emporhebt, ihr eine völlige Unabhängigkeit
vom Menschen gibt,
das bleibt sein Verdienst und sein Recht;
wo
dies Recht irgend unsicher wird, da ist eine Vernichtung der Wissenschaft nicht
zu verhüten.
Aber solange der Abstand
nicht irgend-
welche Überwindung findet und die Wahrheit nicht irgendwie zu
unserer
eignen
Sache
gemacht wird,
behält
sie
etwas Kaltes und
Totes; unerfindlich bleibt, wie sie uns mit überwältigender Kraft zu
bewegen, und wie sie das Ganze des Lebens zu erhöhen vermag.
So sehr wir es ablehnen müssen, die Wahrheit mit dem Pragmatis-
mus nach der
Brauchbarkeit für das Leben zu messen,
haupt nach einem draußenliegenden Ziele zu messen, so
sie
über-
muß doch
das Erfassen der Wahrheit als Entwicklung eines neuen Lebens ver-
standen und die Sache als nicht jenseit, sondern innerhalb des Lebens
befindlich
verstanden
werden.
Es
handelt
sich
schließlich
nicht
Denken und Erfahrung.
die jenseit des Lebens
darum, eine Wirklichkeit zu erfassen,
sondern
Leben
ein
gewinnen,
zu
läßt sich ein innigeres Ver-
ohne daß wir dem Empirismus ver-
hältnis zur Wahrheit erreichen,
der zu keiner Wahrheit gelangen würde,
fallen,
Glauben an
sie
liegt,
das eine Wirklichkeit aus sich
Verfolgung dieser Bahn
In
entwickelt.
123
wenn
er nicht
den
zu seiner Arbeit schon mitbrächte.
dem Empirismus und dem
Rationalismus wirken,
wie wir
sahen, entgegengesetzte geistige Strömungen; von der Art
und Lage
In
der Zeiten wird es abhängen, ob diese oder jene jeweilig das Über-
Wo der Gedankenkreis als im wesentlichen ge-
gewicht gewinnt.
und wohl übersehbar gilt, wie im Altertum und im Mittel-
schlossen
alter,
auch
bei
der deutschen
schätzung
Erfahrung
der
Philosophie, da wird
konstruktiven
man sich zu einer Unter-
die innere Leistung voranstehen, da wird
neigen.
Wo hingegen die Empfindung
der Enge des bisherigen Gesichtskreises vorwaltet und ein Verlangen
nach Erweiterung aufkommt, da wird von der Erfahrung
alles Heil
umwandelnde
Tätigkeit
erwartet
und
weiterbildende,
die
leicht
So geschah
geistiger Art übersehen.
im
19. Jahrhundert
und geschieht
ja
land
19. Jahrhundert vollzog,
wirken,
weil
ihr
hier
ein
Die un-
auch heute.
es vielfach
ermeßliche Erweiterung des Gesichtskreises
die das
Bacon, so geschah es
es bei
in
Natur und Geschichte,
mußte besonders stark in Deutschenergischer
Rückschlag gegen die
zu straffe Zusammenfassung der konstruktiven Systeme zur Seite ging.
Aber je mehr eine solche empiristische Bewegung sich ausdehnt
und
je
einnimmt,
das Feld
ausschließlicher sie
desto notwendiger
Wir sahen den Empirismus zu einem
leidlichen Abschlüsse nur kommen, weil er innerhalb einer vorge-
wird
eine
Gegenwirkung.
fundenen, seinen eignen Begriffen überlegenen,
Gedankenwelt wirkt;
erschüttert
und
diese
aufgelöst
Gedankenwelt aber
werden,
samer jene Richtung wird.
je
ja widersprechenden
muß um so mehr
und
selbständiger
So untergräbt
sie
Fortgang die ihr unentbehrlichen Ergänzungen und
äußeren
Siege
innerlich
wird sonnenklar, sobald
Eine solche
auch
ihre
aus eignen Mitteln
in
jetzt sich
unduld-
muß daher im
Unzulänglichkeit
alles
Wendung sehen wir trotz aller Gunst,
Empirismus noch
erfreut,
zusammenbrechen;
sie
je
durch ihren eignen
bestreiten will.
deren sich der
lebensfremden exaktwissenschaftlichen Kreisen
vorbereiten.
Immer deutlicher wird, daß alle
Aufschichtung und Anordnung von Kenntnissen noch keinerlei Er-
Zum Erkenntnisproblem.
124
kenntnis, keinerlei Ideen, keinerlei
Überzeugungen gewährt, daß aber
ohne solche der Mensch nicht auskommen kann,
bleiben, nicht zu einer Kulturmaschine herabsinken
ein Seelenwesen
Es
soll.
eine Notwendigkeit des geistigen
ist
Lebens, es
ist
im be-
der gegenwärtigen Kultur, welche
sondern die eigentümliche Lage
das
wenn anders er
Denken zwingend über den Empirismus
Ohne
hinaustreibt.
eine Selbständigkeif und Ursprünglichkeit des Denkens kann keine
Aber jene Selbständigkeit
Kultur bestehen.
übersehen und
läßt sich
Bahnen fort-
vergessen, so lange das Leben in vermeintlich sicheren
läuft,
lange es
so
nicht
von schweren Verwicklungen und WiderHeute aber stehen wir ganz und gar unter
sprüchen bedroht wird.
dem Eindruck schwerer Verwicklungen und Widersprüche,
wir er-
kennen die Notwendigkeit einer gründlichen Revision des gesamten
Kulturbesitzes,
alles
welk
die
Notwendigkeit
energischen
einer
unwahr Gewordenen,
und
einer
fassung und Steigerung aller Wahrheitselemente;
Erschütterung, daß die Ungewißheit
greift
und
Wie
sollten
um
das Ganze
des
in
Ausscheidung
kräftigen
ja
so
Zusammengeht die
tief
die letzten Elemente zurück-
Geisteslebens
kämpfen
zu
zwingt.
solchen Aufgaben irgend weiter gelangen
wir nun bei
und ursprünglicher Tätigkeit, ohne
eine Selbstbesinnung und Selbsterweckung des Geisteslebens, ohne
ohne
ein Vermögen selbständiger
Erhöhung und Erneuerung, welche neue Möglichkeiten
und neue Tatbestände erschließt? Das alles aber liegt dem
Empirismus fern. So notwendig daher die Zeit einer Innern Wandlung bedarf, so notwendig muß sie ihn hinter sich lassen.
So sehr
wir nun die Wendung der philosophischen Forschung der Gegeneine geistige
entwirft
wart
dabei
zum Idealismus begrüßen, und so sehr wir die Scheu verstehen,
der alten Metaphysik wiederaufzunehmen:
die Art
wir einer gründlichen Erneuerung und
so gewiß
durchgreifenden Kräftigung
des Lebens bedürfen, ebenso notwendig bedürfen wir eines Idealismus
aufrüttelnder
und vordringender Art,
nicht bloß ein kritischer, er
bedeutend
der
kritische
ein solcher Idealismus aber darf
muß auch ein positiver sein.
Idealismus,
der
philosophischen
Forschung vorherrscht,
und Empirismus
seine
daß
sie
nur
mit
heute
darin
auf der
ist,
Denn so
Höhe der
dem Realismus
Grenze zu weisen, im besondern darzutun,
Hülfe des Gegners ein Ganzes des
versteckter
Lebens und Wissens erreichen, so entschieden er ferner in einzelnen
Hauptrichtungen
des Lebens das Wirken
Ordnung der Dinge zeigt,
und Walten
einer
neuen
er faßt diese Hauptrichtungen nicht ge-
Denken und Erfahrung.
nügend
in
ein Ganzes
zusammen;
ein
125
Ganzes aber ist unentbehrlich,
Bewegung sein geistiges Selbst ergreifen,
den Schwerpunkt seines Lebens dahin verlegen und zugleich eine
Umkehrung seines Leben vollziehen soll. Ohne eine solche, ohne
wenn der Mensch
in
jener
eine Versetzung in ein ursprüngliches Leben von der anderen Seite
her kann aber das neue Leben schwerlich die Kraft gewinnen,
einer andersartigen
sich
gegen die ungeheuren
nächste
Weltlage
intellektuellen
um
Ordnung gegenüber selbständig zu werden und
bietet.
Hemmungen durchzusetzen,
Demnach
treibt
welche die
uns nicht eine Lust an
Abenteuern zur Metaphysik, sondern die unerläßliche
Forderung einer Selbsterhaltung des Geisteslebens.
2.
— organisch.
Mechanisch
(Teleologie.)
I
und Probleme Mechanisch
|ie Begriffe
*-^ eine besonders reiche Geschichte.
und Organisch haben
Sie zeigt nicht nur große
Gegensätze der Weltanschauung und Methodenlehre und einen harten
Kampf um den Charakter der wissenschaftlichen Arbeit, sie ist auch
Nuancen und
Schwingungen und
ergibt damit
einen eigentümlichen Durchblick der Gesamtbewegung.
Die Gegen-
feinerer
voll
leiserer
der Jahrtausende aber erstrecken
sätze
So sei
Gegenwart.
bei diesem
sich
bis
in
Arbeit der
die
Problem unser Augenmerk vornehm-
der Geschichte zugewandt.
lich
a)
Zur Geschichte der Ausdrücke und Begriffe.
Wie
Ausdrücke mechanisch und organisch, so sind auch
die
die Begriffe alt, aber erst spät haben sie einander gefunden.
erscheint, als technische Bezeichnung der
Herstellung von
Aristoteles
spätere Schrift
Jahrtausende
Maschinen,
(A7];^avixrj,
(i^
rä
[A7]x^avix,a. i
und
dient
wie ein eingebürgerter Ausdruck
(XTj^^avtxa),
In
es
Mechanisch
Kunst der Erfindungen, der
dessen
Namen
trägt
bei
eine
diesem Sinne durchlief das Wort die
seit
Descartes
zur
Bezeichnung
einer
Theorie, welche die Bildung der Natur, nach Art der menschlichen
Werke, nicht aus einer Innern Triebkraft des Ganzen, sondern aus
der Zusammenfügung kleiner von Haus aus bewegter Teilchen des
Stoffes erklärt; die Werke der Natur scheinen
von denen des Menschen
nur durch ihre größere Feinheit, also quantitativ, nicht
^
(pUatv
rr,i
In
dieser
::pa^at,
-ziy(yri<i
Siot
Schrift
wird der Ausdruck so erklärt: "OTav
xo j^aXsröv änopiav riapiyji xai oelrai
to Ttpö; ta? xotaux«?
ar.oploic,
ßor]0-ouv
8io
xe'yvy)?.
[lipoc, |jL7])(^av7Jv
qualitativ
oeV,
Tt
Tiapa
xai xaXoü(jL£v
(Arist. 847 a, 16).
Die Kunst erscheint hier als eine Art Überlistung der Natur.
Mechanisch — organisch.
Die technischen Mittel der Erklärung
verschieden.^
Mechanik
theoretische
127
als
Bewegungslehre. 2
In
liefert
dabei die
Umlauf dürfte den
Terminus mechanisch namentlich der Chemiker und Philosoph Robert
liebte und gern auf dem
Werke verwandte. Selbst am Ausdruck „Natur" nahm
Anstoß und wollte ihn durch mechanismus universalis ersetzt
Boyle gebracht haben, der ihn besonders
seiner
Titel
er
wissen.
Die Naturwissenschaft der folgenden Zeiten faßte den Terminus
bald
bald
laxer,
strenger;
Erörterungen darüber waren gang und
Aber gewöhnlich wurde unter mechanischer Erklärung eine
gäbe.
Erklärung körperlicher Beschaffenheiten aus der Figur und Bewegung
verstanden.
Übertragung auf seelische Vorgänge
Eine
mechanisch und materiell gelten
zunächst ganz fern,
dabei
lag
oft als
gleich-
Eine mechanische Erklärung seelischer Vorgänge heißt
bedeutend. ^
daher zunächst eine Ableitung aus bloß körperlichen Ursachen.
In
der
Sache aber unternimmt schon Spinoza, den Bestand des Seelenlebens
aus ,dem Zusammenwirken einzelner Vorstellungen zu erklären, wie
Descartes
^
ipsa
(sc.
facile
percipi
§ 203):
»Nullum
corpora naturalia discrimen agnosco,
et
fadorum operationes
sensu
(principia philos. IV,
sagt:
arte facta)
ut
aliud
nisi
inter
quod arte
plurimum peraguntur instrumentis adeo magnis, ut
hoc enim requiritur, ut ab hominibus fabricari
possint:
queant.
Contra autem naturales effectus fere semper dependent ab aliquibus
organis
adeo minutis,
ut
omnem sensum
effugiant.
Danach bringt
jede
Verfeinerung der Maschinen die Kunst der Natur ein Stück näher.
^ Descartes
princ. phil. IV, §200: Figuras et motus et magnitudines
corporum consideravi atque secundum leges Mechanicae, certis et quotidianis
experimentis continuatas, quidnam ex istorum corporum mutuo concursu
sequi debeat, examinavi.
§ 203 Et sane nullae sunt in Mechanica rationes,
quae non etiam ad Physicam, cujus pars vel species est, pertineant, nee minus
—
naturale est horologio ex bis vel
arbori ex hoc vel
illo
:
illis
quam
Quamobrem
rotis composito, ut horas indicat,
semine ortae, ut tales fructus producat.
ii
qui in considerandis automatis sunt exercitati, cum alicujus machinae
usum sciunt et nonnullas ejus partes aspiciunt, facile ex istis, quo modo aliae
quas non vident sint factae, conjiciunt; ita ex sensilibus effectibus et partibus
corporum naturalium, quales sint eorum causae et particulae insensiles, investi-
ut
gare conatus sum.
' So
stellt Descartes selbst dem Un körperlichen
das mechanicum et
corporeum entgegen (Briefe I, 67), und dieselbe Bedeutung liegt vor, wenn
Wolff (psych, rat. § 395) behauptet, daß die Einsichten der anschauenden
Erkenntnis
nihil
—
mechanice quoque in cerebro absolvi
qua quid in universal! repraesentatur, quod non aeque
(cognitio symbolica)
inesse notioni,
mechanice repraesentatur in corpore.
Zum Erkenntnisproblem.
128
er
denn die Seele eine
Maschine nennt (automaton
geistige
spiritu-
Leibniz verfeinert das trotz seiner Betonung der Einheit der
ale).
und
Wolff
Seele. ^
einer Übertragung des Wortes,
dann
Psychologen
französischen
die
hunderts bilden es näher durch.
Schließlich
des
kommt
18.
Jahr-
auch
zu
und „mechanisch" wird erst bildlich,
das Innere der Seele verwandt.-
lehrhaft für
es
Kant verallge-
meinert den Ausdruck, indem er ihn überträgt auf »alle Notwendig-
Begebenheiten
der
keit
Kausalität,
ob man
der
in
gleich
ihm unterworfen sind, wirklich
In
nach
Zeit
dem
Naturgesetze
der
daß Dinge, die
darunter nicht versteht,
Maschinen sein müßten."
materielle
der Naturphilosophie aber entwickelt er klar und scharf den Gegen-
satz einer
mechanischen und einer dynamischen Erklärung.^
Auch organisch begegnet uns zuerst bei Aristoteles, dem großen
Aber es hat dort einen anderen Sinn als jetzt.
Entsprechend dem Stammwort opyavov, Werkzeug, bedeutet organisch
Bildner der Sprache.
„werkzeuglich",
es
wird
gebraucht
zweckmäßig angelegten Körpers,
teilen,
nämlich
welche
solchen,
zusammengesetzt
vom Ganzen
des
lebendigen,
öfter
aber von einzelnen
aus
ungleichartigen
Körper-
Bestandteilen
So gewiß der Begriff nur Lebewesen zu-
sind.
kommt, so enthält er selbst nicht das Merkmal inneren Lebens,
er
ist
daher auch nicht über das besondere Gebiet hinaus zur Bezeichnung
^
qu'elle
S. z. B.
est,
Erdm. 153:
II
faut considerer aussi
que l'äme,
toute simple
a toujours un sentiment compose de plusieurs perceptions ä
fois;'Ce qui opere autant
la
pour notre but, que si eile etait composee de pieces
comme une machine.
-
man die Übertragung noch im Werden, er sagt im
„Wenn diese Veränderung durch innere Triebfedern, (mich
Bei Lessing sieht
7. Literaturbriefe:
plump auszudrücken) durch den eigenen Mechanismus
des
seiner Seele erfolgt
Mit besonderer Energie hat Herbart die Vorstellung von einer Mechanik
ist."
Innenlebens
durchgeführt;
er
erklärt
Organismus der Vernunft aufzulösen
stellungsreihen,
erklärt
in
es
(III,
255)
als
seine einfachen
Aufgabe,
Fibern,
„den
die Vor-
deren Entstehung nur aus der Mechanik des Geistes konnte
werden."
^
S. IV, 427 (Hart.): „Die Erklärungsart der spezifischen Verschiedenheit
der Materien durch die Beschaffenheit und Zusammensetzung ihrer kleinsten
Teile,
als
Maschinen,
ist
die mechanische Naturphilosophie;
welche aus Materien, nicht als Maschinen,
bewegender
Kräfte,
d. i.
diejenige aber,
bloßen Werkzeugen äußerer
sondern ihnen ursprünglich eigenen bewegenden Kräften
der Anziehung und Zurückstoßung die spezifische Verschiedenheit der Materie
ableitet, kann die dynamische Naturphilosophie genannt werden."
Mechanisch — organisch.
129
eines lebendigen Ganzen, etwa in der politischen Theorie, verwandt;
wo ^pyavijcd;
bei Aristoteles Stellen,
es gibt
mit mechanisch übersetzen
läßt. ^
kaum anders als
Diesen Sinn behielt das Wort unsich
verändert durch Mittelalter und Neuzeit hindurch bis in das 18. Jahr-
Jenen Begriff des Werkzeuglichen konnte sich auch die
hundert. -
neue mechanische Theorie aneignen, unbedenkHch wurden im 18. Jahr-
dem Be-
hundert organische (natürliche) und künstliche Maschinen
der Maschine untergeordnet;
griff
von organischen Maschinen
sprechen hatte damals gar nichts Befremdliches.
zu
^
Erst das Aufsteigen der deutschen Blütezeit mit ihrem Verlangen
und eignen Bewegung der Natur hat dem
Ausdruck organisch das Merkmal des Lebendigen hinzugefügt und
nach einer Beseelung
Besonders hat dahin Kant mit seinen
es zur Hauptsache gemacht.
und Unterscheidungen gewirkt, aber auch Herder,
präzisen Begriffen
Jacobi u.
Von den natürlichen Lebewesen
seien nicht vergessen.^
a.
übertrug sich die neue Bedeutung zunächst auf Staat und Gesellschaft, ^
'
dann auf Recht, Geschichte
Pol. 1259 b,
yevEJEw; xa\ (pO-opä?
rapl
S. Z. B.
Tot; atüjiaat, Bi
a; yEvvtiJsi,
23:
u. s.
w.
Organisch wird nament-
336a, 2: xal xa? Suvatisi? aTioStSdaai
Xiav 6pyavtxt3;, oupaipoüvTe;
aroprJoEtev
av
Ti;,
Tco'xEpov
ettiv
xaxa to eiSo; atttav.
ttiv
ap£Tr'
xt;
SouXou
napa xa;
opyavixa; xat Staxovtxas oXXrj TijJUWTEpa xouxtov.
*
Der letzte bedeutende Ausläufer der Scholastik, Suarez (1548—1617),
anima I, 2, 6): Dicitur corpus organicum, quod ex partibus dissi-
sagt (de
milaribus componitur.
Noch für den Sprachgebrauch der Wolffischen Schule
bezeugt Baumeister: Corpus dicitur organicum, quod
ad peculiarem quandam actionem aptum
^
nannte
Saint-Simon
machine organisee
(s.
noch
Paul Barth,
vi
compositions suae
est.
um 1813
die
Gesellschaft
Vierteljahrsschr.
f.
eine
wissenschaftl.
veritable
Philos.
XXIV, I, S. 72).
Kant definiert (V, 388 Hart.): „Ein organisiertes Produkt der Natur
S. 386
in welchem alles Zweck und wechselseitig auch Mittel ist."
heißt es: „Ein organisiertes Wesen ist also nicht bloß Maschine; denn die
hat lediglich bewegende Kraft; sondern es besitzt in sich bildende Kraft, und
zwar eine solche, die es den Materien mitteilt, welche sie nicht haben (sie
*
ist
das,
organisiert)".
Jacobi
(Hume 172): „Um
die Möglichkeit eines
organischen
Wesens zu denken, wird es notwendig sein, dasjenige, was seine Einheit
ausmacht, zuerst: das Ganze vor den Teilen zu denken." Sachlich wurden
damit nur aristotelische Gedanken wieder aufgenommen und genauer formuliert.
Kant spricht auch von einem „wahren Gliederbau" der reinen spekulativen Vernunft, „worin alles Organ ist, nämlich alles um eines willen
und ein jedes Einzelne um aller willen" (III, 28 Hart.)
' Die Übertragung des Ausdrucks Organisation auf das politische Gebiet dürfte zuerst in den Bewegungen der französischen Revolution erfolgt
Eucken, Grundbegriffe.
4. Aufl.
9
Zum Erkenntnisprablem.
130
ein Lieblingswort der Romantik,
lieh
einzelne Schulen
gebrauch
ein.
es
über
dringt aber zugleich
und Richtungen hinaus in den allgemeinen SprachMechanisch und organisch, am Ursprung ziemlich
gleicher Bedeutung,
bilden
also
schließlich
den schroffsten Gegen-
und bezeichnen jetzt Hauptgegensätze der Weltanschauung.^
satz
b)
Zur Geschichte des Problems.
Der sachliche Gegensatz, den nunmehr
weit
reicht
zurück,
Demokrit und
hat
er
Aristoteles.
im Altertum
Hauptvertreter
wie
sie
in
Kürze heißen mag,
Die Vorstellung
den Ausdruck
— des Organismus
eingebürgert,
von dem auch die Formel stammt, daß
organischen
Wesen das Ganze den Teilen vorangehe. ^
sein,
sich
die
das Ganze vor die Teile, das Lebendige vor
das Leblose und erklärt von dort nach hier.
stellung erstreckt
in
Die künstlerische und synthetische Denk-
entschiedene Oberhand.
stellt
Worte bezeichnen,
Auf der Höhe des klassischen Altertums
gewinnt die organische Lehre,
weise jener Zeit
die
seine
sofort
hat
— nicht
namentlich Aristoteles
bei
dafür
einem
Die Vor-
über ihr nächstes Gebiet hinaus- auf
den innerlichen Sinn aber haben dem Worte erst deutsche Denker und
Kant sagt (V, 387 Hart.): „Genau zu reden, hat die Or-
Dichter gegeben.
ganisation
der Natur nichts Analogisches mit irgend einer Kausalität,
dem
die
Anmerkung hinzu: »Man kann umgekehrt einer gewissen Verbindung, die aber auch mehr in der Idee als in
wir kennen"
und
fügt
in
einer
der Wirklichkeit angetroffen wird, durch eine Analogie mit den genannten
So hat man sich, bei einer neulich
unternommenen gänzlichen Umbildung eines großen Volkes zu einem Staat,
des Worts Organisation häufig für Einrichtung der Magistraturen u. s. w.
und selbst des ganzen Staatskörpers sehr schicklich bedient. Denn jedes
Glied soll freilich in einem solchen Ganzen nicht bloß Mittel, sondern zugleich auch Zweck, und, indem es zu der Möglichkeit des Ganzen mitwirkt,
durch die Idee des Ganzen wiederum seiner Stelle und Funktion nach beunmittelbaren Naturzwecken Licht geben.
stimmt
sein."
In
derselben Schrift (der K. d. Urteilskraft) sagt er S. 364
»So wird ein monarchischer Staat durch einen beseelten Körper, wenn er
nach inneren Volksgesetzen, durch eine bloße Maschine aber (wie etwa eine
Handmühle), wenn er durch einen einzelnen absoluten Willen beherrscht wird,
in beiden Fällen aber
nur symbolisch vorgestellt."
^
So z. B. bei Trendelenburg, s. Log. Untersuchungen (3. Aufl.) II, 142ff.
*
Pol. 1253a, 20 heißt es:
xo
oXov JupoTspov ava^xaiov
etvai
xou
[xtpou?.
xi?
yap xoü oXou oux eaxat noui ouSk X^V^ ^^ Kl' op-wvufxw; tforcEp £i
liycn xT^v Xi'd-ivTiv. 8ta<p^S-ap£toa yap Eoxat xoiauxr). Danach ist der Staat früher
als
das Individuum.
avatpou(xE'vou
Mechanisch — organisch.
den
—
freilich erst nach
und auf das Weltall, bald auch
und am meisten bei den späteren Stoikern — auf das
Staat
Aristoteles
Vom Altertum fließt sie in das Christentum
Ganze der Menschheit.
und gewinnt hier durch die
ein
Innigkeit.
131
Färbung eine besondere
religiöse
Sie befestigt sich später zu der Idee der Kirche als des
^
mystischen Leibes (corpus mysticum) Christi.
im
Sie erhält
Mittel-
und Sinn-
mit seiner untrennbaren Verkettung von Geistigem
alter
lichem eine greifbare Gestalt und beherrscht mit solcher die mittelalterUche Gesellschaftslehre; ^
sie
eigentümlichen Ordnungssystems,
ist
ein Hauptstück
dem
das
des jener Zeit
Einzelnen
Geistig-
alle
von einem Ganzen, und zwar einem sichtbaren Ganzen her zu-
keit
gehen
läßt.
Jene organische Lehre wirkte stark sowohl auf das praktische
Gebiet
als
Dort gebot
auf das wissenschaftliche Verfahren.
eine
sie
unbedingte Unterordnung des Einzelnen unter das Ganze, das ihm
zur Entwicklung seiner Vernunft
erst
sie 'dem
Einzelnen
das Bewußtsein,
und an
verhelfe,
aber
innerhalb
des Ganzen
gab
zugleich
etwas
zu
bedeuten.
Mit besonderer Freude verweilt das spätere Altertum bei
dem Ge-
Eigentümliches
seiner
Stelle
Unersetzliches
(i^ipo?),
sondern ein
ein
Glied
des Ganzen
tröstet
einen
Marc Aurel
danken, daß der Einzelne, nicht bloß ein Teil
Glied
(pi>.o;)
des Weltalls
der Vernunftwesen«,
in
diese
sei.
»Ich
bin
Überzeugung
den Gefahren und Dunkelheiten des Lebens.
aber
entwickelt
namentlich die Vorstellung,
daß
Die
alle
alte
Kirche
Christen
als
Glieder des gottgeweihten Gemeinwesens in Schicksal und Tat auf
einander angewiesen, einander solidarisch verbunden seien.
Nicht minder folgenreich ist jene Denkweise für die wissenschaft-
Denn hier quillt aus ihr die teleologische Betrachtung,
vom Altertum mit mächtiger Wirkung bis in die Gegen-
liche Arbeit.
die sich
wart erstreckt.
War das Ganze das Ursprüngliche und Überlegene,
so bot es den Schlüssel
*
zur Erklärung der einzelnen Glieder und
Bezeichnend für den griechischen Ursprung der Vorstellungsweise ist
die Tatsache,
daß unter den Evangelien nur das Johannesevangelium
sie
bietet (Weinstock und Reben), das stark unter griechischen und philosophischen
Einflüssen steht.
^
Dem entspricht es, daß die Analogie zwischen dem Staat und einem
lebendigen Körper nicht beim allgemeinen Gedanken verbleibt, sondern gern
bis
ins
Einzelne
ausgeführt wird.
So hat
z.
B. Johann
von Salisbury für
jeden Teil des Staates ein entsprechendes Körperglied aufzuweisen versucht
(s.
Gierke,
Das deutsche Genossenschaftsrecht III, 549).
9*
Zum Erkenntnisproblem.
132
Das Ganze
Leistungen.
war
aber
nach
platonisch-aristotelischer
Vorstellung eine unwanddbare Form, ein bei sich selbst befindliches
und
befriedigtes Leben;
so setzte es aller
und Schlußpunkt,
Das
in sich selbst
einen festen Ziel-
i
reicht
des Lebendigen hinaus in das gesamte All.
ein
lebendiges,
fest
Bewegungen
sich gliedartig einfügt; die
über das Gebiet
Die Welt
dem
ineinandergefügtes Ganzes,
vollendetes
sich
in
gilt
hier als
alles
Einzelne
laufen nicht wirr durchein-
um dort in ein
ander, sondern eine jede strebt zu einem Endpunkt,
beharrendes,
Bewegung
Wirken
überzugehen.
(evepysia)
Aber besonders fruchtbar ist jene Betrachtungsweise innerhalb
Heimatsgebietes: bei den Lebewesen.
ihres
Nicht nur werden hier bei den
einzelnen Tierarten alle Organe und Funktionen auf ein allumspannen-
des Leben
bezogen und daraus verstanden,
Mannigfaltigkeit
organischer Bildung als
Normaltypus, der durch
alle
Diesen Normaltypus zeigt
Stufen wirkt.
alle
rein der Mensch, so läßt sich
erscheint auch
es
Entfaltung eines einzigen
von ihm aus das weite Reich durch-
leuchten
und der unermeßliche Stoff unter durchgehende Gedanken
bringen.
Es entstand damit eine Art von vergleichender Anatomie
und Physiologie, sowie eine Entwicklungsgeschichte; auch das Seelenleben der Tiere wird
vom Menschen
her aufzuhellen
gesucht.
So
wenig uns jetzt jene Betrachtungsweise genügen kann, eine gewisse
Zusammenfassung und Anordnung des Stoffes hat sie ihrer Zeit und
langen Jahrhunderten geboten.
Auch im Altertum
aber
Widerständen
nicht an
es
fehlte
dagegen,
gelangten nicht aus bloßer Gegenwirkung zur Leitung der
sie
Das geschah
Arbeit.
in
erst
organische
und
Klarheit wurde.
wo
der
Kampf gegen
des Strebens
nach Freiheit
der Neuzeit,
Lehre ein Hauptstück
jene
Die Befreiung erfolgt zuerst im Gesamtzuge
des Lebens, indem das moderne Individuum die Bindung an eine
Organisation und
greifbare
durch
als
eine
unerträgliche
*
S.
Aristot.
TT?
oüot);
£<jti
?V£xa.
S. femer lQ9a,
8'auTT),
Tou teXou; S'?vexa raXXa,
teles
30: im\
des Geisteslebens da-
Bedrückung empfindet und
Physik 194 a, 28:
xtvr-'<j£(jj?
Vermittlung
abweist,
zum All erstrebt und daraus eine sichere
ein unmittelbares Verhältnis
ouve^^ous
die
r,
ti
i]
Sk
<puais
(pu'at?
auxr)
Sirrr,
av
arj
f|
ou
te'Xo? xa\
{ikv
r ahla.
a;
f,
^vexa.
av yap
touto Eo^^atov xai to ou
teXo? tt^ xtv/jaew?,
uXt)
r
S'w;
jJioptpr',
xeXoi;
Nach Aristo-
-ou ?v£xa.
könnte der Zufall wohl einzelne zweckmäßige Bildungen hervorbringen,
nun und nimmer aber die durchgehende Zweckmäßigkeit;
zweite Buch seiner Physik.
s.
darüber das
Mechanisch — organisch.
Überlegenheit gegen
sichtbare
alle
Ordnung
133
So zuerst
erringt.
in
der Renaissance und in der Reformation, so auch in der politischen
und wirtschaftlichen Befreiung, wie sie namentlich von England ausging.
Mit solcher direkten Begründung des Lebens auf das Indi-
viduum scheint es unermeßlich an Kraft, Vernunft und Wahrheit zu
Alle Zusammenhänge erscheinen hier als das Werk der
gewinnen.
Individuen,
haben nur
sie
Nach Leibniz
gewährt
soviel Recht,
Unendlichkeit
des Alls Und entwickelt sie
wie fem
hier jene organische Lehre!
liegt
Gleichzeitig
erfolgt
das Individuum
als
Individuum
das
trägt
auch eine
sich
in
die
ihnen
ganze
aus sich selbst heraus;
Umwälzung
Wissenschaft
der
Die überkommene Erklärung der Natur aus dem Ganzen und Innern
wird unerträglich,
Deutung,
nicht
sind
die
als
sondern
Bild,
Werke
Lehre,
der scholastischen
Kräften
erscheint sie als ein
rantiae,
s.
B.
als
mit
ihren
ist,
weil es
Erklärung gibt
vollwichtige
jener Zeit voller Klagen über
lichkeit
z.
das als eine durchaus subjektive
das energisch abzuweisen
bloßes Bild,
als ein
sich
man empfindet
die
So
versteckte Bild-
inneren
Formen und
«Asyl der Unwissenheit« (asylum igno-
Oldenburg an Spinoza).
Demgegenüber wird zur
Grundbedingung einer echten Erkenntnis, daß alles Innere aus der
Natur vertrieben und alle Zusammenhänge in kleinste Elemente aufgelöst werden; die Ermittlung
und Verfolgung dieser Elemente ver-
spricht zugleich ein Durchsichtigmachen der bis dahin verworrenen
Wirklichkeit
und
Denn vom
Kleinen her lassen sie sich bewegen
eine
Macht über die sonst unzugänglichen Dinge.
und umgestalten.
Für das Große der künstlerischen Art der Alten fehlt hier aller Sinn;
So
sie
war ja auch durch die Scholastik aufs stärkste verkümmert.
die
mechanische Naturerklärung der Neuzeit; in direktem und be-
wußtem Gegensatz zur
Denkweise macht
älteren
zur Hauptsache, unternimmt
sie
sie
die Elemente
von ihnen her allen Aufbau, zerlegt
überkommene Kontinuum bei Raum, Zeit und Bewegung in
diskrete Größen und gibt sie damit allererst eine exakte Begreifung
Mit der Leugnung aller inneren Zusammenhänge
der Phänomene.
sie
fällt
das
natürlich auch die teleologische Betrachtung; die verschiedensten
Erwägungen verbinden
anthropomorph,
als
sich
unklar,
zu ihrer Verwerfung:
als
unfruchtbar.
sie
erscheint als
Für die Einheit der
Natur haben jetzt statt der Zwecke die Gesetze zu sorgen, die überall
gleichmäßig wirken und als einfache Grundformen
keit
beherrschen.
Das
alles
ergreift
die
Geister
alle
Mannigfaltig-
mit
elementarer
Zum Erkenntnisproblem.
134
echtes Wissen
Gewalt, mit der neuen Denkweise scheint zuerst ein
und zugleich ein Zeitalter der Wissenschaft aufzusteigen, alle bisherige
Arbeit konnte dafür nur als eine Vorstufe gelten.
Daß
neue Denkweise
die
Probleme
hervorrief,
Fragen
viele
offen
ließ
und neue
Der
konnte tieferen Geistern nicht entgehen.
bedeutendste Denker der Aufklärungszeit,
behandelt
Descartes,
die
mechanische Theorie nur als ein Prinzip der exakten Naturbegreifung,
nicht als
metaphysische Lehre von den letzten Gründen
eine
der
Dinge, er zieht zugleich eine scharfe Grenzlinie zwischen sich und
Sein treuer Schüler Robert Boyle verficht eine
Demokrit. ^
zweck-
ein unentbehrliches Gegenstück zu
mäßig wirkende Ursache als
den
mechanischen Ursachen. ^ Berkeley macht geltend, daß die mechanische
Betrachtung nur die Gesetze und Formen (modes)
nicht seine Ursache.
erkläre,
eigentümlichen Typus
entwickelt einen
tief
des Geschehens
Leibniz ein: er
greift
der Weltanschauung, indem
Natur mit ihrem Mechanismus zur Erscheinung einer
er die ganze
Wirklichkeit
geistigen
Besonders
macht,
die
letzten
Einheiten,
die
für
die
mechanische Betrachtung einen bloßen Grenzbegriff bilden, zur Haupt-
und
erhebt
sache
als
sie
Monaden mit innerem Leben
Innerhalb der Natur will er alles
Prinzipien
mechanisch
des Mechanismus aber scheinen
erklärt
selbst
versieht.
wissen,
einer
die
Erklärung
zu bedürfen und diese nur in einer zweckmäßig waltenden Weltvernunft finden zu können. ^ Es sieht aber Leibniz die Zweckmäßigkeit
der Naturgesetze darin, daß sie
alle
dem Ziele dienen, ein möglichst
großes Quantum von Kraft zur Existenz zu bringen.
die
er
*
criti
kürzesten
Wege
eingeschlagen
und
die
Überall findet
einfachsten
Mittel
Die wichtigste Stelle dafür findet sich princ. philos. IV, § 202: (Demo-
philosophandi
ratio) rejecta est,
primo quia illa corpuscula indivisibilia
rejicio; deinde quia vacuum circa
quod ego nullum dari posse demonstro, tertio quia gravitatem iisdem tribuebat, quam ego nullam in ulio corporum cum solum spectatur, sed tantum quatenus ab aliorum corporum situ et motu dependet atque
supponebat, quo nomine etiam ego illam
ipsa esse fingebat,
ad
illa
'
refertur, intelligo
S. z. B.
primordia
rerum
(sive breviore
ff.
de ipsa natura sect. IV:
infinita. sua
sapientia
Hamm autem partium motum sub
ac potestate
ita
direxit,
ut
tandem
tempore sive longiore, ratio definire nequit) in speciosam hanc
ordinatamque mundi formam coaluerint.
*
Omnia in corporibus fieri mechanice, ipsa vero principia mechanismi
generalia ex altiore fönte profluere (S. 161, Erdm.); s. auch 155a, Foucher II, 253.
Mechanisch — organisch.
Auch
gewählt.^
die Schule hält daran
Zusammengesetzte und
alles
während die
Seele,
scharfer Fassung
damit
ganze Körperwelt gehöre,
ihm entzogen
Einfaches,
als
daß dem Mechanismus
fest,
die
I35
In
sei.^
minder
Wolff nach scholastischer Art eine Erklärung
stellt
aus den Wirkursachen und eine aus den Zweckursachen nebenein-
ander und bildet dabei den Ausdruck Teleologie. ^
Natürlich
wurde
Lehre durch jenes
die
überlieferte
organische und
wegs mit einem Schlage
gestürzt;
dazu Stack
und den Methoden der Schule.
Begriffen
teleologische
Aufkommen der mechanischen Erklärung keinessie
viel
Auch
zu
fehlte
tief
den
in
es nicht
an
tüchtigen Männern, welche die Eigentümlichkeit des Lebendigen eifrig
Aber sie fanden
verfochten.*
durfte es erst einer
keit
des
steigen
Ohr der Zeit.
Dafür be-
Das geschah namentlich mit dem Auf-
suchen und sehen hieß.
deutschen
ein Verlangen
nicht das
neuen Lebenswoge, die anderes in der Wirklich-
Humanismus;
siegreich
erhebt sich
nach mehr Unmittelbarkeit des Lebens,
innigeren Verhältnis zu Welt und Natur,
in
ihm
nach einem
nach einem Schauen der
* S. 147b (Erdm.): semper scilicet est in rebus principium determinationis
quod a maximo minimove petendum est, ut nempe maximus praestetur effectus
minimo ut sie dicam sumptu. Den Einwand, ob nicht bloße Naturnotwendig-
keit dasselbe Ergebnis hätte hervorbringen
Cela serait
vrai, si
par exemple
les loix
können, beantwortet er dahiq (605b):
du mouvement, et tout le reste, avait
sa source dans une necessite geometrique de causes efficientes; mais il se trouve
qiie dans la derniere analyse on est oblige de recourir ä quelque chose qui
depend de causes finales ou de la convenance.
So z. B. Baumgarten, Metaph. ed. VI, 1768, § 433: machina est com-
positum stricte dictum secundum leges motus mobile. Ergo omne corpus in
mundo est machina. — Machinae natura per leges motus determinata mechanismus est. At, quidquid non
est
compositum, non est machina, hinc nulla
monas est machina.
dari
* S. philos. ration. sive logica cp. III,
§ 85: rerum naturalium duplices
possunt rationes, quarum aliae petuntur a causa efficiente, aliae a fine.
Quae a causa efficiente petuntur, in disciplinis hactenus definitis expenduntur.
Datur itaque praeter eas
alia adhuc philosophiae naturalis pars, quae fines
nomine adhuc destituta, etsi amplissima sit et utilissima. Dici
posset teleologia.
Der Ausdruck causa finalis ist dagegen scholastisch; ich
rerum
explicat,
finde ihn zuerst bei Abälard.
*
Natur,
Obenan
s.
steht
hier
Cudworth mit
seiner
Annahme einer plastischen
namentlich the true intellectual System of the universe (1678) I, 3, 19.
Von den deutschen Gelehrten ist hier vornehmlich Rüdiger zu nennen, s. z. B.
institutiones eruditionis seu philosophia synthetica pag. 109: physica vel mecha-
nica est vel vitalis.
Zum Erkenntnisproblem.
136
Dinge aus dem Ganzen; was dabei zunächst als
Drang
die
Gemüter
erregte,
das
künstlerischen Lebensgestaltung;
von
da
ein
ungestümer
allmählich
sich
klärte
zu
einer
aus fand sich leicht eine
Rückkehr zu den Alten, die als das Muster
lauterer und edler Natur
erschienen.
So war es kein Wunder, daß,
Renaissance,
auch die organische Denkweise eine Auferstehung
Stück der jüngsten
als ein
daß sie mit wunderbarem Zauberklange die Gemüter
lebte,
er-
ergriff
und gewann.
Eigentümlicherweise hat dieser neuen künstlerischen Denkweise
wenig künstlerische
wissenschaftlich zuerst der seiner Gemütsart nach
Kant die Bahn gebrochen.
mus zu
Er hat es getan, indem er den Mechanis-
bloßmenschlichen Denkweise
einer
herabsetzte
und damit
freien Platz für eine andersartige Betrachtung und Behandlung schuf;
nur mußte für eine solche sich ein zwingender Anlaß finden.
Einen
solchen
aber schien
ihm das Reich der organischen Bildung zu
bieten,
indem
unsere Begriffe nur mit Hilfe der Idee eines
inneren
Ganzen und
es
eines
Zweckes eingehen kann.
leitenden
So
hier die alte Lehre wieder aufgenommen, auch über das nächste
ward
dem Ganzen
Gebiet hinaus
das
in
alles
der Welt zugeführt.
in vorsichtiger Abgrenzung
Betrachtung.
flutete
rasch
gewann
ein
Bei
Kant
selbst
und als eine Sache menschlicher
Aber die vordringende künstlerische Lebenswoge überGrenzen und Dämme, die organische Denkweise
alle
stolzes Selbstbewußtsein
und gab
sich
der Aufklärung
gegenüber als ein Verstehen aus dem eigensten Leben und Wesen
der
Dinge heraus, während die mechanische Lehre nüchtern und
Mit besonderer Energie
schien.
seelenlos
Denkweise zum Ausdruck gebracht und
Idee des
hat
alles
Schelling die
neue
Naturleben unter die
Organismus gestellt^
Begriff
Gebrauch.
und Ausdruck kommen dann
Bei
aller
Festhaltung antiker
rasch in den allgemeinen
Elemente
ist
ein
Einfluß
modemer Denkweise dabei nicht zu verkennen. Die Idee des Organismus liefert hier weniger ein Bild vom Sein als vom Werden; die
Wirklichkeit
ein aus
bildet
Wandlung zunächst
als
hier weniger
ein
geschlossenes
Kunstwerk
eignem Vermögen fortschreitendes Lebewesen.
für die Natur.
weit fruchtbarer für das Reich
So wird
als
die
der Geschichte
Einen bestrickenden Reiz gewinnt der Gedanke,
* Den direkten Gegensatz zu
mechanisch bildet ihm aber gewöhnlich
dynamisch; dort scheint ihm die Welt als etwas Gegebenes, hier als etwas
unablässig Werdendes verstanden zu werden.
Mechanisch — organisch.
daß
alles geschichtliche
Werden
nicht stoßweise, sondern in ruhigem
durch
künstliche
bewußtlosen
Naturtrieb,
nicht
von
sondern
der
eines
Ganzen
Fortgang,
nicht
aus
Kraft
I37
durch
sondern
Reflexion,
einen
dem bloßen Individuum
erfolge.
Indem
her,
diese
sich
Vorstellung auf Staat, Recht, Sprache u. s. w. überträgt, scheint durch-
gängig eine reinere und reichere Tatsächlichkeit, ein größeres Bild
vom Ganzen, ein innerlicheres und ruhigeres Verhältnis des Menschen
Nun soll er sie nicht mehr von draußen
zu den Dingen gewonnen.
her meistern,
sondern von innen
nicht machen,
nun kann
satz
die
engste
her miterleben,
Lebens mit
Stelle
rationalen
verbunden.
das Recht
B.
z.
Erzeugnis des Volksgeistes finden;
und
Wendung zu
zur
ein
als
er die Fülle des geschichtlichen
Art anerkennen
duellen
ist
sondern
an seiner
jedes
aller
So
im Gegen-
einer geschichtlichen Weltansicht,
organischen Lehre
der Aufklärung,
jener
Die
Forschung
historische
indivi-
würdigen.
nahen Freundin die künstlerische Kontemplation;
aufs
aber
hat
es
bezeichnend,
ist
zur
hier
daß^Schelling für den »dritten und absoluten Standpunkt der Historie"
den der historischen Kunst erklärt.
Aber auch die Einseitigkeit dieser historischen Betrachtungsweise
und zugleich die Schranken der organischen Lehre ließen sich nicht
Bedenken mußte schon das erwecken, daß jene
lange verkennen.
organische Lehre von der politischen und kirchlichen Reaktion, von
Männern wie Adam Müller und de Maistre, dem Vater des modernen
Ultramontanismus, mit besonderer
alterlichen Sinne zur
Individuen
der
als
Wärme ergriffen und im mittel-
Unterdrückung der Selbständigkeit sowohl der
lebendigen
Aber
Gegenwart gewandt wurde.
auch über diese besondere Gestalt hinaus kam das Problematische
und Einseitige der organischen Lehre bald zur Empfindung. Jenen
ruhigen Fluß des geschichtlichen Werdens hat sie weit mehr vorausgesetzt als
erwiesen;
was an Objektivität aus den Dingen
gegenzukommen schien, das hat sie selbst in diese
ist
ihr Bild der Geschichte
bilde brachte jene
Augenmerk
Dinge
richtete,
Naturkräfte
Bewegung
das
auf
von
Leben
starker
auch kräftige Antriebe
gab.
Subjektivität.
wertvolle Anregungen,
und den
inneren
ent-
hineingelegt; so
Dem Naturindem
sie
das
Zusammenhang der
zum Suchen der Einheit der
Aber wissenschaftlich fruchtbar sind
diese
An-
regungen nur nach Verpflanzung auf den andersartigen Boden der
modernen Naturforschung geworden; soweit jene organische Denkweise
aus eigenem Vermögen einen Abschluß versuchte, hat
sie sich in ge-
Zum Erkenntnisproblem.
138
Dem Ganzen des Lebens
wagte, oft wunderliche Gebilde verloren.
aber brachte
sie Gefahr,
indem
sie
den Menschen
kontemplativ zur Wirklichkeit verhalten
ließ,
sich
überwiegend
ihn weit
mehr einlud,
willfährig aufzunehmen und sich einzufügen als selbständig aufzutreten
und eigne Wege zu bahnen; das Ganze war namentlich ungeeignet
für eine Zeit, die voll großer Aufgaben war und schwere Verwicklungen
in sich trug.
So kam
die Herrschaft
wieder an
die
andere
die
Seite,
nie
ganz unterdrückt, sondern nur eingeschüchtert war, mit frischer Kraft
neu hervor. Etwas anders gefärbt, aber im Grunde unverändert
trat sie
stieg die
istische
Aufklärung wieder auf, von ihr aus mochte jene human-
Epoche mit ihrer organischen Lehre als eine bloße Episode
erscheinen.
Der Aufbau des Gesellschaftslebens von den Individuen
her kam im modernen Liberalismus und in der modernen Freihandelslehre erst zur vollen Entwicklung, bis in die zweite Hälfte des neun-
zehnten Jahrhunderts sehen wir A. Smiths scharf zugespitzte Theorie
selbst
von hervorragenden Gelehrten
als eine
und einen endgültigen Abschluß behandelt.
ausgemachte Wahrheit
Die Naturwissenschaft
aber unternahm, unter schroffer Ablehnung der naturphilosophischen
Spekulation, 'alle
Reste vitalistischer Theorie aufs gründlichste aus-
nun dahin gestellt, auch die organische
Bildung mit ihrem Leben ganz und gar auf die physikalischen und
zutreiben; die Aufgabe wurde
chemischen Elementargesetze zurückzuführen.
Unter den Philosophen
hat namentlich Lotze eine solche universale Geltung des Mechanismus
verfochten,
Reiche
freilich
seelischen
nicht
ohne ihm, ähnlich wie Leibniz,
Lebens einen
jenes Übermechanische
tieferen
Grund zu
in
einem
geben.
Aber
war Sache der Metaphysik, die Natur verblieb
dem Mechanismus, und in der Zeit kam diese Bejahung mehr zur
Wirkung
richtig
als
jene
verstanden
Begrenzung.
und
— So
umsichtig
Lösung der großen Probleme; so
blieb, das Prinzip
erschien
angewandt,
viel
der Mechanismus,
als
eine
gesicherte
der Ausführung zu tun ver-
dünkte allem Zweifel enthoben.
Da kam ein Widerstand, ein unerwarteter Widerstand,
aus der
eignen Bewegung des modernen Lebens, nicht aus Nachwirkungen
älterer
Zeiten,
und weniger aus
einer
künstierischen
Deutung der
Wirklichkeit als aus wachsender Erfahrung, aus neuen Tatsachen und
Die wirtschaftliche und industrielle Gestaltung des
modernen Lebens treibt die Menschen enger zusammen und vervielfacht ihre Berührungen, sie differenziert und kompliziert die Arbeit
neuen Aufgaben.
Mechanisch — organisch.
13g
und bindet damit den einen weit enger an den anderen,
aber an
Ganzes;
ein
vor der Gemeinschaft,
die
Einzelnen
alle
daraus erwächst,
verschwindet die Isolierung, in welcher der Mechanismus die IndiHatte er alle Zusammenhänge von ihnen her abgeleitet,
viduen sah.
modernen Soziologie der Einzelne von vornherein
so scheint der
einem sozialen Zusammenhange angehörig,
zum bloßen Produkt
den Einzelnen
dahin,
vom
Lehre
die
Milieu
das Unsichtbare der Einflüsse zur Geltung und
bringt auch
seiner
neigt
Umgebung zu
Zugleich wird die Wehrlosigkeit der Individuen gegenüber
machen.
den wirtschaftlichen Verwicklungen und Gegensätzen grell empfunden
und mit
ihr
Notwendigkeit eines Gesamtwillens,
die
wie
der
ihn
Staat verkörpert.
Das alles drängt dahin, den Gedanken des Organismus wieder
aufzunehmen,
Philosophen hat namentlich Comte seine
unter den
und Politik von ihm aus gestaltet.
Ethik
Aber der Begriff wird
bei
ihm gegen die ältere Fassung beträchtlich verändert, er wird, wenigstens
grundlegenden
der
in
Ethischen
ins
aus
Erörterung,
der
Histologie
Fortschritte
eine
anschauliche Ausführung geben.
ist
auch die Gesellschaft ein
Künstlerischen
Es sind
und
namentlich
die dem Grundgedanken
Wie der lebendige Körper,
überaus feines Gewebe aus lauter
die
so
dem
Naturwissenschaftliche versetzt.
(Bichat),
Elementen; so eng sind diese miteinander verschlungen,
einzelnen
daß das Tun und Lassen, das Wohl und Wehe des einen unmittelbar auch die anderen berührt.
Die moderne Arbeitsteilung hat
was von jeher galt, noch weiter gesteigert,
die Anderen
heit des Einzelnen an
Damit scheint
Augen,
ein
sie stellt die
und an das Ganze
leitendes Prinzip
für Ethik
gewonnen, das nur entwickelt zu werden braucht,
dies,
Gebundendeutlich vor
und
Politik
um allem Handeln
bestimmte Bahnen vorzuschreiben.
In
Wahrheit
ist
ein solches Prinzip nicht
sowohl begründet als
durch eine Vermengung antiker und moderner Art erschlichen; unvermerkt
wird
aus
der
Verwebung
ein
Ganzes innerer
Art,
aus
dem Sein ein Sollen hervor.
dem alten Boden, wenn
das Ganze mit Forderungen an den Einzelnen kommt und sie
ihm als Verpflichtung auferlegt.
Das Dunkel, das dem Begriffe
organisch von jeher anhaftete, wird durch solche Vermengung alter
der Tatsache geht ein Wertbegriff, aus
Schließlich
befinden
wir
uns
ganz
auf
und neuer Art bis zu unerträglicher Verworrenheit gesteigert.
Aber
man hält an dem Begriffe fest, weil es drängt, die Abhängigkeit des
Zum Erkenntnisproblem.
140
Einzelnen
vom Gesamtstande irgendwie zu bündigem Ausdruck zu
bringen.
So
moderne Forscher
der
gerät
unter
widerstreitende
Anregungen, und ein Auseinandergehen bis zu schroffem Gegensatz
kann nicht Wunder nehmen.
Gebiete trennen sich
Nicht bloß die Individuen, auch die
Am meisten Anklang hat die organische
hier.
Lehre bei den Soziologen, weit weniger bei den eigentlichen National-
ökonomen gefunden; unter den Juristen sind ihr am ehesten hervorragende Germanisten gewogen.
Mit dieser Bewegung auf sozialem Gebiet geht parallel eine
Bewegung in der Naturwissenschaft; wie sie später einsetzte, so ist
sie heute noch weit mehr mitten im Fluß und Kampf. Unverkennbar
hat dazu an erster Stelle die moderne Entwicklungslehre gewirkt.
Die darwinistische Form,
gelangte,
war
in
der
in
sie zuerst
dem Eigentümlichen
zu allgemeiner Geltung
ihrer
Art der Anerkennung
des Organischen so abgeneigt wie nur möglich, sucht
sie
doch das
ganze Gebiet des Lebendigen den Begriffen des Mechanismus zu
Aber auch
unterwerfen.
massen
oft
ihrer
in der Naturwissenschaft
eignen Absicht entgegen.
wirken Gedanken-
Indem das Reich des
Lebens weit mehr die Aufmerksamkeit und die Arbeit gewann, kam
mehr zur Geltung, und
es erwies
sich
die Zurückführung seiner Erscheinungen auf die physikalischen
und
seine Eigentümlichkeit weit
chemischen Elementargesetze unvergleichlich schwieriger,
Mitte des Jahrhunderts
als
um die
Die Beobachtungen
am
der Entwicklungsmechanik,
das
angenommen
war.
die neuen Ausblicke
Problem der Kontinuität des Lebens, die Mutationslehre mit ihrem
Protoplasma,
Aufweis sprunghaften Entstehens neuer Formen
ander ergibt eine
Scheidung der Geister.
u.
a.,
alles
mitein-
veränderte Lage.
Dabei erfolgt eine
Hoffen die einen von
einer Verfeinerung
wesentlich
der mechanischen Begriffe eine intellektuelle Aneignung der neuen
Tatsachen, so glauben die anderen auf ein neues Prinzip nicht verzichten zu können.*
^
In diesen
Bewegungen erhebt sich von neuem
S. u. a. Rindfleisch, Ärztliche Philosophie 1888 und Neovitalismus 1895.
Roux, Einleitung zum Archiv für Entwickelungsmechanik der Organismen
(1894), wendet sich dagegen, „die organische Form als Unerklärbares und
bloß teleologisch Ableitbares zu bezeichnen" (3. 22), und bemerkt weiter:
„Für den Forscher auf dem Gebiete der Entwickelungsmechanik gilt in hohem
Maße das Wort
»Incidit in scyllam, qui vult vitare charybdim."
Die zu einfach mechanische und die metaphysische Auffassung repräsentieren
Mechanisch — organisch.
141
auch das Problem der Teleologie, ihrer eignen Absicht nach weniger
als ein
Stück der Metaphysik, denn
als
ein Mittel naturwissenschaft-
licher Erklärung, als
„empirische" Teleologie,^ aber auch
von anderen
Rückfall in die Metaphysik bekämpft.
als
ein
solche
als
Wird so die mechanische Lehre vom Gebiet des Lebendigen
her, wenn nicht eingeschränkt, so doch über die herkömmliche Form,
„die
zu einfach mechanische Auffassung"
(Roux),
hinausgetrieben,
Grundbegriffe mannigfach an-
so werden
weiter auch
gegriffen.
Schon die unermeßliche Verfeinerung, welche scheinbar
ihre
eignen
elementare Vorgänge der unorganischen Natur enthüllten, lassen die
Vorstellungen des älteren Mechanismus für das Unterlebendige selbst
als viel
zu roh erscheinen.
Prinzipiell hat namentlich die energetische
Naturlehre das mechanische Weltbild bekämpft, indem
sie die
Grund-
vorstellung von der Materie als einem jenseits der Sinnesempfindungen
befindlichen Sein,
und
als
einem besonderen Träger der
Naturerscheinungen
alle
zurückzuführen suchte. 2
hier
den
auf
Auf die Probleme,
nebenbei einzugehen, verbietet
hat die mechanische Theorie
sie lange zu
haben schien.
sich
Kräfte,
bestritt
der
Energie
Grundbegriff
die
daraus erwachsen,
schlechterdings;
jedenfalls
die Selbstverständlichkeit verloren, die
Selten aber wird etwas neu zum Problem,
ohne sich dabei umzubilden.
So finden wir heute das ganze Gebiet voll Erregung und Streit;
die Scylla
und die Charybdis, zwischen welchen dahin zu segeln in der Tat
ist; und es ist nicht zu leugnen,
schwer und bis jetzt nur Wenigen gelungen
daß die Verführung zu letzterer Auffassung mit der Zunahme unserer Kenntnis
zunächst erheblich zugenommen hat" (S. 23). S. auch W. Roux: „Über die
Selbstregulation der Lebewesen" 1902.
^
Coßmann,
S.
Elemente
der
empirischen
Teleologie
1899,
ferner
König, »Die heutige Naturwissenschaft und die Teleologie". Beil. zur
Es ist eine
Allg. Z. 1900, No. 29 u. 30, sowie „Über Naturzwecke" 1902.
überaus reiche, unablässig wachsende Literatur über diese Probleme entE.
standen, ein deutliches Zeichen, wie sehr sie im Mittelpunkte des Interesses
und der Arbeit stehen.
^
S.
Ostwald, Vorlesungen über die Naturphilosophie S. 153:
„Alles,
was wir von der Außenwelt wissen, können wir in der Gestalt von Aussagen
über vorhandene Energie darstellen; und daher erweist sich der Energiebegriff
Er
allseitig als der allgemeinste, den die Wissenschaft bisher gebildet hat.
umfaßt nicht nur das Problem der Substanz, sondern auch noch das der
Über die Bedeutung des Begriffs Energie aber heißt es S. 158:
»Wir werden allgemein Energie als Arbeit, oder alles, was aus Arbeit entsteht und sich in Arbeit umwandeln läßt, definieren."
Kausalität."
Zum Erkenntnisproblem.
142
die Entscheidung aber liegt nicht bei allgemeinen Reflexionen, sondern
bei
Hauptrichtung,
der
welche Arbeit und Leben
in
Wirklichkeit
So zeigte es die Vergangenheit, so wird auch
einschlagen.
in
Zu-
kunft der eigne Fortgang der Arbeit darüber befinden, wie sich das
Verhältnis der Gegensätze gestalte, welche Weiterbildungen für beide
Grundbegrilfe nötig werden,
auch ob ihnen
aufkommen mögen.
klärungsarten
Das
gegenüber
neue Er-
aber darf die philosophische
Betrachtung erwägen, wie der Überblick der Wirklichkeit heute den
Stand der Begriffe
zeigt,
und welche Aufgaben er nahelegt.
Erwägungen zum Kampf der Gegenwart.
c)
Das Problem im Gebiet der Philosophie.
a.
Die Philosophie muß vor allem darauf bestehen, daß der Mechanismus, selbst wenn er sich der ganzen Breite der Dinge bemächtigen
nun und nimmer einen endgültigen Abschluß zu bilden
könnte,
vermag.
Denn das, was für ihn das Letzte ist,
das Nebeneinander
der Elemente, wird der philosophischen Betrachtung notwendig zu
einem
schweren
Problem.
Stünden
nämlich
Elemente
die
ohne
irgendwelche Verbindung gleichgültig nebeneinander, so wäre schlechterdings nicht zu ersehen, wie eine Wirkung des einen
entstehen können.
sollte
Leibniz
und einen Lotze
die
der Wechselwirkung
Tatsache
das nächste Weltbild gründlich umzugestalten.
trieb,
zum andern
So vor allem in der Natur, wie denn einen
dazu
Und auch der
Gedanke Leibnizens ist nicht wohl abzuweisen, daß kein Ding darin
aufgehen kann, lediglich etwas für andere zu leisten, sondern daß es
auch für sich
selbst etwas sein
muß. daß daher auch das als letztes
Element Ergriffene irgend etwas Eignes zu sein
dieses
hat; die Verfolgung
Gedankens drängt dahin, das Reich des Mechanismus zur
bloßen Erscheinung einer andersgearteten Welt herabzusetzen. Auch
beim Seelenleben würde, wer alles Geschehen auf den mechanischen
Verlauf
von Assoziationen zurückführen
wollte,
ratlos
der
Frage
gegenüberstehen, wie dies ganze Getriebe als ein eignes Leben, als
mein und
Einheit
ist
dein
erlebt
werden könnte.
und Zusammenhang Sorge zu
Überall
tragen,
ist
und
irgendwie für
dieser Aufgabe
der Mechanismus nicht gewachsen.
Läßt so der Mechanismus hinter sich ein ungelöstes Problem,
so darf auch
ganzen
dem
Tatbestande nach er nicht als Beherrscher der
Wirklichkeit gelten,
selbst
wenn
die Natur
ihm ganz ge-
Mechanisch — organisch.
143
Denn zur Natur gesellt sich das Seelenleben, dies aber bringt,
hörte.
namentlich auf der menschlichen Stufe, eine völlig andere Art des
Geschehens mit
Denn soweit das Innenleben über eine bloße
sich.
Begleitung der Naturprozesse
hinauswächst
und
selbständige
eine
soweit in uns Geistesleben aufsteigt, genügt nicht mehr
Art
entfaltet,
ein
Zusammentreten einzelner Elemente, sondern hier befindet sich
jede einzelne Erscheinung innerhalb eines Ganzen,
und hier wird eine
Verkettung nicht direkt zwischen den einzelnen Elementen, sondern
durch ihr Verhältnis
z.
zum Ganzen hindurch gewonnen.
Das Denken
B. verläuft allerdings in einzelnen Vorstellungen, aber es
ein bloßes Aneinanderreihen
und Aufschichten
ist
nicht
dieser, es verfolgt ein
bestimmtes Ziel und wird dadurch innerlich zusammengehalten; es
kann nichts dulden, was diesen Zusammenhang
stört.
diese neue Art bezeichnender als
und
logischen
Widerspruchs.
die Tatsache
Widerspruch
Ein
ließe
Nichts
für
ist
die
Macht des
sich
gar
nicht
empfinden, würde nicht im Denken die Vielheit von einer Gesamt-
umspannt, und er könnte nicht so unerträglich sein wie er
tätigkeit
es
wäre nicht das Verlangen nach Einheit von gewaltiger Stärke.
ist,
Zugleich
erweist der
bindung,
als
Widerspruch eine
Mechanismus
der
völlig andere Art der Ver-
aufbringen
sie
kann:
er
ist
kein
räumlicher Zusammenstoß, sondern eine Unverträglichkeit des Inhalts;
so erscheint
hier
der Begriff
schlechterdings unverständlich
Prinzip der
Anordnung mit
Bedeutung,
lichen
des
Inhalts,
der
dem Mechanismus
ist.
Der Inhalt bringt aber ein neues
sich:
das der Sachlichkeit, der sach-
der gegenseitigen
Determination.
Das
ist
z,
B.
das Verhältnis der Merkmale eines logischen Begriffes; nur gröbstes
Mißverständnis
kann
die
innere
Struktur
eines
solchen
Begriffes
mit dem Nebeneinander einer sinnlichen Vorstellung zusammenwerfen.
Die Grundform der Verbindung
ist
hier die des Systems: jedes Ele-
ment steht innerhalb eines Ganzen, unter dem Einfluß und der Triebkraft eines Ganzen,
Elementen.
in wechselseitiger
Determination mit den anderen
So gehört dem Mechanismus jedenfalls nicht das Ganze
der Wirklichkeit.
Daher verschwindet auch der Zweck
lichen
hat sie
wenn
Denn im mensch-
nicht aus der Welt,
die Natur für ihn keinen Platz mehr bieten sollte.
Leben hat er zweifellos eine Wirklichkeit und eine Macht,
nicht
gemeinsamen
er
bloß in der Seele des Einzelnen, sondern auch im
Leben
Wissenschaft und
bei
Kunst,
den
großen
Zusammenhängen,
wie
sie
Recht und Moral, schließlich das Ganze
Zum Erkenntnisproblem.
144
der Kultur bilden. ^
Damit,
Innenleben wesentlich
ist,
ist
daß
es
gehörig erwiesen, und es
keit
das Bild der Welt unweigerlich
im Oesamtanblick die Sache
schließlich
Entweder
als
daß diese Tatsache verständlich wird.
so zu gestalten,
sich
ist
Handeln dem
zum Ganzen der Wirklich-
zwecktätiges
ein
auch
Ja es
notwendig
— Oder. Wir pflegen heute die 'Welt
als
stellt
auf
ein
ein Stufenreich
aufsteigender Bildung zu betrachten, aber eine große Scheidung der
Geister erfolgt bei der Frage,
ein
ob
diesem Stufenreich das Höhere
in
Ergebnis des Niederen
bloßes
sei
dem Höheren
oder ob in
ganze Erklärung finde,
und daher aus ihm
ein
seine
Neues und
Ursprüngliches durchbreche, das sich nur aus einem tieferen Ganzen
Der Gegensatz erlangt seine höchste
Welt verstehen läßt.
Spannung bei der Frage des Verhältnisses von Natur und Geistesleben. Ist dieses ein bloßes Erzeugnis jener, oder beginnt mit ihm
der
Die Beantwortung dieser Frage
eine neue Stufe der Wirklichkeit?
über Recht oder Unrecht des Zweckes.
entscheidet auch
Geistesleben
mit seiner Innerlichkeit und Ganzheit von
Ist
da§
eigner Art
und eignem Ursprung, so gehört es wesentlich zum Ganzen, so
muß es von vornherein in der Bewegung wirksam sein und ihr eine
Richtung zu sich geben;
dann
erhält
das Weltgeschehen ein Ziel,
und die spekulative Weltbetrachtung wird der Zweckidee nicht entkönnen ;2
raten
Natur, so
aber das Geistesleben ein bloßes Erzeugnis der
alles
entfällt
und mit ihm auch der Zweck; dann
Menschheit sinnlos ins Vage und Leere
Ziel
Welt und die
die
treibt
ist
dahin.
ß.
Das Problem in der Naturwissenschaft.
Im Gebiet der Naturwissenschaft
auf die allgemeinen physikalischen
den Mittelpunkt des
und chemischen Gesetze zurück-
Daß aus dem Zweck „reale Kategorien" hervorgehen, hat Trendelen,
^
bürg
bildet
ob die dem Leben charakteristischen Erscheinungen
Streites die Frage,
einem hervorragenden
in
gezeigt;
s.
Kapitel
der
„Logischen
Untersuchungen"
Kap. XI.
^
So treibt es uns hier wieder zur Metaphysik gemäß jener Überzeugung
Herbarts (Wke. 11, 461): „Im Denken über Natur und Menschheit drängt
sich die Kraft des Geistes unvermeidlich zur Metaphysik hin, welche, ähnlich
den Urgebirgen
menschlichen
schroffen,
selten
hinausragt."
— zugleich die weite,
,
tiefe,
unsichtbare Grundlage alles
und Trachtens ausmacht, zugleich in einzelnen,
erklommenen Spitzen über alle andern Höhen und Tiefen
Dichtens
Mechanisch — organisch.
I45
führbar sind, oder ob in ihnen eine neue Art des Geschehens anerkannt werden muß.
Tatsächlichkeit
Diese Frage
ist
vor allem eine Frage der
und gehört als solche
in
die Fachwissenschaft,
aber
es wirken in die Behandlung auch manche allgemeinere Erwägungen
ein,
an denen wir nicht vorbeigehen können.
bar,
daß die Eigentümlichkeit,
mehr
Lebens wieder
heute
wir
nicht so
den
in
das
So viel
ist
unbestreit-
Problem und Geheimnis
Vordergrund
gerückt
des
und daß
ist,
darüber hinauskommen wie unmittelbar
Mehr und mehr scheint es ausgeschlossen,
bloße Eigenschaft des Stoffes zu fassen, mehr
leicht
vorangehende Zeiten.
das Leben
als
eine
und mehr wird ihm eine Selbständigkeit zuerkannt.
So,
um einige
hervorragende außerdeutsche Namen zu nennen, von Bergson, namentlich in seiner l'evolution creatrice^ (1907),
so von Sir Oliver Lodge.^
Bei solcher Auffassung wird es zur Aufgabe,
ein
wesentliches
und
unterscheidendes Kennzeichen des Lebens herauszuheben; ein solches
Boutroux
findet
in
der Fähigkeit «ein System zu schaffen,
in
dem
gewisse Teile gewissen anderen Teilen untergeordnet sind"; so würde
„agens" und «Organe" geben, die zusammen eine «Hierarchie"
es ein
für die es in der anorganischen
bilden,
Welt keine Analogie
gibt.^
Für Bergsons Fassung des Lebens sind folgende Stellen bezeichnend
La vie est, avant tout, une tendance ä agir sur
*
L'evolut. CTeatrice pag. 105:
la
matiere brüte;
fem er 197: La vie c'est-ä-dire la conscience lancee ä travers
la matiere.
Lodge
*
sagt »Leben
und Materie" (deutsche Übers. 1908, S.
104) zu-
sammenfassend: «Die Anschauung vom Leben, die ich im vorigen auszudrücken versucht habe, ist die, daß es weder Materie noch Energie sei, noch
auch nur eine Funktion von beiden, sondern daß es in eine ganz andere
sei, daß es in einer Weise, die wir zurnoch nicht durchschauen, imstande sei, mit der materiellen Welt in
Wechselwirkung zu treten, aber daß es auch, abgesehen von dieser, seine
wissenschaftliche Kategorie zu setzen
zeit
ursprüngliche Realität habe, wennschon diese sich der sinnlichen Erfassung
entzieht.
Es
Natur und
in
ist
abhängig von der Materie nach seiner Erscheinung in der
uns hier und jetzt und nach seinen gesamten irdischen Wir-
kungen; an und für sich aber ist es davon unabhängig, und sein Wesen ist
und dauernd, während seine Wechselwirkung zur Materie disund zeitlich ist.« Ferner siehe S. 38: »Ich gebrauche das
Wort Leben' in ganz allgemeinem Sinne. Denn wenn ich es nur in dem
kontinuierlich
kontinuierlich
,
engen Sinne, nach dem es nur die Stoffwechselvorgänge bedeuten würde,
genomnien hätte, so wäre es natürlich absurd, ihm Existenz gesondert von
der Materie zuzuschreiben.
leicht
Für diesen engeren Sinn könnte man sich viel-
passend der Bezeichnung
,
Vitalität' statt ,Leben'
bedienen."
O. Beelitz, Die Lehre von Zufall bei E. Boutroux 1907,
Eucken, Grundbegriffe. 4. Aufl.
10
ä
S.
S. 91.
Zum Erkenntnisproblem.
146
Bergson
entscheidenden Beweis für das Wirken des
einen
erblickt
Lebens als einer psychischen Kraft
zu
pflegt
bilden
daß die Natur
der Tatsache,
demnach
und
gleiche
Ziele
auf
abweichenden
Fragen
findet
sich
manche Ver-
zu verfolgen scheint. ^
Wegen
der Behandlung
Bei
schiedenheit zwischen
merkenswert
in
in
oder ähnliche Strukturen bei sehr verschiedenen Organismen
gleiche
ist
dieser
den einzelnen Kulturvölkern, namentlich be»Rolle, die das Prinzip der Diskontinuität
hier die
dem jüngsten französischen Denken
als das
82 ff.) geschehen
ist.
philosophischen Literatur
tritt
(Moderne Philosophen
der französischen
S.
Es läßt sich die-
spielt". ^
selbe mit ihren Motiven nicht besser schildern,
von Höffding
Er sagt dort:
„In
die Diskontinuitäts-
philosophie auf besonders interessante und energische Weise hervor.
Diskontinuitätsphilosophie
Es lassen sich drei verschiedene, für die
maßgebende Motive unterscheiden.
Die Erfahrung
bietet Qualitäts-
verschiedenheiten dar, deren Reduktion weder der Spekulation noch
der
gelungen
Entwicklungslehre
war.
Comtes Positivismus
ausdrücklich die Kluft anerkannt, welche
von
gebiete
einander
die
Für Comte bezeichnete jede
trennt.
hatte
verschiedenen Natur-
neue
Wissenschaft eine besondere, irreduktible Gruppe von Erscheinungen.
— Hierzu kommt, daß
^
selbst
in
jeder
einzelnen
Gruppe von
Er-
«Le pur mecanisme serait donc
oü nous l'entendons, demontrable par
si Ton pouvait etablir que la vie fabrique certains appareils
S. L'evolution creatrice 1907, pag. 59:
refutable et la finalite, au sens special
un
certain cote,
identiques,
gentes.
par des moyens dissemblables, sur des lignes d'evolution diver-
La force de la preuve serait d'ailleurs proportionelle au degre d'ecarte-
ment des lignes d'evolution choisies, et au degre de complexite des strudures
similaires
qu'on trouverait sur elles. Wie sich übrigens auch vom Standpunkt
eines feineren, dabei tiefere Probleme bereitwillig anerkennenden Mechanismus
dem Leben ein eigentümlicher Charakter zugestehen läßt, zeigt namentlich
W. Roux. Er betrachtet als eine universelle elementare Eigenschaft der Lebewesen «die zur Dauerfähigkeit im Wechsel der Verhältnisse nötige Selbstregulation in der Ausübung aller Einzelfunktionen", er sieht darin „diejenige
von allen Leistungen, welche die Lebewesen am meisten von allen andern
Naturkörpern unterscheidet, indem sie die direkte Anpassung an die wech-
Aus der unübersehbar langen, trotz
Umstände zahllose Generationen gleicher Art her-
selnden äußeren Verhältnisse bewirkt.
des Wechsels der äußeren
stellenden
Dauer der
daß auch
die
lationsfähigkeit
einzelligen
Lebewesen
ist
mit Sicherheit zu schließen,
niedersten
Lebewesen außer der Vererbung diese Selbstregu-
haben"
Archiv für Entwicklungsmechanik der Organismen
(s.
XXIV. Bd., 4. Heft [1907], S. 685).
? S. Harald Höffding, Moderne Philosophen (1905) S. 67.
Mechanisch — organisch.
147
scheinungen der Kausalsatz nur unvollkommene Bestätigung zu finden
Man zieht deshalb Hume wieder hervor und stellt seinen
vermag.
Empirismus gegen die Versuche
mus
auf,
die Kant
und der Evolutionis-
— Endlich verweist man
machten, denselben zu überwinden.
auf das Bewußtsein der Initiative, des Vermögens, durch sein Denken
und Handeln etwas Neues
in
Welt zu
die
und man betont
setze i,
mit Stärke die moralische Bedeutung dieses Vermögens. "^
Bei
Mechanismus eine
Neigung bestehen,
Lebens
vielmehr
wird
an
scharfe Kritik
geübt.
er die Welt wie ein gegebenes
daß
verfehlt,
keinerlei
des
Erscheinungen
zurückzuführen,
Kräfte
kann
Denkweise
solcher
charakteristischen
auf
solchem
Versuche
des
Am Mechanismus scheint
und geschlossenes System
im Fluß Befindliches, daß
behandelt, nicht als etwas
die
unterlebendige
er
daher
alle
Bewegung von innen her, sowie alle Möglichkeit eines wesentlichen
daß er den Verbindungen der Elemente nichts
Fortschrittes leugnet, 2
anderes zuschreiben
seine Erklärungen
sammensein
will,
was jedem einzelnen zukommt, ^ daß
als
den Elementen beizulegen pflegen, was ihr Zu-
aufweist,*
daß
er
nicht
genügend
beachtet,
wie
die
genauere Erkenntnis des Lebensprozesses immer mehr die vermeintIsolierung der Elemente aufhebt.^
liche
^
hervorragendsten Vertreter dieser Diskontinuitätsphilosophie sind
Djp.
Renöuvier (f 1903) und E. Boutroux, dessen Werk de l'idee de la loi naturelle
dans la science et dans la philosophie contemporaine (1895) 1907 in einer
deutschen Übersetzung von Benrubi erschien.
'
S.
Bergson, L'evolution creatrice pag. 40: L'essence des explications
mecaniques
est
en
effet
en fonction du present,
^
häufig
de considerer l'avenir et le passe comme calculables
et
de pretendre ainsi que tout est donne.
S. 57: „Man begegnet
dem Satze, daß, was immer an Eigenschaften dem Ganzen zukomme,
Sir Oliver
Lodge, Leben und Materie (Übers.)
auch in den Teilen zu finden sei.
Atomen kann Eigenschaften
Dieser Satz ist falsch.
besitzen,
die den
Ein Aggregat von
einzelnen nicht
zukommen,
auch nicht im geringsten Grade."
„man konstatiert hier einfach das zu Erwägende
dann den Atomen zu, in der Hoffnung, daß damit dem
Fragen ein Ende gemacht sein wird." Bergson hat diesen Gedanken namentlich gegenüber der Entwicklungslehre Spencers verfochten, diese scheint ihm
*
und
(s.
S. Lodge a. a. O. S. 47:
schiebt es
L'evol. creatr. VI)
evoluee,
puis ä
la
darin zu bestehen: ä decouper la realite actuelle, dejä
recomposer avec ces fragments,
et
ä se donner ainsi, par
avance, tout ce qu'il s'agit d'expliquer.
'
S.
Bergson, L'evolution creatrice pag. 205: plus la physique avance,
meme des particules en
plus eile efface d'ailleurs l'individualite des corps et
10*
Zum Erkenntnisproblem.
148
Mit
solcher
Wendung zum Leben und
seiner vordringenden
Bewegung tritt auch die Zweckbetrachtung in ein anderes Licht. Die
völlige Verwerfung der Zwecke in der Natur wurzelte in der lange
vorherrschenden Neigung, das Leben nicht als ein Urphänomen zu
fassen, sondern es vom Leblosen abzuleiten, in schroffem Gegensatz zur
älteren Denkweise, die (Jen ganzen Befund der Natur vom
In einer gewissen
aus erklärte.
Rückkehr zu
Lebendigen
wenn auch in
dieser,
großer Verfeinerung der Betrachtung, treten nun die Tatsachen wieder
mehr
in
den Vordergrund, die eine Richtung der Bewegung auf
ein erst zu erreichendes Ziel, eine «Zielstrebigkeit" (K. E.
sowie ein Zusammenstreben
zu
Die Schwierigkeit,
scheinen.
zeigen
von Baer),
einem Ganzen
einzelner Elemente zu
das irgend vorstellbar zu
machen, ohne die menschliche Art des Erwägens und Überlegens
in
die Natur hineinzutragen, hat schon Aristoteles stark empfunden,
muß sie noch größer erscheinen.
uns Neueren
dazu
nicht
keit dürfte
führen,
Aber alle Schwierig-
Tatsachenkomplexe
zurückzustellen
und geringzuachten, weil sie sich nicht dem Rahmen des Mechanismus fügen. Haben sich doch die Theorien den Tatsachen, nicht
diese jenen anzupassen.
Das Hauptbedenken, das gegen den Vitalismus und die Teleologie auch in der neueren Form erhoben wird, ist, daß das von
ihnen behauptete Gestaltungsprinzip «einfach alles, und zwar auf die-
Weise"
selbe
ohne daß wir von den für die verschiedenartigen
erklärt,
zweckmäßigen Gestaltungen notwendig verschiedenen determinierenden
Faktoren
und deren Wirkungsweisen etwas erfahren können.^
Ausführung dessen
In weiterer
sagt
Roux, der
Probleme" keineswegs leugnet: «Es
letzte
ist
verbleibende
«stets
überaus
leicht,
zweck-
mäßig Erscheinendes von einem wirklich zwecktätigen Agens abzuleiten.
Diese letzte Annahme bleibt uns immer noch, wenn die andere
wirklich als
dem
nicht zureichend erwiesen
scientifique
commengalt par
les
decomposer; corps
corpuscules tendent ä se fondre dans une interaction universelle.
^
xal
was jetzt erst kurz nach
Beginn exakter kausaler Forschung zwar vielfach so scheinen
lesquelles l'imagination
et
ist,
Ttt
y.a,zu
«pu'oiv
äXX(i)v,
xtv£;
199 a, 17: d ouv toc xaxi t:qv -ziyyriv ^vexa tou, SfXov
S. Z, B. Phys.
xaxa
ttjv
6(ao(cü5 yflip
<puaiv.
Jtpos
xa
ouxe
^rjxi^aavxa
xa ucrrepa
a ouxe
TOxepov vw
xi^yri
i]
xivt
ort
^jti Jtpo? aXXrjXa Iv Tot? xata -ziyyriv xai iv toi;
Ttpdxepa.
(AcIXioxa
Se
(pavepov
ouxe ßouXsuga'jjLeva
uotet.
ini
xwv !^towv xuiv
od-ty
SianopoZai
öcXXw IpyaJ^ovxai o'f x'apayvat xa\ oi (Aup[XT]x£i; xai xa xotauxa.
* S. W. Roux im Archiv für Entwicklungsmechanik der Organismen,
XXVI. Bd., 4. Heft (1907), S. 687.
Mechanisch — organisch.
mag, aber doch nicht bewiesen werden kann.
dagegen, solches
»scheinbar
Agentien abzuleiten.
Zweckmäßige" von
I49
Sehr schwer
nicht
es
Erstere Lösung aber läßt alle in den verschiedenen
Fällen verschiedenartigen Determinationen
unbekannt, verlegt
sie
in
Wir aber
Wirkungsweisen nicht aufhellbares Prinzip.
ein in seinen
ist
zwecktätigen
möchten auch diese « Bestimmungsfaktoren « und ihre Wirkungsweisen
erforschen.
Gemeinsam ist uns beiden die Erforschung der physikalisch-
chemischen Ausführungsfaktoren des Determinierten, denn daß das
durch seelisches Wirken „Determinierte" durch physische Faktoren
ausgeführt wird, geben
die Sache
mitten
ja
auch unsere Gegner zu" (S. 688).
So
ist
im Fluß, aber aus Bewegung und Zusammenstoß
Fortgang des Wissens mit Sicherheit hoffen.
läßt sich ein
y.
Das Problem auf gesellschaftlichem Gebiet.
Daß zum Verständnis des gesellschaftlichen Zusammenseins der
Mechanismus nicht ausreicht, ist ohne Mühe zu zeigen, schwierig aber ist,
ähnlich wie bei
der Natur,
ihm gegenüber zu einer positiven Be-
Von den bloßen
hauptung zu kommen.
Einzelelementen her läßt
sich irgendwelches Interesse für das Ganze, irgendwelche innere
Ge-
genwart des Ganzen, irgendwelche Hoheit und Selbständigkeit des
Ganzen, wie
B. des Staates, irgendwelcher geistige Charakter des
z.
Ganzen in keiner Weise begreiflich machen.^ Die mechanische Theorie
müßte die Gemeinschaft in ein seelenloses Räderwerk verwandeln,
in
dem
jeder
nur seine eigenen Ziele
verfolgt;
eine
gemeinsame
Gedankenwelt wäre dabei unmöglich.
Auch die Rechtsidee, worauf
die Anhänger der mechanischen Theorie sich gern berufen, ist von
hier
aus nicht zu erklären,
Gebilde erscheinen.
Vernunft.
ist
könnte hier nur als ein mystisches
Denn der Rechtsgedanke entwickelt sich nie vom
natürlichen Einzelwesen,
solches aber
sie
sondern
nicht möglich
nur
vom Vernunftwesen
her, ein
ohne Begründung in einer Welt der
Das Recht kann vom bloßen Individuum her zu entetehen
nur scheinen, wenn dabei unvermerkt dem natürlichen Einzelwesen
das vernünftige Geisteswesen untergeschoben wird.
So geschah es
namentlich in der englischen Aufklärung, wie denn die politischen und
wirtschaftlichen Systeme eines Locke
gängigen Widerspruch
*
Vortrefflich
in
und eines A. Smith einen durchsie arbeiten mit Größen der
sich tragen:
hat dies neuerdings Gierke ausgeführt: „Das
Verbände", Rektoratsrede.
Berlin 1902.
Wesen der
Zum Erkenntnisproblem.
150
Natur und behandeln
sie
Wer die Ver-
wie Größen der Vernunft.
mengung erkennt, der durchschaut zugleich die Unzulänglichkeit des
hier gebotenen Ganzen.
Aber mit dem Nein
ist
Der
kennen.
Begriff
Gesellschaftslehre
heißt
nicht
des Organischen
überkommen,
andersartigen Kultur
über das Ja entschieden,
nicht schon
mechanische Lehre abweisen
die
ist
organische aner-
uns aus
einer
älteren,
er trägt die Färbung der antiken
und Weltanschauung.
Die Vorkämpfer der organ-
möchfen ihn davon befreien;
ischen Lehre
die
können
sie
sich darauf
mit Begriffen arbeiten, die der Lauf der Ge-
berufen,
daß wir
schichte
über die Enge der anfänglichen Fassung weit hinausgebildet
Aber bei solchen Fragen
hat.
liegt alles
an der besonderen Art des
will
uns nun scheinen, daß jenes Anfängliche dem Begriffe
fest anhafte,
um nicht leicht den Gedanken auf die Stufe älterer
Es
Falles.
zu
oft
Denkart zurückzuziehen.
Die Verwendung des Begriffs Organismus
für das gesellschaftliche Gebiet ist zunächst eine bloße Analogie; mögen
gewisse Übereinstimmungen zwischen einem organischen Lebewesen
und einer gesellschaftlichen Ordnung bestehen ob sie das Wesentliche
und das charakteristisch Geistige treffen, daran läßt sich recht wohl
:
Zunächst bildet der zur Aufklärung herangezogene Bau
zweifeln.
der Lebewesen selbst ein schweres Problem und
eben
wieder ein Gegenstand
heute
härtesten
ist,
wie wir sahen,
Streites;
von
seinen
philosophischen Definitionen, wie sie Aristoteles und, fügen wir hinzu,
auch Kant und seine Nachfolger gaben, hat Lotze mit Recht gesagt,
daß
sie
mehr das Rätselhafte des Eindruckes wiedergeben als eine
Beim Begriff des Organischen bringt uns nicht,
Erklärung enthalten.
wie es wohl scheinen kann, die Natur die Sache sicher und
fest
entgegen, sondern es wird von uns eine eigentümliche Vorstellungs-
weise in die Natur hineingetragen und wird, dort ins Anschauliche und
Körperliche
Umweg?
gestaltet,
dem
Geist wieder zugeführt.
Warum
dieser
Enthält er nicht die Gefahr eines Einströmens naturhafter
Größen in das Geistesleben, oder doch des Einsetzens eines bloßen
Bildes für eine Erklärung?
Das Hauptbedenken aber ist das zähe Fortleben der griechischDenkweise
mittelalterlichen
in
diesem
Begriff;
es
droht,
den not-
wendigen Gedanken an eine innerlich überwundene Stufe zu binden.
Die organische Theorie der älteren Zeit betrachtet den Einzelnen ganz
und gar
ihn
als
völlig
ein
in
Glied des Ganzen,
das
Verhältnis
sie
läßt,
zum Ganzen
bei präziser Fassung,
aufgehen,
sie
kennt
Mechanisch — organisch.
Recht
Selbständigkeit, keinerlei
keinerlei
des
151
gegen
Einzelnen
das
Das war von vornherein nur möglich durch eine Vermengung von Staat und menschlicher Gemeinschaft überhaupt; was
Ganze.
immer das Zusammenleben für den Menschen bedeutet, das ward für
den Staat in Anspruch genommen; so konnten Ethik und Politik, individuelles
und gesellschaftliches Lebensziel als völlig gleichartig gelten.
In Wahrheit ist jene organische Lehre nicht einmal
der treue Aus-
druck des Staatsl6bens auf der Höhe der griechischen Kultur, sondern
sie ist ein Gebilde der
Auflösung
Philosophen, ein Unternehmen, der beginnenden
in lauter individuelle Lebenskreise
zu widerstehen,
sie
ist
der Versuch einer Restauration, erfolglos wie alle derartigen Versuche.
Ja
die
Philosophen
selbst
haben
am
meisten
dazu
getan,
ihre
Forderung unmöglich zu machen, indem sie vor allem den Menschen
über
den
eines
neuen Lebensideals der wissenschaftlichen Forschung hinaus-
bloßgesellschaftlichen
Lebenskreis
durch
die
Eröffnung
hoben.
Derselbe Aristoteles, der den Staat für früher (d. h. begrifflich
früher)
erklärt
als
den Menschen,
sieht
nur
dem
in
theoretischen
Leben mit seiner Richtung auf das große All ein wahrhaft glückseliges Leben.
Und damit formuliert er nur die durchgehende Über-
zeugung der gesamten griechischen Philosophie, zu deren Hauptverdiensten die Befreiung des Individuums von der gesellschaftlichen
Umgebung gehört.
Mittelalter.
Die Hauptstätte der organischen Lehre
ist
das
Hier ward in der Kirche das gesellschaftliche Ganze dem
Individuum unbedingt überlegen, hier erhob es den Anspruch, dem
Menschen alle Geistigkeit zuzuführen, hier bemaß sich alle Bedeutung
der Einzelnen nach der Stellung im Ganzen, hier ward das Ganze
zum Gewissen der Menschheit.
Auch die wirtschaftliche Gestaltung
des Mittelalters bildet ein Ordnungssystem, das aus sicherer Überlegenheit
dem
Einzelnen
seine
Stellung
Gedankenwelt hat eine hierarchische
zuweist
Gestalt,
indem
Ja
die
gesamte
von gewissen
Zentralwahrheiten der Religion und Metaphysik den einzelnen Gebieten
ihre Richtlinien vorgeschrieben werden,
die sie
nicht zu prüfen oder zu ändern haben.
Für diesen Stand mag Vor-
stellung
und Ausdruck «organisch"
Für die Neuzeit aber
ist
als
nur weiterzuführen,
angemessen erscheinen.
nichts charakteristischer, als
daß
sich
das Leben von solcher Bindung an einen sichtbaren Mittelpunkt befreit
und über die ganze Fläche des Daseins gleichmäßig ausgedehnt
hat:
die Individuen
sind
geistig
wie gesellschaftlich selbständig ge-
worden, und die einzelnen Lebensgebiete wollen ihre Probleme selb-
Zum Erkenntnisproblem.
152
sie wollen, jedes an seiner Stelle, auch um das
Dem an die mittelalterliche Denkweise Gewöhnten
behandeln,
ständig
Ganze kämpfen.
muß das als ein kecker Abfall, als eine eigenwillige Auflösung aller
Zusammenhänge erscheinen, wie es denn selbst freier nnd universaler
denkenden Katholiken recht schwer zu werden
Art und ein eigentümliches Recht des
liche
erkennen.
In
die
die eigentüm-
Wahrheit bedeutet jene Abwendung vom
nicht die Preisgebung aller,
hänge;
pflegt,
Protestantismus anzuMittelalter
sondern nur die sichtbarer Zusammen-
Größe der Neuzeit
liegt
in
der
Entfaltung
und Ver-
fechtung der Überzeugung, daß das Geistesleben als Ganzes an jeder
gegenwärtig
Stelle
und sich zu
sei
voller Tätigkeit bringen lasse; so
braucht der Mensch Zusammenhänge nicht erst von draußen her zu
empfangen, sondern sie umfangen ihn von innen her; mit ihrer vollen
Aneignung aber erhält er eben durch die innere Bindung eine sichere
Überlegenheit gegen
Bindung
solche
sich
eigne Entscheidung
nie
und menschliche Ordnung.
von außen erzwingen
erst solche
läßt,
Da
sondern eine
und Zuwendung verlangt, so ist sie kein Gegen-
sondern eine Zwillingsschwester der
satz,
keit,
alle sichtbare
Freiheit.
Auch
verleiht
Wendung dem Leben den Charakter der reinen Innerlich-
während es unvermeidlich ein Moment des Äußeren und Äußer-
lichen behält,
Ordnung
solange der Einzelne an erster Stelle einer sichtbaren
angehört.
Persönlichkeiten
wie Luther und Kant
stellen
uns deutlich genug vor Augen, wie sehr diese Umwälzung, dieser
Fortgang von einem sichtbaren zu einem unsichtbaren Ganzen, diese
Möglichkeit und diese Forderung, an jeder Stelle ursprüngliches und
unendliches Leben zu erwecken, den Anblick der menschlichen Wirklichkeit umgestaltet
Solche Umwälzung aber trägt einen Bruch mit der organischen
muß nun als zu eng und beengend empfunden
Der Mensch geht nicht auf in das Verhältnis zur gesellschaftlichen Umgebung und noch weniger in das zur politischen
Lehre
in sich; diese
werden.
Gemeinschaft; auch hat das Ganze, das uns umfängt, einen geistigen
Charakter nicht
aus
als
einen festen, aller Gefährdung entzogenen Besitz,
dem die Individuen mühelos schöpfen, sondern was immer sich
im geschichtlich-gesellschaftlichen Leben an gemeinsamen Vorstellungen,
Einrichtungen u.s.w. gebildet hat, das verliert seinen geistigen Charakter
sofort,
wenn
es
nicht
durch die Arbeit der Individuen, namentlich
durch die großer Persönlichkeiten, immerfort mit neuem Leben erfüllt
wird; wie überhaupt, so erhält sich auch im gesellschaftlichen Ganzen
Mechanisch — organisch.
die Geistigkeit
nicht
153
durch ihr bloßes Dasein, sondern nur durch
eine fortwährende Erneuerung, durch ein unablässiges Schaffen.
scheint uns die Hauptgefahr der organischen Lehre,
für
allemal
vorhanden
betrachtet,
daß
Das
sie als ein
was immer neu aus
freier
Tat
muß; sie will im Gegensatz zum Naturalismus dem
Zusammenleben einen ethischen Charakter geben, aber sie gerät dabei
in Gefahr, das Ethische selbst als ein Ruhendes und damit naturhervorgehen
zu fassen.
haft
erlegen
die
ist:
Es
ist
das dieselbe Gefahr,
der die Romantik oft
der Rückschlag gegen die bloße Reflexion führt unter
Macht von Naturbegriffen.
Warum
sollen
wir also
die
not-
wendige Wahrheit einer so problematischen Form verketten, warum
nicht
für
die
Eigentümlichkeit
geistiger
Zusammenhänge Formen
suchen, die der modernen Stufe des Lebens entsprechen?
3. Gesetz.
Zur Geschichte.
a)
|er Begriff des Gesetzes steht heute im Mittelpunkt der wissen-
I
*—^ schaftHchen Arbeit; man streitet über seine Ausdehnung und
man streitet über seinen Inhalt, besondere Fassungen kommen auf und
und Feinde,
finden Freunde
die Auseinandersetzung zwischen Natur-
und Geisteswissenschaften ist hier besonders lebhaft, ja erregt.
weniger scheint hier gekämpft zu werden
nichts
als
Um
um die Eigen-
tümlichkeit der einzelnen Wissenschaften wie um den Gesamtcharakter
der wissenschaftlichen Arbeit.
So breitet sich eine schier unermeßliche
Fülle von Erörterungen aus; unsere Betrachtung kann an die Probleme,
die dabei in Frage stehen, mehr nur erinnern als sie zu fördern hoffen.
Der Begriff des Gesetzes
ist
vom Bereich
des Menschen
Natur gewandert, hat hier eine neue Gestalt gewonnen
mit ihr
zum Menschen
in
neues
ein
der
Erscheinung,
zurück,
zu
Licht
daß
um auch sein Leben und Handeln
rücken.
der
zur
und kehrt
Er
Mensch
ist
sein
ein
sinnfälliges
eignes
Bild
in
Beispiel
das All
hineinsieht und es, erweitert wie umgewandelt, aus ihm zurückempfängt.
Dem
einen dünkt das ein bloßer Zirkel
der andere erhofft von
und Anthropomorphismus,
solchem Ausgehen und Zurückkehren eine
innere Erweiterung des Menschen.
Zu einem Hauptpunkt der Arbeit machte den Begriff des Gesetzes
erst
die
Neuzeit,
auf die
bemerkenswerte Anfänge
reichen aber in
Der Ausdruck Naturgesetz geht zuerst nicht
das Altertum zurück.
Außenwelt, sondern auf die eigne Natur des Menschen, er
bezeichnet das ungeschriebene Gesetz im Gegensatz zum geschriebenen.
Über den geschichtlichen Ursprung des Terminus Naturgesetz handelt
„Über Begriff und Begründung der sittlichen Gesetze" 1883 (Abh.
der K. Pr. Akad. d. W.) und mit besonderer Umsicht und Gründlichkeit
*
F..
Zeller
R.
Hirzel «ypacpo?
vojj.o?
K. Sachs. Gesellschaft
a'Ypacpo?
vo'[jlos
(Abhandlungen der
der Wissenschaften,
philologisch-histor.
20.
Bd.).
Klasse der
Nach ihm bedeutet
zunächst die altüberlieferte Sitte und Gewohnheit, diese Be-
Gesetz.
155
Für die Natur als Außenwelt verwenden Plato und
Ausdruck nur an vereinzelten
Aristoteles
den
und geben ihm dabei keine
Stellen
technische Zuspitzung, ^ andere Bezeichnungen sind ihnen geläufiger
den
für
Öfter haben den Ausdruck Naturgesetz zuerst die
Begriff.^
wobei
verwandt,
«der Begriff der göttlichen Gesetze war
dem
zu
konnte
religiöse
der Naturgesetze
hinüberleitete"
Vermittlung
zur
Vorstellungen
Stoiker
dienten;
welcher zuerst
es,
Den
(Zeller).
Stoikern
von der Gottheit begründete Gesetz zugleich
das
die
als
Ordnung der Dinge gelten, weil ihnen die Gottheit nicht so-
eigne
wohl eine jenseitige Macht als
Der Ausdruck
bedeutete.
unter den
Schule;
die der Welt
sich
erhielt
die
Grenzen der
Römern hat ihn öfter gleich der erste Philosoph
Der Einbürgerung des Aus-
Lucrez (foedera, foedus, leges naturae).
deutung
innewohnende Vernunft
dann bald
überschritt
durch das ganze Altertum.
Aber daneben kam
(seit
Thukydides) die andere der göttlichen, der ins Herz geschriebenen Gesetze
auf.'
40 heißt es hier von der Revision und Reform der solonischen Ge-
S.
setzgebung, welche Kleisthenes ausführte: «Wie diese Reform nur durch die
Macht des Demos gelang, so diente sie auch dessen- Zielen und Absichten,
und es ist begreiflich, daß von an das Volk von Athen in seinen Gesetzen
das Bollwerk des jungen Staates von Athen erblickte. Von hier an datiert
der Kultus, der mit dem Gesetz und seinem Namen getrieben wurde. Die
Weihe gaben ihm die Taten und der siegreiche Ausgang der Perserkriege"
S. 50.
«Wahrscheinlicher ist, daß der Name (i'Ypatpo? vo[i.O(;) erst im Gegensatz
zum
Y£Ypa(jL[x£voi;
und
aufkam,
Xo'yo;
daß er erst durch diesen
Über den Gegensatz von vo'(jlo?
noch genauere Untersuchung in
sicher,
Gegensatz seine schärfere Bedeutung erhielt."
und «pu'ai;
s,
dort S. 82 ff.
S.
ferner
die
»Themis, Dike und Verwandtes" S. 386
vöaoiv opyava ye'Yovev, oxav atjxa
aXX'
ivavTitüv
15
caelo 268a,
T015
—411.
Die einzigen Stellen sind Plato Timaeus 83 E:
^
xptatv
tov
10 ff:
ü'pKTcat.
[i.^
yäp
xtijv
tou?
izapa
o-fxov
xa^^-a'^p
Ix
ipaut
«puaew?
TTfi
xa\
o{
Xa[xßavT|
8t navta
jj.kv
vdfxou?.
to
HuÖ'aYopEioi,
xeXeuxr yap xal [xeaov xa\ ap/71 xov
xauxa 8k xov xtJ; xpia'8os.
xal xaCta
oiTtwv xa\ ttotcTv nXrj'S'uiJTi xata cpüatv,
Ttäv
ciptO-iiov
Sio x:apa xtJ? ou'aew; £?X7)<pdxei; tSorcep
Arist.
de
xol xa Travra
iyti
xoü rtavxd?,
vo'jjlou? Ixe(vr)?,
xal
Wie vo'fxo; bei den
Philosophen leicht einer- künstlichen Zubereitung angenähert und dem Wesen
Ttpo?
xa?
iytaxeia;
/ptüpieO-a
xwv
9'Etöv
entgegengestellt wird, zeigt u. a.
aXXa $uXov, w? xo
xXfvTjv
xa\
X£')(^V7]V,
x:^v
8'
ou(Tiav
[lIw
xaxa
ouaav
xw
aptö-jAW xouxw.
Aristot.,
au[j.ߣßr]xd(;
IxeJvtjv,
.ri
Phys.
193a,
Ü7:a'p)(^ov,
14:
xi^v
oOx ay yeviid-on
xaxa vdfxov Sta'O^atv
xa\ 8ta[i£V£t Jtolvxa
7i:au)(^oucra
<7\iVf)(iZ<;.
Namentlich gehört hieher avccYxrj (meist im Plural), das sowohl in der
ältp?i-:ri medizinischen Literatur als bei Demokrit, Xenophon (z. B. Memor. I,
*
1,
11),
Plato
(z.
B. leges 967 A), Aristoteles nicht selten
vorkommt.
Es dürfte
der griechischen Forschung der Begriff des Naturgesetzes zuerst in der Astro-
nomie und in der Medizin aufgegangen sein.
Zum Erkenntnisproblem,
156
drucks war förderlich die im späteren Altertum übliche Personifikation
der Gesamtnatur, indem
Regelmäßigkeit ihres Geschehens als
sie die
den Ausdruck eines ordnenden Willens verstehen ließ.
Einen tieferen
Einfluß auf die wissenschaftliche Arbeit erlangte aber der Begriff des
Naturgesetzes im Altertum nicht, namentlich wohl wegen des Über-
gewichts
einer
und
künstlerischen
welche nicht zur Zerlegung
in
Naturbetrachtung,
teleologischen
elementare Vorgänge und zur Er-
mittelung ihrer Regelmäßigkeiten trieb.
Die Kirchenväter nahmen den
Ausdruck auf und verstärkten seine religiöse Färbung, einem Augustin
sind die Naturgesetze bloße Gewohnheiten göttlichen Handelns,
Ge-
wohnheiten, die zu Gunsten besonderer Zwecke jeden Augenblick
verlassen
ander
So stören Wunder und Naturgesetze
werden können.
Im
nicht.
Mittelalter
tritt
ein-
der Ausdruck sehr zurück; Natur-
gesetz (lex naturae) bezeichnet hier das innere Moralgesetz, nicht die
Ordnung der Außenwelt. ^
in
Um so mehr hat die Neuzeit «Naturgesetz"
den Vordergrund gerückt, an kaum irgendwelchem anderen Begriff
hat sie so sehr ihr Selbstbewußtsein gefunden und ihre eigentümliche
Art erwiesen.
Gestaltung
Ein Allgemeineres der Denkweise und die besondere
der Arbeit verstärken einander dabei gegenseitig.
Das
Naturgesetz, als Ordnung des Geschehens, nicht des Sollens, als Aus-
druck
der
Wirkformen
einfachen
der
Elemente,
Neigung der damaligen Menschheit für sich,
hatte
die
volle
weil es ein Verstehen
der Wirklichkeit nicht aus jenseitiger Ordnung,
sondern aus ihrer
eignen Natur versprach, und weil es diese Natur ohne
alle mensch-
und Verfälschung bei sich selbst zu erschließen schien.
Dazu kam das der modernen Wissenschaft eigentümliche Streben nach
liche Zutat
einer neuen, einer exakten Begreifung der Natur durch eine Zerlegung
der Wirklichkeit in kleinste Elemente und eine Durchleuchtung von
Der Ausdruck
^
leges naturae
war so fremdartig geworden, daß der
Beginn der Aufklärung ihn glaubte rechtfertigen und verteidigen zu müssen.
So z. B. Clauberg op. omn. 103
:
Est qui hie
nodum in scirpo quaerat, quod
quae imperant, non efficiunt, quae materiae,
non possunt. Causa autem hujus appellationis
leges sint tantura causae morales,
utpote rationis
experti,'
(Naturae legum)
est in
ferri
propatulo.
ditis Deus leges imposuit morales,
peccant,
ita
voluit res
omnes
Quemadmodum enim rebus ratione praequas abservando bene agunt, transgrediendo
naturales certo semper ordine,
moveri ac quiescere, quas quidem leges ipsae
sariae,
ligiösen
zutage.
non possunt non
observare.
Auch
illae res,
certis legibus
utpote causae neces-
hier liegt die
Bedeutung der re-
Denkweise für die Bildung und Verwendung des Begriffes deutlich
Gesetz.
157
Die völlige Umwandlung des Weltbildes, die daraus hervor-
daher.
ging, hatte drei Hauptsttifen: Analyse, Gesetz, Entwicklung; das
ist
Ge-
der einfachen Wirkformen der Elemente
setz mit seiner Ermittelung
das Rückgrat des Ganzen, erst mit ihm wird eine Präzision der
Erkenntnis erreicht und eine vollständige Unterwerfung der Wirklich-
den Gedanken angebahnt
keit unter
auf
einfache Vorgänge
so
gewährte
die
sie zugleich die
Elemente zu Gunsten
wo
Punkt,
Natur
Wie aber die Zurückführung
zu machen schien,
durchsichtig
Möglichkeit neuer Kombinationen der
Das Gesetz
menschlicher Zwecke.
ist
der
das Streben nach engster Verbindung von Theorie und
Praxis, das der neueren
Forschung von Anfang an innewohnt sich
Erkennens
Denn hier wird der Endpunkt des
zum Ausgangpunkt des Handelns; aus vereinzelten und
zufälligen
Funden
in
wirksame Arbeit umsetzt
ist
die
Technik
eine
selbständige
und
lebens-
umspannende Macht nur mit Hilfe der Gesetze geworden. So laufen
bei ihnen alle Fäden zusammen als dem Mittelpunkt der geistigen
Arbeit;
sie
bilden den klarsten Ausdruck des modernen Verlangens
nach immanenter und sachlicher Erklärung, nach analytisch-präziser
Begreifung, nach einem aktiveren Verhältnis des Menschen zu seiner
Naturumgebung.
Aber zugleich war das Gesetz in dem neuen Sinne eine schwere
Aufgabe und eine Sache mannigfachster Verwicklung.
Im Streben
nach Gesetzen verschlingt sich eng Erfahrung und Vernunft
Regel-
mäßigkeiten werden entdeckt, und es entsteht eine große Freude daraus, daß was im ersten Eindruck wirr durcheinander läuft, einer schärferen
Betrachtung geordnete Reihen zu erkennen gibt
Aber man verbleibt
mehr oder weniger verwickelter
Tatsachen, man möchte diese zerlegen und auf einfache, letzte, allnicht
bei der bloßen Konstatierung
gegenwärtige Elemente zurückführen, zugleich aber statt eines bloßen
Nach- und Nebeneinander einen kausalen Zusammenhang erreichen;
man
strebt
von empirischen zu
rationalen,
erklärenden Gesetzen, die notwendig
und
von beschreibenden zu
allgemeingültig sind.
Nur
solche rationale Gesetze dürfen ausschließlich herrschen wollen, sie
können keine Ausnahmen,
also auch nicht das Wunder dulden.
werden nach möglichster Einfachheit streben und
keit
als
suchen.
Ausdruck eines durchgehenden
alle
Sie
Mannigfaltig-
Geschehens zu verstehen
Auch werden diese Gesetze auf einem
präzisen Ausdruck,
einer bestimmten Formel bestehen, da nur eine solche zur Beherrschung
des Tatbestandes zu führen vermag.
Diesen präzisen Ausdruck gibt
Zum Erkenntnisproblem.
158
dem Naturgesetz vornehmlich
So konnte Newton
die Mathematik.
Aufgabe echter Naturforschung darin
die
unter Verzicht auf
setzen,
Formen und die verborgenen Qualitäten die Naturerscheinungen auf mathematische Gesetze zurückzuführen, ^ und Kant
die substantiellen
behaupten, „daß in jeder besonderen Naturlehre nur so viel eigentliche
Wissenschaft angetroffen werden könne, als darin Mathematik anzutreffen
ist"
(IV,
360 Hart).
Aber solche Wendung
schwere
stellt
Aufgaben und legt manche Irrungen nahe. Oft wird bloß empirischen
Verallgemeinerungen
beigelegt,
was
den
nur
Gesetzen
strengster
Fassung zukommt; kaum hat jemand mehr von der Ausnahmslosigkeit,
der Unveränderlichkeit der Gesetze gesprochen
dem
sie
nur
eine
Beschreibung
derselbe Comte,
als
Erfahrung
der
bedeuten
sollten.
Zugleich gibt die bloße Regelmäßigkeit sich leicht wie einen völligen
Abschluß, die Aufgabe scheint gelöst,
wo
sie
eben nur bezeichnet
Der Begriff des Gesetzes hat oft dogmatisierend gewirkt, nirgends
ist.
dem Gebiet
der Biologie,
wo
Erscheinungskomplexe sich
den Anschein
strenger Gesetze
mehr
als
auf
und deren Forderungen
sehr
oft
verwickelte
gaben
stellten.
Zu solchen Verwicklungen aus der Durchführung der weiteren
kommen Störungen aus einem mehr oder minder versteckten
Fassung
Fortwirken des älteren
Gesetzesbegriffes
ein überlegendes Wollen.
1 8.
mit seiner Beziehung auf
Das geschieht, wenn Denker des
1 7.
und
Jahrhunderts aus der Gesetzlichkeit der Natur eine gesetzgebende
Gottheit glauben
wenn
in
erschließen
zu können.
umgekehrter Richtung
eine
Aber
es
geschieht auch,
pantheistische
Denkweise die
Gesetze wie lebendige Mächte behandelt und
sie
als
einen Gegen-
stand andächtiger Verehrung an die Stelle der Gottheit
setzt, ^
Auch
darin steckt ein Fortwirken der älteren Art, daß das Gesetz oft wie
über den
eine
^
S.
den
einzelnen
Beginn
der
Vorgängen schwebende Macht behandelt
philosophiae
naturalis
Missis formis substantialibus et qualitatibus occultis
principia
mathematica:
phaenomena naturae ad
leges mathematicas revocare.
So zieht sich von Giordano Bruno durch die Neuzeit bis zur GegenBruno sucht das Höchste in
inviolabili intemerabilique naturae lege, in bene ad eandem legem instituti
^
wart ein gewisser Kultus des Naturgesetzes.
animi religione (de universo
et
immenso 653).
Je skeptischer sich heute die
Menschen zur Religion verhalten, desto blindere Verehrung pflegen sie dem
Naturgesetz zu zollen.
Gesetz.
wird,
EndUch gehört auch das
ihnen die Bahn vorschreibt. ^
die
daß,
hieher,
I59
kecker
je
Gesetzesformel
auftritt,
Behauptung eines Gesetzes und
die
sie
desto leichter Eingang findet.
an einem Gegegen
den
Geist der
Sünde
eine
sache pflegen wir zu prüfen, ehe wir sie anerkennen
zu
setze
zweifeln,
das scheint
So überträgt
Wissenschaft.
ein praktisches Gebot hat, zu
einer
Eine Tat-
die
sich
Autorität,
die
;
das Gesetz als
Unrecht auf das Gesetz des Geschehens:
auch für dieses wird schleunige Zustimmung verlangt und
keinerlei
Widerspruch geduldet. Wie hätte ohne solchen kritiklosen Respekt das
berüchtigte «eherne Lohngesetz" eine so große Rolle spielen können!
Namentlich die Formel wirkt dabei Wunder.
Mahhus die Geister bewegt, wenn
rungszunahme
nicht
von der Bevölke-
mathematische Formel gegeben
bekannte
die
Wie viel weniger hätte
er seiner Lehre
«Man liebt die Sicherheit", so klagte schon Pascal; man hält
hätte!
aber leicht für sicher, was keck und selbstbewußt
Aber
so
mißlich
das
sein
alles
mag,
auftritt.
menschliche
solche
Irrungen sind eine unvermeidliche Begleiterscheinung jeder großen
Wendung, an den Gesetzen
selbst
Überblicken
die
wir
also
rasch
dürfen
sie
intellektuelle
nicht
irre
Bewegung,
machen.
der
die
Kampf um das Gesetz in der Neuzeit hervorgerufen hat und unablässig hervorruft.
b)
Der Kampf um das Gesetz in der Neuzeit.
Die
Naturgesetze
haben
ihre
eigentümliche
Gebiet der unorganischen Natur gefunden;
Ausprägung im
so war es eine Über-
tragung der Größen und Methoden dieses Gebietes, welche das Vordringen des Gesetzesbegriffes
mußte aber
selbst
— früher oder
in
andere Disziplinen begleitete.
später
— bemerklich werden, was
an Problemen und auch Schranken
liegt.
Dabei
ihm
in
Beim Gesetze
ist
alle Aufmerksamkeit den Formen des Geschehens zugewandt, die Kräfte
und Ursachen bleiben im Hintergrund; wird solche Zurückstellung
überall
^
möglich
sein,
und wird das Hervortreten
dieses
Mit Recht bemerkt darüber Sigwart (Logik II,', 512):
Problems
„Eine
leere
von Naturgesetzen so zu sprechen, als ob die bloße
Formel eine magische Macht über die Erscheinungen übte und ihnen etwas
zumutete, was nicht aus ihrer eignen Natur von selbst folgte. Gesetze können
rhetorische Phrase
ist
es
Gründe des wirklichen Geschehens sein, sondern nur die konstante Art
und Weise ausdrücken, wie reale Dinge sich verhalten."
nie
Zum Erkenntnisproblem.
160
Ganzen verändern? Beim Gesetze wird die
Wirklichkeit in lauter einzelne Vorgänge zerlegt und alles beherrschende
nicht den Anblick des
Ganze entfernt;
jedes
bildet
das für
sollte
alle
Beim Gesetze
Gebiete taugen?
Geschehen nur einen Sonderfall
einzelne
eines
allge-
meinen, alle Individualität wird hier für die Wissenschaft ausgeschaltet;
wird das Individuelle sich überall mit einem so bescheidenen Platz
begnügen, wird es nicht
aller
versuchten Gleichmachung eine
namentlich
will,
wenn es nicht bloß
das Geschehen
als
auch erklären
beschreiben, sondern
determiniert
völlig
un-
Endlich erscheint beim Gesetze,
vergleichliche Art entgegensetzen?
und unweigerlich
fest-
gelegt; für freie Entscheidung, für eine Wahl zwischen verschiedenen
Möglichkeiten
gibt
keinen
hier
es
Platz.
Werden
dem
alle
einzelnen
hin-
sich
Lebensgebiete fügen?
Probleme also
Hülle und Fülle, durch
in
alle
durch aber das Gesamtproblem, wie weit die mechanischen Naturbegriffe
die
ganze Wirklichkeit unter sich
zu
bringen
vermögen.
Der Widerstand gegen den Gesetzesbegriff kann dabei eine schroffere
und eine mildere Form annehmen: entweder wird der Gesetzesbegriff für ein besonderes Gebiet ganz und gar abgelehnt, oder er
wird, unter Ablösung von der naturwissenschaftlichen Fassung, dessen
Aus beidem zusammen erwächst eine höchst
Besonderheit angepaßt.
lebhafte Bewegung, die nicht
wenig dazu beigetragen hat, die Eigen-
tümlichkeit der einzelnen Gebiete in klares Licht zu stellen.
Es beginnt aber die Ausbreitung des Gesetzesbegriffes über die
Natur hinaus schon im
1 7.
Jahrhundert, namentlich findet er schon
Das 18. Jahrhundert setzt die
Bewegung fort und führt sie tiefer in die einzelnen Gebiete ein.^
Ihren Höhepunkt aber erreicht jene erst im 1 9. Jahrhundert, nament-
damals Eingang
lich
in
die Psychologie.
in seiner zweiten
Hälfte.
Es wirkte hier manches dahin zusammen, das Gesetz
Mittelpunkt der wissenschaftlichen Arbeit zu rücken.
^
Mit besonderer Energie
fochten.
Er sagt gleich
la signification la
la
in
hat
Montesquieu den
les
lois,
ont leurs
les intelligences
lois,
superieures ä
l'homme a ses lois.
den
les lois,
ver-
dans
rapports necessaires qui derivent de
nature des choses; et dans ce sens tous
a ses
Gesetzesbegriff
den Anfängen seines esprit des lois:
plus etendue, sont
in
Vor allem das
les etres
ont leurslois:
l'homme ont
leurs
lois,
la divinite
les
betes
Und etwas weiter: il y a donc une raison
primitive, et les lois sont les rapports qui se trouvent entre eile et les diffe-
rents etres, et les rapports de ces divers etres entre eux.
Gesetz.
Selbständigwerdeii
fernerhin
von
wollten, desto
der
der
Wissenschaften.
einzelnen
Philosophie
151
und
Prinzipien
weniger
Je
Regeln
sie
entlehnen
mehr mußten sie darauf bedacht sein, in ihrem eignen
feste Zusammenhänge zu finden.
Gebiet durchgehende Begriffe und
In
den Gesetzen aber hofften
sie
solche zu finden;
schien der unermeßliche Stoff sich ordnen
schien auch
mit ihrer Hilfe
und gliedern zu
lassen,
eine Vergleichung verschiedener Reihen und Gruppen
des Geschehens möglich zu werden.
Diese
Bewegung
erhielt eine
besondere Spannung durch das Verhältnis von Natur- und Geisteswissenschaften.
Die glänzenden Erfolge der Naturwissenschaften haben
auch ihre Expansionskraft, ihr Streben nach Beherrschung des ge-
samten
intellektuellen
Reiches gesteigert, namentlich scheint die Ent-
wicklungslehre Weltbegriffe zu
liefern,,
denen kein Gebiet
sich ent-
ziehen kann; so dringt die naturwissenschaftliche Denkweise mit ihren
Begriffen tiefer und tiefer in die anderen Gebiete hinein.
Aber zugleich
werden diese zum Widerstände gereizt und zur Besinnung auf ihre
Eigentümlichkeit getrieben,
es
entsteht
ein
Kampf, dessen
lebhafter
Fortgang die Unterschiede immer mehr zum Bewußtsein gebracht hat.^
*
Ein anschauliches Bild von der Bewegung in der Sprachwissenschaft
liefert B.
Delbrück in der Abhandlung ,,Das Wesen der Lautgesetze" (Annalen
der Naturphilosophie I, 277 ff.).
Nachdem schon Fr. Schlegel und Bopp die
Sprachwissenschaft in Vergleich mit
der
Naturwissenschaft
gestellt
hatten,
ohne jedoch sie zu den Naturwissenschaften zu rechnen, gab Schleicher der Sache
Sein Bekenntnis geht dahin (Die Darwinsche
eine schärfere Zuspitzung.
Theorie und die Sprachwissenschaft,
S. 7): „Die Sprachen sind Naturorganisohne vom Wollen des Menschen bestimmbar zu sein, entstunden,
nach bestimmten Gesetzen wuchsen und sich entwickelten, und wiederum
altem und absterben; auch ihnen ist jene Reihe von Erscheinungen eigen,
men,
die
die,
man unter dem Namen „Leben" zu
verstehen pflegt.
Die Glottik, die
demnach eine Naturwissenschaft,
ihre Methode
im Ganzen und Allgemeinen dieselbe, wie die der übrigen Naturwissenschaften."
Demgegenüber haben, wie Delbrück näher ausführt, andere
Forscher, unter ihnen namentlich Whitney, zur Geltung gebracht, daß sich
bei der Entstehung und Veränderung von Sprachen nirgends dem Sprachstoff
innewohnende Lebensgesetze, sondern immer nur menschliche Handlungen
finden. Als ein solcher Ausfluß menschlichen Handelns und Wollens ist die
Wissenschaft der Sprache,
ist
ist
Sprache kein Naturorganismus, sondern eine Institution,
stitutionen,
welche die menschliche Kultur ausmachen.
eine von den In-
Damit muß auch
das Sprachgesetz anders gefaßt werden
als das Gesetz eines Naturorganismus.
Delbrück kommt in seiner Untersuchung der Lautgesetze zu dem Ergebnis,
daß, so sehr diese sui generis sind, kein Grund vorliegt, ihnen deswegen
den
Namen von Gesetzen abzusprechen.
Eucken, Grundbegriffe. 4. Aufl.
„Denn wir
verstehen
H
auch
bei
Zum Erkenntnisproblem,
162
Von der Welt des Leblosen ausgegangen, hatte das Naturgesetz
zunächst des Reichs des Lebendigen zu bemächtigen; auf wie
sich
viel
in
"Widerstand es dabei
vollem Gange
ist,
stieß,
und wie der Kampf eben jetzt wieder
das hat uns im vorigen Abschnitt beschäftigt.
Die Übertragung des Naturgesetzes auf die Seele war schon durch
nahegelegt
Descartes
geführt;
und
Seelenleben
alles
wurde
durch
verwandelt
Spinoza
großartig
hier
sich
in
ein
aus-
Gewebe
von Einzelvorgängen, die ganz nach Art der mechanischen Natur
sich
bewegen und wirken.
Monade ihren eignen
Leibniz läßt jede
und unterscheidet von den «physiko-mechanischen"
Gesetzen folgen
Gesetzen der Körper die »ethiko-logischen" der Seelen (736 b, Erdm.).
Psychologische Gesetze in strengerem Sinne haben
namentlich die
Engländer aufgebracht,
in
so
die
Assoziationsgesetze;
Deutschland
geht die Bewegung über Wolff zu Herbart fort, der die mathematische
auch
Formel
in
das
Innere
des
einführen
Seelenlebens
möchte.
Zugleich aber fehlt es nicht an Männern, welche die eigentümliche
Art des Seelenlebens mit seiner inneren Einheit, seiner Beweglichkeit,
und damit auch dem,
seiner Individualität zur Geltung bringen
was hier
weisen
sich
an Gesetzen aufstellen
läßt,
deutliche
eine
Entscheidend für die Behandlung der inneren Welt
Stellung der Gesetze in ihr
eine
Grenze
1
neue Stufe
wird oder
nicht.
ist
die Frage,
ob
in
und
die
dem Geistesleben
und selbständige Art der Wirklichkeit anerkannt
Wo jenes geschieht, kann über einen wesentlichen
Unterschied von allem naturgesetzlichen Geschehen keinerlei Zweifel
sein.
Die Naturgesetze sind die rein gefaßten Wirkformen des Ge-
schehens, schlichte und einfache Tatsächlichkeit.
Nun muß auch das
Gesetz geistiger Art in irgendwelcher Tatsächlichkeit wurzeln; haltlos
anderen Wissenschaften unter Gesetzen nichts weiter,
als
den Ausdruck für
Gleichmäßigkeiten, welche zwar im Einzelfalle nicht rein hervortreten, von
denen wir aber annehmen, daß sie stets rein hervortreten würden, wenn im
Einzelfalle alle anderswoher kommenden störenden Einwirkungen entfernt
werden könnten" (308).
^ So bemerkt Sigwart, dessen Untersuchungen über alle diese Probleme
besonders klar und eindringend sind, über die Assoziationsgesetze (Log. IP,
553): „Die Assoziationsgesetze deuten nur bestimmte Richtungen an, in denen
unsere Reproduktionen verlaufen können,
oder in vielen Fällen
verlaufen,
bestimmte Tendenzen der wirklichen Aneinanderreihung von Bildern oder
Wörtern u. s. w.; Gesetze, aus denen jeder wirkliche Vorstellungsverlauf als
notwendig nachgewiesen werden könnte, vermögen sie nicht darzustellen."
Gesetz.
in
der Luft schwebende Gesetze, die doch eine Wirkung tun sollen,
Aber das Geistesleben, das den unentbehrlichen
sind ein Unding. ^
Halt gewährt,
es
I53
wenn auch zu seinem
für ihn,
ist
gleich
keineswegs ein voller Besitz des Menschen, sondern
ist
hohes
ein
Damit werden
Ideal.
gehörig, zu-
die Gesetze zu Normen, die nicht wirkungslos
die aber Widerstand finden
sind,
Wesen
innersten
schwere Aufgabe, zugleich Natur und
eine
Ziel,
und
gegen ihn durchsetzen
sich
müssen. ^ Je nach der Eigentümlichkeit des intellektuellen, des ethischen,
des ästhetischen Gebietes gestaltet sich der Widerstand und überhaupt
der Lebensprozeß verschieden, wie wir
Nur
Sittengesetz
Dies Problem
hier
sei
mit
einigen
er
die
Moral
über
Worten
alles
das
Sittliche
und
eines sicheren
trieb
ihn,
einseitigen
Aber diesen berechtigten Gedanken hat
dadurch abgeschwächt.
rechnet, der gibt
und
Wer die
das
gestellt
das
Moral zur Natur
Schleier-
der
Charakteristische
des
Moral
Menschen
dem Begriff der Natur einen neuen Sinn und hat
von allem bloßen Dasein zu scheiden; so
Kant in besserem Recht als Schleiermacher.*
*
Anblick
Haltes in der menschlichen Natur beraubt;
überspannt
stark
sie scharf
einen
vom Natur-
Einem Schleiermacher
den engen Zusammenhang von Natur- und Sittengesetz
zu verfechten. 3
macher
unter
Getriebe
seelische
hinaushob, mußte sich das Sittengesetz mit seinem Soll
gesetz bis zu schroffem Gegensatz abheben.
schien damit
gestreift.
den Vordergrund ge-
namentlich durch Kant in
ist
Denn indem
bracht.
wissen.
das vielbehandelte Problem des Verhältnisses von Natur-
und
gesetz
alle
ist
schließlich
Die unmittelbare Gleich-
Mit Recht macht Husserl bemerklich, daß „jede normative und a fortiori
jede praktische Disziplin eine oder mehrere theoretische Disziplinen als Funda-
mente voraussetzt,
ablösbaren
in
dem Sinne nämlich, daß sie einen von aller Normierung
Gehalt besitzen muß, der als solcher in irgend-
theoretischen
welchen, sei es schon abgegrenzten oder noch zu konstituierenden theoretischen
Wissenschaften seinen natürlichen Standort hat" (Logische Untersuchungen 1, 47).
S.
auch
S.
164:
„Der Gegensatz von Naturgesetz
Regel eines tatsächlichen Seins und Geschehens
als
ist
empirisch begründeter
nicht das Normalgesetz
sondern das Idealgesetz im Sinne einer rein
als Vorschrift,
(Ideen, reinen Gattungsbegriffen)
in
den Begriffen
gründenden und daher nicht empirischen
Gesetzlichkeit."
' Von neueren Untersuchungen darüber sei namentlich erwähnt Windelbands Abhandlung ,, Normen und Naturgesetze" in den „Präludien".
» S. Werke zur Philos. II, 397-417.
*
Zeller
„Über Begriff und Begründung der
sittlichen
Gesetze" (1883)
kommt hinsichtlich der Ethik zu dem Schlußergebnis: „Ihre Sätze sind nicht
11*
Zum Erkenntnisproblem.
164
Setzung von Natur- und Sittengesetzen entspricht
antiken Ethik; sie
hinfällig
ist
Geistesleben hat schwere Verwicklungen erkennen lassen.
zu zeigen, daß,
leicht
faßten,
der Verlauf
immer wieder zur Anerkennung
zwungen hat.i
sie
Die dem
liche
Auch wäre
wo immer moderne Denker die Sittengesetze
Naturgesetze
prinzipiell als
Stande der
nachdem das Verhältnis des Menschen zum
Lage,
geschichtlichen
dem
geworden und widerspricht der welt-
einer
der Untersuchung
abweichenden Art ge-
Jahrhundert eigentümliche geschichtlich-gesellschaft-
1 9.
Denkweise mußte das Streben erzeugen, sowohl das gesellschaftgeschichtliche Gebiet
liche als das
zu unterwerfen.
festen Gesetzen
Das Drängen auf Präzision vornehmlich unterscheidet
Gesellschaftslehre, die Soziologie,
großen Zahl
der
Hilfe
wird
die
moderne
von allen früheren Versuchen.
Mit
der
Er-
das Zufällige
individuellen
scheinungen ausgeschieden, werden Durchschnitte ermittelt und die
Grenzen
abgesteckt,
innerhalb derer etwaige Abweichungen liegen,
werden Regelmäßigkeiten innerhalb des sozialen Gebietes aufgedeckt ^
Je
mehr aber
welche das
Forschung die Überraschung überwunden
die
erste Gewahren
hat,
von Regelmäßigkeiten innerhalb eines bis
dem Zufall preisgegebenen Gebietes bewirkte, desto
man gegenüber dem Begriff der Gesetze geworden, desto
dahin scheinbar
kritischer
ist
deutlicher hat sich der Unterschied zwischen bloßen
gesellschaftlichen
Tendenzen des
Lebens und eigentlichen Naturgesetzen herausgestellt.
Noch mehr Bewegung hat der Begriff auf dem wirtschaftlichen
Gebiete engeren Sinnes hervorgerufen; nirgends hat der Streit darüber
mehr Folgen
für
das
Handeln
und Leben
als
hier.^
Denn das
der Ausdruck dessen, was irgendwo als Recht oder Sitte besteht, sondern
der Forderungen, die als Normen der menschlichen Willenstätigkeit aus der
Idee des Menschen hervorgehen." S. auch Siebeck: „Über das Verhältnis
von Naturgesetz und Sittengesetz" (Philos. Monatshefte 1884, S. 321 ff.).
^ Comte liefert dafür ein hervorragendes Beispiel.
Der große Empirist,
der grundsätzlich
will,
die Gesetze
nur
als
Beschreibungen
verstanden
wissen
meint bei der Wendung zur Gesellschaft (cours de phil. pos. IV., 466):
Cette generalite empirique, qui en toute autre science pourrait dejä avoir
une valeur süffisante, ne saurait pleinement convenir ä la nature propre de la
sociologie.
'
In
dem
allen
nimmt
bekanntlich Quetelet eine hervorragende Stel-
lung ein.
'
Zur Geschichte des Begriffes bemerkt Neumann, dem wir besonders
verdienstliche Untersuchungen über diesen Gegenstand verdanken (Jahrbücher
für Nationalökonomie
und Statistik, 3. Folge 1899, S. 152/3): „Geforscht hat
Gesetz.
Problem des Gesetzes
steht
165
unmittelbarem Zusammenhange mit
in
wie sich der Staat zu den wirtschaftlichen Bewegungen
der Frage,
zu verhalten habe, ob bloß zuschauend oder selbsttätig eingreifend.
der
Bildete
Prozeß
wirtschaftliche
bloßes
ein
Gewebe
sich
selbst
regulierender Einzelbewegungen, so erschien alles Eingreifen als eine
Störung,
das laissez
scher
Weisheit
selbst
etwas
In
laissez
faire,
Wahrheit
anderes
als
ist
ein
aller
ein
bloßer
wurde zur Summe
solches
Denn neben
Naturprozeß.
jenem Gehenlassen stehen andere Möglichkeiten,
Boden der Geschichte gegen
politi-
Gehenlassen an sich
muß auf dem
es
andersartige Zustände erst durchgesetzt
werden, es wirkt, wenn eingeführt, nicht wie selbstverständlich
sondern es läßt sich zurücknehmen, es
Willen getragen werden.
fort,
muß von einem fortlaufenden
Dazu ist der Glaube an
eine Selbstregu-
lierung der wirtschaftlichen Verhältnisse durch die natürlichen Triebe
und Kräfte der Individuen nicht möglich ohne einen
Glauben an
optimistischen
die Vernunft der gesellschaftlichen Verhältnisse; jede Er-
schütterung dieses Optimismus untergräbt auch den Glauben an das
Allvermögen der Naturgesetze.
Nun haben die wirtschaftlichen Ver-
wicklungen des 19. Jahrhunderts jenen Optimismus aufs schwerste erschüttert, sie drängen
immer stärker zum Eingreifen des Staates in den
wirtschaftlichen Prozeß, sie entwinden damit das Gebiet
den bloßen
Naturgesetzen und verstärken die Bedeutung der ethischen wie der
historischen
historischen
man nach
Elemente.^
Die Anerkennung dieses ethischen und
Moments verhindert keineswegs die Anerkennung wirt-
wirtschaftlichen
zeigen versucht
ist
und
sozialen Gesetzen, wie an
(So zuletzt in
dem Aufsatz:
anderem Ort zu
Wirtschaftl.
Gesetze nach
und jetziger Auffassung. Jahrb. für Nationalökonomie und Statistik.
N. F. 1898, Bd. 16), schon im Altertum und sodann, nach den allerdings auf
anderen Gebieten erzielten Erfolgen Bacons und Newtons, namentiich seit
der zweiten Hälfte des 17. Jahrhunderts, seit Locke und Hobbes, von denen
der Erstere auch bereits den Ausdruck law hierfür gebrauchte, während
gerade die Physiokraten da sie diesen Vorgängen folgten, nicht ganz frei
von der Schuld zu sprechen sind, daß sie unter dem Einfluß damals allmächtiger naturrechtiicher Voretellungen die in Rede stehenden Gesetze des
früherer
,
Geschehens von
solchen des Sollens oder den ethischen Gesetzen nicht
ausreichend zu trennen wußten."
^
Erwähnung verdient
dabei,
daß nicht bloß auf
individualistischer,
sondern auch auf sozialistischer Seite Neigung bestand, den Begriff des Gesetzes auf
Kosten der freien Tat zu überspannen.
seit
alles
Nur erhielt die Sache hier
daß eine Gesamtbewegung des weltgeschichtlichen Lebens, jenWollens und Tuns der Individuen, mit unabwendbarer Dialektik
die Gestalt,
Zum Erkenntnisproblem.
166
Aber sie entsprechen dann
schaftlicher Gesetze.
den
einfach
nicht
sondern sind nach Neumanns Definition
Naturgesetzen,
»der Aus-
druck für eine infolge der Macht wirtschaftlicher Zusammenhänge
aus gewissen Motiven sich ergebende regelmäßige Wiederkehr wirtschaftlicher
Erscheinungen (Tendenzen oder Vorgänge)".^
Am meisten Bewegung hat in neuester Zeit das Problem der
mehr und mehr
geschichtlichen Gesetze hervorgerufen,
um
Mittelpunkt des Kampfes
geworden.
Je
mehr
zum
es
die Gesamtauffassung der Geschichte
Beginn
seit
ist
der
Neuzeit
überkommene
die
supranaturale Vorstellung von der Geschichte wich, desto mehr mußte
dazu
es
drängen,
innerhalb
ihres
Bereiches durchgehende
eignen
Bewegungen und feste Regelmäßigkeiten aufzuweisen. Die Aufklärung
solchem Verlangen ihren eigentümlichen Stempel
prägte
Ihre
auf.
Geschichtsforschung „zertrümmerte das bisherige Geschichtsbild, wie
es
an den
Monarchien,
danielischen
Augustin orientiert war,
sie
der Apokalypse
an
oder
an
deckte eine bisher ungekannte oder un-
beachtete Welt auf, eröffnete unberechenbare Zeiträume der Geschichte,
den Sündenfall von der Spitze
verwies
konstruierte einen
weg und
der Geschichte
ganz anderen Urzustand
Ausgangspunkt.
als
—
Indem aber diese von Wunder und Vorsehung absehende Erklärung
ein
unendlich
verworrenes
Spiel
menschlicher
aufdeckte,
Kräfte
empfand man zugleich doppelt das Bedürfnis nach einem einfachen,
normalen Gehalt der Geschichte, den man
rechts,
den Ideen des Natur-
in
der natürlichen Moral und Religion fand."^
Hatte die Philo-
sophie zunächst die Neigung, die Geschichte der Vernunft entgegenso
zusetzen,
erwuchs bald ein Streben,
in
nunft und zugleich eine Gesetzlichkeit der
ihr
eine
gewisse Ver-
Bewegung aufzudecken. ^
Leibniz namentlich verficht die Idee einer durchgehenden Kontinuität
große Wendungen und Umwälzungen hervortreibt.
So hat es namentlich
Marx in engem Anschluß an die Hegeische Geschichtsphilosophie ausgeführt.
Aber auch hier entsteht der Widerspruch, daß eben das, was aus
Karl
gesetzlicher
Notwendigkeit
hervorgehen
soll,
zu
seinem
vollen
Siege
der
Anerkennung durch den Menschen, der Aufnahme in die eigne Überzeugung
Nicht zu
bedarf.
ruhiger Kontemplation,
sondern
zu energischer Aktion
wird eben hier der Mensch aufgerufen.
^
S.
«Naturgesetz
und Wirtschaftsgesetz"
Staatswissenschaft 1892, Heft
*
S.
Tröltsch,
(Zeitschrift
für
die
gesamte
3).
Real-Encyklopädie für Theologie und Kirche.
3.
Aufl.
unter „Aufklärung" S. 231.
*
in der
S.
über diese Fragen meine Behandlung der Philosophie der Geschichte
„Kultur der Gegenwart".
Gesetz.
167
der geschichtlichen Be^x'egung, Vico den Gedanken einer regelmäßigen
Folge bestimmter Stufen in der Entwicklung der Völker und Epochen,
immer stärker wird das Verlangen nach einer durchgängigen VerDas
kettung der Ereignisse zum Zusammenhang eines Ganzen.
19.
indem
Jahrhundert führt die Sache erheblich weiter,
Typen der Gesamtauffassung der Geschichte ausprägt,
scharfe
unermeßlichen Ausdehnung der Forschung
der
sowohl
es
mäßigkeiten aufdeckt.
gesetzten
Jenes geschieht
empirische
den
in
als
in
Regel-
entgegen-
zugleich
und nahe verwandten Systemen Hegels und Comtes.
Dort
eine allumspannende Logik, hier eine langsame Anhäufung, der ein-
zelnen Elemente,
dort
eine
Bewegung durch
schroffe
Gegensätze,
hier ein ruhiges Aufsteigen, das aber deutlich drei Hauptstufen (trois
erkennen
etats)
hier wie dort ein
läßt,
Ausschluß
aller
Willkür, ein
sicheres Fortschreiten, eine völlige Bestimmtheit aller Mannigfaltigkeit
durch den jeweiligen Stand der Gesamtbewegung.
Geschichte
überströmenden Reichtum
leicht
ihren
preßte
und alle Irrationalität wegzudeuten
der anderen Seite her
Forschung
Wurde
so
der
von der Philosophie eine Gesetzlichkeit zugeführt, die
beflissen
Durchdringung des
die
zu engen
einen
in
Rahmen
war, so hat
Stoffes
mittels
von
der
reichem Maße empirische Regelmäßigkeiten aufgedeckt.
in
Dabei wirkten
tief
in die
Forschung hinein die großen Gegensätze
Den gesetzlichen, ja speziell naturgesetzlichen
des modernen Lebens.
Charakter der Geschichte verstärkte die wachsende Einsicht in die
Abhängigkeit des menschlichen Befindens und Tuns von äußeren Bedingungen, es verstärkte ihn weiter die Erkenntnis der Abhängigkeit
vom Ganzen, vom sozialen „Milieu"; es widersprach
der Individuen
ihm die der Aufklärung entgegengehaltene Individualität und Positivität
der Geschichte,^ es widersprach ihm nicht minder die Betonung der
So vertritt
*
zeugung,
ihr
»daß
seinen
in
z.
B. Steffensen
in der
(Gesammelte Aufsätze,
S. 278)
die Über-
Geschichte das durch und durch Indixaduelle, das in
Formen,
höchsten
in
Persönlichkeiten
willenskräftigen
und
großen Taten und
Leiden eines wahren Werdeprozesses offenbar wird, den unvergleichlichen
Gesellschaften,
zu oberst in der Menschheit
selbst,
in
Reiz hervorbringt, den das geschichtliche Wissen für den menschlichen Geist
hat";
sowie daß
„nicht die
pirischer Naturgesetze,
Bestätigung von der Geltung allgemeiner em-
sondern
sammenstoßen der höchsten
idealen
Gesetzen,
besser
Gottes widerspiegeln",
es
vielmehr
das
so
irdischen
Natur,
mit
idealen
Mächten,
ist,
„was uns
in
des
ganz unverkennbare Zuinneren Menschen, mit
welche
dem
die
Unbedingtheit
dramatischen,
Gang des geschichtlichen Lebens die Seele erschüttert."
tragischen
Zum Erkenntnisproblem.
168
großen Persönlichkeiten, wie sie in Carlyle einen besonders prägnanten
Ausdruck gefunden hat.
Es ist nicht bloß die verschiedene Schätzung
von Natur und Geist
unserer Wirklichkeit,
dem Inhalt
die Frage nach
Problem
das
in
der
des Geisteslebens,
oder
Rationalität
es
es
Irrationalität
ist
minder
nicht
ist
im besondern
unseres
Daseins,
welche die Frage nach der Gesetzlichkeit der Geschichte verschieden
beantworten lassen.
Diese
Gegensätze
methodologischen
und
vielfach
Klarheit
neuerdings Windelband
hat
der
Behandlung des
die
Forscher
und
hier das Einzelne
den Unterschied
geschichtlicher
von
Behandlung zum
in
natur-
Ausdruck
der geschichtlich bestimmten Gestalt gesucht,
immer sich gleichbleibende Form,
der einmalige,
hier
bestimnite Inhalt des wirklichen Geschehens
wissenschaftliche
bilden
bewegt
Dort wird das Allgemeine in der Form des Naturgesetzes,
gebracht. ^
sich
in
heute
auch
welches
Mit besonderer Energie und einleuchtender
entzweit
wissenschaftlicher
dort die
erscheinen
Problems,
—
darf
idiographisch
Denken
ist
dem
einen
in
— wenn man neue Kunstausdrücke
Falle
26)"; ,;den festen
(S.
in
«Das
betrachtet.
nomothetisch, in dem andern
Rahmen unseres Weltbildes gibt
jene aligemeine Gesetzmäßigkeit der Dinge ab,
welche,
über allen
Wechsel erhaben, die ewig gleiche Wesenheit des Wirklichen zum
Ausdruck
und innerhalb
bringt;
dieses
Rahmens
sich
entfaltet
der
lebendige Zusammenhang aller für das Menschentum wertvollen Einzelgestaltungen ihrer Gattungserinnerung" 2 (S. 38).
Diese Überzeugung
in geistvoller und selbständiger Weise weitergeführt;^
ist
von Rickert
sie
hat überhaupt viel literarische Bewegung hervorgerufen.
Gegen-
über solcher Richtung auf das Individuelle verficht Lamprecht die
Überzeugung, daß das Individuelle nur der künstlerischen Erfassung
zugänglich
so
auch
sei,
in
und daß das
der
Geschichte
S.
^
Es hatte auch schon Paul
dort
Denken wie
überall,
nur auf das Typische gehen könne;
„Geschichte und Naturwissenschaft", Rektoratsrede 1894.
^
„Geschichtswissenschaften"
sagt
wissenschaftliche
(S.
1):
in seinen Prinzipien der Sprachgeschichte
und „Gesetzeswissenschaften" unterschieden;
er
„Wie jedem Zweige der Geschichtswissenschaft, so muß
auch der Sprachgeschichte eine Wissenschaft zur Seite stehen, welche sich
mit den allgemeinen Lebensbedingungen des geschichtlich sich entwickelnden
Objektes beschäftigt, -welche die in allem Wechsel gleichbleibenden Faktoren
nach ihrer Natur und Wirksamkeit untersucht."
^
S.
„Grenzen der naturwissenschaftlichen Begriffsbildung"
I
und II.
Gesetz.
von hier aus entwickelt
169
von
die Lehre
er
wicklungsstufen, die in bestimmter
sozialpsychischen
Eine nähere Erörterung an dieser Stelle
Hauptproblem wird
Hier
nur zweierlei bemerkt.
sei
Würdigung
der
ist
nicht wohl möglich;
der Artikel Geschichte
auf das
Tatsächlichkeit
Ent-
Ordnung verlaufen.^
Zunächst,
und
daß
zurückführen.
auch
Einmaligkeit der
eine
volle
Geschichte
nicht verhindert, gewisse Regelmäßigkeiten in ihr anzuerkennen.
Art
z. B.,
und wie sich
Kunst
u.
Die
wie sich der Ablauf der Entwicklung eines ganzen Volkes,
s.
die
Bewegung von
einzelnen Gebieten
in
verschiedenen
Kulturepochen
w,
wie Religion,
vollzieht,
kann
als
Erzeugnis der bleibenden Art des Menschen sehr wohl etwas verwandtes, ja gleichmäßiges haben.
Insofern könnten unbedenklich Ge-
setze der Geschichte anerkannt werden.
Aber sie würden dann nur
die Form des Geschehens betreffen, der Inhalt würde der Individualiät
der einzelnen
entziehen.
Epochen verbleiben und
— Wie
der "Geschichte
viel
sich
damit
aller
Ableitung
Selbständigkeit ferner die Einzelvorgänge in
haben,
Einzelvorgang zu fassen
und wieweit
ist,
ganze Geschichte
die
das wird sich
als
ein
verschieden gestalten je
nach der prinzipiellen Fassung des Geisteslebens und seines Verhältnisses
über,
ob
zur menschlichen Lage.
Persönlichkeiten
geschichtlichen
Bewegung
oder
bilden.
Diese Fassung entscheidet dar-
Massenwirkungen
Das
alles
den
Kern
der
aber weist über das
methodologische Problem hinaus und wird uns später zu beschäftigen
haben.
*
Über den Streit um Gesetze der Geschichte orientiert vortrefflich
Bemheim, Lehrbuch der historischen Methode und der Geschichtsphilosophie.
3. u. 4.
Aufl.
S.
91 ff.
Zum Weltproblem.
C.
Monismus und Dualismus.
1.
Auch bei der Wendung zu
•
den Weltproblemen
Problem des Lebensprozesses,
uns das
stets
Denn auch dort liegt die Entscheidung nicht
gegenwärtig bleiben.
abstrakt-begrifflichen
bei
wird
des Geisteslebens,
speziell
Erwägungen,
sondern
beim
Tatbestande
der Wirklichkeit; dafür aber ist nichts wichtiger als die Frage, welchen
Inhalt das Geistesleben zeigt
hier liegt das Zentrum,
bringen sind, und
wo
und welche Stellung es damit gewinnt;
wohin
seine
besonderen Untersuchungen zu
alle
letzte
Würdigung
fahrung an der Breite der Dinge ermittelt
Kämpfen
hat;
findet,
was die Er-
sucht doch in allen
um die Weite der Welt schließlich der Mensch sich selbst,
den Kern seines eignen Wesens. Auch die geschichtliche Betrachtung
bestätigt das
durch den Nachweis, daß es überall die eigentümliche
Gestaltung des Geisteslebens war, welche die Theorien erzeugte und
zur Macht erhob.
a)
Zur Geschichte und Kritik der Begriffe.
Die Ausdrücke Monismus und Dualismus entstammen den letzten
Jahrhunderten.
1700
(z.
B.
das
Dualismus verwandte zuerst Thomas Hyde
erschienenen
Kap. IX,
dem guten
S.
Schrift
164)
Historia
religionis
(s.
der
in
Persarum
zur Bezeichnung eines religiösen Systems,
Prinzip ein böses als gleichewig zur Seite
diesem Sinne ward das Wort durch Bayle
und Leibniz
veterum
(s.
stellt;
in
den Artikel Zoroastre)
Theodicee II, 144, 199) an weitere Kreise gebracht.
Als Gegenstück zu
„Monismus" gebrauchte
es
zuerst Wolff,
aber
zugleich übertrug er die Ausdrücke auf das Verhältnis von Körper
und
Geist:
Monisten
— das Wort
nunmehr, die nur eine Art des
ist
Seins,
von Wolff gebildet — heißen
sei
es
Körper
sei
es Seelen,
Monismus und Dualismus.
171
annehmen, also sowohl die Idealisten
als
die Materialisten, Dualisten
welche Körper und Seelen
als
voneinander unabhängige
dagegen,
Substanzen
Wolff
betrachten.^
selbst
Dualist
wollte
Beide
sein.
Ausdrücke blieben auf die Schule beschränkt, namentlich Monist
das 19. Jahrhundert hinein äußerst selten.
scheint bis in
er-
In weiteren
Umlauf brachten das Wort zuerst Hegelianer als Bezeichnung ihrer
eignen Denkweise, so erschien 1832 eine Schrift von Göschel »Der
Dann folgte wieder eine Ebbe, bis die
Monismus des Gedankens".
und
darwinistische Entwicklungslehre (Häckel
druck ergriff und ihn sich anpaßte.
System,
das
und
Körper
Weiter aber bezeichnet er jedes
und Geist
Natur
Seele,
den Aus-
Schleicher)
nicht
einander,
sondern beide einem überlegenen Dritten unterordnen und einfügen
diesem Sinne werden
In
will.
oft
Monismus und Spinozismus als
gleichbedeutend genommen.
So führen
uns
das
'
von
und
Natur
Ausdrücke
die
jetzt
Körper und Seele oder
—
Der Gegensatz,
Geist.
erhält eine besondere Schroffheit dadurch,
eignen Wesens
und daß
betrifft,
eröffnet sich
uns
in
er
wachsen
daß
dabei
entsteht,
den Kern unseres
er
scheint.
Die Welt, so scheint
von außen her durch
Empfindung, von innen her durch
ein Reich
von sinnlichen Eindrücken und
wird
der
von
— auf
im Fortgang der weltgeschicht-
sinnliche
Gedankengrößen;
Verhältnis
zwiefacher Weise:
lichen Arbeit unablässig zu
es,
das
auf
Ausdehnung über das Weltall
in
die
nehmen können, oder auch
eine
ein
als ein
als
Reich unsinnlicher
Reihe die andere
tieferes
Denken,
selbsttätiges
sich
in
auf-
Eindringen den Gegensatz
in einen
bloßen Schein verwandeln? Zum Gegensatz der Betrachtungs-
weisen
kommt
der wachsende Abstand
des
Inhalts
Aus der Natur hat die Wissenschaft zu Gunsten
beider Welten.
einer präzisen Be-
greifung und einer sicheren Beherrschung immer mehr alles seelische
Element vertrieben,
zugleich
aber hat
das Seelenleben
im
eignen
immer weiter über die bloße Natur hinausgehoben und
immer mehr zu einem selbständigen Reiche ausgebildet. So
Kreise sich
sich
*
Es ergab sich danach bei Wolff folgendes Scliema philosophischer
Parteien
Dogmatiker
Skeptiker
Monisten
Idealisten
Egoisten
Dualisten
Materialisten
Pluralisten
Zum Weltproblem.
172
der Verlauf der Geschichte das Körperliche immer seelenloser,
läßt
immer
das Seelische
Das müßte den Dua-
erscheinen.
spiritueller
zum Monismus hin, da nicht nur die exakte Forschung den Zusammenhang von
lismus empfehlen, aber zugleich wachsen die Antriebe
immer deutlicher aufweist und immer weiter
Seelenleben und Körper
sondern auch ein zunehmender Drang nach
ins Einzelne verfolgt,
dem Menschen
Einheit
So
verbietet.
Nebeneinander
ein
verschiedener
Welten
unseren Begriffen immer weiter auseinander, was
tritt
Erfahrung immer enger verschlungen zeigt; so treibt es uns
immer zwingender zur Umwandlung des widerspruchsvollen Bildes.
die
Die Hauptrichtungen solches Strebens
was sie aber zeigt,
lichem Bude;
erloschen,
und
sondern es bleibt
fordert
immer
ist
zeigt die Geschichte in deut-
nicht mit der besonderen Zeit
eine Möglichkeit stets gegenwärtig
als
neuem zur Entscheidung
von
auf;
bei
allen
Wandlungen der Begriffe behaupten sich charakteristische Denkund Lebenstypen durch den Lauf der Zeiten bis in die Gegenwart.
Es geht aber die lebendige Geschichte,
die
die eigne Arbeit hineinreicht,
in
Frühere
alles
ist
für
uns zur
bloßen
befaßt,
Jahrhunderts
17.
nicht
Auseinandersetzung
Seelischen enthielt
positive
aber
zu
der
alle
Arbeit
Geschichte,
einer
präzisen
geführt.
bis
sich
Denn
viel
mit
zu Beginn des
Fassung und deutlichen
Die
Vorstellung
des
mehr eine Verneinung des Körperlichen als eine
Behauptung; 1 so war es nicht zu vermeiden, daß immer
in
sie
bewegt von
belebt.2
Mittelalter
hatte
Begriffe
wieder das Bild einer nur feineren,
lichkeit
die
geworden.
Historie
wohl haben schon das Altertum und das
dem Probleme
d. h.
nicht hinter Descartes zurück;
subtileren,
luftartigen
Der Körper aber schien
eindrang.
seelenartigen
Kräften,
die
belebt,
Körpergebildet,
ganze Natur war innerlich
Bei solchem Stande der Begriffe bediente sich die Erklärung
der Natur fortwährend seelischer Größen und verschloß sich damit
eine
exakte
Begreifung
Psychologie unter den
es
erregte
*
der
Vorgänge.
Einfluß sinnlicher
Andererseits
geriet
die
und räumlicher Begriffe,
keinen Anstoß, Wirkungen von außen in die Seele ein-
So konnte Descartes mit gutem Grunde sich dessen rühmen, daß er
d. h. bewußte Tätigkeit, positiv be-
das Ganze der Seele als Denken,
zuerst
stimmt habe.
*
Bezeichnend dafür
ist
die
aristotelische
Definition
was das «Prinzip der Ruhe und Bewegung in sich
satze zur Kunst, die es außer sich hat.
dessen,
der Natur als
trägt",
im Gegen-
Monismus und Dualismus.
173
fließen
und Willensimpulse in räumliche Bewegungen übergehen zu
lassen.
Es war ein chaotischer Stand, der weder der Natur noch der
Seele ihr Recht gewährte.
überwunden, namentlich
Erst die Aufklärung hat diesen Stand
ist
der eine durchgreifende Scheidung und
Descartes,
es
Klärung
Nun erst wird jedem Gebiet eine volle Eigentümlichkeit
vollzieht
Das Seelenleben wird
zuerkannt.
reines
ein
als
Beisichselbstsein
verstanden, dessen Einheit des Wesens (unitas essentiae) sich scharf
von
Einheit
aller
geht
hier
bloßen
einer
Zusammensetzung
wie solche die Außenwelt
scheidet,
sitionis)
einen seelischen Charakter; die seelische
sich
zurück und
selbst
schendes
einfließen,
kettet
und verleiht ihr erst
Bewegung kehrt immer zu
Mannigfaltigkeit an
alle
sondern
eignen Grunde
Anregung kann
alle
auch
in scheinbarer
sich
selbst.
nur
es
aus seinem
reizen,
hervorzubringen;
Leistungen
gewisse
beherr-
ein
kann nichts von draußen
In ein solches Seelenleben
Ich.
compo-
(unitas
das Bewußtsein
voran
besonderen Tätigkeit
aller
bietet;
so
Abhängigkeit von draußen im Grunde
bleibt
es
stets
bei
Solcher Selbständigkeit der Seele entspricht ein Selb-
ständigwerden der Natur.
nach Austreibung
Die Massen und Bewegungen,
alles seelischen
Elementes verbleiben,
ihr
die
bilden eine
eigne Welt; die Bewegung, für die bis dahin eine Seele unentbehrdünkte,
lich
gelegt;
wird nun den kleinsten Teilchen von Haus aus bei-
das macht es möglich, aus der Zusammensetzung kleinster
— bewegter, aber seelenloser — Teilchen
faltigkeit
alle
der Natur hervorgehen zu lassen.
Strebungen verschwinden damit aus ihr.
sinnlichen
Eigenschaften,
mit
der
in
den Dingen
geborgt, als
So
ein
selbst angehörig,
Tönen
u. s. w.
die
sie
erscheint
nun
nicht
als
ihnen von der Seele
Farben
sondern
sich
scharf
beide
Abschluß dabei verbot
Seiten,
Aber so
Verwicklungen diese Spaltung hervorrief,
schritt,
sie
der fruchtbarste Anregungen brachte.
beiden Gebiete ihre eigentümlichen
voller Klarheit
lisch,
,
vom Menschen in sie hineingelegt
scheiden
letzter
und
Ja, die ganze bunte Fülle der
menschliche Fassung die Natur umkleidet,
als
unermeßliche Mannig-
Alle inneren Kräfte
herausarbeiten,
nun
so
scharf,
viele
ist
daß
sich
neue Fragen und
ein gewaltiger Fort-
Nun erst konnten die
Prinzipien
und Methoden mit
erst ließ sich
das Seelische see-
das Körperliche körperlich verstehen, eröffnete sich eine exakte
Physik und eine erklärende Psychologie.
jetzt von
den Dingen, nun
Wie
ein Schleier
erst schien die Wirklichkeit sich
fiel
es
uns völlig
Zum Weltproblem.
174
aufzuhellen.
Dazu brachte jene Scheidung nicht bloß
der Begriffe,
sie
der Kultur,
die
seits
Gedankengrößen,
in
lichkeit
von jener Zeit an durch die Neuzeit geht.
erhöhte Tätigkeit des Denkens,
eine
der Vernunft, ein
kühn
eine
Klärung
über
ein
Einer-
Umsetzen der Wirk-
Rationalisierung aller Verhältnisse,
Bindung vordringende
bisherige
alle
ein
Messen des Daseins an Forderungen
ein
nach
Streben
andererseits
kultur;
eine
entsprach der zwiefachen Richtung des Lebens und
Selbständigwerden
volles
Intellektual-
Außenwelt
der
gegenüber dem Menschen, eine engere Verflechtung seiner mit der
Umgebung,
unermeßliches Wachstum der Erfahrung, eine
ein
Bedeutung
höhte
der
Faktoren,
materiellen
ein
immer
er-
stärkeres
Anschwellen einer Realkultur; wer könnte leugnen, daß diese zwei"
Ströme durch das Leben der Neuzeit gehen,
liche
Spannung
versetzen
und durch
in
Gegensätze auseinandertreiben?
ist
die
tiiefste
die
es in eine unaufhör-
ganze Breite der Dinge
Dieser Gegensatz
des
Lebens
Wurzel des Dualismus der Begriffe und Lehren, aus
ihm schöpft dieser immer neue
Kraft,
so sehr das Verlangen nach
Einheit die Geister über ihn hinaustreiben mag.
solches Verlangen
Ein
konnte
schließen
die
war allerdings unabweisbar, denn ab-
Denkarbeit
bei
hatte eine kräftige Analyse vollzogen
jenem Dualismus
und
schärft,
aber von der Analyse drängte es
welcher
Synthese,
Einheit.
Auch
vom Gegensatz
fehlte es nicht
Zerspaltung der Wirklichkeit,
zu
die Begriffe
nicht.
immer wieder zu
irgendwelcher
Er
dauernd verirgend-
umfassenden
an gewichtigen Gründen gegen jene
Ihr
widerspricht nicht nur der un-
mittelbare Eindruck einer engen Zusammengehörigkeit
von Seele und
Körper, sowie die wachsende Erkenntnis der Abhängigkeit des Seelenlebens von körperlichen Bedingungen, ihr widerspricht auch die philo-
sophische Forderung einer Einheit der Wirklichkeit, ihr widerspricht
endlich
die
Tatsache der Kunst mit ihrer engen Verflechtung und
fruchtbaren Wechselwirkung von Sichtbarem und Unsichtbarem, von
Äußerem und Innerem. Alles zusammen ließ den Dualismus als
einen bloßen Durchgang zur Einheit erscheinen; diese Einheit fand
sich allerdings nicht fertig vor, sie
gewinnen,
und
diese
durfte
Eindruck nicht scheuen.
kühner geworden
Es sind
als je
aber
auseinanderging,
drei
So
war durch geistige Arbeit erst zu
dabei einen
ist
Gegensatz
zum
nächsten
das Einheitsstreben in der Neuzeit
zuvor.
Hauptrichtungen,
die Richtungen
in
welche
des Materialismus,
dies
Streben
Spiritualismus,
Monismus und Dualismus.
175
Monismus: entweder wird das Körperliche, oder es wird das Seelische
zum allumfassenden Sein, oder es wird beides zu Seiten, Erscheinungen,
Ausdrucksweisen einer
Wie
es
tiefer
gegründeten Wirklichkeit.
strengem Sinne
Materialismus in
einen
durch Descartes erfolgten Klärung der Begriffe
damals
eine
erst
durchlief
feste
Bezeichnung
tüchtigste,
bei
der
Der Materialismus
den Franzosen die
den Deutschen die derbste Gestalt gefunden;
seit
so hat er auch
gibt,
erlangt. ^
Kulturvölker und
nacheinander die großen
Engländern die
erst
bei
den
geistreichste,
bei
hat
und zu
widerlegt
oft
Boden geworfen, ist er immer von neuem erstanden, und hat er
immer von neuem weite Kreise gewonnen. Das bekundet doch
mehr in ihm steckt als naive Gemüter wähnen,
wohl, daß
die ihn
durch scharfsinnige Widerlegungen endgültig abgetan glauben und
daß immer wieder Menschen dem längst durchschauten
sich wundern,
Irrtum verfallen.
In
zwingen, wenn
bei
Denn
ständen.
hängigkeit des
Vorzug
den
Wahrheit wäre der Materialismus
Seelenlebens von
großen
einer
Bedingungen,
körperlichen
Einfachheit
und
und problematischeren
Gemeinverständlich-
zu
Begriff als
steht
und
es,
Gerade die
lassen.
und Seelischen,
Körperlichen
schaft
lange, als eine
den der Materie, er
ent-
wo wir ihn zu fassen glauben; je präziser wir ihn aber
nehmen, desto unmöglicher wird
vom
Ab-
sowie
Denn kaum gibt es einen schwie-
Analyse der Begriffe unterbleibt.
vorgehen
Frage
die Abhängigkeit läßt sich auch in anderer
Weise verstehen, und die Einfachheit verbleibt nur so
weicht uns,
in
so gewiß der Materialismus die unbestreitbare
keit für sich anrufen kann,
rigeren
zu be-
leicht
ihm bloß theoretische Erwägungen
fällt,
unmöglich gemacht;
hat
mit
aus ihm seelisches Leben her-
moderne Klärung der
Begriffe
mit der die exakte Naturwissen-
den Materialismus
Recht sagt daher
F.
als
Weltanschauung
daß den
A. Lange,
Materialismus scharf denken ihn widerlegen, heiße.
Aber
es
sind
nicht
wissenschaftliche
Erwägimgen,
es
sind
Kultur- und Lebenslagen, welche dem Materialismus eine Anziehungs-
und Überzeugungskraft geben.
reich
in
^
solchen
Zeiten,
Der Ausdruck
wo
Materialist
Wir
erscheint
Naturphilosophen Robert Boyle (so
in
stark
und
sieg-
Kulturformen
ihre
volle
finden
überlieferte
zuerst
ihn
bei
dem Chemiker und
der 1674 erschienenen Schrift:
The
and grounds of the mechanical philosophy), der eine Vorliebe für
feste Termini hatte.
Noch Giordano Bruno verwandte den älteren Ausdruck
excellence
»Epikureer".
Zum Weltproblem.
176
Wahrheit eingebüßt haben und von vielen als ein tyrannischer Druck
empfunden werden; der Materialismus
dann
erscheint
das beste Mittel zur Befreiung von drückenden
sowohl
als
Fesseln wie als ein,
Rückgang auf einfache Grundlagen des Lebens,
er
scheint
eine
natürlichere und wahrere Gestaltung aller Verhältnisse zu versprechen.
Auch
bringt er
die
Bedeutung der materiellen Lebensbedingungen
für das
Ganze der Kultur mit besonderem Nachdruck zur Geltung.
So
er
riß
Geister fort
die
in
den Bewegungen
vor
und
in
der
französischen Revolution, so auch beim Sozialismus der Gegenwart.
Was
das Leben hervorgebracht
so
Leben widerlegen,
es
nur das
auch
negativ durch den inneren Widerspruch,
positiver Weise,
dem eine
durch ihre eigne Entwicklung
materialistische Gestaltung der Kultur
durch die Gegenwirkung einer andersartigen Kultur.
positiv
verfällt,
kann
hat,
widerlegt es aber sowohl in negativer als in
Jener Widerspruch wurzelt darin, daß dort den materiellen
Größen
als eigne Leistung beigelegt wird, was ein überlegenes Geistesleben
ihnen macht;
aus
wie dieses
der
in
Welt uns
sichtbaren
unver-
gleichlich mehr wahrnehmen läßt als die Sinne direkt zeigen, so macht
es die materiellen Güter wertvoll als ein
Werkzeug für die Betätigung
und Entwicklung vernünftiger Lebewesen; wie vom Materialismus
der
dort
Zuschauer,
eignis
behandelt.
das Streben
so
wird
unvermerkt
hier
eine
zwecktätige
hinzugedacht und ihr Erlebnis wie ein äußeres Er-
Persönlichkeit
und
Indem aber
die Arbeit
die
materialistische Lebensgestaltung
von dem Träger des Lebens
ablenkt,
überliefert sie ihn einer wachsenden Verkümmerung und Leere; hat
nun zugleich jenes Wachstum der Beziehungen nach außen einen
gewaltigen Lebensdurst erzeugt, so muß ein schreiendes Mißverhältnis
zwischen unserem Verlangen und unserem Besitz entstehen; das Mißbehagen, das daraus hervorgeht, wird schließlich mit Sicherheit über
eine materialistische Lebensführung hinaustreiben.
Die weltgeschichtliche Arbeit zeigt das
vollzieht
durch
Materialismus.
das Ganze
ihres
Verlaufs
in
großen Zügen und
eine
Überwindung des
Der durch jahrtausendlange Arbeit, durch fruchtbare
Erfahrungen und schmerzliche Enttäuschungen zu einem Innenleben
geweckte Mensch kann unmöglich mit
in
dem Kinde und dem Wilden
der materiellen Welt seine ganze Wirklichkeit sehen und in ihren
Gütern seine Befriedigung finden.
für ihn durch
wonnen.
jene
Bewegung
ein
Die materielle Welt
wesentlich
selbst
hat
anderes Ansehen ge-
Aus dem bunten Reich der sinnlichen Eindrücke ist jetzt
Monismus und Dualismus.
177
Gewebe von Kräften, Gesetzen, Beziehungen geworden; nicht
mehr die Handfestigkeit der Sinnesempfindung verbürgt uns die
Wirkhchkeit des Ganzen, sondern die kausale Ordnung mit ihrer
ein
Verkettung
schehens
und
Mannigfaltigkeit
aller
unter
ihrer
Auch
Gesetze.
einfache
Einfügung
die
Ge-
alles
Außenwelt
ist
ins
dem Denken entsprungene Größen, ideelle
Größen, bilden ihren Kern. Wohl bleibt hier die geistige Tätigkeit
Unsinnliche verwandelt,
an einen ungeistigen Vorwurf gebunden, aber auch so
verschieden von
noch so fortgebildeten
aller,
sie weit
ist
Welcher
Sinnlichkeit.
Abstand ist zwischen der Welt des Naturforschers und der des Natur-
menschen mit noch so geübten Sinnesorganen!
Nicht minder verwandeln sich dem Kulturmenschen die äußeren
Was sie heute ihm wertvoll macht, ist weniger der sinnliche
Güter.
Reiz
und Genuß, als die Herrschaft über die Dinge, das Vermögen,
diese nach
eignem Wollen zu bewegen und damit das eigne Leben
Unbegrenzte zu
ins
sowohl die Dinge
So
steigern.
als sich
genießt der
selbst in
Kulturmensch nicht
den Dingen, sein Denken gibt
dem Sinnlichen Wert und gestaltet es zu Gedankengrößen. Welcher
Abstand liegt hier zwischen der Lust des Wilden am Glanz des
Goldes und dem Selbstgefühl des großen Geschäftsmannes, dessen
wirtschaftliche
Macht den Erdball umspannt und
von den
sich dabei
sinnlichen Wertzeichen ganz emanzipiert hat!
So
wirken
in
geistige
Kräfte,
die
der
Gestaltung
eignen
der
materiellen
Welt
kann.
Aber
der Materialismus nicht verstehen
zugleich leuchtet ein, daß, was dabei an Lebensentfaltung aufgeboten
wird, als Abschluß unmöglich ist;
leistet,
was so viel an einem fremden Stoffe
muß notwendig auch bei sich selbst etwas sein;
werfung des Äußeren und
alle
vor peinlicher Leere, wenn
Unter-
alle
Ausdehnung der Macht schützt nicht
dem
Geistesleben
nicht irgendwelcher
Diesen aber kann ihm alle Steigerung materieller
Inhalt gegeben wird.
und wirtschaftlicher Macht unmöglich gewähren. So muß der Versuch,
das Glück von
außen
her
zu
begründen,
schließlich
große
eine
Enttäuschung und Erschütterung ergeben. Die materialistische Lebensgestaltung wird
hat,
aufs
So muß
härteste
sich
dem Glückverlangen,
mit
zusammenstoßen
möglich
ist
sie
selber
angefacht
und dabei zusammenbrechen.
auch praktisch der Materialismus durch seine
Entwicklung widerlegen.
lösung
das
Aber alle solche Kritik und
noch keine positive Überwindung.
durch
eine
Eucken, Grundbegriffe.
kräftige Entfaltung
4. Aufl.
Eine solche
selbsttätigen
eigne
kritische Aufist
nur
Geisteslebens;
12
Zum Weltproblem.
178
wo
mit
dies
Aufgaben
seinen
die
Gemüter
da wird
erfüllt,
es
kaum begreiflich dünken, wie der Mensch das ihm innerlich Nächste
und die Quelle seiner eigentümlichen Größe mit dem Materialismus
als
etwas Abgeleitetes behandeln, seine eigne Existenz auf den Kopf
sein
stellen,
Glück von außen her suchen kann.
Vermag der Materialismus mit seiner Sinnfälligkeit und scheinbesonders
baren Selbstverständlichkeit
auf
die
breiten
Massen zu
wirken, so ist der Spiritualismus eher eine Sache einzelner vornehmer
und auserlesener
Geister
Eindruck gegen
sich,
Denn
er
hat
den unmittelbaren
geistige Energie
ist
der von ihm ver-
Kreise.
ohne
Weg nicht zu Ende zu gehen.
suchte
den Spiritualismus
entweder
ein
als
in
Es zeigt aber
Reich von
lauter
einzelnen Seelen,
Leben und Sein eines Gesamtgeistes, jenes
neueren deutschen Spekulation,
wie da
soll
die
am
großartigsten
Außenwelt sich gänzlich
als
eine Stufenfolge innerhalb
Sinnliche wird hier
niedere,
statt
oder
als
das
bei Leibniz, dieses in der
in
bei
Hier
Hegel.
Innenleben verwandeln,
das Verhältnis von Geist und Natur wird nicht
sondern
die Neuzeit
zwiefacher Gestalt: die Wirklichkeit erscheint
als
ein
Gegensatz,
des Geistes verstanden;
das
einer in sich selbst gegründeten Welt eine
noch nicht zur vollen Bewußtheit gelangte Art des seelischen
oder geistigen Lebens.
Diese Denkweise braucht nur ausgedacht zu werden,
minder wunderlich darzustellen,
mag;
um sich
beim ersten Anblick scheinen
uns doch das Innenleben die nächste und gewisseste Wirk-
ist
lichkeit,
als sie
und zeigt die einfachste Besinnung, daß wir diesen Kreis nie
und uns in ein anderes Sein versetzen können,
daß auch, was Außenwelt heißt, nur eine besondere, eine eigentümlich
gänzlich verlassen
gebundene Art des Innenlebens bedeutet.
Aber so berechtigt und überzeugend der allgemeine Gedanke
sein mag,
leicht
Die
beim Versuch einer strikten Ausführung überspannt
sich
das menschliche Vermögen und wird unser Besitz überschätzt.
Spiritualisten
können nicht unternehmen, die Natur ganz und
gar in Geist umzusetzen, ohne unser Geistesleben als Geistesleben
schlechthin, als absolutes Geistesleben zu behandeln.
Zu einer Stufe
menschlichen Geisteslebens aber läßt die Natur sich nun und nimmer
herabsetzen; jenem gegenüber erweist sie eine viel zu selbständige
Art und verfolgt
sie
einen viel
zu
sie
viel
zu sehr ihren eigenen Weg, ihm
hartnäckigen Widerstand.
leistet
Solcher Selbständigkeit
der Natur und solcher Härte ihres Widerstandes hat sich der Spiri-
Monismus und Dualismus.
lyg
tualismus nur gewachsen fühlen können, indem er das Geistesleben
in
bloßes Denken
und Erkennen verwandelte und das Ungeistige
als
etwas verstand,
das noch nicht voll aufgeklärt
der Stufe der Unbewußtheit verharre.
der
Intellektualismus,
dem
aus
Weltleben
das noch auf
ein
kecker
bloße Weltansicht
eine
machte, der damit die Wirklichkeit verflüchtigte
Inhalts beraubte.
sei,
Aber das war
und alles lebendigen
Begreiflich ist solche intellektualistische Überschätzung
des menschlichen Vermögens nur aus der Eigentümlichkeit beson-
wo das Bewußtsein geistiger Kraft und das Voll-
derer Kulturlagen,
gefühl
geistigen
den Menschen
Schaffens
sich
den Mittelpunkt
als
der Wirklichkeit fühlen ließ und ihn über alle Schwere der Dinge in
kühnem Fluge hinaushob.
Aber jene Schwere wird bald bemerklich
der Kultur
werden,
jene Art
aber
aller Spiritualismus,
fällt
sich
als
zu
erweisen.
flach
der sich als ein
fertiges
Damit
System
gibt.
Das Mißlingen der Versuche, eine der beiden Lebensformen
ausschließlich durchzusetzen, mußte dem Monismus zu Gute kommen.
Aucfi er will eine Einheit, aber er will sie nicht durch Aufopferung
der einen Seite an die andere, sondern durch Einfügung beider in
ein umfassendes Drittes;
lichkeit voll
entfalten
hier scheint jedes Gebiet seine Eigentüm-
zu können, ohne aus der Gemeinschaft heraus-
zutreten, hier entfällt die Schwierigkeit einer Wechselwirkung zwischen
Körper und
Seele,
da dem Vorgehen auf der einen Seite unmittel-
auf der anderen
Zu Gunsten
bar
eins
als
Lebensgestaltung aber wirkt namentlich
vermeintlich
entspricht.
des
Monismus
das hier erstrebte und
gewonnene Gleichgewicht zwischen Natur und
Geist,
zwischen Äußerem und Innerem, zwischen Sinnlichkeit und Denken,
zwischen
realistischer
und
idealistischer
Kultur;
solches
Gleich-
gewicht scheint besonders geeignet, das Leben ins Weite und Große
zu heben, den Menschen der Enge eines Sonderkreises zu entwinden
und ihm an der ganzen
Fülle
der Wirklichkeit Anteil
So hat der Monismus, namentlich
seit
ihm Spinoza
zu geben.
eine klassische
Verkörperung gab oder doch zu geben schien, eine gewaltige Anziehungskraft geübt auf Dichter
und Denker, auf Naturforscher und
religiöse Naturen, er schien die Zauberformel, die überallhin Frieden
bringe.
Aber eine solche Zauberformel
Seine zu denken
in
gestattet,
ist
er
nur,
weil er
weil jeder den allgemeinen
jedem das
Gedanken sich
eigentümlicher Weise zurechtlegt; so gewiß in jenem Gedanken
eine unangreifbare Wahrheit liegt, in der Ausführung stellt sich alsbald
12*
Zum Weltproblem.
180
der Gegensatz wieder ein, den es zu überwinden galt; es zeigt sich,
daß auch
Entweder
diesem Problem
bei
— Oder
menschliche Streben
das
vor
ein
gestellt ist, nicht friedlich die Gegensätze zusammen-
zuschließen vermag.
zeigt,
Nach der Absicht des Monismus, wie ihn der Spinozismus
müßten beide Gebiete in vollem Gleichgewicht stehen. So
will
es
der neuerdings
auch der «psychophysische Parallelismus",
In Wahrheit ist eine
jene Absicht zu genauerer Durchbildung bringt.
nähere Ausführung des Grundgedankens gar nicht möglich ohne der
einen Seite ein Übergewicht über die andere zu geb^.
ist,
Spinoza selbst
genauer angesehen, an keiner Stelle Monist, sondern bald Materialist,
bald Spiritualist, jenes in der Grundlegung, dieses im Abschluß seiner
Denn
Lehren; so zeigt es namentlich seine Ethik.
zu Beginn die Natur
Wirklichkeit,
erscheint
dort
das Hauptgeschehen und als das Maß
als
während das Seelenleben
zu
einer
Beim Ab-
erscheinung, einem Reflex des Naturprozesses herabsinkt. ^
dem Materialismus ein Spirituawenn ein göttliches
schluß des Systems aber wird aus
Oder
lismus.
ist
es
kein
aller
bloßen Begleit-
Spiritualismus,
Leben die ganze WirkHchkeit durchdringt und zusammenhält, die
der Mensch
Natur zur Entfaltung dieses Lebens wird,
Gott an
intellektuelle
Liebe
zu
teilgewinnen
soll?
Und
der
hinaus in den Kern des
Unendlichkeit und
der
Zwiespalt
über
die
Begriffe
es
nicht
ein
einziges,
ist
sondern ein zwiefaches Leben, das aus Spinoza wirkt:
dann
werden mag,
die erstrebte Einheit
Ewigkeit
reicht
Lebens hinein,
Naturalismus,
durch die
einmal ein
Wie immer Spinoza beurteilt
ein Mystizismus.
hat
er
nicht
erreicht.
Ebenso-
wenig ist es späteren Versuchen gelungen, Natur und Geist
Gleichgewicht zu bringen.
hat es nicht erreicht;
er
Auch der psychophysische
in
ein
Parallelismus
macht entweder das Seelenleben zu einem
bloßen Reflex der Naturvorgänge, oder diese zu einer Erscheinung
des Geisteslebens; er verläßt damit die Neutralität und nähert sich
entweder
^
dem Materialismus oder dem Spiritualismus.
Mit vollem Recht bemerkt dagegen Herbart
Metaphysik (Wke.
aus Bestimmungen
III,
198):
„Da überdies
alles
in
seiner Allgemeinen
Psychologische bei Spinoza
man wenig
vom Ausgedehnten bestehen
des Körperlichen gefolgert wird: so merkt
davon, daß nach ihm das Denken unabhängig
und wie könnte es anders sein in irgend einer Lehre, die ursprüngGedanken als Bilder des Ausgedehnten betrachtet? Eine solche
unterwirft immer notgedrungen den Geist der Masse, vermöge des Verhältnisses der Abbildungen zu ihrem Vorbilde."
sollte;
lich
die
Monismus und Dualismus.
Noch weniger
ergibt
jg!
von jenem vermeintlichen Gleich-
sich
Denn
gewicht aus eine charakteristische Gestaltung der Kulturarbeit.
die
Ausgleichung von Natur und Geist, welche namentlich künst-
lerische Seelen anzog, erfolgte nicht zwischen
als gleichberechtigten
Außen- und Innenwelt
Größen, sondern sie erfolgte gänzlich auf dem
Boden des Innenlebens; wenn z. B. im Schaffen
um
zur Darstellung drängte,
Innere
eines
damit sich
so erhielt zugleich das Äußere ein inneres Leben;
wurde hier durch
gestaltet,
ein
befreit,
menschliche Sein zwischen Geist und Natur
deutlicher
verläßt
die
nicht aber
Denn
Monismus
der
unterscheidet
vom
der
ihn
Materialismus nur dies, daß er das Seelenleben nicht
däres,
sondern
als
von -Haus aus
primäres
ein
beilegt,
Punkten entstehen
läßt.
feineren Materialisten;
es
wurde das
zerteilt.
Neutralität
Entwickluilgslehre.
und
zum Weltall
innigeres Verhältnis
von der Enge kleinmenschlicher Art
Noch
zu finden,
das Geistesleben
ein kräftigeres Erfassen der Natur bereichert
im besondern durch
darwinistischen
Goethe alles
selbst
als'
ein sekun-
Phänomen erachtet, es aller Materie
nicht
erst
nachträglich
an
besonderen
Aber kaum anders dachten von jeher die
wie ihnen,
so wird auch
Wahrheit die sinnliche Natur zum
All,
den Monisten
bemächtigen sich
in
Natur-
begriffe der gesamten Wirklichkeit und wird alles selbständige Geistes-
leben
geleugnet.
So muß,
bei
konsequentem Verfahren, auch die
Lebens- und Kulturgestaltung ganz in die Bahnen des Materialismus
geraten.
Anders würde die
Sache auslaufen,
wenn mit der Be-
seelung aller Elemente der Wirklichkeit voller Ernst gemacht würde;
denn das müßte
ein
dem leibnizischen ähnliches Weltbild ergeben.
Aber so weit pflegt der materialistische Monismus nicht vorzudringen,
er glaubt den Elementen die Seele wie eine Eigenschaft, neben anderen
beifügen zu können, ohne daß sie dadurch etwas wesentlich anderes
werden.
In
Wahrheit läßt sich Seele nicht haben, sondern nur sein.
Wird demnach der materialistische Monismus von allen Bedenken
getroffen, die
dem ausgesprochenen Materialismus entgegenwirken, so
sind einem spiritualistischen Monismus bessere Aussichten zuzuerkennen.
Ein solcher Monismus wird die Tatsache zum Ausgangspunkt nehmen,
daß Innenleben nicht bloß an einzelnen Punkten, zerstreut und
splittert,
erscheint,
zer-
sondern daß es sich zu einem umfassenden Zu-
sammenhange verbindet, daß sich auf der menschlichen Daseinsstufe
ein den Individuen überlegenes Geistesleben und mit ihm eine Innenwelt voll eigentümlicher Größen und Aufgaben erschließt. Der Wende-
Zum Weltproblem.
182
punkt der Wirklichkeit wird hier nicht zwischen Natur und Seele, sondern
zwischen Ungeistigem und Geistigem gesucht; das Seelenleben hat an
beiden Stufen teil, sofern es zunächst ein Stück der Natur bildet, dann
Die Frage, wie sich Körper-
aber ein Gefäß des Geisteslebens wird.
liches
und Seelisches zu einander verhalten,
tritt
hier zurück vor der,
wie Eine Welt Geistiges und Ungeistiges miteinander umfassen kann.
Diese Frage findet aber in diesen Zusammenhängen die Antwort, daß
das Ungeistige nur etwas Untergeistiges bedeutet, daß dasselbe Sein,
das die Natur und
das natürliche Seelenleben im stände der Ver-
einzelung und als ein Gewebe bloßer Beziehungen zeigt, im Geistesleben sich zu einem Ganzen zusammenzufassen
entwickeln
beginnt;
erst
und einen Inhalt zu
damit scheint die Wirklichkeit ein Inneres
Solche Erhebung von
und zugleich ihre eigne Tiefe zu gewinnen.
Untergeistigem zu Geistigem ist nicht eine bloße Forderung der Spekulation,
sondern eine Aufgabe, die das ganze Menschenleben durch-
dringt,
denn
ethische
Bewegung,
alle
eigentümlich menschliche Leistung, vor allem die
ein Aufstieg
ist
von der Natur zum
Erhebung des Seins von der natürlichen zur
wird hier das Problem
vom
bloßen
Intellekt
geistigen
Geist, eine
So
Stufe.
das Zentrum des
in
Lebens versetzt.
Wenn aber dabei das Geistesleben zugleich als der Grundbestand
und
als
das Ziel der Wirklichkeit erscheint, so besagt das keineswegs,
daß die Gestalt, worin
es dem Menschen vorliegt,
im stände sei,
sich
der ganzen Welt zu bemächtigen und in der Natur sich einfach wiederzufinden,
Denn so gewiß
wie das der reine Spiritualismus wollte.
das Geistesleben auch
dem Menschen
als
etwas Übermenschliches
und Allgemeingültiges irgend gegenwärtig sein muß, in seine nähere
Gestaltung dringt unablässig Bloßmenschliches ein; wir haben nicht
das Geistesleben an sich, sondern ein menschliches Geistesleben, d. h.
ein Geistesleben, dessen übermenschlicher Kern uns immer nur durch
eine menschliche Hülle zugänglich ist. Streben wir daher vom menschlichen Geistesleben her die ganze Wirklichkeit zurechtzulegen, so geraten
wir unvermeidlich in eine zu enge,
in eine
Fassung; eine unentbehrliche Hülfe dagegen
Unendlichkeit und ihrer Überlegenheit gegen
Zwecke;
sie
bewahrt
eine Sonderart,
sie
ist
anthropomorphe
die Natur mit ihrer
alle
kleinmenschlichen
den Menschen vor einem Sicheinspinnen
treibt
in
immer von neuem dazu, den Allgemein-
gedanken des Geisteslebens von der bloßmenschlichen Daseinsform
abzuheben.
Aber
alle
diese
Wirkung
liegt
innerhalb des Geistes-
Monismus und Dualismus.
und nur darin
lebens,
besteht
die
183
Abweichung vom dogmatischen
nun innerhalb des umfassenden Ganzen zwei
verschiedene Ausgangs- und Angriffspunkte anerkannt werden. Eben
daß
Spiritualismus,
dies aber ergibt einen anderen Typus des Kulturlebens, als ihn jener
Das Geistesleben erscheint nun nicht bloß als die begründende
vertrat.
Tatsache, sondern auch als eine sich immer erneuernde Aufgabe; das
menschliche Leben wird weit mehr zwischen Gegensätze
erscheint
weit
mehr
weit
unfertiger,
gestellt,
es
im Aufstreben begriffen;
erst
das ruft den Menschen
mehr zu eigner Tat und Entscheidung auf
und
zur
von ihm,
fordert
so
wird
Fortbewegung des Weltalls
nicht nur es sich in seinen
mitzuwirken,
nicht
das
Intellektuelle,
ein Ethisches weitester Fassung,
immer
die
keinen
Zweifel
Kämpfen
liegt,
dieses,
das
Ethische,
wo
sich
das näher gestalten
freilich
— Wie
mögen,
der Hauptstreitpunkt bei diesen
und an welcher Stelle vornehmlich die Geister sich
Das nämlich
scheiden'.
sondern
zum Kern seines Strebens.
Menschen und Zeiten
leidet
selbsttätig
Gedanken zurechtzulegen;
ist
die Frage,
ob
selbständiges Geistes-
ein
leben und mit ihm eine neue Stufe der Wirklichkeit anerkannt wird
oder
nicht.
Alle Verneinung oder auch nur Zurückhaltung gibt einem
gröberen oder feineren Materialismus die Oberhand, mit dem Ja wird
dagegen
eine
Überwindung und
sichere
Bahnen gewonnen.
Wohin
und
nicht bloß an Scharfsinn
ein
Einlenken
aber die Entscheidung
Intelligenz,
fällt,
in
neue
das hängt
sondern vornehmlich an
der Kraft und der Klarheit, mit der das Geistesleben Manschen und
Zeiten gegenwärtig
ist,
das greift damit in
das persönliche Leben
und Sein zurück.
b)
der
Der Monismus der Gegenwart.
Wer die geistigen Strömungen der Gegenwart kritisch erörtert,
muß sich notwendig auch mit dem Monismus der Gegenwart
befassen;
nisses
ist
dieser
doch über das besondere Problem des Verhält-
von Natur und Geist hinaus eine mächtige, ja stürmische Be-
wegung geworden, auf deren Beleuchtung und Würdigung sich nicht
wohl verzichten
läßt;
je
heftiger
aber der
Kampf an
dieser
Stelle
entbrennt, desto mehr hat die philosophische Betrachtung nach ruhiger
Abwägung zu streben.
Der Monismus der Gegenwart ist nicht verständlich ohne eine
Beachtung der breiteren Grundlage, von der aus er sich
gestaltet hat.
Diese
die
ist
das
Vordringen
des
Bildes
der
Natur
in
Begriffe
Zum Weltproblem.
184
vom All und in die Gestaltung des Lebens.
Es vollzog sich damit
ein notwendiger Rückschlag gegen die ältere, einseitig religiöse
welche die Natur wie etwas untergeordnetes
transzendente Denkart,
und
wenn
nebensächliches,
und
nicht
zu
gar bedenkliches
behandeln
Das energische Vordringen der Naturerkenntnis und auch
pflegte.
Umwandlung des Lebens gaben diesem RückWucht und siegreiche Kraft. Innerlich fiel dabei
die dadurch bewirkte
schlag gewaltige
besonders ins Gewicht die Präzision der Naturbegriffe und ihr Zu-
sammenstreben zum Ganzen einer Gedankenwelt; den Einflüssen, die
davon ausgingen, konnten sich selbst die nicht entziehen, deren Streben
der
Das
Hauptrichtung
ist
nach
eine
entgegengesetzte
z. B. augenscheinlich bei Leibniz.
Richtung verfolgte.
Sein unablässiger
gegen den „Naturalismus" hat nicht verhindert,
an Hauptstellen in seine Gedankenwelt eindrangen und
brachten.
Oder
zum
ist
kein
es
wenn
solcher Einfluß,
Kampf
daß Naturbegriffe
sie unter sich
er
die Lebens-
wenn er alle
Gegensätze in Unterschiede des Grades verwandelt, wenn ihm der
Begriff der logischen Möglichkeit mit dem der gehemmten Kraft zusammenfließt?
Im 19. Jahrhundert ging diese Bewegung weiter,
mit lautem und stillem Wirken unterwirft die naturwissenschaftliche
Denkweise sich mehr und mehr die Begriffe wie die Überzeugungen;
steigerung
allbeherrschenden Wertbegriff macht,
wir empfinden gar nicht, wie eigentümliche Bejahungen
neinungen das mit sich
bringt.
So
idee die Gestalt eines Naturprozesses
Verkettung, die daraus erwächst,
weise
alle
Gegenwart,
erhält
und
z.
B.
und Ver-
die Entwicklungs-
zerstört mit der strengen
alle Selbsttätigkeit, ja
ohne daß uns das irgend
konsequenter-
aufregt.
In
der
Natur hat sich das Beharrungsgesetz, das sog. Gesetz der Trägheit,
immer weiter bewährt; unbedenklich wird es auf
und geschichtliche Gebiet übertragen, während hier
der Forschung
das
geistige
der Lebensstand
gehen muß,
immer neu aus ursprünglichem Schaffen hervor-
um nicht sofort zu sinken.
Glück die sinnliche Lust, ohne weiteres
Glück als eine, wenn auch
Die Naturstufe zeigt
gilt
als
vielen auch das geistige
feinere Art der Lust.
Wenn so unauf-
haltsam von außen und innen die Natur auf uns eindringt, so kann
wundernehmen, wenn mehr und mehr die Natur als Welt
und Wirklichkeit schlechthin behandelt wird, und wenn eine „naturwissenschaftliche Weltanschauung" sich unbedenklich und siegesgewiß
es nicht
nicht als einen besonderen Ausschnitt, sondern als das erschöpfende
Bild des
Ganzen der Wirklichkeit gibt.
Monismus und Dualismus.
185
Alles Vordringen ließ indes diese Bewegung keinen vollen Sieg
erringen, so lange noch der Mensch eine privilegierte, ja einzigartige
Diese aber hat nun die schwerste Erschütterung
Stellung einnahm.
durch
Entwicklungslehre
die
dem
aufs engste mit
ganz und gar
Leben
verkettete
tierischen
sie
Menschen
den
und ihn dadurch
von
um so mehr mit
Es gewann das die Gemüter
ließ.
überwältigender
indem
die Natur hineinzog, ihn als ein bloßes Stück
in
erscheinen
ihr
erfahren,
weil
Kraft,
es
in
der
Wendung
zur Arbeit,
emsiger und fruchtbarer Arbeit, eine unermeßliche
zu
von Tat-
Fülle
sächlichkeit eröffnet, sonst zerstreute Daten verknüpft und als Ganzes
Wirkung
zur
langem
gebracht
zurückzukehren,
eine
Der Mensch schien nunmehr von
hat.
Wahn und eitler Selbstüberhebung zu seiner echten Heimat
sein
Leben aber damit
schlichtere,
frischere,
festere
Grundlagen,
sowie
Art
gewinnen.
Altes
wahrhaftigere
zu
erschien damit als neu. Neues als alt, eine durchgreifende Wandlung
Diese Strömungen und Stimmungen sind es, welche
Monismus ergreift und zusammenfaßt, die Naturbegriffe scheinen ihm nur einer gewissen Ergänzung in der Richtung
ward eingeleitet.
der -moderne
um den ganzen Umkreis der Wirklich-
des Seelischen zu bedürfen,
keit
in
sich
aufnehmen und
alles
Leben beherrschen zu können.
und Aussichten würden jedoch aus eignem
Vermögen schwerlich der Bewegung zum Monismus eine solche
Stärke verliehen und eine solche Macht über die Gemüter geAlle solche Vorteile
geben haben, wie jene
noch
etwas
anderes,
in
sie
das
Wahrheii
und große Massen zu erregen vermag,
entschiedenes
ein
dem Stande
Zwischen
der
bestand
leben
überbrücken
Nein.
der Religion,
Ein
Religion
vornherein
versucht,
solches
gesellte
Nein
sich
entzünden
nicht
entspringt
zum Ja
aber
und dem modernen
eine weite
mehr und
Kluft,
mehr aber
ist
lange
das
dazu
nicht
zu
wie ihn die kirchliche Gestaltung
überlieferten
von
käme
besitzt,
Leidenschaften
direkter
ist
aus
bietet.
Kultursie
zu
Unmögliche
dessen ersichtlich geworden, mehr und mehr hat sich der Abstand
in
einen
vollen
und schroffen Gegensatz verwandelt.
Und wenn
Probleme lange Zeit nur die obere Schicht der Gesellschaft
diese
zu berühren schienen,^ so sind
^
So erschien es z. B. einem
sie
immer mehr auch in die Massen
Pierre Bayle gänzlich ausgeschlossen,
die Aufklärung je die Massen gewinne.
daß
Er hält für die Interessen und die
Bedürfnisse der Gesellschaft, die nach seiner Überzeugung im wesentlichen
zu allen Zeiten dieselben bleiben, einen gewissen Aberglauben unentbehrlich
Zum Weltproblem.
186
gedrungen und erregen
trotz aller Wandlungen
offiziell
aufrecht
auferlegt
wird,
erhalten
so
immer
und
entsteht
ein
Wenn
stärker.
Druck und
schwerer
Der nun müßte
Psycholog und ein kurzsichtiger Staatsmann
viel
aber
zugleich
namentlich der Schule gebieterisch
Un Wahrhaftigkeit.
lähmenden
einer
sie
und Erschütterungen die alte Art der Religion
sein,
die
Gefahr
ein schlechter
dem entginge, wie
Unwille, wie viel verhaltener Zorn sich dadurch ansammeln
und
irgendwelchem Ausbruch drängen muß. Der
Monismus aber zeigt einen Weg, einen scheinbar nahen und leichten
Weg, auf dem ein solcher Ausbruch erfolgen kann; ist es ein Wunder,
wenn er wie ein brausender Strom die Gemüter ergreift und un-
schließlich auch nach
widerstehlich fortreißt?
Bewegung durchaus verständlich; sie würde nicht
wenn nicht ihr Ja wie ihr Nein Wahrheitsmomente enthielte. Aber es ist etwas anderes, eine Bewegung aus
der geschichtlichen Lage heraus zu verstehen und Berechtigtes in ihr
zu würdigen, etwas anderes, ihr die Führung des Lebens zu geben.
So
die
ist
wirken, was sie wirkt,
Was
zunächst die
Religion
betrifft,
so
dem Stande der
Art zu befreien und
jetzt
ist
bei
allen Kultur-
Bewegung im Gange, sie von der veralteten
völkern eine wachsende
weltgeschichtlichen
Evolution
des Lebens gemäß zu gestalten; solche Bestrebungen sind minder einaber sie sind auch
fach,
fruchtbarer
und
aussichtsreicher,
summarische Verwerfung der Religion, wie
sie
der
als
die
Monismus zu
Es kommt darauf an, ob die Religion über alle
Form hinaus in inneren Notwendigkeiten unseres Wesens
und unserer Stellung zum All gegründet ist oder nicht. Sollte sie
vollziehen pflegt
kirchliche
es sein, so könnten alle Schäden des gegenwärtigen Standes in keiner
Weise
das
Preisgebung einer Lebensmacht
die
des
Verhältnis
Menschen zum
rechtfertigen,
die
sich
Ganzen der Wirklichkeit zur
Aufgabe macht, die seinem Leben Größe, seiner Seele einen Selbstwert und eine reine Innerlichkeit zu geben unternimmt.
In dem
das Priestertum und die Kirche zu schädigen, pflegen die
Gegner der Religion zu übersehen, daß sie durch jene Verneinung
Eifer,
mit ihrer
Preisgebung
sich selber schädigen.
„Les besoins dont
et
des tenebres,
ils
je parle
ne sont point sujets aux vicissitudes de la lumiere
ils sont les memes sous un siecle
un siecle de science (s. den Artikel Franz von Assisi im
sont de tous les tems;
d'ignorance, et sans
Dictionnaire).
alles selbständigen Innenlebens am meisten
Man gedenkt dabei unwillkürlich der Anekdote
Monismus und Dualismus.
187
von dem Knaben, der keine Handschuhe erhalten hat, und dem nun in
„Es geschieht
arger Kälte die Hände erfrieren, der dazu aber meint:
meinem Vater ganz recht, daß mir die Hände erfrieren; warum gibt
er mir keine Handschuhe?"
Sachlich
des
Hauptfrage,
die
bleibt
Monismus entscheidet,
die,
über Recht oder Unrecht
die
ob die von ihm ausschließlich
ver-
wandten Naturbegriffe zur Umspannung der Wirklichkeit genügen.
Namentlich von zwei Seiten erheben sich dagegen Bedenken, die
einen von der Erkenntnislehre, die anderen
vom Gehalt des Geistes-
die weltgeschichtliche Arbeit
lebens her, wie
Erwägung
kenntnistheoretische
uns nicht von außen her
zufällt,
ihn zeigt.
— Die
er-
einzuwenden, daß das Weltbild
hat
sondern daß wir es von seelischen
Vorgängen aus und nach den Gesetzen unseres eignen Geistes zu
vom Subjekt pflegt sich zunächst
dem Forschen jene
Solches Ausgehen
bilden haben.
auf Kant zu berufen, dessen überlegene Energie
Richtung zwingend vorgeschrieben
hat.
Aber
es
nicht bloß Kant,
ist
es ist überhaupt kein einzelner Philosoph, sondern es ist die Gesanitart
des modernen Denkens,
Denn
nichts
ist
ja
Lebens, welche diesen
Weg befiehlt.
der modernen Kultur
dem modernen Leben und
eigentümlicher, als die Befreiung des Subjekts
von der Gebunden-
heit an die Umgebung, seine Befestigung im eignen Leben.
nicht auf
dabei
mit
Aufgebot
einen
Besitz
Kraft
aller
der Welt verzichtet,
leidenschaftlich
dieser
begehrt,
Wird
vielmehr
nimmt das
so
vom Objekt zum Subjekt,
von der Welt zum Menschen, geht es jetzt vom Subjekt zum Objekt,
vom Menschen zur Welt. Eine solche Umwälzung muß auch den
Leben eine
völlig andere Richtung:
ändern und sich damit in alle einzelnen
Inhalt des Lebens wesentlich
So auch
Gebiete erstrecken.
statt
in
wird
Ergebnis aus
dem Werden verstanden wird,
sich
Das
das des Erkennens.
Wirklichkeit
verfeinern,
beleben,
Bild der
wenn das
wenn volle Anerkennung
vergeistigen,
daß wir jenes Bild von innen her zusammenfügen, daß nicht
Außenwelt, sondern unsere geistige Organisation den Umriß
findet,
die
wie die Grundlinien
dazu
liefert,
daß eine
oft
sehr
komplizierte
Arbeit in Größen steckt, welche die naive Ansicht für einfach erachtet
Zugleich wird
klar,
daß wir mit aller Arbeit nur einen menschlichen
Durchblick
der Wirklichkeit gewinnen,
Forschung
selbst
wieder
der
einer
tieferdringenden
zum Probleme wird und seinen Wahrheits-
gehalt erst zu erweisen hat.
Neue Tragen und neue Sorgen tauchen
damit
unfertiger
auf,
unvergleichlich
müssen wir uns fühlen
als
Zum Weltproblem.
188
zuvor, aber in aller Unfertigkeit gewinnen
Dies
wir eine Vertiefung der
und auch unseres eignen Lebens.
Wirklichkeit
aber
alles
vorhanden; ihm
ist
ist
für
den Naturalisten und Monisten
die Welt, wie sie sinnlich
und handfest auf uns
einzudringen scheint, die ganze und echte Wirklichkeit.
heißt, erkenntnistheoretisch angesehen,
und
festhalten
sich
der
kopemikanischen
Denkweise der Neuzeit
wie ihn die
huldigen,
mittelalterliche Scholastik besaß, für die gerade der
Geringschätzung zu haben
Das aber
den ptolemäischen Standpunkt
dem naiven Realismus
verschließen, es heißt
nicht
Naturalismus nur
So vertritt die Philosophie gegen
pflegt.
den Naturalismus das Recht jener modernen Wendung zum Subjekt,
damit eine Wahrheit, die sich wohl verdunkeln,
vertritt
sie
aber aufgeben
Tiefer noch greift das Zweite, bei
Inhalt
ihm geht die Frage auf den
Diesen Inhalt faßt der Naturalismus und
der Wirklichkeit.
Monismus viel zu knapp,
er übersieht
das Hauptstück erscheint:
als
nicht
läßt.
das uns anderen
ein Stück,
das Geistesleben.
Alles
Innere
der
Natur einfügen kann jener nur, indem er das Seelenleben lediglich
ein
als
Vorgehen an den einzelnen Individuen behandelt
nämlich
mag
er
sich
darauf
wie fließend
berufen,
Dann
Grenzen
die
zwischen Menschem und Tier sind, wie langsam sich auf dem Boden
was früher als
der Geschichte emporgearbeitet
hat,
des Menschen
auch
galt,
wie
Macht der Naturtriebe
in
seiner
sehr
Das alles
bleibt.
ein Stammbesitz
der Kulturmensch
sei
unter
der
vollauf anerkannt
und
Bedeutung keineswegs abgeschwächt
Aber
es
nicht
ist
Denn das menschliche Seelenleben verbleibt nicht wie
das Ganze.
das tierische bei jener Zerstreuung an einzelne Punkte, bei ihm erfolgt
ein
Zusammenschluß zu einem gemeinsamen Leben, und
meinsame Leben entwickelt eine unermeßliche
wesentlich
dies ge-
Tatsächlichkeit,
neue Züge gegenüber der bloßen Natur erweist
Zusammenschluß
erst
macht Geschichte und Gesellschaft im
die
Jener
aus-
zeichnend menschlichen Sinne möglich, auf diesem Boden entsteht
Gedankensprache und Kultur, hier erwächst eine reiche Verzweigung
eigentümlicher
Lebensgebiete
Wissenschaft
Wie das Ganze,
biete ihre
eignen Gesetze,
in
Recht und Moral,
in
Kunst
und
so haben auch diese einzelnen Ge-
Probleme,
Menschen vor schwerste Aufgaben,
sie
Erfahrungen,
ziehen ihn
sie
stellen
den
immer mehr an
und machen zugleich unvergleichlich viel mehr aus ihm: aus
einem bloßen Stück der Natur wird er mehr und mehr ein geistiges
sich
Monismus und Dualismus.
Wesen,
das
die
von
Unendlichkeit
innen
189
her miterlebt
und
als
Persönlichkeit die Welt in eigne Tat zu verwandeln vermag.
sittliche
Eine so eingreifende Wendung kann sich nicht vollziehen ohne einen
neuen Anblick der Wirklichkeit; mit jenem allen verkündet sich deutUch
eine neue Stufe der Welt, deren Anerkennung das Gesamtbild des
erweitern
Alls wesentlich
und
muß.
vertiefen
Dies
alles
ist
keine
bloße Theorie, es hat im geschichtlich-gesellschaftlichen Leben der
Menschheit
es
Die
Wirkung.
Hauptaufgabe,
zur
Wirklichkeit
viel
eingearbeitet,
sich in alle Einrichtungen hin-
entfaltet,
umfängt uns mit bildender Kraft
deutsche
jenen
inneren
tausendfacher
darin
ihre
Zusammenhang des Menschenlebens
Anerkennung zu bringen,
vollen
in
fand
Philosophie
spekulative
Aufklärung weit überlegen, daß
sie
sie
fühlte
sich
dadurch der
die geistigen Inhalte
und Werte
aus jenem, nicht wie diese von den bloßen Individuen her erklärte.
Der Naturalismus aber übersieht jene Wendung zum
Geistes-
Kulturstandes,
Ausbildung eines eigentümlichen
jenes
leben;
jene
innere
Wachstum des Menschen durch die Arbeit der Jahrtausende,
der ganze Reichtum der dabei erschlossenen Wirklichkeit,
für
ihn
nicht
hängend
von allem
allem
vorhanden
gewürdigt;
er
charakteristisch
Lebensgehalt.
Was
oder
gibt
ihre
Größe
anderes
ein Verzicht
suchte.
ein
ein
Bild
aber
vom
sind
ist
auf alles das,
Alls
All
unter
Absehen
von allem Geistigen, von
Verwerfen
Das Bild des
sie
werden doch nicht zusammen-
Menschlichen,
Verengung und Verarmung,
der Geschichte,
sie
dies
alles
die
stärkste
inneren
Ertrages
als
worin die Menschheit
wird ohne den Menschen
und dann der Mensch unter Abschleifung aller
Wir sprechen von
überwundene
innerlich
Reaktion, wenn das Leben auf eine ältere,
Phase zurückgeschraubt werden soll. Aber wie bescheiden sind alle
fertig abgeschlossen,
Eigentümlichkeit möglichst in es hineingepreßt.
Versuche, es an einen Höhepunkt innerhalb der geschichtlichen Bewegung zu binden, gegen das Unternehmen, es ganz und gar an
die vorgeschichtlichen Anfänge zu ketten und ihm alle Möglichkeit
einer inneren Erhöhung, einer wirklichen Entwicklung zu nehmen?
Von hier aus angesehen, wird die ganze eigentümlich menschliche
Geschichte ein großer Irrtum, ein Abweg von der Wahrheit, indem
sie dem Menschen immer stärker eine Innenwelt vorspiegelt, die doch
nur eine
leere
Einbildung
ist.
Dabei erfahren wir das Verdrießliche, daß jene Verneinung eines
selbständigen Geisteslebens oft wie etwas Selbstverständliches auftritt,
Zum Weltproblem.
190
das nur Unverstand verkennen
oder böser Wille verwerfen könne.
Verneinende Richtungen waren
Begreifen läßt sich das freilich wohl.
besonderer Gefahr
eines
starren
Fanatismus gegen Andersgesinnte.
Nichts
in
stets
kritisches Verhalten
gegen sich
ja
eines
notwendiger für ein
ist
eine gerechte Würdigung
und
selbst
Dogmatismus,
anderer als die Fähigkeit, sich in fremde Denkweisen hineinzuversetzen
und von ihnen aus die eigne zu
alles Jenseitige als
als
nicht
betrachten.
Dies Vermögen aber
wo rasch der Kreis geschlossen und
wird besonders dort gefährdet,
vorhanden erklärt wird.
Hume war gewiß
Denker und Forscher groß und im Leben allem Fanatismus so
fern wie möglich,
aber gibt es einen krasseren Ausdruck eines
Fanatismus,
tellektuellen
als
jenes berühmte Autodafe
aller
in-
anders-
gerichteten philosophischen Literatur?
Das Gleichgewicht des Geisteslebens war lange genug von der
und der Theologie bedroht,
Religion
in
einem Rückschlag dagegen
wird es nun von dem ausschließlichen Herrscheranspruch der Naturwissenschaften gefährdet.
Naturwissenschaften
wie
Diesen Anspruch aber erheben weniger die
selbst,
ihre
als
Gestaltung
im Naturalismus und Monismus
sie
sehr daran zweifeln, ob der
vorliegt.
erachten, die Einheit der Gedankenwelt,
läßt sich
etwas großes
als
ob er nicht, indem er die
Begriffe gewaltsam zusammenschweißt, das
Leben als Ganzes inner-
Seine Begriffe und Lehren folgen der Natur, wie die
spaltet
lich
Philosophie,
Dabei
Monismus eben das erreicht, was ihm
Hauptsache gilt, und was auch wir anderen
als die
zur
mechanische Denkart
sie
faßt,
das
bloßen und blinden Tatsächlichkeit,
All
in
wird damit ein Reich der
dem es kein Handeln, sondern
nur ein Geschehen, keinen inneren Forttrieb, sondern nur ein Nebeneinanderlagern, keine wahrhaftige Einheit, sondern nur eine Zusammen-
setzung
gibt.
Bei
und Werte
halte
^
konsequenter Denkweise
verschwinden,
Jene Stelle (am Schluß der
hätte
auch
müßten
hier
alle
In-
der Wahrheitsbegriff
12. Sektion d. Enquiry conc. h. u.) lautet:
When we run over libraries, persuaded of these principles, what havock must
we make?
If
we take in our band any volume of divinity or school meta-
let us ask: Does it contain any abstract reasonings
conceming quantity or number? No. Does it contain any experimental
reasonings conceming matter of fact or existence. No.
Commit it then
physics
for
to the
flames.
instance;
—
For
it
can obtain nothing but sophistry and
Illusion.
—
Würde ähnlich ein spekulativer Philosoph urteilen, so würden alle es Borniertheit
oder Fanatismus nennen; geschieht es von der anderen
vielen als Zeugnis eines starken
Seite, so gilt es
und unerschrockenen Geistes!
Monismus und Dualismus.
191
und damit eine Wissenschaft keinen Platz. Was an geistiger Regung
das hätte ruhig und urteilslos das Weltgeschehen über
aufkäme,
sich ergehen zu lassen.
einem eifrigen Kampf
Hoffnung auf
ein
dessen
Statt
finden wir den
Monismus in
um die Wahrheit begriffen und von freudiger
Vordringen der
Menschheit
erfüllt,
er
hält
in
der Gestaltung des Menschenlebens die alten Ideale des Guten und
Wahren fest,
er schöpft
den Hauptantrieb seines wissenschaftlichen
Strebens aus der Überzeugung, dadurch
mehr Wahrheit und mehr
Vernunft in das menschliche Dasein zu
bringen,
hier
ganz und gar die
Wege
des
kurz er wandelt
Gibt es nun wohl
Idealismus.
einen schrofferen Dualismus, als Materialist in der Weltanschauung
und
Idealist
im Handeln zu sein?
Beispiel der alten Erfahrung,
So haben wir hier
daß der Mensch
das Gegenteil der eignen Absicht erreicht.
in
ein
neues
seinem Streben
oft
2. Entwicklung.
Zur Geschichte des Ausdrucks.
a)
|ie Ausdrücke für Entwicklung hat, wie den Begriff selbst, erst
I
-^ die Neuzeit
allgemeinen Umlauf gebracht.
in
erscheint in unserer Sprache erst gegen das Ende des
und findet erst in der zweiten Hälfte des
Älter
es
ist M
zuerst
Grimm
»Entwicklung"
1 7.
Jahrhunderts
18. eine weitere Verbreitung.
Auswicklung" (auch „sich auswickeln«), philosophisch hat
wohl Jakob Böhme verwandt.
Lexikograph
der
zuerst
„Entwickeln"
(„der
Stieler
hat
deutschen
nach
Sprache
Stammbaum", 1691) „sich entwickeln" Haugwitz (im Soliman, 1684),
sowie Hagedorn.
Die Gelehrten des
1 8.
Jahrhunderts sprachen oft
von einem Entwickeln und einer Entwicklung eines Begriffes, Beweises
und Satzes; „das Verfahren, wodurch ein Begriff ausführlich gemacht
wird,
heißt
die Entwicklung
Entwick-
(Lambert).
eines Begriffes"
lung im Sinne eines Sichentwickeins,
als Selbstentwicklung,
gewann
Boden erst mit dem Aufsteigen des deutschen Humanismus, dessen
Verlangen nach einer inneren Beseelung der Wirklichkeit und nach
Anerkennung bildender
Kräfte in der Natur darin einen bezeichnen-
den Ausdruck fand. Es genügen dafür die Namen Herders und Goethes.
Im Titel
1777
eines Buches erscheint Entwicklung bei Tetens, in seinem
veröffentlichten
Hauptwerk „Philosophische Versuche über die
menschliche Natur und ihre Entwickelüng".
nun „Auswicklung«, das in den älteren
wiegt,
gänzlich
zurück; auch
„Auswicklung" zu begleiten
„Entwicklung" drängt
noch vor-
Schriften Kants
„Einwicklung",
das
pflegte, verschwindet
als
aus
Gegenstück
dem Sprach-
gebrauch der Philosophie.
Der deutsche Ausdruck war
teils
verdrängte,
involutio
und
lateinischen
teils
eine Übersetzung fremder, die er
neben sich duldete.
Die Termini
evolutio-
explicatio-complicatio oder implicatio entstammen den
Klassikern,
aber
sie
werden dort nur methodologisch.
Entwicklung.
1Q3
Werden verwandt.^ Ähnlich verbleibt es im
Thomas von Aquino erscheinen nur explicitus und
implicitus, und zwar in jener formalen Bedeutung. Nur die mystische
das reale
für
nicht
Mittelalter,
bei
Spekulation, die von
den Schriften des Pseudo-Dionysius ausging,
um ein inneres Verhältnis von
verwendet die Wörter und Begriffe,
zum Ausdruck zu bringen.
So hat Scotus Eriugena
involutus, convolutus, complicatio, replicatio.
Wie Nikolaus von Kues,
Gott und Welt
der Philosoph an der Schwelle
knüpft, so gebraucht
er
Neuzeit, an jene Denkweise an-
d*er
fortwährend die Ausdrücke explicatio und
complicatio.
Wenn er evolutio gebraucht, so glaubt er es erläutern
zu
Der Verlauf der Neuzeit macht die Ausdrücke immer
sollen. 2
geläufiger.
Evolutio und involutio
enveloppement Lieblingswörter
des
18.
Jahrhunderts
sind neben
übernahm
sie
genannten »Einschachtelungstheorie".
in
developpement und
auch
Leibniz;
bei
Physiologie
die
dem Sinne
der später so-
Im Gegensatz dazu hieß dann
die besonders glänzend von C. F. Wolff in der theoria generationis
vertretene Lehre von einer Neubildung im Werden die der Epigenesis;^
die
»Evolution"
wurde
hier
verstanden und abgelehnt
als
eine
bloß
quantitative
Steigerung
Aber daneben behält Evolution auch den
weiteren Sinn von Entwicklung überhaupt, so
ist
es namentlich bei
außerdeutschen Völkern die gebräuchlichste Bezeichnung der neuesten
Form
der Entwicklungslehre geworden.
b)
Zur Geschichte des Begriffs und Problems.
Es gibt kaum eine Überzeugung und Lehre, bei der die
alte
und die neue Denkart so weit auseinandergehen; die Beharrungs^
Bei Cicero heißt es
z.
B. (Top. 9)
:
tum definitio adhibetur quae quasi
involutum evolvit id, de quo quaeritur.
'
Niliolaus sagt (Pariser
evolutio.
Ausgabe von
1514,
I,
89a):
linea
- Quomodo intelligis lineam puncti evolutionem? -
est
puncti
Evolutionen!
id est explicationem.
'
C. F. Wolff hat sich über jene Begriffe namentlich in der deutschen
Ausgabe und in der zweiten lateinischen Auflage von 1774 deutlich geäußert.
Praemonenda § 50 heißt es in dieser: evolutio phaenomenon est, quod, si
essentiam ejus et attributa spectes, omni quidem tempore, at inconspicuum,
exstitit, denique vero, speciem prae se ferens ac si nunc demum oriatur, quomodocunque conspicuum redditur. S. auch Kant, Krit. d. Urteilskraft (V,
436 Hart.): „Das System der Zeugungen als bloßer Edukte heißt das der
individuellen Präformation oder auch
die Evolutionstheorie;
das der Zeug-
ungen als Produkte wird das System der Epigenesis genannt."
Eucken, Grundbegriffe.
4. Aufl.
13
Zum Weltproblem.
194
den Idealen jener ebenso eng zusammen, wie die
lehre hängt mit
Entwicklungslehre mit denen der Neuzeit.
So
läßt
wiederum der
besondere Punkt einen Durchblick der gesamten weltgeschichtlichen
— Wohl
Bewegung erwarten.
zeigt die älteste griechische Philosophie
Höhe
bedeutende
Ansätze
klassischen
Kultur hat der Beharrungslehre das entschiedene Über-
Entwicklungslehre,
einer
aber
die
der
gewicht gegeben, da diese der künstlerischen Art jenes Volkes weit
mehr
in
und
entspricht
Wirklichkeit
gilt
Maße
feste
Grundbestande nach
und
gefügtes
von
Die
zu fassen geeignet war.
in -Begriffe
sie
hier ihrem
als ein
unwandelbaren
lebendiges,
Ordnungen
be-
herrschtes Kunstwerk; aus der wirren Flucht der sinnlichen Eindrücke
dies
und
klar
zum Hauptziel
das wird
kräftig herauszusehen,
der
Diese Aufgabe ist nicht zu lösen ohne die Anerkennung
Wissenschaft.
eines zeitüberlegenen Seins; aus der Übereinstimmung mit ihm stammt
Wahrheit der Begriffe, die
alle
Handeln
und
mitteilt
Wissenschaft
Werdens in
die
allem
des Seins,
Versetzung
eine
eines
freilich
So
aus der
ist
die
Welt des
lebendigen Seins; durch-
dem Werden voran.^
gängig geht hier das Sein
dem
aus sich auch
beharrende Ziele richtet
auf
es
vor
hier
vom Denken
Eine nähere Aus-
führung gibt dieser Denkart die Formenlehre, die Plato schafft und
Zeitlos wirken die Formen als Urbilder und
Aristoteles weiterführt.
Grundkräfte der Dinge, durch den Weltprozeß, der keinen Anfang
und kein Ende hat, gehen unverändert die Gestalten. Alle Veränderung
kommt vom Stoffe, der, wenigstens im Bereich des Erdenlebens,
sich der Form nicht
ihr ergriffen
Gestalt
und
verliert.
Stoff ergreifen
dauernd verbindet, sondern, eine Zeit lang von
immer wieder entweicht und seine
Immer von neuem muß daher die Form den
gebändigt, ihr
und bezwingen; das erklärt den unablässigen Wechsel,
Werden und Vergehen.
das rastlose
einzelnen Lebewesen.
geleugnet,
gilt
es
zunächst von den
Auch jenseit ihrer werden Verschiebungen nicht
vielmehr Wandlungen
Wandlungen
So
auch- im Auf-
in
den Stellungen
der Gestirne,
und Absteigen der Völker
bereitwillig
anerkannt. Aber auch dieses verwandelt sich einer näheren Betrachtung
*
Es sei dafür nur eine Stelle aus Aristoteles angeführt (de part. anim.
640a, 18): ^
b,
der
(S.
1:
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^vexa
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S'eoTt toioÜtov, -n^v ye'vEOtv üS\
Ausdruck für Wissenschaft wird
PhyS. 244 b, 9):
r
o
i^
«px.'i? ?>^'^'?
xcl^
a.\X
mit
ttj;
oOy^ f
ouata l'veza
-rf?
^evetew?.
zoiauTTjV au[j.ßaiv£tv ivaYy.atov.
Stehen
ir.iTzr^rii
^pefifaat xa\ arrivat ttv Siavotav eTiioraaö-ai xai (ppovetv
Selbst
Beziehung gebracht
Y£v£at; oüx eoriv tw y«P
in
Xe'yo(j.ev.
Entwicklung.
jg5
Denn aus aller Veränderung
in eine Bestätigung der Beharrungslehre.
ihrer Stellungen kehren die Gestirne nach Ablauf eines grofkn Welt-
um die alte Bewegung
jahres wieder an den Ausgangspunkt zurück,
neu zu beginnen, der Wechsel
Ähnlich
im Grunde
verändert also
nichts.
auch die Geschichte eine endlose Folge von Kreis-
bildet
bewegungen wesentlich gleichen
Denn alles Aufsteigen der
Inhalts.
Völker geht nur bis zu einem gewissen Punkt,
um
Sinken umzuschlagen,
von
bis
Elementarkatastrophen
dann
in
Feuer
ein
oder
Wasser wieder eine Verjüngung bewirken, so daß nun dieselbe Bewegung von neuem anheben kann. So eine ewige Wiederkehr der
Dinge; was wir jetzt erleben, geschah schon unzähligemal und wird
Demnach die Welt kein starres Sein,
noch unzähligemal geschehen.
sondern voller Bewegung, aber diese Bewegung, nach der Art von
Tages- und Jahreszeiten,
Rhythmus verlaufend und inmitten
in festem
alles
Wandels
feste
Grenzen gebannt, kein echtes Streben geht
es
voll
keinen
gibt
Ruhe.
sicherer
Fortschritt
Überall
Unendliche,
ins
das Leben
hier
ist
keine
in
Unbestimmte,
in?
Hoffnung einer
wesentlich besseren Zukunft.
Dafür aber gilt die Überzeugung, daß
die unmittelbare Gegenwart
Ewiges erfassen und mit ihm das Leben
erfüllen
kann.
nehmen,
sie
Die Tätigkeit
tut
das,
selbst hat hier die
indem
sie
sich selbst befriedigtes
und
des Aristoteles,
Eine derartige Tätigkeit
Stelle
wird.
Ruhe in
sich aufzu-
aus einem bloßem Streben
gesättigtes
ein
in
Wirken, „Energie" im Sinne
gewährt
an
erster
Anschauung des Wahren und Schönen, aber auch das
die
Handeln wird hier auf seiner Höhe zum Darstellen einer beharrenden Art und Natur.
Denkweise
Solche
Gutes,
in\
sieht
Veränderlichen ein
durchgängig im Unwandelbaren
Schlechtes.
ein
Das Hauptmerkmal der
Gottheit bildet hier die Ewigkeit, das Unberührtsein vom Strom der
Zeit.
Dem Handeln aber wird ein
gehalten, an
unwandelbarer Idealstand vor-
dem es sich messen und
auf den es sich richten
So vornehmlich im Entwerfen von Idealverfassungen,
schichtlichen Wandlung entzogen sind.
soll.
die aller ge-
Die Überzeugung, daß unser
Leben auf festen Grundlagen ruhe und sich innerhalb fester Grenzen
bewege,
selbst
gestaltet die Arbeit aller
Gebiete eigentümlich und verleiht
der Logik und wissenschaftlichen Methode einen besonderen
Charakter.
Die Grundwahrheiten
Urteilen gegeben
zustellen,
und abgeschlossen,
zu einander
in
dünken
es gilt
hier
in
Begriffen
und
nur sie deutlich heraus-
Beziehung zu setzen,
in ihre
Konsequenzen
13*
Zum Weltproblem.
196
zu
Die
verfolgen.
ward damit zum Hauptstück der
Schlußlehre
während der Neuzeit sich diese
Arbeit,
in die
Urteile
und
Begriffe
zurückverlegt hat.
Dabei floß
in die
Lehren der Philosophen von Anfang an auch
Stimmung des ganzen Menschen mit ein, das Verlangen, gegenüber dem bunten und ermüdenden Wechsel der kleinstaatlichen
Verhältnisse einen beharrenden und bedeutenden Lebensgehalt zu
gewinnen; die Wendung vom menschlichen Kreise zum All war
zugleich ein Suchen nach innerer Erhöhung und Befestigung des
'Daseins.
Solche Strömung verstärkte sich gegen den Ausgang des
Die Sache
Altertums und gewann neue Nahrung im Christentum.
die
ward nun vom Künstlerischen
ins Religiöse
gewandt; verlangte die
Höhe des griechischen Lebens ein Ruhen innerhalb der Bewegung,
so galt es jetzt, eine Ruhe in Gott im Gegensatz zu dem unsteten
und sinnlosen Treiben der Welt zu finden, sich dahin als in einen
Nicht
sicheren Hafen aus den Stürmen des Lebens zu flüchten.
streben wollte man, sondern fest
dere Kraft und
Kern
Innigkeit
Weisheit
aller
und sicher besitzen.
Eine beson-
gab solchem Verlangen die Mystik, den
konnte
sie
darin
finden,
die
ganze Zeit zu
einem bloßen Scheine herabzusetzen und durch ein wachsendes Aufgehen
ewige Sein jeden Tag »jünger" zu werden.
in das
ergriff in
Dies Ideal
den Zeiten des ausgehenden Altertums und des beginnenden
Mittelalters
um so mächtiger das Gemüt, als es der gesamten Lage
der Kultur entsprach.
Denn eine alte Art hatte sich hier ausgelebt,
ohne daß fruchtbare Keime einer neuen erkennbar waren;
selbst
den
Besten konnte es daher als Hauptaufgabe erscheinen, den vorhandenen
Besitz treu zu
wahren und gewissenhaft den künftigen Geschlechtern
mitzuteilen.
Unwandelbar schien vor allem
als göttliche
Offenbarung; aber auch auf den übrigen Gebieten,
Philosophie und Medizin, in Recht und
die religiöse Wahrheit,
Staat,
in
schien alles erreicht,
was dem Menschen irgend erreichbar ist; die Autorität eines Aristoteles
und
eines
Galen
kaum
stand
hinter der
des kirchlichen
Dogmas
zurück.
Diese Überzeugungen tragen
Mittelalters,
herstellt,
das
große Ordnungssystem des
das überall unwandelbare Normen und feste Verkettungen
innerlich wie äußerlich, namentlich auch in
lichen Verhältnissen, das
Leben
in sichere Bahnen
Trieb zur Veränderung aufkommen
es fern, die Natur als ein
läßt.
den wirtschaft-
bringt und keinen
Solcher Denkweise
liegt
Reich allmählichen Werdens zu verstehen.
Entwicklung.
X97
vielmehr hat diese nur die Formen zu bewahren, die der Schöpfer
ihr zu
Anfang
mitgeteilt hat.^
Die Neuzeit stand von vornherein zu einer solchen Beharrungs-
Denn
lehre feindlich.
ohne einen Glauben an
sie
konnte keine selbständige Art entfalten
die Bewegung und das Recht der Bewegung,
im Eintreten dafür mußte sie ihrem eignem Streben Boden gewinnen.
In
Wahrheit war gegenüber dem Ausgang des Altertums die Lage
der Menschheit erheblich verändert
Neue Völker voll
strotzender
Jugendkraft waren entstanden, die langen Jahrhunderte des Mittelalters
Vermögen gesammelt, das immer stärker nach Betätigung
die Welt mit eignen Augen zu
sehen und nach eignen Zielen zu gestalten; aus dem bloßen Empfangen und Überliefern treibt es zum Weiterbauen und Erneuern,
ein verändertes Lebensgefühl eröffnet neue Ausblicke und Aufgaben,
die Idee einer fortschreitenden Bewegung ergreift immer mehr das
hatten viel
drängte und sich zutrauen durfte,
Leben wie die Arbeit
Aber solchen Lebensdrang in eine sichere Bahn zu
keineswegs
leicht;
die
Geschichte der Entwicklungsidee
das nur im Anschluß an ältere Bestrebungen gelungen
es verschiedene Stufen durchlaufen hat.
reicht
in
das Christentum selbst zurück.
leiten,
war
zeigt,
wie
ist,
und wie
Die Anregung zum Neuen
Mochte seine kirchliche
Gestalt fest an der Beharrungslehre haften, seine Gedankenwelt enthielt auch fruchtbare Antriebe entgegengesetzter Art.
wird die Geschichte weit mehr
Zeit
in
als
dem
Dem Christentum
Mitten
Altertum.
in
die
war nach seiner Überzeugung das Göttliche eingetreten, nicht
mattem Abglanz, sondern mit ganzer Fülle seiner Herrlichkeit*
beherrschender Mittelpunkt des Ganzen mußte es
auf sich beziehen
und
alle
artigkeit dieses Geschehens
Zukunft aus sich
litt
alle
als
Vergangenheit
entfalten.
Die Einzig-
keinen Zweifel, Christus konnte nicht
immer von neuem kommen und sich kreuzigen lassen; so entfielen
die unzähligen Perioden, die ewige Wiederkehr der Dinge; aus einem
gleichmäßigen Ablauf von Rh>1hmen wurde die Geschichte ein um-
spannendes Ganzes, ein einziges Drama; daß der Mensch hier zu
^
Um nur eine bezeichnende Stelle dafür anzuführen, so läßt Alanus
de insulis (s. Baumgartner, die Philos. des A. d. J., S. 79) die Natur sprechen
Me igitur tamquam sui vicariam rerum generibus sigillandis monetariam destinavit,
ut ego
in
propriis incudibus
rerum effigies commonetans ab incudis
forma conformatum deviare non sinerem.
Zum Weltproblem.
198
einer völligen
Umwandlung aufgerufen wurde, das gab seinem Leben
größere Spannung, als wo es nur eine vor-
eine unvergleichlich
handene Natur zu
entfalten
So
galt.
Wurzeln
die
liegen
einer
höheren Schätzung der Geschichte und des zeitlichen Lebens nirgend
anders
im Christentum.
als
Aber nur langsam geschah es, daß solche Wandlungen prinzipiell
und deutlich ausgesprochen wurden;
gefaßt
es
ist
das vornehmlich
von der Spekulation her geschehen, die damals mit einem Verlangen
nach
gemütvoller
Aneignung Hand
Hand
in
Vor allem
ging.
suchte sie die Welt zu Gott in ein engeres Verhältnis zu
Was
und bedeutet
als
der naive Glaube es
all
ihrem Geschehen, von Gott aus betrachtet?
tat.
ist
die
bringen,
Welt mit
Sie kann, so lautet
die Antwort eines Augustin, nichts anderes als eine Selbstdarstellung
des göttlichen Wesens
einen
ihr
keit in
sein.
Damit aber gewinnt alle Mannigfaltig-
Zusammenhang, auch
inneren
die
verschiedenen
Punkte der Geschichte können nun kein bloßes Nacheinander bleiben,
sie
werden Stücke einer durchgehenden Bewegung,
weltumspannenden Tuns; auch
das,
was später
Früheren irgend schon gegenwärtig sein; so
ja
eines einzigen
eintritt,
mußte beim
sich
der gesamte
läßt
Weltprozeß der Entwicklung eines Baumes aus seinem Samen vergleichen. ^
Die mystische Spekulation eines Dionysius, Scotus Eriu-
gena
w.
u.
s.
steigert
dies
«Auswicklung" dessen zu
als eine
dahin,
fassen,
Ganze der Welt
das
was
„eingewickelt"
Entfaltung der Ewigkeit zu einem Leben
unsichtbaren Einheit zu sichtbarer Vielheit.
uns
sollten
freilich
nahe zu rücken.
*
lehre
in
als
eine
bei
Gott
ist,
der
Zeit,
der
Ihre Ausdrücke und Bilder
nicht verführen, diese Denkart der
Das begründende Sein und
die
modernen zu
bewegende Kraft
Bei dem allen ist Augustin der führende Geist; für seine Entwicklungs-
ist
namentlich bezeichnend folgende Stelle (op.
III,
148 D): Sicut in ipso
omnia simul quae per tempora in arborem surgerent:
ita ipse mundus cogitandus est, cum Deus simul omnia creavit, habuisse
simul omnia quae in illo et cum illo facta sunt, quando factus est dies, non
solum caelum cum sole et luna cum sideribus
sed etiam illa quae aqua
et terra produxit, potentialiter atque causaliter, priusquam per temporum moras ita exorirentur, quomodo nobis jam nota in eis operibus, quae
Deus usque nunc operatur. Wie er sich aber die Entwicklung eines Baumes
aus dem Samen vorstellt, zeigt V, 714 E: In illo grano seminis exiguo, vix
visibili, si consideres animo, non oculis, in illa exiguitate, illis angustiis et
grano
invisibiliter erant
—
radix latet
et
robur insertum
apparebit in arbore, jam
est
,
est et folia futura alligata
praemissus
in
semine.
sunt
et fructus,
qui
Entwicklung.
199
die Kette
blieben gänzlich jenseitiger Art;
Stufenfolge des Nacheinander war nicht
des Geschehens und die
dem eignen Boden der Zeit
entsprungen, sondern in ihr hatte die göttliche Einheit sich zeitlos
Wie
auseinandergelegt.
bedingt Höhere
Fülle der
Einheit.
diese
mit ihrer ewigen
Bewegung,
Zerstreuung
aller
und
hat
Unruhe überlegenen
mystisch-spekulative
die
Gedankenwelt die moderne Entwicklungslehre
gegeben.
mit solcher
hat
Größe gegenwärtig sein,
Forschung nicht
hätte nicht die Gottesidee, die
Leben und Glanz geliehen.
ihr
und Welt
Allerdings mußte im Verhältnis von Gott
Verschiebung
Unend-
Richtung auf Ewiges und
die
Die Welt würde der modernen
Idee des absoluten Wesens,
beträchtliche
Sie
eingeleitet.
der Erscheinung göttlichen Wesens, höher denken
als
und ihrem Leben
gelehrt
liches
das un-
als
ihr zurück zu jener aller Viel-
Aber trotz solches Abstandes
von der Welt,
Ruhe
so strebte auch das Leben letzthin nicht in die
Welt hinein, sondern aus
und
heit
galt,
erfolgen,
ehe
das
sicher
erst
eine
fortschreiten
konnte: die engere Verbindung von Welt und Gott durfte nicht dazu
dienen, die Welt gänzlich in Gott verschwinden
mußte dahin wirken,
sie
höheren Wert zu
zu lassen, sondern
ihr als dem Ausdruck göttlichen Seins einen
Diese Verschiebung
geben.
aber
ist
dem
bei
bahnbrechenden Philosophen der Renaissance, bei Nikolaus von Kues
(1401
—
— 1464), erfolgt.
Als die Auswicklung des unendlichen Lebens
die neuere Spekulation
pflegt
durch lebendig
sein, ja
eben damit aber einen Trieb zu
in sich
tragen.^
Was Gott besitzt,
un-
das hat
Nur ein Fortschritt ins Unendliche kann die Lebensfülle des absoluten
*
Seins zum Ausdruck bringen,
1514,
der Ewigkeit
muß sie an jeder Stelle nach Teilnahme am
unendlichen Leben dürsten,
begrenztem Fortschritt
beim Gottesbegriff
— muß ihm die Welt durch und
die Unendlichkeit voranzustellen
II,
aeternae sapientiae, et ex hoc
creatori
s.
sine fine.
II,
B. Nikolaus
z.
188a): Posse semper plus
et
elice,
von Kues (Paris. Ausg. von
plus intelligere sine fine, est similitudo
quod est viva imago, quae se conformat
187b: semper vellet id quod
intelligit
plus intelligere
mundus totus non sufficit ei, quia non replet
desiderium intelligendi ejus. — Der Begriff des Fortschritts ist dem Altertum
et
quod amat plus amare,
et
trotz der Vorherrschaft der Beharrungslehre keineswegs fremd, Plato und Aristoteles
haben dafür die Ausdrücke smSoai; und iTrtötSo'vai, weit mehr aber trat
r.poy.oTu hervor, das wir (z. B. bei Polybius) ganz ähnlich ver-
das stoische
wandt finden wie das heutige »Fortschritt". Der Gedanke eines Fortschritts
ins Unendliche hat seine Wurzel bei den Piatonikern und Mystikern, er ist
aber zur vollen Entwicklung
erst in
der neuen Philosophie gelangt.
Seinen
Zum Weltproblem.
200
die
Kreatur
Wachstum annähernd zu
allmählichem
in
der ganzen Welt ein Streben nach aufwärts eingepflanzt.
winnt,
in
erreichen.
Wendung wird die Bewegung wesentlich gehoben und
Mit solcher
dem
neben
Künstlerische
Indem
zurückzudrängen.
zum Ausgang
Gegensatz
direktem
Religiösen
Welt
die
Platz,
sich
ja
Zugleich ge-
des
Altertums,
es
beginnt
mehr und mehr
das
dieses
ein
als
lebensvolles Kunstwerk darstellt, in dessen Harmonie sich alle Gegen-
des ersten Anblicks auflösen, scheint
sätze
alle
Bildung von innen
hervorzubringen;
das
sie
die
Bewegung wie
durch Entfaltung ihres eignen Wesens,
her,
Absolute
bedeutet
nun weniger
ein
eignes
Reich, als es der Welt eine Tiefe oder doch einen Hintergrund gibt.
Solche Verschiebung
vom Theismus zum Pantheismus kommt zum
Siege in Giordano Bruno; die immanente und künstlerische Gestalt
der Entwicklungslehre hat von nun an die Oberhand.
Form, welche
bis zur
Gegenwart
in
Das
ist
die
den Ausdrücken und Bildern
von der Sache überwiegt, das stille und stetige Wachsen der Pflanze
wird hier
zum Vorbilde für das Wirken und Aufsteigen der Natur
von innen heraus.
in seelenlose
Die Aufklärung mit ihrer Zerlegung der Natur
Elemente
ist
dieser Gedankenrichtung
minder günstig,^
dagegen bringt der Rückschlag gegen die Aufklärung, wie ihn der
deutsche Humanismus enthält, sie voll zu Ehren. Nicht die bloße Be-
wegung, sondern das künstlerische Gestalten wird hier zum Haupt-
Höhepunkt
bildet
Leibniz,
s.
z.
B. 150a (Erdm.):
In
cumulum
etiam pul-
operum divinorum progressus quidam
universi est agnoscendus, ita ut ad majorem
chritudinis perfectionisque universalis
perpetuus liberrimusque totius
semper cultum procedat ff.; femer deutsche Schriften II, 36: «Der Kreaturen
und also auch unsere Vollkommenheit bestehet in einem ungehinderten
starken Forttrieb zu neuen und neuen Vollkommenheiten."
Bei Wolff und
seiner Schule galt als das höchste Gut perpetuus sive non impeditus ad
majores, per fection es progressus.
— Der Ausdruck Fortschritt dürfte
erst in
der
zweiten Hälfte des 18. Jahrhunderts ein fester Terminus geworden sein.
*
Es
fehlt
hier aber keineswegs an
wenig beachtete
Richtung.
S.
z.
(Nouv. ess.
III,
cp. VI, pag. 317a,
B.
die
anregenden Gedanken nach dieser
Stelle
in
Leibnizens
Hauptwerk
Erdm.): Peut-etre que dans quelque tems
ou dans quelque Heu de l'univers ies especes des animaux sont ou etaient
ou seront plus sujets ä changer, qu'elles ne sont presentement parmi nous,
et plusieurs animaux qui ont quelque chose du chat, comme le Hon, le tigre
et le
lynx pourraient avoir ete d'une
comme
meme race et pourront etre maintenant
des sousdivisions nouvelles de l'ancienne espece des chats.
Ainsi
que j'ai dit plus d'une fois que nos determinations
des especes physiques sont provisionelles et proportionelles ä nos connaisances.
je
reviens toujours ä ce
Entwicklung.
201
werke der Natur; damit wird ihre Veränderung zu einer Entwicklung
von innen
und
her,
Mannigfaltigkeit ihrer Gestalten scheint auf
alle
Über die Natur hinaus
einen einzigen Grundtypus zurückzukommen.
bemächtigt sich der Entwicklungsgedanke dann des Menschenlebens
und des ganzen Alls;
was
„alles,
in
der Wirklichkeit vorkommt«,
erscheint nun als „Entwicklung einer absoluten Vernunft" (s. Schelling I,
Die nähere Durchführung dessen zeigt verschiedene Färbungen;
481).
Romantik das ruhige Werden und Wachsen in den Vorder-
stellt die
grund, so gibt Hegel mit seiner kosmischen Logik der Entwicklungsidee
mehr
Immer aber
Selbsttätigkeit.
erfolgt
Bewegung von
die
innen heraus, immer wirkt, auch an der einzelnen
mit über-
Stelle,
legener Kraft das Ganze.
Das eben
es,
ist
was
die
der strengwissenschaftlichen
tümlich
stellt
künstlerische Entwicklungslehre
unterscheidet,
die
der Neuzeit
von
eigen-
Denn diese gibt alle inneren Zusammenhänge auf und
ist.
das Problem ganz und gar auf den Boden des
unmittelbaren
Zusammenwirken der Elemente soll den
lehren, und es soll aller Fortder
Natur
verstehen
Gesamtstand
Daseins:
das empirische
schritt in
einem
wird
Entwicklungsidee
die
Diese
Wissenschaft.
Nacheinander erfolgen.
zeitlichen
zu
einem
Hauptstück
das nächste Bild
setzt
diesem Sinne
In
modernen
der
der Dinge
zu
einer
bloßen Erscheinung herab, von der es zu den echten Beständen
durchzudringen
Elemente
Das
gilt.
aufsucht,
geschieht,
Gesetz
das
die
indem
Analyse
einfache
Wirkformen
dieser
ermittelt,
dann aber die Entwicklung die Welt neu
schichtlichen
macht
sie
die
und vom
ge-
begreiflich
der neueren Wissenschaft zur Hauptvertreterin
der Synthese, und erscheint
sie als
der gesamten Arbeit der
stein
aufbaut
Werden aus den vorgefundenen Bestand
So wird
erst
die
Gunst des Denkens,
ja
die
Vollendung und der Prüf-
Forschung;
kein
Wunder,
daß ihr
des modernen Menschen gehört
Es
erscheint aber die neue Entwicklungslehre gleich beim entscheidenden
Durchbruch der modernen Denkart; schon Descartes hat, wenn auch
nur
als
eine
Möglichkeit,
den
Gedanken,
nach und
nach
daß
der
gegenwärtige
(cum tempore, successive)
Weltstand in der
Zeit,
entstanden
Im Lauf der Jahrhunderte bemächtigt
^
sei.^
sich
dieser
Clauberg beschrieb im wesentlichen zutreffend (op. philos. 755) die
Methode folgendermaßen: Hanc methodum Cartesiana physica
cartesianische
omnes res naturales non statim quales sunt in statu perfectionis suae absoluto (ut vulgo fieri solet ab aliis), sed prius agit de quibusdam
tenens considerat
Zum Weltproblem.
202
Gedanke aller einzelnen Gebiete und gräbt sich immer tiefer in den
Stand des Wissens ein.^
Kosmologie weicht der antike Ge-
In der
danke einer Unveränderlichkeit des Himmelsgewölbes dem des allmählichen Werdens der Weltkörper und Weltsysteme (Kant, Laplace),
der Befund der Seele wird nicht nach alter Art als fertig hingenommen,
beschrieben und
seit
das
Locke,
sondern die moderne Psychologie strebt
eingeteilt,
Werden
und Wachsen der Seele von
Lebenserscheinungen
ner genetisch
Geschichte
als
erscheint
zu verstehen;
allmähliches
ein
schwindenden Anfängen her zu
Überall
befindlich behandelt.
Bestände und ein
bleibenden
einem
festen
Ganzen,
in
als
Fluß
von
ver-
und ähnlich
und Wandel
gegen die
ältere
Art der Anblick
war sie dort
ein
Herausheben der
ist
der Wissenschaft umgewandelt:
menschliche
die
Aufklimmen
unbegrenzter Höhe,
werden auch die einzelnen Kulturwelten
einfachsten
unmittelbares Zusammenschliefkn
künstlerisches
ein
Zusammenschauen
zu
der
nun das nur scheinbar Feste in Fluß,
dringt unermüdlich zu kleineren und kleineren Elementen vor und
Mannigfaltigkeit, so bringt sie
verwandelt die Wirklichkeit
einen
in
noch mitten im Werden be-
griffenen Prozeß.
Dabei scheint sie zu einer weit engeren Berührung
mit den Dingen
zu
trachtete;
so
heißt
kommen,
etwas in
die sie früher
die Entwicklung
mehr von außen
be-
hineinziehen,
voll
es
der Erkenntnis gewinnen.
Aber so wirksam
war,
die
moderne Entwicklungsidee schon lange
zum Siege für das Ganze
erst Darwin
gebracht.
des Lebens und der Arbeit hat
sie
Zunächst hat seine Leistung eine große Lücke
ausgefüllt: die organischen Formen widerstanden bis dahin einer gene-
tischen
Erklärung
hartnäckig,
so
blieb
eine
unüberbrückte
Kluft
zwischen den Weltbegriffen und den Erfahrungen des menschlichen
Kreises, die beide
der Entwicklung schon gewonnen waren.
Wohl
waren, namentlich durch Lamarck, bedeutende Anfänge zur Aufhellung
gemacht, aber den Versuchen fehlte die Verbindung zu einem durchgebildeten
Ganzen und zugleich
eine zwingende Kraft.
Darwin
hat
earundem principiis valde simplicibus et facilibus, deinde explicat, quomodo
paulatim ex Ulis principiis, suprema causa certis legibus opus dirigente,
oriantur et fiant, aut carte oriri aut fieri possint, donec tandem tales evadant,
quales esse experimur dum consummatae et absolutae sunt.
'
Einen wichtigen Abschnitt dieser Bewegung behandelt in vortrefflicher
Weise H. Heußler: „Der Rationalismus des
ziehungen zur Entwicklungslehre", 1885.
17. Jahrhunderts
in
seinen
Be-
Entwicklung.
203
von Deszendenz- und Selektionslehre jene
mit seiner Vereinigung
Lücke ausgefüllt und das Schlußstück des Ganzen
Lehre
namentlich dadurch stark, daß
ist
sie
seine
geliefert;
aus genauester Durch-
forschung ihres besonderen Gebietes Begriffe gewinnt, die einer unermeßlichen Ausdehnung nach allen Richtungen fähig scheinen; er
wie Helmholtz sich ausdrückt
hat,
II,
»alle
204),
—
(s.
popul. wissensch. Vortr. 2. Aufl.
vereinzelten Gebiete aus
Wunderlichkeiten
dem Zustande einer An-
den Zusammenhang einer
häufung
großen Entwicklung erhoben und an die Stelle einer Art von künsträtselhafter
lerischer
in
Solches Verdienst
Anschauung bestimmte Begriffe gesetzt."
erfährt keinen
Abbruch dadurch, daß
die Schranken der Selektions-
und ihrem Kampfe ums Dasein zu immer klarerer Einsicht kommen; hat doch Darwin selbst
diese Lehre nicht als die einzige Erklärung der organischen Formen
lehre mit ihrer natürlichen Zuchtwahl
Es bleibt dabei, daß durch ihn das Problem
ausgegeben.
neue Lage gebracht
ist,
eine
in
daß die Entwicklungsidee mit der Sicherung
auf dem Gebiet des organischen Lebens
zugleich die Fähigkeit er-
zum Ganzen einer Weltanschauung zu erweitern.
Für solche Wendung muß vornehmlich Spencers gedacht werden,
langt
hat,
sich
der zuerst von einer realistischen Denkweise aus die Entwicklungslehre ein eigentümliches Weltbild hat erzeugen lassen.
ist
ihm
ein
gilt
Stoffes
ihm Entwicklung,
als
Zusammenschluß
und Zerstreuung der Bewegung;
ihr
von Bewegung
in
Aufnahme
zunehmende Spezialisierung und Differenzierung, vom
als
Ganzen
die
(ab-
vom Gleichartigen ins Ungleichartige,
eine
in
des
aber bei ihm in
und Lockerung (disintegration) des Stoffes.
Dort erfolgt eine Verschiebung
bis
einen
(Integration)
folgt
endlosem Wechsel eine Periode der Auflösung
sorption)
in
Als die allgemeinste Tatsache der
zusammenhängenderen Zustand.
Welt
Entwicklung
Übergehen aus einem zusammenhangsloseren
einzelnen
Weltkörper,
die
in
Weltall
menschliche
Gemeinschaft, die Kultur, das Einzelwesen hinein; die Periode der
Auflösung verfolgt die umgekehrte Richtung.
Eine Verwandtschaft
solches Rhythmus mit Gedanken der ältesten griechischen Philosophie,
namentlich mit Empedokles,
Lehre, die im Grundriß
ist
nicht zu verkennen.
zeitlich
Wenn Spencers
voranging, der Darwins einen uni-
versalen Hintergrund gegeben hat, so hat sie durch die Verbindung
mit dieser unermeßlich an Fülle, Anschaulichkeit
keit
und Eindringlich-
gewonaen.
Trotz solches Vordringens der Bewegungslehre ist die Beharrungs-
Zum Weltproblem.
204
lehre zu
tief
und zu
eingelebt
mit den Überzeugungen wich-
fest
tiger Lebensgebiete verschmolzen, als daß sie nicht vielfachen
Im besonderen
stand leisten müßte.
nur einzelne Bestandteile
die Religion,
es
ist
Wider-
die nicht
herkömmlichen Vorstellungskreises,
ihres
sondern auch den ihr unentbehrlichen Gedanken einer ewigen Wahr-
Aber auch
heit
bedroht
die
Überzeugung, daß
sieht.
hier befestigt
als
selbst,
mit Überzeugungen material-
oder doch naturalistischer Art
istischer
mehr und mehr
weniger die Bewegungslehre
es
ihre, keineswegs notwendige Verquickung
lichen Gegensatz
sich
ist,
zur Religion enthält. ^
welche einen unversöhnAlles
zusammen hebt die
Entwicklungslehre weit über die Stellung einer besonderen Theorie
neben anderen hinaus, sie hat die Führung des Ganzen übernommen
und erzeugt einen neuen Lebenstypus, der ebenso unser Grundverhältnis
lichste
zur Wirklichkeit
verändert.
Nun
als
Art unseres Tuns aufs wesent-
die
mehr eine fertig vorhandene
heißt es nicht
Wirklichkeit anzueignen, sondern einer erst werdenden zur Vollendung
zu verhelfen; die Tätigkeit verschlingt sich dabei enger mit der Um-
gebung und gewinnt ihre nähere Gestalt erst aus der Berührung mit
den Dingen.
Sie
darf sich
der Welt zu stehen
fällt
sagen,
und zu
die alte Flucht aus
nicht
neben, sondern inmitten
ihrer Bildung mitzuwirken.
Nun ent-
dem Strom der Zeit zu einer wandellosen
Ewigkeit, sowie die Vorhaltung eines Idealstandes als eines unverrück-
baren Zieles, vielmehr heißt
Zeit zu
folgen
Es
*
es,
ganz und gar der Bewegung der
und das Handeln den Forderungen der
jeweiligen
dafür nur eine Stelle des hervorragenden französischen Theo-
sei
Er sagt in seiner vielbeachteten Rede
Methode der Theologie (s. Bulletin de litterature ecclesiastique
Nov. 1901, pag. 272): Vous savez avec quelle defiance justifiee fut regue
logen Erzbischof Mignot angeführt.
über
die
l'idee d'evolution, qui paraissait liee par
il y a trente ans,
de graves compromissions avec la philosophie pantheiste; depuis que l'analyse
en a precise le contenu, on est ä peu pres unanime ä reconnaitre qu'une
dans nos ecoles,
d'entendre
une conception
germe dans saint
avec
certaine
fagon
religieuse
et
chretienne
Augustin,
et
on deeouvre, avec Vincent de Lerins, qu'appliquee ä l'histoire
religieuse,
eile
seraient
restes
forschung und
peut
de
l'evolution
est
conciliable
l'univers;
on
en
apporter
insolubles.
de grandes
Auch
evolutionistische
trouve
clartes
dans des problemes qui
Reischle: „Wissenschaftliche Entwicklungs-
Weltanschauung
in
Christentum" (Zeitschrift für Theologie und Kirche,
scheidet scharf zwischen
lungstatsachen.
innert sein
(s.
le
Evolutionismus
Auch an Newman's
als
theory
darüber Lady Blennerhassett
,
ihrem Verhältnis
zum
Jahrgang,
Heft)
12.
1.
Weltanschauung und Entwickof development mag hier er-
Kardinal
Newman S- 125 ff).
Entwicklung.
Das
Lage möglichst genau anzupassen.
aus träger Starrheit auf und bringt
z.
Von
wart.
in
der lebendigen Gegen-
Ergreifung des unmittelbaren Augenblicks und der Gestaltung
nach seinen Bedürfnissen, eine Elastizität des Lebens,
Wandlungen
bereitwillig
entgegenkommt.
dem Ausdruck Hegels das Werden
.,die
so müssen auch die Ideale die Beweglichkeit
temporis
(veritas
völligen
älteren
Denkweise hat
Hauptziel
Das
filia).
einem
Relativismus,
ein
nicht
gilt ja
Bildet
so werden auch
»Kind der Zeit"
ein
unterwirft augenscheinlich
aber
mit
dem
so
Wahrheit des Seins",
teilen,
die Wahrheit
die Ziele veränderlich, so wird
als
Lebensgebiete
alle
frischesten Fluß, das gibt
hier aus erwächst ein eigentümlicher Begriff des Modernen,
weiteren
nach
rüttelt
mit den Aufgaben
erfüllt sie
aller Verhältnisse
die
sie
Gesetzgebung wie der Erziehung eine engere Beziehung
B. der
zur Zeit und
als der
205
das
Leben
Hinfälligwerden
solcher allen Schrecken verloren.
mehr die Aneignung
einer
der
Denn
um uns vor-
handenen und fertig abgeschlossenen Wahrheit, sondern die Erzeugung
eines möglichst reichen Lebens
mehr
jene
im eignen Kreise; dafür aber scheint
Art mit ihrer unbegrenzten
relative
Anpassungsfähigkeit besonders geeignet.
bei
Bestrebungen
inneren
Beweglichkeit und
alles
und Wandlungen, auch
modernen Lebens
staltung des
Das
hat jene
verbleibt
die
nicht
äußere Ge-
Verwandlung des Daseins
in eine rastlos fortschreitende Bewegung aufs wirksamste unterstützt.
Die Technik hat den Lebensprozeß
in
ungeahnter Weise beschleunigt,
den Augenblick bedeutender gemacht,
die
Berührungen
und mit
ihnen die Wandlungen der Dinge unermeßlich gesteigert; bis in ihre
Werkzeuge hinein
griffen. ^
Mit
ist
jetzt
die Arbeit in unablässiger Wandlung be-
dem allen scheint der Sieg der Bewegungslehre end-
gültig entschieden
und mit ihm
ein
kräftigeres,
freieres,
frischeres
Leben errungen. ^
*
Die Konsequenzen dessen für die sozialen Probleme hat namentlich
K. Marx
in eindringlicher Weise dargelegt; er sagt (Das Kapital, I, 479):
„Die moderne Industrie betrachtet und behandelt die vorhandene Form eines
Produktionsprozesses nie
während
revolutionär,
als
definitiv.
Ihre technologische Basis
ist
die aller früheren Produktionsweisen wesentlich
daher
kon-
servativ war."
*
Merkwürdig genug ist, daß in derselben Zeit, wo die Entwicklungslehre
so siegreich vordringt, der Forschung ernste Besorgnis über den bleibenden
Fortbestand
Wärme
des
nur von
Lebens erwachen;
das
aber von
daß
und daß
dem das Leben auf-
der Tatsache
heißeren zu kälteren Körpern übergehen kann,
damit das Weltall einem Gleichgewicht zustrebt, bei
aus,
Zum Weltproblem.
206
Die Verwicklungen und Schranken der bloßen
Entwicklungslehre.
c)
Das alles
und
hat seine Wahrheit
Recht; töricht wäre
sein
es,
sich einem solchen Strom von Tatsächlichkeit entgegenzuwerfen, kleinlich, an ihm einzelne Irrungen
aufzusuchen und bei ihnen zu verweilen.
Aber daß Welt und Leben darin aufgehen, daß der Kampf zwischen
Bewegungs- und Beharrungslehre schon endgültig ausgekämpft
ist,
das dürfte mit allen jenen Wandlungen noch keineswegs entschieden
sein.
Es müßte wunderbar zugehen, wenn die Idee der Entwicklung
selbst
ohne
wäre, wenn eine Strömung, welche die
alle Verwicklung
Zeit so überwältigend fortreißt, nicht viel Ungeklärtes enthielte, wenn
das
Sehen
ausschließliche
Menschen
des
nach
einer
Richtung ihn nicht nach anderen Seiten vieles übersehen
besonderen
ließe,
seien
es Ergänzungen, seien ts Widerstände. Nach dem Plane unserer Arbeit
unsere Untersuchung namentlich dieses erwägen, was die Be-
soll
hauptungen und Wandlungen dem
Lebensprozesse
wie er
leisten,
unter ihrem Einfluß gestaltet, im besonderen, ob er dabei einen
sich
geistigen Charakter zu
des Geisteslebens
keit
bewähren
Daß
wahren vermag.
ist
es,
dem
alles
Unternehmen
sich
zu
hat.
der modernen Entwicklungslehre verschiedenartige Ten-
in
zusammenwirken,
denzen
an
Das Problem der Möglich-
das
verraten
schon
die
Bezeichnungen.
Wer von „Entwicklung", von »Evolution" spricht, scheint anzunehmen,
daß
in
sich die
Dinge von innen
her,
nach einem Gesetz des Ganzen,
sicherer Richtung auf ein Ziel entfalten.
Das aber will der über-
Zug der Gegenwart nicht, vielmehr erwartet er allen
Fortschritt von dem Zusammentreffen von Elementen, die von Haus
wiegende
aus
gegeneinander
gleichgültig
sind,
Summierung kleiner Verschiebungen,
hören
müßte.
besteht,
und
werden, wie
Indessen
hier
sie
mag
fragt
auf
sich,
die
sowie
von
er verwirft
einer
alle
langsamen
inneren Ziele
ob nicht eine Gegenwirkung dagegen
Lehre
vom
Strahlungsdruck
hingewiesen
namentlich durch Arrhenius zu jenem Problem
in
frucht-
So kommt Arrhenius zu dem Ergebnis (s. das
Werden der Welten, deutsche Übers. S. 190): »Durch dieses kompensierende
Zusammenwirken von Schwerkraft und Strahlungsdruck, sowie von Temperaturausgleich und Wärmekonzentration, wird es möglich, daß sich die Weltentwicklung in einem fortwährenden Kreislauf bewegt, bei dem wir weder Anfang
noch Ende wahrnehmen können, und bei dem auch das Leben Aussicht hat,
beständig und unvermindert weiter zu bestehen."
bare Beziehung gesetzt
ist.
Entwicklung.
207
und Richtungen, er verwirft alles Wirken aus einem Ganzen. Aber
wozu dann der andersartige Ausdruck, der unvermeidlich den Schein
einer von innen her sicher und ruhig fortschreitenden Bewegung
erzeugt?
Gibt er nicht einem seelen- und sinnlosen Weltbilde ein
freundliches Ansehen,
zu
viel
und Zerstörungen,
versteckt
er
nicht
Verneinungen
die
die jenes Weltbild enthält?
Solche Bedenken stören indes die Durchschnittsmeinung wenig.
dem Gedanken
Berauscht von
Endlose,
ins
Dinge
einer Entwicklung,
fühlt sie kein Bedürfnis
das Woher und Wohin
geringer die Präzision,
je
Je
ist
die Begeisterung.
die Tatsache nicht zu verkennen, daß in den leiten-
Jedenfalls
ist
Systemen
der
Fassung
überwiegt
erscheint.
Die
modernen Entwicklungslehre
und
die
als
ist
dadurch
mit
ihrer
des Problems
künstlerischen
weit zurückgedrängt;
mag er im Verborgenen stärker fortwirken als die
die Herrschaft auf
mechanische
eine
endgültige Lösung
Entwicklungslehre
ältere
oder logischen Art
ist
machen.
zu
vager die Fassung, desto sicherer scheint
die Sache, desto summarischer
,
Viele
ohne über das Was und Wie,
irgendwelche Sorge
sich
aller
nach einer präziseren Fassung.
begeistern sich heute für Entwicklung,
den
eines Fortschritts
unbegrenzten Besser- und Besserwerdens
eines
von Hegel,
meisten
denken,
Darwin übergegangen.
Bei Darwin und im Darwinismus sind die beiden Hauptgedanken
der
Deszendenz
und
Die Deszendenzlehre
der
ist
Selektion
deutlich
auseinanderzuhalten.
von so verschiedenen Seiten her
bestätigt
und hat sich so unermeßlich fruchtbar erwiesen, daß über sie in
kaum noch ein Streit besteht. Die Selektionslehre
der Wissenschaft
dagegen,
die
zeitweise
die
Forschung überwältigend
fortriß,
hat
mehr und mehr Widerstand gefunden. Daß sich das ganze Reich
der Formen lediglich aus einer Ansammlung zufälliger individueller
Variationen,
Beharren,
durch
ein
blindes Zusammentreffen
das hatte den überwiegenden
gegen
^
E.
und
tatsächliches
ohne irgendwelche innere Gesetzlichkeit, aufbauen
solle,
Zug der Philosophie von Anfang
an
sich,^
das hat sich mehr und mehr auch der Naturwissenschaft
Es
hier
ist
namentlich der unermüdlichen und scharfsinnigen Arbeit
von Hartmanns zu gedenken, der ebenso von der Spekulation wie von
den Tatsachen her die Unzulänglichkeit jener Lehre mit überlegenen Gründen
dargetan hat. In seiner neuesten Behandlung der Frage: »Die Abstammungslehre seit Darwin" (in den Annalen der Naturphilosophie II, 3) faßt er S. 354
das Ergebnis der Forschungen der letzten Jahrzehnte dahin zusammen: «Die
Zum Weltproblem.
208
als
Gerade auf dem eignen Boden der Ent-
unzulänglich erwiesen.
wicklungslehre findet diese besondere Fassung mehr und mehr Widerspruch.
Es sei hier, wo sich ein näheres Eingehen auf diese Probleme
verbietet,
nur an die Weismannschen Theorien, an die Entwicklungs-
mechanik, an die Mutationslehre erinnert.
die eigentümlichen Züge
und
Dieselbe Bewegung, welche
die Probleme des Lebens wieder mehr
muß auch dem Abschluß bei einer mechanischen Ent-
hervortreibt,
wicklungslehre
und
widerstehen
eine
dynamische empfehlen.
So
geschieht es vielfach in Wiederaufnahme und Weiterführung Lamarck-
scher Gedanken, so geschieht es in gleichzeitiger scharfer Kritik einer
bloßmechanischen
Entwicklungslehre,
die
her und aus dem Ganzen leugnen muß.
daß jene Lehre mit ihrer Verneinung
gemacht,
triebs alles wesentliche Fortschreiten
Grunde
im
Entwicklungsgedanken
dieses,
alle
Bildung
von innen
Es wird dagegen u. a. geltend
alles
inneren Fort-
des Lebens und zugleich
Nicht
aufgebe.^
den
minder auch
daß jene Lehre nur dadurch zu einem leidlichen Abschluß
daß
gelange,
sie
schon den Elementen beilege,
der Bildung sichtbar
vorliegt. ^
was auf der Höhe
Es stehen hier große Linien neben-
Selektion kann überhaupt nichts Positives leisten, sondern nur negative, aus-
schaltende Wirkungen entfalten.
Die Entstehung neuer Arten durch minimale
Abänderungen ist möglich, aber nicht erwiesen und, seit man den undulatorischen Charakter der minimalen Abänderungen kennt, weniger wahrscheinlich
geworden die sprunghafte Abänderung ist jetzt in den Vordergrund getreten.
Die Zufälligkeit weicht einer bestimmt gerichteten, planmäßigen Entwicklungstendenz aus inneren Ursachen, und diese bekundet sich ebensowohl in den
Der Anspruch des Darkleinsten wie in den sprunghaften Abänderungen.
;
winismus, zweckmäßige Resultate aus rein mechanischen Ursachen erklären
zu können,
^
ist
ganz unhaltbar."
Bergson bemerkt (L'evol. creatrice, pag. 40)
mecaniques
est
en
effet
:
L'essence des explications
de considerer l'avenir et le passe comme calculables
en fonction du present et de pretendre ainsi que tout est donne.
verficht
die idee d'un elan
originel
de
la vie,
Er selbst
passant d'une generation de
germes ä la generation suivante de germes par l'intermediaire des organismes
developpes qui forment entre les germes le trait d'union. Cet elan, se conservant sur les lignes d'evolution entre lesquelles
il
se partage, est la cause
profonde des variations, du moins de Celles qui se transmittent regulierement,
En general, quand des
commence ä diverger ä partir d'une souche commune, elles ac-
qui s'additionnent, qui creent des especes nouvelles.
especes ont
centuent leur divergence ä mesure qu'elles progressent dans leur evolution.
Pourtant,
sur
des
points
definis,
elles
pourront
et
devront
meme evoluer
identiquement si l'on accepte l'hypothese d'un elan commun.
'
S. Oliver Lodge,
fach das zu Erklärende
Leben und Materie S. 47: „Man konstatiert hier einund schiebt es dann den Atomen zu, in der Hoff-
Entwicklung.
209
und der Kampf schwankt noch hin und
einander,
dünkt heute die Lage minder
als begeisterten
einfach,
als
sie
her,
jedenfalls
weniger Darwin selbst
Jüngern erschien.
merkwürdig gfenug, gewinnt dieselbe Selektionslehre,
Aber,
die
dem Gebiete ihres Ursprungs mehr und mehr kritisch behandelt
auf
und eingeschränkt wird,
darüber hinaus,
in
der allgemeinen
trachtung menschlicher Dinge, noch immer an Boden.
auf
überall
weitverbreitet,
ist
zugreifen, die den
möglichst
einfache
Be-
Die Neigung
Anfänge zurück-
Menschen dem Tiere nahe verwandt zeigen, und
Bewegung aufwärts nicht auf einen inneren Trieb, sondern auf
die
Notwendig-
ein allmähliches Weitergestoßenwerden durch die äußeren
keiten
zurückzuführen,
gebungen und
sie
als
eine
Lebensbedingungen
bloße Anpassung an
zu
Um-
die
anderes
Nichts
verstehen.
scheint dabei in Frage zu stehen als das natürliche Dasein, der Sieg
im Kampf gegen die Mitbewerber.
Das vermeintlich Höhere bringt
dann nichts wesentlich Neues, sondern nur Kombinationen und Variationen der elementaren Lebenserscheinungen; damit
daß dem Geistesleben
Die Wandlung der Begriffe, die damit
gebührt.
ist
auch gesagt,
keinerlei Selbstständigkeit gegenüber der Natur
erfolgt,
reicht tief in
wo alle Lebensentfaltung
im Kampf ums Dasein zurückkommt, wo alle
die Gestaltung der einzelnen Gebiete hinein;
Erhaltung
auf die
geistige Betätigung ein bloßer Anhang des physischen Daseins wird,
da wird
zum Wert
eines an
sich
Wahre nur
der Werte das Nützliche,
Guten zu einer leeren
in
dem
Sinne
einer
Illusion,
da sinkt der Begriff
da
der Lebenserhaltung
Gruppierung der Vorstellungen bestehen bleiben.
auch
kann
das
förderlichen
Ethik,
Ästhetik,
Erkenntnislehre müßten sich damit völlig verwandeln, sie alle müßten
Lösung ihrer Probleme von den\ Ergreifen und Festhalten der
die
ersten
Anfänge erwarten.
Das Ganze wirkt mit der frischen Kraft einer eben errungenen
Einsicht, es läßt in alten
zerstreute
seiner
Daten
in
Erfahrungen Neues sehen, es bringt sonst
einen aufhellenden
Zurückwendung des
Blickes
Zusammenhang,
es
mancher Entdeckungen
ist
mit
fähig.
Die Naturbedingungen unseres Daseins, das Fortwirken elementarer
daß damit dem Fragen ein Ende gemacht sein wird. Bergson findet
den Fehler des Evolutionismus Spencers darin: Ȋ decouper la realite actuelle,
dejä evoluee, en petits morceaux non moins evolues, puis ä la recomposer
nung
avec
ces fragments,
et
ä se donner ainsi,
par avance,
tout ce qu'il s'agit
d'expliquer."
Eucken, Grundbegriffe.
4. Aufl.
14
Zum Wcltproblem.
210
Triebe inmitten
und
Langsamkeit
die
Kultur,
aller Verwicklung
und scheinbaren Vornehmheit der
der
Schwerfälligkeit
geschichtlichen
kommen nunmehr zur vollen Geltung; das Bild
unseres Daseins scheint mit dem allen mehr Naturfarbe und LebensBewegung,
sie
wahrheit zu erreichen, zugleich aber gewinnt das Wirken zur Hebung
der menschlichen Lage festere Angriffspunkte.
Aber das alles ist in ein größeres Ganzes aufzunehmen und aus
seinen
Zusammenhängen zu würdigen, um
Vernunft zu dienen
;
rein
der Wahrheit und
und
sucht es bei sich selbst den Abschluß
will
es aus eignem Vermögen die Gedankenwelt gestalten, so sind schwere
nicht
zu
vermeiden.
Sie
kommen
zurück,
die
besondere
Art,
in
Irrungen
Fehler
der
namentlich
beim
sich
auf
den
iMenschen
Geistesleben und Vernunft entwickelt, als den schaffenden und treiben-
den Grund des Geisteslebens selbst zu behandeln; wird aber dieses
so von vornherein zu einer bloßen Erscheinung
gesetzt
und
aller Selbständigkeit
beraubt,
von der bloßen Natur keine Mühe
kreise dieser
so
bereiten.
am Menschen herab-
kann
seine Ableitung
Wer nicht im Bann-
Denkweise steht, wird alsbald den Zirkel
in jener
Er-
klärung gewahren und die dort vollzogene Umwandlung des Geistes-
Vernichtung
lebens als eine
erkennen.
schließlich 'das Geistesleben selbst
Die
geistigen
Liebe,
Treue
und
Ein Gutes
unter das Nützliche nicht sowohl verändert als zerstört
Recht, Ehre,
Werte
werden durch jene Unterordnung
— das wegen
,
seiner
—
Nützlichkeit, d. h.
als ein
bloßes Mittel für die physische und soziale Lebenserhaltung
erstrebt
wird,
wird
Charakter des Guten.
damit
innerlich
verwandlelt
und
verliert
den
Ebenso müßte es dem Begriff der Wahrheit
wenn er zu einer bloß zweckdienlichen Anordnung unserer
ei gehen,
Vorstellungen sänke;
Wahrheit
ist
er
mag dann alles mögliche andere
er nicht mehr.
werden,
Einer solchen Erniedrigung des Lebens
widersteht aber das innere Erlebnis, das Gewisseste von allem, was wir
Denn mag über die nähere Fassung des Guten und Wahren
viel Streit sein, und mag der Einzelne noch so wenig teil
an jenen Größen haben, als bloße Lebensmöglichkeiten sind sie
kennen.
noch so
Tatsachen, die sich schlechterdings nicht wegerklären lassen,
aus dem Ganzen der Wirklichkeit etwas anderes machen.
kommt hier in
die
Rede
nisches
Frage,
sein kann.
Getriebe
und
die
Schließlich
ob überhaupt noch von einem Geistesleben
Wird unser ganzes Seelenleben
elementarer
Kräfte
verwandelt,
so
in
ein
gibt
mechaes
kein
Leben aus dem Ganzen, kein Denken, kein erlebendes Subjekt, so
Entwicklung.
müßte der Urteilende auch
und
alle geistige
und
nicht tut
zum Verschwinden
selbst
Arbeit für ein Wahngebilde erklären.
Verneinung
selbst,
bringen
Solange er das
Form der Aussage ihren
nicht tun kann, widerlegt die
bestätigt die
Inhalt,
sich
211
die als wissenschaftliche
und
allgemeingültige Wahrheit vorgetragene Verneinung, das Wirken eines
dem Naturprozeß überlegenen Geisteslebens.
Zu diesem Widerspruch, durch die geistige Arbeit ihre eignen
Grundbedingungen aufzuheben,
gebung
aller
bloße
Natur
genauer
selbständigen
betrachtet,
unseres Lebens.
wie
der Menschheit,
Verzweigung,
sie
und
die
Denn,
auftritt.
Aufbau der Kultur mit seiner
die physische Existenz zu
Dasei;!,
Befreiung
keine andere Aufgabe,
hätten
der
daß jene Preis-
Wendung allen Sinn und Wert
Mühe und Arbeit des Menschen
jene
unsägliche
aller
Verwicklungen
allem,
und jene Bindung an
Geistigkeit
zerstört
Alle
sich
vor
ist
Erhöhung
eine
als
gesellen
Wunderlich
näheren Ausführung.
erhalten,
das
als
auf einem
reichen
sinnliche
ungeheuren
Umwege zu leisten, was das Tier so viel leichter und einfacher erreicht.^
Allts,
was der physischen Existenz gegenüber einen Selbst-
zweck und einen Selbstwert behauptet, müßte als unhaltbar verschwinden, irgendwelchen
Nun
Inhalt
könnte
ein
aber sind wir einmal denkende
solches Leben
nicht
und
Wesen,
urteilende
bieten.
nun
haben wir den Mittelpunkt eines Selbst und müssen darauf alle Erfahrung beziehen und
jene
Inhaltslosigkeit
sie
als
von da aus messen.
So werden wir denn
schmerzliche
empfinden,
eine
Leere
eine
um so unerträglicher wird, als diese Zusammenhänge nicht
Leere, die
die mindeste Hoffnung einer Wandlung gewähren, uns vielmehr das
Getriebe des Naturprozesses
sinnlose
es
trostlosere Lebensgestaltung
eine
unablässiger Arbeit ohne allen
Aufbietung
aller
inneren
Es
sei
hier an
erinnert (V, 65 Hart.):
aus
soll,
der
was
festhält.
Gibt
mit ihrer Forderung
Gewinn,
ihrer fieberhaften
bei Tieren
vermeintliche
Seite
gerät diese Denkweise
das Wort Kants aus der Kritik der prakt. Vernunft
Ȇber
die
bloße Tierheit erhebt ihn (den iMenschen)
das gar nicht, daß er Vernunft hat,
dienen
diese
Kräfte zur Erringung des nackten Daseins?
Auch nach der methodologischen
^
unbarmherzig
als
wenn sie ihm nur zum Behuf desjenigen
Ja es müßte von hier
der Instinkt verrichtet."
Fortschritt
in
Wahrheit
als
ein
Rückschritt
gelten.
Denn ist es nicht ein Rückschritt, wenn für die Erreichung desselben Zieles
immer kompliziertere Mittel, immer mehr Mühe und Arbeit aufzubieten sind?
Mit der Anerkennung neuer Gehalte und Werte aber wird die mechanische
Evolutionslehre verlassen.
14*
Zum Weltproblem.
212
in
arge Verwicklungen,
sobald
Wir sehen dann
zieht.
Ästhetik
u.
s.
zurückstreben
Fehler,
und
Nun
suchen.
eine
w. entstehen,
evolutionistische
die
gewiß
beging
die
begonnen
Anfängen
Anschauungsweise
ältere
den
Daß
das Geistesleben nicht
vom
sondern von geringen, halbtierischen Anfängen
ist,
daran
hat,
Rechtslehre,
Ethik,
tierischen
Anfänge hineinzusehen und diese
die
in
damit fälschlich zu idealisieren.
Himmel gefallen
den
zu
alle
ihnen den Schlüssel aller weiteren Bildung
in
höhere Stufe
die
auch das Geistesleben an sich
sie
kaum mehr zu
heute
ist
Aber
zweifeln.
"müssen jene Anfänge maßgebend für die Gesamtbewegung bleiben,
könnte nicht der Lebensprozeß sich bei sich selbst erhöhen, könnten
neue Kräfte
nicht
die
Bindung an
ihm hervorbrechen?
in
die
Sind ferner die ersten Anfänge so einfach
leugnet
als sie dieselbe
und klar, daß von ihnen
Vermögen wir
fiele?
Licht
hellstes
sie
notwendig
heute einnehmen?
So führt jener
es
ist
auf
sonst
dunkle Gebiete
uns unmittelbar vor Augen zu
gestalten wir nicht
hinein;
Wahrheit bekräftigt
In
Anfänge die Entwicklung nicht sowohl,
ersten
wir
Weg uns erst recht ins Dunkel
Weg, sondern
nicht ein gerader
stellen,
dem Stande, den
nach
ihr Bild
ein
Umweg, wenn
wir von hypothetisch ausgedachten Anfängen aus höhere Stufen zu
erklären suchen.^
Das alles wendet sich gegen
und
die mechanische
Art der Entwicklungslehre, sofern sie nach ihren
Leben
gestaltet.
naturalistische
Maßen das ganze
Aber auch der Gesamtgedanke der Entwicklung,
wie er die Neuzeit durchdringt, enthält mehr Probleme,
wärtig zu sein pflegt.
*
Zunächst wird
viel
zu
leicht,
als
gegen-
wo irgend Be-
Neuerdings hat dies für das Gebiet der Ästhetik ebenso scharfsinnig
in der Abhandlung «Die entwickelungs-
wie überzeugend Volkelt dargelegt
geschichtliche Betrachtungsweise in der Ästhetik" (Zeitschrift für Psychologie
und Physiologie der Sinnesorgane, Bd. 29). Dort heißt es (Abdruck S. 7):
»Es gilt zu bedenken, daß sich für die Beantwortung der Frage, was es
heiße, sich zu den Dichtungen dichterisch, künstlerisch, ästhetisch verhalten,
nur vom Standpunkte des gereiften gegenwärtigen Menschen aus eine sichere
—
Grundlage gewinnen läßt.«
S. 8: »In Wahrheit ist
um diesen kurzen
Ausdruck zu gebrauchen
die Ästhetik der Naturvölker kein methodisches
—
Mittel,
sondern
vielmehr
eine
der
Sonderaufgaben der Gesamtästhetik."
lungsgeschichtlicher Grundlage"
Verhältnisses."
ist
allerdunkelsten
S.
11:
demnach
und unzugänglichsten
„Eine „Ästhetik auf entwickeine
Umkehrung des richtigen
Entwicklung.
wegung vorliegt, ein
213
Fortschreiten, eine Entwicklung
im Sinne eines
Daß unsere Welt, namentlich der Bereich unseres Handelns, voller Bewegung sei, das stand
unablässigen Aufsteigens angenommen.
auch den Alten deutlich vor Augen; ihnen
niedere Stufe,
weil
sie
Vordringen
sicheres
enthielt als ein
die Religion
als eine
Die
moderne Überzeugung dagegen
Hauptstück den Glauben an ein solches Aufsteigen;
ihn
hat
weiter gestützt
aber diese
nur ein wirres Durcheinander, kein
ihr
in
sahen.
galt
und
aufgebracht,
die
Philosophie ihn
spekulative
und Spekulation sind
Religion
durcTigebildet.
heute verblaßt und vielen gänzlich entschwunden, ihr Erzeugnis aber,
der Fortschrittsglaube, ist verblieben; ist er, nach Wegfall jener Grundlagen,
noch genügend fundiert, gibt ihn die bloße Erfahrung als
eine unumstößliche Tatsache, kann sie mit ihrer Begrenztheit über-
haupt einen unablässigen
Fortschritt
erweisen?
im
Es
viel
subjektive- Stimmung
alle
Veränderung als einen Fortschritt zu deuten.
Lauf des Lebens an
bringt,
liegt
und
Sicherlich
ist
selbst
Streben
als
alles
den
hier
dem Menschen nahe,
Er sieht, was der
was zu-
vergißt darüber,
an Altem verloren geht; von da aus fühlt jede Zeit
gleich
sich
Neuem
Spiel.
Höhepunkt des Ganzen, weil
Übrige mißt; eine künstlerische Zeit
sie
am
pflegt
leicht
eignen
nach der
Kunst, eine technische nach der technischen Leistung zu schätzen.
Zu diesen dauernden Ursachen kommen zeitliche: nichts ist günstiger
für den
Fortschrittsglauben
gefühl,
wie es aufsteigende Zeiten durchdringt, wie es namentlich
den Hauptzug der Neuzeit
ergriffen,
als
ein
starkes Kraft-
Von
erfüllt.
und Gegenwarts-
hier aus wird alles freudig
was eine Mehrung des Lebens verspricht, dahingehende
Erfahrungen einzelner Gebiete werden verallgemeinert. Vereinzeltes
und
Zerstreutes wird ergänzt
und verbunden. Hemmendes dagegen
übersehen oder zurückgestellt, selbst der Widerstand
in
durch den
umgewandelt.
inneren
Lebenstrieb
als
ein Antrieb
dem allen die bloße Erfahrung
zu weiterer Tätigkeit verstanden,
Einer
solchen
Be-
trachtung und Behandlung des menschlichen Daseins droht schließlich
notwendig ein Rückschlag, eine kühlere und
art
wird manches Stück jenes Fortschrittsglaubens zerstören,
retardierende
Momente vor Augen
stellen,
kritischere
wird manches
Denk-
als
wird
eine
vorübergehende Erscheinung erkennen lassen, was jener Glaube. zu
einem bleibenden Gesetz erhob. So ging z. B. durch die letzten
Jahrhunderte die Lehre von
Bevölkerungszahl,
einem
ihr Stillstand
bei
unablässigen
Anschwellen
einzelnen Nationen
der
wurde wie
Zum Weltproblem.
214
Wie jung aber
Noch Montesquieu meinte, die Bevölkerung Europas
habe gegen das Altertum abgenommen, und es empfehle sich, die
eine
ist
merkwürdige Ausnahme behandelt und erörtert.
diese Lehre!
Vermehrung des Menschengeschlechts durch besondere Gesetze zu
Dann siegte die entgegengesetzte Annahme, und die Gefahren einer übergroßen Vermehrung kamen bei Malthus zu starkem
fördern.
Die Zahlen
Ausdruck.
gaben
einstweilen
Annahme
dieser
recht,
immer mehr Anzeichen, daß auf einer
Höhe der Kultur die Zunahme langsamer wird und zum
Stillstande, ja Rückgange kommt; das macht die Frage unabweisbar,
aber neuerdings erscheinen
gewissen
ob
sie
nur für besondere Lagen
vielleicht
der Kultur,
nicht aber
Wie sehr aber muß die Verfolgung dieses Gedankens
dauernd gelte.
den Gesamtanblick der Geschichte verändern!
Ferner greift das Problem
über
hinaus
das Quantitative
ins
Wachstum der
geistigen
Minder
des
Vermögens?
sie
die
Summe
Menschheit, erhöht
zuversichtlich, als die durchschnittliche Meinung darüber denkt, stimmt
Qualitative.
Bringt
die
Geschichte
ein
geistiges
der von
Lorenz nachdrücklich verfochtene Antagonismus zwischen
geistiger
Leistung
und
Tätigkeit
eine
geringere
nach würde sich
Fortpflanzungsfähigkeit
—
es
und stärkere geistige
schließt«
sich
in
(Lehrbuch der Genealogie" S. 486/7), und meint,
lichkeit
nennt
Lorenz
Fortpflanzungsfähigkeit.
«eine sehr beachtenswerte Tatsache, daß höhere
»aller Wahrschein-
eine Erfahrung, die
man
anderweitig
beobachtet hat, auch genealogisch bestätigen lassen, daß der männliche
Keim
eine Wanderung von unten nach
oben
den oberen Ständen, oder wie man nach heutiger
Organisation sagen könnte, in
und Kulturvölker
Überwältigungen, sondern vielmehr
stellt
nicht
als
sich
als
und
in
gesellschaftlicher
den höheren Berufen
Weiterführung dieses Gedankenganges
höherer Kulturen
vollzieht
abstirbt."
der
eine
,;
In
Untergang
Folge
äußerer
die natürliche Abnahme der
Fortpflanzungspotenzen des höheren, kultivierten Individuums" dar;
es erscheint ein
«Unvermögen der Natur, das Geistige
— um diesen
gebrauchen
— schlecht-
Ausdruck nur im Sinne der
hin
fortzupflanzen"
(S.
487).
Kausalität zu
So würde die Bewegung
sich
bei
sich selbst erschöpfen, die Kulturen müßten sich ausleben und greisenhaft werden, eine Stagnation erfolgen, bis
vor allem aber frische Menschen
dann aus einem unablässigen Aufsteigen
verschiedener Phasen verwandeln.
wieder neue Anregungen,
kommen;
das Ganze würde sich
in ein
Auf- und
Abwogen
Was es dabei etwa an Fortschritt
Entwicklung.
215
gäbe, das würde sich jedenfalls anders ausnehmen als im gewöhnlichen Fortschrittsglauben.
eine
Auch das gehört hierher, daß die verschiedenen Lebensgebiete
verschiedene Art der Bewegung zeigen, und daß das Über-
wiegen eines dieser Gebiete die ihm innewohnende Schätzung zur
erheben
Allgemeingültigkeit zu
es
Einen unaufhödichen Fort-
pflegt.
am meisten Technik und exakte Wissenschaft, obschon
schritt zeigen
auch hier an Verlusten und Rückgängen keineswegs fehlt; geistiges
Schaffen im Sinne einer inneren
findet
eine
volle
Erhöhung des menschlichen Lebens
Verkörperung nur an einzelnen ausgezeichneten
Punkten, um dann rasch wieder abzunehmen;
in
scheint die Menschheit sowohl im Guten als
im Bösen,
wie
Gegenwirkung
in
moralischer Hinsicht
der Gegensatz
fortzuschreiten,
schroffer zu werden; die Religion endlich gibt
als
allem
zeitlichen
Wahrheit
Wandel überlegen,
sie
ihre
in
Wirkung
immer
also
Grundwahrheit
betrachtet
leicht
diese
an irgendwelchem früheren Zeitpunkt schon erreicht
als
und -kettet damit das Streben an
die Vergangenheit.
Jede dieser
Arten hat aber die Neigung, von sich aus ein allumfassendes Ge-
und Weltbild zu entwerfen.
schichtsschritts
voller Verwicklung,
gemeingültig
auftritt,
Lage vorübergehender
Endlich
ist
oft
So
ist
das Problem des Fort-
und was als selbstverständlich und allnur
Erzeugnis
das
besonderen
einer
Art.
muß die Entwicklung auch in der Richtung Bedenken
erwecken, daß sie leicht dazu führt, die Bewegung ausschließlich als
ein
Werk
ein
zu kontemplatives und
der Notwendigkeit zu verstehen
Der
bringen.
Fortschritt
passives
scheint
und
den
Menschen
in
Umgebung zu
mehr an dem Menschen als
Verhältnis
hier
zur
durch ihn zu erfolgen, er scheint nicht eignen Eintretens und eigner
Entscheidung zu bedürfen.
So geschah es z. B. in der Entwicklungs-
idee der Romantik, die ein
stilles
heraus
alle
und
Gestaltung hervorbringen
sicheres Wachsen
ließ
zu eigner Tätigkeit lähmte; so kann es auch geschehen,
wegende
Kraft in sinnliche Naturtriebe
von innen
und damit den Antrieb
wo die be-
und äußere Notwendigkeiten
gesetzt wird.
Hier wie da gefährdet die Entwicklung den ethischen
Charakter des
Lebens und
zerstört
sie
die
Grundbedingung
einer
echten Geschichte: ein immer neues Einsetzen ursprünglichen Lebens,
ein
Verwandeln
Gegenwart.
alles
Empfangenen
in
eigne
Tat
und lebendige
Während das menschliche Geistesleben seine Spannung
durch den Zusammenstoß von
und seinen Charakter vornehmlich
Zum Weltproblem.
216
und
Schicksal
Freiheit
opfert
erhält,
eine
dem Schicksal auf.
die Freiheit gänzlich
das solche Probleme übersehen
ein Durcheinander-
Entwicklung wird
läßt.
Bewegung genannt, ohne daß dabei der Ur-
oft alle fortschreitende
sache des Fortschritts nachgefragt wird;
ein
ist
und einer strengeren Fassung des Entwicklungs-
laufen einer laxeren
begriffes,
Entwicklungslehre
solche
Es
so könnte
einen Naturprozeß, der aus zwingender Not-
zeichnet Entwicklung
wendigkeit vorwärts treibt
—
sei
es
durch ein Zusammenschießen
—
einzelner Elemente, sei es durch eine Bewegung des Ganzen
dann
und zugleich auch
entfällt alle Freiheit
zeichnend
wohl
hier ganz
Im strengeren Sinne dagegen be-
Platz für Freiheit verbleiben.
menschlichen
Alsdann
Sinn.
alle
nur etwas
geht
,
als-
Geschichte im ausvor,
es
ist
ge-
wird aber nicht gehandelt; bei dieser Bedeutung des Wortes
Unding.
schichtliche Entwicklung ein
Ja der Zweifel greift noch
tiefer,
er kehrt sich überhaupt
gegen
die Vorherrschaft der Bewegung, gegen die Verwandlung der ganzen
Wirklichkeit in einen Prozeß.
dem Flüssigwerden
hältnisse,
An dem Beweglichmachen aller Veraller starren
Größen sah die Neuzeit
zunächst nur den Gewinn: die Steigerung des Lebens, das Wachs-
tum an
Freiheit
bleiben,
daß auch
und
Schließlich aber kann nicht verborgen
Kraft.
vieles
damit verloren geht, etwas verloren geht,
ohne das geistiges Leben schlechterdings
in
Grundformen
elementarsten
seine
nicht bestehen
hinein
verlangt
kann.
und
Bis
erweist
nämlich das Geistesleben eine beharrende Art, ein Beharren nicht
innerhalb der Zeit, sondern
heute oder morgen
ist
ein
gegenüber der
Zeit.
Ein Wahres für
Unding; was irgend wahr ist, das
für alle Zeit oder vielmehr ohne alle Beziehung zur Zeit;
gilt
mag die
Behauptung unter besonderen Umständen nur auf eine Zeitspanne
ist immer zeitloser Art,
Wahre eine Befreiung von aller
gehen, die Art, wie sie ausgesprochen wird,
als
geistiges Erlebnis
enthält
alles
Auch was wir als gut erachten und schätzen, das hat seinen
Wert nicht aus dem Gesichtspunkt einer besonderen Zeitlage, sondern
Zeit
unabhängig von
So gewiß
aller Zeit,
aus einer zeitlosen
die Begriffe der Zeiten vom
Guten
Ordnung der Dinge.
sich ändern:
Zeit als gut ergreift, das erklärt sie damit für schlechthin
gültig.
Aller Wandel menschlicher Verhältnisse
was eine
und dauernd
vermag solche innere
Überlegenheit des Geisteslebens über die Zeit nicht zu zerstören.
Auch
Begriffe
wie Persönlichkeit, Charakter, geistige
Individualität
Entwicklung.
bekunden
eine
Denn
sie
fordern die Gestaltung
treues
Festhalten
des
Überlegenheit
gegenüber
217
beharrenden Art
Bewegung;
aller
gegen
Geisteslebens
einer
die
Zeit.
und
ihr
Mannigfaltigkeit
alle
des Handelns hat jene Art zum Ausdruck zu bringen und zu fördern.
heißt,
es
von Grund aus zerstören.
Bewegung selbst
Ja die
Beharrens.
entbehrlichkeit
des
Ganzes zusammenfassen,
Sie
Sonst nämlich
mögen die Seele wohl
erfüllen
und
zerfällt sie
Je
sich
als
ein
nämlich
nicht
über-
Ganzes erleben, ohne
in
lauter einzelne Punkte; diese
mit kaleidoskopisch wechselnden Eindrücken
ergötzen, ein
ihr nicht.
läßt
und ohne eine von daher bewirkte
einen ihr überlegenen Standort
Synthese.
Un-
bezeugt, innerlich angesehen, die
blicken, in ein
sie
Bewegung verwandeln,
das Geistesleben ganz und gar in
So
Ganzes und zugleich einen
Inhalt
geben
Bewegung überlegene Kraft
mehr drängt das Leben zur Oberfläche und
mehr daher
verschwindet, desto
eine der
Beisichselbstsein.
verliert alles
Diese zeitüberlegene Art des Geisteslebens
erweist
besonders
deutlich der Aufbau einer Geschichte, d. h. einer eigentümlich menschlichen
und
Sinne
ist
Denn Geschichte im menschlichen
geistigen Geschichte.
bloßes Nacheinander von Ereignissen, ein
keineswegs ein
Dahintreiben des Menschen
dem Strom
mit
der Zeit; das würde
nie über die äußere
Anhäufung von Wirkungen hinausführen, wie
die Natur,
der Bildung der Erdrinde,
ist
alle
z.
B. in
sie
zeigt.
Vielmehr
Geschichte menschlicher Art eine Gegenwirkung gegen die
Flucht der Erscheinungen, ein Versuch, den Strom irgend zum Stehen
Auch die primitivsten
zu bringen, ein Kampf gegen die bloße Zeit.
Versuche, Vorgänge und Taten
überliefern,
sie
dem Gedächtnis
der Nachwelt zu
im Bewußtsein der Menschheil festzuhalten, zeigen
einen solchen Widerstand gegen die Zeit; je mehr aber die Geschichte
dem Menschen wird,
je
mehr
seines Wissens, sondern eine
desto
mehr
Selbsttätigkeit
ist
sie
ihm
nicht
nur eine Erweiterung
Erhöhung seines Lebens bringen soll,
dabei aufzubieten; das aber verlangt
notwendig einen zeitüberlegenen Standort.
Um die Vergangenheit
innerlich mitzuerleben, müssen wir uns von
der Zufälligkeit
der
Gegenwart befreien, wenigstens nach solcher Befreiung streben; sonst
würden wir in
alles
Frühere lediglich die heutige Art hineinsehen
und in aller äußeren Erweiterung innerlich bei uns selbst verbleiben;
ein Verständnis anderer Epochen aus ihren eignen Zusammenhängen
wäre
uns gänzlich
versagt.
Dazu möchten wir
die
Vergangenheit
Zum Weltproblem.
218
zum eignen Leben in Beziehung
setzen, ihren Reichtum in eignen Besitz verwandeln, an dem Großen
sondern
nicht bloß erkennen,
in ihr
sie
Dafür aber gilt es nicht nur eine Gemein-
uns selber heben.
Epochen zu gewinnen, sondern auch an ihrem
schaft mit früheren
Bestände Wesentliches und Zufälliges, Wertvolles und Gleichgültiges
zu scheiden;
sollte
das möglich sein ohne irgendwelche
dem Wandel
der Zeiten überlegenen Maßstäbe, ohne eine Versetzung der Arbeit
auf einen zeitlosen Standort?
für uns
sofern
Wert,
zusetzen vermögen; das
Letzthin
wir
als
in
sie
die Geschichte
hat
eine
zeitlose
ihr Hauptertrag,
ist
nur in-
Gegenwart um-
uns aus der engen und
armen Gegenwart des bloßen Augenblicks zu
einer weiteren,
überlegenen und zeitumspannenden Gegenwart zu führen.
zeit-
Es gibt
keinen gefährlicheren Gegner einer echten Gegenwart als die Hin-
gebung an den bloßen Augenblick.
Bei
solcher Lage der Dinge
verbietet
sich schlechterdings die
Auslieferung des ganzen Lebens an die Bewegung;
gewicht
in
extremsten
So haben auch
irgendwelchem Bleibenden gesucht.
die
Vorkämpfer der naturwissenschaftlichen Bewegungslehre
irgendwelche Ergänzung der Bewegung anerkannt.
der Lehre
mag das Bewußt-
von dieser erfüllt sein, die Arbeit hat stets ein Gegen-
sein lediglich
vom Beharren
Das sowohl
des Stoffes oder der Energie als
in
in
der
Unterordnung aller Erscheinungen unter unwandelbare Gesetze. Ohne
solche Befestigung hätte ihre Arbeit den Charakter der Wissenschaft
vedoren und wäre
statt
kausaler Begreifung eine
bunte Erzählung
geworden.
Auch die Philosophen haben die Entwicklung nicht zur Zentralidee ihrer Gedankenwelt rpachen können, ohne ein der Veränderung
überlegenes, ja
umspannendes Beharren anzuerkennen.
sie
Hegel wäre sein System
in
lauter einzelne
Punkte zerbrochen, und
der Wechsel der einzelnen Phasen hätte ihm
hätte
ihm
nicht
eine
zeitüberlegene
Einem
alle Wahrheit zerstört,
Betrachtung eine Zusammen-
fassung zu einem Ganzen geboten, alles Nacheinander in ein Selbst-
Ganzen verwandelt und zugleich über den zeitlichen
Ablauf hinaus in eine zeitlose Gegenwart gehoben. Ob das erstrebte
leben
Ziel
dieses
bei
Hegel vollauf erreicht
dem Streben
aber
ist
ist,
das
ist
nicht zu zweifeln, alle
eine
andere Frage, an
Größe des Hegeischen
Systems hängt eng mit ihm zusammen.
Auch
bei
Comte, dem großen
steht es ähnlich: zu
realistischen
Gegenstück Hegels,
einem wissenschaftlichen System gelangt er nur
Entwicklung.
219
Wohl
durch Ausbildung und Voranstellung beharrender Elemente.
bringt er alle bisherige Geschichte in Fluß und gewährt den früheren
nur
Stufen
eine
relative
Aber
Wahrheit.
Positivismus scheint die absolute
Wendung zum
der
in
und endgültige Wahrheit erreicht;
die Zukunft
mag diese weiter entfalten, der Kern scheint unwandel-
bar für
alle
Zeiten
schichte
rückwärts erfolgt
gesichert.
Auch
die Durchleuchtung
von diesem
gänzlich
als
der Geerachteten
fest
So wird inmitten aller Bewegung eine bleibende Wahr-
Höhepunkte.
heit festgehalten.
Für das gemeinsame Leben war freilich mit solcher versteckten
Anerkennung eines Beharrens wenig gewonnen, der fortschreitenden
Verwandlung des modernen Lebens in einen bloßen Prozeß wurde
von da aus kein
Gewicht
ins
fiel
genügender Widerstand
das tatsächliche
Weit
geleistet.
stärker
beharrender Größen
Fortwirken
und Mächte aus der älteren Lebensführung.
In
ihnen,
die
tief
in
den Bestand des Daseins eingebildet waren und den Menschen wie
umfingen,
seltjstverständlich
hat die
Bewegung stillschweigend bald
Aber eine solche Lage
einen Halt, bald eine Ergänzung gefunden.
mit
Unausgeglichenheit
ihrer
nicht auf die
entgegengesetzter
Strömungen
Dauer verbleiben, im Vordringen befindet
unverkennbar die Bewegung, so wird
sie
kann
aber
sich
mehr und mehr das Feld
einnehmen, ihre Konsequenzen hervortreiben,
Feste
alles
auflösen,
das ganze Leben in einen ruhelosen Prozeß verwandeln.
Zugleich aber werden auch die Folgen eintreten, die das Ver-
schwinden
beharrenden Größen
aller
vornehmlich der Wegfall
lebens
aus
aller
dem Ganzen,
selbständigen
Geistigkeit,
zugleich
aber
das Sinken
einen völligen Sieg sowohl
in
mit sich
Kräfte
bringt,
die
Verkümmerung
aller
inneren
Er-
der Arbeit
zur
Der Triumph der bloßen Bewegung bedeutet
höhung des Daseins.
eine Preisgebung
und
inneren Zusammenfassung, alles Er-
alles
des Relativismus als
Lebensinhalts,
eine
des Sensualismus,
Auflösung des Daseins
einzelne Augenblicke, einen Verzicht auf alle wahrhaftige Gegen-
wart,
Auch muß sich damit die Menschheit in lauter einzelne Lebens-
kreise
zersplittern
und eine gemeinsame Gedankenwelt
der und erhöhender Art mehr und mehr
befestigen-
verlieren.
Läßt sich leugnen, daß der Anblick der Gegenwart uns die zerstörende
stellt,
Kraft
dieser
Wendung schon
deutlich
genug vor Augen
und daß die daraus erwachsenden Fragen und Zweifel
die Grundlagen des
modernen Lebens zurückgreifen?
bis in
Ja wir haben
Zum Weltproblem.
220
ein bunteres
Autorität
und
ein bewegteres Leben
und keine
Tradition, wir
gewonnen, uns beengt keine
können mit voller Frische jedem
Tempo des Lebens
Eindruck folgen, den Augenblick ergreifen, das
Aber in
beschleunigen.
und Geschäftigkeit droht
aller Beweglichkeit
das Leben sich uns an die bloße Oberfläche zu verlegen und seiner
nach immer leerer zu werden;
seelischen Art
uns
entweicht
eine
innere Einheit des Wesens und damit der einzig mögliche Halt gegen-
dem Strom der Dinge; unfähig, unsere Selbständigkeit an ihnen
über
zu erweisen, werden wir wehrlos von ihnen hin- und hergeworfen.
Zugleich zerrinnt uns alle wahrhaftige Gegenwart, da
Lebens
des
bloße Zeit
sich
in
unvermeidlich
und
eine
sie ein
Ruhen
Erhebung über
die
Dafür bekommen wir bloße Augenblicke, deren
enthält. ^
bunter Wechsel
verlangt
selbst
das Leben in eine rastlose Flucht verwandelt und
dem Streben die Richtung auf das unmittelbar Wirk-
same, das Sinnfällige, das äußerlich Vorteilhafte gibt.
Als notwendige
Folge dessen jenes Haschen nach immer Neuem, Blendendem, Auf-
regendem, jenes Spähen nach Sensation, Effekt
dienerei gegen die
u. s. w.,
jene Liebe-
Launen und Stimmungen des Massenpublikums,
dieses geringen Durchschnitts der Menschheit, jene unwürdige «Aktua-
die den schönen Begriff des Aristoteles in sein volles Gegen-
lität«,
teil
verkehrt hat!^
Der Zeit unserer Klassiker war das mit voller Deutlichkeit gegenwärtig.
*
Es sei nur an jenes Wort Goethes (aus den Gesprächen mit Eckermann) erinnert: „Jeder Zustand, ja jeder Augenblick
er
ist
ist
von unendlichem Wert, denn
der Repräsentant der ganzen Ewigkeit"; auch des Wortes eines neueren
feinsinnigen Geistes (W. Gidionsen) sei gedacht:
»Nicht vom Tage sollst du leben,
Auf und nieder schwankt die Welle
Laß dein Inn'res fröhlich weben.
—
Stets verjüngten Daseins Quelle.
Ist
Ursprünglichkeit dir eigen,
Darfst sie hegen, darfst sie zeigen.
So nur spürst du in der Zeit
Vorgefühl der Ewigkeit."
^
Der Ausdruck
actualis
ist
eine
Schöpfung des späteren Altertums
(Augustin, Macrobius), im Mittelalter gewannen, von griechisch-lateinischen
Übersetzungen des Aristoteles
her, actus, actualis, actualitas
- namentlich seit
Duns Scotus — eine weite Verbreitung und gelangten von dort zur Neuzeit.
Das Wort diente zur Wiedergabe des aristotelischen Begriffes der Energie
oder Entelechie, der in sich selbst ruhenden und bei sich selbst befriedigten
Tätigkeit
im Gegensatz zu der noch unfertigen, erst anstrebenden Bewegung.
Entwicklung.
mehr uns aber damit
Je
die
221
Gegenwart
entschwindet, desto lebhafter wird ein Sehnen
unter
den Händen
eine
unbestimmte
in
Zukunft, ein Erhaschen und Vorausnehmen dessen, was dort erwartet
»Nie
wird.
als
es die
wie
jetzt
beeifert
ist",
so sagt Lotze
Gegenwart
in
einer
noch weit ruhigeren Zeit
(Mikrokosmus 2. Aufl. II, 281), «so lebhaft
der Widerspruch aufgetreten, das ganze Leben, das man
ist
und emsig mitlebt, doch im Grunde
das wahre
für
nicht
zu halten und von einem anderen schöneren zu träumen, das
man
leben
möchte und leben wird, sobald uns jenes Zeit lassen
und
einen
Zugang zu ihm öffnen wird."
So zerfällt uns in einer Überspannung und Überstürzung der
Bewegung das Leben von innen her, aus einem wahrhaftigen Leben
wird es mehr und mehr ein bloßes Lebenwollen, eine Anweisung
auf Leben, ja ein Schein des Lebens. Das kann unmöglich so weiter
gehen, jener Verwandlung des Daseins in bloße Bewegung muß als
einer
Zerstörung
völligen
widerstanden
so gewiß das Verlangen
keit ihrer innersten
Die Menschheit
werden.
muß jene gefährliche Krise überwinden und
wird
sie
übei'winden,
danach aus einer zwingenden Notwendig-
Natur hervorgeht.
Aber sie wird
winden ohne eingreifende Wandlung des Daseins,
sie nicht über-
ohne
nicht
die
Ausbildung eines neuen Lebenstypus, nicht ohne den Mut und die
Kraft zu einem neuen geistigen Aufschwung.
d)
Forderungen für einen neuen Lebenstypus.
So wenig das Problem,
hier näher erörtern läßt,^
Richtung
des
in
das unsere Untersuchung auslief, sich
ohne irgendwelche Orientierung über die
Weges würde unsere Betrachtung
einzuschlagenden
aller Kürze
ins Leere zu verlaufen scheinen; so seien
in
einige Umrisse entworfen. — Vor allem
gilt
wenigstens
jener drohenden
es,
Verflüchtigung des Lebens einen festen Halt^ entgegenzusetzen.
solchen kann
nicht
Außenwelt
die
mittels unserer Seele erleben
uns
beweglich werden
Bewegung
gehörte.
^
und
in
da wir
und daher auch das
müßte,
Denn
hier
Einen
immer
sie
Festeste
wenn das Seelenleben
Eine Festigkeit aber gewährt
unmittelbare Seelenleben.
einander,
bieten,
gänzlich
auch
erst
draußen
nicht
der
das
wogt Mannigfachstes durch-
buntem Wirbel verdrängt
die
eine
Erscheinung
Es sei dafür auf meine «Grundlinien einer neuen Lebensanschauung"
(1907) verwiesen.
Zum Weltproblem.
222
die
zu
Es bleibt also nur die Hoffnung,
andere.
geistigen Tätigkeit vorzudringen,
auch das übrige Leben
die,
sich
fest in
irgendwelcher
gegründet,
selbst
Das haben große
zu befestigen verspräche.
Denker der Neuzeit in verschiedener Weise versucht: den archimedischen Punkt suchte Descartes im reinen Denken, Kant im sittlichen
Handeln; beider Unternehmen aber wurzelt
Bewegungen
in weiteren
des modernen Lebens, indem einerseits die wissenschaftliche Arbeit,
andererseits ein ethisches Schaffen dem rneijschlichen Dasein einen festen
Grund zu geben und seiner Verflüchtigung in bloße Erscheinungen
entgegenzuwirken bemüht war.
geleistet
Beide
Bewegungen haben Großes
und fahren fort das zu tun; trotzdem wächst der Zweifel,
ob sie den tiefsten Punkt erreichen und von hier aus das ganze Leben
zu umfassen vermögen.
Einmal nämlich treiben
eine besondere Richtung
und geben ihm
intellektual istische,
dort eine
Färbung.
Für unser
moralistische,
eine
hier
das Leben in
sie
eine besondere,
Problem aber fällt noch mehr ins Gewicht, daß die Festlegung eines
besonderen Punktes immer wieder von anderen Punkten
zweifelt
und
bestritten
werden kann;
gegen dieses der
das Handeln,
gegen den
Intellekt
wenden,
Intellekt
her be-
kann sich
der Skeptizismus
kann die Wissenschaft zu einem bloßen Vorstellungsgewebe herabzudrücken,
Moral
der
Naturalismus
die
Naturtriebe
zu
verwandeln
suchen.
Die
Gewißheit
kann
nicht
besonderes
Gebiet,
ein
Zusammenfassung zu einem Ganzen
nicht eine der
Erzeugnis
ein
in
uns
mögliche
sondern
nur eine
höchste
bieten;
bloßer
liegt
im Geistesleben
Verzweigung überlegene Einheit, und bricht nicht in
dieser Einheit ein ursprüngliches Leben hervor, so kann unser Leben
und Streben
nie eine Festigkeit erlangen.
Daß aber der Gedanke einer allumfassenden Einheit mehr als
eine
bloße
Einbildung
ist,
das bezeugt die
Bewegung zu einem
Denn mag unser
Persönlichsein, wie sie die Menschheit durchdringt.
menschliches Persönlichsein noch so
viel
Bloßmenschliches an sich
tragen
und mannigfachsten Bedingungen und Einschränkungen unter-
liegen,
eine neue Art des Lebens, eine größere Tiefe der Wirklich-
beginnt
damit
das Geistesleben
erscheint
hier nicht als eine besondere Betätigung, sondern als eine
neue Art
keit
Wirklichkeit,
sich
als
eine
mit
Handeln,
sittliche
Demnach entsteht
neue Stufe des Seins,
ihnen
Betätigungen,
das
aufzuarbeiten;
sowohl
sich
der die
besonderen
das wissenschaftliche Denken
als
unterzuordnen und einzufügen haben.
eine Befestigung nur durch
ein Vordringen
des
Entwicklung.
223
damit wird
gesamten Lebens zu einer wesenbildenden Geistigkeit;
auch
der
Kultur
Theorie und Praxis überlegen
dem Gegensatz von
das
vorgehalten,
ideal
ein
und das jede von ihnen
ist,
eine
in
wesenhafte und wesenlose Stufe zerlegen muß.
So
rung des Daseins
energische Aufrüttelung,
eine
gestattet lediglich
Umkeh-
ja
Vordringen zu einem festen Punkte und ein
ein
Aufnehmen des Kampfes mit der Flucht der Zeit und der SinnlosigOhne ein Gegründetsein des Menschen
keit der bloßen Bewegung.
in
einer dem nächsten Dasein überlegenen
unmittelbar
wäre die Sache völlig aus-
gegenwärtigen Geisteswelt
und
sichtslos
selbst das Streben
und doch im Lebensprozeß
danach nicht zu begreifen.
Jene Zurückverlegung aber enthält die weitere Forderung, daß
das Geistesleben
nicht
der Mensch
sondern
eine
als
als
einem ihm überlegenen Geistesleben
an
teilhabend gelte, daß das Geistesleben
dem Menschen
ständig gegenüber
geistiges
als
Leben
und
in
seiner Substanz
in
anerkannt werde.
als
selb-
Wenn
damit
Dasein weiter auseinandertreten
menschliches
der durchgehenden
Menschen,
Eigenschaft des bloßen
Fassung,
wird
so
zugleich
Ver-
eine
und Bewegung und die .Ausbildung
dem Gegensatz überlegenen Lebenstypus ermöglicht. Der Sub-
ständigung zwischen Beharren
eines
stanz des Geistesleben
die
ist
Veränderung und mit
eine Ent-
ihr
—
Der Begriff der Wahrheit
dem Gegensatz des Theoretischen und des
duldet kein Werden und keine Veränderung,
wicklung schlechterdings fernzuhalten.
auch dieser Begriff
ist
—
Praktischen überlegen
die Zugehörigkeit zu einer zeitlosen Ordnung
der Zeit und
ist
für ihn unerläßlich.
einen Lebensinhalt
nur innerhalb
durch allmähliche Erfahrung erringen;
dazu aber be-
Der Mensch hingegen kann
Auch was c;r an Wahr-
und der Beweglichkeit.
darf er der Freiheit
ihm
nicht
heit
erreicht,
sich
des Besitzes ruhig
ist
sich
ein
für allemal
erfreuen
könnte,
gewonnen,
sondern
es
so
daß er
will
immer
neu gewonnen werden, es wird immer wieder zum Vorwurf des
Kampfes. Auch in die Grundlagen unser geistigen Existenz
Ungewißheit
neuem
immer von neuem zurück und
So
entstehen,
immer von
Überwindung.
eine kräftige
Lebens: die eine
verlangt
greift die
deutlich
ist
geschieden,
drei Arten
und Typen des
ausschließlich auf ein Beharren, ja einen ewigen
Bestand gerichtet und sucht das
menschliche Sein möglichst
aller
Bewegung zu entwinden; die andere ist gänzlich von der Bewegung
erfüllt
und
will
ihr
nichts
entzogen wissen;
die
dritte
strebt
über
Zum Weltproblem.
224
den Gegensatz hinaus und möchte aus innerer Überlegenheit jeder
zweite
die
Die
gewähren.
Recht
ihr
Seite
erste
moderne Gestaltung des
beherrscht
Lebens,
die
die
antike,
die
dritte
wirkt
von
Alters her innerhalb der geistigen Arbeit, aber prinzipiell
anzuerkennen, sowie
führen.
Hier
als
ist
sie erst
Lebenstypus zu voller Kraft und Klarheit zu
liegt die Aufgabe
Die
der Zukunft.
alte Art
war stark
dem Geistesleben Festigkeit und Ruhe zu geben, es als eine
unantastbare Ordnung über alles Mögen und Meinen der Einzelnen
darin,
wie der Massen hinauszuheben.
sie
veränderlich,
sondern auch
behandelte,
die
Ins Problematische aber
Wahrheit nicht nur als
dadurch, daß
den Menschen
als für
geriet sie
ihrer Substanz un-
in
fertig
vorhanden
daß sie Substanz und menschliche Existenzform
in
Eins
So gilt der antiken Welt und mehr noch dem
zusammenschob.
Mittelalter die wissenschaftliche Wahrheit als endgültig abgeschlossen,
so kennt auch das kirchliche Christentum keine Weiterbewegung der
Damit wird aber der Besitzstand einer be-
religiösen Gedankenwelt
sonderen Zeit für immer festgelegt,
alles
Weiterstreben gehemmt, der
Menschheit ein starres Joch auferlegt, das der Lauf der Zeiten immer
drückender machen muß.
Auch
Wahrheit
die
selbst leidet Schaden,
Wesen gesetzt
Dagegen mußte ein Rückschlag kommen, die Bewegung erstritt
indem Zufälliges der Zeiten und Menschen
wird.
ihr
ins
sich die Anerkennung ihres Rechts, der Mensch begann seine Schran-
ken und die Bedingtheit seiner Leistungen zu empfinden, es begann
modernen Lebens, dessen Größe, aber auch dessen
jene Entwicklung
Selbstverzehrung uns beschäftigt hat.
Hatte die Beharrungslehre die
menschliche Existenzform unmittelbar mit der Substanz des Geisteslebens
zusammenrinnen
umgekehrt
Art;
jenes
das
lassen,
Geistesleben
ergibt
eine
so
den
Erstarrung,
unterwirft
die
Bedingungen
dieses
Bewegungslehre
der
menschlichen
eine Verflüchtigung
des
Geisteslebens.
An Versuchen zu Kompromissen hat es nicht gefehlt, das Ganze
des Lebens half und
hilft
sich
vornehmlich dadurch, daß das Neue,
was der Lauf der Zeiten bringt, möglichst in das Alte hineingedeutet,
in
den geschichtlichen Bildungen Kern und Schale unterschieden,
jener nach Kräften festgehalten,
ist
diese abgestreift
wurde.
Aber das
nur eine Ausflucht, und zwar eine Ausflucht, der die historische
ihrer Hervorkehrung der Eigentümlichund Unvergleichlichkeit der einzelnen Zeiten immer mehr den
Denkweise der Neuzeit mit
keit
Boden
entzieht.
Wollen wir also nicht zwischen den Gegensätzen
Entwicklung.
225
und uns von ihnen zerreiben lassen, so ist von innen
und unter wesentlicher Umwandlung des Wirklichkeitsbildes
Das aber wird erst möglich bei Anüber sie hinauszustreben.
des Geisteslebens und einer schärferen
Selbständigkeit
erkennung einer
stehen bleiben
her
Abhebung des menschlichen Daseins von ihm. Denn nur so lassen sich
Der Mensch muß
Beharren und Bewegung miteinander festhalten.
im tiefsten Grunde seines Wesens in einer unwandelbaren Geisteswelt gegründet sein, und es müssen von da aus bewegende und
Aber zugleich
richtende Wirkungen ausgehen.
Dasein höchst unsicher und unfertig, langsam
wegung in
weiter
Fluß,
zum
und nur
sein unmittelbares
kommt eine Be-
erst
der Zeit läßt sich weiter und
inmitten
Aber die Bewegung
vordringen.
Ziele
ist
verliert
sich,
dank jener Grundlage, nicht ins Vage und Fremde, es vollzieht sich
in
ihr
ein Erringen
ist
sie
keine
inmitten
aller
Wandlung
Vom Menschen
aus
angesehen
eignen Wesens,
des
bloße Veränderung.
verlangt eine solche Überzeugung eine Zurückverlegung des Lebens
Denn diese"
hinter die Fläche der einzelnen seelischen Betätigungen.
zeigen uns die Sache mitten im Fluß, namentlich
Gedankenwelt als
in
unablässiger
Wandlung
erscheint hier die
Aber
begriffen.
aller
solchen Veränderung kann eine charakteristische Art des Grundlebens
überlegen
bleiben
und
hindurch
behaupten,
entfalten.
So
steht
der Mensch zu-
über der Zeit;
sein
Leben
durch
sich
zeitüberlegene Wahrheit darin
gleich
Art,
in
der Zeit und
indem
es sich einmal einer zeitüberlegenen
Tatsache zu versichern
innerhalb
sie
der
Zeit
und
eine
in
immer
ihr
zu
Wahrheit als einer
zugleich
aber
Herausarbeitung
und
deutlichere Entfaltung jener Wahrheit zu erstreben
hier die
zweiseitiger
ist
begründen,
kräftigere
ihre
Daher
hat.
ist
Wahrheit zugleich Besitz und Problem, jenes im innersten
Grunde des Wesens,
dieses
bei
der Verwandlung des
Daseins in
volle Selbsttätigkeit.
Von hier aus wird ein Verhältnis zur Geschichte möglich, das
den Gegensatz von Beharren und Bewegung in sich aufnimmt und
Betrachten wir
zugleich überwindet.
z.
B. unsere Stellung zu
geschichtlichen Religion, etwa der christlichen.
die
menschlich-geschichtliche
Zeiten behaupten.
Form,
die
sie
Unmöglich
erhalten
hat,
einer
läßt sich
für
alle
Bei der gewaltigen Veränderung unseres äußeren
und inneren Daseins würde
unser Gefühls- und
Überzeugungsleben
wahrhaftigkeit geraten,
Eucken, Qnindbegriffe.
nicht nur unser Denken, sondern auch
wenn
4. Aufl.
sie
in
mit aller
die
Gefahr einer Un-
Gewalt auf jene* ältere
15
Zum Weltproblem.
226
Art gestimmt werden
leicht
sollte;
möchten wir unserer eignen Zeit
Unrecht tun, wenn wir nur darauf bedacht wären, anderer Zeiten
Recht
wahren.
zu
— Aber die Entfernung von der unmittelbaren
Lebensform braucht keine Preisgebung der Substanz zu bedeuten.
Es kann
unzulänglich gewordenen Existenzformen eine wahrhaf-
in
tige Art des Geisteslebens durchgebrochen sein,
die gesamte Geschichte erfüllende Tatsachen
leben fortfährt,
eine Art,
und nimmer losreißen
welche zeitüberlegene,
belebt
und zu be-
hat
von der sich das menschliche Leben nun
Dies Ewige aber würde zugleich seiner
darf.
menschlichen Gestaltung nach eine fortwährende Aufgabe
es
würde
durch
die
alle
Zeitüberlegenheit nicht durch
seine
ein
starres
bleiben,
Beharren
sondern vielmehr dadurch erweisen, daß es
Zeiten,
eingehen
Eigentümlichkeit aller Zeiten
kann,
ohne
sich
in
selbst
daß es jede Zeit auf das ihr innewohnende Ewige
zu verlieren,
zu bringen und damit von der bloßen
zu
befreien
vermag.
Die Zeit aber würde gegen die antike Fassung dadurch
gewaltig
Zeit
gehoben, daß innerhalb ihrer ein Fortschritt im Ewigen möglich wird.
Wie sich
auch
und
der Weltanblick
die Stellung
des
Menschen zur Wirklichkeit verwandelt, wenn das Werden an
die
zweite Stelle
weiter
tritt,
das läßt sich
ohne
hier
in
die antike Geringachtung zurückzusinken,
nicht weiter
Nur
verfolgen.
zum Schluß noch erwähnt sein.
ein
Punkt möge
Jene Grundüberzeugung mit ihrem
Ausgleich von Beharren und Bewegung kann nie den Tatsachen der
Entwicklung widersprechen, wohl aber
muß sie mit einer alleinselig-
machenden Entwicklungsphilosophie, einer naturalistischen Evolutionslehre, hart
Die
zusammenstoßen.
der Gesamtauffassung des
eignen Wesens.
letzte
geistigen
Entscheidung
Lebens
liegt hier bei
und zugleich unseres
Wie die Entwicklung im Ganzen der Wirklichkeit
zu verstehen sei, das hängt am meisten davon ab, ob im Geistesleben
eine neue Stufe des Lebens anerkannt, oder eine bloße Fortführung
der Natur gesehen wird.
Ist
jenes
der
Fall,
so gewinnt die Ent-
wicklung das Ansehen, daß nicht der erfahrungsmäßig vorliegende
Prozeß allen Fortgang aus sich
selbst hervortreibt, nicht das
ein bloßes Erzeugnis des Niederen bildet,
sondern daß
Höhere
in
die Be-
wegung neue Kräfte aus weiteren Zusammenhängen eintreten.
Damit
erhält
unsere Wirklichkeit einen
Hintergrund
und eine Tiefe,
sie
Bewegung aber ist
dann nicht mehr ein Weiter- und Weiterhasten ohne Ziel und ohne
Sinn, sondern sie wird getragen und umfaßt von einem Reiche
hat einem größeren Ganzen sich einzufügen; die
Entwicklung.
ewiger Wahrheit.
Ist
227
dagegen das Geistesieben ein bloßes Neben-
ergebnis der Natur, so
entfällt
alle
der
Möglichkeit,
Bewegung
ein
Gegengewicht zu geben und dem Leben einen Gehalt zu erringen,
dann
treibt
die Menschheit
Leere hinein.
So
ist
wie
die
ganze Welt
es auch hier, wie an allen
Untersuchung, die Stellung
zum
Geistesleben,
unaufhaltsam
ins
Hauptpunkten der
namentlich die An-
erkennung oder Verwerfung einer Selbständigkeit des Geisteslebens,
welche über die Richtung der Gedankenarbeit entscheidet.
15-
D. Zu den Problemen des Menschenlebens.
1.
Kultur.
I m eignen Gebiet des Menschen bildet den beherrschenden Mittel* punkt der Probleme die Kulturidee. Sie treibt aus sich eine
reiche
Verzweigung hervor, deren Gestaltung auf den Haupt- und
Gesamtbegriff zu näherer Bestimmung zurückwirkt.
Kultur führt
Das Wie der
den Problemen von Geschichte und Gesellschaft,
zu
Was zu denen der Moral, Kunst u. s. w. Als eine Einleitung
zu dem allen sei zunächst der Kulturbegriff im bloßen Umriß
das
erörtert
Zur Geschichte des Ausdrucks und Begriffs.
a)
vom Ausdem heute üblichen Sinne ist neueren Ursprungs.
Unserer Gewohnheit gemäß beginnen wir auch hier
Kultur in
druck.
Denn so nahe die Übertragung des Bildes von der Bestellung (colere)
dem späteren Altertum wie der
des Ackers auf den Stand der Seele
Renaissance
einen geschlossenen und abgegrenzten Begriff bildet
lag,
Die Kultur oder Georgik des Geistes wird ihm
daraus erst Bacon.
Aber dieser Versuch
ein Hauptteil der Ethik. ^
Folge,
er
ward unmittelbar
nicht
Eine ausgedehntere Bewegung hat wohl
des
17.
Jahrhunderts
ließ sie sich
^
S.
selbst
de augm.
hervorgerufen.
viel
zu deutlich
scient. VII, cp.
1:
hatte zunächst keine
aufgenommen und
weitergeführt.
erst die französische
Ihr
stolzes
Kultur
Selbstbewußtsein
von niederen Stufen abheben,
Partiemur igitur ethicam in doctrinas
principales duas, alteram de exemplari sive imagine boni, alteram de regimine
cultura animi,
et
quam etiam partem georgica anirai appellare consuevimus.
haec regulas de animo ad illas conformando
Der Ausdruck Georgik zeigt, wie stark das Bildliche des Ausdrucks empfunden wird.
lila
naturam boni
praesCTibit,
s.
describit,
auch cp. 3.
Kultur.
um nicht allgemeinere Reflexionen
229
über verschiedene Zustände der
Menschheit anzuregen; das 18. Jahrhundert mit seinem Streben nach
einer natürlichen Begreifung der Geschichte verfolgt solche Richtung
mehr und mehr mit dem Gegensatz
Aber so wenig es an Ausdrücken
eines Natur- und Kulturstandes.
für das Weiterkommen der Menschheit fehlt, verschiedene Bilder
und Vorstellungen laufen hier neben- und durcheinander: Kultivieren,
und
weiter
einen
Polieren, Polizieren, Aufklären; ^
Zivilisieren,
für
das Ganze
mit
»Zivilisation"
der
Latein
sich
beschäftigt
des dadurch
geschaffen
Standes dürfte
haben. ^
In
Ausdruck
den
Renaissance
Ausdruck
festen
erreichten
Deutschland
civilisatio, ^
Turgot
erst
das
besaß
auch
civilitas
wird
in ähnlicher Bedeutung
verwandt,* aber die lebendige Sprache
blieb
davon unberührt und
hatte bis in die Anfänge der klassischen
Die
Literaturepoche hinein nur schwankende Bezeichnungen.^
Aus der
*
(s.
ent-
Bayle
schier endlosen Fülle sei hier nur einiges angeführt.
oeuv. div. Haag 1727,
453a) hat cultiver leur esprit et leur raison; wenn
I,
ebenda 407a von toutes les societes, ou l'on cultivait I'esprit redet, so
würden wir das kaum anders als mit „Kulturvölker" übersetzen. Aber zu-
er
gleich
hat er civiliser
lisees
im Gegensatz zu
(z.
B. dictionn. 1465 se civiliser, 1472 b
barbares).
Leibniz (398a Erdm.)
nations
les
eclaire;
wo wir „Naturmensch" und „Kulturmensch"
plus
„Wilder"
und
peuples
grossiers
eclairees,
„Europäer";
gegenüber,
auch
öfter
nations civi-
Bossuet hat in ähnlicher Bedeutung
Montesquieu
le
stellt
siecle
les
qui passe pour
sagen würden, sagt er
peuples
eclaires
aber hat er poli oder police
(z.
den
B. les
les polis, la Grece seul polie au milieu des barbares, un pays police,
un royaume aussi police comme la France, les peuples polices, peuples bien
Polices). Auch in England fehlt ein fester Ausdruck; so gebraucht A. Smith
bunt durcheinander civilized und polished nations (s. z. B. the theory of moral
peuples
sentiments V, cp.
*
2).
S. Barth, die Philosophie der
Geschichte als Soziologie,
S. 253.
Nach Paulsen (Gesch. des gelehrten Unterrichts in Deutschland,
S. 78 u. 131) wurde zu Beginn des 16. Jahrhunderts von Wittenberg gesagt,
daß es in termino civilisationis liege.
'
Es bildet z. B. bei Kepler (II, 730) den Gegensatz von barbaries.
In seiner
zeigt z. B. der gediegene und gedankenreiche Iselin.
„Geschichte der Menschheit" pflegt er dem „Stande der Natur" den „Stand
*
^
Das
der Sitten" entgegenzusetzen und spricht demgemäß von „gesitteten" Völkern.
Aber nicht minder oft hat er „Polizierung" und
„poliziert", er unterscheidet
aber dabei, die spätere Sonderung von Kultur und Zivilisation vorausnehmend,
zwei
Arten
der Polizierung:
„die eine,
durch welche der Gesellschaft die
äußerliche Gestalt gegeben vcird", „die andere verbessert die Geister und die
Gemüter"
keit
(7.
Buch, 21. Hauptstück). Auch
stellt
er Barbarei und Menschlich-
einander entgegen und verwendet „Milderung" (auch
„Milderung der
Zu den Problemen des Menschenlebens.
230
scheidende Wendung brachte für Deutschland die klassische LiteraturIhr Verlangen nach einer
epoche.
und
einer
künstlerischen
Gestaltung des Daseins
zu selbständiges Kulturideal,
dem
hätten
ein
enthielt
viel
daß nicht auch die Ausdrücke sich
als
So
müssen.
anpassen
Belebung des ganzen Menschen
denn auch geschehen.
es
ist
Kultur wird nun zu einem festen und
zum herrschenden Begriffe,
Zivilisation grenzt sich davon als eine niedere Stufe ab,
»Aufklärung"
kaum durchgedrungen, die allgemeine Bedeutung und sinkt
verliert,
zur Bezeichnung der besonderen Art des 18. Jahrhunderts, zu einer
historischen Kategorie; dafür hebt sich »Bildung", die bisherige Be-
deutung
ins
Innere
und gewinnt den Affekt der
wendend,
diese Verschiebung der Ausdrücke etwas näher
Es
sei
sie
den deutschen Sprachgebrauch
ohne
»Kultur"
wohl erscheint
aber er befestigt sich schon
abzugeben.^
dargelegt, da
Gegenwart beherrscht.
begegnet uns zuerst bei Herder;
allen Zusatz
hier der neue
bis zur
Zeit.
Gebrauch noch
genug,
Neben Kultur steht,
als in
um einen
Fluß begriffen,
bündigen Terminus
so auch bei Goethe,
lange noch
Geisteskultur, aber allmählich gewinnt Kultur schlechtweg die Ober-
hand.
Die weitere Verwendung des Begriffes nimmt eine zwiefache
gemäß den beiden Hauptströmungen im deutschen Idealismus: der künstlerischen und der ethischen. Bei den Dichtern und
Humanisten überwiegt die ersk, Kunst und Wissenschaft in ihrer
Richtung,
Verbindung
zum
Schaffen
literarischen
sicheren Träger der Kultur,
erscheinen
hier
als
die
das unterscheidende Merkmal eines
als
Kant und mehr noch Fichte dagegen machen zur
Kulturstandes. 2
und „Erleuchtung" (auch „Erleuchtung der Geister") als unserem
Goethe hat in seinen Jugendschriften „polierter"
Mensch und „polierte" Nationen, und Kant spricht von „geschliffenen"
Sitten")
«Kultur« gleichbedeutend.
—
Volks klassen.
^
Namentlich wichtig ist für den Ausdruck die Stelle Ideen zur Philos.
der Gesch. IX,
1
:
„Wollen wir diese zweite Genesis des Menschen, die sein
ganzes Leben durchgeht, von der Bearbeitung des Ackers Kultur oder vom
Bilde des Lichts Aufklärung nennen
:
so stehet uns der Name frei
;
die Kette
der Kultur und Aufklärung reicht aber sodann bis ans Ende der Erde." Die
Kultur hat als beherrschendes Ziel die „Humanität", die für Herder die volle
und Harmonie aller Kräfte bedeutet, gemäß einer Überzeugung,
enge Verbindung von Leben und Schönheit als Ideal verehrt.
Das Unterscheidende des Menschen gegenüber der bloßen Natur aber ist
Entfaltung
welche
die
die Freiheit; so gehört diese wesentlich
siehe bei
^
zum Kulturbegriffe. Näheres darüber
Genthe „Der Kulturbegriff bei Herder".
S. die gleich
anzuführende Stelle aus F. A. Wolf.
Kultur.
und geben
die Freiheit
der Kultur
Seele
einen moralischen Charakter.
«Die Hervorbringung der
231
Also kann nur die Kultur der
Kultur.
vornehmlich
vernünftigen Wesens
Tauglichkeit eines
zu beliebigen Zwecken überhaupt (folglich
der Natur in Ansehung
damit
ihr
Kant definiert Kultur folgendermaßen:
in seiner Freiheit)
letzte
der Menschengattung
Ursache
beizulegen
wohl gar bloß
hat (nicht seine eigne Glückseligkeit auf Erden, oder
Ordnung und
das vornehmste Werkzeug zu sein,
Einhelligkeit
der vernunftlosen Natur außer ihm zu stiften") (V, 464, Hart.).
hat dies weiter ausgebaut
Ihm wird
die Freiheit,
der Kultur.
die
ist
Zweck sein, den man
in
Fichte
und gemäß seiner Art kräftig durchgesetzt
zum Inhalt
«Übung aller
die volle Selbsttätigkeit, zugleich
So bedeutet ihm diese (Wke.
VI,
86):
Kräfte auf den Zweck der völligen Freiheit, der völligen Unabhängigkeit
ihm
von allem, was nicht wir selbst, unser reines Selbst
diese
Aufgabe alles Übrige
in sich
der Sinnenwelt, nichts von unserem Treiben,
Erscheinung betrachtet, einen Wert,
Religion,
Wissenschaft
ist".
Wie
hat «nichts in
so
schließt,
Tun oder Leiden, als
als insofern es
auf Kultur wirkt"
und Tugend werden ausdrücklich
zu
den
höheren Zweigen der Vernunftkultur gerechnet (VII, 166); auch den
dem Denker vor-
Staatszweck bildet die Kultur, und der Staat, der
schwebt, wird als Kulturstaat bezeichnet^
Die beiden Nuancen der Kulturbewegung stimmen aber darin
zusammen, Kultur,
als ein
Bilden von innen her und eine Erhöhung
des ganzen Menschen, von
aller
bloßen Ordnung der Gesellschaft
deutlich abzuheben; zur Bezeichnung dieser
so
unterscheiden
sich
Zivilisation
und
dient
Kultur
nun
wie
Zivilisation;
Niederes
und
Höheres, wie Beginn und Vollendung.^
^
Der Begriff des Kulturstaates widerspricht zunächst der Fassung des
Staates als eines bloß „juridischen
Institutes".
Auch zum
nationalen Staat
Welches ist
denn das Vaterland des wahrhaft ausgebildeten christlichen Europäers? Im
allgemeinen ist es Europa, insbesondere ist es in jedem Zeitalter derjenige
bildete der Kulturstaat anfänglich einen Gegensatz;
Staat in
Europa, der auf der
s.
VII, 212:
Höhe der Kultur steht."
,,
Später hat gerade
und Vaterland zu Ehren gebracht, aber nie war es
das sinnliche Dasein, sondern immer der geistige Gehalt, der sie ihm beFichte die Begriffe Volk
deutend machte.
" Das erscheint schon deutlich genug bei Kant, s. namentlich IV, 152:
„Wir sind in hohem Grade durch Kunst und Wissenschaft kultiviert. Wir
sind zivilisiert bis
und Anständigkeit
zum
Überlästigen,
zu
Aber uns für schon
allerlei
gesellschaftlicher Artigkeit
moralisiert zu halten,
daran
fehlt
Zu den Problemen des Menschenlebens.
232
In
engem Zusammenhange mit
jener Steigerung
des
Kultur-
Aufkommen von «Bildung"; erst in der zweiten
1 8. Jahrhunderts wird es vom Äußeren aufs Innere, vom
begriffes steht das
Hälfte des
Mit besonderer Lebhaftig-
Körperlichen aufs Seelische übertragen.^
keit
bemächtigten sich seiner die Romantiker,
den Ausdruck «die Gebildeten"
fester
namentlich dürften
Bei
wie das Wort aus anfänglicher
Fichte läßt sich deutlich verfolgen,
Unsicherheit ein
sie
Umlauf gebracht haben. ^
in
Terminus wird.
„Bildung"
wie
„gebildet«
haben sich dabei insofern eigentümlich gestaltet und von den anderen
Ausdrücken abgezweigt, daß
von ganzen Völkern
nicht sowohl
sie
oder der Menschheit als von der höheren intellektuellen Schicht innerhalb eines Volkes gebraucht werden; bei „Bildung" wird
mehr die
eigne Tätigkeit, die selbständige Aneignung seitens des Individuums be-
noch sehr viel. Denn die Idee der Moralität gehört noch zur Kultur; der
Gebrauch dieser Idee aber, welcher nur auf das Sittenähnliche in der Ehrliebe und der äußeren Anständigkeit hinausläuft, macht bloß die Zivilisierung
aus." Pestalozzi XII, 154 sagt in ähnlicher Tendenz: „Die kollektive Existenz
unseres Geschlechts kann dasselbe nur zivilisieren, sie kann es nicht kultiDie
vieren."
spezifisch
literarische
Wolf verfochten, namentlich
F. A.
in
Kultur
hat
mit
besonderer
Energie
der berühmten Abhandlung, die das
„Museum der Altertums-Wissenschaft" einleitet (1807). Der Unterschied von
Kultur und Zivilisation wird ihm zum Mittel, die Griechen und auch die
Römer über alle anderen Völker hinauszuheben. Als Hauptmerkmal echter
Kultur erscheint
Kultur
ist
dabei
der Besitz
einer
allen
der durch Ausbildung von Literatur
Stand der Gesellschaft.
S,
S. 16:
gemeinsamen Literatur; die
und Kunst hervorgebrachte
„Eine der wichtigsten Verschiedenheiten
und diesen Nationen ist die, daß die ersten gar nicht oder nur
wenige Stufen sich über die Art von Bildung erheben, welche man bürgerim Gegensatze höherer eigentlicher
liche Polizierung oder Zivilisation,
Geisteskultur, nennen sollte."
S. 17, ,,jene höhere Kultur, die geistige oder
literarische."
S. 18, „Asiaten und Afrikaner werden, als literarisch nicht
kultivierte, nur zivilisierte Völker, unbedenklich von unseren Grenzen ausgeschlossen." Jener ganzen Zeit sind „Europa" und „Kultur" eng assoziiert.
Dieser Unterscheidung von Kultur und Zivilisation folgt auch W. v. Humboldt.
* S. darüber Imelmann, Ausg. von Klopstocks Oden, S. 86;
Paulsen
Bildung
in Reins Enzyklop. Handbuch der Pädagogik; Biese in d. N.
Art.
unter jenen
Jahrb. für das klass. Altertum, Jahrgang 1902, S. 241.
Der Ausdruck besagte aber weit mehr als heute nach der abschleifenden
Wirkung des Jahrhunderts; das will auch bei Schleiermachers „Reden über
'
die Religion an die Gebildeten unter ihren Verächtern" beachtet sein.
über den Sinn des Ausdrucks bei den Romantikem
mantische Schule",
S. 420, 430.
s.
bei
Näheres
Haym, „Die ro-
Kultur.
So wird
tont. *
sie
233
wohl der Kultur als etwas Innerlicheres entgegen-
— Die Abgrenzung von Kultur und
gehalten.
dings sehr ins Unsichere
und zwar
geraten, 2
Zivilisation
ist
neuer-
insofern
nicht
ohne
einen sachlichen Grund, als jene innere Kultur, die unseren großen
Dichtern und Denkern vorschwebte, und die sich deutlich von aller
bloßen
Zivilisation
abheben
Boden mehr hat.
keinen
unserer Zeit
in
wollte,
Auch gehen
die Nationen
festen
auseinander;
hier
wo
wir Deutschen von „Kultur" sprechen, sagen die Engländer und Fran-
zosen „Zivilisation ".3
den
über
folgen;
Doch das
allgemeinen
sich
hier
von
„Kultur"
Zweifel, die nähere Fassung aber
weiter
ver-
besteht
kein
nicht
läßt
Sinn
Kräftige
völlig herrenlos, jeder
ist
mag ihr seinen Stempel verleihen.
Mag
aber der Begriff der Kultur heute
sein,' sicherlich
bezeichnet
er
ein
altes
noch
Problem.
so unbestimmt
Auch
die antike
Welt konnte sich der Anerkennung eines großen Gegensatzes zwischen
den Völkern, sowie der verschiedener
eines Volkes
nicht
entziehen;
die
intellektueller Stufen
Höhe
innerhalb
des attischen Lebens aber
mußte sowohl das Selbstbewußtsein der griechischen Kultur steigern
als innerhalb des griechischen Lebens eine schroffere Scheidung erEiner vollen
zeugen.
Würdigung des Kulturproblems
wirkte hier
freilich manches entgegen: die nationale Abschließung ließ den höheren
Stand leicht
als
bloße Naturgabe eines besonderen Volkes erscheinen,
zugleich setzte die geschichtliche Ansicht von einem endlosen Kreislauf der
Dinge allem Fortschreiten enge Grenzen und hemmte leicht
eine unbefangene
viel
Neigung,
erkennen
;
'
Erforschung der Anfänge.
Aufsteigen aus
Andererseits
bestand
einem rohen Naturstande anzu-
der Scheidung der Menschheit in Griechen und Barbaren
mußte aber
fels,
ein
die Erweiterung
des Horizontes und die engere Ver-
Über die Probleme im Begriff der Bildung s. neuerdings O. Weißen-
„Die Bildungswirren der Gegenwart."
^
Näheres darüber s. Barth, die Philosophie der Geschichte als Sozio-
logie, S. 253.
*
Bei
der Übersetzung eines Artikels von mir „Religion und Kultur"
No, 3, pag. 114) wird zu Kultur angemerkt:
in der Liberte Chretienne (1907,
Nous n'avons guere l'habitude, en fran^is, d'employer ce mot Sans quelque
d^terminatif:
,,la
culture intellectuelle", „la culture des lettres".
Zu den Problemen des Menschenlebens.
234
bindung der Völker entgegenwirken,
In derselben
Zeit
wo
aber,
die
mit Alexander begannen.*
der Völker
der Gegensatz
verblaßte,
Welt der Gegensatz von
verschärfte sich innerhalb der griechischen
Gebildet und Ungebildet, indem nun lediglich ein gelehrtes Studium
den ererbten Kulturgütern vollauf teilnehmen
an
heit ist das spätere
Im
ließ. 2
Wahr-
In
Altertum voller Betrachtungen zum Kulturproblem.
christlichen Altertum
und im
Mittelalter
tritt
diese Frage zurück,
um in der Renaissance mit verstärkter Kraft wieder aufzuleben.
Seit-
dem steht die Kultur im Mittelpunkt der geistigen Arbeit; am Kampf
um sie sind alle Gegensätze der Neuzeit beteiligt:
der Idealismus
von innen her aufbauen, der Realismus sie von außen her
will sie
zusammenfügen; künstlerische, intellektuelle, ethische Fassungen durchkreuzen sich und bestreiten einander die Oberhand;
nicht an
auch
Im Verlauf des
Mischungen mannigfacher Art.
fehlt es
19. Jahr-
hunderts hat ein Zusammenwirken von Geschichte und Naturforochung
die
spekulative
ältere
Behandlung dieser Fragen mehr und mehr
weichen lassen; zugleich werden die
exakt-wissenschaftlichen
einer
Bedingungen
seelischen
des
Kulturlebens genauer
erforscht,^
und
während der Stoff massenhaft anschwillt, erzeugt das Bedürfnis eines
Gesamtbildes neue Versuche einer Kulturphilosophie.
Von den zahl-
und Kontroversen
reichen dadurch erzeugten Problemen
seien hier
nur diejenigen ausgewählt, welche das Lebens- und Geistesproblem
unmittelbar berühren.
^
Das
gibt
nicht
nur der
Philosophie
einen
kosmopolitischen Zug,
sondern verwandelt auch sonst die Denkweise. Bemerkenswert ist, was Strabo
(Geographica,
Sk
Tou
am Schluß des 1. Buches) von Eratosthenes berichtet: in\ -reXei
oux ETratveaa? tou? Stya SiaipoGvxa; areav to twv avO'paxwv
ü7:o{jLvr'|jLaT05
te "EXXrjva? xa\ ßapßa'pou;,
— ßeXTiov etva(
TcXfd-o?
e'i?
Taura.
noXXou^ yap xa\ tcTv 'EXXTJvtüV
etvat
cprjaiv
xaxou? xai
«pexf y.ai xax(a: Staipetv
tiIjv
ßapßaptov aoTeiou?.
Strabo verteidigt dagegen den Vorrang der Hellenen damit, daß dort gesetzliche Ordnung
Tots
(AEv
'
und Bildung überwiege, bei den anderen aber das Gegenteil:
iTzixpaztl
TO
vo[JLt{j.ov
xa\ TO :iatS£(a; xa\ Xo'ytov o^xtiov, Tot; Se TavavT(a.
Schon bei Plato und Aristoteles hat 7rai5e(a neben der Bedeutung der
Erziehung auch die weitere der Bildung.
aristotelische
Zusammenstellung:
(«XoÜTos, euye'veta, aptTr^, TcatSeta), Pol. 1291b,
xttl
euyevEta,
1296b, 18: iXsudepfa,
Bezeichnend dafür
Reichtum,
tiXoütoc,
Adel,
ist
Tüchtigkeit,
z.
B.
die
Bildung
28 (s. ähnlich 1293 b, 37 natSeta
TtatSeta,
euyevsta,
1317b, 39: y^^°S
ihm entsprechen TOnaiSeufievo; und a7:a(8euTos durchaus
unserem „gebildet" und „ungebildet".
* S.
darüber das wertvolle Buch von Vierkandt, „Naturvölker und
jcXouTo;, TtatSet'a).
Kulturvölker.
Bei
Ein Beitrag zur Sozialpsychologie", 1896.
Kultur.
b)
235
Kritische Erwägung.
Das Problem des Wesens und Wertes der Kultur.
a.
sich um so mehr verDenken mit ihnen beschäftigt.
Der
Begriff soll alles zusammenfassen, was den Menschen und die Mensch-
Die Kultur gehört zu den Größen, die
mehr
wickeln, je
heit
unser
sich
über die bloße Natur hinaushebt, aber worin besteht dies Mehr
Gelangt der Mensch nur zu einer größeren Selb-
gegenüber jener?
und Macht innerhalb eines gegebenen Daseins, und ver-
ständigkeit
mag er seine Umgebung nur weiter zu überschauen wie geschickter
für sich zu verwerten, oder erscheint bei
Art des Lebens,
eröffnen
sich
ihm eine wesentlich neue
neue Tiefen und gestatten ihm, ein
neues Reich der Wirklichkeit aufzubauen?
Dort würde nur eine
Außenkultur, hier eine Innenkultur erreicht,
dort eine bloße Zivili-
sation,
über jene kann kein Zweifel
echte Geisteskultur,
eine
hier
sein, 'die
Möglichkeit dieser wird hart bestritten.
Wie aber der Inhalt der Kultur, so ist auch ihr Umfang keineswegs sicher.
Zweifellos
liegt
in
eine Versetzung des mensch-
ihr
lichen Lebens in größere Tätigkeit, ja ein Gründen seiner auf eignes
Tun, wie das auch schon der Ausdruck
indem
anzeigt,
er
an das
Bestellen eines Ackers gegenüber der wildwachsenden Natur erinnert.
Aber umfaßt diese Tätigkeit alles, was irgend dem Menschen eigentümlich
ist,
oder
Möglichkeiten
ist sie
nur eine Seite des Lebens, neben der andere
Daß
bleiben?
hier
ein
Problem
vorliegt,
bekundet
schon die Unsicherheit über das Verhältnis der Religion zur Kultur:
bald wird jene zu
ihr gerechnet,
Stande der Kultur zu
und
hängen; bald
es scheint
die
Religion
am
scheinen
sie
Gegensätze, die
einander durchkreuzen und hemmen, wie denn
oft
genug die einen
von der Religion aus die Kultur, die anderen von der Kultur aus
die Religion bekämpfen.
Nicht
Kultur.
viel
Dient
anders steht es mit der Frage nach
sie
dem Werte der
zur Bezeichnung alles dessen, was den Menschen
über den Stand der rohen Natur hinaus zu
dem der Gesittung und
muß sie als der höchste aller Werte erscheinen,
muß sich innerhalb ihrer alles begründen, was uns irgend
Bildung führt, so
und
es
schätzbar sein
will.
Aber zugleich
ist
die Geschichte voller Klagen
über Schäden und Gefahren der Kultur,
sie
so sehr, daß
Danaergeschenk
die
ganze Kultur wie
ein
steigern sich zuweilen
erscheint.
236
Zu den Problemen des Menschenlebens.
Namentlich
in
drei
Richtungen
ist
die
Kultur von alters her ein
die
Kultur
Gegenstand harter Angriffe gewesen.
Von
der Religion
menschlicher
Kraft
aus
konnte
als
eine
und
Steigerung
als
Stärkung
eine
menschlichen
Selbst-
bewußtseins schwere Bedenken erregen.
Ein frommer Sinn sah in
dem kühnen
Aufstreben
ein
Vermögens,
ein
der Menschheit
Überschreiten
Überspannen eignen
naturgewiesener
Mangel an frommer Gesinnung.
Schranken,
einen
Die Mißstände und Rückschläge
des Kulturlebens erschienen dann als eine Strafe für solchen Frevel.
Eine derartige Überzeugung wirkt von Babylon her in der Erzählung
vom Sündenfall und vom Turm, der bis zum Himmel reichen soll,
sie
erscheint
in
den
Prometheussagen,
Zuspitzung gegen einen
sie
ist,
in
der besonderen
übermäßigen Wissensdurst, auch
in
den
Faustlegenden unverkennbar.
Auf dem eignen Boden der Menschheit aber griff oft der Zweifel
um
ob die Kultur dem Menschen das Glück
sich;
bringe, das sie
ihm
Verwicklung des Lebens,
sie
bildet
in
Wahrheit
Sie erzeugt eine große
zuversichtlich verheißt.
künstliche Bedürfnisse aus, sie
bindet den Menschen mehr und mehr an seine Umgebung, sie schafft
ihm Arbeit und Mühe,
sie
erweckt unerreichbare Wünsche und wilde
Leidenschaften, sie mag mit dem allen als ein Losreißen des Menschen
von
seiner
natürlichen Grundlage
erscheinen,
äußeren Glanz innerlich unglücklich mache.
das
Auch
ihn
bei
allem
derartige Stim-
mungen sind uralt, sie tauchen z. B. bei den alten Juden auf, wie
Hosea und Jesaias zeigen. * Besonders voller Zweifel war das spätere
griechische Altertum, ein Widerwille gegen das Raffinement der da-
maligen Kultur, eine Sehnsucht nach einfachen Zuständen und schlichter
Lebenshaltung griff weiter und weiter um sich.
Stimmung namentlich
diese
in derberer, die Stoiker in
Literatur gerät unter
ihren
Zum Ausdruck haben
die Philosophen gebracht,
die Kyniker
etwas feinerer Art; aber auch die schöne
Einfluß
breitung im gemeinsamen Leben. ^
und bekundet damit
ihre Ver-
In der Neuzeit stellt namentlich
Rousseau das ProJDlem aufs deutlichste vor Augen, mit seiner sensitiven,
aufgeregten
und aufregenden Art
hat
er es der
modernen
Menschheit zwingend auferlegt.
^
S.
darüber Budde „das nomadische Ideal im alten Testament" (Preuß.
Jahrbücher, Bd. 85.)
"^
Anziehende Ausführungen darüber gibt E. Rohde „Der griechische
Roman und seine Vorläufer".
Kultur.
237
Das drohende Entweichen des Glücks hätte sich etwa ertragen
lassen,
wenn dabei
Wachstum der
ein
Tüchtigkeit
Menschen
des
Aber das war es nicht, vielmehr pflegen
außer Zweifel gewesen wäre.
den Klagen über das sinkende Glück solche über eine Minderung
der Kraft und Tüchtigkeit durch den Fortgang der Kultur zur Seite
zu
Die Kultur,
gehen.
indem
sie
seines Handelns
sie
stellt
hören
so
wir,
schwächt
den
Menschen,
ihn von anderen abhängig macht, sie erhebt die Wirkung
im
Zusammensein zur Hauptsache,
gesellschaftlichen
damit die Leistung vor die Gesinnung und droht bis in
und UnMehr und mehr spielt der Einzelne nur eine ihm
von der Gesellschaft zugewiesene Rolle, und es wird sein Leben mehr
und mehr ihm selbst etwas fremdes, es hängt ihm nur äußerlich an;
wie könnte er dabei eine Größe der Seele bewahren, ein wahrhaftiger,
die innerlichsten Gefühle hinein das Leben ins Scheinhafte
wahre zu führen.
kräftiger,
ganzer Mensch sein?
Wohl fehlt es der Kultur nicht an Anwälten gegen solche Beschuldigungen.
Jene
Schäden,
so
heißt
es,
bloße
seien
Begleit-
erscheinungen, Schatten, ohne die kein Licht besteht; nur der Mensch
ziehe ins Kleine herab
und mache damit
groß und unangreifbar
sei.
menschlichen
Lebenskreises;
bunden, wird
sie
sich
— Indes
vom
nicht
sie
ist
zweifelhaft,
was an
sich
die Kultur liegt innerhalb des
an
seinen
Zustand
kleirmienschlichen Getriebe
ge-
irgend ab-
heben können unter deutlicher Scheidung von wesentlichem Gehalt
und menschlicher Zutat, von Recht und Unrecht? So bleiben jene
Bedenken einstweilen unwiderlegt, und zugleich bleibt die Frage offen,
ob die Kultur ein Segen oder
ß.
Fluch für die Menschheit
ein
sei.
Das Problem des Inhalts der Kultur.
Darüber ist kein Zweifel, daß die Kultur das Dasein des Menschen
in
höherem Grade auf seine eigne Tätigkeit
allgemeinen Begriff der Tätigkeit
ist
Umgebung
stellt.
Aber mit dem
noch recht wenig gewonnen,
an sich ziehen und sich
umwandelnd in sie ergießen, ohne sich selber näher zu determinieren,
die Tätigkeit kann
ihre
ohne dem Leben einen
festen Mittelpunkt, eine beherrschende Haupt-
eigentümliche
richtung,
eine
aus
Antwort sofort
der
die damit entsteht,
ist
nicht
eine
Durchbildung zu geben;
Frage
hervor.
wächst
so
Die Aufgabe
von der Arbeit der Weltgeschichte
in
aber,
recht
verschiedener Weise gelöst, mannigfache Arten der Kultur sind entstanden,
von denen keine einzelne
voll
und dauernd zu genügen
Zu den Problemen des Menschenlebens.
238
scheint,
und
die
und Schätzungen
mit ihren widerstreitenden Zielen
unmöglich aneinanderlegen
sich
lassen.
Es heben sich aber aus der Bewegung unseres gesamten Kulturmit ausgeprägter Gestalt namentlich
kreises
drei Arten
der Kultur
hervor: eine künstlerische, eine ethische, eine dynamische; im Griechen-
tum, Christentum, modernen Leben sind
Im Griechentum
langt.
bildet
sie
zur Verkörperung ge-
den Kern der Kulturarbeit die Ver-
bindung der Elemente, welche die Natur entgegenbringt, zu einem
harmonisch geordneten, von innerem Leben erfüllten Ganzen.
Verbindung, Ordnung,
eigne Tätigkeit bereiten,
Belebung kann dem Menschen
sie
entringt
der Zerstreuung
der sinnlichen Eindrücke ein beharrendes
Weltbild,
sie
schlossenen
die
stellt
Individuen in
Gemeinschaft,
Triebe der Seele,
sie
Diese
nur seine
und Flucht
und zusammenhängendes
das feste
Gefüge einer ge-
verbindet die einzelnen
Kräfte
und
ohne irgend etwas davon aufzugeben oder abzu-
schwächen, zu einem Gesamtwerk des Lebens,
sie
vollzieht an allen
Wendung vom Chaos zum Kosmos.
Durch solches
Wirken sind Natur und Geist in eine enge und fruchtbare Beziehung
Stellen
eine
gesetzt,
ist
ein
kräftiges,
tätiges,
freudiges Leben geschaffen,
ganze Umkreis des Daseins veredelt und durchgebildet.
Fragen und Zweifel blieben nicht aus.
ist
der
Aber auch
Das Ganze ruht auf der
Überzeugung, daß das Leben von Grund aus eine sichere Richtung
besitzt, und diese Überzeugung geriet mehr und mehr
Wanken; die Form, welche hier das Leben beherrscht, konnte
zur Vernunft
ins
solche Stellung nur wahren, solange sie eine Seele in sich trug,
die schien
sie
nicht
und
dauernd bewahren zu können; schließlich ge-
wannen die Verwicklungen des Lebens so sehr die Oberhand, und
schien der Mensch im innersten Kern seines Wesens so schwer bedroht,
daß
sein Grundverhältnis
zur Welt und die Rettung seiner
Seele zur dringendsten aller Aufgaben wurde.
Dieser Aufgabe unterzog sich das Christentum, bei vollster An-
erkennung des Nein unternahm
es
den Menschen zu einem über-
legenen Ja zu führen, inmitten ungeheurer Erschütterung
feste
Pole zu wahren.
Das
dem Leben
forderte eine unbedingte Konzentration
auf eine Aufgabe ethischer Art, ein völlig neues Leben galt es gegen-
über
dem
nächsten
losigkeit jenes
Dasein zu gewinnen, der Härte und Seelenwurde ein Reich barmherziger Liebe und kindlicher
Hingebung entgegengehalten.
In Entwicklung dessen ist eine gewaltige
Vertiefung des Lebens erfolgt, unsichtbare Zusammenhänge taten sich
Kultur.
239
eine große Weichheit der Empfindung und ein großer Ernst der
auf,
Gesinnung gingen Hand
in
Hand miteinander; ZeitUches und Ewiges,
Endliches und Unendliches, Menschliches und Göttliches traten hier
Aber auf dem Boden
engsten Kontakt.
in
der Geschichte
blieb
Denkweise überwiegend transzendent und gewann kein sicheres
diese
Verhältnis zur Weltumgebung, neben einem Kreise reiner Innerlichr
keit
verblieb daher die übrige Welt unergriffen und ungeläutert, die
Flucht
in
die
Welt des Gemütes
ließ
leicht
die
Arbeit
an
den
Widerständen des Daseins als nebensächlich erscheinen und gefährdete
damit die männliche Kraft des Ganzen.
zum Kern alles Strebens.
Überwindung der Hemmung, der Aus-
Diese Arbeit aber wurde der Neuzeit
Der Gedanke einer
treibung
alles
vollen
Dunkels
tritt
hier in
den Vordergrund, die eigne Be-
wegung des Lebens, seine Steigerung ins Unbegrenzte wird hier
zum Ziel aller Ziele, zum vollgenügenden Glück. Den Menschen
scheint
alles
'
hier
vornehmlich auszuzeichnen seine Überlegenheit gegen
gegebene Maß,
sein Vermögen, die eigne Kraft weiter und
immer neue Wege zu bahnen, immer neue An-
weiter zu steigern,
fänge zu setzen.
Die Bewegung, die daraus hervorgeht,
gängig neue Bilder
läßt durch-
vom All, dem menschlichen Zusammensein, der
Seele des Einzelnen entstehen, sie schafft eine neue Art der Arbeit,
in
welcher diese zuerst das Bewußtsein einer Weltüberlegenheit ge-
winnt.
Mehr
als
irgend
sonst wird
seines Daseins, überall erfolgt hier
heit, ein
Leben ein
Unermeßliche Mut und
in
der Mensch
zum Herrn
Beleben alles Schlummernden, ein Befreien alles Gebundenen,
überall wird das
uns
hier
ein Aufrütteln aus träger Starr-
rastloses Vorwärtsstreben,
Kraft.
Aber wenn
die
schwellen ins
Ergebnisse dessen
tausendfachen wohltätigen Wirkungen vor Augen stehen, vor
Augen stehen uns auch die zahllosen Verwicklungen, die jene Belebung und Befreiung gebracht hat, an die freudig vordringende
Vernunft hat sich soviel Unvernunft angeschlossen, mit den Erfolgen
des Wachstums des Geisteslebens ist so viel kleinmenschliche Irrung
und Leidenschaft aufgeschossen, daß die alleinseligmachende Kraft
Immer
der modernen Kultur uns sehr ins Unsichere geraten ist.
weniger läßt sich auch der Zweifel unterdrücken, ob selbst beim Gelingen der Bewegung der Mensch ganz und gar in sie aufgehen kann.
Denn als denkendes Wesen überschaut er die Bewegung, faßt er sie
ein Ganzes und muß er von ihr eine bleibende Förderung seines
Wesens verlangen; von hier aus wird ihm eine Kultur, die immer
in
Zu den Problemen des Menschenlebens.
240
nur ungestüm vorwärts drängt, nie einen zeitüberlegenen Besitz ergibt,
und unerträglich werden.
sinnlos
Dies
nacheinander entwickelt, aber es hat einander
alles hat sich
nicht einfach abgelöst; was äußerlich versank, behält eine innere Gegen-
Nun
wart und behauptet einen Einfluß auf das menschliche Leben.
aber ist die Grundrichtung und der Gesamtcharakter jener Gestaltungen
so verschieden, daß nur eine flache Denkart eine unmittelbare Ver-
bindung für möglich erachten kann.
Sie
ist
um so weniger mög-
das geschichtliche Bewußtsein der Gegenwart uns die Unter-
lich, als
schiede mit vollster Schärfe gewahren läßt. So stehen die verschiedenen
Lösungen wie fremdartig gegeneinander und führen, wenn auch meist
miteinander
versteckt,
Kampf und
Die künstlerische Kultur
Krieg.
eng und düster, die dynamische für form-
ethische für
erklärt die
und ruhelos; der ethischen muß die künstlerische als flachoptimistisch
und
naturgebunden,
gelten; die
die
dynamische
dynamische wird
wegung und Forttrieb finden.
steht der
als
selbstbewußt und trotzig
den anderen Formen zu wenig Be-
in
Und zwischen all diesen Gegensätzen
Mensch der Gegenwart; wird
und geistig herabgedrückt werden?
er nicht
von ihnen zerrieben
Er kann jene verschiedenartigen
Kulturtypen weder verbinden, noch auch zu Gunsten eines auf die
anderen verzichten; er müßte,
um jedem sein Recht zu geben und
von seinem Unrecht zu befreien, eine sichere Überlegenheit gewinnen,
aber er ermangelt nicht nur einer solchen, er sieht nicht einmal, in
welcher Richtung
y.
Die
Frage,
sie
zu suchen
sei.
Das unsichere Verhältnis des Menschen zur Kultur.
Verwicklungen
wie sich
Mensch
steigern
weiter
sich
und Kultur
bei
Erwägung
der
Nur
zueinander verhalten.
zweierlei scheint hier möglich: entweder hat die Kultur dem Menschen
Nun aber ist keines von
Mühe ersehen läßt.
Wäre die Kultur ein bloßes Mittel für das Wohl und den Zu-
oder der Mensch der Kultur zu dienen.
beiden möglich, wie sich ohne
stand
des Menschen,
so
müßte
ihr
Wachstum
sein
Leben
immer
angenehmer gestalten, so müßte das Mehr der Kultur zugleich
Mehr des Glückes
besagen.
Das aber
tut
es
nicht.
menschlichen Behagen scheint die Kultur mehr schädlich
zu
sein.
Sie
erzeugt
Mühe und Arbeit,
sie
ein
Denn dem
als nützlich
unbegrenzte Wünsche und kostet unsägliche
verwickelt
in
Sorgen
und Aufregungen,
sie
umfängt uns mit festen Bindungen, sie verlangt Gehorsam und Opfer;
Kultur.
daß das
241
die Lust, die Annehmlichkeit des Lebens erhöhe, läßt
alles
behaupten.
sich schwerlich
Jene befindet sich weit besser, und es
wird sich der Mensch weit eher zufrieden fühlen auf niederen Stufen
der Kultur, auch werden weit eher Individuen geringer
geistiger Regung
dazu gelangen.
hoher
als
Wäre zufriedenes und angenehmes
Dasein das höchste Ziel, wie sehr müßten wir Kulturmenschen die
brasilianischen
Neger mit ihrem sorglosen Lebensgenuß beneiden!
So wäre auch
leicht
zu zeigen, daß geistige Bewegungen, welche das
Glück zum höchsten Ziele machten, wie der Epikureismus und der
für den inneren
Utilitarismus,
geleistet
sie
haben.
Innerhalb
Aufbau der Kultur verzweifelt wenig
eines
gegebenen Kulturstandes mochten
manche Härte mildern, mancher Not entgegenwirken: das Leben
zu heben, Neuem die Bahn zu brechen, geht über ihr
wesentlich
Vermögen.
So
nur der andere Weg: die Kultur als Selbstzweck an-
bleibt
zuerkennen und den Menschen zu einem
Fortgang zu machen.
bloßen Mittel für ihren
Eine solche Fassung hat für sich den Ein-
druck innerer Größe: unvergleichlich wächst die Kultur, wenn
sich bei
jenem Selbständigwerden
in ein
sie
Ganzes zusammenfaßt und
mit der Kraft einer eignen, inneren Notwendigkeit wirkt; der Mensch
aber scheint bei aller äußeren Unterordnung innerlich nur zu wachsen,
wenn
er alle
Sorge
um den eignen Zustand ablegt und
sich
ganz
dem Strome des Weltlebens hingibt. Hegels System hat dieser Denkweise eine
großartige Verkörperung gegeben.
hinaus übt
sie
Aber weit darüber
im modernen Leben eine nicht geringe Macht.
In-
menschlichen Verhältnisse und inWerdens und Vergehens der Geschlechter gibt heute
Überzeugung vielen einen Halt und Trost, daß durch alles
mitten alles Unerquicklichen der
mitten
die
des
Mühen hindurch die Kultur ihren sicheren Weg verfolge, und daß
Gewinn auch dem Leben und Wirken des um sie bemühten
ihr
Menschen einen Sinn und Wert, sowie eine bleibende Dauer verleihe.
«Viele werden
vorbeiziehen und die Wissenschaft wird
wachsen."
Aber so anziehend dieser Gedanke, zum Siege vermag er nicht
zu gelangen.
Denn
es gibt keine Kultur freischwebender Art; eine
Kultur, die sich gänzlich
bloßen
Mittel
vom Menschen ablösen und ihn zu einem
herabsetzen
wollte,
würde
selbst
ins
Leere
fallen.
Immer liegt die Kultur innerhalb des menschlichen Lebens, und muß
sie
diesem etwas
sein,
der Mensch
behaupten haben, wenn
Eucken, Grundbegriffe.
sie
4. Aufl.
seine
muß in ihr ein geistiges Selbst zu
volle Kraft
gewinnen, ihn durch
16
Zu den Problemen des Menschenlebens.
242
Hemmungen hindurch hohe Ziele erreichen lassen soll. Eine
unpersönliche, vom Menschen völlig abgelöste Kultur wäre ein Ge-
alle
und
spenst ohne Fleisch
Blut; soweit dies in unseren Vorstellungen
eine Wirklichkeit erlangte, würde es uns ins Irre locken, unbekannten
uns aufzuopfern heißen,
Zielen
das Leben
einer Seele
berauben.
Und wie könnte die Hoffnung auf die Zukunft uns in den Mühen
und Kämpfen der Gegenwart aufrecht erhalten und freudig stimmen,
wenn diese Zukunft niemandes Sache, niemandes Freude, niemandes
Förderung wäre?
Unsere eigne Zeit
immer
uns
stellt
klarer
daß
vor Augen,
jene Selbstaufopferung des Menschen an die Kultur sich schlechter-
dings nicht vollziehen
Denn immer stärker
läßt.
bricht
aus allem
und lärmenden Kulturbetriebe wieder das Verlangen nach
Entfaltung und Förderung des lebendigen Menschen, nach Bildung
hastigen
der Seele, nach Rettung eines geistigen Selbst hervor; wir erkennen
zugleich,
nicht
daß dies für die eigne Wahrheit und Tiefe der Kultur
zu entbehren
ist
solchen Erfahrungen
In
tritt
ihn dieses nicht nach
seinen
vor
deutlich
Augen, daß der Mensch kein bloßes Gefäß des Kulturlebens
Bedürfnissen wie weiches
daß
ist,
Wachs
so
oder anders formt, sondern daß er ihm eine selbständige Art entgegenzusetzen
Kultur
hat,
alt
nicht
schreitet
sicherm Zuge
Die
die nicht auf Befriedigung verzichten kann.
fort,
aus
einem
vielmehr
ihr
scheint
in
innewohnenden Zwange
ihr
alle
in
besondere Gestalt
zu werden und sich auszuleben; immer wieder bedarf es neuer
Anfänge, eines Hervorbrechens ursprünglichen Lebens, vor allem aber
So erging es dem späteren Altertum; das Kulturleben kam erst wieder in Fluß, als neue Völker es aufnahmen und
durch frische Kräfte verjüngten.
Sollte auch der Gegenwart eine
neuer Menschen.
solche Verjüngung notwendig
sein,
sei
es
durch neue Völker,
sei
es durch neu aufsteigende, geistig noch minder verbrauchte Klassen?
Wie dem
sein
mag, der lebendige Mensch
behauptet
seine
Selbständigkeit gegen allen Versuch, ihn zu einem bloßen Werkzeug
herabzusetzen. Aber auch die Kultur, so sahen wir, darf nicht zu
einem bloßen Mittel sinken, wenn sie nicht einer Auflösung verfallen will.
So befinden wir uns
in
einem schweren Dilemma, wir
müssen darüber hinaus, aber zunächst sehen wir nicht, wie das möglich sein sollte.
bald
nach
Im Durchschnitt des Lebens aber werden wir heute
dieser, bald
nach jener Seite getrieben, zwischen leerer
Subjektivität und seelenloser Arbeit schwanken wir ratlos hin und her.
Kultur.
Alle diese Verwicklungen
243
treffen in der Gegenwart zusammen
Vor allem peinlich wirkt die Unsicher-
und
steigern sich gegenseitig.
heit,
die über unser eignes Verhältnis zur Kultur besteht, das Fehlen
umfassenden und leitenden
eines
zur
eignen
zur
Sache,
Zieles,
Erhaltung
zwingenden Notwendigkeit macht und
dem
menschliche Getriebe hinaushebt,
das uns die
unseres
sie
Kulturarbeit
geistigen
zugleich
Selbst,
über das
zur
klein-
wir sonst wehrlos verfallen
Schon das verhindert uns, nach einer neuen charakteristischen
sind.
Art der Kultur gegenüber den verschiedenen Gestaltungen zu streben,
die
von naher oder ferner Vergangenheit auf uns eindringen und
uns einnehmen, ohne uns
voll
die daraus entstehen,
uns schließlich der Wert und das Wesen
aller
fällt
zu befriedigen.
Kultur ins Ungewisse, notdürftig genug verdeckt gewandte Remit schönklingenden
flexion
Reden und ausgeklügelten «Gesichts-
punkten" den Mangel eines Kernes des Ganzen.
schließlich
zwischen
dem, was
solches erstrebt wird,
immer
ausgeht,
weiter
wird
der Abstand
Ziel verkündet und was in Wahrheit als
immer größer wird damit die Unwahrhaftig-
als
Dem muß widerstanden werden; die wachsende
des Lebens.
Unzufriedenheit
zeigt
schon im Gange
ist.
c)
Unerträglich wird
jene aufgeputzte Scheinkultur, wie sie namentlich von
all
unsern Millionenstädten
keit
den Wirren,
In all
genug, daß eine
deutlich
solche
Bewegung
Forderungen für ein wahrhaftiges Kulturleben.
a.
Die Notwendigkeit einer tieferen Begründung.
Die Philosophie
mag
in
solchen
Bewegungen und
Erschütter-
ungen eine noch so bescheidene Rolle haben, der Aufgabe entziehen kann
sie sich
nicht.
Ihre Sache wird
es vor allem sein, die Richtungen
herauszuarbeiten, die das Streben einzuschlagen hat, um uns das Leben
wieder aus einem «Geschäft" zu einem «Dasein« zu machen (nach
J.
Burckhardt).
Dazu aber bedarf
Kultur ganz unser eigen
sei
es
vornehmlich dessen, daß die
und zur zwingenden Notwendigkeit
unserer Selbsterhaltung werde, ohne unter die Kleinheit der bloßen
Lust zu geraten.
lebens
Dafür aber
bietet
unsere
Fassung des Geistes-
und seines Verhältnisses zum Menschen einen gangbaren Weg.
Denn mit dem Selbständigwerden des Geisteslebens, wie wir es vertreten, wird die Kultur, die seiner Entfaltung dient, von dem flachen
Menschengetriebe befreit und auf eine tiefere Grundlage aufgetragen,
16'
Zu den Problemen des Menschenlebens.
244
aber sie wird dabei nicht dem Menschen entfremdet, da er seiner eigentümlichen Art nach im Ganzen des Geisteslebens allererst sein echtes
Wesen, die Möglichkeit eines wahrhaftigen Beisichselbstseins
Bei
Fassung
solcher
arbeitet
er
in
der
Kultur nicht
für
findet.
fremde,
sondern für eigne Zwecke, und vermag er auch in der weitesten
Ausdehnung seines Strebens einen beherrschenden Mittelpunkt festSo
zuhalten.
ist
Menschen und
es
das Geistesleben
unmittelbar zu verschmelzen
ohne
sie
die
fertige Tatsache,
uns bequemer Weise
welches das ganze Leben
Ideal,
einander
und damit das eine dem andern auf-
Die Verbindung nämlich erscheint hier nicht
zuopfern.
hohes
unserem Sinne, das den
in
die Kultur aufs engste verbindet,
aufregt
und
in
eine
als
sondern
zufällt,
ein
als
Bewegung
Zusammenhängen erscheint die Kultur als unsere
Mitarbeit an einer großen Bewegung des Alls, welche die Wirklich-
versetzt.
keit
In solchen
höheren Stufe, der Stufe des Beisichselbstseins, zuführt.
einer
Hinter unserer Arbeit steht und innerhalb unserer Arbeit wirkt damit die Kraft des Ganzen.
Es
keineswegs bloß eine
ist
leise
Verschiebung oder gar eine
bloße Veränderung des Namens, wenn so die Kultur
von Forderungen,
Erfüllung
eine
die
allem
als
Entfaltung
Denn das gestattet
eines selbständigen Geisteslebens verstanden wird.
echten
Kulturstreben
wesentlich sind, denen aber die durchschnittliche Fassung in keiner
Weise genügt.
So
erst
wird eine Selbständigkeit der Inhalte und der Werte
Wäre die Kultur bloß ein
möglich, welche die Kulturarbeit erfüllen.
innermenschlicher Vorgang, so würde der Stand des Menschen zu
ihrem ausschließlichen Maße, so gäbe es keine Zerlegung und Scheidung
des uns umfangenden Chaos, so könnte nicht die Kultur dem menschlichen Dasein Ideale
alle
aufrüttelnde
mit zwingender Stärke vorhalten, so fehlte ihr
und vorwärtstreibende
Kraft.
Ganz anders
sich die Sache, wenn in der Kultur eine dem bloßen
stellt
Menschen über-
Bewegung anerkannt wird, die ihm den Kern seines eignen
Wesens erst aufzuschließen vermag.
Ferner kann erst die Begründung auf ein selbständiges Geistesleben der Kultur eine Größe geben. Denn wo das Leben ganz und
legne
gar auf den
bloßen Menschen beschränkt bleibt,
über seinen Zustand
keit
führt,
geistern,
da
in
ein
Leben
mit
mag der Mensch noch
raffinierte
Unterschiede
ihn
nicht irgend
dem Ganzen der Wirklichso
ersinnen,
sehr sich für
in
Größe
be-
Hochmut und Eitel-
Kultur.
keit sich
245
oder seinen Stand über andere hinausheben,
bleibt alles
klein,
Erhabenheit,
klein
vornehmlich
Größe, etwas,
echte
in.
der Sache
der Einbildung einer Größe.
in
Ehrfurcht gebieten
das
und im
Unterordnen zugleich erheben könnte, entsteht innerhalb dieses bloßmenschlichen Kreises
Dazu muß im Menschen etwas Mehr-
nicht.
dem er zugleich eine volle Überlegen-
alsmenschliches durchbrechen,
zuerkennen muß, und das er doch als irgendwie zu sich selbst
heit
gehörig betrachten darf;
erst
von da aus wird eine wahrhaftige Er-
höhung seines Wesens möglich, auch die größte aller Befreiungen,
die Befreiung des Lebens von der Enge des bloßen Menschen. Wie
dies Übermenschliche im Menschen den Quell aller echten Größe
bildet,
so bewahrt es allein die Kultur davor, ein bloßer Menschen-
dienst
zu werden, Menschendienst gegen Einzelne,
Menschendienst
auch gegen Massen.
Möchte uns stets jenes Kantische Wort gegen-
wärtig
auch
«Alles,
sein:
Erhabenste,
das
den Händen des Menschen, wenn
verkleinert
sich
unter
Idee desselben zu ihrem
sie die
Gebrauch verwenden."
Auch zur Ursprünglichkeit des Kulturlebens ist die Gegenwart
einer
ist
neuen
Stufe
der
Wirklichkeit
Kultur nicht mehr als ein
nicht
zu
entbehren.
Denn
menschlicher Zusatz zur Natur, so
muß ihre Bewegung sich immer weiter von ihrer Basis entfernen,
so muß ihr Bestand immer künstlicher, komplizierter und raffinierter
Die Kultur wird dann das Leben immer starrer festlegen,
ihm immer mehr Möglichkeiten verschließen^ es immer gebundener
werden.
machen.
Mit dem allen würde
und aller Ursprünglichkeit
heit das in
Ist
sie
die Zerstörerin aller Jugendfrische
es ein Wunder, daß, wenn die Mensch-
besonderen Lagen mit besonderer Stärke empfindet,
sie
dagegen sich aufbäumt und sich mit ganzer Seele zur Natur, zu den
einfachsten
Anfängen zurücksehnt, wie
Kindheitsalter mit seiner Frische
das Individuum in das
oft
und der Fülle seiner Möglichkeiten?
Aber eine wirkliche Rückkehr zur Natur
ist
der Menschheit ebenso
versagt wie dem Individuum die zur Kindheit, die Geschichte mit ihren
Wirkungen
läßt sich
unmöglich
streichen.
So müßten wir uns also
darin ergeben, daß die Kultur immer greisenhafter und starrer würde,
daß die Menschheit im Großen
erläge,
demselben
öden Spießbürgertum
wie die meisten Individuen im Kleinen, wenn nicht etwas
Neues ursprünglich durchbrechen, nicht frische Kräfte einsetzen, nicht
neue Möglichkeiten aufgehen könnten.
Sie
können aber nur
auf-
gehen, wenn es eine geistige Tiefe des Lebens gibt, die inmitten
Zu den Problemen des Menschenlebens.
246
und Abgelebten
Vergriffenen
alles
neue Anfänge
einer
bloßmenschlichen
Kultur
einfache Größen erzeugt, im Einfachen eine neue
setzt,
Wenn es heißt, daß alles Große einfach ist, so
erschließt.
muß das wohl eine andere Einfachheit sein als die der natürlichen
Welt
Anfänge.
wenn
Endlich entbehrt die Kultur auch der nötigen Triebkraft,
sie
nur einer gegebenen Welt etwas beifügt, nicht eine neue, uns
unentbehrliche Welt eröffnet
Kräftig
erregen
und zwingend
be-
wegen kann uns nur die Erfahrung und Empfindung eines Widerspruches im eignen Leben, die Unmöglichkeit, bei ihm abzuschließen.
Einen
derartigen
Widerspruch aber kann eine bloße Zusatz- und
Luxuskultur nun und nimmer erzeugen.
Das von ihr gewünschte
Mehr könnte man ruhig ablehnen oder etwa wie Wind und Wetter
geduldig über sich ergehen lassen, wie
schnittsleben
ja in
Wahrheit das Durch-
gegen die Kultur recht gleichgültig
innerlich
ist
und
Zwang denn als eine eigne Freude
empfindet. Wenn es auf der Höhe des Schaffens anders stand, und
wenn überhaupt solches Schaffen möglich war, so kam das daher,
sie
mehr
als
einen
daß hier die Arbeit
sozialen
als
die Erringung eines wahrhaftigen
Lebens und damit eines Beisichselbstseins galt, und daß,
einmal
Verlangen
aufgegangen
war,
die
vorgefundene
geistigen
wo solches
Lage
als
schlechterdings unerträglich, als Hemmung einer notwendigen Selbst-
erhaltung
erschien.
Mit solchem
Verlangen
nach
Selbsterhaltung
fuhr eine leidenschaftliche Glut in das Streben, die keine Rücksicht
auf
Menschen kannte und zu jedem Opfer
bereit
war, vor keinem
Hemmnis zurückwich.
Durch alle Fragen zieht sich ein und dasselbe Problem, ein
und derselbe Gegensatz: der einer echten und einer Scheinkultur.
Echt ist die Kultur nur soweit, als sie den Zusammenhang mit dem
begründenden Geistesleben wahrt und seiner Entfaltung dient, unecht
wird sie, sobald sie unter die Zwecke des bloßen Menschen sinkt
Der Kampf beider
und auch das Geistesleben dahin herabzieht.
durchdringt die ganze GeFormen
hie Geist, hie Mensch
schichte und läßt in ihr eU,'as anderes sehen als einen reinen Triumph
—
—
des Geistes.
Heute aber
heit deutlicher erfaßt,
tut es
besonders not, daß die
alte
Wahr-
die notwendige Bedingung echter Kultur klarer
herausgestellt, die Scheidung der Geister für dieses oder jenes kräftiger
vollzogen werde.
Kultur.
ß.
247
Die Notwendigkeit einer inneren Weiterbildung der Kultur.
Daß wir
einer Weiterbildung
welcher Richtung
wir
der
zu suchen
sie
Erörterung zur Genüge
bisherige
bedürfen,
Kultur
haben,
das
ließ
auch
schon
in
die
Verschiedene Haupt-
erkennen.
gestaltungen wirken zu uns von der Geschichte her, von denen wir
keine aufgeben
und die wir auch
nicht unmittelbar zusammenfassen
können; was anderes bleibt da übrig,
als
ob nicht eine Lebensbewegung vorhanden
uns danach umzusehen,
ist
und sich
weiter ver-
stärken läßt, die über den Gegensatz hinaushebt
und ihm entgegen-
genug ist,
um sich über das
zuwirken
gestattet,
die dabei universal
Ganze des Lebens zu erstrecken und seinen Befund in ein Für oder
Wider zu scheiden, und zugleich charakteristisch genug, um allem,
was sie ergreift, eine eigentümliche Gestalt zu geben. Ein Urphänomen
müßte
das jedem Einzelnen gegenwärtig
in ihr ergreifbar sein,
und das zugleich
das Ganze des Lebens
ist,
und bildender Wirkung über
mit aufrüttelnder
reicht.
Ein solches beherrschendes Urphänomen ist nun nicht dieses oder
am Geistesleben,
jenes
ist
nicht
diese
oder jene Leistung, sondern es
das Geistesleben selbst, wie wir es verstehen, die Bewegung der
Wirklichkeit zu einem Beisichselbstsein des Lebens.
Erst mit solchem
Beisichselbstsein wird überhaupt ein wahrhaftiges Sein
Übrige
nur ein Schein davon; ein solches Sein kann nicht außer-
ist
nur innerhalb ihrer liegen, es
halb der Tätigkeit, sondern
indem jene
Ganze
ein
in
sich zu
die
Aufstieg
einem beharrenden Ganzen
einzelnen Betätigungen
eines
Lebens
bloßen
oder vielmehr es wird damit
erst
im Begriff des Lebens
sonst
spruch, daß
einem
Wendung wird
Selbstleben
Oder
ist
es
nicht
diese
Lust.
eine durchgehende
*
Für
einer
und
muß
ich
erst
deutlich
einer wesenlosen, einer
gebundenen Art, und
Aufgabe daraus,
alles weitere
an
Erst
Von hier aus tritt alle Tätigkeit
unter den Gegensatz einer wesenhaften
und
Widerstets
der Begriff von Lebensinhalten verständ-
von der niederen Stufe der
selbständigen
ein
aber
eine Selbständigkeit erlangt?^
auch der Begriff des Wertes scheidet sich hier
lich,
erreicht,
der Widerspruch überwunden, der
liegt.
nie
bleibt,
entsteht,
und dieses
Damit erst wird
vertieft
hineinlegt.
zu
eine gewisse Innerlichkeit entsteht,
Fremdes gebunden
mit jener
erreicht, alles
die
es
entsteht zugleich
übhche Vermengung, die
auf meine systematischen Schriften, zu-
nächst auf die „Grimdlinien einer neuen Lebensanschauung" verweisen.
Zu den Problemen des Menschenlebens.
248
ineinander
ungeschieden
beides
verfließen
auszutreiben,
läßt,
Forderungen der Wesensbiidung scharf herauszuarbeiten und
was die Bildung eines Wesens
können,
als
Von der Kultur wird damit als echt nur
unerläßlich durchzusetzen.
gelten
die
sie
was eine
fördert,
Weiterbildung der geistigen Wirklichkeit und zugleich unseres echten
Selbst
enthält;
damit
zu
prunkend
andere, so
alles
bloßen Menschenkultur,
einer
Soweit aber jene Wesensbildung gelingt,
Befestigung
und Vertiefung des Daseins
auftreten
es
zu
einer
mag,
sinkt
Kulturkomödie.
muß eine durchgreifende
erfolgen;
der Hauptaffekt
des Lebens wird damit das Verlangen nach Wahrhaftigkeit, nach Befreiung von allem Schein.
So
entsteht ein eigentümlicher
Typus des Lebens mit strengen
Forderungen und mit aufrüttelnder Kraft; daß aber innerhalb dieses
Kreises für mannigfache Bewegungen Platz verbleibt, das bringt die
Tatsache mit
sich,
daß jene Wendung zu einem Beisichselbstsein des
Hemmungen des mensch-
Lebens sich unter den Bedingungen und
muß;
lichen Daseins vollziehen
wird dadurch möglich,
ja
eine Mehrheit
von Angriffspunkten
Wir Menschen
unentbehrlich.
sind an
das unmittelbare Dasein gebunden und bleiben auch für den Fort-
Wir können uns
gang des Lebens darauf angewiesen.
nicht
ein-
uns der wesenhaften Einheit bemächtigen
fach von jenem trennen,
und von dort aus die ganze Wirklichkeit entwickeln, sondern auch
wenn wir uns dorthin versetzt haben, bedarf es einer unablässigen
Dabei stößt
Befassung und Auseinandersetzung mit jenem Dasein,
das geforderte Wirken aus dem Ganzen für das Ganze, das Getriebenwerden durch die innere Macht der Wahrheit, wie es echt-
und Schaffen innewohnt,
geistigem Leben
Naturtriebe der Selbsterhaltung,
zu
Kräften
einem
zusammen mit dem
hart
den die Verflechtung mit geistigen
grenzenlosen
Egoismus
eine
steigert;
völlige
Wandlung der Gesinnung wird damit unerläßlich und erweist sich
als Grundbedingung alles Geisteslebens echter Art; das hebt die
ethische
Aufgabe
hauptet
das
über
alles
Wert.
Aber zugleich
hinaus.
be-
seiner
Formgebung einen
Was im Menschen an
Geistigkeit aufstrebt,
künstlerische
eigentümlichen
andere
Wirken
mit
das hat zunächst ein rohes und seelenloses Dasein neben sich und
verbleibt
daher
leicht
in
einem Stande der Halbwirklichkeit;
erst
weit über die eigentliche Kunst
hin-
das künstlerische Bilden, das
ausreicht,
bringt die verschiedenen
wirkung, vermag in
Seiten
und Stufen
der Berührung das Innere
zu
in
Wechsel-
gestalten,
das
Kultur.
Äußere zu beseelen, das Leben
bei
249
sich
selbst
zusammenzuführen.
So gibt es keine volle Durchgeistigung des Lebens ohne die Kunst,
ohne ihr bildendes und veredelndes Wirken vermag aller Eifer
ethischen Aufschwunges es nicht vor Barbarei zu behüten.
Endlich
aber behauptet auch die Aufgabe der Lebenssteigerung ein
Zum Geistesleben gehört Unbedingtheit, Unend-
greifbares Recht.
lichkeit,
volle
unan-
Beherrschung der Wirklichkeit; der Mensch des un-
mittelbaren Daseins aber steht unter zahlreichen Bedingungen
Einschränkungen, er
ist,
Enge und Schwäche.
und
an jener Aufgabe gemessen, von kläglicher
So
bedarf
notwendig einer Steigerung
es
seiner Kraft, einer Erweiterung seines Daseins, einer
Schlummernden; ist es verwunderlich, daß
Belebung alles
dies besonderen
Epochen
das Ganze der Kultur zu bedeuten schien?
Aus solchem Nebeneinander verschiedener Lebensrichtungen
müssen schroffe Spannungen und harte Zusammenstöße erwachsen, und
zwar keineswegs durch bloßes Irren und Mißverstehen der Menschen.
Denn keine der Aufgaben läßt sich mit voller Hingebung ergreifen
und mit voller Kraft verfolgen, ohne als Selbstzweck aufzutreten und
im Augenblick des Handelns als die Hauptsache zu fühlen; so
sich
wird es begreiflich, daß im Ganzen des Menschenlebens nicht bloß
ethische, künstlerische, dynamische Antriebe wirken,
sondern daß sich
eigentümliche Kulturtypen ausbilden und um die Herrschaft kämpfen.
Abschwächungen und Kompromisse vermögen dagegen nichts, sie
Aber wenn der Kampf
drücken leicht das Niveau des Lebens herab.
nicht zu vermeiden,
ist,
ja
sein Nachlassen
nicht einmal zu
wünschen
so wird um so wünschenswerter, daß etwas dem Kampf überlegen
bleibe
und einen Kampf gegen den bloßen Kampf unternehme.
Das
aber vermag nur die Belebung eines wesenhaften Seins, das durch
alle
Verschiedenheit hindurch sich selbst eriebt, das die verschiedenen
Leistungen auf eine überlegene Einheit zurückbezieht, von da aus
von dort aus zusammenzufassen
mißt,
winnen nun
selbst
alle
Jene Bewegungen ge-
strebt.
eine Richtung auf die Entwicklung eines bei sich
befindlichen
wesenhaften
Wirklichkeit; es wird hier ein
und
Geisteslebens
Lebensraum geboten,
einer
in
geistigen
dem sie sich
begegnen und auseinandersetzen können; etwaigen Konflikten stehen
wir hier nicht wehrlos gegenüber, wir können zur Ausgleichung
wirken,
wir
können
der Festlegung
bloßer
die
Ent-
damit vor ein
Ent-
Teilkulturen
wicklung einer Gesamtkultur entgegensetzen.
Die Teilkulturen mit
ihrer
Arbeit
treten
Zu den Problemen des Menschenlebens.
250
weder— oder:
dem Ganzen
Ganzes
ein
finden
— oder lösen
sich
sie
,
immer mehr
fallen
den Zusammenhang mit der Tiefe und
sie
— nur mit der Wendung zur
Tiefe wird
Leben
das
vom Lebensgrunde ab und ver-
einer Vereinzelung?
nach der Entscheidung
Je
Dort eine
dorthin oder hierher ergibt sich ein schroffer Gegensatz.
wesenhafte, hier eine wesenlose Art der Kultur;
dort ein Insichauf-
nehmen der Erfahrungen und Schicksale des ganzen Menschen und
zugleich eine konkrete Gestaltung, hier eine freischwebende Betätig-
ung und damit eine große Vagheit; dort eine Erhebung über alles
zum
Kleinmenschliche,
mindesten ein tapferer Widerstand dagegen,
hier eine Wehrlosigkeit der Geisteskultur gegen die bloße Menschendie
Gegen-
wesenhaften Geisteswelt bloßes Gesetzes- und
Formel-
So droht
kultur.
wart
einer
die
Lebensbewegung ohne
ethische
wesen zu werden, zur Einengung und Bedrückung zu wirken, auch
Pharisäismus auszulaufen; die künstlerische
in einen selbstgerechten
Gestaltung führt, auf sich allein
das Leben
gestellt,
unvermeidlich
ins Genießende, Weichliche, Spielende, die dynamische ins Selbstische,
So hängt auch die Wahrheit der Teilkulturen daran,
Wilde, Brutale.
daß
sie eine Wesens-
und Gesamtkultur hinter sich haben, daß jene
Zurückverlegung. der Kultur erfolge, die nur durch Anknüpfung an
ein selbständiges Geistesleben möglich wird.
Was die Idee einer zugleich wesenhaften und universalen Kultur,
in
ihrem Gegensatz zur ersten Kulturlage, an Folgen und Forder-
ungen mit sich
schäftigen.
wird uns bei den nächsten Artikeln be-
bringt, das
Die Kultur wird sowohl
in ihre Mittel
in ihren
Inhalt zu verfolgen sein: dort sind die
schichte
und
Gesellschaft,
hier die
und Träger
als
Probleme von Ge-
von Kunst und Moral
in
ihren
mannigfachen Beziehungen zu erörtern; Punkt für Punkt wird sich
zeigen,
daß
bei
eine neue Sache
jener
Idee
nicht
und Aufgabe
in
bloß
ein
Frage steht
neues Wort,
Hier
sei
sondern
nur noch
das eine erwähnt, daß die gegenwärtige Lage die Forderung einer
Zurückverlegung
und
dringlich macht.
Diese Lage
treffen
zweier
festeren
Tatsachen
ist
kritisch
Begründung
der
Kultur
namentlich durch ein
geworden.
überaus
Zusammen-
Einmal sind die ge-
schichtlich überkommenen Grundlagen und Inhalte der Kultur, soweit
sie
das Ganze und Innere des Menschen betreffen, sehr ins Unsichere
sind das vornehmlich deshalb,
geraten;
sie
Art
als
zu anthropomorph, zu kleinmenschlich empfinden und aus
solcher
Empfindung in Zweifel kommen, ob der Mensch überhaupt
weil wir jetzt die ältere
Kultur.
251
das sinnlich -natürliche Dasein irgend
ob alles,
überschreiten kann,
was er an Mehralsmenschlichem zu erfassen glaubte, nicht ein bloßes
Trugbild, ein Erzeugnis menschlichen
tief
in das
der Welt
Leben
ein,
dem
das eine mit
fällt
Das greift sehr
weit tiefer als diejenigen meinen, welche aus
alle Geistigkeit
entfernen
und zugleich dem Menschen
eine
Denn in Wahrheit
und
wahren zu können wähnen.
Idealität
Wahnes sei.
steht
Punkt
andern; es läßt sich unmöglich im
und Subjekt erhalten, was für das Ganze und im Wesen aufgegeben
wird.
So sind wir aller unserer Ideale, ja unseres eigenen Wesens
geworden,
unsicher
nicht
mehr schöpfen wir aus einem gemein-
samen Grundstock von Überzeugungen zusammenhaltende,
erhöhende Kräfte; bei
Verfall
des
aller
subjektiven Regsamkeit
ist
richtende,
ein innerer
Lebens unvermeidlich, wenn jene Erschütterung weiter
und weiter greift.
Und in diese wankende und schwankende Zeit fällt hinein das
Drängen der Massen nach vollem Teilhaben an Kultur
und Glück, samt dem Anspruch, über das, was an der Kultur ge-
stürmische
und
halt-
wertvoll
scheiden nach
kraft
sei,
dem
mit eignem Urteil zu
entscheiden,
zu ent-
unmittelbaren Eindruck und nach der Fassungs-
der Individuen, die von
den weligeschichtlichen Bewegungen
und Erfahrungen der Menschheit kaum irgend berührt worden sind.
Nun macht jene innere Unsicherheit der bestehenden Kulturkomplexe,
im
besondern
ihre
schwere Belastung
und Überunerschütterliche Wahr-
mit Veraltetem
lebtem, sie unfähig, solchem Verlangen eine
und es damit in sichere Bahnen zu leiten; so
Bewegung alles fortzureißen, wie sie schon jetzt vergröbernd und verflachend, verengend und verneinend wirkt. Über
heit entgegenzuhalten
droht
jene
eine solche Krise kann schlechterdings
als ein
nichts
anderes hinausführen
neuer Aufschwung des Lebens, eine Vertiefung des Geistes-
lebens in sich selbst, die Entdeckung innerer Tatsachen
Zusammenhänge.
und innerer
Von draußen kann uns das Heil nicht kommen;
was an Stützen und Hülfen dort unwiderbringlich verloren ging,
das können wir nur durch eine Verstärkung des
Innern
ersetzen,
nur dadurch, daß wir bei uns selbst zu einer überlegenen Welt gelangen, uns darin befestigen,
von daher unserem Leben einen Inhalt
geben,
von daher eine neue Kultur erbauen.
tiefung
und Befestigung, so kann
die
Gelingt solche Ver-
bedrohliche Krise zu
Erneuerung und Verjüngung des Lebens führen
und durch
einer
alle
menschliche Irrung hindurch dem Dasein einen größeren Wahrheits-
252
Zu den Problemen des Menschenlebens.
gehalt verleihen.
Besteht dagegen
keine Möglichkeit einer solchen
Vertiefung und eines Hervorbrechens ursprünglicher Kräfte,
ist
im
menschlichen Dasein keine wesenhafte Geisteswelt neu zu beleben,
so
entfällt
alle
Hoffnung eines glücklichen Ausgangs,
so
müssen
der menschlichen Selbstsucht und Leidenschaft Vernunft und Kultur
unterliegen.
2.
Zur Entwicklung des Problems.
a)
I
Geschichte.
|as Verhältnis des heutigen Menschen zur Geschichte
ist
voller
-^ Verworrenheit: wir hängen an der Geschichte und zehren von
aber zugleich fühlen wir unser Leben
der Geschichte,
aufs stärkste bedrückt
durch
sie
und möchten uns dieser Last entledigen;
in-
dem wir aber das unternehmen, drohen wir der Leere des bloßen
und
Augenblicks zu verfallen,
flüchten
einen und
Handeln
solcher
Gefahr doch
dem anderen hin und her, eine Lage, die zielbewußtes
und glückliches Schaffen unmöglich gedeihen läßt. Bewas
trachten wir etwas näher,
Das
vor
So schwanken wir zwischen dem
wieder zur Geschichte zurück.
19.
Lage führte.
in eine solche
Jahrhundert wird
in
seinem Verhältnis zur Geschichte
vom Gegensatz zur Aufklärung mit ihrem Rationalismus beherrscht.
Aus verworrenen Verhältnissen hatte die moderne Menschheit einen
Ausweg durch ein Zurückgehen auf eine allen innewohnende Vernunft gesucht; nur ihre kräftige Belebung schien das menschliche Dasein
von
Veraltetem
und
Irrigem
gründlich
sowie
befreien,
das
Leben von kindlicher Befangenheit und dumpfer Gebundenheit zu
voller Mündigkeit
und Klarheit erheben zu können.
Die Vergangen-
heit mit ihrer Autorität versank gegenüber der Forderung, das Leben
und Wirken
in
eine
zeitlose
Gegenwart des Denkens zu
unbeirrt durch die Überlieferung, meist in
ihr,
schuf die Vernunft eine »natürliche" Religion, eine
Moral,
«natürliches"
ein
Wirtschaftsleben,
stellen;
bewußtem Gegensatz zu
eine
;,
natürliche"
«naturgemäße« Er-
Gemüter überwältigend fortgerissen und in
die Gestaltung des Lebens tief gewirkt; was es daraus an Frische,
ziehung.
Freiheit
Das
hat die
und Selbständigkeit gewann, das
konnte
trotz
fehdung und Verdunklung nicht wieder verloren gehen.
Anfang
an
trug
jenes
Streben
Problematisches
in
sich,
aller
Be-
Aber von
das
im
Zu den Problemen des Menschenlebens.
254
Laufe der Zeiten wuchs und endlich einen
Das
jugendliche Kraftgefühl, mit
Rückschlag hervorrief.
dem die Aufklärung einsetzte, gab
ihr die freudige Zuversicht, einer absoluten
den überkommenen Verhältnissen konnte
Wahrheit nahe zu
sie
sein;
siegesgewiß
so
nicht
entgegentreten ohne ein festes Vertrauen auf das unmittelbare Walten
und
der Wirklichkeit
im
Menschenwesen.
So
einer Vernunft
in
schien denn
in
jedem Einzelnen die Vernunft angelegt und durch
eine
Selbstbesinnung leicht erreichbar;
kräftige
Erhebung zu
Wahre zur
voller Bewußtheit dünkte
Herrschaft zu
eine Klärung,
Damit ward
bringen.
vornehmlich auf Denken und Erkennen
gestellt,
eine
um das Gute und
genügend,
die
Lebensarbeit
die Kultur
erhielt
einen einseitig intellektuellen Charakter; beim Nachlassen des ersten
Aufschwungs zog
Räsonnement das Leben
verstandesmäßiges
ein
mehr und mehr an
sich,
zwischen den Menschen und die
stellte
Dinge sein Erwägen und seine Zwecke und gefährdete damit immer
einen
stärker
inneren
Zusammenhang
ward
schließlich als
drang schlug
bildet die
und eine Un-
zu eng und seelenlos empfunden, der Lebens-
um und verlangte mehr Inhalt sowie mehr Betätigung
ganzen Menschen;
des
mit der Welt
Die Wirklichkeit, die daraus hervorging,
mittelbarkeit des Lebens.
Hauptstück dieses neuen Lebens aber
ein
Wendung zur Geschichte.
Denn was zu
ein Durst nach mehr
Grunde des Daseins, nach mehr
Anschauung und mehr Lebensfülle, auch nach mehr Verbindung der
Mannigfaltigkeit zu großen Zusammenhängen.
Wie viel gesättigter
ihr
trieb,
war vor allem
Wirklichkeit, nach einem breiteren
das Leben
und
dadurch ward, das zeigen
Religion,
die sonst ungenutzt verblieb,
lichkeit,
einzelnen Gebiete,
alle
Kunst und Wissenschaft; unendlich
ist
hier
viel
Recht
mehr Wirk-
dem eignen Tun ver-
Das Ganze der Arbeit erzeugt eine historische Denkweise
und verändert damit den Charakter des Lebens. Es reißt sich hier
bunden.
nicht der Mensch, wie in der Aufklärungszeit,
los
und stellt sich
zu
beherrschen,
damit ihr Leben
freie.
ihr schroff gegenüber,
sondern
in
er
ersehnt
ihn überströme
von seiner Umgebung
um sie wie etwas Fremdes
eine
innere
Einheit
mit
ihr,
und ihn von aller Kleinheit be-
Damit gewinnt sein Dasein wie mehr Weite, so auch mehr
Ruhe, aus den Dingen wächst dem Menschen eine Vernunft entgegen,
deren Führung
er
getrost
sich
anvertrauen darf.
auch die früheren Zeiten nahe und
Verwandtschaft finden;
läßt in
Das
rückt
ihm
ihnen die mannigfachste
die eigne Zeit erscheint als die Spitze eines
Geschichte.
255
Gesamtbaus, der alle Zeiten umfaßt; von solcher Spitze aus erscheint
Frühere
alles
als ein
wird
Niederen
allmähliches Ansteigen zur Höhe, auch in
dem
sowohl der Abstand und Gegensatz,
die
nicht
als
Annäherung und Vorbereitung gesehen; es kann mehr Verständnis
und mehr Liebe finden, wenn die Schroffheit einer absoluten
Schätzung, wie
sie
ung eigen war,
dem Reformationszeitalter und auch der Aufklär-
einer universaleren
Keine Zeit hat solchen
weicht.
und versöhnlicheren Denkweise
Umschwung der Behandlung stärker
erfahren als das Mittelalter.
Diese mehr relative Behandlung besagte aber zu Anfang keines-
soluten Wahrheit.
zum Relativismus und eine Preisgebung einer abDenn ein stolzes Selbstbewußtsein ließ die geistige
Kraft sich allem
Zustrom des
wegs
ein
Sinken
Stoffes
gewachsen und seiner Assimi-
Die Vernunft zog, wenigstens
lierung fähig fühlen.
in
der Denk-
weise der Philosophen, unter eigner innerer Erweiterung weit mehr
die
an
Geschichte
sich,
als
daß
sie
sich
ihr
unterworfen
hätte.
Diese Denkweise hat den großartigsten und durchgebildetsten Aus-
druck
in
der Geschichtsphilosophie Hegels gefunden;
zwischen Vernunft
indem
und Geschichte
scheint
Spannung
überwunden,
und gar zur Entwicklung der Vernunft
die Geschichte ganz
wird, diese aber in solcher Entwicklung ihr
Welche Bedenken
alle
glücklich
Konstruktion
diese
Wesen findet.
der Geschichte
erregen
mag, die Überlegenheit der Vernunft und damit die geistige Aktivität
wurde dabei
kräftig
gewahrt.
Minder besorgt war dafür eine Be-
handlung der Geschichte aus der Denkweise der Romantik.
Denn
hier schien ein
unbewußtes Walten und Weben die Bewegung her-
vorzubringen,
aus
der
Vergangenheit schien
dem Menschen
ein
Strom der Vernunft mühelos zuzufließen und ihn sicher mit fortzutragen; mit der Hingebung an diesen Strom schien sein Leben und
Das schwächte die Aktivität
in sichere Bahnen geleitet.
und verkümmerte das Recht der lebendigen Gegenwart; indem man
sich in vergangene Zeiten einlebte und diese dabei idealisierte, verSchon hier
schloß man sich leicht den Aufgaben der eignen Zeit.
Streben
erscheint die Gefahr, daß der Erweiterung des Gesichtskreises durch
die Geschichte nicht die Kraft des Zusammenhaltens
entspricht,
und daher der Mensch
seines Lebens
Schaden
Dann kam
von
den
die
Problemen
bei
und Aneignens
äußerem Gewinn im
Kern
erleidet.
dem
des
19. Jahrhundert
inneren
eigentümliche
Wendung
und
geistigen
Menschen
des
Zu den Problemen des Menschenlebens.
256
Schaffens zur Arbeit mit ihrer Richtung auf das Gegenständliche der
Dinge; bei der Geschichte besagte das ein
der
exakten
Forschung gegenüber
Gesamtbildern.
rissenen
Wendung
Diese
Deutschland gewonnen, da
wußtheit in
siegreiches Vordringen
Konstruktion
der
hat
sie
eine
in
nur um-
besondere Be-
hier gegen
Über-
das
wiegen einer spekulativen Behandlung ihr Recht erst erstreiten mußte
Namentlich erhob sich gegen die Hegeische Konstruktion der Geschichte
erschien
Jene
nach,
kreise
nach mehr Weite,
ein Verlangen
vidualität.
indem
ihre
Tatsächlichkeit
und
Indi-
zu
eng schon dem äußeren Gesichts-
Begriffe
im Grunde nur die europäische
als
und sich namentlich um den Gegensatz von
Kulturwelt umspannten
Altertum und Neuzeit bewegten;
sie
litt
aber auch an einer inneren
Enge, indem sie die Individualität und Positivität der einzelnen Er-
scheinungen aufs stärkste abschwächen mußte,
tischen
Gefüge einordnen zu können.
um sie ihrem dialek-
Das neue Verlangen
nach
reiner und unbegrenzter Tatsächlichkeit sah darin eine Vergewaltigung
und eine Verfälschung der Dinge; mit um so glühenderem Eifer
wurde die historische Forschung als eine Befreiung davon ergriffen.
Diese Forschung hat jenes Verlangen nach Weite und Tatsächlichkeit in vorzüglicher Weise in Arbeit und Leistung umzusetzen
verstanden, sie hat für diese Arbeit neue
hat
Methoden ausgebildet,
sie
durch Inhalt und Form eine eigentümliche Denkweise erzeugt
und wirkt damit
stark
auf das Leben der Neuzeit.
Diese Forsch-
ung will keineswegs Philosophie sein, geht doch ihr Hauptanliegen
dahin, die Geschichte von aller Bevormundung durch die Philosophie zu befreien und allein auf die eigne Kraft zu stellen; aber
die Arbeit hätte unmöglich so siegreich vordringen und die Hingebung des ganzen Menschen gewinnen können, ohne bestimmte
Überzeugungen in sich zu tragen und anzuregen. Die Forschung
kann nicht das Verlangen nach einem reinen Tatbestande entwickeln
und verfechten, ohne zu gewahren, wie viel zwischen dem Menschen
und jenem Tatbestande
der Überlieferung
ein
energischer
nommen,
heit
als
liegt,
ohne
vielfacher
Subjektivität
sowohl
der eignen Auffassung innezuwerden; so wird
Kampf zur Austreibung
Leben
sein
Gelingen
läßt
das
gewinnen.
Indem
jenes
Streben
Subjektivität
unter-
mehr Ruhe und
Klar-
dieser
nach
Tatsächlichkeit
eine
grenzenlose Fülle individueller Bildungen aufdeckt, und indem sich
ihm der Lauf der Zeiten
mehr an Einen Faden reihen läßt,
Gewebe durcheinanderlaufender, kaum
nicht
vielmehr ein unermeßliches
Geschichte.
Fäden
entwirrbarer
kann
wird,
ersichtlich
257
Unvermögen des
das
von innen her zu durchschauen und in einfache
Menschen, das alles
mehr zweifelhaft sein; so wird ihm eine
geboten, nicht mehr darf er von sich
aus die Tatsachen zurechtlegen und abrunden wollen. Aber indem
Begriffe umzusetzen, nicht
bescheidene Zurückhaltung
herrschen zu dienen
er statt zu
meßliche Bereicherung,
alter
Enge.
Gewinn und ohne VerAber Verwicklungen erschienen gar bald, und aus dem
Das
zunächst ein
schien
alles
wicklung.
Gewinn des Wissens drohte
Die Objektivität,
sagt
Befreiung von
gründliche
eine
Leben eine uner-
erfährt sein
hat,
Beziehung zum Subjekt,
werden.
keineswegs einfach.
der Dinge nur das, was
ohne
sie
Bealle
Wirken des Denkens sind, so
ihnen zu verzichten, denn dieses ist einmal
ohne
wäre auf alles Innere bei
nicht zu fassen
ist
Lebens zu
des
Verlust
gefordert wird,
die
reine Tatbestand
der
ein
reiner
alles
ohne ein Aufgebot eignen Denkens, ohne
ein Nach-
Auch eine Scheidung von Großem und
Kleinem, von Wesentlichem und Nebensächlichem in der Geschichte
die aus dem Ganzen einer
ist nicht wohl möglich ohne Maßstäbe,
erleben
und Miterleben.
Überzeugung hervorgehen müssen.^
Eine
Geschichte
aber ohne
und ohne Abstufung müßte ein chaotisches Neben- und
Durcheinander werden, das kaum mehr Wissenschaft heißen könnte.
Innerlichkeit
Wie wenig die Geschichte, bei aller Abweisung der Philosophie, gewisser Grundüberzeugungen entbehren kann, das bekundet mit voller
immer stärker wogende Streit über den
Aber woher
Überzeugungen nehmen, nachdem alle Philosophie verneint
Deutlichkeit der neuerdings
Hauptinhalt und die bewegenden Kräfte der Geschichte.
diese
ist?
Die Zeit hat sich
Einmal
beholfen.
und abgeschwächt,
ein
zwiefacher Weise geholfen oder vielmehr
in
erhält
sich
unverkennbar,
als Ganzes aufs schroffste abgelehnt wurde.
aber
führende
innere
Stellung
unbedenklich
fest.
Notwendigkeit des
der
Es
Intelligenz
ist
das
bei
ein
scheinung im Leben der Gegenwart.
*
S.
versteckt
die
Hegel wird perhorresziert,
Innewohnen der Vernunft
irgendwelches
irgendwelche
wenn auch
Einfluß derselben spekulativen Denkart,
in
der
Fortschritts,
Geschichte,
irgendwelche
diesem Prozesse
hält
Stück einer allgemeineren
man
Er-
Die pantheistische Denkweise,
darüber die vortrefflichen, besonnen abwägenden und selbständig
Werke von Arvid Grotenfelt „Die Wertschätzung in der Ge-
urteilenden
schichte" 1Q03,
und „Geschichtliche Wertmaßstäbe
in
sophie bei Historikern und im Volksbewußtsein", 1905.
Eucken, Grundbegriffe. 4. Aufl.
der Geschichtsphilo-
17
den Problemen des MenschenleDens.
2[u
258
Bewegung der Neuzeit hervorging, und die früher eine
die aus der
feste
Überzeugung, auch eine freudige Lebenstimmung hinter
hatte,
erhält sich in
Grundlage
unsicher
wurde.
wie
Begriffe
und
Humanität u. s.w. verbleiben
schritt,
digen
wurden
Kräften
und
blaß
alles
Vernunft,
Geist,
schattenhaft
geworden, aus leben-
aus fruchtbaren Ideen
Schattenbilder,
um so unwahrer werden,
Das Ganze muß
Fort-
dem Denken ge-
liefern
Nur ist nach Erschütterung
wisse Richtungen wie Wertschätzungen.
jener Grundlage
sich
mannigfachen Wirkungen, auch nachdem ihre
leere
mehr
Grundwelche
jener
gewinnen,
Seele
solche Überzeugungen unsere
überzeugung schnurstracks widersprechen. Der kräftigste Vertreter
Redensarten.
Widerspruchs
solches
Pessimismus,
der
ist
immer
19. Jahrhunderts
weiter
im
der
je
des
Verlauf
um sich gegriffen hat.
Indem er
das Dunkle und Unvernünftige unserer Welt und mit ihr der Ge-
Empfindung bringt,
schichte voll zur
unbarmherzig den
zerstört er
verklärenden Glanz, den der Pantheismus
dem Dasein' verlieh; viel
zu energisch hat er uns allen neue Gruppen von Tatsachen vor die
Augen gerückt, neue Durchblicke des Ganzen eröffnet,
Glaube an
alte
als
daß der
die Vernunft unserer Wirklichkeit schlechthin
Ein merkwürdiger Widerspruch
bleiben könnte.
ver-
liegt hier vor: die
Stimmung der Menschheit verdüstert sich, Menschen wie Schicksale machen einen trüberen Gesamteindruck, die Widersprüche des
Daseins
fallen
der Zeit jene
Dinge
fest,
um
Halt,
sie
uns
grell
ins
pantheistische
Aber zugleich
Auge.
hält die Arbeit
Denkweise mit ihrer Idealisierung der
klammert sich daran wie an den einzig möglichen
Der Pessimismus
nicht völliger Nichtigkeit zu verfallen.
und der Optimismus hier gewähren ein Beispiel für jene Entzweiung von Seele und Arbeit, für jene Zerklüftung des ganzen
Menschen, woran das moderne Leben leidet
Aber es gibt noch eine andere Weise, in der die Gegenwart
dort
des
sich
Problemes erwehrt:
sie
setzt
den Zeiten gar keine be-
stimmte Denkweise entgegen, sondern sucht ein Urteil über
ein
Maß
selbst
jene
für sie
lediglich
zu entwickeln und zu erweisen; so ganz möchte
versenken
und
daß
einfühlen,
sie
lediglich
Denkweise verstanden und gewürdigt werden.
viel
Bedeutendes
geleistet;
geschickt,
fremden
innerstes
Wollen
Zeiten
und
sie
aus ihnen selbst zu gewinnen, an ihnen
nie
abzulauschen,
volles
Recht
ihnen einen
in
eignen
In dieser Richtung ist
war eine Zeit so
ihr
aus
sie sich
ihrer
zu
bereit
und auch so
geben,
ihnen
Zusammenhang
ihr
nicht
Geschichte.
von draußen
aufzudrängen,
her
259
sondern
ihn
ihrem
aus
eignen
Wirken und Wollen hervorzulocken, dabei mit gleicher Liebe
in das Verschiedenste und Widersprechendste hineinzuversetzen.
wir
in
Wahrheit
bei
ob
trotz
äußern,
nicht
tivität
unsere
Zeiten
besser urteilen
solchem Streben
Abwehr
aller
Subjektivität
die
in
Ob
uns
eignen Art
ent-
vermeintliche
Objek-
werden
spätere
darüber
miteinfließt,
wir
als
aller
sich
Eine Gefahr aber und eine
selbst.
Schädigung erfahren wir selber deutlich genug:
es ist die Abschwäch-
ung unseres eignen Wollens und Wesens durch jene Beflissenheit,
fremder Art uns anzuschmiegen und anzupassen.
Indem bei der
unermeßlichen Erweiterung des Horizontes Mannigfachstes und Verauf uns
schiedenartigstes
verleiht,
eindringt,
uns bewältigt und seine Farbe
wird unsere Seele zu einer Bühne, auf der
alle
möglichen
Personen auftreten und ihre Rolle. hersagen; wir vergessen, daß die
Ausdehnung unseres Vorstellungskreises keineswegs schon
ein Weit-
werden unseres Lebens bedeutet, und neigen dahin, gelehrtes Wissen
für geistiges Leben einzusetzen; wir geraten in Gefahr, über jenem
Mitleben, das schließlich
immer ein Halbleben bleibt,
ein eignes und
Leben, ein klares Denken und festes Wollen mehr und mehr
volles
einzubüßen; die geistige Synthese, die wir an allen früheren Zeiten
aufzusuchen bemüht sind, vermögen wir an
der eignen
nicht
zu
vollziehen.
Am
meisten
wird unsere Schwäche bei
ersichtlich
fremdem Streben zu unserem eignen zu
heit
uns sicher zu Hause,
wir
fühlen
gekommen
dem
Ver-
Übergang von der Vergangenheit zur Gegenwart, von
suche, einen
In der Vergangen-
finden.
wir durchschauen,
wir verstehen, wie das eine aus
ist,
wie
alles
dem anderen ent-
sprang, entspringen mußte, wir verfolgen solche Betrachtung bis an
die Schwelle
die
sich
der Gegenwart;
Verbindung
uns
mitteilen
würdig genug
verbleibt,
ist
hergestellt,
und
in
nur ein einziger kleiner
eignes Leben verwandeln.
— dieser kleine
Schritt,
der Ertrag der langen Arbeit
und
kann
Aber merk-
Schritt will nicht gelingen, die Kluft
Wissen und Leben kommen nicht zusammen. Ja der FortWissens hemmt die Verbindung der Geschichte
schritt des historischen
mit
dem
Leben.
Denn
je
deutlicher
die
Wissenschaft die Eigen-
tümlichkeit fremder Zeiten herausstellt, desto
mehr zeigt sie die Ge-
bundenheit ihres Gehalts an besondere Lagen, desto schärfer zieht
sie
ihre
Grenzen
gegen
andere
Zeiten
und
Denkweisen,
desto
zwingender verbietet sie ein einfaches Überströmen fremden Lebens
17»
Zu den Problemen des Menschenlebens.
260
auch darin
Diese Spannung zwischen Wissen und Leben kommt
zum Ausdruck, daß die Forschung besondere Liebe weit-
entlegenen
Zeiten
in das eigne.
Denn
feiert
Leben
eignen
und dort
spendet
steht;
mehr wir uns aber der Gegenwart nähern,
je
unabweisbarer wird diese Frage und desto peinlicher unsere
desto
Nirgends
Unsicherheit.
solche Kluft zwischen Wissen und Leben,
ist
Lebens und
Vorbedingungen geistigen
zwischen
Religionsforschung
wie
fortgeschritten,
Leben
geistigem
Wie weit ist heute
selbst schroffer als auf dem Gebiete der Religion.
die
Triumphe
glänzendsten
die
kommt minder zur Sprache, wie die Sache zum
dort
stehen
präziser
viel
namentlich die großen Religionen und innerhalb ihrer die einzelnen
Phasen
Augen,
unseren
vor
wie
über dieser Tatsächlichkeit,
unsere
eignen
Leben,
wie
bleiben,
wie
ist
hier
solange
wir
nicht
Fülle
eine
hier
wenig gewinnen wir aus
Überzeugungen,
religiösen
groß
uns
umflutet
Aber wie wehrlos sind wir gegen-
anschaulicher Tatsächlichkeit!
unser
eignes
Und
unsere Hilflosigkeit!
die
zu
Kraft
eigner
ihr
für
religiöses
wird es
sie
Gestaltung
des
Lebens finden; daß wir sie aber finden, daran hindert uns vor allem
indem
Geschichte,
die
indem
haften läßt,
sie
uns
am
bloßen
Schein
Besitzes
eines
uns durch unablässige Befassung mit fremden
sie
Dingen eignen Denkens und eigner Verantwortung entwöhnt, indem
sie
gelehrtes
Kein
Wissen für Leben
Wunder
ist
schaftliche Bewegung
wieder
ein
es daher,
einsetzt.
daß von
Zeit zu Zeit eine leiden-
gegen die Geschichte aufwallt, daß auch heute
Zorn gegen den entnervenden Historismus mit
Verstrickung in ein Halbleben
um sich greift.
Vergangenheit von euch und
stellt
Gegenwart; dann wird es wieder
seiner
„Werft die Bürde der
das Leben ganz und gar in die
und wahrhaftig, dann erst
frisch
wird es euer eignes Leben werden."
Ja,
wenn solches Abschütteln
so einfach wäre und nicht mit der erstrebten Befreiung vieles verloren ginge,
auf das sich nicht wohl
hält die Geschichte
uns weit fester
uns fest auch gegen unseren Willen.
ein
Erzeugnis
einer
geschichtlichen
eigentümliche Färbung, ihr Nein
verzichten
als jene
trifft
auch
empfunden,
sobald
hat
Wie
rasch
Lage
und
erhält
B.
die
sie
hält
daraus eine
Solche geschichtliche
Bewegungen wird
der Verlauf der Dinge über
z.
Wahrheit
besondere Schäden, und auch
geschichtsfeindlicher
ist
In
Denn die Opposition ist selbst
ihr Ja trägt das Gewand der besonderen Zeit.
Bedingtheit
läßt!
Gegner meinen,
Aufklärung,
die
deutlich
sie
hinausgeführt
alle
geschichtliche
Geschichte.
Bindung
abstreifen
gestalten
wollte,
und das Leben gänzlich aus zeitloser Vernunft
selbst
zu einer geschichtlichen Größe, einer histo-
geworden, wie
rischen Kategorie
261
viel
berührt uns
ihr
in
wie
jetzt
Wer das Ganze
eine ferne Vergangenheit, ein vergilbtes Dokument!
der Geschichte überschaut, dürfte eine Art Wellenbewegung zwischen
Anschluß und Verwerfung gewahren und zugleich sich überzeugen,
daß
Verneinung genau
die
gut zur Geschichte gehört wie die
so
Bejahung, daß der leidenschaftliche Ansturm gegen die Geschichte
mit seiner Neigung, das Gegenteil des Vorgefundenen zu behaupten,
weniger eine
echte
Abhängigkeit
verändert.
Unterschied,
ob das bewußte Streben des Menschen mit der Ge-
erheblichen
Beim Widerspruch wird das
sie geht.
aus
lediglich
als
Empfinden jedes Einzelnen
der
unmittelbaren
überzeugend
Aber wird
dartut.
das Leben nicht eng und arm?
solcher Absteckung
zur Oberfläche gedrängt und
legt,
nur die Art der
einen
es
Gegenwart zu
wahr nur anzuerkennen, was sich dem Denken und
gehießen,
schöpfen und
als
- Immerhin macht
oder ob es gegen
schichte,
Leben
Unabhängigkeit gewährt,
Erscheinungen zer-
lauter einzelne
in
bei
Wird es nicht
wenn das dem bloßen Individuum Gegenwärtige als Maß alier
Dinge
dabei
Leidet
gilt?
die
liefert
gewachsenen Geisteswelt
diejenigen
unter ihren Denkern,
legenheit
des Geisteslebens
Ausbau
ihrer
ist
sie nicht gelangt;
die Natur
verfochten,
begriffen
erlegen,
gewiß mit aus dem Grunde,
mit ihrem
reichen
vertiefenden
Gegen
die
mächtig
sind
im
Gedankenwelt immer wieder dem Einfluß von Natur-
schichte
ihren
auch
mit größtem Eifer die Über-
die
gegen
Die Aufklärung
Denn zu einer festgegrün-
ein anschauliches Beispiel davon.
deten, der Natur
der geistige
innere Selbständigkeit,
Charakter des Lebens nicht schwersten Schaden?
Erfahrungen
unbegrenzte
umfängt,
Gehalt,
scheint
ihren
glaubten
uns
Welt,
die
das
innere
festen
weil sie die GeZusammenhängen,
geringachten
zu
von
her
draußen
dürfen.
über-
Leben nicht aufkommen zu
können, wenn es nicht selbst einen Zusammenschluß übersubjektiver
Art gewinnt; dazu aber bedarf es notwendig der Geschichte.
ferner
wart zu
stellen,
Muß
das Leben
ganz und gar in die Gegen-
sich selbst zerstören,
da die Gegenwart unablässig
nicht der Versuch,
und
Ganze in ein Nichts zu verflattern und verwehen
eine andere wird, das Heute alsbald in ein Gestern umschlägt,
so schließlich das
droht?
Vor dem Äußersten behütet uns freilich der Umstand, daß,
wie sich
zeigte,
die
Geschichte den Menschen auch
gegen seinen
Zu den Problemen des Menschenlebens.
262
Willen
festhält;
aber steht es dann
nicht so,
daß,
soweit wir uns
der Geschichte entledigen, wir das Leben verflüchtigen, soweit es
uns aber einen Gehalt bewahrt, wir dieselbe Geschichte widerwillig
bejahen,
Abschüttelung
deren
zur
Kraft
und Wahrhaftigkeit
des
Lebens unerläßlich schien?
So befinden wir uns in einer höchst verworrenen Lage, ja einem
unerträglichen
Dilemma: wir können die Geschichte weder festhalten
noch entbehren; wir geraten ins Leere, wo wir sie abschütteln, wir ver-
einem Schattenleben, wo wir uns ihr unterwerfen.
fallen
schnittsart
Die Durch-
mag sich demgegenüber mit Kompromissen behelfen und
an einem Mittelding von Freiheit und Knechtschaft Gefallen finden, eine
energischere Denkweise wird die Unmöglichkeit eines Kompromisses
durchschauen und eine Überwindung des Gegensatzes verlangen.
Ist
von der Geschichte möglich, die zugleich eine
aber eine Befreiung
Versöhnung mit der Geschichte bedeutet, kann das Leben eine Übergeschichtlichkeit erreichen und zugleich der Geschichte einen Wert belassen,
ist
ein Lebenstypus denkbar,
der nicht haltlos zwischen
dem
Rationalismus des 18. und dem Historismus des 19. Jahrhunderts hin-
undherschwankt, sondern
Ausbildung einer selbständigen Art das
in
Recht eines jeden anzuerkennen und zugleich zu begrenzen vermag?
Ohne eingreifende Umwandlungen des ersten Anblicks und energische
Fortbildungen des Lebens
wir,
ist
das sicherlich nicht zu erreichen; sehen
ob der Zusammenhang unserer Untersuchung Anhaltspunkte
dafür gewährt. 1
b)
Forderungen und Ausblicke.
Die nächste Frage
Geschichte
irgend
zu
ist
die,
ob das menschliche Leben
entwinden
und
ihr
zutreten vermag; die Beantwortung dieser Frage aber
^
sich der
gegenüber-
selbständig
hängt daran.
Alle Erörterung hier soll sich auf diesen einen Hauptpunkt beschränken,
für das Weitere sei auf
meine Darstellung der Philosophie der Geschichte in
der „Kultur der G^enwart" (Band: „Systematische Philosophie") verwiesen.
Das Auftauchen mannigfacher Streitfragen und die Leidenschaft, welche ihre
Behandlung hervorruft, bekundet jedenfalls deutlich, daß uns der Inhalt der
Geschichte wie unser Verhältnis zu ihr wieder unsicher geworden ist; wie
hätte
sonst
in
emporsteigen
abgetan galt?
neuester Zeit
können,
die Geschichtsphilosophie wieder
die vor
kurzem noch den meisten
als
so mächtig
erledigt
und
Geschichte.
263
wie über das Ganze des menschlichen Lebens gedacht wird,
notwendig ein Bekenntnis
hält
vom Kern
der Mensch ganz und gar zur Natur
tut,
steht
Überschritte er
sie ent-
Gehört
zum guten Teil
er es
,
zu einem
nie
eignen
Leben
gelangen.
ferner die Natur nur durch einzelne Eigenschaften,
Ganzen
die nicht im
— daß
Ganzen.
— so bleibt er unrettbar dem Strom der Zeit
außer Frage
und kann daraus
verfallen
dieses
und Seins gegründet wären, so
eines Lebens
käme er vielleicht zu irgendwelchem Weiterstreben, aber nie zu einer
wahrhaftigen Befreiung von der Zeit.
gewährt
das
und
Bestehen
Die einzige Möglichkeit dessen
Anerkennung
die
einer
selbständigen
den Hauptvorwurf unserer ganzen Untersuchung
Geisteswelt, wie sie
bildet.
Denn wie schon beim Problem der Entwicklung ersichtlich
wurde,
ist
die
Erhebung über
dem
Ordnung
loser
Durchgängig wird
die Zeit
Geistesleben
hier
dem
und
ein
Streben
Wirken aus zeit-
und
wesentlich
unentbehrlich.
die Richtung auf ein zeitlos
Gültiges gegeben, nie kann hier die Wirkung und Anerkennung auf
dem Boden der Geschichte eine Wahrheit und ein Recht begründen,
sondern die Wahrheit
lichen
will
Leben her dargetan
die Vergangenheit die
hier unmittelbar,
So kann
sein.
von einem ursprüngin
diesem Gebiete nie
Gegenwart ersetzen, und
Gestern wie eine Frucht der Blüte entwachsen.
Heute dem
Denn was frühere
nie das
Zeiten an geistigem Leben erzeugten, das besteht keineswegs dadurch
fort,
daß
es einmal
da war; es
gilt
hier nicht das Beharrungsgesetz
der Natur, wonach jedes Ding den vorhandenen Zustand einhält, bis
es von außen her darin verändert wird.
Ordnung, daß
von neuem
zugleich,
in
daß
Vielmehr gilt hier die andere
und immer weiter sinkt, was nicht immer
eignes Leben und Tun verwandelt wird. Das besagt
sofort sinkt
alles Geistesleben
aus unmittelbarer Gegenwart hervor-
gehen muß, daß jede Verdunklung dessen eine Abschwächung seines
Auch innerhalb der menschunterscheidenden Charakters bewirkt.
lichen Erfahrung
ist
über die Gegenwart
deutlich genug, daß weniger die Vergangenheit
als
diese über jene entscheidet, daß sich
dem-
nach mit der geistigen Art der Gegenwart das Bild der Vergangenheit unablässig verschiebt.
Wie verschiedenes wurde am klassischen
nachdem, was das eigne Leben
Altertum gesehen und geschätzt,
je
an Aufgaben und Bedürfnissen
enthielt.
Die Scholastik
ihm eine weltliche Kultur zur Ergänzung einer
suchte in
religiösen
Lebens-
ordnung, die Renaissance eine Unterstützung ihres Verlangens nach
Leben und Schönheit, die Aufklärung schätzte an ihm, soweit sie es
Zu den Problemen des Menschenlebens.
264
schätzte,
die Klarheit
flüchtete
sich
und Nützlichkeit/ der deutsche Humanismus
zu ihm aus der Verwicklung modernen Lebens
So erschloß das Alter-
zu einer lauteren, einfachen, großen Natur.
tum jedem verschiedene Seiten, aber
es
als
gab und gibt auch
viele,
denen es inmitten aller Emsigkeit gelehrter Beschäftigung geistig gar
nichts erschließen kann,
nichts erschließt,
weil sie
Leben entgegenbringen.
Daran also
entscheidende Hauptsache
immer der Besitz
anders vollziehen
geistige
bildet sein,
auch
uns nicht zu,
fällt
die
bleibt
einer Gegenwart,
einer
sie
will
selbst.
Eine
von uns selbst ge-
bloßer Augenblick, sondern eine Be-
sie kein
ist
festigung gegenüber
ihm kein eignes
und so
wir Lebenden und Handelnden
als
Gegenwart
alles,
Die Prägung aber kann niemand
Gegenwart
geistig ausgeprägten
liegt
dem Augenblick, ein Leben zeitloser Art.
Ein solches Leben wäre aber nun und nimmer erreichbar,
selbst
ja
das Streben danach würde eine Torheit, bestände nicht eine
ewige Ordnung
als
neue Art der Wirklichkeit, und wäre
eine
sie
nicht auch innerhalb unseres Lebensbereiches irgendwie gegenwärtig.
Denn was hülfe uns jene Ordnung, wenn sie nicht auch in uns wirkte?
Ohne das gibt es also keine Befreiung von der Geschichte, während
die Wendung dahin einen sicheren Standort ihr gegenüber erreichen
läßt.
Da es uns zu solcher Anerkennung einer selbständigen Geisteswelt bei allen Problemen drängte, so kann ihre Forderung hier nicht
Aber zugleich
überraschen.
erscheint eine
muß,
steht
bei
ihm
in
Befunde des Daseins.
zerfällt
in
Das Geistesleben
und stellt alle Mannigfaltigkeit
Menschenleben
ungeheure Verwicklung
dem er letzthin irgendwie wurzeln
schroffem Widerspruch mit dem nächsten
beim Menschen: jenes Geistige,
in
in
ist
vor allem ein Ganzes
umfassende Zusammenhänge, das
individuelle
Kreise,
innerhalb derer die
einzelnen Erscheinungen bunt durcheinander wirbeln; dort treibt die
und Freude der Sache das Handeln, hier herrscht die
Berührung mit geistiger Kraft
innere Kraft
natürliche Selbsterhaltung, die in der
sich leicht zu
einem unbegrenzten Egoismus steigert; der dort gefor-
derten Ewigkeit widerspricht die strenge Gebundenheit des Menschen
an die Zeit, der unaufhörliche Fluß
aller
Lebenserscheinungen samt
dem raschen Versinken der Individuen; im Geistesleben gewinnt die
Welt einen
selbstseins,
^
und gestaltet sich zu einem Reiche des Beisichder Mensch hingegen scheint geistig leer mid wehrlos
Inhalt
Leibniz
liebte die Alten
Foucher de Careil, lettres et opuscules II introd. XXXIII)
wegen la clarte dans l'expression et l'utilite dans les choses.
(s.
Geschichte.
265
Wie läßt sich eine so schroffe Kluft
gegenüber der Unendlichkeit.
überwinden?
Das Erste
ist
Umkehrung
innere
jene
sicherlich
des
Lebens,
die Erhebung über die bloßmenschliche Art, die Versetzung auf den
Standort;
geistigen
das tut
in
Wahrheit
Arbeit,
alle
die
auf
das
Ganze geht und zum ganzen Menschen wirkt; es braucht hier nur
als Ganzes verstanden und in volle Tätigkeit aufgenommen zu werden,
Aber
was mit tausendfachen Wirkungen unser Leben durchdringt.
Umkehrung und
solche
neue
solcher
Standort
führt
Es war eine
das neue Leben zu genügender Entfaltung.
weiteres
ohne
nicht
Überspannung menschlichen Vermögens, wenn man von
hier aus
unmittelbar durch ein möglichst energisches Kraftaufgebot alle Geistigkeit
hervorbringen wollte; diese Überspannung hat sich durch die
viel
zu blasse und schattenhafte Gestaltung der Welt gerächt, die
Nachdem
Schranken
des
Menschen
daraus
hervorging.
deutlich
genug vor Augen gerückt sind, werden wir nicht so
die
uns
leicht
wieder die Wirklichkeit aus freischwebender Tätigkeit zu konstruieren
versuchen.
So bedarf
nach Entfaltung
unser Streben
einer
zeit-
überlegenen Geistigkeit einer wirksamen Unterstützung; eine solche
liefert
ihm aber die Geschichte.
Nicht
freilich
die Geschichte, wie
sie als ein
ungeschiedenes Ganzes an uns kommt; denn dies Ganze als
ein Reich
von
lauterer Vernunft, eine reine
Entwicklung des Geistes-
lebens zu verstehen, darauf haben wir einstweilen verzichtet.
das schließt nicht aus,
daß sich
Aber
innerhalb der Geschichte irgend-
welche Eröffnung des Geisteslebens vollzieht, daß sich eine esoterische
Geschichte von einer exoterischen
bloßmenschlichen abhebt;
in
,
jener
eine Geistesgeschichte
mag
leben hervorbrechen, das durch allen
ein
von der
selbständiges Geistes-
Wandel der Zeiten hindurch
auch zu uns spricht und unser eignes Streben zu fördern vermag.
Am sinnfälligsten erscheint solches Geistesleben an einzelnen Höhepunkten, die klassisch heißen, weil an ihnen das Schaffen der bloßen
und dem bloßen Menschen überlegen wird. Das wahrhaft
Große waren dabei nicht einzelne Gedanken und Bestrebungen,
sondern eine neue Art des Lebens gegenüber den Zwecken und
Meinungen des Alltages; es vollzog sich dabei eine Umkehrung und
Zeit
mit ihr eine Erschließung geistiger Lebensquellen, geistiger
Kräfte
und Notwendigkeiten, eine Befreiung des Menschen von bloßmenschohne einen Zusammenhang
mit dem übrigen Leben, nicht ohne mannigfache Vorbereitung und
licher
Art.
Gewiß geschah das
nicht
Zu den Problemen des Menschenlebens.
266
eine enge Beziehung zur geschichtlichen Lage, aber nun
und nimmer
war jenes Klassische mit dem, was sein Wesen ausmacht, eine bloße
Summierung und Weiterbildung vorhandener Elemente. Vielmehr
ihm immer ein Bruch und eine Umkehrung, eine Versetzung auf einen neuen Standort, ein Gewinn eines neuen Lebens-
erfolgte in
raumes, ein Aufbauen einer geistigen Wirklichkeit.
So
pflegte
denn
auch sein Hervorbrechen schwere Erschütterungen mit sich zu bringen,
und soweit es siegreich wurde,
es das durch
ist
Kampf und Schmerz
geworden, es hat seine Vorkämpfer auch da zu Märtyrern gemacht,
wo das Märtyrertum nicht gerade durch Blut besiegelt wurde. Auch
besagt die äußere Anerkennung, die das Große schließlich zu finden
keineswegs
pflegt,
einen
Sieg und
reinen
eine Umgestaltung
der
Denn jene Anerkennung ist zugleich ein Herabzum menschlichen Dasein und ein Anpassen an die klein-
menschlichen Lage.
ziehen
menschliche Gesinnung; jedenfalls gelangt es hier nur mit einzelnen
Wirkungen, nicht mit dem Ganzen seines Wesens zur Geltung.
wird im Grunde
So
der Gegensatz nicht aufgehoben, sondern nur ver-
und durch die ganze Geschichte bleibt echte Geistigkeit und
steckt,
bloßmenschliche Lebensführung
in
hartem
Streit miteinander.
Nun aber erscheint selbständiges Geistesleben nicht nur an vereinzelten Punkten,
sondern diese Punkte suchen einen Zusammen-
hang und möchten
sich schließlich zum Aufbau eines allumfassenden
Reiches verbinden.
Dabei entstehen
und
harte
Zusammenstöße.
schwere Verwicklungen
freilich
Unter menschlichen Verhältnissen hat
jede Eröffnung des Geisteslebens bemessene Schranken; wie sie das
Problem nur an besonderer Stelle
und nur
angreift
in
besonderer
löst, so wird sie das Ganze des Geisteslebens, das aus dem
Grunde des Menschen wirkt, nicht voll befriedigen können;
schließlich wird eine Gegenbewegung entstehen und neue Entfaltungen
hervortreiben. Wiederum bringt das nicht bloß neue Ansichten und
Richtung
tiefsten
Bestrebungen, sondern Erweiterungen und Vertiefungen des Lebensprozesses; es
ist
der Lebensprozeß und mit ihm die geistige Wirk-
welche durch die Bewegung der Jahrtausende wächst;
lichkeit selbst,
es vollziehen sich in ihr
Erzeugnis der bloßen
Offenbarungen geistigen Lebens, die kein
Reflexion
der Tatsächlichkeit sprechen,
daher
erst
sind,
freilich
sondern die mit der Macht
einer Tatsächlichkeit
durch Selbsttätigkeit anzueignender
So wenig
menschlichen
diese
Daseins
Eröffnung geistigen
einnimmt,
Art.
Lebens
innerhalb
geistiger,
der
die
Breite
geistigen
des
Arbeit
Geschichte.
267
Höhe vor, ohne deren Erreichung
voll zu befriedigen vermag. Was
diesem weltgeschichtlichen Stande zurückbleibt, das mag die
übt sie Macht und hält ihr eine
sie
und
nicht wahrhaft zu fördern
hinter
Menschen
und
zeitweilig aufregen
wird es auf
fortreißen, schließlich
überlegenen Widerstand stoßen und als unzulänglich befunden werden;
indem er gewisse Lösungen
jener weltgeschichtliche Stand wirkt negativ,
als
unzulänglich verbietet,
gaben
wirkt positiv,
er
und gewisse Anregungen
stellt
indem
So kann dem Ganzen
gibt.
der Menschheit keine Lebensgestaltung
genügen, welche nicht die
und den moralischen Ernst
seelische Vertiefung
er gewisse Auf-
in
sich
aufnimmt,
die das Christentum brachte, aber auch keine, welche die Befreiung
des Subjekts und den Gewinn einer inneren Unendlichkeit verschmäht,
welche die Neuzeit errungen
Demnach
hat.
liegen in der Geschichte, geistig angesehen, Anweis-
ungen, Aufforderungen, Möglichkeiten;
belebt
sie
können
Zeit es
ins
sie
wollen angeeignet und
um zu vollen Wirklichkeiten für uns zu werden, aber
sein,
insofern jenes Geistige, so sehr besondere Nöte der
das,
Dasein hoben, seinem Kern nach zeitloser und daher
bleibender Art
es gilt
ist;
Eigentümlichkeit
zu
nur dieses Zeitlose
dann
ergreifen,
kann
seiner Kraft
in
es
uns zur
und
lebendigen
Gegenwart werden, dann ist die Geschichte kein bloßes Nacheinander
und das Frühere keine bloße Vorbereitung eines Späteren, sondern
dann
hat jedes
Große wie einen Selbstwert so eine unvergängliche
Wahrheit, auch läßt sich dann über die Vielheit hinaus ein Ganzes
Wenn so die Geschichte
erstreben.
aus einem
bloßen Strom der
Ereignisse zur allmählichen Eröffnung einer Geisteswelt, zum Gewinn
einer zeitüberlegenen
Gegenwart wird, so kann das Verlangen nach
einem gehaltvollen Geistesleben durch (sie die kräftigste Unterstützung
finden.
und
Nur
es
gilt
vom
eine Geistesgeschichte
Zeitlichen
von
dem
zum Ewigen durchzudringen
sonstigen
Chaos abzuheben.
Dieses aber hat seine festen Bedingungen; es fordert zunächst,
daß eine Tiefe des Lebens
ein
Ganzes
können
in
jenseit
den
jenseit
der nächsten Existenzform, und
Denn nur so
der einzelnen Funktionen wirke.
Bewegungen
Lebenstypen, mächtige
der
Weltgeschichte
charakteristische
Lebensströme aufkommen, die
nicht an die
Besonderheit des sichtbaren Ursprungs gebunden bleiben, sondern
darüber hinaus ins Ganze wirken, wirken mit ausgeprägter Art, nicht
mit vager Allgemeinheit.
Nur so läßt im Wandel der Erscheinungen
sich eine innere Einheit
erkennen und
in
die
Gegenwart
überleiten.
Zu den Problemen des Menschenlebens.
268
Zu solcher Überleitung und Aneignung gehört aber weiter, daß
die
eigne Zeit
muß
sich
sie
ein
Geistesleben
selbständiges
aber
ihre eigne welt-
was in ihr an Selbständiggeistigem und Zeitüberlegenem
erfassen, das
ist,
energisch herausarbeiten
Wandel der Erscheinungen
bloßen
dazu
Punkt des eignen Strebens
geschichtliche Aufgabe, den springenden
erreichbar
entfalte;
kräftig bei sich selbst konzentrieren,
und sich zugleich über den
sicher
Unser
erheben.
selbst
um bei anderen Charakteristisches zu geselbst Ewiges entdecken, um andere Zeiten und
müssen wir inne werden,
wahren,
bei
uns
das Ganze der Geschichte auf ihr Ewiges zu bringen.
sonders wird nur
Hier be-
dem gegeben, der da hat; hier ist besonders klar,
daß die Vergangenheit die Gegenwart wohl zu erhöhen, nie aber
zu ersetzen vermag.
Soweit derart
in
Bewegung der Geschichte eine durchallen Wandel der Zeit hin-
der
gehende Aufgabe entdeckt und durch
muß sich der
Nun treiben wir nicht mehr
durch zu einer zeitlosen Wahrheit vorgedrungen wird,
Gesamtanblick des Lebens verändern.
wehrlos mit
dem Strom
der Zeit dahin, sondern wir können ihm
an ewiger Wahrheit eine Ruhe und
Der Lebensprozeß wird sich nun durch die
Erfahrungen der Geschichte hindurch immer gehaltvoller und immer
gegenüber durch Teilhaben
Festigkeit gewinnen.
konkreter gestalten, das Geistesleben selbst in anschaulicherem Bilde
vor Augen stehen, die besondere Art und Lage der Menschheit sich
aufhellen,
in
dem
ein
allen
Typus und eine be-
charakteristischer
harrende Art unseres geistigen Seins zur Entfaltung kommen.
die damit das
Festigkeit,
gibt
erlangt,
selbst
in
ihm
eine
Die
menschliche Leben im innersten Grunde
Überlegenheit gegen die bloße Bewegung;
dem Wandel wird
es
nun vor allem
sich selbst erleben
Mögen die Erimmer wieder in die letzte
Grundlage zurückgreifen und den Menschen von neuem zum Probleme machen, was schon gesichert schien; mag, was an Ewigem in
uns wirkt, in die Besonderheit der Zeiten eingehen und ihr gemäß
und
sich
in
seiner
eigentümlichen Art bestärken.
schütterungen des geschichtlichen Lebens
sich gestalten müssen, trotzdem besagt es eine Wendung fundamentalster
Art,
wenn durch das Teilhaben
an
einer
zeitüberlegenen
Geisteswelt ein Ewiges im Kern unseres Lebens gesichert
Aufgabe vor allem dahin
gestellt
wird,
dieses
in
aufzunehmen und, was unser geistiges Wesen uns zuweist,
Besitz zu verwandeln.
und die
unsere Tätigkeit
in vollen
Denn nunmehr läßt sich danach streben, in
Geschichte.
269
der Geschichte Vergängh'ches und Unvergängliches zu scheiden und
eine geistige Gegenwart zu entringen; sie erscheint nun nicht
mehr als das Ganze, das seinen Zweck in sich selbst hat, sondern
als eine bloße Seite des Lebens und Seins, die nur in Zurückbeziehung auf eine zeitlose Ordnung einen geistigen Gehalt und
ihr
irgendwelchen Sinn gewinnt.
Nun und nimmer läßt sich demnach zu der älteren Art zurückwelche
kehren,
Ewige glaubte
das
uns
und
ergreifen
als eine Verleugnung der lebendigen
als eine Erstarrung,
wart zugunsten
Einem Zuge
in
können; die damit gewonnene Ruhe erscheint
völlig durchbilden zu
Gegen-
Aber wir brauchen deshalb
Vergangenheit.
toter
keineswegs der modernen Verflüchtigung alles festen Bestandes 2u verfallen,
das Leben in einzelne Augenblicke aufzulösen, damit aber allen
inneren Zusammenhang, alle überlegene Einheit preiszugeben.
wenn unsere Verbindung mit
einer
umzukehren
im Ewigen
nehmen,
Wirklichkeit vordringen,
Beharrende
werden
alle
inmitten
sie
Wahrheit einen
sich
unser Hauptstandort
hindurch
zu einer zeitlosen
läßt
Zeit
aller
Bewegung
Die Vergangenheit
festhalten.
bloße Vergangenheit,
wart
so
gestattet,
durch
zeitloser
und durch eine Wesensbildung
geistigen Charakter herauszuarbeiten
das Leben
Welt
Denn
ist
ein
überlegenes
dann nicht mehr eine
kann ein Stück einer zeitüberlegenen Gegen-
und damit eine Sache eignen Lebens, unablässiger
Arbeit bleiben.
muß
Die Wissenschaft
Überzeugung eine eigen-
aus solcher
tümliche Behandlung geschichtlicher Erscheinungen entwickeln,
die
am
und
Zeitlichen
das Bleibende,
So geschah
sucht.
es
z.
B.
am
in
Einzelnen
Iherings
das
Ganze
sieht
großem Werk über den
Geist des römischen Rechts, und zwar mit voller Klarheit über die
Art des Verfahrens.
Das Augenmerk ist hier «nicht das römische,
sondern
erforscht
(3.
das Recht,
Aufl.,
Einl. IX),
und
und veranschaulicht am römischen"
es wird
demgemäß zur Aufgabe, „das Ver-
und rein Römische von dem Unvergänglichen und Allgemeinen zu scheiden" (I, 15). Gewiß kann eine derartige philosophische Behandlung nur den Endpunkt einer langen wissenschaftgängliche
lichen
Arbeit bilden,
aber wer
sie
in
kleinmütiger
Besorgnis vor
ihren Gefahren abweisen wollte, den
würde Hegels bekanntes Wort
von der Metaphysik
einen Tempel ohne ein Aller-
treffen:
er will
heiligstes.
Auch
in
das Leben
des
Individuums erstreckt sich
die
neue
Zu den Problemen des Menschenlebens.
270
Nur so lange ist für den
Art und gibt ihm eine neue Beleuchtung.
Einzelnen das Dasein
eine
Flucht von
rastlose
Erscheinungen,
als
er eines selbständigen Innenlebens entbehrt, nicht irgendwie zu einem
Ganzen
persönlichen
und
Seins
Individualität
geistiger
gelangt.
Denn wo das geschieht und sich damit das Ereignis in ein Erlebnis
zu verwandeln, der Mensch in Werk und Schicksal ein geistiges
Selbst
zu erleben vermag, da
was uns irgend bewegt, nicht
zieht,
wie ein Schatten vorüber und versinkt
sondern da vermag es Wurzel
entfalten wie zu fördern,
in
den Abgrund des Nichts,
uns- zu schlagen.
in
einer zeitüberlegenen
Bleibendes zu
Gegenwart sich
ein-
zuordnen.
Dem Leben eine gehaltvolle Gegenwart zu sichern und
es damit
gegenüber dem Augenblick zu befestigen, das bleibt immer
das Hauptziel;
Liebe und
solcher
in
Leid,
in
Gegenwart
Daher haben
Lebens wurde.
fährt
Glück und Unglück
von jeher
fort
je
ein
zu wirken, was
in
Stück selbsteignen
geisteskräftige
Menschen
über die Flüchtigkeit des Lebens zu klagen verschmäht, da es doch
bei
uns selber steht, uns jener Flüchtigkeit zu entwinden und unser
Leben
ins
Unvergängliche zu
und sich
in
«Ich bedaure die Menschen",
stellen.
«welche von der Vergänglichkeit
sagt Goethe,
viel
Wesens machen
Betrachtung irdischer Nichtigkeit verlieren; sind wir
eben deshalb
da,
ja
um das Vergängliche unvergänglich zu machen."
So kann uns auch das bekannte Woil Dantes nicht als richtig gelten,
daß das größte Elend darin
ist
bestehe, sich
im Unglück vergangenen
Denn war das Glück wahrhaftiges Glück,
Glücks zu erinnern.
es gar nicht zerstörbar, so
so
muß es durch alles Unglück hindurch
mit lebendiger Gegenwart wirken.
Auch
die
natürlichen Phasen
scheinen in solchem
ander.
eine jede
sie
bleibt
darin
auf,
spätere
dem Leben
seinen Gesamtstand.
eine Jugend
er-
Zusammenhange nicht als ein bloßes Nachein-
Diese Phasen leben sich weder gänzlich
noch gehen
Jugend,
des Lebens, die Lebensalter,
Daher
ist
in
sich selber aus,
Phasen vorzubereiten, sondern
innerlich
gegenwärtig und wirkt auf
so wichtig eine frische
wahrhaftiger Art;
sie ist
und freudige
mehr als eine Sache
sentimentaler Erinnerung, sie kann ein Stück einer weiteren Gegen-
wart bleiben, eine unversiegliche Quelle frischen Lebens.
Demnach ist der Mensch keineswegs ein bloßzeitliches Wesen;
mehr Recht meinten tiefsinnige mittelalterliche Denker, daß er
an der Grenze, dem Horizont von Zeit und Ewigkeit stehe und
mit
teil
an ihnen beiden habe.
Die Zeit
ist
für uns weniger ein starres
Geschichte.
271
Schicksal als ein Problem; wie weit aber das Leben sie überwindet
und
Gegenwart erreicht, das hängt vor allem an
eine überzeitliche
der geistigen Kraft, die es aufzubieten vermag; bei uns selbst steht
es
ob der Schwerpunkt unseres Seins
schließlich,
oder ins Unvergängliche
fällt.
Allerdings
hat
und
entbehrlichen Voraussetzung die Wirklichkeit
ins Vergängliche
dieses
Tun zur un-
die innere Gegen-
wart einer geistigen Welt, auch die leidenschaftlichste Erregung des
bloßen Subjekts kann nie einen geistigen Inhalt und mit ihm eine
Zeitüberlegenheit erreichen,
Schaffen
zugleich
und
es
Empfangen
ein
bleibt
für
den Menschen
unsichtbaren
aus
alles
Zusammen-
hängen.
Nach dem allen bedeutet unsere Abweisung des zerstreuenden
und erschlaffenden Historismus keinen Rückfall in den Rationalismus.
Das
beiden
gestellt,
das von
ist
eine
freilich
ihm
gestehen wir gern, daß, vor eine Wahl zwischen
wir den Rationalismus vorziehen würden; denn mag
entfaltete
Leben noch so eng und
einseitig sein,
es
und Wollen, während dem Historismus
Nachbildung fremden Lebens genügt. Aber es bleibt genug,
doch
eignes Leben
ein
was uns vom Rationalismus scheidet.
verleitete ihn, die
Sein überspanntes Kraftgefühl
Aufgabe zu unterschätzen; seine Verkennung des
weiten Abstandes zwischen
dem nächsten Dasein und der Tiefe des
Menschenwesens
von einer unmittelbaren Aufraffung
ließ
ihn
warten, was in Wahrheit durchgreifende Vertiefungen
ungen
von einer Klärung konnte er nicht wohl
verlangt;
erhoffen, hätte er nicht die Vernunft schon in
und
vorhan/Jen
er-
und Umwälzalles
Heil
unserem Daseinskreise
lediglich einer Freilegung bedürftig geglaubt.
Das
war überhaupt, weit über den Rationalismus hinaus, der Irrtum der
Neuzeit,
Wesen des Geisteslebens
das
Daseins zur Bewußtheit zu setzen;
um
in
die
bloße Erhebung des
uns schien dasselbe wirksam,
und dunkel, was in uns zur vollen Freiheit und
Klarheit gelangt. Denn es wurden dabei die großen Widerstände und
nur gebunden
V^erwicklungen unseres Weltanblicks ebensowenig gewürdigt, als der
Überwindung zur nötigen Tiefe vordrang.
Ganz anders stellt sich die Sache, wenn das Geistesleben nicht als
Lebensprozeß
in
ihrer
eine bloße Aufhellung der Natur, sondern mit seinem Beisichselbstsein
als
eine
wesentlich
neue
Art
des
Lebens
verstanden
wird.
Wenn damit eine weit größere Spannung entsteht, so gewinnt auch
die Geschichte an Bedeutung, nur wird
dürfen,
sie
in
ein Reich
man nicht darauf ausgehen
der reinen Vernunft zu verwandeln, son-
Zu den Problemen des Menschenlebens.
272
dern zufrieden sein müssen,
ihr irgendwelche
in
Eröffnung der Ver-
nunft zu entdecken.
Auch
schichte
war die Aufklärung einer Anerkennung der Ge-
insofern
wenig günstig,
des Lebens
der Wahrheit
welche dort die Führung
als die Intelligenz,
einen
hatte,
zu engen und unduldsamen Begriff
viel
Eine bloß intellektuelle Wahrheit will unmittel-
hat.
bar als ausschließlich gelten, es verträgt sich hier nicht Verschiedenes
nebeneinander; der Gegenwart Recht geben, heißt hier die ganze
Wie sich die Lage völlig verwenn das Intellektuelle auf das Geistige aufgetragen wird,
und wenn in der Geschichte nicht bloß Lehren und Meinungen,
Vergangenheit ins Unrecht setzen.
ändert,
sondern Lebensentfaltungen und
in
ihr
lichkeiten
selbst
Lebenskomplexe zusammentreffen,
um Bilder der Wirklichkeit,
bloß
nicht
sondern
um Wirk-
gekämpft wird, das dürften die früheren Ausführ-
ungen zur Genüge dargetan haben.
Entscheidend
ist
immer der Gewinn einer zeitüberlegenen
hier
Gegenwart mit ihrer Umkehrung des Lebens. Denn nur dadurch kann
die
mehr
Geschichte
nur dadurch
Sache gelehrter Forschung werden,
eine
als
läßt sich verhindern,
daß die unbegrenzte Ausdehnung
des Werdens und damit der geschichtlichen Betrachtung einen zer-
Der Sieg einer geschichtlichen Be-
störenden Relativismus bewirke.
trachtung
wohl
der
größte
Nicht
nur
beim
ist
Forschung.
Triumph
Weltbau
gesamten
der
und
den
bei
neueren
organischen
Formen läßt diese Betrachtung allen vorgefundenen Bestand aus dem
Werden verstehen, sie erstreckt sich bis in die elementarsten Vorgänge
der
weithin
das
leblosen
Natur,
Geschehen
in
umkehren
indem
einer
selbst
im
Gebiet
bestimmten Folge
der
Physik
verläuft,
sich
Das menschliche Dasein aber erscheint in weit klarerem Bilde, seitdem die Gegenwart als das letzte
nicht
beliebig
Glied
einer
langen
Kette
läßt.
gewürdigt wird,
Hauptrichtungen des Strebens manches
was sonst als ein
der Mensch bis
fester Einsatz galt,
in
und
nicht
als veränderlich
nur in den
erkannt
ist,
sondern auch einleuchtet, wie
seine seelische Art hinein
von der Besonderheit
seiner Zeit abhängt, wie verschiedene Zeiten verschiedene Menschen
hatten.
Ein
unermeßlicher Reichtum
des Lebens
geht damit auf,
das Verständnis wird weit präziser, indem es solchem Reichtum sich
anschmiegt^
*
Wir dürfen das alles als eine wesentliche Erweiterung
Wir erinnern nur an Diltheys geistvolle Zeichnungen der Menschen
verschiedener Jahrhunderte; auch Lamprechts Forschungen seien hier nicht
Geschichte.
273
unseres Gesichtskreises, eine Befreiung von der Gebundenheit einer
Aber der Gewinn an Wissen kann zu
besonderen Zeit begrüßen.
einem Verlust für das Leben werden, wenn
Erweiterung eine Befestigung und
Ewigen entgegenzusetzen.
stärkung des
das Zweite bleiben,
sie
nehmen,
als er
jener
muß uns
Die Geschichte
darf nie das Erste werden.
Überzeugung unser Daseinskreis
dieser
es nicht gelingt,
dem Wachstum der Zeit eine Ver-
sich
weit
Wohl wird bei
unfertiger
aus-
dem Rationalismus und der konstruktiven Geschichts-
philosophie erschien, aber woher sind wir denn dessen gewiß, daß
bei
uns der Kreis des Lebens sich schließt, und was schadet die
bescheidnere Fassung,
wenn dem Kleinerwerden des Menschen ein
entspricht und das Leben an Tiefe
Größerwerden der Wirklichkeit
gewinnt, indem es sich minder einfach darstellt?
Anhang: zum Begriff des Modernen.
Der Begriff des Modernen bewegt und entzweit
die
Gemüter
heute so sehr, daß einige Erörterung und Aufklärung nicht zu
um-
gehen
des
ist.
Ausdrucks,
Eine
Aufklärung
über
die
im Schwange
zunächst
fordert
sehr unklare,
wenn
die
nicht
Geschichte
irrige
Meinungen
sind.
Das sachliche Problem
reicht
natürlich weit über die
Prägung
Den seelischen Stand der Gegenwart behandelt u. a. R. Baerwald,
vergessen.
„Psychologische Faktoren des modernen Zeitgeistes" (Publikation der Gesellschaft
für
psychologische Forschung).
Es hat aber das Problem der Ab-
hängigkeit des Menschen von seiner Zeit von alters her die Gedanken beschäftigt
und
sich schon
Da sich für uns ein
im
17.
Jahrhundert zu einer Streitfrage zugespitzt.
näheres Eingehen auf die Sache verbietet, so
sei
nur
Walchs Philos. Lexikon angeführt (schon in der ersten Aufl.
von 1726) Art. Sitten, S. 2377: „^t\\ nun solche Veränderung (nämlich der
Sitten) fast unvermerkt geschieht, und wir es gemeiniglich nur gewahr
werden, wenn es vorbei ist, daß diese und jene Sitten zu der und jener
Zeit Mode gewesen, so pflegt man solches der Zeit zuzuschreiben. So haben
einige einen genium seculi statuieren wollen, welcher nach den Zeiten die
Gemüter der Menschen lenke und die Sitten der Menschen verändere. Dieser
Meinung ist Barclajus, welcher in icone animor., pag. 505 (John Barclays icon
animorum erschien 1614) sagt: omnia secula genium habent, qui mortalium
animos in certa studia solet inflectere. Mit diesem stimmt überein der ungenannte Autor, der Germaniam milite destitutam geschrieben, und der sogenannte Pater Firmianus, von dem ein besonderes Buch unter dem Titel
eine Stelle aus
seculi genius, Paris 1663,
Eucken, Grundbegriffe.
12 heraus(kam)".
4. Aufl.
18
Zu den Problemen des Menschenlebens.
274
des Ausdrucks zurück;
wo immer daran lag, das Eigentümliche der
Gegenwart abzugrenzen, da werden sich irgendwelche Bezeichnungen
gefunden haben. ^
In
,/
modern" aber entstand
beharrender Ausdruck, dessen
ein
Das Wort
Geschicke etwas näher zu verfolgen sich lohnen dürfte.
(abgeleitet
modo
von
eben,
jetzt)
wird
besonders
geläufig
sein,
wenn innere Verschiebungen zum Bewußtsein kommen und die
Menschen entzweien.
Dann nennt sich der Freund des Neuen
modern, um seine Überlegenheit gegen solche zu bekunden, die zähe
am Alten haften; der Gegner aber stempelt den Ausdruck zu einem
Schmähwort, das einen Menschen kennzeichnen
und ohne
Pietät
Die Geschichte des Wortes
Höhe
sondere
soll,
der ohne Halt
den flüchtigen Anregungen des Augenblickes
erreichte,
läßt ersehen,
wann der Streit
und welcher Punkt
es
folgt.
eine be-
vornehmlich war,
der die Geister schied.
Der Ausdruck erscheint im Übergang vom Altertum zum Mitteldem Grammatiker Priscianus, der im 6. Jahrhundert lehrte,
alter bei
dem Beamten Theodorichs (f um 575).^ In
den folgenden Jahrhunderten findet es sich hie und da. ^ Zu einem
eigentlichen Partei- und Streitwort wurde «modern« seit dem Ende
des 11 Jahrhunderts, das aber auf dem Gebiete der Logik; es diente
und
bei Cassiodorius,
.
nämlich zur Bezeichnung der Nominalisten, d. h. solcher, welche den
Begriffen des Denkens keine objektive Realität zuerkannten.*
„Moderne"
werden aber auch andere genannt, auch einfach die Gelehrten der
*
So hat z. B. Aristoteles wiederholt den Ausdruck o! vGv.
bezefchnet
vGv
1^
er deutlich
(piXodo^fa,
^iQ-taaiv
Toc
die Platoniker seiner Zeit: y£y°^^^
ebenso auch lü69a, 26:
y*P T^^T xaO^oXou, a
(paaiv
ol
[ilv
"^^
Met. Q92a, 33
(xaÖTri'jjiaTa
tot;
vüv Ta xaO-o'Xou oüaia? jiocXXov
«px*? "O" ouafa^
etvai
(xaXXov
Sia
to
Xo^ixtü? ^T)T^v Ol 5c Jva'Xat ta xa^ ^xaorov, oTov nup xa\ ytjv, äXX'oü xo xotvov aöJjAa.
*
Cassiod. Variarum 4, 51 wird ein Architekt empfohlen als antiquorum
imitator,
*
modemorum institutor.
Ein Artikel der Historisch-politischen Blätter (139, 5 vom Jahre 1907)
erwähnt einen Brief des Abtes Benediktus Avianensis (geschrieben zwischen
modemos scholasticos, maxime
apud Scotos [ijste Syllogismus delusionis, ut dicant trinitatem sicut personarum
ita esse substantiarum.
Mon. Germ. bist. Epist. Carol. Aevi, Tom. II, 563.
* Prantl, Geschichte der Logik im Abendlande II, 82, führt die älteste
Stelle an, wo die Nominalisten als modemi bezeichnet werden (non juxta
quosdam modemos in voce, sed more Boethii antiquorumque doctorum in
re discipulis legebat, nämlich Otto, seit 1106 Bischof von Cambray).
800 und 821), worin es heißt: Unde apud
Geschichte.
eignen
275
Einen bedeutenderen Gehalt und eine schärfere Zu-
Zeit. ^
spitzung erhielt die Sache, als seit Johannes von Salisbury die Aristoteliker des 13. Jahrhunderts (also
namentlich die großen Dominikaner,
Thomas von Aquino, derselbe Mann,
an den jetzt sich alles Antimoderne hängt) Moderne genannt werden,
im Gegensatz zu der mehr durch Plato und Augustin beherrschten
wie Albert der Große und
Denkweise, die von der franziskanischen Theologie^ vertreten wurde.
Diese «moderne" Denkweise erschien den Gegnern
als eine
Über-
flutung der Theologie durch dialektische Sorgen und Spitzfindigkeiten.'^
Später überträgt sich der Begriff wie der Ausdruck Modernität auf
Ockam und seine Schule, »Ockams Lehre blieb die ,moderne' Theologie bis in
Luthers Zeit",^ auch Luther
hat
sich
Aber es findet sich das Wort auch
in
vom gemeinsamen Leben
eine devotio
standen
darunter eine solche,
stark die
den
trägt
vertraten
»Innigkeit"
Titel
Aber
Brüder
modema und ver-
welche neben der äußeren
Haltung
Werke von Johannes Busch
über de origine devotionis modernae. ^
es dauert, lange,
für das Bewußtsein
lich
ihr bekannt.
anderer Bedeutung.
betonte, eines der
Das Mittelalter versinkt und
Welt.
zu
klärt
und
bis
die Renaissance eröffnet eine neue
was
feste
in
den Geistern vorgeht,
Bezeichnungen annimmt.
sich
Natür-
konnte der Renaissance „modern" nicht eine neue Art gegen-
über
dem Altertum, sondern nur eine neue Art in der Behandlung
des
Altertums
bedeuten;
da zugleich
der
mittelalterliche
Sprach-
gebrauch verblieb, so lief verschiedenartiges bunt durcheinander. Die
epistolae
^
obscurorum virorum
stellen
das deutlich
vor Augen. ^
Eine nähere Darlegung dessen hat hier kein Interesse, es
auf Prantl verwiesen sein
(s.
z.
B. II,
Je
mag dafür
116ff, 195, 241).
^ Von Roger Bacon werden Alexander von Haies
und Albert genannt
duo modemi gloriosi (s. den Artikel Scholastik von Seeberg in Herzogs Real-
enzyklopädie).
Der päpstliche Legat Simon de Brion, der tief in die Bewegungen
welche damals die Pariser Universität aufregten und beinahe zerrütteten, erwähnt tadelnd die modema curiositas, quae plus solito innumeras
multiplicat quaestiones (s. das ausgezeichnete Werk von Mandonnet: Siger de
Brabant et l'Averroisme latin au XIII siede [1899} CCVIII, Anm. 1).
* S. Seeberg, Herzogs Realenzyklopädie, 3. Aufl., XIV, 279.
* S. den Artikel von Gustav Boerner:
„Die Brüder des gemeinsamen
Lebens in Deutschland" in den Deutschen Geschichtsblättem von 1905, Juni.
^
eingriff,
S.
namentlich
S. 244/5.
modernus bedeutet hier bisweilen nichts anderes als neu (modernus
episcopus, modernus imperator), auch die ältere Bedeutung aus dem Streit
*
18*
Zu den Problemen des Menschenlebens.
276
mehr aber die Neuzeit mit dem Beginn des 17. Jahrhunderts eine
Selbständigkeit und ein Selbstgefühl gewann, desto stärker mußte es
von allem Früheren ab-
die Gelehrten drängen, das Eigne deutlich
zuheben und damit die Weltgeschichte anders zu ordnen und
zuteilen
als
bisher
geschehen
Es
war.
wohl
ist
ein-
namentlich
der
Aufschwung der Naturwissenschaften und die Blüte der französischen
Literatur gewesen, welches der damaligen Zeit das Bewußtsein verliehen,
Das ergab
etwas neues und allen früheren Zeiten überlegenes zu sein.
vor allem den Gegensatz der anciens und der modernes.
kanntes
Buch:
des
Parallele
anciens
des
et
modernes
behandelt die Ausdrücke schon als eingebürgert, es
so
Hälfte
lagen Reflexionen
des Modernen
War
erfüllte.^
einmal
(1688
der
ff.)
aber bezeich-
ist
17. Jahrhundert
nend für das Selbstgefühl, welches das
zweiten
Perraults be-
Gegensatz
seiner
in
geschaffen,
über die eigentümliche Art des Antiken und
nahe; wir wissen, wie
Bewegung das
viel
hervor-
gerufen und wie namentlich Schiller einer näheren Bestimmung dieser
Begriffe hingebende
und eindringende Arbeit gewidmet hat.
Moderne
Andererseits hatte das
abzugrenzen,
Das aber ist
erst gebildet sein.
bemerkt darüber
trotz
sich
und dazu mußte der
(a. a.
gegen das
Begriff
spät
Mittelalterliche
des Mittelalters
genug geschehen.
selbst
Bernheim
O. S. 69): «Der Bann der Tradition dauerte
doch noch lange
einzelner Angriffe
fort.
Noch
ein
Sleidan,
der bekannte Historiker des Zeitalters Karls V., nennt seine Chronik
»De quattuor monarchiis" und hält trotz aller von ihm aufgeführten
Zeichen der Auflösung des heiligen Römischen Reiches den Glauben
an dessen Fortbestehen
fest,
weil eine fünfte irdische Weltmonarchie
nach Daniels Prophezeiung unmöglich
sei.
Im 17. Jahrhundert kam
et modemi), meist aber bezeichnet
Anhänger der neuen humanistischen Denkart, z. B. poetae modemi;
ex quo in Erphordia sumus modemi artista de via modemomm. Oft findet
sich der Ausdruck nicht.
^ Es seien aus dem
ersten Dialog jenes Werkes nur ein paar Stellen
dafür angeführt: je pretens que nous avons aujourd'hui une plus parfaite
der logischen Schulen erhält sich (antiqui
es die
;
connaissance de tous les arts et tous les sciences, qu'on ne l'a jamais eue.
—
Weiter spricht er von dem progres prodigieux des arts et des sciences, depuis
cinquante ou soixante ans.
Ferner:
il
ne faut que
lire
les
joumaux de
yeux sur les beaux ouvrages des academies
de ces deux grands royaumes pour etre convaincu que depuis vingt ou trente
ans il s'est fait plus de decouvertes dans la science des choses naturelles, que
France et d'Angleterre
et jetter les
dans toute l'etendue de la savante antiquite.
Geschichte.
man zuerst
lich
277
zu einer brauchbaren Einteilung des Stoffes.
den Philologen und Literaten drängte
Nament-
das Bedürfnis auf,
sich
den augenfälligen Unterschied zwischen der klassischen und
alterlichen
Sprache und Literatur
Bildung
literarischen
seit
der Renaissance andererseits zum stehenden
Ausdruck zu bringen, und
oder
oder
von den Antoninen
für
die Literaturepoche
von Augustus
bis ins
15. Jahrhundert.
Der Hallenser
— 1707) war
der in seinen
Professor Christoph Cellarius (1634
Kompendien
dieses
Bezeichnung media
es bildet sich so die
medium aevum
aetas
mittel-
der letzteren und der
einerseits,
auf
Einteilungsprinzip
gemeinen anwandte, indem
er
die
unterschied
es,
Geschichte
Historia
im
all-
antiqua bis zu
Konstantin
dem Großen, und zwar so weit, nicht nur bis Augustus,
weil, wie
er ausdrücklich
die innere
erklärt,
römischen Reiches noch weit über A
und äußere Blüte des
o^ustus' Zeit hinausreicht; Historia
medii aevi bis zur. Eroberung Konstantinopels durch
Historia nova.
Und
diese Einteilung
nicht' ohne lebhaften Widerstand,^ durch."
dem
Mittelalter
die
Türken;
drang allmählich, wenngleich
So war „modern" auch
gegenüber abgegrenzt; die weiteren Geschicke des
Ausdrucks auf dem Boden der Neuzeit zu verfolgen, das würde ins
Unbegrenzte führen und
ist
für
unsere Aufgabe keineswegs nötig.
So viel haben wir gesehen, daß der Ausdruck „modern" weit weniger
^
Wie langsam aie Sache sich durchgesetzt hat, und wie sie
bis
zum
wenig bekannt. Ein Artikel der Revue
des deux mondes vom 15. Januar 1907 über den hochverdienten belgischen
Historiker Godefroid Kurth von Georges Goyau bemerkt über das Verhalten
heutigen Tage Streit erweckte,
der französischen Akademie
ist
zum Ausdruck Mittelalter folgendes:
premieres editions du dictionnaire de l'Academie
frangaise
Les cinq
contiennent
au
mot „moyen age" l'article suivant: „On appelle autheurs du moyen äge les
autheurs qui ont ecrit depuis la decadence de l'empire romain jusque vers
le
X siecle ou environ."
C'est seulement dans la 6e edition (1835)
qu'on
„Moyen äge, le temps qui s'est ecoule depuis la chute de l'empire romain,
en 475, jusqu'ä la prise de Constantinople, par Mahomet, en 1453." Kurth
selbst bekämpft den Begriff und Ausdruck Mittelalter sehr entschieden.
In
der Schrift Qu'est ce que le moyen äge tritt er dafür ein, einen einzigen
Hauptschnitt mit dem Eintritt des Christentums zu machen, und so behandelt
sein großes Werk Les origines de la civilisation moderne (3. Aufl. 1898) das
Mittelalter als den Beginn der modernen Welt.
In jener kleineren Schrift
heißt es: Loin que le moyen äge soit intermediaire entre la civilisation
antique et la civilisation moderne, le moyen äge est lui-meme le commencelit:
ment de la
civilisation
moderne.
Loin qu'il
faille
de depart de celle-ci aussi bas que l'epoque de
stater
au contraire qu'elle sort du christianisme.
la
faire
descendre
Renaissance,
il
le
point
faut con-
Zu den Problemen des Menschenlebens.
278
modern
ist,
als
elastischer Art
Soviel
man gewöhnlich
und daß der Begriff sehr
denkt,
ist.
zur Geschichte des Ausdrucks, nun einige Worte zur
Daß der Begriff des Modernen so viel Bewegung und Streit
kommt letzthin darauf zurück, daß es zum glücklichen
Sache!
hervorruft,
Fortgang
sowohl
der Kulturarbeit
eines
Festhaltens
des Alten als
von Neuem bedarf. Wir würden schwerlich viel
kommen, wenn wir immer von neuem beginnen müßten,
wenn unsere Arbeit nicht geeignete Werkzeuge und leichteste Bahnen
zu sicherem Besitz gewänne, wenn nicht vieles, was zunächst die
volle Anspannung bewußter Tätigkeit fordert, ins Unbewußte und
Gewohnheitsmäßige gebildet würde und damit freie Zeit für vordringendes Schaffen ließe.
Wie nützlich, ja unentbehrlich ist z. B.
der Philosophie der reiche Schatz von Begriffen und Kunstauseines Aufbringens
weiter
drücken,
den
die
Arbeit
vereinte
Aber die Sache geht noch
tiefer.
von Jahrtausenden
bereitet
hat!
Was an Wahrheit und überhaupt
an geistigem Gehalt erreicht wurde,
das konnte die Überzeugung
und Hingebung der Menschen nur gewinnen, indem es sich über
allen Wandel der Zeit hinaushob und alle Veränderung abwies;
soweit wir
Wahres echter Art besitzen, stehen wir über dem Fluß
der
Solcher Denkweise entsprang das Wort:
Zeit.
„Die Wahrheit war schon längst gefunden,
Hat edle Geisterschaft verbunden;
Das alte Wahre, faß' es an!"
Das rechtfertigte die Hochschätzung des Alten und
ung,
die
eigne Arbeit eng
damit zu
verketten,
allen
die Forder-
und jeden
schroffen Bruch zu vermeiden.^
^ In der Philosophie ergab das den Gedanken einer philosophia perennis,
der, schon in der Scholastik angelegt, mit besonderem Nachdruck von Agostino
Steuco verfochten wurde (er schrieb de perenni philosophia libr. X, Bas. 1542);
Leibniz
nahm den Ausdruck auf, gab ihm aber aus der Idee einer stetig fort-
schreitenden Entwicklung einen veränderten Sinn.
Neuerdings hat namentlich
Trendelenburg, wiederum in einer eigentümlichen Weise, den Gedanken der
Stetigkeit der philosophischen Arbeit verteidigt:
«Die Philosophie", so sagt er,
»wird nicht eher die alte Macht wieder erreichen,
als bis sie Bestand gewinnt,
und sie wird nicht eher zum Bestände gelangen, als bis sie auf dieselbe Weise
wächst wie die anderen Wissenschaften wachsen, bis sie sich stetig entwickelt,
indem sie nicht in jedem Kopfe neu ansetzt und wieder absetzt, sondern geschichtlich die Probleme
aufnimmt und weiterführt" (Vorwort zur 2. Aufl. der
Log. Untersuchungen, S. Vlll).
Geschichte.
279
Aber der Vertreter des Neuen hat dem manches entgegenzuwie äußere Dinge es tun,
zur anderen,
und anerkannt
angeeignet
und
sein,
in
es will
vermeiden
der äußere
wird
Bestand
und
hältnis
derselbe
bleibt,
immer von neuem
der Aneignung wird eine
gewisse Verschiebung sich schwerlich
in
von der einen Zeit
Geistiges überträgt sich nicht so einfach
halten.
Auch wo
lassen.
sich
das
leicht
dem Alten
etwas
Ver-
man wird
die Bewertung der einzelnen Teile verändern,
anderes sehen und etwas anderes an ihm zur
Dazu kom.men neue Lagen und
Hauptsache
machen.
Probleme;
der Mensch
stellen
neue
kann diesen nicht gewachsen werden ohne
auch innerlich weiterzustreben; Kulturen leben sich aus, neue Völker
mit neuer seelischer Art; sollte das alles den Stand des
erscheinen
Geisteslebens gar nicht berühren?
daß das überkommene Leben
daß der eingeschlagene
es
ferner zweifellos ausgemacht,
und
steht,
Weg uns gerade zum Ziele führt? Ja gibt
wahrhaftiges Leben
ein
Ist
unantastbarer Wahrheit
in
ohne
eignes Entscheiden ohne Zweifel
ein
eignes Entscheiden,
und
ein
und Kampf, ohne ein Umwandeln
und Weiterbilden?
Was an Verschiebungen erfolgt, mag zunächst innerhalb einer
unantastbaren Welt zu liegen scheinen, ja die Veränderung wird lange
Dann aber kommt ein
wo die Spannung übergroß und eine Losreißung vom Alten
hindurch überhaupt nicht empfunden.
Zeit
Punkt,
für die Frische
und die Wahrhaftigkeit des Lebens unerläßlich wird,
wo die geistige Selbsterhaltung einen Bruch mit der Tradition und
aus unmittelbarer Gegenwart zwingend verlangt.
Ob
und wann solche Umwälzungen nötig werden, darüber kann
allein
ein Schaffen
die
Erfahrung der Geschichte
die
radikalste
von
zeigt
sie
sie
aber jedem
Denn eine solche Umwälzung, viel-
Unbefangenen deutlich genug.
leicht
belehren,
allen,
bildet
das Eintreten
des Christen-
tums mit seiner von Grund aus neuen Schätzung der Dinge; das
gute Recht
und
die
ainer solchen
Umwälzung dürfen auch die Reformation
neue Wissenschaft für sich
in
Anspruch
religiöse Leben der Neuzeit hätte seine Kraft
nicht gefunden
ohne
nehmen.
und seine
ein selbständiges Neueinsetzen
Das
Innerlichkeit
und ein Hervor-
brechen ursprünglicher Kräfte, ebenso wenig hätte die neue Wissenschaft mit ihren völlig
neuen Ausgangspunkten und Methoden sich
allmählich aus der Scholastik heraus gestalten können.
liche
Leben
darf es
Das mensch-
bedarf gewiß einer Kontinuität, aber nicht minder be-
einer Diskontinuität,
um
in frischem
Fluß zu bleiben und
Zu den Problemen des Menschenlebens.
280
Nur das kann bei den Be-
seine ganze Tiefe entfalten zu können.
wegungen Sache des Streites sein, ob in ihr geistige Notwendigkeiten
wirken und walten, oder ob dabei nur ein menschliches Bedürfnis
nach Abwechslung in Frage steht.
Das nämlich ist nicht zu leugnen, daß nicht alle Wandlung aus
daß im menschlichen
solchen geistigen Notwendigkeiten entspringt,
namentlich im gesellschaftlichen Zusammenleben, auch ein
Dasein,
mehr
Müdewerden des Menschen am
subjektives
Alten,
ein
Be-
dürfnis nach Veränderung wirkt, wie es besonders greifbar die Tat-
sache der
Mode zeigt.
einander,
die
Dabei entfernen die Zeiten sich weit von-
einen fühlen
sich
indem
wohl,
ruhig die alten
sie
Wege verfolgen, andere zeigen eine eigentümliche Unruhe, ein Unbehagen an allem, was vorgefunden wird, eine Vorliebe für alles,
was an Neuem
auch
sich
dem
mit
Diese verschiedene Art hängt sicherlich
regt.
des Geisteslebens zusammen, jene Unruhe
Stande
bezeugt eine Kluft zwischen inneren Notwendigkeiten und äußerem
aber solcher Lage bemächtigt sich leicht die Neuerungssucht
Besitz,
des bloßen Menschen und entwickelt eine Neigung, das Alte zu verwerfen, weil es
alt,
das Neue zu feiern, weil es neu
ist.
Demnach giU es zwischen Modernem echter und unechter Art
zu scheiden, einem Modernen, in dem eine geistige Notwendigkeit
wirkt,
und einem Modernen, das bloßmenschlicher Lust und Laune
Grundverschieden sind die Wirkungen und
verdankt.
sein Dasein
Aussichten beider.
Menschen
nach
Erzeugt die Bewegungen nur ein Verlangen der
Veränderung,
und
Hin-
ein
Herschwanken
der
Stimmung, so mögen sie die Oberfläche noch so sehr erregen, sie
dringen nicht durch bis zur Tiefe und gewinnen keine Kraft des
Schaffens, derselbe Wind, der sie brachte, wird sie bald auch wieder
verwehen, und der rasche Wechsel, der
einem Extrem
starke
der
ins
andere
umschlagen
Ermüdung bewirken;
Zeit,
anders
steht
es,
wenn
die
muß
läßt,
traurig das
das an solchem Modernen
Völlig
leicht
Menschen von
schließlich
eine
Leben des Menschen wie
haftet.
ein
Modernes echter Art eine
Wendung des weltgeschichtlichen Lebens vertritt und ihrem Wahrheitsgehalt zur Anerkennung verhelfen möchte.
in
sich
eine
geistige
Notwendigkeit,
Dauer kein Widerstand
solcher
und
Art
wohnt
eine
zu
deren
hindern vermag.
wunderbare
Kraft.
Denn dann trägt sie
Durchdringen für die
In
einem Modernen
Scheinbar
vereinzelte
zerstreute Vorgänge weisen dann nach derselben Richtung, die
Geschichte.
neue Denkweise, der Geist der Zeit
dem
an
Wirkung, der
eine
Meinungen,
Eingewurzelte
ergreift die verschiedensten Ge-
Weg in die entlegensten Winkel und übt auch
einen
findet
biete,
281
sich
ja
als
einen schroffen Gegner fühlt.
Interessen
selbstische
verlieren
ihre
Macht gegenüber einer solchen Bewegung. — Schwierigkeit macht
daß im nächsten Anblick der Dinge Echtes und Unechtes wirr
nur,
durcheinanderzugehen
pflegt,
des Flachmodernen auch
verneinen
zu
Laune
in
daß
auch
das
für
der
geistige
während
dürfen,
schrittsgedankens
die
der
Abweisung
der
glaubt
Recht
des
Fort-
flüchtigster
Lage
und
andere
das
Erzeugnis
Der Freund des Alten pflegt sich dann
Anspruch nimmt.
der Ordnung, der des Neuen
als Vertreter
mit
eine
Bewegung der Zeiten
als Vertreter
zu fühlen, jener dünkt sich moralisch, dieser
der Freiheit
intellektuell überlegen,
jener glaubt besonders das Interesse der Gesellschaft, dieser das des
Individuums zu wahren.
Dabei
eigentümliche Dialektik.
Was alt ist, war einst neu;
von Aquino
galt
einst
kommen und
Alter
Sache in sich selbst eine
trägt die
auch Thomas
«modern"; was heute neu
als
eines
sich
erwehren
anderen
ist,
wird ins
müssen.
Das
Moderne, das aufstrebt, zieht einen guten Teil seiner Kraft aus seiner
oppositionellen
Stellung;
im Siege büßt
und gerät in
diese ein
es
Nachteil gegen neue Bildungen.
Die Verwicklung, die aus solchem Kampfe hervorgehen kann,
hat in der Gegenwart eine besondere Höhe erreicht.
gegen alles Neue, ihn
Seite findet sich der hartnäckigste Widerstand
vertritt
vornehmlich
eine
Auf der einen
das römische System,
große Weltmacht,
dem Worte nach katholisch nennt, das aber in der Sache
ist.
Denn es setzt allen Eifer daran,
die Bewegung der Menschheit in partikulare Bahnen zu leiten und
das sich
so unkatholisch wie möglich
bei einer mittelalterlichen Gestalt endgültig festzulegen.
aber erscheint
die
weiteste
Verbreitung
neueste Gestaltung des Kulturlebens
schub.
Das Tempo des Lebens
eines
leistet
ihr
Andererseits
Flachmodernen,
die
den vielfachsten Vor-
hat sich in unheimlicher Weise be-
immer mehr Menschen drängen sich in die Groß- und
Weltstädte zusammen; was dort zu Gehör kommen will, muß dreist,
schleunigt,
laut,
ja
schreiend auftreten, er
zu bringen
sich
scheinen.
auf der
Neues,
muß Neues,
dem
«Höhe der Bildung"
sich
fühlt.
Prickelndes, Unerhörtes
keiner entziehen darf,
So
der
ein Sichüberspannen
und Überbieten des Neuen, ein Schätzen des Neuen, bloß weil es neu
ist,
mag es an sich noch so leer oder töricht sein, dabei unendlich
Zu den Problemen des Menschenlebens.
282
viel
Schein, eine
eitler
des Lebens,
Abneigung gegen
keckem Verneinen,
Lust an
eine
elende Talmikultur,
ein
allen Ernst
und alle Tiefen
in
alles
Versuch des Bildungspöbels,
allem eine
die
geistige
Bewegung der Menschheit unter sich zu bringen und sich zum
Richter über gut oder böse, wahr oder unwahr zu machen.
So wird das Moderne echter Art unmöglich vordringen können,
ohne
sich
von
Flachmodernen
derartigem
aufs
zu
allerschärfste
und einen heftigen Kampf dagegen aufzunehmen. Sein
eignes Recht aber können die Verirrungen des Flachmodernen nicht
scheiden
im Mindesten antasten.
Unsere Zeit
samtverlauf unserer Untersuchung,
kommene Bahnen
suchen
hat;
so zeigt es auch der Ge-
eine
solche
und
sie
Selbstvertiefung des
Lage stempelt
schluß an das Alte zu einem bloßen und
tum; für uns
daß
nicht über-
ruhig weiterverfolgen kann, sondern daß
energischer Selbstbesinnung
zu
ist,
so geartet,
gilt
es auf
allen
sie
in
Lebens neue
willfährigen
An-
unfruchtbaren Epigonen-
uns selber zu stehen und den geistigen
einen offnen Weg zu schaffen.
Notwendigkeiten, die
jetzt aufstreben,
Dann wird sich mit
frischer
wart ein
ewiger Wahrheit verbinden lassen, dann wird
Festhalten
und freudiger Ergreifung der Gegen-
das eine das andere zu fördern vermögen.
und Individuum.
Gesellschaft
3.
(Sozialismus.)
a)
Das Verhältnis von Gesellschaft und Individuum.
a.
Geschichtliches.
t^eim Problem der Gesellschaft steht es heute ähnlich wie bei
*-^ dem der Geschichte.
Das 19. Jahrhundert hat einen Rückschlag
gegen die Aufklärung gebracht, dieser aber
voller
hat,
obschon noch
in
Wirkung, einen neuen Rückschlag hervorgerufen; so durch-
kreuzren
sich
Wirkungen mit Gegenwirkungen und
höchst verworrene
Lage;
sich
ihr
7u
entwinden
erzeugen
wird
eine
keineswegs
leicht sein.
Einige Worte seien zunächst den Ausdrücken gewidmet, soweit
sie
Aufklärung fordern.
eine
und
Individuum
Bildungen, aber die Neuzeit erst hat
ältere
sie
Individualität
geführt.
Individuum hieß anfänglich etwas, das sich nicht
trennen
läßt;
(XTOftov.
die
Mittelalter;
spaltig."
so
ist
bei
Dieser Sinn
älteste
Cicero Individuum
sind
weiteren Kreisen zu-
eine
teilen
oder
Übersetzung von
überwiegt wie im späteren Altertum^ so im
deutsche
Übersetzung
(bei
Notker)
ist
Aber schon dem Ausgang des Altertums bedeutet
«un-
indivi-
duum auch das Einzelne als einzigartiges, von anderem unterschiedenes,
in seiner Besonderheit
'
So
z.
B.
nur einmal vorhandenes;- das Mittelalter ver-
Seneca de provid. 5: quaedam separari a quibusdam non
possunt, cohaerent, individua sunt.
'^
Bemerkenswert ist hier namentlich der höchst einflußreiche Boethius,
aus dessen Kommentar tu Porphyrius
(edit. Bas.
1570, pag. 65) folgende Stelle
angeführt sein mag: Individuum autem pluribus dicitur modis.
Dicitur In-
dividuum quod omnino secari non potest, ut unitas vel mens; dicitur Individuum quod ob soliditatem dividi nequit, ut adamas; dicitur Individuum
(üujus praedicatio in reliqua similiü non convenit, ut Socrates: nam cum illi
sunt caeteri homines similes, non convenit proprietas et praedicatio Socratis
in caeteris, ergo ab iis quae de uno tantum praedicantur genus differt, eo
quod de pluribus praedicetur. Bei Porphyrius lautet die Hauptstelle (siehe
Zu den Problemen des Menschenlebens.
284
wendet diese Bedeutung weiter und prägt auch
1 2.
Jahrhundert) die Ausdrücke individualis
(jedenfalls
schon im
Dem
und individualitas.
allgemeinen Leben führt diese erst Leibniz zu, auch hier ein Vermittler alter
und neuer Zeit.
Die Anfänge der Kultur zeigen das Individuum
Stück
als ein
einer engeren oder weiteren Gemeinschaft, das in seinem
Tun und
Lassen durch solchen Zusammenhang wesentlich bestimmt wird.
Die
Weiterbewegung des Lebens pflegt das Individuum mehr und mehr
zu verstärken, es gewinnt an Selbständigkeit, es beginnt nach einem
Recht der überkommenen Ordnung zu fragen und ihre Vernunft zu
dividuum
endlich an einen Punkt,
die Sache gelangt
bezweifeln,
Bindung abzuwerfen sucht und
alle
wo das In-
eigne
die
Meinung
zum Maßstab der Wahrheit, das eigne Wohl zum Ziel des Handelns
Das erscheint solchen, die auf den Stand des Ganzen bedacht
macht.
sind, als eine verderbliche Zerstörung; so wird von ihnen ein kräftiger
Widerstand geleistet und der Versuch unternommen, das Individuum
bei
Zuerkennung
eines
gewissen Rechtes wieder einem Zusammen-
hange einzufügen und für seine Ziele zu gewinnen; geistige Arbeit soll
was als natürlicher Besitz verlören ging. Zu solchem
wiederherstellen,
Zweck der Wiedereinfügung des Individuums ward zunächst der Begriff
dem
des Organismus verwandt, mit
wir uns schon befaßten.
Er scheint besonders geeignet, die Ansprüche des Einzelnen und der
Gemeinschaft glücklich miteinander auszugleichen.
digen Körper
kräftiger
es
aber steht
ist
sich
in
Eigentümlichkeit ausbildet.
seiner
daß
Glieder innerhalb des Ganzen zu verbleiben
Glied aus allem Leben und
aller
vom Ganzen
So
kein
Recht
sich
ablöst.
gegen
des Einzelnen
auf das Weltall
übertragen,
wie
von der Stoa
geschah,
mag
Prantl,
so
Twv xaxä
?xa(JTOv,
(le'po?.
wv to
a'9-potajjia
alle
hat,
Das
Betätigung der
daß das einzelne
Gestaltung herausfällt, sobald es
duldet
organische
diese
Wird
Ganze.
das
es
Fassung
diese
Lehre
Bewußtheit zuerst
mit
voller
ein
besonderer
Vorzug der
Toiaüx«,
oti I? JStoxrJTwv
es
Gesch. der Logik l. 629): axop-a
ouv£(rn|xev
Das Ganze
um so höher, je differenzierter seine Teile sind.
bildet eine unüberschreitbare Grenze,
freilich
Denn im leben-
um so wertvoller für das Ganze, je
jedes Glied
Xe^e-cat
oux av in
-zä
i'XXou Ttvo? izore ro auTo yivoixo
Diese Definition geht durch die Kette der Jahrhunderte bis
zu Leibniz, noch Jakob Thomasius, sein Lehrer, definierte: Individuum
quod
constat ex proprietatibus
esse potest.
quarum
collectio
est
numquam in alio eadem
Gesellschaft und Individuum.
Einrichtung dünken, daß auch
in
285
den kleinsten Dingen nicht zweies
einander völlig gleicht, daß nicht zwei Haare, zwei
Körner, zwei
zusammenstimmen. ^
Blätter gänzlich
Zu dieser organischen Lösung des Problems
aber gesellt sich
eine hierarchische, die, dem Ausgang der griechischen Philosophie ent-
dem Boden des Christentums und im Mittelalter zur
sprungen, ^ auf
vollen Entfaltung gelangt ist und noch heute mächtige Wirkungen übt.
Hier erscheint das All
als eine fortlaufende Stufenfolge,
das Leben von oben her in sicherem Fortgang nach
jede Stelle hat
von der nächsthöheren zu empfangen,
fangene der niederen zuzuführen.
eigentümlichen
Wert und
ein
davon losreißt
Werk, aber
ins Leere, sobald
fällt
mitteilt;
um das Emp-
hier hat jeder Teil
eigentümliches
nur im Gefüge des Ganzen; er
sie
Auch
bei der sich
unten
einen
er
hat
er sich
Dieser Gedankengang hat eine geschichtliche Ver-
körperung nicht nur
in
der kirchlichen Hierarchie, sondern auch
in
der mittelalterlichen Lehnsordnung gefunden, indem hier jede Gewalt
als
von einem Höheren an den Niederen verliehen
gilt.
Die organische wie die hierarchische Fassung lassen dem Indivi-
duum
allen
Wert aus dem Verhältnis zum Ganzen kommen,
Wert für sich selbst wird ihm abgesprochen.
ihm
erst
die Fassung,
die
sich als die
ein
Einen solchen gewährt
mikrokosmische bezeichnen
Hier wird das Individuum aus einem bloßen Stück der Welt
läßt.
zum Ganzen einer Welt, zu einer eigentümlichen Konzentration der
Wirklichkeit, zu einer Stätte, wo die Unendlichkeit des Lebens unDas All wird damit eine Welt von Welten,
Auch dieses
aller umgrenzenden Zusammenfassung.
mittelbar gegenwärtig
es entzieht sich
entstammt
Bild
dem
der dabei voransteht.
ist.
spätesten
Bei
ihm
Altertum,
wiederum
ist
es
Plotin,
zuerst erscheint mit voller Klarheit
der Gedanke, daß jeder Mensch eine eigne Welt bedeute und das All
eigentümlich spiegle,
«jeder einzelne sind wir eine geistige Welt.«
Vom Neuplatonismus aus ist auch der Ausdruck Mikrokosmos, der
bis auf Demokrit und Aristoteles zurückreicht, erst recht in Umgekommen. Im Mittelalter hat namentlich die mystische SpekuGedanken festgehalten, von hier kam er durch verschiedene Zwischenglieder (Nikolaus von Kues, Giordano Bruno) an die
lauf
lation jene
^
S. Cicero acad. quaest. II; dicis nihil idem quod sit aliud; Stoicum
quidem nee admodum credibile, nullum esse pilum omnibus rebus talem,
qualis sit pilus alius, nullum granum etc.
est
'
Am meisten hat dafür Plotin gewirkt.
Zu den Problemen des Menschenlebens.
286
moderne Philosophie,
um in Leibnizens Monadenlehre eine präzisere
gewinnen.
Gestalt zu
In
engem Zusammenhange damit steht die
und Persönlichkeit in unserer
hohe Schätzung von
Individualität
klassischen
So führt auch
Literatur.
hier
fortlaufende
eine
Kette
vom Ausgang des Altertums bis zur Höhe der Neuzeit.
Zwischen der organischen und der hierarchischen Fassung einerder mikrokosmischen andererseits besteht ein voller Gegensatz,
seits,
der einen
Kampf unvermeidlich
ein bloßes Glied, hier
an
ist
es
macht.
Dort
ein selbständiges
ist
das
Individuum
Ganzes, dort hat es
den geistigen Gütern nur durch die Vermittlung einer Gemein-
schaft teil, hier vermag es sie unmittelbar bei sich selbst zu erreichen.
Damit
muß
fallen.
Dort wird zur Hauptsache die Leistung
schaft,
hier
auch
der
Lebens
des
Inhalt
grundverschieden
für
die
aus-
Gemein-
die Entfaltung des Individuums selbst in seiner Inner-
und
lichkeit.
Auch
Kraft erst
von den Individuen empfangen und ihnen nie als Selbst-
die
Gemeinschaft wird hier
zweck entgegentreten dürfen.
ihre
Es wird aber bei solcher Befreiung
das Individuum seinem Leben einen
lich
ihre Gestalt
bedeutenden Inhalt vornehm-
durch ein unmittelbares Verhältnis zur Utiendlichkeit, zu den
sei es, daß sich das mehr
mehr künstlerisch oder mehr religiös gestaltet.
So verschlingt sich mit dem Kampf zwischen Individuum und Ge-
schaffenden Quellen des Lebens, geben,
wissenschaftlich oder
sellschaft
das Problem,
ob wir die Hauptaufgabe unseres Lebens
mehr beim menschlichen Zusammensein oder im
All zu suchen haben,
ob an erster Stelle eine soziale oder eine kos-
mische Lebensführung zu erstreben
Wandel der Zeiten zu
zum
Verhältnis
verfolgen,
ist
ist.
Diese Fragen durch den
hier nicht wohl möglich; wenden
wir uns daher sofort zur Lage und Stimmung der Gegenwart.
Es unterliegt aber unsere Zeit
dem Einfluß dreier Strömungen
von verschiedener Stärke und Breite: es sind das die Gesamtbewegung
der Neuzeit
zum Individuum, der Rückschlag des
19.
Jahrhunderts
zu gunsten der Gesellschaft, die Wiederbelebung des Individualismus
gegen Ende des
19. Jahrhunderts.
Die Emanzipation des Individuums ist wohl der hervorstechendste
Zug des gesamten modernen Lebens. Der Einzelne erstrebte und
gewann dabei sowohl ein unmittelbares Verhältnis zur Gottheit und
zum
All,
Ganzen.
als
eine
selbständige
Stellung
gegenüber dem
sozialen
Allmählich hat sich das von der Renaissance und von der
Reformation aus über das ganze Dasein verbreitet, sich immer
tiefer
Gesellschaft und Individuum.
eingegraben,
darin
es
durchgängig
frischer,
287
bewegter
kraftvoller,
Wie die neue Wissenschaft in ihre aufsteigende Bahn nur
gekommen ist unter Zerlegung der überkommenen Größen, wie Zeit,
Raum, Masse u. s. w., in diskrete Elemente, so ist auch dem modernen
gemacht.
Leben eine wachsende Selbständigkeit und Sonderung der Individuen
Von der Behandlung der
wesentlich.
den Äußerlichkeiten von
Sitte
Fragen
innerlichsten
und Verkehr^
sich
hat
das
bis
zu
immer
mehr durchgesetzt. Es will das keineswegs alle gegenseitige Beziehung
aufheben, aber die Verbindung
werden,
her aufgedrängt
und
freien
den Individuen nicht von draußen
soll
sondern
aus
Vereinbarung hervorgehen.
Individualisierung des Daseins
sammenhänge.
Vielmehr
ist
ihrer
einen Verzicht
auf der
eignen
Entscheidung
Noch weniger bedeutet die
auf alle inneren Zu-
Höhe der geistigen Arbeit, bei
Männern wie Luther und Kant, das Selbständigwerden des Menschen
gegen die Menschen nur die eine Seite des Lebensprozesses, dessen
andere die unbedingte, aber
Gewalten
bildet..
Unterwerfung unter unsichtbare
freie
Wer solche Männer als Verfechter bloßer Willkür
lobt oder tadelt, der zeigt
nur
wie wenig er den Kern der Sache
erfaßt hat.
Im
breiteren
Strome
der Zeit
Verwicklungen und Bedenken.
zum Individuum
seit
ist
die Sache
minder
frei
von
In Deutschland gewann die Bewegung
der Sturm- und Drangzeit vorwiegend einen
künstlerisch-literarischen
Charakter;
wie das
Individuum
jener Zeit
durch künstlerisches Schaffen über den Durchschnitt hinaushob
sich
und als »Genie" allem «Philistertum" weit überlegen fühlte, so hat
sich öfter die selbstbewußte Erhebung des künstlerischen Individuums
wiederholt.
So zunächst
in
der Romantik, welche eben
darin die
Größe des Menschen fand, eine Individualität zu sein (Schleiermacher),
und in Überspannung dieser Denkweise das unumschränkte Recht
der «unendlich freien Subjektivität" verkündete, zugleich geneigt Kunst
und Wissenschaft dem
'^
politischen
Leben weit voranzustellen,- ähn-
Z. B. findet Ihering (Der Zweck im Recht II, S. 43Q) bei der ästhetischen
Gestaltung des gem.einsamen Mahles einen höchst beachtenswerten Fortschritt
der
modernen
Zeit
gegenüber der Vergangenheit
Kommunismus zum Individualismus".
genossen
gemeinsam waren,
in
der
»Erhebung vom
Während früher die Geräte der Tisch-
erhält sie heute jeder für sich
allein
zu aus-
schließlichem Gebrauch.
'
tische
Bezeichnend dafür sind die Worte Fr. Schlegels: „Nicht in die poliWelt verschleudere du Glauben und Liebe, aber in der göttlichen Weil
Zu den Problemen des Menschenlebens,
288
Erscheinungen zeigt später das junge Deutschland, zeigt der
liehe
heutige Individualismus.
Die Hochschätzung des Individuums teilt
unsere klassische Zeit, und die leitenden Pädagogen, Pestalozzi wie
Herbart, übertragen diese Denkweise auf die Erziehung. ^
das Individuum
strebt
nicht
Gegensatz zur Welt und sozialen
in
Aber hier
um in einem
zur Selbständigkeit auf,
Umgebung zu bleiben und
sich
das Bewußtsein stolzer Überlegenheit einzuspinnen, sondern es
dehnt seinen Lebenskreis weiter und
kehrt freudig dorthin zurück,
weiter aus,
wächst schließlich in Versöhnung mit
zur weltumspannenden
So
Persönlichkeit
stellt
aller
es
Umgebung
namentlich die
und das Lebenswerk Goethes vor Augen,
geistige Art
Der erste Widerstand gegen den Vorrang des Individuums ent-
dem
sprang
Idealismus selbst,
indem der Gedanke eines weltum-
spannenden, durch seine eigne Bewegung getriebenen Prozesses den
Schwerpunkt des menschlichen Daseins von den Individuen
Ganze der Menschheit verlegte.
Wendung zur anschaulichen Welt.
seiner
in
das
Dann aber kam der Realismus mit
Damit erschien eine un-
absehbare Fülle von Aufgaben, deren Lösung eine Verbindung der
zerstreuten Kräfte verlangte,
die
Menschen aus der bisherigen Ver-
einzelung heraus zu engerem Zusammenschluß trieb und ein Arbeiten
in
Reih und Glied ihnen auferlegte.
politischer Freiheit,
Dahin wirkte das Verlangen
das Streben nach einer von eigner
Gesinnung der Bürger getragenen Ordnung, dahin
Kraft
und
die Ausbildung
nationaler Kreise, die alle Individuen mit überlegener Art umspannen
und zu großen Aufgaben verbinden, dahin der ungeahnte Aufschwung
der Technik, der die Verkettungen der Arbeit ausdehnt und die
Arbeiter fester zusammenhält, dahin endlich das moderne Wirtschafts-
leben mit seinen Riesenbetrieben, seiner Erzeugimg schroffer Gegenseiner
sätze,
Ansammlung
gewaltiger Massen.
Auch
die
moderne
Beschleunigung des Lebens, das Einandernäherrücken der Menschen,
der Wissenschaft und der Kunst opfere dein Innerstes in den heiligen Feuer-
strom ewiger Bildung".
*
Namentlich Pestalozzi verficht mit großer Energie die Überlegenheit
des Individuellen gegen das bloß Kollektive, er spottet über „Kollektivhand-
lungen",
meint
:
über ein „Kollektivgewissen",
„Die
sieren, sie
über „R^mentsbekenntnisse" und
kann dasselbe nur zivili-
kollektive Existenz unseres Geschlechts
kann es nicht kultivieren" (Wke. XII, 154).
zuerst den Gegensatz
herausgestellt hat.
Hier muß immer der
Ganze des Lebens
von Individuum und Gesellschaft mit voller Klarheit
große Einfluß Rousseau's gegenwärtig
sein,
der für das
Gesellschaft und Individuum.
289
die tausendfache Verschlingung der Lebenskreise trägt viel dazu bei,
die individuellen Züge abzuschleifen und der Summierung zu Massen-
Im Zeitalter
erscheinungen eine überwältigende Macht zu verleihen.
der Telegraphen und
der Presse,
öffentliche
eine
Meinung und gewinnt
umfängt das Individuum schon
sie
eine größere Stärke; sie
seinem Werden und
in
als eignes Werk erscheinen, was
zugeführt
Eisenbahnen bildet sich rascher
in
ihm so
läßt
Umgebung ihm
Wahrheit die
hat.
Endlich steigert, empfangend und zurückwirkend, auch die Theorie
die Abhängigkeit des Individuums.
Denn
die neuere Gesellschafts-
die „Soziologie" (Comte, Quetelet u. s. w.),
lehre,
bemüht,
eifrigst
ist
Umgebung,
die völlige Bedingtheit des Menschen durch seine soziale
das „Milieu",^ zu zeigen; bis in seine Wünsche und Träume scheint
er
durch
ihr
selbst ein
den Bedürfnissen
in
lich
das beherrscht,
des öanzen.
schnitts,
was die Gesellschaft an ihn
heftiger Kampf des Individuums
Zugleich
des
tritt
bringt;
gegen sie wurzelt schließ-
Ganzen und
damit
liegt
innerhalb
der Begriff des gesellschaftlichen Durch-
des mittleren Menschen, in den Vordergrund; es wird nach-
gewiesen, daß die Abweichungen des Individuums, soweit meßbar,
sich innerhalb weit engerer Grenzen
bewegen,
als der erste
Eindruck
mehr
bei der Gleichheit als bei der Verschiedenheit der Individuen,^ und
uns annehmen
Analyse
die
großen
läßt.^
des
Dichter,
individuellen
weicht
Statistik ein handliches
*
Milieu in
So verweilt die Aufmerksamkeit
Seelenlebens,
diese
der Massenbeobachtung,
weit
unserer
Stärke
die
sich
der
in
Werkzeug schafft.
präziser Zuspitzung dürfte zuerst
Philos. zoologique verwandt sein,
von Lamarck
in
seiner
Comte hat es von der Zoologie der Qesell-
schaftslehre zugeführt, Taine aber es hier «mit besonderer Vorliebe verwandt.
von ihm aus ist es in Deutschland zu einem Modewort geworden.
Erst
^
Hierfür sei namentlich Quetelets „Anthropometrie" erwähnt.
*
Der Gedanke der Gleichheit samt dem des gleichen Wertes und
Rechtes aller Menschen hat ältere Wurzeln, zur vollen Entwicklung aber ist
er erst in
den
letzten
Jahrhunderten gelangt.
Dem klassischen Altertum ist
und auch was sich zu seinen Gunsten im späteren Altertum regte,
kam gegen die tatsächlichen Unterschiede der Menschen nicht auf. Die Wurzel
er fremd,
der Gleichheitsidee liegt in der Religion, für unseren Kulturkreis im Christen-
tum.
Es war das Verhältnis zu Gott,
in
dem alle Abstände der Menschen
verschwanden, es war die Unendlichkeitsidee, der gegenüber alle endlichen
Unterschiede gleichgültig wurden. Aber Konsequenzen für das irdische Dasein
wurden daraus zunächst recht wenig gezogen, und in der weiteren Ge-
schichte des Christentums trat der Gedanke des allgemeinen Priestertums weit
Eucken, Grundbegriffe.
4. Aufl.
19
Zu den Problemen des Menschenlebens.
290
Das
auch
alles
eine
war nicht bloß eine äußere Verschiebung,
innere
es
war
Denn nunmehr wurde
Wandlung des Lebens.
zur Hauptsache an ihm, was wir für die Gemeinschaft leisten, nicht
was wir im eignen Bereiche denken und
alle
Kraft
zur Betätigung aufgerufen,
Energischer wird
tun.
deutlicher
die
Bindung des
Individuums an das Ganze hervorgekehrt.
Auch
eine eigentümliche
davon
aus.
Die Verbesserung
Gestaltung
des
Geisteslebens
geht
der gesellschaftlichen Lage wird das allüberragende Ziel.
Die Moral
zum Wirken für die Gesellschaft, zum Altruismus,
die Kunst
wird
findet keine höhere Aufgabe als die eindringliche Vergegenwärtigung
der
gesellschaftlichen
Hebung
Zustände,
die
Erziehung
des gemeinsamen Bildungsstandes
dividueller Art.
Einen
Zusammenhalt
gibt
mehr
die
Entwicklung
in-
erstrebt
als die
dem Individuum
hier
namentlich die Arbeit, die Arbeit, welche weite Komplexe und feste
Methoden
entwickelt, damit aber stark
genug wird,
um einen Kampf
mit aller Unvernunft des Daseins aufzunehmen und die Bedingungen
des Daseins wesentlich zu
verbessern.
Daß das Ja auch ein Nein
sich trug, der Gewinn von einem Verluste
kam einstweilen kaum zur Empfindung.
in
begleitet war,
das
Wirkt solches Zusammenstreben der Kräfte mit seinen engeren
Verschlingungen gegen das Individuum
tut es offensichtlich die
des 19. Jahrhunderts
in
versteckterer Weise,
so
mächtige Verstärkung, die der Staat im Lauf
erlangt
zurück hinter dem der Hierarchie.
hat.
Am zwingendsten trieben dazu
Was von einzelnen Nebenströmungen das
Mittelalter mühsam genug bewahrt hatte, das gelangte zu vollerem Durchbruch
in
der Reformation, und es war namentlich ihr calvinistischer Zweig,
der
daraus energische Folgerungen für die Gestaltung des Gemeindelebens zog.
Von hier aus fand sich auch der Übergang in das politische Gebiet: unter
Cromwell zuerst enthält ein Verfassungsentwurf das Verlangen des allgemeinen
Stimmrechts (1647). Weiter wirkt dann für die Idee der Gleichheit die Aufklärung mit ihrer Berufung auf die allen Menschen gleiche Veniunft. So
sagt z. B. Descartes (de methodo zu Anfang): Rationem quod attinet, quia
per illam solam homines sumus, aequalem in omnibus esse facile credo.
Rousseau hat endlich mit besonderer Energie den Gedanken der Gleichheit
ins allgemeine Leben geworfen; der Gedanke der Menschenrechte dürfte aus
Amerika stammen. Die Formel von der „Gleichheit alles dessen, was MenschenDas
gesicht trägt", hat Fichte aufgebracht, s. z. B. IV, 423, VII, 673.
18. Jahrhundert bringt auch die Zusammenstellung von Freiheit und Gleichheit, und zwar wohl zuerst für das Gebiet des geselligen Verkehrs.
So sagt
z. B. schon Montesquieu in seinen lettres Persanes (erschienen zuerst 1721),
—
Bch.
II:
A Paris regne la liberte et l'egalite.
Gesellschaft und Individuum.
wirtschaftlichen
die
strengung
Entwicklungen,
da ihnen
Individuums
verloren
des
bloßen
der Höhepunkt
291
gegenüber
An-
alle
Aber dieser
dünkte.
durchgehenden Erscheinung.
Punkt
ist
nur
Es
die
wachsende Komplikation, die technischere Gestaltung der
ist
einer
mehr Ineinandergreifen der einzelnen Kräfte und
mehr Organisation des Ganzen verlangt, damit aber nach einer leitenden Spitze ruft. Das erzeugte z. B. mit Notwendigkeit eine stärkere
Zentralisation im Unterrichtswesen. Und es fehlte dieser Bewegung
welche
Kultur,
des
Kulturlebens
spricht
zum
Staates
einer
Kraft
Gedankenwelt.
Hauptträger der Kulturarbeit ent-
der modernen Überzeugung von einem Innewohnen abso-
Vernunft
luter
beseelende
die
nicht
Die Erhebung des
leitenden
unserer Wirklichkeit;
in
Systematiker
es
Pantheismus,
des
kein Zufall,
ist
Spinoza
daß die
und Hegel,
ent-
schiedenste Vorkämpfer der Staatsidee waren, daß Spinoza nicht bei
sondern beim Heil des Vaterlandes geschworen haben wollte,
Gott,
Hegel 'aber den Staat »wie ein Irdisch-Göttliches" verehrte.
bünden
Macht des
So ver-
gegen die Selbständigkeit des Individuums die sichtbare
sich
und
Staates
die
unsichtbare
der Gesellschaft;
einen entflieht oder zu entfliehen glaubt, verfällt leicht
wer der
um so mehr
der anderen.
Aber wie der
volle'
einen Rückschlag ergibt,
Menschen durch
Sieg leicht eine Überspannung und damit
hunderts eine Neuerhebung des Individuums
sich
davon
die
Selbstvergötterung
subjektiver
hervordrängt,
geflissen
unechter
Stimmung zu
ist
vermeintlicher
Denn
ist
Was
hervorgerufen.
unerquicklich genug,
oft
Genies
mit der Verspottung jener Auswüchse
ledigt.
Umklammerung des
so hat auch hier die
und Gesellschaft gegen Ausgang des 19. Jahr-
Staat
so
und das Sichaufbauschen
Aber
Weltüberlegenheit.
nicht schon die Sache er-
hinter allem Problematischen
steckt eine
Gegenwehr
des Individuums und Subjekts gegen die drohende Einengung und Ver-
kümmerung; was jene Bewegung zur Gesellschaft an Begrenzungen
und Verneinungen
enthält,
lichem Bewußtsein.
das bringt der Widerspruch jetzt zu deut-
Eine Abschleifung der individuellen Züge, eine
Gefährdung der Selbständigkeit,
und Schaffens,
verbunden.
drückte,
sie
Ähnlich
eine kleine
Großes mehr
sieht,
der Individuen
ein Stocken ursprünglichen
Lebens
scheinen mit jener Gesellschaftskultur untrennbar
wie
die
Geschichte
Gegenwart aber auch
die
in
Gegenwart
unter-
der Geschichte nichts
so scheint die Gesellschaft mit der Verkleinerung
unvermeidlich auch bei sich selbst zu sinken.
19»
Ge-
Zu den Problemen des Menschenlebens.
292
wahren wir nicht deutlich genug, wie inmitten
glänzenden
aller
Triumphe technischer Arbeit uns ausgeprägte Persönlichkeiten mehr
und mehr entschwinden, zugleich aber das Niveau des gemeinsamen
Lebens niedriger wird?
gestaltung,
daß
muß
das
und
sie
den
Zugleich sucht das Individuum sich
lassen.
Bindung möglichst abzulösen,
und gänzlich „ausleben",
entfalten
viel
von der
gesellschaft-
es will sich mit voller Freiheit
es
kehrt
sein
Unterscheidendes
vom Durchschnitt irgend abzuheben.
hervor und strebt sich
Wie
schwächt, ja unter-
Gegenwehr reizen, sie der GesellWert ihrer Erfolge bestreiten
die Seele zur
widerstehen
schaftskultur
lichen
nun kommt zur Empfindung,
mit ihrer riesenhaften Entwicklung
sie
drückt;
Die Arbeit, der Kern der neuen Lebens-
die Seele kräftigen;
sollte
dem allen überspannt und verkehrt sein mag, es
Zeit; mag es arm an positiver Leistung
in
hat eine
Macht über unsere
sein, in
der Kritik
und den Glauben an
es stark,
ist
die Allgenug-
samkeit einer bloßgesellschaftlichen Kultur hat es schwer erschüttert.
Aber trotz solcher Erschütterung geht die Arbeit mit ihrer Richtung
auf den Stand
der Gesellschaft
Individuum und mehr noch
fort,
so wächst ihr Druck auf das
unsere Empfindung dafür.
Demnach
werden wir nach widerstreitenden Richtungen gezogen: die gesellKultur
schaftliche
behei"rscht
verlangt
unsere Seele.
ergeben,
oder
Arbeit,
ihr widerstehen
sich
läßt
unsere
eine
Individualkultur
Müssen wir uns solcher Spaltung wehrlos
und nach irgendwelcher
Einheit des Lebens streben?
Die Probleme der Gegenwart.
ß.
Die Unzulänglichkeit einer bloßgesellschaftlichen Kultur.
aa.
Etwas
anderes
ist
die
es,
Bedeutung einer
gesellschaftlichen
Kultur anzuerkennen, etwas anderes, in sie das ganze Dasein des
Menschen zu
setzen.
Für das
Mannigfachste zusammen.
erstere
wirkt in
unserer Zeit das
Deutlich steht uns vor Augen, wie von
Anfang an der Mensch die eigentümlichen Züge seines Wesens nur
in
der Gemeinschaft entwickeln konnte, wie auch später
finden wesentlich
deutlich
auch,
wie die Wirkung des Zusammenseins weit
das Leben des Einzelnen und den
als
man
früher
anzunehmen
schaftliches Wesen
funden.
alles
Be-
von der Gestaltung des Zusammenseins abhing,
ist,
Grund
Daß der Mensch
pflegte.
tiefer in
seiner Seele hineinreicht,
ein gesell-
das hat erst jetzt seine volle Anerkennung ge-
Aus den neuen Einsichten aber erwachsen sofort Aufgaben
Gesellschaft und Individuum.
Art
fruchtbarster
293
Sind wir so sehr auf die Gesellschaft angewiesen
und hängt unser Glück so sehr an ihrem Gedeihen, so wird
es be-
sonders wichtig, den Stand der Gesellschaft zu heben und
alle
ihr
vorhandene Kraft zur vollen Wirkung zu bringen.
Zusammenschluß
die Menschheit
hat
im Kampf gegen die Unver-
vordringen laäsen und mehr Glück
nunft unaufhaltsam
sein gebracht, die straffere Organisation
das
Handeln
Hebel
bei
fälligkeit
hat
festere
in
Der engere
in
ihr
Da-
hat jedes Einzelne gehoben,
Angriffspunkte gewonnen,
den allgemeinen Verhältnissen
ansetzt,
indem
es
den
nicht auf die
Zu-
der bloßen Individuen angewiesen bleibt.
Die engere Ver-
bindung im unmittelbaren Zusammensein hat reiche Quellen moralischer Gesinnung erschlossen, die Teilnahme für einander gesteigert,
ein
Bewußtsein
durchgängiger
Miteinanderarbeiten,
Notwendigkeit,
die
mehr
und zu einander zu fügen,
Mannheit und Kraft
in
Solidarität
erzeugt.
sich
Disziplin
Auch
hat
das
gegenseitig zu
halten
und zugleich
mehr
das Leben gebracht, das in der Vereinzelung
leicht verweichlicht.
War es bei solchen Erfolgen ein Wunder, daß die Hoffnungen
und Gedanken
die
tatsächliche Leistung
soviel geleistet hatte, alles leisten
Indem
und
sie
daß was
Lebensführung das ganze Dasein des Menschen aus-
sellschaftliche
zufüllen
weit überflogen,
zu können vermeinte, daß die ge-
alle
Wünsche zu
das versuchte, hat
befriedigen
sie allen
eigentümliche Gestalt gegeben.
sich
zutrauen
konnte.
einzelnen Lebensgebieten eine
Zum Inhalt der Ethik wird hier die
Umgebung, der Altruismus, zum Ziel der
Leistung für die soziale
Erziehung die Bildung des Einzelnen für die Zwecke der Gesellschaft,
die Kunst macht die gesellschaftlichen Zustände
vorwurf ihrer Arbeit und
die Wissenschaft sucht
sondern
will
zum Haupt-
mit ihr den weitesten Kreisen dienen,
den Menschen nicht als isoliertes Individuum,
vom Ganzen der Gesellschaft her «sozialpsychologisch" zu
zum Maßstab der Wahrheit macht der Pragmatis-
verstehen; selbst
mus die Leistung für das Wohlbefinden der Menschheit. Indem in
dem allen das Leben und Handeln direkter auf den lebendigen und
empfindenden Menschen
als
Ganzes bezogen wird, scheint es eine
größere seelische Nähe zu gewinnen, wird es mehr ins Unmittelbare,
Frische,
liegen
je
ja,
hier
so scheint
alle
unsicherer
es.
Wahrhaftige
gestaltet;
im Hintergrunde
Verwicklungen der Religion wie der Metaphysik;
die
Neuzeit
über
diese
Fragen wurde,
kommener muß ihr solche Befreiung sein.
desto
will-
Zu den Problemen des Menschenlebens.
294
Aber mag diese Bewegung noch so fruchtbare Ausblicke und
Aufgaben eröffnen, nur so lange kann
Zustimmung rechnen,
zur Seite
vertretenen Ja
ihr
Das Leben
stimmter Art.
gänzlich
geht;
läßt
ist
dem von
sehr
be-
wie es dort geschieht,
nicht,
Umgebung,
das Verhältnis zur
in
das
bleibt,
Nein aber
dieses
sich
und freudige
auf volle
sie
das Nein unbeachtet
als
die Entwicklung der
in
gegenseitigen Beziehungen verlegen, ohne daß zugleich die Selbständig-
ist
Nun aber
wo geistiges Leben ursprüng-
des einzelnen Punktes aufs stärkste herabgesetzt wird.
keit
das Individuum die einzige
während das
aufquillt,
lich
mehr
Zusammensein
nicht
Die
und
gesellschaftliche
und verwerten kann.
verbinden
als
Stelle,
Freihaltung
Kräftigung dieses ursprünglichen Lebens wäre nun minder bedeut-
sam und eine Einengung ließe sich eher ertragen, wenn das menschliche Leben auf fester Grundlage stünde und nur eine naturgewiesene
Richtung ruhig zu verfolgen brauchte.
Wahrheit steckt es nicht
In
nur voller Probleme im Einzelnen, sondern, zwischen bloßer Natur
und
Welt befindlich, hat es eine durchgehende Wendung,
geistiger
einen Aufstieg von halbgeistiger zu echtgeistiger Lebensführung erst
zu
vollziehen;
so
große Entscheidungen
liegen
ohne eine Aufrüttelung und
standes
können,
erfolgen
Tiefe zurückgreifen
Die
kräftige
die
notwendig
auf
die nicht
ursprüngliche
eine
und damit auf das Individuum kommen.
Lebensführung dagegen
soziale
ihm,
in
Belebung des gesamten Seelen-
vorwiegend auf eine
ist
sie hebt und fördert,
wenn sie das Leben angenehmer und
Verbesserung der äußeren Verhältnisse gerichtet,
sie
und
mildert
freundlicher
macht,
sinkt
alle
geistige
Be-
wo sie nicht als völliger Selbstzweck behandelt wird;
der
behandelt.
Feind
aller
Leben
als
ein
Mittel
Denn unvermeidlich
Form er auch annehmen mag, ist ein unversöhn-
echten Geisteskultur.
für
den Menschen
erlangen,
nie
Als ein bloßes Mittel kann
bei
aller
innere
die
Daher kann auf diesem
äußeren Erweiterung keine innere Erhöhung des
Menschen
erfolgen,
Verhältnis
zum All, alle innere Selbständigkeit.
hier
fehlt
alle
Urerzeugung,
kann nie etwas wesentlich Neues bringen,
an
denen
sich
Notwendigkeit
mit solcher die Seele ergreifen und
zu ursprünglichem Schaffen treiben.
Wege
den
menschliche
einer Selbsterhaltung
sie
sie
das
Utilitarismus, welche
licher
indem
wirkt sie zerstörend,
Lebens
für
Wohlsein
geistiges
am Kern
des
Gehalt
geistigen
tätigung,
aber
glättet,
das
Dasein
nie
alles
unmittelbare
Ein derartiges Leben
hohe Ziele weisen,
heben könnte, sondern es bannt den
Gesellschaft und Individuum.
295
Menschen an seine eigne Zuständlichkeit und macht ihn zum Sklaven
selbst, es läßt ihn sein Dasein zieren und schmücken, aber
seiner
Niederem,
es
vermag daher
des
Durchschnitts
und
Geist,
Scheidung
gründliche
keine
vollzieht
es
nicht
den Menschen aus der Trägheit
und Allgemeingültigem
von Kleinmenschlichem
entgegenzuwirken,
die
das
Vermengung von Natur
der
nicht
aufzurütteln,
Höherem und
zwischen
gewöhnliche
Dasein
kräftig
kennzeichnet;
bei
unermeßlicher Emsigkeit und Betriebsamkeit fehlt diesem Leben der
rechte Tatcharakter, fehlt ihm ein unerbittliches Entweder - oder,
Dies Leben der bloßen
ein wahrhaftiger Gehalt und Sinn.
ihm
fehlt
mag erträglich scheinen, solange der Blick und
das Streben nur vom Einzelnen aufs Einzelne geht und von der
bunten Fülle wechselnder Anregungen eingenommen wird. Aber man
denke darüber hinaus und frage nach dem Ertrage des Ganzen, dann
muß die Dürftigkeit, die Leere dieses Lebens augenscheinlich werden.
Menschenkultur
Wenn die Sozialkultur solcher innern Leere entrinnen zu können
so pflegt sie das in der
glauhjt,
bindung der Elemente
in
der Vereinzelung vorliegt;
Einzelnen
das
des
Durcheinander
der
sellschaft
öffentlichen
Überzeugung zu
daß die Ver-
tun,
etwas wesentlich Höheres entstehen läßt als
so scheint
individuellen
Meinung sich
zu
weit
B.
z.
Meinungen
ein Träger
das
überragen,
Wohl
der Ge-
dem
so scheint
in
der
bilden.
In
gegenüber
der Wahrheit zu
Wirklichkeit kommt der Schein einer inneren Erhöhung nur zu
Stande, indem aus andersartigen Zusammenhängen Neues zugeführt
wird; aus dem bloßen Durch- und Nebeneinander könnte nun und
nimmer eine neue Stufe des Lebens hervorgehen; zu Unrecht wird
hier,
wie es
tives
unvermerkt
Ziel
als
in
freilich
liegt,
Gibt es
umgesetzt.
Qualitatives
in
das der
der Neigung der Gegenwart
natürlichen Selbsterhaltung,
kein
Quantita-
anderes
gibt es keine geistige
Wesensbildung, so kann auch die Verflechtung der einzelnen Kreise
im
Zusammensein
gesellschaftlichen
stehen
lassen,
auch
bei größter
nichts
wesentlich
Neues
ent-
Ausdehnung nähert sich das Nütz-
und Angenehme der natürlichen Lebensstufe in keiner Weise
dem Guten. Ebensowenig ergibt die Bildung gewisser Durchschnitte
der Meinungen, mögen sie sich noch so festlegen und noch so
liche
selbstbewußt auftreten,
griff
einer
echten
nicht die
Wahrheit,
die
mindeste Annäherung an den Bealles
menschliche Streben
Gutes
und Wahres werden immer schon
glaubt,
sie
aus
der Verbindung
vorausgesetzt,
mißt.
wo man
der Elemente ableiten zu können.
Zu den Problemen des Menschenlebens.
296
Solche Überzeugung zwingt zu starker Skepsis gegenüber der
bekannten Lehre von einer Summierung der Vernunft
Sie
sellschaft.
ist
philosophisch
zuerst
von
der Ge-
in
vertreten.
Aristoteles^
Die Gesamtheit erscheint ihm befähigter zu politischem wie zu künstder andere
lerischem Urteil als die Individuen, weil der eine dies,
anderes besser beurteile, im Zusammentreten aber eine gewisse Aus-
dem Zorn
gleichung erfolge; auch scheint ihm das Ganze minder
und anderen Affekten unterworfen als das Individuum.
den begrenzten Kreis eines Stadtstaates, den gemeinsame
er aber an
Ordnungen auch innerlich zusammenhalten,
und unübersehbare Masse. So bleibt er
Überlieferungen
und
nicht
an
jede
beliebige
auch
in
seinem Demokratismus
weit geschieden.
— Für
dern durch
das
eines
die Sache spricht zunächst die
daß hervorragende
Rousseau
alte,
große Publikum
zur Anerkennung
auch
literarische
nicht durch das Urteil der Techniker,
Leistungen gewöhnlich
gelangt
sonsind,
wegen eines moralischen Defektes jener, sondern weil sie sich
nicht
fest in
Die
vom Volksglauben
gegenwärtige Erfahrung,
Aristoteles
zu
Dabei denkt
einen geschlossenen Gedankenkreis eingesponnen hatten.
Unbefangenheit
weiterer
Bildung.
— Weiter
Kreise
wiegt
namentlich
ungewöhn-
gegenüber hier mehr als die höhere technische
lichen Erscheinungen
steht in einer gewissen Verwandtschaft mit jener
Lehre von der Summierung der Vernunft die Überzeugung, daß es
eine Berufung
von der Zufälligkeit der Augenblicke und der
im besonderen auch
viduen,
Ganze der Menschheit
von der Enge der Parteien,
das Vertrauen
gibt,
Indi-
an das
auf irgendwelchen Sieg
des Guten auch innerhalb des menschlichen Kreises.
Ohne solchen
Glauben müßte
Wirken nach
ja,
außen als zwecklos
das
lich
wer
in
der Minderheit
ist,
einstellen; so durchdringt jener Glaube
politische
Auch
Streben.
liefert
anfänglicher schroffer Verfolgung,
es,
minder
als
festgelegt
Er-
zum Siege kam
trotz
der Stein, den die Bauleute ver-
oft als ein Eckstein
das dazu verhalf
sich
nament-
geschichtliche
die
fahrung Zeugnisse dafür genug, daß Großes
warfen, hat sich
alles
erwiesen; was anders aber war
das größere Ganze, die weiteren Kreise, die
hatten
und neuen Anregungen zugänglicher
waren? Aber solches Durchdringen des Wahren vollzog sich kaum
durch eine bloße Summierung menschlicher Meinungen, sondern
dem Zwange
unter
*
S. 62 ff.
einer
S. Pol. 1281b, 8, 34.
geistigen
Näheres
s.
Notwendigkeit,
in
welche
jenes
meinen gesammelten Aufsätzen
Gesellschaft und Individuum.
Höhere immer
So
machte.
ist
abhob und
deutlicher
297
es schließlich unwiderstehlich
Glaube an die Masse, sondern an eine
es nicht der
innerhalb der Menschheit waltende geistige Notwendigkeit,
welcher
Hoffnung auf einen Sieg der Vernunft auch im Bereich des
Menschen rechtfertigt. Erst in Berührung mit einer solchen geisti-
jene
gen Notwendigkeit und
Meinung
ein
kann
leicht hinter
sie
als
Recht
gutes
ihre Vertreterin
und eine
gewinnt die öffentliche
sichere
Überlegenheit;
sonst
dem Stande der einzelnen Individuen zurück-
und weniger zur Vernunft als zur Unvernunft wirken. Es
Zeiten, wo der Durchschnitt den Menschen erhöht, es gibt
bleiben
gibt
aber auch solche,
Masse
wo er ihn herabdrückt.
Jedenfalls tut die bloße
es nicht.
Dasein
und gar in das unmittelbare
die Sozialkultur sich ganz
Indem
hineinstellt,
wird
zum Haupt-
unvermeidlich die Masse
sie
träger des Lebens machen, wird sie wohl oder übel der Art huldigen,
mit der jene die großen Kulturfragen betreibt: hastig und aufgeregt,
maßlos im Ja wie im Nein, haftend
lichst starke
geworfen,
am sinnfälligen Eindruck, mög-
Erregung suchend, zwischen Gegensätzen hin- und her-
aller
Zugleich
Besonnenheit und Gerechtigkeit abgeneigt.
wird das Individuum mehr und mehr zurückgestellt; selbst wo es
unbestreitbar Großes wirkt, gilt es hier als ein bloßes Werkzeug
der Gesellschaft,!
als
gleichgültig
allem,
in
was
an
es
Eigentüm-
Nun hat gewiß auch die größte Leistung ihre geschicht-
lichem
hat.
lichen
und gesellschaftlichen Bedingungen und Zusammenhänge,
alles
Atmosphäre
her-
Schaffen
aus,
es
wächst aus
damit
trägt
einer
besonderen
unvermeidlich
geistigen
Färbung seiner
die
Zeit;
ein
Augustin wäre nicht möglich zur Zeit Kants noch ein Kam zur Zeit
Augustins,
ein
heißt
nicht
das Ganze für
und das Individuum zu einem
gleichgültigen Werkzeuge machen.
erklären
Kraft
nach völlig
Aber solche
Goethe nicht inmitten der Kreuzzüge.
Bedingtheit anerkennen
die erzeugende
seiner Besonderheit
Bei
Zusammenhang stand das Große zum Durchschnitt
allem
inneren
seiner Zeit ge-
wöhnlich im Verhältnis des Gegensatzes, meist hat es seine Größe in der
Art entu'ickelt, daß es eine Notwendigkeit seines eignen Wesens dem
Ganzen der
Zeit
gegenüber siegreich
durchzusetzen wußte,
sieg-
reich nicht in der Breite des Daseins, aber siegreich in der Sphäre
^
S. z. B.
Comte, cours de
phil. pos.
IV, 269: Les
ne se presentaient essen tiellement que conime
les
hommes de genie
organes d'un niouvement
predetermine, qui, ä leur defaute, se tut ouvert d'autres issues.
Zu den Problemen des Menschenlebens.
298
Was dabei seine Leistung auszeichnet, ist vor
der geistigen Arbeit.
allem das Individuelle, Unvergleichliche
ward
Hilfe dessen
und aufquoll, das aber
regte
und Unableitbare.
möglich, das Geistige, das
es
in
ihrer
in
Nur mit
der Zeit sich
mit Niederem
Breite
und
Fremdartigem untrennbar zusammenfloß, davon abzulösen und zu
wie kräftiger Gestalt zu bringen,
klarer
und erhöhenden Kraft zu erheben.
welche
Individualisierung,
eine
eigentümliche Bahnen
trieb.
zu einer aufrüttelnden
es
Dabei erfuhr das Geistige selbst
Geschicke
die
Nirgends
ist
Menschheit
der
Denn das steht wohl außer Zweifel, daß ein
Gebiete der Religion.
Augustin und ein Luther nicht bloß zusammenfaßten, was die
gebung ihnen
darbot,
sondern daß
Lage
gleich
geistige Art
haben.
in
sie
die
Um-
Probleme der weltge-
durchaus eigentümlicher Weise gelöst und zu-
schichtlichen
ihre
in
das deutlicher als auf dem
ganzen Jahrhunderten zwingend auferlegt
Jede Zeit von kräftigerer geistiger
Regung enthält verschie-
dene Möglichkeiten; welche von ihnen zur Wirklichkeit wird, das
Schon das verbietet
hängt vor allem an den führenden Individuen.
eine Konstruktion der Geschichte aus einer Formel.
War das Große einmal da, so konnte es alles irgend Entgegenkommende an sich ziehen, Aufstrebendes verstärken, Zerstreutes verGesamtbewegung erzeugen. Aber es war dabei nicht
der Summierung, vielmehr hat es seinerseits die
Summierung erst möglich gemacht. Denn die Summierung, das
binden, eine
Ergebnis
ein
Sichzusammenfinden, das den Vertretern der gesellschaftlichen Kultur
so leicht erscheint,
als
blem.
In
liegen
und
ist
in
Wahrheit ein überaus schweres Pro-
einer Zeit kann viel Verschiedenes,
eine
Summierung
in
sehr abweichender Höhenlage möglich
in
an einzelnen Stellen
und
ist
aufstrebt,
nicht
sein;
Richtung,
zusammen
Daß die Verbindung der
gelingen
will,
das kann
einem schweren Druck belasten; ein solcher Druck
eignen Zeit.
Das eben
ist
das
auch
was an Tüchtigem
das findet sich oft nicht
daher für das Ganze wie verloren.
aufstrebenden Kräfte
Widersprechendes
ja
mannigfacher
eine
liegt
Zeit
mit
auf unserer
Werk der Großen, durch glückliche
Ausprägung eines geistigen Charakters und mutiges Vordringen eine
Summierung in bestimmter und erhöhender Richtung anzubahnen
und durchzusetzen; so waren sie die Herren, nicht die Diener der
Zeit.
des
in
Sprechen wir von einer Goethezeit, weil
18.
Jahrhunderts
eine
humane und
in
der zweiten Hälfte
künstlerische
Denkweise
der Art Goethes massenhaft verbreitet war, oder weil seine über-
Gesellschaft und Individuum.
299
ragende Persönlichkeit Gestalten schuf und Ziele vorhielt, an denen
minder Ausgeprägtes
sich
die
in
Höhe hob und
zugleich mitein-
ander zusammenfand?
Wenn dem gegenüber die Sozialkultur die Abstufungen zurückstellt
und nach möglichster Gleichheit strebt, so
sicht
der Besten,
möglichst
alle
das Gesamtniveau
die
steigern,
möglichst
viele,
auf die Höhe zu führen, ohne daß diese irgendwelche
Minderung erfährt.
als
zu
gewiß die Ab-
ist
Absicht
Aber auch
hier
ist
die Natur der Dinge stärker
Unvermerkt wird der Stand des
Menschen.
(ier
Maß der geistigen Bewegung, und es sinkt
damit unvermeidlich die Höhe des Ganzen, es läßt sich die Arbeit
Aufnehmenden zum
vorwiegend auf die Wirkung bei anderen
nicht
sie
Einbuße
eine
Denker
in
sich
bei
selbst
ohne daß
richten,
Schopenhauer
erfährt.
teilte
die
solche ein, die für andere, und solche, die für sich selber
denken, und erklärte nur diese für Denker echter Art;
hat.
er damit
Recht, wie wir meinen, so kann über die Gefahr einer vorwiegend auf
Mitteilung
und Wirkung
aus der Verbreiterung
gerichteten
muß
Arbeit
keinerlei
eine Verflachung
werden,
Urerzeugung erfolgt, welche jener die Wage
eine
Dazu
gesellt sich
anzunehmen, was
auf
lichst
sein;
wenn
nicht
hält.
die Neigung, sich nicht nur des Schwächeren
und
ist,
sondern sich mög-
seinen Standort zu versetzen und das
Ganze des Lebens
sicherlich
edel
nach seinen Interessen einzurichten.
weise
Zweifel
recht
Zeiten harter und weicher Denk-
miteinander abzuwechseln,
pflegen
heute herrscht unzweifel-
weiche Art und erzeugt die Neigung, den Schwachen
haft
die
gut,
den Starken
als schlecht
als
zu denken, daher beim Zusammenstoß
jenem ohne weiteres Recht zu geben.
So nimmt eine weitverbreitete
Zeitströmung Partei für die Kinder gegen die Eltern, für die Schüler
gegen den
selbstischer
die
als
und
finden.
auf
Auch
sei
alle Ordnung
und
brutaler Gesinnung.
Gerechtigkeit
Menschen
durchgängig für den Untergebenen gegen den
Lehrer,
Gebietenden,
untergeht,
Erden
leben"
der drohende
so
hat
würde
alle Strenge
Der
keinen
es
hier
nur ein Ausfluß
kantische
Satz:
Wert
schwerlich
«Wenn
mehr,
daß
Zustimmung
Feminismus hängt damit zusammen,
der nicht nur den Frauen zu ihrem gebührenden Recht verhelfen,
sondern die Erziehung wie die ganze Kultur möglichst ihren Interessen
gemäß
»Coeducation"
gestalten
möchte.
So wird
die höchst problematische
vornehmlich deshalb empfohlen,
nur ja genau so
viel
damit die Frauen
und genau dasselbe erhalten wie die Männer!
Zu den Problemen des Menschenlebens.
300
Auf solchem Wege der Qleichmacherei muß die Kultur unvermeidlich ins' Welke und Matte geraten, sie wird vor aller kräftigen Art
und aller ausgeprägten
recht zurückscheuen,
Individualität wie vor
einem Übel und Un-
was noch schwerer wiegt, das
wird,
sie
was nach Ooethe's Wort »niemand mit auf die Welt
lieren,
und worauf doch
Seiten zu ein
ankommt,
alles
Mensch sei": die
Bewegungen
Derartige
eine Zeit lang ertragen, der
der Mensch
damit
nach
in die Breite und Fläche lassen sich
überkommene Lebensstand liefert einst-
Aber
zuweisen;
erschöpft
schließlich
die Frage der
reichste Kapital,
allen
Ehrfurcht.^
vom er-
weilen eine Ergänzung, für einige Zeit läßt sich ganz wohl
erbten Kapitale zehren.
ver-
bringt,
sich
auch das
Urerzeugung ist nicht dauernd ab-
sobald sie aber erscheint, werden die Schranken der ge-
sellschaftlichen
Kultur
Die gesellschaftliche
unverkennbar.
konnte das Geistesleben auf den Menschen nicht
stellen,
Kultur
ohne ihn
innerlich zu erhöhen; sie konnte der Gesellschaft die höchsten Güter
ohne mehr aus ihr zu machen.
nicht anvertrauen,
Aber mit eignen
Erhöhung nicht bewirken, vielmehr
zerstört sie mit der Schwächung und Stagnation des Geisteslebens
die Bedingungen echter Größe und kann daher nicht verhindern,
daß eine bloße Menschen- und Massenkultur eine wesenhafie Geisteskann
Mitteln
kultur überflutet
Treten
eine
sie
solche
und unterdrückt
schon
nicht
schmerzlich hervor?
und im Ganzen
heute
sehen,
deutlich
und
wie hervorragende Geschichtsschreiber uns
frühere Zeiten sehen lassen,
Außenkultur ein
einer
Erfahrungen
solche
Würden wir unsere eigne Zeit von innen her
so möchte sich
bewegendes
inmitten
vor Augen
Bild
alles
Glanzes
stellen.
Die
Menschheit wollte durch einen engeren Zusammenschluß und eine
Entfaltung der Kräfte das Leben
vollere
genug,
stark
Wirken
alles
Geistesleben
wollte sie einen
selbst
erhöhen, sie dünkte sich
zu
erzeugen,
in
rastlosem
Turm bis zum Himmel führen. Nun aber
erfährt sie bei allen äußeren
Triumphen einen inneren Niedergang,
Widerspruch gegen die Mattheit und Schlaffheit jener
Macht über die Seelen
gibt.
„In die Höhe will es sich bauen mit Pfeilern und Stufen, das Leben
selber: in weite Femen will es blicken und hinaus nach seligen Schönheiten,
^
Der
kräftige
Qleichmacherei
ist
es vornehmlich, welcher Nietzsche
—
darum braucht es Höhe! Und weil es Höhe braucht, braucht es Stufen
und Widerspruch der Stufen und Steigenden! Steigen will das Leben und
steigend sich überwinden". (Also sprach Zarathustra.)
Gesellschaft und Individuum.
vermag
ja sie
sich selbst nicht
zu verstehen, es droht ihr ein innerer
und
eine
Streit,
lösung
in
wachsende
Parteien, ein
301
mehr zusammenzufinden, sich selbst
Überall Gegensatz
Zerfall.
des
Leidenschaft
Kampfes,
Wo wir unter Zurückstellung aller Weltprobleme bei
werden
einig
und
wollten
eine Auf-
Entschwinden gemeinsamer Ideen und Ziele.
von
Einigung
solcher
uns selber
die
schönsten
Früchte hofften, da ist eine Sprachverwirrung erfolgt, und wir werden
immer mehr der Zerstreuung und Zerstückelung verfallen, wenn es
Bewegung Einhalt zu gebieten, das menschliche
Dasein wieder in größere Zusammenhänge zu stellen und ihm einen
nicht gelingt, jener
festeren
Grund zu geben.
Bis dahin beschäftigte uns das Problem der Sozialkultur übermit- einigen
Worten der Stellung des Staates
im modernen Geistesleben gedacht.
Die stärkere Machtentwicklung
haupt, es sei
nun auch
des Staates steht heute uns allen vor Augen, und es drängen nament-
Verwicklungen zu noch weiterer Steigerung; damit
lich die sozialen
aber erwächst die Gefahr, daß mehr und mehr das ganze Geistes-
und seine Stemplung er-
leben unter den Einfluß des Staates gerät
fährt,
und
diese Gefahr,
nennen möchten,
des Staates
ist
ist
und
die
Gefahr
ist
Politismus,
sittlich"^ indifferent,
sie
wir wir
Entwicklung der Macht; nun
ist,
sie
Die leitende Idee
nicht gering anzuschlagen.
bleibt die
Macht keineswegs, wie wohl gesagt
sie
eines
ist
die
an sich etwas böses, aber
kennt kein höheres Ziel
als sich selbst.
Sie hat aus ihrer Natur das Streben, alle geistige Betätigung als ein
bloßes Mittel für ihre Zwecke zu
Selbständigkeit
anderer
behandeln,
Wenn
Lebensgebiete.
sie
anerkennt keine
aber
diese
Gebiete
vor allem daraufhin angesehen und danach gemessen werden, was
sie
für
das staatliche Leben
wert und
leisten,
so
verlieren
sie
ihren Selbst-
müssen zugleich an der Ursprünglichkeit ihres Schaffens
schweren Schaden erleiden.
Zugleich
muß bei einer Beherrschung
des ganzen Lebens durch die Staatsidee die eigne Art und die
freie
Bewegung des Individuums aufs stärkste eingeengt werden. Wenn
der Mensch sein Sinnen und Denken vor allem darauf richtet, einen
Platz im Staatsgefüge zu erringen und in diesem weiter zu kommen,
wenn sein Wert daran hängt, was er in ihm bedeutet und leistet,
so
wird damit der Schwerpunkt seines Lebens nach außen verlegt,
und Ursprünglichkeit des Lebens muß unvermeidlich Schaden erleiden. Ob das Gefüge des Staates mehr demo-
die
Selbständigkeit
Zu den Problemen des Menschenlebens.
302
oder aristokratischer Art
kratischer
dabei wenig.
Wie
Hand in Hand gehen
besagt
ist,
politische Größe mit geistiger Unproduktivität
dafür
kann,
liefert
die
römische Geschichte das schlagendste
Bei-
Denn höchst merkwürdig ist und bleibt es, daß bei so viel
politischer FCraft, Weisheit und Disziplin jenes Volk nicht einen
spiel.
einzigen großen philosophischen Gedanken, nicht eine einzige große
künstlerische Tat aus eignem
Auch wir Deutschen
welche
Jene
tritt
unsere
Vermögen hervorgebracht hat.
müssen der Gefahren
zurückdrängende Macht der
alles
eingedenk
jener Politismus,
Staatsidee,
uns namentlich im preußischen Staat entgegen.
zeitweilige
Unterordnung
zwingende
Notwendigkeit,
schichtliche
Aufgabe
aller
Aufgaben unter
wenn
lösen
dieser
sollte;
Staat
dabei
hat
danke der Macht aufs engste mit dem der
damit
innerlich
sein,
Entwicklung bei diesem Probleme mit sich bringt.
veredelt.
Gewiß war eine
die Staatsidee
eine
seine
große weltge-
sich
hier
Pflicht
der
Ge-
verbunden und
Die Verbindung beider Ideen hat herr-
liche Leistungen hervorgebracht, die das Deutschland der Gegenwart
erst
haben entstehen
lassen.
Aber alles dies darf die Gefahr einer
Unproduktivität, auch
geistigen
eines Einschnürens
der Individuen,
und schablonenhaften Gestaltung des Lebens nicht
lassen.
Geistiges Schaffen und ursprüngliche Lebens-
einer typischen
übersehen
führung wollen schlechterdings als Selbstzwecke behandelt sein, der
Politismus aber,
mag er noch so veredelte Formen annehmen,
seinem Utilitarismus
setzt sie
mit
unvermeidlich zu bloßen Mitteln und
Werkzeugen herab.
ßß.
Die Unzulänglichkeit einer bloßen Individualkultur.
Die Gegenbewegung gegen die gesellschaftliche Kultur, die das
moderne Individuum
Sorge
vollzieht,
entsprang
zunächst
weniger einer
um den geistigen Gehalt des Lebens, als sie die Schädigungen
abwehren wollte womit das Vordringen jener Kultur das Individuum
bedrohte; doch standen tiefere Probleme im Hintergrunde und verschärften
den Gegensatz.
das Individuum
es
lediglich
als
— Die gesellschaftliche Kultur behandelt
ein Stück ihres
großen Räderwerkes,
nach seinen Leistungen,
sie
muß
sie schätzt
es für ihre
Zwecke
beengen und einschränken.
Dazu wirkt sie mit ihren Verzahnungen der Elemente, mit ihrer Anhäufung von Massen, ihrem
lauten und fabrikmäßigen Getriebe übermächtig zur Unterdrückung
vielfach
und Abschleifung der
individuellen Züge,
sie
gewährt keine
stille
Gesellschaft und Individuum.
Ruhe zur Bildung eigentümlicher Art,
Durch-
erzeugt gewisse
sie
selbst zum Maßstab für Gut und Böse, für Wahr
die sich
schnitte,
303
und Unwahr machen.
Gegen
solche
Bindung und Gleichmachung
erhebt sich schließlich das Individuum kräftigerer Art und tritt dafür
daß keineswegs der Mensch
ein,
lichen
und
heit
das Verhältnis zur gesellschaft-
die
des
Innerlichkeit
Lebens,
jenes Verhältnisses
jenseit
Dabei kann es das Zeugnis der Weltgeschichte dafür anrufen,
liegt.
daß
in
Umgebung aufgeht, daß vielmehr das Beste an ihm: die Ein-
vorwiegend
alle
der
Gestaltung
gesellschaftliche
Kultur
eine
Veräußerlichung und Mechanisierung bewirkte, und daß es keines-
wegs bloß
trotziges Selbstgefühl
ein
der Individuen,
sondern das
unabweisbare Verlangen nach mehr Seelenhaftigkeit des Lebens war,
was einen Bruch damit erzwang.
bringt
Religion
die
dem Gebiet der
Namentlich auf
zur Kirche unver-
gesellschaftliche Gestaltung
meidlich die Neigung mit sich, die Leistung (Gottesdienst,
Werke,
vor die Gesinnung,
pflichtungen)
das
des
Beisichselbstsein
Interesse
der Religion
Kirche nötig gemacht.^
Individuums
des
überhaupt
Bekenntnisse,
korrekte
Ver-
vor das persönliche Leben,
vor
zu
Innern
fromme
religiösen
die
sog.
so
stellen;
hat
das
eigenste
immer von neuem einen Kampf gegen die
Zu solcher Verfechtung der Selbständigkeit
gesellt
ein
sich
flammender
Protest
gegen
die
gleichförmige und schablonenhafte Gestaltung der Kultur, womit die
Gesellschaft das Leben bedroht.
Sind die Durchschnitte, die dabei
entstehen, nicht recht geringer Art,
Festlegung
des
Sind
in
nicht
Lebens
Wahrheit
und bedürfen
sie
Wirkung
das
auf
auf
einer
geistige
nicht zu
aber
So gab
wesentlichen
Scheidung
werden,
der
um das
Menschheit,
übrige
nicht leicht zur
und
edle
Mittelhöhe?
Gesinnung
selten,
einer
scharfen
Ausprägung
und
kleinen Kreisen einer engeren Jüngerschar?
in
keinen
Kraft
sie
beträchtlichen
ihrer Ausbildung voller Freiheit, zu einer
Ganze
sicheren Befestigung
es
und führen
nicht
ein
Fortschritt
Höheres
der Kultur ohne eine
mußte
vorausgeworfen
nach sich ziehen zu können,
eine Feuer-
mußte dem großen Haufen voranleuchten und ihm den Weg
säule
Trotz aller Bedenken und Verwahrungen
immer wieder ein Gegensatz esoterischer und exoterischer Lebens-
durch die Wüste zeigen.
ist
'
Beim Begräbnis eines Zentrumsführers wurde seitens eines hohen
rühmend hervorgehoben, der Verewigte habe die Sorge für seine
Prälaten
—
Seele gänzlich den
Händen der Kirche überlassen.
daß eine
innere Preisgebung des Lebens auch noch belobt wird?
derartige
Ist
es nicht entsetzlich,
Zu den Problemen des Menschenlebens.
304
führung entstanden, auch die
radikalste politische Verfassung hat die
Bildung schroffer sozialer Unterschiede nicht verhindert,
Menschen
sind,
Einzelnen
alle geistige Tätigkeit
wenn
sie
sich
damit
selbst
nicht
mit
individuell
eignes Wesen kämpft?
Und bleibt nicht für jeden
matt und wie von außen angeweht,
seiner
gestaltet,
Art
individuellen
wenn
er
nicht
verflicht
und
um
sein
bei
ihr
Bilden heißt scheiden, differenzieren, indivi-
war die Scheidung überall
ein unentbehrliches Mittel
Bewegung und Weiterkommen.
Aus solchen Erwägungen gehen die Individuen von der Abwehr
für
zum
vor
Angriff
und rücken
Schranken deutlich vor Augen.
ist
bis in
der Ehrgeiz, es Höherstehenden gleichzutun, eine
unentbehrliche Triebkraft der Bewegung.
dualisieren; so
ja
und der Sitte ist, wie einmal die
die Äußerlichkeiten des Anstandes
der
gesellschaftlichen
Kultur
unmittelbaren Verhältnisses zur Wirklichkeit fähig,
eines
ihre
Der Mensch, als denkendes Wesen,
er
ist
kein bloßes Glied einer Verkettung, er kann sich der Unendlichkeit
gegenüberstellen und mit ihr ringen, er wird der Enge der bloßen
und kann von ihr zur eignen Wahrheit der
Gewiß stößt solches Streben auf Hemmungen über
Zuständlichkeit bewußt
Dinge streben.
Hemmungen, aber schon als Streben erweist es eine Überlegenheit
des Menschen über den Kreis der bloßen Gesellschaft.
Ist
es
nun
nicht ein Widersinn, einem solchen Weltwesen das Geistesleben erst
Maß dessen
durch die Gesellschaft vermitteln und es dabei an das
binden zu wollen, was der Zusammenschluß der Kräfte an Geistighat?
keit erreicht
Soll das
Wesen, das aus seinem Grundverhältnis
zur* Geisteswelt einen unendlichen Wert besitzt, sich seinen Wert erst
von menschlicher Schätzung zusprechen
der Menschen leben und damit
verlieren?
Existenz
Soll
erst
alle
und sicher fühlen, nachdem
Güter, wenn nicht auf
In
und
dem
von Gnaden
der Mensch sich einer Wahrheit, ja einer geistigen
froh
ihm mit Brief und Siegel verbürgt hat?
erfolgen,
lassen, soll es
Unabhängigkeit der Gesinnung
dem Markt
die Gesellschaft sie
Soll die Erzeugung geistiger
des Lebens,
so doch für ihn
sollen damit jene Güter zu bloßen Marktwaren sinken?
allen
wegte Individuum,
erscheint das Individuum,
als
der Vertreter
d. h.
das geistig be-
der Geisteskultur
gegenüber
einer bloßen Menschenkultur, einer inneren Unendlichkeit gegen alle
äußere Begrenzung, es erscheint
widersteht,
das
aus der Erstarrung
der Verflachung
als eine
Kraft, die
aufrüttelt,
notwendige Ziele vorhält,
menschliche Streben immer neu auf seine wahren Grundlagen
Gesellschaft und Individuum.
305
Und wenn solche Schätzung des geisterfüllten Individuums notwendig eine Abhebung vom gesellschaftlichen Durchschnitt
zurückführt.
mit
sich
so
bringt,
Unduldsamkeit gegen
auch die diesem Durchschnitt eigne
wird
sie
alles
irgend Überragende mit sicherem Stolze
Es gibt einen gemeinen Neid und Haß des Mittelmäßigen
abweisen.
gegen das Höhere als gegen eine Überschreitung und Herabsetzung
der eignen Dürftigkeit; dies Höhere wird leidlich geduldet nur dann,
wenn
es
Äußerung des
alle
bittet,
Kraftgefühls behutsam unter-
Daher steht die Tugend der Bescheidenheit beim
drückt.
um Ent-
bescheiden duckt, für sein Dasein höflichst
sich
schuldigung
Philister-
Nahe verwandt ist dem das Unterordnen
des Verschiedenartigsten und Verschiedenwertigsten unter die gleichen
tum so hoch
Ehren.
in
von Lob und
Begriffe, das Operieren mit nichtssagenden Schablonen
Tadel, jene laue
und matte Art,
die keiner kräftigen Liebe und keines
kräftigen Hasses fähig ist, der Licht und Dunkel in einen grauen Nebel
Demgegenüber zur Kräftigung des Empfindens,
zusammenfließen.
zur Scheidung der Geister
zur 'Schärfung des Urteils,
das
zu wirken,
ein gutes Recht, ja eine heilige Pflicht des Individuums.
ist
kann das Individuum
Freilich
rechtem Sinne überlegen nur
in
werden, wenn es eine Geisteswelt hinter sich hat und aus ihrer Kraft
zu schöpfen
vermag.
Dies aber
modernen Individualismus, wie
Er
stellt
und
das Individuum
heißt
es
sich
im Durchschnitt ausnimmt.
gänzlich
auf
sein
Zusammenhänge zu
Bindung an Menschen, sondern auch
streifen.
So
ihm
verbleibt
seelische Zustand,
unmittelbares
da das Leben gestalten;
unsichtbaren
alle
flissen,
von
keineswegs die Meinung des
ist
er
er
Dasein
besonders be-
ist
lockern, nicht
nur die
die an eine Geisteswelt abzu-
anderes
nichts
das subjektive Befinden;
als
indem
der
unmittelbare
dies
zum Kern
Lebens wird, verschmilzt der Individualismus mit dem Sub-
alles
damit eine gewisse Art von
jektivismus.
Augenscheinlich
Wirklichkeit.
Jene Zuständlichkeit läßt sich fixieren
entsteht
und
steigern.
Eigentümliches vermag sich schrankenlos auszubilden, Leben immer
So
neu
aufzuquellen,
eine
große Leichtigkeit, Frische und Flüssigkeit, das Leben scheint
hier
ganz
Stand
sein
auf sich
selbst
sich
gestellt,
unablässig
und
zu
verändern.
bei solcher Freiheit feiner,
Auch der Begriff der
Denn als
Starrheit.
wahr gilt nunmehr nur das, was die Seele des Einzelnen erlebt und
was sie eben jetzt erlebt; so weicht der Begriff einer einzigen Wahr-
zarter,
intimer
Wahrheit
geworden
verliert
seine
Eucken, Grundbegriffe.
4.
als
irgendwo
sonst.
sonstige Schwere und
Aufl.
20
Zii
306
den Problemen des Menschenlebens.
dem unzähliger Wahrheiten, jeder Mensch hat hier seine eigne
Eine besondere Freude und Bewußtheit gibt dem der
Wahiheit.
heit
Kontrast
zur Gesellschaft,
dem Lebensgefühle
deren
denn demgegenüber das Leben
das
halten,
und Anordnungen
Einrichtungen
des Individuums
oft
widerstreiten;
in
Freiheit
stets
so heißt es
und Fluß zu
möglichst zu stärken und deutlich
Eigentümliche
er-
her-
vorkehren.
Das alles konnte aber den Stand einer formlosen Erregung und
unklaren Bewegung nicht überschreiten ohne sich irgend in geistige
Arbeit
Wendung
umzusetzen, es fand solche Umsetzung durch die
und
zur Kunst
Die Kunst
Literatur.
mannigfachen Ver-
ihrer
in
zweigung wird hier zum Hauptmittel, das sonst unstet wogende und
wallende Leben irgend zu fassen und festzulegen, durch die Fassung
aber
es
zu
verstärken,
sich
bei
es
selbst
nach außen hin unabhängig zu machen.
voll
durchzubilden und
Konzentration des Lebens
in sich selbst und Steigerung seiner Kraft, das wird damit zur Haupt-
aufgabe der Kunst.
Sie
die
weit überlegen
fühlt;
über
vornehmere einer
die Kunst kann
Zweckmäßigkeit
bloßen
aller
wird die Seele einer individuell -aristokra-
sich als die
tischen Kultur,
liegt
wiegend auf sein individuelles Vermögen
weil
sie
aus
schnittslebens
aller
die
Verworrenheit
und
weil
sie selbst
und den Menschen vorstellt,
weiter aber deshalb,
Vergriffenheit
des
Durch-
Grundzüge menschlichen Seins heraus-
einfachen
sehen. Ewigjunges in
praktisch-sozialen'
das werden,
ihm ergreifen und es damit aus aller Erstarr-
ung im Konventionellen aufrütteln kann.
Leicht
überträgt
das Bewußtsein
Bahn.
schüssige
sich
aber solche Abstufung des Lebens
auf
der Individuen und gerät dabei rasch auf eine abNicht nur,
wer an der neuen Art mit eignem
Wirken teilnimmt, sondern auch, wer sich bloß dazu bekennt, glaubt
sich der übrigen Menschheit
legen;
entsteht
es
und der gesellschaftlichen Kultur über-
eine Neigung,
den Abstand hervorzukehren, das
in der Absonderung ein GeGröße zu suchen. Und zugleich greift der Anspruch um sich, unbekümmert um alles, was da gilt, um Sitte und
des Landläufigen zu tun,
Gegenteil
fallen,
eine
ja
Gesetz, die eigne Art nach Lust
sichtslos
M auszuleben".
und Laune zu entfalten, sich rück-
Die Individualkultur mag das
wollen, unter menschlichen Verhältnissen
ist
alles
nicht
solche Folge schwer zu
vermeiden.
Derartige subjektivistische Strebungen und Stimmungen spielen
Gesellschaft und Individuum.
307
bekanntlich eine große Rolle in der neuesten Gestaltung des Lebens.
Neu ist dabei freilich nur der Name, die Sache ist alt, uralt. Denn
in periodischer Wiederkehr kommen immer wieder Lagen, wo
wie
das
unmittelbare
befriedigt
Lebensgefühl von der dargebotenen Kultur nicht
war und nun als Hülfe und Heil die völlige Emanzipation
des Individuums verkündet wurde, wo seine unmittelbare Empfindung,
selbstgefundenes
sein
Urteil,
sein
künstlerischer
Geschmack
eine
Wendung zum Besseren bringen sollte. Wer Piatos Gorgias kennt, der
kennt auch die nahe Verwandtschaft der Sophisten mit den heutigen
Subjektivisten; in Deutschland brachte zuerst die Geniezeit, die Vorläuferin der klassischen Literaturepoche, eine derartige Emanzipation
des Einzelnen; damals waren »Genie", »Kraftgenie«, «Originalgenie"
Modewörter wie heute «Übermensch"; auch »schöne Seele"
steht in
Dann kam eine neue Welle
der Romantik, deren nahe Verwandtschaft mit dem ästhetischen
Verwandtschaft mit dieser Bewegung.^
in
Subjektivismus der Gegenwart deutlich zutage
liegt.
Das Ganze gerecht zu beurteilen ist schwer,
um
eine Übergangserscheinung handelt,
die
weil es sich offenbar
um so mehr Vernunft
und Recht besitzt, je mehr sie sich weiteren Zusammenhängen einfügt
und über sich
selbst hinausweist, die
um so mehr ins Unrecht gerät,
Über das Aufkommen und die Schicksale des Ausdrucks Genie handelt
und erschöpfender Weise Hildebrand in Grimms deutschem
Wörterbuch. Hinzufügen möchten wir nur eine Stelle aus dem kürzlich veröffentlichten Briefwechsel zwischen Goethe und Lavater, die für die schärfere
Abgrenzung von „Genie" gegenüber „Talent" von Bedeutung ist. Goethe
^
in mustergültiger
schreibt (Schriften der Goethe-Gesellschaft, Bd. 16, S. 125) 24. Juli 1780:. „Bei
Gelegenheit von Wielands Oberon brauchst du das Wort Talent,
der Gegensatz von Genie wäre,
subordiniertes.
sein
kann
als
als
wenn es
wo nicht gar, doch wenigstens etwas sehr
Wir sollten aber bedenken, daß das eigentliche Talent nichts
die Sprache des Genies."
Lavater antwortet darauf (unterm
August 1780) mit einer längeren Auseinandersetzung über den Unteischied von Talent und Genie (S. 130ff.), woraus nur folgendes angeführt sei:
5.
„Nur Ein Wort von Talent und Genie.
Zwei Worte, die ihrem Sinn und
mögen wie schön und erhaben.
Gehalte nach ungefähr so verschieden sein
macht mit Leichügkeit, was tausend andere nur mit äußerster
oder es macht mit Frohmut und
Grazie, was andere nur gerecht und korrekt machen; Genie macht, was niemand machen kann. Alle Werke des Talentes erregen bewunderndes Wohlgefallen; Genie erweckt Ehrfurcht, erregt ein Gefühl, das der Anbetung
nahekommt."
Über „schöne Seele" bringen die beste Aufklärung die
neuesten von Ippel besorgten Auflagen von Büchmanns Geflügelten Worten.
Talent, mein' ich,
Mühe und Langsamkeit machen können;
—
20*
Zu den Problemen des Menschenlebens.
308
je starrer sie
sich festlegt
hindert
die
hier
und
Verpönung
Dazu ver-
in sich selber einspinnt.
aller
bindenden Normen
alle
Höherem und Niederem, zwischen
Scheidung zwischen
scharfe
geistiger
Notwendigkeit und menschlicher Willkür, bunt wirbelt Mannigfachstes
kaum
durcheinander,
vermeidlich
Gefahr,
die
ist
in
Anerkennung
des Höheren nachgiebig gegen das Niedere, in Abwehr des Niederen
ungerecht gegen das Höhere zu werden.
Trotzdem
sich
läßt
auf
Würdigung nicht wohl verzichten.
irgendwelche.
Warum
kann eine auf das bloße Individuum und seine Zu-
Aus zwei Hauptgründen
das Individuum des unmittelbaren Daseins — und
standlichkeit gestellte Kultur nicht genügen?
nicht:
1.
weil
das allein steht hier in
genugsam
ist,
2.
und hohler wird,
je
von ihm entwickelte Leben
mehr
Individuum
empirische
— weder unabhängig noch selbst-
Frage
weil das
ist
in
um so leerer
— Das
Folgen hervortreibt.
seine
es
Wahrheit
eher als unabhängig.
alles
Denn Vererbung, Umgebung, Erziehung bedingen es nicht nur aufs
mannigfachste, sie scheinen es gänzlich hervorzubringen; sie flechten
ein
daß ihm weder
so dichtes Netz,
kann.
Seele,
Sicherlich
der
das
Jedenfalls
ist
reicht
Individualismus
es nicht deshalb
List
noch Gewalt entrinnen
Bindung auch
diese
für
schon
druck keine Bindung empfindet.
völlig
frei,
in jenes Innerste
freischwebend
der
ausgibt.
weil der unmittelbare Ein-
Denn mag sich der Individualist
der Welt noch so keck entgegenwerfen und sich völlig von ihr abzulösen scheinen, er bleibt doch im Schatten
und Bannkreis dieser
Welt
gewöhnlich nur eine
Seine vermeintliche Unabhängigkeit
ist
Der Indi-
andere Art der Abhängigkeit, eine indirekte Abhängigkeit
vidualist
ist
geneigt, das Gegenteil dessen zu sagen
die Umgebung sagt
ung
vorschreibt;
und zu tun, was
und tut, so ist es diese, we'.he ihm
die
Kette
ist
nicht zerbrochen.
die Richt-
Der Individualist
fühlt sich der Umgebung überlegen, aber den Abstand ermessen und
genießen kann er nur,
bleibt er
auch hier an
sofern er die anderen
sie
gebunden.
der Unabhängigkeit, aber er
Zuschauer
im Auge
behält;
so
Er wonnt im stolzen Gefühl
muß dabei unablässig die anderen als
und Bewunderer solcher Größe denken.
Das Leben
kommt also nicht zu einer festen Ruhe und einem freudigen Schaffen
bei sich selbst, es steht nicht auf seinen eignen Notwendigkeiten.
So
muß
es
kann es die Beziehung zum Menschen nicht aufgeben, so
vom Kontraste leben, vom Kontraste zehren,
den Stand einer inneren Abhängigkeit
so überwindet es nie
Gesellschaft und Individuum.
Auch
gerät
Wendung
solcher
bei
von
in Gefahr, einen Zusatz
das Bewußtsein der
aufzunehmen.
Eitelkeit
und Wesens sind
schiede des Lebens
309
Größe
Starke Unter-
der Grad der Belebung
da,
der Geistigkeit zeigt weiteste Abstände, die ordinäre Gleichmacherei,
deren Stumpfheit
alles
zusammenwirft,
Auch
die
Individualität
ja
sei
nicht
dem
schwächt!
Denn
Wahrheit
und
Durchbildung
einzelnen
Stelle
nicht
sie
ist
wird mit Recht verworfen.
verdunkelt oder abge-
irgend
Schaffen
geistigen
unentbehrlich;
zu seiner vollen
gelangt
es
an
der
auf den Punkt seiner eigentümlichen Stärke,
wo es vollauf seine eigne Natur entfaltet, so wird es nie der Widerstände Herr werden.
Aber in dem allen muß eine überlegne Notwendigkeit
des
Menschen
treiben
und wahr.
Lebensprozesses
und
walten,
ein
geistiger
Zwang den
nyr dann verbleibt die Sache gesund
leiten;
Sie verfällt sofort ins Künstliche
und Ungesunde, wenn
das Individuum darauf ausgeht, sich möglichst an jeder Stelle indi-
und groß zu zeigen, wenn es den Abstand geflissentlich
wenn es gar zur Sache reflektierenden Genusses macht,
viduell
hervorkehrt,
dessen Ausführung reine Hingebung und selbstlose Liebe verlangt.
Jedes Zurücktreten hinter die Sache, jedes
Aufkommen eitler Selbst-
bespiegelung schwächt die geistige Kraft und lockert den Zusammen-
hang mit den inneren Notwendigkeiten, an dem alles Gelingen liegt.
muß man haben, nicht danach streben" (J. Burckhardt).
Gewiß kämpft unter der Fahne des modernen Individualismus
«Originalität
vieles,
was solchen reflektierenden Subjektivismus mit seinem epikure-
ischen Selbstgenusse
und der
bare
Eifer der
weit überragt;
Größe ihrer Leistung nur verstehen aus dem Erscheinen neuer
sachlicher
Aufgaben,
frischer
Antriebe
Wirklichkeit neue Seiten abgewinnen
zu
ihr eröffnen.
setzt sie
sie
namentlich läßt sich der Ernst
modernen bildenden Kunst, sowie die unverkenn-
in
innere
des Schaffens,
welche
der
und ein innerlicheres Verhältnis
je bedeutender die Arbeit, desto mehr verZusammenhänge und Notwendigkeiten, unterwirft
Aber
das Schaffen einer überlegenen Wahrheit, befreit sie vom bloßen
Subjektivismus und Individualismus.
Unvermerkt wird hier aus einem
Individuum gegenüber der Geisteswelt ein Individuum mit der Geisteswelt;
einen
einem solchen aber kann der heutige Sturm und Drang nur
Übergang zu
Ähnlich
einer
bewahrt auch
reinen Individualismus
höheren Stufe
bei
der Wahrheit
der Frage des
bedeuten.
Lebensinhalts den
und Subjektivismus nur eine unablässige Er-
gänzung vor unerträglicher Leere.
Streng
genommen muß er die
Zu den Problemen des Menschenlebens.
310
Seele in lauter einzelne Vorgänge, schließlich in bloße Stimmungen
Da
und vertreiben.
auflösen, die in rascher Flucht einander jagen
jeder Augenblick genau so viel Recht wie der andere hat, so hätte
jeder seine eigne Wahrheit;
mag,
das
was dabei zunächst ein Gewinn scheinen
schließlich
sich
stellt
als
Das menschliche Leben erschöpft
schwerer Verlust
ein
keineswegs ganz
sich
heraus.
in
lauter
einzelne Augenblicke.
Die Augenblicke mit ihren Erlebnissen ver-
sinken nicht völlig,
kehren zurück,
so
Seele;
urteilen
und messen, so
Daher muß
er
heute als wahr
keit
sie
sie stellen
sich
uns vor die
muß der Mensch sie vergleichen und verbinden,
er
steht
auch das Unwahrwerden
gilt,
dessen erleben, was ihm
daher empfindet er die Flüchtigkeit und Nichtig-
des ganzen Getriebes, daher überzeugt er
heit für gestern
be-
sie
über den bloßen Augenblicken.
sich,
daß eine Wahr-
oder heute überhaupt keine Wahrheit
ist,
daß
sein
und jede Wahrheit verliert^ wenn es an die bloßen Augen-
Leben
alle
blicke
gebunden bleibt
niederdrückt,
als
Stimmungen, das
der
Gibt es etwas, das mehr ermüdet und tiefer
unablässige
Umschlag der Meinungen und
eifrige Verketzern dessen,
was eben noch begeistert
verehrt wurde, die Herabsetzung aller geistigen
Sache bloßer Laune und
Bewegung zu einer
Mode?
Der Individualismus möchte dem menschlichen Leben zur vollen
Entwicklung seiner Kraft verhelfen und ihm möglichst den Charakter
der
Größe geben.
und würdigen läßt.
Das
ist
ein Streben,
Steht der
das sich vollauf verstehen
Mensch an einem Wendepunkt des
Alls,
beginnt in ihm eine höhere Stufe der Wirklichkeit, so
dies
Höhere zu ergreifen und gegen
durchzusetzen, es
gilt,
einem Berge zu leben.
um
gilt
es
allen Widerspruch des Alltages
mit Marc Aurel
So hat sich von
zu sprechen, wie auf
alters her,
wo immer der
Abstand zwischen den Forderungen des Geisteslebens und der Durchschnittslage der Menschheit zur deutlichen
Empfindung gelangte, mit
zwingender Notwendigkeit der Gedanke einer höheren Art des Lebens,
Größe des Menschen entwickelt; er läßt sich von der
Höhe der griechischen Kultur durch mannigfaltigste Wandlungen hin-
einer inneren
durch bis zur Gegenwart verfolgen. ^
^
Aber wird der moderne Indi-
Eine solche Verfolgung des Problems durch die verschiedenen Zeiten
hindurch wäre eine anziehende Aufgabe.
Den wissenschaftlichen Ausgangs-
punkt würden dabei die eindringenden Untersuchungen des Aristoteles über
den Großgesinnten (jjLEYaXdlu/o;) bilden. Hier sind die Begriffe noch mitten
im Fluß, der Gedanke einer Größe innerhalb des menschlichen Kreises ver-
Gesellschaft und Individuum.
311
Größe gelangen, wenn er alle
Zusammenhänge und damit alle Möglichkeit einer ErweiterEs gibt kaum
ung des Menschen zu einem Weltwesen aufgibt?
vidualismus zu einer wahrhaftigen
inneren
einen härteren Widerspruch, als den Menschen zu einer überlegenen
und zugleich eine selbständige Innen-
Innerlichkeit führen zu wollen
und
welt schroff
ja
an erster Stelle jene selbständige Innenwelt ver-
vielfach unerfreulich sein,
tritt,
Mag der heutige Stand
zu bekämpfen.
erbittert
der Religion, die
wir sollten doch als
freie
Menschen
unsere Begriffe und Überzeugungen von höchsten Dingen nicht nach
dem bilden, was
die
Umgebung uns zuführt, sondern nach dem,
verlangt. Ohne eine Um-
was die Notwendigkeit des eignen Lebens
kehrung der ersten Lage, ohne Metaphysik gibt es keine selbständige
Innenwelt, keine wahrhaftige
Wo immer daher
Größe des Lebens.
dem Gemenge des modernen Lebens Gestalten merklich hervorragen, da ist eine Wendung zur Metaphysik mit im Spiel. So z. B.
aus
bei
Nietzsche..
seinen
In
Begriffen
drücklichst bekämpft, in seinen
Welt
dieser
hinreißende
es
des nächsten Anblicks,
die
als
Bildner
Welt,
er
Metaphysik
alle
und eben
als
modernen
nach-
künstlerischer
Metaphysiker der Stimmung,
Gemüter gewonnen.
Gewalt über die
gesamten
der
als
hat
Stimmungen wirkt eine völlig andere
hat er die
Ähnlich
Strömung zur Romantik.
geht
Die bloße
Stimmung aber reicht nun und nimmer aus, eine Größe gegenüber
den verflachenden und niederdrückenden Wirkungen der Umgebung
auszubilden und durchzusetzen,
die bloße
sie gibt
Aus
ung, nicht der Wirklichkeit.
nur eine Größe der Mein-
nichts läßt sich nichts bauen,
und
Stimmung hat nichts hinter sich.
Nicht anders steht es mit dem Verlangen nach Kraft. Ja gerade
heute bedürfen wir gegenüber schweren Verwicklungen
Aufgaben
des Qesamtlebens
viel
mehr
Kraft,
Kraft,
gesellschaftliche
Kultur zu bereiten vermag, aber
wandelt sich
unvermerkt
Menschlichen.
fast
in
und großen
als
die bloß-
durch ein bloß-
den einer Größe im Gegensatz zu allem
Die alte Weit denkt bei der Größe namentlich an eine dem
Ruhe und Selbständigkeit, die Neuzeit
und eine geistige Schöpfungskraft; so auch hier der Gegensatz der Ideale von Beharren und Bewegung.
Das viele Gerede von Größe dürfte namentlich aus der Zeit Ludwigs XIV.
menschlichen Getriebe
mehr an
ein
überlegene
überlegenes Leistungsvermögen
stammen, wenigstens berauschen sich die Schriftsteller jener Zeit besonders
Die bedeutendste neuere Untersuchung über historische
an jenem Begriffe.
Größe
dürfte
die
Betrachtungen" sein.
von Jakob Burckhardt
in
seinen
„Weltgeschichtlichen
Zu den Problemen des Menschenlebens.
312
subjektives Sichemporheben,
ein
Sicheinreden
der
Kraft,
Sichr
ein
distanzieren von den anderen Menschen kommen wir nun und nimme:
Die eigne Erfahrung der Gegenwart
zu wahrhaftiger Kraft
das deutlich genug.
Mehr reden von
läßt sich schwerlich; sind
literarisches
und unser
Kraft,
als
zeigt
wir es heute tun,
wir dadurch kräftig geworden, zeigt unser
politisches
Leben eine Fülle
starker,
selbst-
wüchsiger, ausgeprägter Persönlichkeiten, eine Fülle großer, erhöhender
Schöpfungen ?
Die Notwendigkeit einer inneren Überwindung des Gegensatzes.
XX-
Wenn weder die bloßgesellschaftliche noch die individualistische
Kultur den Aufgaben gewachsen
wenn
ist,
keine von
beiden
dem
Leben einen wesenhaften Inhalt gibt, und wenn zugleich außer Zweifel
daß nur eine klägliche Stumpfheit einen direkten Kompromiß
steht,
zwischen beiden versuchen, das Leben zwischen hier und dort verkann,
teilen
so
müssen wir unbedingt dem Gegensatz überlegen
Individuum und Gesellschaft sind notwendige Seiten und
werden.
Erscheinungsweisen des Geisteslebens, zu seiner Ursprünglichkeit bedarf es der Individuen, zu seiner Befestigung der Gesellschaft; Indi-
viduum und Gesellschaft aber ziehen
aus sich
selbst,
umfangen.
ihre Kraft
und Wahrheit nicht
sondern aus den geistigen Zusammenhängen, die sie
Das Verhältnis von Individuum und Gesellschaft wird
dem Boden der Geschichte verschieden gestalten; die Gesellschaft hat für sich den Zug des Lebens, wo es nach Auflösungen
sich auf
und Erschütterungen vor allem einer Befestigung bedarf, wie
beim Ausgang des Altertums.
Individuen
das
stellt
zwingend
zur
uns Augustin
Was
damals auch
Anlehnung an
mit voller
die
die
Gemeinschaft
Klarheit vor Augen.
z.
B.
kräftigsten
trieb,
Die Be-
wegung zum Individuum erhält dagegen die Oberhand, wo frisch
aufstrebende Kräfte die überkommenen Ordnungen als zu eng und
starr
empfinden und nur
in
Befreiung von ihnen neue Bahnen zu
Das war der Haupttrieb der Neuzeit bis ins
Daß dann ein Rückschlag kam, und daß in
19. Jahrhundert hinein.
der Gegenwart zugleich die Gesellschaft und das Individuum eine
Verstärkung verlangen, daß eine praktisch-soziale und eine künstlerisch-
suchen
vermögen.
individuale Art
um den Menschen kämpfen, das zeigt mit besonderer
Deutlichkeit die innere Zerklüftung unserer Zeit, das
muß aber zu-
gleich als ein zwingender Antrieb zur Erhebung über jenen Gegensatz, zur Wendung von einer bloßen Menschenkultur zu einer
Gesellschaft und Individuum.
Nur durch ein
inneres Vordringen des Lebens läßt
klüftung begegnen;
das
sich jener Zer-
denn was überall von den echten Problemen,
daß nicht Meinungen gegen Meinungen,
besonders hier,
gilt
313
umspannen vermag.
Geistes- und Wesenskultur wirken, die jenen zu
sondern Lebensentfaltungen gegen Lebensentfaltungen stehen.
Die sozialdemokratische Bewegung.
b)
Von geistigen Strömungen der Gegenwart läßt sich nicht wohl
handeln, ohne der Sozialdemokratie zu gedenken;
so
da aber über
sie
bis zum Überdruß verhandelt und geschrieben wurde, so
viel
empfiehlt
Beschränkung auf
die strengste
sich
was die
das,
sophische Betrachtung an Eigentümlichem vorzubringen
philo-
hat.
muß aber für die sozialdemokratische Bewegung am meisten
Ihr
daß
charakteristisch scheinen,
drei verschiedene
sie
Strömungen zu-
sammenfaßt und zur Wirkung verbindet: die demokratische, die ökonomische, die politistische;.es handelt sich einmal um die- Verlegung
Lebens
des Schwerpunktes des gemeinschaftlichen
um
sodann
die
Erhebung des
herrschenden Seele jenes Lebens, endlich
Staates
den
dem
zu
der Massen
gunsten
alle
einzelnen
namentlich
schöpft
Massen,
um die Anerkennung des
Durch
ökonomische Umwälzung
eine
herbeizuführen und aufrecht zu erhalten, das
bei
die
des einzigen Trägers von Vernunft und Macht.
als
Staat
in
Problems zur be-
wirtschaftlichen
ist
der Zentralgedanke,
Fäden zusammenlaufen.
Das Ganze aber
daraus
Kraft,
daß
die
einzelnen
Bewegungen
schon vor ihrer Vereinigung die Menschheit geweckt und begeistert
hatten,
und daß
sonst
bei
ihre
Verschmelzung nur zu vollenden
Konsequenzen zurückschreckt.
Strömungen
Bei
scheint,
was
unbestimmter Fassung verbleibt und vor seinen eignen
sei
in
Kürze
Demokratie
Der weltgeschichtliche Verlauf jener
betrachtet.
denken
wir
keineswegs
bloß
an
den
Staat,
Zusammenleben der Menschheit und an alles Verhältnis der Individuen zu den gemeinsamen Lebensgütern. Die Neu-
sondern an
zeit
ist
alles
schon deswegen jener Richtung günstig, weil schwere Hemm-
ungen früherer Epochen
einer
in
Wegfall kamen.
Im Altertum widerstand
die Beschränkung
Anerkennung der Gleichheit aller Menschen
der Kultur
auf
einzelne
Völker,
Besten keinen Anstoß erregte; was
eines unmittelbaren
so
daß
die
Sklaverei
auch
den
dem Christentum die Eröffnung
und eines gleichen Verhältnisses aller Individuen
Zu den Problemen des Menschenlebens.
314
zu Gott an Demokratischem einpflanzte,
die bis in die
das wurde sowohl
durch
Anfänge zurückreichende hierarchische Gestaltung als
durch die transzendente Lebensstimmung recht weit zurückgedrängt.
Erst
gehen.
kam
Zweigen der Reformation
einzelnen
in
Entfaltung,
es
zu stärkerer
um dann aber bald in die moderne Bewegung überzu-
Die Neuzeit
richtet
den Menschen immer mehr und immer
ausschließlicher auf das Diesseits,
und zugleich macht ihr Hauptzug,
die Aufklärung, zur Hauptsache an ihm etwas, das jenseit aller Unter-
schiede der Individuen liegt: die abstrakte Vernunft, das reine Denken.
Je
mehr sich das zu voller Bewußtheit erhebt und auch in die Über-
zeugung der Individuen
eindringt, desto unwiderstehlicher wird es;
immer mehr läßt damit das Menschsein alle gesellschaftlichen Unterimmer unabweisbarer wird die Anerkennung der
schiede verblassen,
was Menschengesicht
Gleichheit alles dessen,
trägt.
Wohl
enthält
auch die Neuzeit Gegenwirkungen zu gunsten eines Aristokratismus.
Große Unterschiede von
von Besitz und von
politischer Stellung,
Bildung übermittelt die Geschichte; aristokratischer als alle Geschichte
erweist sich bleibend die Natur mit ihren Unterschieden der körperlichen
und
schafft
auch die moderne Kultur mit der technischen Gestaltung und
seelischen
Ausstattung;
eine
der wachsenden Verzweigung der Arbeit.
eigentümliche Aristokratie
Denn je mehr diese
nimmt, desto mehr Gliederung und Abstufung erzeugt
zu-
desto
sie,
mehr Anordnung und Beherrschung bedarf sie, desto stärker wirkt
zu
sie
stände
gunsten eines neuen Aristokratismus.
Dinge
der
hindern
demokratischen
Strömung
Abstufung
bald
wird
bekämpft, bald
sie
nicht wie
als
in
jedoch
nicht
Alle
das
der menschlichen
künstlich
oder doch
solche
Wider-
Vordringen
der
Überzeugung;
die
künstlich
geworden
als
nebensächlich beiseite geschoben; jedenfalls wird
ein
starres
Schicksal
hingenommen, sondern durch
menschliche Gegenwirkung tunlichst verringert.
Mögen
bei
dieser
Bewegung die kleinen Wogen vielfach zurücklaufen, die große Woge
immer in der Richtung der Demokratie.
Auch die Selbständigkeit und das Übergewicht der wirtschaft-
geht noch
lichen Fragen
ist
wiß war
Sorge
allen
die
erst auf
um
dem Boden der Neuzeit erwachsen.
das Mein
Ge-
und Dein den Individuen zu
Zeiten die alles überragende Hauptsache; nur ein arger Fehl-
schluß
konnte früher das antike Leben lediglich idealen Aufgaben
zugewandt denken, weil die Philosophen das übermächtige Verlangen
nach Geld und Gut aufs stärkste zu brandmarken suchten.
Aber
Gesellschaft und Individuum.
eine
Würdigung fand das wirtschaftliche Gebiet in der
prinzipielle
Lebensordnung
antiken
315
volle Glückseligkeit
Es fand es einmal
nicht.
nicht,
weil
die
von der Entfaltung einer festen und begrenzten
Natur erwartet wurde, diese Entfaltung aber nur eines beschränkten
Aufwandes äußerer
bedarf;
Mittel
es
fand
auch deshalb
es
viduum auf
und auch
die Gemeinschaft übertragen
nicht,
vom Indi-
weil jenes ethisch-künstlerische Lebensideal unbedenklich
bei dieser jene
Grenze gezogen wurde.
Das Christentum mit seiner Richtung der
Gedanken auf eine übersinnliche Welt widerstand noch mehr einer
Schätzung der wirtschaftlichen Güter.
Die Theorie aber blieb bei
ihm durchaus unter dem Einfluß des Altertums.
Die Neuzeit da-
gegen mit ihrem Verlangen nach Entwicklung aller Kraft und ihrem
Streben
zur unmittelbaren Welt stand von vornherein anders zur
Sache.
Die
Güter gelten hier
materiellen
Hebel zur Bewegung der Kräfte,
einzuleiten als weiterzuführen.
Dazu wird das wirtschaftliche Streben
Bedenken verblassen, indem
ökonomie
gestaltet.
Schon
die
sich die
in
die
die
die
veränderte
den Über-
zeigt
So war es
in
den
all-
wenn schließlich die Theorie in
Bewegung zum Kern und zum maß-
vorbereitet,
wirtschaftliche
gebenden Typus des gesamten
Haupttriebkraft aller
die
17. Jahrhunderts
Politik eines Großstaats findet.
gemeinen Verhältnissen
A. Smith
veredelt,
Ökonomie zur National-
Renaissance
Schätzung, die dann im Frankreich des
gang
unentbehrlicher
ein
als
scheinen den Fortschritt sowohl
Bildung nationaler Einheiten verstärkt und
die
durfch
alten
sie
machte
Kulturlebens
Bewegung auch
in
Wissenschaft,
und
als
die
Kunst,
Er-
ziehung, Religion das Streben nach besserer Lebenshaltung erklärte.
An entschiedenem Widerspruch gegen
Ökonomischen
lässige
Wachstum
deutung der
solche
Führerstellung
des
hat es nicht gefehlt, aber andererseits hat das unab-
einer technischen
materiellen
und
verfeinerten Kultur die Be-
Güter immer weiter
gesteigert;
auch das
Anschwellen des Realismus, der die Abhängigkeit des Geisteslebens
von Naturbedingungen deutlich vor Augen
stellt
und
von außen her ableiten möchte, unterstützt das weiter.
in
alles
Innere
Erzeugt nun
vollem Gegensatz zu A. Smiths Optimismus die neueste Gestalt-
ung der Arbeit schwere Verwicklungen auf wirtschaftlichem Gebiet,
so kann es nicht Wunder nehmen, wenn von der Lösung dieser
Verwicklungen, von der Herstellung einer neuen ökonomischen Ord-
nung Heil
Die
für das ganze
dritte
Strömung
Leben erwartet wird.
ist
der
Politismus,
die
Schätzung
und
Zu den Problemen des Menschenlebens.
316
Wie viel im
Überschätzung des Staates.
1 9.
Jahrhundert dazu
trieb,
das hat uns schon früher beschäftigt; die Neigung, in allen Dingen
den Staat voranzustellen und ihm die Leitung aller Kulturarbeit zuzuweisen,
noch immer im Wachsen.
sichtlich
ist
und abgeschwächt,
verblaßt
jedenfalls nicht,
daß
Auch hier bringt
nur zu vollem und starkem Ausdruck,
Sozialdemokratie
die
in
die meisten beherrscht
was,
Zufällig
ist
es
Deutschland mit seiner Neigung zur Staats-
omnipotenz die Sozialdemokratie besonders rasch vordrang, während
bei
sie
den angelsächsischen Völkern langsamer Boden gewinnt.
Die Verbindung des Demokratismus, Ökonomismus, Politismus
an sich keineswegs nötig,
ist
Frage
ja die
liegt
nahe, ob sie nicht
ob im besondem nicht die vom Demo-
harte Widersprüche enthalte,
kratismus verfochtene Freiheit der Individuen mit der Zwangsgewalt
Wie immer es aber mit
des Staates unversöhnlich zusammenstoße.
der Berechtigung jener Verbindung stehen mag, zunächst
geschichtliche Tatsache,
ist sie
eine
und mit der Macht einer Tatsache ergreift sie
die Zeitgenossen. Auch haben jene Hauptrichtungen trotz aller Differenz
eine
innere Verwandtschaft, die namentlich in
merklich
Abgelehnt wird
wird.
durchgängig
der Verneinung beJenseitige
alles
und
Metaphysische, also auch eine selbständige Qeisteswelt; das Ganze
will
durchaus immanent,
will
bloße Diesseitskultur sein
und wird
damit bloße Menschenkultur.
Diese Grundüberzeugung spricht aus
dem Glauben
sie spricht
an die Masse,
aus der Voranstellung der
wirtschaftlichen Güter, sie erscheint in der Erhebung des Staates zum
Daher
Träger der Vernunft.
Gedankenwelt eine
in
religiöse
ist
die
Meinung
Überzeugung verbinden,
das religiöse Fahrwasser überleiten zu können.
bloßmenschliche Charakter
ihr
verfehlt,
ist
Bewegung
jener
mit jener
sie
wohl gar
Denn der säkulare,
wesentlich,
keineswegs von den Individuen nur nebenbei
er
angehängt.
ist
Es
um partielle Theorien, die so oder so zu
wenden sind, sondern um eine Gesamtgestaltung des Lebens und
handelt sich
hier
nicht
eine allumfassende Gedankenwelt, die zum ganzen
und
seine
ganze Seele verlangen.
Menschen sprechen
Das vornehmlich
gibt
der Be-
wegimg heute ihre Kraft, daß sie den ganzen Menschen fordert und
alle
Mannigfaltigkeit
seines
Strebens
einer
allbeherrschenden
Idee
unterwirft
Lebensentfaltungen sind nur Lebensentfaltungen gewachsen,
bloße Kritik,
sich
ihnen
alle
mag sie noch so geschickt und geistreich sein, verhält
gegenüber wie der flüchtige Schatten zu einem
festen
Gesellschaft und Individuum.
So
Körper.
sei
auch hier die Kritik auf das Notdürftigste dessen
was besonders
beschränkt,
317
und die Lebensanschau-
die Philosophie
ung betrifft. — Augenscheinlich ist zunächst der schroffe und unversöhnliche Gegensatz des Ganzen unserer Überzeugung mit dem in
Bewegung wirksamen
jener
Wir widersprechen
Lebensideal.
bloßen Menschenkultur;
entschiedenste aller
aufs
wir tun das, weil wir
im Menschen zwei Welten zusammentreffen sehen, und weil nur die
Ergreifung der höheren
unserem Leben einen Sinn und Wert zu
geben und es
Bewegung zu
rechte
in
greifung aber verlangt eine energische
Weltanblicks,
ersten
Menschen,
so
ist
setzen
vermag.
Er-
Jene
Umwandlung nicht nur des
mehr noch des eignen Wesens des
sondern
kräftige Aufrüttelungen,
Erhöhungen, Erneuerungen; nur
zu einer Geistes- und Wesenskultur zu gelangen und dadurch
dem Menschen eine innere Größe zu geben. Aus solcher Überzeugung widerstehen wir dem Demokratismus, weil er eine fälschliche
Idealisierung des sinnlichen
Menschen
vollzieht
und
die Geisteswelt
dem bloßen Menschentum unterzuordnen geneigt ist, widerstehen
wir ferner dem Ökonomismus, weil sein Bauen von außen nach
innen eine Leugnung selbständiger Probleme des Innenlebens ent-
und
hält,
weil er mit der Herstellung eines sorgenfreien, behäbigen
Zustandes das volle Glück des Menschen gesichert glaubt, verwerfen
wir endlich den Politismus, weil er die Selbständigkeit der Persönunterdrückt
lichkeit
und damit
Ursprünglichkeit des geistigen
die
Schaffens gefährdet, auch den Selbstwert der geistigen Güter bloßen
Zweckmäßigkeiten
aufzuopfern
bereit
Überall
ist.
hier
in
allem
äußeren Vordringen ein inneres Sinken, eine Behandlung der Hauptsachen
als
So
Aber
Nebendinge, ein geistiges Kleinwerden des Menschen.
ein
die
voller
und
Gegensatz
eine
entschiedene
Macht über die Menschheit gewinnen konnte, wie
Leidenschaften
viel
sondern auch
erregen,
edle Gemüter gewinnen konnte.
Zuspitzung
nähere
Verneinung.
bloße Verneinung läßt unerklärt, wie das Ganze so
Leben
das
viel
viel
es nicht bloß die
Aufopferung erzeugen,
Es müssen hinter dem, dessen
gefährdet,
allgemeinere
Probleme
wirken, die auch wir anderen nicht ablehnen können, die uns nicht
ruhen und rasten
lassen,
bis
sie
irgendwelche,
wenn nicht Lösung,
so doch Beschwichtigung gefunden haben.
Ein solches allgemeineres Problem steckt
mus;
es
ist
die Frage
in
dem Demokratis-
einer größeren Verbreitung
der Kultur und
einer gleichmäßigeren Verteilung ihrer Güter, einer kräftigeren Teil-
Zu den Problemen des Menschenlebens.
318
nähme der einzelnen Individuen am Geistesleben. Nach aller Arbeit
der Jahrtausende steht es damit noch immer kümmerlich genug.
Wie gering ist trotz aller Fortschritte noch immer das, was der weit
überwiegenden Mehrheit von den Schätzen der Bildung
dünn
die Schicht,
ist
innerlicheren Kultur teilnimmt!
lichem
wenig
wie
Christentum,
das
zufällt,
wie
Bewegung zu einer höheren und
Über tausend Jahre wirkt bei uns
die an der
es
ist
uns
dieser
in
nach
mensch-
Maße langen Zeit zu einer umgestaltenden Macht, zu festem
inneren
Besitz,
Viel zu sehr
ist
zu wesendurchdringender Überzeugung geworden!
es bei allem
Gerede von Fortschritt und Geistesleben
ein bloßer Überwurf über ein
Dasein geblieben,
zu
viel
von bloßen Naturtrieben beherrschtes
wenig sind die großen Gegensätze und
Spannungen, aber auch die großen Möglichkeiten, die unser Leben
für das Bewußtsein der Einzelnen herausgearbeitet.
Nunmehr beginnen wir - schon das ist eine Wendung zum Besseren —
enthält,
es
als
eine
Unwahrheit zu empfinden,
in
der
Lebens
Mehrzahl
Menschheit
wohl
daß eine höhere Art des
der Individuen innerlich fem
solches Gefühl einmal
wirksam
irgend
und fremd
ist,
aber
bleibt;
ist
der
ein
wach geworden, so wird es irgendwie zu be-
wenn im Kampfe nach solchen Zielen die
Schranken menschlichen Vermögens sich noch so bemerklich machen,
friedigen
es
sein;
selbst
besagt einen gewaltigen Unterschied,
ein Schicksal
Teilnahme
aller
ob die bisherige Lage wie
Kampf für eine stärkere
aufgenommen und damit die Menschheit der Schuld
hingenommen, oder ob
ein
möglichst entlastet wird.
Diese Erwägungen werden durch eine Wahrnehmung verstärkt,
der sich kein Unbefangener verschließen kann.
manche Zeichen
mus
des Lebens
Unsere Zeit enthält
eines Oreisenhaftwerdens: ein raffinierter Epikureis-
weiter
greift
Führung berufene Kreise zeigen
und
weiter
um sich, manche zur
sich geistesträge
ohne ihrem Leben einen wertvollen
und abgestumpft;
Inhalt zu geben, halten sie hohe
ist
es verwunderlich, daß die Überzeugung
immer mehr Boden gewinnt, daß es heute fast mehr noch als neuer
Ansprüche aufrecht;
Ideen neuer Menschen, frischer und einfacher Individuen, aufstreben-
der
und
geistesdurstiger
anerkennt,
braucht
noch
bekennen und die von
halten, das
Bevölkerungsschichten
keineswegs
sich zur
bedarf?
Wer
das
Sozialdemokratie zu
ihr erstrebte Erneuerung für die richtige zu
Verlangen nach Erneuerung aber wird er verstehen.
Der Ökonomismus droht mit seinem Anspruch auf die Führung
Gesellschaft und Individuum.
des Lebens
es
in
319
abschüssige Bahn
zu
wo ein selbständiges
kann nur da die Gemüter gewinnen,
er
leiten,
problematische und
eine
wo seelische Probleme fehlen. Aber die wirtschaftHebung ließe sich nicht als eine Befreiung von aller Not be-
Innenleben und
liche
grüßen,
lastete
Schwere
licher
die Sorge
nicht
um die Lebenserhaltung mit pein-
Menschen.
auf vielen
Es wäre gewiß kein Glück,
wenn der Tisch des Lebens den Menschen fertig bereitet würde und
sie nur zu genießen brauchten, wenn alle Sorge und aller Kampf
Aber
entfiele.
es
bleibt
um
daß diese eine Sorge
tieftraurig,
die
Lebenserhaltung so sehr überwiegt und das menschliche Sinnen und
Denken so zwingend
kommt
festhält,
wie
gewöhnlich
es
Es
geschieht.
damit ein schwerer Druck auf das Leben, der zu innerer
Kleinheit
lichkeit
und Erniedrigung wirkt und mit seiner stumpfen Alltäg-
und
frische
alles
freie
Großes geboren.
Not
viel
gibt
eine Armut,
Aufstreben
Aber
mit
hemmt.
Recht
sagt
Gewiß
hat die
Pestalozzi:
zur Emporbildung der menschlichen
die
»Es
Kräfte
und 'zur Grundlage seines Glückes und seiner inneren Größe dient.
Aber
es
VIII, 98).
gibt
auch eine Armut, die zur Verzweiflung führt" (Wke.
Vieles
in
ist
der Neuzeit zur Bekämpfung solches Druckes
und solcher inneren Erniedrigung geschehen; dürfen wir behaupten,
viel mehr geschehen könnte, daß nicht bloß in den
daß nicht noch
Gesinnungen der Individuen, sondern auch
hältnissen nicht
manches
sich anders
den allgemeinen Ver-
in
und besser gestalten ließe?
Das Bedenkliche des Politismus kam wiederholt zu Erwähnung;
nicht
nur der Freiheit der Einzelnen, auch der Seele des Gesamt-
lebens drohen von daher Gefahren.
gehen
sollte,
erträglich
Jahren.
so
„Wenn alles nach Vorschriften
müßte das Leben, das so schon schwer,
völlig un-
werden", so meinte schon Plato vor mehr als zweitausend
Aber warum dringt denn
vor, gerade
auch
die Staatsidee heute so mächtig
Kreisen, welche besonders
in
um die Freiheit be-
Doch wohl, weil das Individuum bei der Zerreibung
der überkommenen Zusammenhänge und dem völligen Unsichersorgt sind?
werden der eignen Stellung irgend welchen
es
seine
irgendwie auch
Existenz
schützt sehen will.
hinaus
auch
in
Das
das
greift weit
Innere
festen Halt ersehnt, weil
vom Ganzen
über
alle
geschätzt
und ge-
wirtschaftliche
und Ganze des Lebens.
Ein
Fragen
solches
Verlangen nach mehr Halt und Schätzung hat beim Zusammenbruch
des Altertums nicht wenig dazu beigetragen, der christlichen Kirche
die
Herzen
zu
gewinnen;
heute aber scheint es sich
wieder mit
Zu den Problemen des Menschenlebens.
320
Hüten wir uns, solche Bewegungen gering
und verborgen im Untergrunde des Lebens
neuer Kraft zu erheben.
zu achten, weil
sie still
Denn
erfolgen.
bereiten, die
sie
sind
in
es,
denen sich die seelischen Lagen
dann plötzlich mit unwiderstehlicher Kraft hervorbrechen
und das Ganze des sichtbaren Lebens in völlig neue Bahnen treiben.
Verschiebungen,
Innere
Ausdruck
gestattet
Umwandlungen, wenn
molekulare
heute
sind
ist,
im
Gange;
welche
menschlicher Verhältnisse daraus hervorgehen wird, das
dieser
Gestaltung
liegt
einst-
weilen in tiefem Dunkel.
Auch die Einheit der Gedankenwelt, die in der sozialdemokraBewegung wirkt, sei nicht zu gering angeschlagen. Wohl
muß bei unserer Abweisung aller bloßen Menschenkultur die be-
tischen
sondere Art dieser Einheit mit ihrer Vergötterung des Menschen
sie allein
macht es möglich, daß die Verzweigungen der Arbeit sich
gegenseitig unterstützen,
Mensch
als
Aber Einheit bleibt Einheit,
eine verhängnisvolle Irrung erscheinen.
in Tätigkeit
tritt.
und daß an jeder einzelnen Stelle der ganze
Das einzige System, das außerdem heute
eine allumfassende Einheit bietet,
ist
der kirchliche Katholizismus, der
aber mit seiner strengen Bindung an die mittelalterliche Denkweise
unvermeidlich in einen immer schrofferen Gegensatz zu den Beweg-
ungen der Zeit und den Bedürfnissen des modernen Menschen,
zu den inneren Notwendigkeiten des Geisteslebens gerät.
ja
Auf dem
eignen Boden der Neuzeit besaß die Aufklärung eine Art von Lebens-
und zugleich
einheit
ein allumfassendes Ideal;
seit ihrer
Erschütter-
ung befinden wir uns in starker innerer Zerklüftung, die immer
Im besonderen pflegen die, welche von der
unerträglicher wird.
Freiheit aus das Leben gestalten möchten,
spruch zu verfallen, daß
fähigkeit
gegen
des Menschen
eine
allein
und
ihrer
sich
leerer sich ihre
Freiheit
*
gegenbringen,
um
welche
eine
so
solche
sie
da-
Schätzung
des
mit Feuereifer bekämpfen
sicherer dünken,
Gedankenwelt gestaltet.
in
Würde, Leistungs-
genug rühmen können, daß
zu begründen vermag,
ung und Verflachung
trägt.
nicht
Weltanschauung,
Menschen
dem wunderlichen Wider-
praktisch die Größe,
sie
Indem
je
negativer,
je
sie so aller Vernein-
der Weltanschauung ihre Sympathie ent-
untergraben
sie
selbst
den Boden, der ihr Streben
Solche Unklarheit oder vielmehr Gedankenlosigkeit
ist
keiner
durchgreifenden Wirkung fähig.
So
ist
entscheiden
eine
schwere Krise nicht zu verkennen;
müssen,
ob die heutige Kultur und
es wird sich
Gesellschaft
die
Gesellschaft und Individuum.
Kraft enthält,
Zusammenfassung und
geistige
Erhöhung
zu vollziehen und damit der Auflösung Widerstand zu
des Lebens
leisten,
eine innere
321
oder ob
sie
eine solche nicht aufzubringen vermag.
Ersteren
könnte der Angriff nur dazu dienen, die Kultur auf ihre eigne
Falls
Tiefe zu bringen
und sie der Kleinheit des menschlichen Getriebes
zu entwinden; im anderen Falle müßte die heutige Kultur und Gesellschaft
untergehen, und sie würde solchen Untergang dann ver-
dienen.
Die
heit
Welt
geistige
und
steht sicher
fest
selbst, .sowie
stirne
über
den Wandlungen
könnte
sein,
daß
Geistigkeit
und
erst
eine
bloßen
der
krasse
Wirken zur MenschErdoberfläche.
Verneinung
eine Auflösung aller
erforderlich wäre,
ihr
über solchen Wandlungen, wie die Ge-
aller
unsichtbaren
es
Zusammenhänge
um der Menschheit auf dem Wege eines indirekten
zum Bewußtdem Leben wieder zu dem Wahr-
Beweises ihre Unentbehrlichkeit wieder nachdrücklich
sein
Ja
selbständigen
zu bringen und dadurch
heitsgehalt zu verhelfen, der heute schmerzlich vermißt wird.
Eucken, Grundbegriffe.
4. Aufl.
21
4.
a)
I
Probleme der Moral.
Die unsichere Stellung der Moral in der Gegenwart.
|ie Fassung wie
'-^ Sicherheit.
die Schätzung der Moral
ist
Un-
heute voller
Einmal erscheint gegenüber der Erschütterung der
religiösen und philosophischen Überzeugungen die Moral als ein fester
dem sich alles zusammenfinden kann, was sonst auseinanderfällt.
Denn wenn alles ins Wanken gerät, es bleibt der
Mensch und das Verhältnis zum Menschen, unser Zusammensein
Punkt, an
stellt
uns Aufgaben, die niemand bestreiten kann.
Bewegung zur ethischen
findet sich viel
So entstand eine
Kultur, über den besonderen Kreis hinaus
Bewegung, das Wohlsein der Mitmenschen zu fördern
und zugleich dem eignen Leben einen wertvollen
Inhalt zu geben.
Als Moral erscheint hier der Altruismus, das Wirken für andere, das
Zurückstellen der eignen Interessen vor denen der anderen.
Diese
Bewegung bildet ein Hauptstück der gegenwärtigen Kultur, breite
Wirkungen zur Hebung von Not und Leid, zur Milderung von
Strenge und Härte, zur Humanisierung des menschlichen Daseins
sind von ihr ausgegangen und gehen unablässig von ihr aus.
Aber inmitten solcher Leistungen
Bedenken
und Widerspruch.
Moral
Altruismus nur deshalb
als
tieferen
verbleibt
Vielleicht
so
im Prinzipiellen viel
uns
bei
der
zusammen,
weil
die
finden
leicht
wir
Probleme der Sache zurückgeschoben, wenn
werden.
Ist
es
nicht geleugnet
denn ausgemacht, daß Moral mit Altruismus,
mit
einem Handeln für andere zusammenfällt? Der Ausdruck Altruismus
stammt aus der Gedankenwelt Comtes, das heißt aus einem Systeme,
das alles Beisichselbstsein der Seele
und gar
in
das Verhältnis
die hier der Moral
anderen gelten?
setzen
die
In
zur
preisgibt
Umgebung
läßt;
sollte
gegebene Fassung ohne weiteres auch für uns
Wahrheit heißt Moral und Altruismus gleich-
Moral im Umfang einengen und
Erschöpft denn
und das Leben ganz
aufgehen
das soziale und
im
Inhalt
verflachen.
humane Wirken den ganzen Um-
Probleme der Moral.
kreis der Moral, tragen wir nicht
auch große Aufgaben in uns selbst,
Bildung der eignen Seele, auch
in der
323
in
dem Verhältnis zur Welt
und zu den Dingen ? Durch das Ganze unseres Lebens geht das Ent-
— oder, ob das Handeln unserer eignen
weder
Es kann
sachlichen Ziele dient.
oder ob es einem
Lust,
B. das Schaffen eines Künstlers
z.
durch Beweggründe verschiedenster Art geleitel sein.
Der Schaffende
kann Vorteil, Ruhm, Anerkennung suchen, er kann die Wünsche
und Launen des ihn umgebenden Publikums zu befriedigen trachten,
er
kann endlich
gegen
und allein der inneren Notwendigkeit des
lediglich
wenn es sein muß, diese mit heroischer Kraft
Schaffens folgen und,
Widerspruch der Umgebung
allen
durchsetzen;
ist
bis
Treue gegen sich
solche
nicht auch ein Handeln moralischer Art?
zu eigner Gefährdung
selbst
und
Sache
die
Ähnlich kann den Forscher,
ähnlich den religiös Überzeugten die geistige Selbsterhaltung in den
härtesten Widerspruch zur Umgebung führen und ihn ihre Ruhe, ihr
Behagen gründlich zu erschüttern zwingen;
ja es
mag die ganze Be-
wegung zur Geistigkeit mit ihren Ansprüchen, Sorgen und Zweifeln
als
und
eine Störung des Gleichgewichts
Wohlseins
mittelbaren
erscheinen;
moralische Aufgabe anzuerkennen?
ist
als
eine Feindin
trotzdem
Trifft
nicht
in
des uneine
ihr
aber dieses zu, so
ist
die
Moral sicherlich etwas tieferes und besseres als der bloße Altruismus.
Auch das
spricht wider
diesen,
daß er die Moral
nicht aus
der eignen Tiefe der Seele zu begründen, nicht sie zu einer Sache
machen versteht
geistiger Selbsterhaltung zu
Zu Gute kommt ihm
aber bei den Menschen die Unsicherheit, die heute über einer solchen
Begründung
Von
waltet.
der Vergangenheit her wirken
auf uns
zwei Gedankenwelten mit einer innerlicheren Art der Moral: die der
und
Religion
Verhältnis
die
immanenten Idealismus.
eines
Dort war es das
zu einem weltüberlegenen Sein, hier war es die eigne
Vernunft des Menschen, woraus lebenumspannende Aufgaben und
eine
ethische
Bewertung
Beide Arten aber sind
in
ihrer
Grundlage
all
unseres
Handelns hervorgehen
dem Geistesleben
erschüttert,
sondern
sollte.
der Gegenwart nicht nur
auch
ihrem
Inhalt
nach
vielfach
bedenklich geworden.
gänzlich
verschwunden, aber auch die einer immanenten Vernunft
ist
Die Welt der Religion
ist
für viele
mehr und mehr verblaßt; zugleich aber erscheint dem Hauptzuge
der Zeit die religiöse Moral als zu weich und zu passiv, die Ver-
nunftmoral aber
auch
als
zu rauh.
als
So
zu abstrakt und mit ihrer strengen Pflichtidee
bleibt als der einzige unbestrittene Punkt die
21*
Zu den Problemen des Menschenlebens.
324
soziale Moral mit
ihrem Altruismus, dieser aber hat sich uns als viel
Demnach bleibt zunächst nur die Tatsache
zu eng und flach erwiesen.
festzustellen,
daß unsere Zeit überhaupt keine Moral
dem
vorhandenen Stande der weltgeschichtlichen Evolution
in
ihr
besitzt,
welche
des Geisteslebens entspricht, daß sie einer charakteristischen,
ihre
Ihrem innersten
innersten Bedürfnisse befriedigenden Moral entbehrt.
Wesen nach angesehen, ist die Moral uns heute mindestens ebenso
unsicher als die Religion. ^ Wie sehr solcher Mangel an einer eignen
Moral die Kraft der Moral in unserer Zeit herabsetzt, wie sehr er
es
den Gegnern der Moral
werfen und
Karikaturen von ihr zu ent-
erleichtert,
und
mit der Verspottung dieser sie selbst für widerlegt
abgetan zu halten, das
stellen
uns zahlreiche Eindrücke der Gegen-
wart mit voller Klarheit vor Augen.
Wir werden solchen Verwick-
lungen nicht gewachsen werden, wenn es nicht gelingt, durch Selbst-
besinnung und Selbstvertiefung des Lebens wieder zu einer eignen,
selbsterlebten
Moral
zu
Hier
gelangen.
liegt
das
vielleicht
alier-
dringendste Bedürfnis der Zeit.
b)
Moral und Metaphysik.
Weithin reicht heute die Neigung, die Moral von den Problemen
der Weltanschauung
zu
selbst
erfassen.
gänzlich
abzulösen
Viele sehen
darin
zugleich eine Vereinfachung des Lebens,
bilder
ist
und unmittelbar
eine
bei
sich
große Befreiung
und
manche geschichtliche Vor-
werden zur Unterstützung dessen herangerufen, im besondern
es der große Name Kants, der dies Unternehmen sanktionieren soll.
Gewiß bestand von altersher die Neigung,
Verwicklungen
alle
der Weltprobleme von sich zu werfen und sich auf ein rechtschaffenes
Leben zurückzuziehen; der Einzelne mag
ein
gutes
Recht haben,
das zu tun, aber hat es auch das Ganze der Menschheit ?
die
Setzt nicht
Wendung des Einzelnen voraus, daß unabhängig von ihm eine
Moral gesichert und anerkannt
^
Trotz solcher Unsicherheit
ist?
Im besondern kann nur eine
im Fundament
bücher und Kompendien der Moral in Hülle und
bietet
Fülle.
unsere Zeit Lehr-
Und warum auch
wenn er zu dem Worte Hamlets, es gäbe
Menge Dinge im Himmel und auf der Erde, wovon nichts in unseren
Kompendien stünde, bemerkt: «Gut, aber dafür stehen auch wieder eine
Menge von Dingen in unseren Kompendien, wovon weder im Himmel noch
nicht? Lichtenberg hat wohl recht,
eine
auf der Erde etwas
vorkommt« (s. Vermischte Schriften [1801] II, 356).
Probleme der Moral.
gänzliche
Verkennung Kants ihm
probleme von
die Absicht zuschieben, alle Welt-
sich zu werfen und
tischen Arbeit zu
in
Denn
flüchten.
325
den sicheren Hafen der prak-
Denken bewegt
sein
sich
nicht
um den Gegensatz von Theorie und Praxis, sondern um den von
wo
und praktischer Vernunft;
theoretischer
Frage kommt, da handelt es sich immer
aber die Vernunft in
um Weltzusammenhänge;
vom Ganzen der
so verzichtet Kant nicht auf letzte Überzeugungen
anderen
einer
den Punkt, der über sie entscheidet, an
er sucht nur
Wirklichkeit,
Stelle
als
die
alte
Spekulation,
nicht zum Mittelpunkt seiner Gedankenwelt,
Ordnung der Dinge, eines intelligiblen Reiches der Ver-
einer neuen
nunft zu machen.
Kant
Metaphysiker
durch
ein
macht die Moral
er
ohne sie zur Erscheinung
ist
ein Metaphysiker eigner Art, aber er
und durch,
die Alltagsweisheit
ist
von dem
Vorrang der praktischen Arbeit über das Denken hat an ihm keinen
Bundesgenossen.
In
Wahrheit brauchen wir das Phänomen der Moral nur etwas
um zu gewahren, daß es mit dem nächsten
genauer zu betrachten,
Weltanblick aufs
schroffste
Lebens und Strebens
immer
versteckt
wieder
verschieden
eine
Sobald wir an einer
Handlung entdecken,
die
Ablösung des
daß
als
moralisch ge-
Gesinnung,
die
wenn auch
und auf Umwegen, zu den Zwecken der Selbsterhaltung
zurücklenkt,
Nun
nichtet.
enthält sie
vom bloßen Ich, ein Hinauswachsen über die
natürliche Selbsterhaitung.
priesenen
Wie
zusammenstößt
Moral gefaßt werden mag,
zeigt
gilt
ihr
freilich
Charakter
moralischer
uns
als
ver-
schon die Natur gewisse Ansätze einer
Befreiung des Lebens von der bloßen Selbsterhaltung, aber diese
bleiben zerstreut
Wendung,
und mit fremdem
verquickt;
so bedeutet es eine
Umwälzung, wenn die neue Art des Handelns
eine reine Ausbildung erlangt und die Herrschaft über das Leben
fordert Neue Größen, neue Werte treten damit ein; bedarf es nicht
einer neuen Welt, um ihnen Kraft und Zusammenhang zu geben?
ja
eine
Einen moralischen Charakter kann ferner das Handeln nur behaupten,
liches
wo es aus freiem Entschlüsse hervorgeht und ein ursprüng-
Leben
betätigt.
Wo irgend sich daher herausstellt, daß ein
vermeintlich moralisches Tun aus bloßer Gewöhnung, mechanischem
Zwange, autoritativem Drucke hervorgeht,
scheidung und Zuwendung
auszeichnende Art
Nun
und
in sich trägt,
fällt
aus
der
nicht
eine
eigne
Ent-
da verliert es sofort seine
moralischen Sphäre
heraus.
aber bietet jener Selbsttätigkeit und jener freien Entscheidung
Zu den Problemen des Menschenlebens.
326
die natürliche
Welt mit
den mindesten
Platz,
ihrer
ihr
durchgängigen Kausalverkettung nicht
Gefüge widerstrebt jeder Lockerung.
Be-
steht also nur ein Reich der Natur, und bleiben seine Ordnungen
maßgebend auch für das Geistesleben, so ist kein
Platz für irgend-
welche Moral, die mehr als eine Polizei des gesellschaftlichen Lebens
sein will.
Wo die moralische Forderung volle Selbstbewußtheit gewinnt,
da erhebt
den Anspruch,
sie
allen übrigen
überlegen zu sein, da "verwirft
mäßigkeit und
stellt
Leben
Sie steht
hinein.
Zwecken unvergleichlich
Erwägung der bloßen Zweck-
sie alle
unmittelbar in das menschliche
ein Absolutes
und fällt mit dem Wort:
«Was
dem Menschen, wenn er die ganze Welt gewönne und
an seiner Seele?"
Aber wie
ist
eine neue Art der Wirklichkeit
sich
alle
hülfe es
Schaden
das möglich, und bedeutet es nicht
eine ungebührliche Überspannung,
müssen
litte
wenn hinter jener Schätzung nicht
Denn im nächsten Dasein
steht?
Zwecke zueinander fügen und nacheinander be-
messen; hier gibt es nichts Absolutes, das sich unvergleichlich über
andere hinaushöbe.
alles
So
enthält
trachten, die
die
Moral,
von welchen Seiten her wir
Forderung einer neuen Welt,
sie
be-
es liegt in ihr eine
Um-
kehrung des nächsten Anblicks der Dinge und damit eine Metaphysik.
Wir werden also die Metaphysik nicht los, wenn wir uns
zur Moral begeben;
machen wir Ernst damit, von
physische fernzuhalten, so drücken
licher Flachheit herab.
ihr alles Meta-
wir sie unvermeidlich zu kläg-
Andererseits hat es gewiß einen guten Grund,
die Moral nicht an die verwickelten Gedankengänge der älteren Spekulation
zu binden,
jenseit ihrer
Einen
bietet
sie
nicht als ein sekundäres
Phänomen von einer
begründeten Weltanschauung abhängig zu machen.
Weg zwischen solcher Scylla und Charybdis hindurch
uns wiederum unser Begriff des Geisteslebens als der Wendung
dem Zusichselbstkommen des Weltprozesses, der Erringung eines Wesens und Sinnes
der Wirklichkeit zu einem Eigen- und Innenleben,
gegenüber allem
erhaltungen
der
sinnlosen
Gewebe der Beziehungen und
Selbst-
Mit Anerkennung
neuen
bloßen Punkte.
dieser
Welt sinkt die Natur notwendig zur zweiten, zu einer niedrigeren
Form des Seins. Wie aber die höhere von
unablässiger Selbstttätig-
keit
getragen werden muß, so
erst
zu erwecken und von ihr selbsttätig anzueignen.
aber
als solche selbsttätige
ist
sie
auch an jeder einzelnen Stelle
Nichts anderes
Aneignung der Geisteswelt ist die Moral,
Probleme der Moral.
327
sie ist
damit ein Durchdringen des Lebens zur Wesen- und Wahrhaf-
tigkeit,
das Gewinnen eines neuen, unendlichen Selbst, ein Unendlich-
verden von innen
Denn das erkannten wir als der Geistesstufe
her.
daß hier jedes Einzelne unmittelbar am Gesamtleben
wesentlich,
teil
hat, nicht erst durch die Vermittlung des Punktes es zugefiihrt erhält.
Bei solcher Fassung
ist
die Moral an erster Stelle eine Beweg-
ung innerhalb des eignen Lebenskreises,
jetzt
Weltcharakter
einen
selbst unmittelbar eine
erweist,
Streben zu unserem
ein
Aber indem
eine Erringung des eignen Wesens.
Selbst,
erscheint
dies Wesen
der Arbeit an uns
in
Bewegung der Welten.
Das ist es, was uns
die engste
Verbindung von Moral und Metaphysik verlangen
was uns
eine
läßt.
Moral
ohne Metaphysik
ein
als
heißt,
Unding erscheinen
Die Moral verlangt nicht etwa bloß zu ihrer Erklärung Welt-
begriffe,
sondern durch ihr eignes Dasein entwickelt sie unmittelbar
eine neue Welt
und umfängt uns damit
in
einleuchtender Gegen-
Die Verbindung der Moral mit der Metaphysik verwerfen
wart.
kann nur, wer entweder unter Metaphysik jene alte Schulmetaphysik
versteht, die aus vermeintlicher Denknotwendigkeit
zur vorhandenen
Welt eine neue hinzu ersann, oder wer die Moral zu einer bloß-
Ordnung, zu einer Polizei des Lebens herabdrückt.
gesellschaftlichen
Denn dafür bedarf es allerdings keiner neuen Welt, aber es ist dann
auch jene Lebenspolizei nur dem Worte nach Moral. Nach unserer
Überzeugung sinkt alle Moral zu einem bloßen Schein, wenn nicht
das Geistesleben,
Aneignung
dessen
sie
vollzieht,
den Kern
der
Wirklichkeit bildet.
Die dargebotene Fassung der Moral
ist
den Problemen und
Schwierigkeiten
gewachsen, mit denen die Moral zu tun hat, und
an die sich
Irrung und Mißverständnis knüpft.
hier
viel
Die Moral
ist
an erster Stelle Lebenserhöhung, Gewinn eines wahren Selbst
gegenüber einem nur scheinbaren, Aneignung der ganzen Unendlichkeit.
Aber diese Erhöhung entwickelt sich nicht vom unmittelbaren
Dasein aus durch eine bloße Steigerung der Natur, sondern
im Gegensatz zu jenem
sie erscheint
ergriffen sein,
Aufgabe, eine Forderung, ein Gebot.
grenzungen und Verneinungen
damit
sie will
als
eine
Aber was dies Gebot an Be-
enthält,
dient
letzthin
der Lebens-
bejahung; der Pflichtgedanke, der hier entsteht, entspringt dem eignen
Wesen,
nicht
von außen
her.
So eine Bejahung des Lebens, die
keine Vergötterung der bloßen Natur und Selbstheit bedeutet, sondern
sich
durch ein entschiedenes Nein davon abgrenzt.
Zu den Problemen des Menschenlebens.
328
Bei dieser Fassung besagt die Moral nicht eine bloße Leistung
gegebenen Welt, sondern das Erringen einer neuen Welt,
in einer
nicht
einen
Kampf
sondern einen Kampf um
um eine neue Art des Wirkens,
innerhalb der Welt,
ganze Welten; es handelt sich nicht
um eine neue Art des Seins, die sich freilich unablässig
sondern
Wirken umsetzen muß.
Im Menschen
nunmehr verschiedene Stufen der Wirklichkeit, ja
ganze Welten, und seine Entscheidung gehört dazu, welche davon
für ihn zur Hauptwelt werde. Ja, indem er nunmehr an seiner beein
in
entsprechendes
begegnen
sich
sonderen
Stelle die
höhere Stufe der Wirklichkeit zu behaupten
hat,
indem die neue Welt hier nur durch seine Tat zur vollen Verwirklichung kommt,
Handeln
wächst sein
über
den
einzelnen
Punkt
hinaus und gewinnt Bedeutung auch für den Weltstand.
sicherste
Befreiung
von der bloßen
Ichheit,
ein
So die
Weitwerden der
Seele, die Erhebung über alle bloße Zweckmäßigkeit, eine unvergleich-
Größe und Würde des Menschen.
liche
Mit der Größe zusammen
freilich
auch schwere Verwicklungen.
Denn in jener Weise läßt sich die Aufgabe nicht steigern, ohne daß
in
der menschlichen Lage weiteste Abstände
und
härteste
Wider-
Vor allem hält die natürliche Welt den
Menschen fest beim bloßen Ich, die Bewegung zur Geistigkeit dringt
stände ersichtlich werden.
demgegenüber wenig durch,
sie
zu bloßem Scheine zu sinken.
droht ein bloßer Vorsatz zu bleiben,
Deutlich erhellt, wie, an der Kraft
des bloßen Menschen gemessen, etwas Unmögliches gefordert wird;
muß der Mensch mehr werden als bloßer Mensch; wie könnte
Wendung im Weltleben erfolgen ohne die Kraft einer Welt?
muß
eine Weltkraft von vornherein im Menschen wirken, es
So
muß dem Tun ein Empfangen, dem Aufsteigen ein Gehobenwerden
entsprechen, in der Freiheit selbst muß eine Gnade ersichtlich werden.
In dem allen vollziehen sich große Wandlungen des ersten Anblicks;
so
eine
das anfängliche Ja wird unerträglich, aber aus
dem Nein erhebt sich
Große Forderungen und schwere Erschütterungen,
ergreifend und verwandelnd,
viel Unfertigkeit und viel Unsicherheit, viel starrer Widerstand und
lähmende Hemmung. Aber inmitten der Zweifel und Widerstände
ein
neues
Ja.
gewaltige Lebensfluten den Menschen
ein Aufrechterhalten des Lebens, die Eröffnung größerer Tiefen, das
Gewißwerden einer inneren Unendlichkeit.
zeigt
die Moral,
Wenn irgend etwas, so
daß unser Leben nicht gleichgültig
deutendes in ihm vorgeht.
ist,
daß Be-
Probleme der Moral,
c.
(Ethische
Moral und Kunst.
und ästhetische Lebensanschauung.)
Daß Kunst und Moral von
und
Zusammenstoß
vielfachem
menschlicher
Folge
329
Irrung,
Spannung
altersher in feindlicher
stehen,
einen
hat
es
keineswegs
bloß
Grund
der
ist
in
eine
Sache.
Beide Gebiete scheinen das Leben unter entgegengesetzte Aufgaben
Die Moral verlangt eine Unterordnung
und Schätzungen zu stellen.
unter allgemeingültige Gesetze,
dagegen die freieste Ent-
die Kunst
faltung der eignen Individualität; die Moral spricht mit dem strengen
Gebote der Pflicht, die Kunst beruft zum
freien Spiel der Kräfte;
die Moral hat ihre Stätte in der reinen Innerlichkeit
und
ist
geneigt
von der sichtbaren Leistung gering zu denken, die Kunst
schätzt
Weg zur Verkörperung findet
Zur
richtigen
Wür-
und
es
vorteilhaft
sein,
nur,
was den
digung dieses
Gegensatzes
Streites
ein^n Blick auf die geschichtliche
weil
halb,
zur Ablösung
das
augenblicklicher
wird
Bewegung zu werfen, schon des-
des Problems
von der
Zufälligkeit
Stimmungen dienen mag.
a.
Die Geschichte des Problems.
Merkwürdigerweise hat das alle anderen Völker in künstlerischer
Leistung überragende griechische Volk in seiner Gedankenarbeit der
Kunst keinen
bedeutenden Platz
die Anklagen
gegen die Kunst
Denkern:
gegeben.
als
der größte Künstler unter den
Die verschiedensten
Plato.
Kein Geringerer führt
Richtungen
verbinden sich ihm zur Bemänglung der Kunst.
einem
zum
wesenhaften
seines
Strebens
Sein Verlangen nach
und unsinnlichen Sein drückt ihm
die
Kunst
Schein des bloßen Scheines herab, anstößig wird ihm femer
der bunte Wechsel ihrer Gestalten, wie ihn vornehmlich das Drama
zeigt,
anstößig
kreises,
die
Unlauterkeit
des
mjrthologischen Vorstellungs-
der sie beherrschte, anstößig endlich die fieberhafte Erregung
des Gefühlslebens, die er
immer
weiter vordringen sah.
durch solche Anklagen ging die Kunst ihren
Führung des antiken Lebens.
Virtuosität
verlief,
liche Spielerei,
bald
in
Unbeirrt
Weg und behielt die
Aber je mehr sie sich
in subjektive
barocke Überspannung, bald in weich-
und je mehr die Glätte der Form einen bedeutenden
Inhalt verdrängte, desto stärker
wurde der Rückschlag einer herben
und harten Moral, desto mehr wurden Kynismus und Stoizismus
Zu den Problemen des Menschenlebens.
330
Zuflucht
eine
dem Genüsse
des Schönen
zu
selbständigen
Wertes
die
Verbindung mit der aufsteigenden
religiösen
die
Seelen,
stolzer
huldigen verschmähten.
Die
volle
Kunst
erst
ung,
hat
in
die
Kunst
vertieft.
Wirklichkeit
Nach
Plotin
dabei
Plotin
des Niedern
wältigtwerden
die Seele,
eines
namentlich bei
sie
welche
Anerkennung
vollzieht
den
sich
auch
die
Aufgabe der
beim Schönen
durch das Höhere,
Be-
ein
des Körpers durch
den Gedanken; das Schaffen versenkt
des Stoffes durch
sich nicht in
hat
Beweg-
Verinnerlichung,
die
erreicht;
erfuhr,
hat
sondern es bleibt bei sich selbst und geht
Stein,
von Seele zur Seele; nur
als
Übermittlung des seelischen Standes
Werk einen Wert. Die Kunst ist nicht eine bloße
Nachahmung der Natur wie bei Plato, sondern sie sucht die in
hat das sichtbare
jener
wirksame höchste Vernunft abzubilden und kann dabei ganz
wohl
mehr
stimmung
erreichen
läßt
solche
die
als
religiöse
Grund-
Schätzung des Schönen weit mehr der Be-
trachtung des Alls zugute
kommen,
Eine künstlerische
Schaffen triebe.
Aber die
Natur.
Ganze des Lebens, aber
sie
als
daß
sie
zu künstlerischem
Stimmung durchwaltet
hier
das
scheut vor greifbarer Gestaltung eher
zurück, als daß sie dieselbe suchte.
Das Christentum konnte den Schwerpunkt des Lebens
nicht
vom Künstlerischen ins Moralische verlegen, ohne daß die Schätzung
und Stellung der Kunst zunächst den schwersten Abbruch erfuhr.
Auch die besondere Art der Kunst des ausgehenden Altertums konnte
die Abwendung von ihr nur unterstützen.
Aber was in der Breite
des Daseins sich oft recht unerquicklich ausnimmt und nicht selten
in eine Geringachtung aller Form verfallen läßt,
das hat auf den Höhe-
punkten die entschiedenste Gegenwirkung gefunden; die Verinnerlichung des Seelenlebens durch die Religion hat auch hier die Kunst
in
neue Bahnen geführt.
So vornehmlich
bei
Jesus selbst.
überhaupt von den Begründern der geschichtlichen Religionen
Was
gilt,
daß nur das Vermögen einer hervorragenden schöpferischen Phantasie
sie
einer
unsichtbaren
Gegenwart geben,
das
gilt
Welt eine anschauliche und
ja sie
überwältigende
zur Hauptwelt des Menschen machen
ließ,
ganz besonders von Jesus; bei ihm gewinnt jenes Bilden eine
besondere Wärme, Zartheit und Innigkeit.
Daß er der Menschheit ein
Reich Gottes als ein Reich echter Liebe und kindlichen Vertrauens mit
deutlichen Zügen vorhielt, damit schlummernde Gefühle erweckte und
den Gemütern eine
tiefe
Sehnsucht einpflanzte, das hat dem mensch-
Probleme der Moral.
Dasein
liehen
solche
künstlerische
eine
Verklärung gebracht;
eine
dem Entdecken
der
mit besonderer Klarheit hervor in
tritt
Reinheit,
auch
331
Unschuld und Hingebung des kindlichen Lebens und
dem wunderbaren
Gestalten
einfachster
nissen menschlicher Seelenstände.
So ist hier bei Zurückstellung aller
Kunst der seelischen Kunst eine sichere Bahn
sinnlichen
in
Naturvorgänge zu Gleich-
bereitet.
vom GriechenSo bei Gregor von Nyssa, so auch bei Augustin. Wohl
Später wirkte der Schönheitsgedanke immer stärker
tum
her.
haben über diesen kunstfeindliche Stimmungen eine große Macht
gewonnen, seine Wendung zum Christentum ward zum guten Teil
durch einen
tiefen
Überdruß an einer
formalliterarischen
Bildung,
durch ein Verlangen nach einem Inhalt des Lebens hervorgetrieben.
Aber
der
in
eignen Gedankenwelt hält er
am Schönen
einen Aufstieg zur allumfassenden Einheit bildet
als
Werk und Zeugnis
ein
staltet
sich
ethischer
ihm
Art,
schließlich
dieser Einheit
und
verstehen
fest,
das
alle
Vielheit
lehrt.
So ge-
das ganze Weltall zu einem Kunstwerk
zu einer Ordnung, die Gerechtigkeit und Liebe zu
voller Ausgleichung bringt;
zugleich
ist
Augustin selbst ein hervor-
ragender Meister des Worts, seine Sprache hat die ganze Kraft und
Weichheit eines zwischen den Gegensätzen des Daseins hin- und her-
bewegten Geistes
in
sich
aufgenommen, durch ihn hat die lateinische
Sprache einen wunderbaren musikalischen Klang erhalten, und ist sie
zu einem angemessenen Gefäß einer
lichkeit
in
sich selbst vertieften Inner-
geworden.
Wie das mittelalterliche Kirchensystem durchgängig eine gewisse
Ausgleichung der großen Gegensätze brachte, so hat es auch dem
Schönen einen Platz innerhalb seiner Ordnung gegeben. So im
Gesamtbau des Lebens, indem die Sorge für die Ordnung und
Harmonie des Ganzen
Gestaltung,
indem
hier sehr hervortritt, so auch in der näheren
die Kunst der Verherrlichung
der Religion und
der Kirche in mannigfachster Weise zu dienen hat.
Die Neuzeit mit ihrer größeren Lebensenergie und ihrer Schärf-
ung aller Gegensätze zersprengt die mittelalterliche Ausgleichung,
während ihres ganzen Verlaufes hat der Kampf und Gegensatz nicht
aufgehört.
Ihr Beginn in Renaissance und Reformation gibt dem
Gegensatz sofort den stärksten Ausdruck.
In
der Renaissance ge-
winnt eine ästhetische Welt- und Lebensanschauung zuerst eine volle
Bewußtheit.
Das Schöne wird
hier
zum Hauptwerkzeug
der Ent-
wicklung des Lebens, das wichtigste Mittel zur Herausarbeitung aller
Zu den Problemen des Menschenlebens.
332
zur Selbstaneignung und Selbstgenießung des Menschen; die
Kraft,
Höhe
Kunst lehrt das Leben sich selber finden, seine eigene
Zugleich wirft das Leben
reichen.
überwiegend der nächsten Wirklichkeit zugewandt, erstrebt es
sich;
in
er-
unsichtbaren Bindungen von
alle
ihrer äußeren
kenloses Glück.
man
empfindet
und inneren Unterwerfung ein volles und schranBei starkem Lebensdurst und stolzem Selbstgefühl
leicht
die
Moral
als
von draußen auferlegte
eine
Fessel, als eine starre Satzung" und widerwärtige
tiger die
Individualität,
desto berechtiger scheint
Hemmung; je kräfsie,
Hemm-
solche
ung abzuschütteln und allein der eignen Neigung zu leben. So der
Immoralismus der Renaissance, ein Hauptgrund ihres Zusammenbruchs
als
nicht
es
und denen
die Kunst,
sich
zugleich
dafür
eines
ihrer
Höhe aber
fehlt
mit Einsetzung des ganzen Wesens erfaßt,
zu einem ethischen Lebenswerke
Michelangelo
pflanzt die künstlerische
fort,
Auf
weltbeherrschender Macht.
an Gestalten, welche den Gegensatz überwunden haben,
zu
gedenken.
Bewegung sich
—
in
gestaltet;
Von
der
es
genügt
Renaissance
das Barock und Rokoko
auch zieht die Renaissance die Geister immer von neuem zu
sich zurück.
Die Reformation hat ihre Stärke
in
der entschiedenen Hervor-
kehrung der Moral und der Steigerung der persönlichen Verantwortlichkeit, weit
über den Kreis ihres eignen Bekenntnisses hinaus
War
hat sie damit einen gewaltigen Ernst in das Leben gebracht.
schon solche Verinnerlichung unmittelbar der Kunst nicht günstig,
so
kam
dem
dem ein
hinzu, daß jene mit der Fülle ihrer sinnlichen Gestalten
Menschen den Zugang zu Gott zu erschweren
schien, zu
unmittelbares Verhältnis zu gewinnen die allüberragende Hauptsache
So konnte ein heftiger Zorn gegen alles Bild und allen
Schmuck als gegen eine Verdunklung der lebendigen Gegenwart Gottes
in der Seele und als eine Veräußerlichung und Verweichlichung des
Lebens entstehen. Aber wenn damit kunstfeindliche Stimmungen
weithin Boden gewannen, so hat auch hier auf der Höhe des
wurde.
Schaffens die Kunst in anderer Richtung mannigfachste Förderung
sie ist nur mehr von der sinnlichen Anschauung abgeund tiefer in die Seele zurückverlegt. Die Namen eines Luther
empfangen,
löst
und eines Bach genügen,
um das zu zeigen.
Die verstandesmäßige Art der Aufklärung mit ihrem Dringen
auf logische Klarheit
und kluge Zweckmäßigkeit, auch mit ihrer un-
geschichtlichen Denkweise
war der Kunst wenig günstig und
stellte
Probleme der Moral.
sie
weit hinter die Moral zurück.
Diese selbst aber gewann dabei
Um so stärker wurde das Verlangen nach
keine besondere Tiefe.
Schönheit, als durch das Aufsteigen des
Epoche begann.
333
Neuhumanismus eine neue
Es strebt aber auf seiner Höhe, bei den führenden
deutschen Dichtern und Denkern, Gutes und Schönes zusammen und
fördert sich
stein
gegenseitig.
Daß Kant
Selbständigkeit
zum Eckdem Schönen eine
die moralische Idee
des Lebens machte, hat ihn nicht verhindert,
und einen Selbstwert zuzuerkennen; er zuerst hat es
gegen das Gute wie das Angenehme mit voller Deutlichkeit abgegrenzt, er hat es
allen
ein
im eignen Innern der Seele begründet und es über
bloßen Nutzen und Genuß sicher hinausgehoben.
Goethe «die großen Hauptgedanken der
So konnte
Kritik der Urteilskraft
seinem bisherigen Schaffen, Tun, Denken ganz analog" finden. Goethe
war bei aller Größe seines künstlerischen Schaffens von
selbst aber
einer Geringschätzung der Moral
und zugleich von dem Bekenntnis
zu einer ästhetischen Weltanschauung weit entfernt
viel
Dazu war ihm
zu sehr sein künstlerisches Schaffen ein ernstes und mühsames
Suchen seines eignen innersten Wesens,
ein gewissenhaftes Arbeiten
an sich selbst; eine laxe Auffassung dieser Probleme hat kein Recht,
sich auf Goethe zu berufen, wenn man ihn im Ganzen seines
Wesens nimmt, nicht auf einzelne Äußerungen festlegt So wenig
er die Kunst und eine künstlerische Kultur durch »konventionelle
Sittlichkeiten",
wollte,
er
durch
„Pedanterie
gab dem Menschen
und Dünkel" beschränkt wissen
in der
Forderung, die Ordnung der
Welt mit Freiheit zu ergreifen und sich
setzen,
eine
selbst
Grenze zu
seine
moralische Aufgabe, die sein ganzes Leben
und ihm überall eine hohe Aufgabe stellt
umspannt
Schiller endlich, zwischen
dem Dichter und dem Denker stehend, war unablässig um eine Versöhnung von Gutem und Schönem, der „Freiheit in der Erscheinung" bemüht; „die hohe Reinheit des
sittlichen
Standpunktes bei
der vollsten Anerkennung des künstlerischen Lebens in seiner Selbständigkeit
ist
das
Eigenartige,
Denkweise" (Kühnemann).
Die
ja
Einzigartige
Ströme haben
der
sich
Schillerschen
dann
wieder
geschieden, die Romantik gab dem Vorrang der Kunst und der ästhe-
Lebensanschauung einen besonders zugespitzten und selbstbewußten Ausdruck, Fichte und die anderen Führer der nationalen
tischen
Bewegung wirkten kräftig zur Verstärkung der Moral. Die soziale
und technische Kultur, die im Verlauf des 19. Jahrhunderts immer
mehr zur Herrschaft kam, ist mit ihrer Richtung auf die gesellschaft-
Zu den Problemen des Menschenlebens.
334
Wohlfahrt und auf die Zweckmäßigkeit des Lebens die Kunst
liehe
erhebt
Anhang zu behandeln
nebensächlichen
als einen
nun
sich
die
moderne Kunst,
dem
das Ganze des Lebens,
sie
mehr
Dagegen
geneigt.
einen
erstrebt
Einfluß
auf
mehr Freude,
Leichtigkeit,
mehr Individualität verheißt, und hält oft der Moral eine ästhetische
Lebensanschauung
als
allein
So hat zur Zeit
entgegen.
berechtigt
einmal wieder die Entzweiung der beiden Gebiete die Oberhand.
Die geschichtliche Betrachtung zeigte, daß der Gegensatz durch
die Jahrtausende
geht,
nicht
eine
Stimmung des bloßen Tages be-
Immer wieder rückte die Moral der Kunst eine Verweich-
deutet.
lichung, Ermattung, Auflösung des Lebens vor, aber zugleich mußte
sie
dafür den Vorwurf der Härte, Schabion enhaftigkeit, Seelenlosig-
keit
entgegennehmen.
Aber auch davon überzeugten wir uns, daß,
Höhe sich
suchte; bei den schaffenden Geistern war der Gegensatz, wenn auch
was
der Breite des Lebens einander
in
aufgehoben, so
völlig
nicht
deutlich,
daß das
geistige
floh,
auf seiner
doch sehr gemildert;
daß die Schuld des Zwistes weniger an der Sache
In
liegt.
ihnen ward
in
Leben keiner Seite entbehren kann, und
und Kunst
Wahrheit können Moral
als
die
am Menschen
eigne Aufgabe
großem Sinne nehmen, ohne sich gegenseitig als bedeutend,
ja unentbehrlich anzuerkennen, sie können jenes nicht, ohne sich in
nicht in
umfassendes Ganzes des Geisteslebens hineinzustellen und dort
ein
eine Verständigung anzustreben.
Wo die Moral unmittelbar das ganze Leben einnehmen wollte,
da
pflegte
sie
zu einem System von Regeln und Vorschriften zu
zum Menschen sprachen, ihm zugleich aber für
werden, die streng
Erfüllung
ihre
auch
viel
einen
straffe
Lohn
Konzentration
überwiegende Fassung
voller
innerer
entzünden.
Leicht
zu
hohen
als
ist
verhießen.
damit
eines Gebotes
Viel
geleistet
ließ
Aufrüttelung,
worden, aber die
hier die Moral nicht
Aneignung kommen, sowie Liebe und
Freude
ward hier der Mensch zwischen einem Bewußt-
Schwäche
sein
hülfloser
hin-
und hergeworfen.
und einem
In der Tat
selbstbewußten
Pharisäismus
war es immer nur eine gewisse
Mittelhöhe des bürgerlichen oder kirchlichen Lebens, der die bloße
Moral genügte; weder der Durchschnitt der ersten christlichen Jahrhunderte noch der der Aufklärung hatte bei allem moralischen Eifer
einen
bedeutenden geistigen Gehalt.
Gefahr eines
sie
in
weitere
Die Moral
selbst
konnte die
und Äußerlichwerdens erst überwinden, wenn
Zusammenhänge hineingestellt wurde. Geschah dies
Starr-
Probleme der Moral.
aber und wurde
keit,
eine
Weg zur Aneignung einer neuen Wirklich-
sie ein
von Geboten, son-
ihr nicht eine korrekte Erfüllung
galt es in
dern
335
innere
ursprünglichem
Erneuerung des
Leben,
Menschen,
konnte
so
Vordringen
ein
zu
keiner Weise
der Kunst in
sie
Denn jenes Neue ließ sich nur mit Hülfe einer künstlerals Ganzes fassen und in lebendige Gegenwart
entbehren.
ischen
Tätigkeit
stellen,
auch konnte es die Breite des Lebens nicht gewinnen ohne
das bildende, Inneres und Äußeres zusammenflechtende Wirken der
Kunst.
wenn
Ja
daran
alles
neue Welt und ein neues
eine
liegt,
Leben gegenüber den kleinen Zwecken des bloßen Menschen und
dem Alltagsleben zu erreichen, so muß die Kunst mit ihrem stillen
und
Wirken aus der inneren Notwendigkeit der Dinge,
sicheren
mit ihrer inneren Befreiung der Seele, mit ihrem Vermögen, uns die
ganze
Unendlichkeit
zu
nahe
innerlich
und zu eignem
bringen
Leben zu machen, unmittelbar als moralisch
gelten.
Ebenso wenig kann eine Kunst, die groß von
ihrfer
Schwerlich war
Aufgabe denkt, die Moral verachten.
und
sich selbst
ein
je
schaffender Künstler ersten Ranges einer ästhetischen Lebensanschau-
ung
und zwar deshalb nicht,
zugetan,
ein Sondergebiet vom übrigen
Schaffen seine ganze Seele,
und weil
legen mußte,
Schaffens
In
um
Dilettanten;
Genuß
vom Ganzen
den reflektierenden und
als bei
haben
diese
die,
sie
genug
oft
den
auch
theoretischen Erörterungen wenig hold,
kaum empfinden, daß jene Ablösung
ihnen gegenüber wehrlos,
der Kunst
daraus einen bloßen
Wahrheit hat die ästhetische Lebensanschauung ihre
Künstlern aufgedrängt,
ja
das
nicht nur eine gewisse Technik hinein-
Heimat weniger bei den Künstlern
genießenden
in
er die Mühen, ja die Unzulänglichkeit dieses
zu stark empfand,
viel
zu saugen.
weil er nicht die Kunst als
Leben ablösen konnte, weil er
des Lebens
sie
nicht
sowohl erhöht
als
erniedrigt.
Das Angewiesensein von
namentlich
Kunst und Moral aufeinander wird
da zu voller Anerkennung gelangen,
wo
unsere Welt
nicht als eine fertig abgeschlossene, sondern als eine erst im Werden
befindliche,
ja
als
eine
solche
gilt,
in
der nicht nur Vorhandenes
auszubauen, sondern eine neue Stufe der Wirklichkeit zu erreichen
ist.
Dazu bedarf
rüttelung des
es
eines selbständigen
Entscheidens,
einer
Auf-
Wesens, eines energischen Tuns, welches das ganze
Dasein umfaßt; deutlich
ist
hier,
daß wir nicht an
erster Stelle
zu
behaglichem Genießen, zur Kontemplation, sondern zu Wirken und
Zu den Problemen des Menschenlebens.
336
Schaffen, zur Aktivität berufen sind.
und glücklichen
kräftigen
Aber zugleich bedarf es eines
künstlerischen
wenn
Bildens,
die
neue
Welt uns nicht in vagem Umriß verbleiben, und wenn sie das Ganze
unserer Seele gewinnen
soll;
auch die Kunst
ist
eine unentbehrliche
zum Aufbau eines neuen Lebens.
Helferin
Die Probleme der Gegenwart,
ß.
Der moderne Ästhetizismus.
aa.
Wie
wir
prinzipiell
über
den
denken,
Ästhetizismus
bedarf
Den Ästheti-
nach diesen Darlegungen keiner weiteren Erörterung.
zismus der Gegenwart aber klagen wir einer inneren Unwahrhaftig-
und das Leben viel zu viel
Dunkel und viel zu viel Unvernunft, und es regen uns die großen
Uns
keit an.
zeigen
heute die Welt
als daß wir aus tiefster
und mit voller Hingebung unser Dasein überwiegend in Genuß verwandeln und eine Harmonie des Ganzen mit reiner Freude
Widersprüche des Daseins viel zu sehr auf,
Seele
Der Ästhetizismus
erleben könnten.
druck des heutigen Lebensgefühls
als
weniger ein wahrer Aus-
ist
ein Versuch,
seiner
Schwere
und seinem Ernst zu entfliehen. Das aber kann er nur, indem er
sich mit dem modernen Subjektivismus verbindet und in solcher
Verbindung Stimmungen erzeugt, die gewiß
beachtenswert sind,
alle
als
Zeichen der Zeit
denen aber alles schöpferisches Vermögen und
seelenerhöhende Kraft gebricht.
und Ästhetizismus
Ein Zusammenwirken von Individualismus
ließ das Schlagwort einer «neuen Ethik" entstehen,
in weiblichen Kreisen viel
Einfluß
gewonnen hat.
das namentlich
Auch
eine der-
Bewegung sei nicht von vornherein verschrieen, sondern in
Gründen unbefangen gewürdigt. Was die Gesellschaft Moral
nennt, ist nicht mehr als eine gewisse Ordnung des Zusammenlebens,
der Sitte und Gewöhnung einen Heiligenschein verliehen haben, die
artige
ihren
daher bei
auftritt,
der
aller
Unzulänglichkeit leicht mit starker Selbstbewußtheit
wie Diener
leicht arroganter als
Lauf
geschichtliche
der
Herren
sind.
Nun ändert
Dinge die Art des Zusammenseins,
Änderungen können notwendig werden, die starre Festhaltung der
überkommenen Art kann einen peinlichen Druck erzeugen, kann Recht
in Unrecht,
Unrecht
in
Recht verwandeln.
den gegenseitigen Verhältnissen und
verschoben,
daß
eine Revision
jener
in
Die Neuzeit aber hat in
der Art der Arbeit so viel
gesellschaftlichen
Ordnungen
Probleme der Moral.
337
und damit der konventionellen Moral in verschiedener Richtung notwendig ist.
Aber dieses anerkennen heißt nicht der hastigen und summarischen Art
beipflichten,
mit der von den Vertretern und vielleicht
noch mehr den Vertreterinnen eines ästhetischen Subjektivismus die
schwierigen und verantwortungsvollen Fragen erledigt werden.
nächst
Moral
ist
Zu-
etwas anderes als ihre sichtbare Vertreterin,
selbst
die gesellschaftliche Ordnung, und moralisches Verhalten etwas anderes
Wie gering wurde auf der Höhe des moralischen
als soziale Korrektheit
Schaffens von jener Korrektheit gedacht, wie entschieden wurde hier
abgelehnt, das bloße Mittel
zum beherrschenden Zweck zu machen!
Aber auch
bei
das
ist
nicht
alle
Ordnung wie
gesellschaftliche
Druck verrufen werden.
ist
keineswegs
Unzulänglichkeit
aller
Weil gewisse Einrichtungen problematisch geworden
wertlos.
sollte
Mittel
sind,
ungebühriicher
ein
Denn wie die menschlichen Dinge stehen,
unentbehrliches Mittel, das Leben auf ein gewisses Niveau
sie ein
zu heben und den zerstörenden Kräften,
am Werke
die unablässig
genügenden Widerstand zu leisten. Nur ein grenzenloser, man
möchte sagen kindlichnaiver Optimismus, den man liebenswürdig
sind,
nennen
möchte,
wenn
mit seiner die Halbgebildeten be-
er nicht
man dem
stechenden Flachheit gefähriich wäre, kann wähnen, daß
Menschen nur schrankenlose Freiheit zu gewähren brauche, um das
ganze Leben zu seliger Harmonie zu führen. Daß der Mensch zur
Zügelung seiner
Begierden
ist
wer über
müßte auch
sie
klagt,
angenehm schmeckt;
Ordnungen
sozialer
klagenswert sein, es
mag
bedarf,
be-
aber doch nicht die Schuld der Ordnungen;
Medizin verwerfen, die nicht
alle
würden wir
nicht
leicht
das
Gegenteil
der
wenn wir einem verschönernden Bilde zuliebe
alle Schranken einreißen wollten?
L'homme n'est ni ange ni bete;
Absicht
et
erreichen,
malheur veut que qui veut
le
Wenn von
Tange
faire
fait
bete (Pascal).
la
»neuer Ethik" geredet wird, so müssen wir auch
einem solchen Mißbrauch des Wortes Ethik widersprechen.
gleichgültig
sind
die
Wörter
nicht,
ihr
Mißbrauch
Ganz
kann zu einer
Verdunklung echter Probleme wirken.
Unter Moral hatten wir uns
gewöhnt an
willkürentzogenen
Ordnung,
und Gewissen zu denken.
Was aber
die
die
Anerkennung
Hochhaltung von
Pflicht
einer
der ästhetische Subjektivismus mit seiner „neuen Ethik"
nichts
anderes als ein
feinerer
Epikureismus,
des Individuums, das sich von aller
Eucken, Orundbegriffe. 4. Aufl.
als
ein
bietet,
ist
Selbstgenuß
Hemmung befreit; wer darin
22
Zu den Problemen des Menschenlebens.
338
sein
Genüge findet, der sollte die Ethik genau so gut wie die Reli-
gion
Bausch und Bogen für eine Verirrung erklären und aus
in
seiner Gedankenwelt streichen, nicht aber sollte er durch ihren Namen
Denkweise einen schimmernden Aufputz geben.
seiner andersgearteten
Ein schroffer Gegensatz
ist
hier unverkennbar.
Ist
der Mensch nicht
mehr als die Summe seiner natürlichen Neigungen und kommt alle
Lebensweisheit
darauf
hinaus,
unser Dasein
Herr zu werden?
Ist
rezeptiver oder
es aktiver Art?
Höchste
ist
in
ein
leidliches
uns eine geistige
Kraft,
unser Verhältnis zur Wirklichkeit vorwiegend
oder
Güter,
aller
in
Tat zu verwandeln und zugleich unser selber
freie
in
Neigungen
diese
oder steckt
Gleichgewicht zu bringen,
über solchen Abschluß
hinaus?
unser subjektives Glück das
Ist
uns
treibt
innere
eine
Dieser Gegensatz
Notwendigkeit
steht
den
seit
Stoikern
und den Epikureern mit voller Klarheit vor Augen, und er
schließt
jede
Die
Vermittlung aus.
modernen,
präziser als die
alten
Epikureer aber dachten
haben sich nicht als Vertreter einer
sie
neuen Ethik gegeben.^
Der Punkt, an dem der moderne Subjektivismus mit anderen
Überzeugungen am
zusammenstößt,
härtesten
ist
die
Behandlung
des Sinnlichen, besonders in der geschlechtlichen Sphäre.
Verwicklungen
namentlich
Christentum,
vielfach
keit
vorliegen,
niemand
wird
in
seinen
noch eine Geringachtung,
fort,
die
in
katholischen
ja eine
Im
Fassungen,
wirkt
Verachtung der Sinnlich-
den Stimmungen des ausgehenden Altertums und
seinem Kampf gegen eine entartete Sinnlichkeit wurzelt;
hier ein Stück
Manichäismus
in das
*
ist
es,
es
ja
ist
Christentum eingedrungen und
wirkt bei aller äußeren Strenge zu innerer Verflachung.
Verflachung
Daß hier
können.
bestreiten
Denn eine
wenn die Hauptsorge des Lebens darin gesetzt
Mit besonderer Kraft und Tiefe hat sich über diese Probleme der aus-
gezeichnete schwedische Philosoph
Vitalis
Norström
in
seinem Buch
tausendjährige Reich" (deutsch 1907) ausgesprochen; er sagt z. B. S. 31
:
„Das
„Selt-
same Schwärmerei, die auf der Ersättigung der Sinne ein stabiles seelisches
Gleichgewicht zu begründen meint! Armselige Weisheit, die kein höheres
Ziel kennt, als sich dauernd,
wenn man so sagen kann, einen guten Tag zu
—
wofern sie überhaupt
machen! Diese Welt des allgemeinen Genießens
würde das Beste nicht aufkommen lassen, dessen der
möglich wäre
Mensch fähig ist: die Erhebung über den Genuß, die Selbstüberwindung.
Sie würde das ausschalten, was unsrem gewiß verbesserungsbedürftigen
Dasein doch eine gewisse Weihe gibt; das, was ein Landsmaim Zolas, der
—
edle Alfred de Vigny, gefeiert hat als die „majeste dessouffrances humaines.«
Probleme der Moral.
Kampf
einen
wird,
dem
mit
Sinnlichen
339
zu führen,
das Sinnliche
wenn wer es
möglichst abzuschwächen, zu erniedrigen, zu kasteien,
in
solchem Ertöten der Sinnlichkeit weit gebracht
ein
noch so
als
Vorbild erwählt wird.
handlung
harter
Mensch,
als
Held
ein
Denn was wird denn
und
sei
gefeiert
er
und
mit solcher Miß-
und
Dazu muß jene Unter-
der Sinnlichkeit für eine innere Läuterung der Seele
Aufbau
für den
und
leerer
hat,
geistigen
Lebens gewonnen?
drückung der Sinnlichkeit, wie alles Naturwidrige, schlimmere Schäden
herbeiführen, als sie zu heilen unternimmt; die Natur pflegt sich für
Aber mit der Abweisung einer
eine Mißhandlung schwer zu rächen.
derartigen Askese
die Sache nicht schon erledigt,
ist
und
sinnliche
geschlechtliche
Aber zugleich
fest.
Seite
zeigt
den Menschen der Natur
mehr als irgendwie sonst hält hier die Natur
aufs engste verwachsen,
ihn
sie liegt nicht
dem ästhetischen Subjektivismus scheint. Die
so einfach, wie es oft
hat er sich mit der Entwicklung geistigen
Lebens weit über sie erhoben und damit die schlichte Naivität auch
auf jenem Gebiete verloren; so wird ihm das Sinnliche zum Problem,
vom
das
es
sich
in
ein
voller
Freiheit,
verschieden
beantworten
läßt;
den Zwecken des Geisteslebens unterzuordnen und
Maß
zu
Wer
finden?
soll
unbekümmert um höhere Zwecke des
und Laune des Einzelnen ausleben, oder
nach Lust
Geistes,
es sich
Geistesleben aus sich
unter
hat
in ihnen
Berufung auf das unantastbare
Recht der Natur sich für jenes entscheidet, der pflegt zu übersehen,
daß wir bei unserer gesteigerten und
vielfach
überspannten KuUur
nicht mit der reinen Natur zu tun haben,
daß die heutige Sinnlichkeit
vielfach eine raffinierte, ja entartete ist
So können wir schon zur
Scheidung dessen, was an der Natur echt oder unecht
ist,
geistiger
Arbeit nicht entbehren, eine einfache Kapitulation vor dem, was sich
heute sinnlich nennt,
ist
schlechterdings ausgeschlossen.
bb. Die Stellung der Kunst im modernen Leben.
Siegreich dringt im
modernen Leben die Kunst wieder vor und
mächtig bewegt sie die Geister; so
ist
es kein Wunder,
daß
sie alle
Abhängigkeit verschmäht und auf voller Selbständigkeit besteht.
solches Verlangen
l'art
pour l'art.*
^
S. 326)
findet
In dem
Der Ausdruck
hatte,
(s.
wie ihn
einen Ausdruck
in der
Ein
bekannten Formel
was sie an Verneinung enthält, wird ihr kein
darüber Büchmanns Geflügelte Worte, 21. Aufl.,
zuerst Victor Cousin 1818
Vorlesungen an
in
Sorbonne gebrauchte, einen ganz unverfänglichen Sinn
:
II
faut
der
de la religion
22»
Zu den Problemen des Menschenlebens.
340
Freund der Kunst widersprechen. Die Kunst soll nicht fremden Zwecken
dienen, sie darf nicht der Moral,
reichung
und damit zu
leisten
der Politik, der Religion Hand-
einer Tendenzkunst sinken,
die nur
für einen Augenblick blenden mag, nie aber wahre Förderung bringt
Schwieriger ist es, jenem Wort einen positiven Sinn zu geben.
geschieht
das
nicht selten
gleichgültig gegen
allen Stoff
und Inhalt
sein
der Vollendung ihrer Form befassen; so allein
und könne
selbst
völlig
Heute
mit der Behauptung, die Kunst müsse
und
sei
ungehindert ihre eignen
sich
sie
allein
mit
ganz bei sich
Wege
verfolgen.
Aber liegt eine solche Ablösung vom übrigen Leben in ihrem eignen
Interesse,
ist?
kann
bei
sie
Die Gefahr
liegt
ihr
das Höchste leisten,
dessen
sie
fähig
nahe, daß mit dieser Wendung die Kunst zur
bloßen Formbeherrschung, zu virtuosenhafter, glänzender, blendender
Technik wird, daß
sie
so weder den ganzen Menschen hinter sich
hat, noch zum ganzen Menschen und zur Menschheit zu wirken
vermag.
Eine solche Kunst mag weit mehr in der sinnlichen Erfahrung entdecken, sie mag unser Empfinden in ungeahnter Weise
bereichern und verfeinern, sie mag in Überwindung von Widerständen
schwelgen, aber seelisch kann sie uns wenig bieten, und das Geistesleben wird
kaum erhöhen.
sie
War nicht dies den großen Kunst-
werken, die dauernd zur Menschheit sprechen,
aller
Gegensatz von Inhalt und Form
haben
sie
nicht
in
in
der Vollendung der
eigentümlich,
Form
zugleich zu vollem
Ausdruck gebracht, was das Innere des Lebens erfüllte?
die
daß
ihnen überwunden wurde,
Muß nicht
Kunst die Probleme der Menschheit aufnehmen und
in
ihrer
Weise zu lösen suchen ?i Auf jene Innerlichkeit kann am wenigsten
pour la religion, de la morale pour la morale, de l'art pour l'art. Erheblich
wurden die letzten Worte zum Bekenntnis einer Schule und zu
einem Zankapfel der Parteien. — Hinzufügen möchten wir nur, daß auch
Comte sich einmal mit jenem Schlag\i'ort befaßt, aber es sehr äußerlich
wendet. Cultiver l'art pour l'art lui-meme bedeutet ihm nichts anderes als
ne se proposer habituellement d'autre bout reel que de divertir le public
(cours de phil. pos. VI, 167).
^ Auch
über dies Problem hat sich Norström in vorzüglicher Weise
später erst
So bemerkt er z. B. über das oft als Beispiel eines bloßen SchönGriechentum (Das tausendjährige Reich, S. 73): »Man
bildet sich oft ein, die Grundkraft des griechischen Lebens sei ein unwiderstehlicher, zwangloser Trieb zur schönen Form gewesen, also ein Bedürfnis,
das an sich schon schöne Dasein noch mehr zu verschönern. Statt dessen
geäußert.
heitskultus angeführte
aber dienten
sittliche
bei
Kräfte
zu
ihnen
die Gebilde der Kunst wesentlich dazu,
befreien,
dunkle
Bewußtseinsinhalte
gefesselte
aufzuhellen
und
Probleme der Moral.
Er
der nordische Mensch verzichten.
341
besitzt
nicht die
natürliche
Leichtigkeit der sinnlichen Darstellung wie der Bewohner des Südens,
er
das
der
Innerste
sich
selbst
unausgesprochen
Seele
Daher wird ihm die Kunst
Tiefe verschlossen.
Mittel,
Weg von innen nach außen; so bleibt ihm
schwer den
findet
leicht
zu
finden,
Eigentum
sein
und
ein
in
seine
eigne
unentbehrliches
vollen
Besitz
zu
nehmen, die Kluft im eignen Wesen irgend zu schließen. So kann
ihm auch
die vollendetste
Form
als
bloße
Form nun und nimmer
genügen.
Wer einen Inhalt der Kunst als etwas ihr fremdes und gefähran eine Gedankenleistung, eine
liches verschmäht, der pflegt dabei
Idee
abstrakte
Aber
zu denken.
fällt
denn Geistesleben mit dem
Denken zusammen, gibt es keinen anderen geistigen Inhalt als eine
Gedankengröße? Der alte Intellektualismus mochte das meinen, heute
aber wollen wir. nicht mehr Intellektualisten sein;
also jetzt die veralteten
wie könnten uns
Maße binden und uns hindern, in tieferem
und' weiterem Sinne nach einem Inhalt für den ganzen Menschen zu
streben?
Ja wir sehen sogar in der Zurückstellung des Inhalts eine Ge-
fahr für eben die Selbständigkeit der Kunst,
zu deren Gunsten
sie
verlangt wird. Mit dem Unabhängigwerden vom Stoffe wird nicht eine
Unabhängigkeit schlechthin
wird
ist,
leicht
wenn
nicht anderen,
zu zeigen;
erreicht.
Eine überwiegende Formkunst
zu einem Virtuosentum, das vor allem darauf bedacht
das
aber
so doch sich selbst das eigne
Vermögen
eine Vorliebe für das Abweichende,
erzeugt
Paradoxe, Überspannte, und mit dem Suchen dessen gerät man leicht,
indem man seine Freiheit beteuert, nur
in eine
andere Art der Ab-
hängigkeit, eine Abhängigkeit von anderen und auch von der eignen
Stimmung.
Schaffen
Nur da
besteht
wahrhaftige
Unabhängigkeit,
wo
das
ganz und gar aus einer innem Notwendigkeit des eignen
Wesens hervorgeht; dazu aber muß der Mensch etwas auszusprechen,
zersplitterte
Arbeit
weiter: „Tatsächlich
um gemeinsame praktische Ziele zu sammeln." - Und
gilt
von
aller
großen, echten Kunst, daß
sie die
Form
dem Inhalte, den sie wiederzugeben sucht, mehr oder weniger unterordnet
und gegen die Lust oder Unlust, die sie — bei dem Künstler und anderen
— erweckt, sich gleichgültig verhält, wenn nur die Bilder den Inhalt mitteilen.
eher,
Die echte Kunst öffnet uns die Tiefe der schöpferischen Phantasie
um uns von dem ,heiteren' Dasein zu befreien, als um uns tiefer in
dasselbe hineinzuführen.«
Zu den Problemen des Menschenlebens.
342
ja
Der bloßen Virtuosenkunst aber fehlt eine
zu offenbaren haben.
solche Notwendigkeit.
Mit
Worte
einem
zur
nicht
von
ihr
kann
ihr
nur eine
oft
des Verhältnisses der modernen
auch
sei
Daß
geschlechtlichen Sphäre gedacht.
Kunst
die Kunst sich
zurückzieht, sondern sich eingehend mit ihr befaßt,
aber jenes
unkünstlerische
Denkweise verargen.
Daß
sie
mit sichtlicher Vorliebe in den Vordergrund rückt
und möglichst lange bei ihm
und so raffimehr ein Zeichen sitt-
verweilt, es so reflektiert
niert bis zur Eckelhaftigkeit ausmalt, das
ist
Könnens, das kann durch keine ästhe-
licher Fäulnis als technischen
tische Theorie verteidigt werden.
Wie viel bei der bildenden Kunst im Fluß und im Streit sein
mag, an hervorragenden Persönlichkeiten und an glänzenden
ungen
auf
fehlt es hier wahrlich
dem Gebiete
nicht.
der Literatur.
Minder günstig steht die Sache
Anregungen und Aufgaben
und
und neue
über seine Stellung im All
steigen
auf,
Alte
ist
der Mensch
höchst unsicher geworden, der eigne Kreis der Menschheit
Bewegungen und Wandlungen, dazu
bietet
Qedankenmassen versinken
hier die Zeit in Hülle
Fülle.
Leist-
gestattet
ist
voller
die wachsende Hast
des Lebens keine genügende Selbstbesinnung; so
ist
uns das Dasein
Hätte hier
verworren und wir selbst sind uns vielfach ein Rätsel.
nicht das literarische Schaffen die Aufgabe einer Klärung, hätte es
nicht,
was auf uns eindringt und
Aussprache zu bringen, aus
sich in
uns
regt,
zur deutlichen
dem Chaos der Erscheinungen, das uns
umwogt einfache Grundlinien herauszuheben, uns das Leben mögGanzes zu fassen und zugleich es weiterzuführen? Dazu
lichst in ein
aber bedürfte es einer inneren Überlegenheit über die Gegensätze
der Zeit, einer energischen Synthese, die nicht aufnähme ohne auch
auszuschließen,
Schaffens.
ist
An
eines
mutig
und
kräftig
vordringenden
geistigen
Ansätzen dazu fehlt es nicht, aber im allgemeinen
zu sagen, daß unsere Literatur, die Literatur eines der größten
Kulturvölker, nicht auf der Höhe der Zeit steht, daß sie dem modernen
Menschen
in
dem Kampf um eine geistige Selbsterhaltung und um
einen Sinn seines Lebens wenig Hilfe gewährt.
dies offen auszusprechen.
Es
ist
eine Pflicht,
5. Persönlichkeit
Persönlichkeit.
a)
a.
I
und Charakter.
Zur Geschichte des Ausdrucks.
|en Ausdruck Person, einen der wenigen Termini
lateinischen
*-^
Ursprungs, von seinem Ursprung an in seine schon im Alter-
tum
reiche Verzweigung zu verfolgen, seine Bedeutung im römischen
Recht wie in der christlichen Theologie darzulegen, das würde uns
zu weit von unserer Aufgabe entfernen.^
So halten wir uns an die
Philosophie und streben auch hier möglichst rasch zur Gegenwart.
Die neuere Philosophie empfing den Ausdruck von der Scholastik,
aber folgte der Definition des Boethius,
diese
nünftiges Einzelwesen.2
die Trinitätslehre
Person
sei
ein ver-
Aus der Anwendung dieser Definition auf
entstanden
ernste Verwicklungen (Roscellin),
aber
Über das Technische des Ausdrucks berichtet näher die RealenzykloÜber seine Anfänge und seine Schicksale bis ins Mittelschrieb einen anregenden Aufsatz Max Müller in Good Words, Juni
^
pädie von Pauly.
alter
1866, bedeutender ist eine Untersuchung Trendelenburgs, die ich aus seinem
Nachlaß in den Kant-Studien Bd. XIII, Heft 1 u. 2 veröffentlicht habe.
^ Näher
lautet die Definition (in der Schrift de duabus naturis, ed.
R. Peiper,
pag. 193, 4):
persona
est
rationalis
naturae individua substantia.
Im früheren Mittelalter ward Person wohl etymologisch erklärt als per se
una.
Über die verschiedenen Fassungen des Begriffes bei den wichtigsten
Denkern s. Baumgartner, die Philosophie des Alanus de
Indem Thomas, namentlich in seiner Untersuchung über die
Dreieinigkeit (im 1. Buch der summa theologiae), die Lehre des Boethius
weiter entwickelt, betont er, daß Personen non solum aguntur, sed per se
agunt.
Er verteidigt die Anwendung des Ausdrucks auf Gott, obwohl sie
mittelalterlichen
insulis, S. 45.
Wie andere Scholastiker, so hat auch Thomas
dem schon Eckhardt in personlicheit ein deutsches Gewand gab,
sich nicht in der Bibel finde.
personalitas,
wie er auch personlich öfter verwendet.
gebräuchlichste Definition
In der späteren Scholastik
von Person suppositum
intelligens,
war die
suppositum
aber bedeutete dabei eine substantia singularis viva. Zesen übersetzte persona
mit Selbstand
(s.
Paul Piur, Studien zur sprachlichen
Würdigung Christian
Wulffs, S. 58), Clauberg (Wke. 1691, S. 321) mit „selbständig verständig Ding."
Zu den Problemen des Menschenlebens.
344
haben ihren Gebrauch
sie
in
der Scholastik nicht gehindert.
Philo-
sophische Probleme bedeutender Art werden dabei nicht aufgenommen.
Erst die Neuzeit brachte die Sache in regeren
Fluß.
Der Be-
dem Zuge nach
einer gemeinsamen und gleichförmigen Weltordnung dem Menschen
Person wird hier ein Hauptmittel, gegenüber
griff
eine ausgezeichnete
übernommenen
Stellung zu wahren.
Begriffe
Person
Die von der Scholastik^
und Persönlichkeit werden dabei
psychologisch genauer durchgebildet.
Leibniz führt diese Bewegung,
indem er das Wesen der Persönlichkeit
in
das Selbstbewußtsein,
das Bewußtsein der Identität in den verschiedenen Zeitpunkten
d. h.
eignen Daseins, setzt und daraufhin die Unsterblichkeit des
Menschen von der Unzerstörbarkeit der niederen Wesen abhebt^
des
Wolff wie die Aufklärungsphilosophie nehmen diese Fassung
Herbart führt
sie
bis ins
auf,
19. Jahrhundert hinein.^
Das auszeichnende Merkmal der Persönlichkeit war bis dahin
^
Wie eng in der ersten Hälfte des 18. Jahrhunderts der Zusammenhang
mit der Scholastik noch war, und wie sehr die Ausdrücke als bloße Termini
der Schule galten, zeigt das weitverbreitete philosophische Lexikon von Walch.
Es heißt dort (auch noch in der vierten, von Hennings besorgten Ausgabe
von 1775) unter Person: «Person heißt in der Metaphysik eine besondere,
vollkommene und vernünftige Substanz, welche ihr Wesen und ihre Subsistenz
vor sich hat. Das Abstraktum davon oder die Subsistenz eines solchen Wesens
ist
Personalitas genannt worden."
^
Theodic^e I, § 89: l'immortalite, par laquelle on entend dans l'homme,
non seulement que l'äme, mais encore que la personalitesubsiste: c'est-ä-dire,
en disant que l'äme de l'homme est Immortelle, on fait subsister ce qui fait
que
c'est
vant
la
la
meme personne, laquelle garde ses qualites morales, en conser-
conscience ou
le
sentiment reflexif interne de ce qu'elle est: ce qui
Ja rend capable de chätiment et de recompense. Femer in dem Briefe an
Wagner (de vi activa corporis et de anima brutorum) S. 466 b der Erdraannschen
Ausgabe: Itaque non tantum vitam et animam, ut bruta, sed et conscientiam
sui et memoriam pristini status, et, ut verbo dicam, personam servat.
' So sagt Wolff (psych, rationalis,
§741): Persona dicitur ens, quod
memoriam sui conservat, hoc est, meminit, se esse idem illud ens quod ante
in hoc vel isto fuit statu.
Femer in den »vemünftigen Gedanken von Gott,
der Welt und der Seele des Menschen" § 924 (angeführt von Trendelenburg):
»Da man nun eine Person nennet ein Ding, das sich bewußt ist, es sei eben
dasjenige,
was vorher
in
diesem oder jenem Zustande gewesen: "?o sind die
Tiere auch keine Personen.
daß
sie
Hingegen weil die Menschen sich bewußt sind,
die vorher in diesem oder jenem Zustande
eben diejenigen sind,
gewesen: so sind sie Personen."
Selbstbewußtsein,
Herbart aber sagt: (Wke. III, 60): »Persön-
worin das Ich
lichkeit
ist
faltigen
Zuständen als Eins und Dasselbe betrachtet."
sich
in
allen
seinen mannig-
Persönlichkeit und Charakter.
die
Nun
Intelligenz.
345
aber beginnt eine neue, eine ethische Phase.
Nach mannigfacher Vorbereitung hat Kant mit seiner Voranstellung
der praktischen Vernunft
BPersönHchkeit«
durchgesetzt
sie
ist
einer
der Hauptpunkte, welche die neue Denkweise zu greifbarem Ausdruck
bringen,
aus,
in
wächst bei
es
Kant weit über die bloße Intelligenz hin-
ihr erscheint eine wesentlich höhere,
Freiheit gegründete
in
Persönlichkeit bedeutet nämlich »die Freiheit
Ordnung.
und Unab-
hängigkeit von
dem Mechanismus der ganzen Natur", das »was den
Menschen über
sich
selbst
was ihn an eine Ordnung
einen Teil
(als
der Sinnenwelt) erhebt,
der Dinge knüpft, die nur der Verstand
denken kann, und die zugleich die ganze Sinnenwelt, mit ihr das
empirisch -bestimmbare Dasein des Menschen in der Zeit und das
Ganze
Zwecke, unter sich hat« (V, 91 Hart).
aller
Als Personen
und dürfen nie als bloße Mittel
gebraucht werden. Ja es wird in uns neben der Tierheit und
Menschheit noch die Persönlichkeit unterschieden; der Mensch ist
zunächst ein lebendes, dann ein lebendes und zugleich vernünftiges,
sind Vemunftswesen Zwecke an sich
als Persönlichkeit endlich
ein vernünftiges
nung fähiges Wesen
120).
Denker,
Spätere
(VI,
wie der
ältere
und zugleich der Zurechauch
Schelling,
H. Fichte,
J.
haben diese ethische Fassung des Persönlichkeitsbegriffes nach der
metaphysischen Seite zu ergänzen und vertiefen gesucht,^ im großen
und ganzen aber ist man
seit
ihm
fest,
daß das
bei
allen
Kant verblieben.
einzelnen
Subjekt
Zum mindesten steht
Handlungen überlegene,
auch
mit
praktischer
als
Vernunft
Persönlichkeit
bezeichnete
auszustatten
daß zu seinem Wesen nicht bloß das Selbstbewußt-
sein,
ist,
sondern auch die Selbstbestimmung gehört
ß.
Zur Geschichte des Begriffs.
Die letzterrungene Fassung des Begriffs als des selbstbewußten
und
selbsttätigen Subjekts
^
ist
es,
deren Geschichte wir
nunmehr in
Dne Geschichte des Persönlichkeitsbegriffes im 19. Jahrhundert wäre
manchen Ertrag verspricht. Über den Gebrauch des Ausden verschiedenen Völkern bemerkt Alexander Chamberlain in
eine Aufgabe, die
drucks bei
Harpers Monthly (1003, Juli, pag. 281): The word personality is not a natrve
English
term,
but has been borrowed, ultiraately frora mediacN-al Latin and
from the lau^-ers. The corresponding French term,
was admitted to the Academy's dictionary so recendy as 1762.
subsequently rescued
personnalite,
The German Persönlichkeit was once entirely in the possession of the mystics.
Zu den Problemen des Menschenlebens.
346
Kürze aufrollen möchten.
Die griechische Philosophie gelangte nicht
zu einem deutlichen Begriff der Persönlichkeit, sowohl weil die Frage
der Einheit des Seelenlebens nicht im Vordergrunde stand,
das Denken
dem vorwaltenden Intellektualismus
das wahre Selbst des Menschen
als
Immerhin
galt.
großen Durchforschern des Menschenwesens,
als
weil
der Kern
und
sich
den
arbeitet
im Widerspruch
fast
mit ihren Hauptlehren, ein gewisser Begriff der Persönlichkeit her-
So bei Plato, so mehr noch
genug über einzelne Handdas sich selbst im Handeln er-
aus und wirkt in ihrer Gedankenwelt.
bei
dessen
Aristoteles,
Ethik
deutlich
lungen zu einem Wesen vordringt,
Das spätere Altertum
lebt.
hat
mehr und mehr den Menschen auf
und auch den
eignes Innere gestellt
sein
Begriff des Selbstbewußt-
seins entwickelt,^ aber zu einem vollen Begriff der Persönlichkeit ist
es
nicht
Eine
gelangt.
unserem
der
Begriff
Persönlichkeit
ent-
sprechende Fassung der Gottheit wurde von hervorragenden Denkern
nachdrücklich abgelehnt.^
1
S. Siebeck,
Geschichte der Psychologie
ausbildung des Bewußtseinsbegriffes."
I,
2,
S.
331—342: „Die Her-
Trendelenburg hat in dem angeführten
Aufsatz näher dargelegt, wie sehr die Stoiker die Entwicklung dieses Begriffs
gefördert haben; er zeigt, wie »wir bei ihnen, die im Leben auf die Über-
einstimmung mit
sich
selbst,
Charakters gerichtet waren, das
auf
die
Konsequenz des mit
r.po'jujxo^j ,
die
persona
sich
einigen
zum Ausdruck
des
Ethischen werden sehen"; und weiter: »die gut gedichtete Rolle ist der Natur
gemäß, wie der
leben,
die
d. h.
gut
erste
Grundsatz der Stoiker verlangt, der Natur gemäß zu
der Vernunft, die der Natur
gedichtete Rolle
individualisiert
zum Grunde liegt, zu folgen, und
ferner das
Allgemeine der eigen-
tümlichen Natur des Einzelnen gemäß und gründet es in einem vernünftigen
Mittelpunkt".
*
Zuerst
tat das,
so weit bekannt, der Akademiker Karneades (213/4 bis
129 V. Chr.), später mit größter Kraft und Schärfe Plotin.
großem Werk
S.
darüber neben
Grundriß der Gesch. d. griech. Philosophie.
Karneades suchte zu zeigen (s. Grundr. 6. Aufl., S. 242 ff.): „daß man sich
die Gottheit nicht als ein lebendes, vertiünftiges Wesen (Cwov Xoyizo'v) denken
Zellers
seinen
könne, ohne ihr Eigenschaften und Zustände beizulegen, die ihrer Ewigkeit
und Vollkommenheit widerstreiten." Besonders energisch aber kämpft Plotin
dem Ganzen einer Weltanschauung dagegen, dem schlechthin Unendlichen und aller Besonderheit überlegenen Urwesen, wie er es faßt, Denken
oder Wollen und weiter auch Selbstbewußtsein zuzuschreiben (s. Zeller a. a. O.,
S. 293 ff.).
„So tritt die von Karneades vorbereitete Leugnung der Persön-
aus
lichkeit Gottes
hier zuerst mit grundsätzlicher Entschiedenheit auf« (Zeller).
Gründe sind für die spekulative Ablehnung einer Persönlichkeit
Gottes maßgebend geblieben, auch Spinoza hat ihnen kaum etwas neues
Plotins
hinzugefügt.
Persönlichkeit und Charakter.
Die
Belebung und größere seelische Nähe, welche
kräftigere
die Gottesidee
im
347
Christentum gewann,
alten
gestattet hier weit eher,
von einer Persönlichkeit Gottes und einem persönlichen Verhältnis
des
Menschen zu Gott zu
eines
Anthropomorphismus
nicht unbemerkt:
das zeigt schon
dem höchsten Wesen ein Affekt
der heftige Streit darüber, ob sich
wie der Zorn zuschreiben
Die darin liegende Gefahr
sprechen.
blieb
Eine Lösung fanden die Probleme
lasse.
im Gottesbegriff unter Augustins Führung schließlich
in der Weise,
daß eine menschlich -persönliche Fassung auf die Grundlage einer
Persönlichkeit
Mittelalters
absolutes
empfand
Widerspruch.
ein
und
Gott
aufgetragen wurde.
spekulativ-mystischen
Sein.
Die
zugleich sittliche
ist
mattere
Denkweise
des
dem Nebeneinander beider Fassungen keinen
in
Der Neuzeit verwandelte
Entweder -oder, was das
Mittelalter
sich
auch hier alsbald
als
ein
in
Sowohl — als auch
ruhig, ja willig ertragen hatte.
So
erfolgt
denn
in
der Neuzeit eine energische Scheidung der
Geister beim Gottesbegriff.
folgt
Was immer dem Zuge zur Immanenz
und auf universalen Weltbegriffen
besteht,
das bekämpft die
persönliche Fassung der Gottesidee als einen unerträglichen Anthro-
pomorphismus.
Der Widerstand und Widerspruch gegen den Pan-
theismus dagegen stützt sich in seinem Verlangen nach einem lebendigen göttlichen Sein auf die Idee der Persönlichkeit und gibt auch
dem Wort einen besonderen Nachdruck. Bis dahin hatte man viel
von dem Verhältnis der drei Personen im göttlichen Wesen, aber
wenig von der Persönlichkeit Gottes gesprochen.^ Nun aber wird
Persönlichkeit ein Bekenntnis
und Lieblingswort der Antipantheisten.
So beteuert z.
dem
B. Jakobi
in
bekannten Gespräch mit
Lessing
seinen Glauben an eine «verständige persönliche Ursache der Welt"
und vermißt an der Substanz Spinozas „eine eigne, besondere, indiVon da aus
viduelle Wirklichkeit", „Persönlichkeit und Leben".
zieht sich der Streit
durch das
19.
Jahrhundert bis zur Gegenwart.
Wo immer der Lebensprozeß sich vorwiegend künstlerisch oder intellektuell
und zu
dagegen
gestaltet,
klein,
da dünkt leicht die Idee der Persönlichkeit zu eng
um das Ganze der Wirklichkeit zu beherrschen; wo
der ethische
Zug die Führung
hat,
da möchte
man den
* Wiederum sei hier Walch als Zeuge angeführt, der im Artikel Person
wohl von Personen der Dreieinigkeit, nicht aber von einer Persönlichkeit
Gottes spricht und in einer ausführlichen Erörterung des Wesens Gottes
»Persönlichkeit" mit keinem Worte berührt.
Zu den Problemen des Menschenlebens.
348
missen und strebt nach einer Fassung, die auch die
Begriff nicht
Gottesidee
umspannen vermag
zu
Lotze und
(so
Ritschi).
— Die
neueste Zeit Spricht besonders viel von Persönlichkeit, die Wendung
zu
dieser
erscheint
oft
als
ein
Heilmittel
sicheres
aller
Schäden.
Eine kräftigere Belebung der Persönlichkeit will die Kunst, die Redas allgemeine Leben; sie dünkt eine unentbehr-
ligion, die Moral,
liche Hilfe
gegen die drohende Entseelung des Daseins, ein Mittel
zur Abstoßung von Veraltetem und
Überlebtem, der einzige
Weg
zur ersehnten Verjüngung und Vereinfachung des Lebens; mit der
Persönlichkeit hofft man gegenüber der Erschütterung der Weltbegriffe
einen
Halt
sicheren
im
eignen
Innern
gewinnen,
zu
auch einen
Punkt, auf den sich die Menschen inmitten unerträglicher Spaltung
und Zersplitterung einigen können.
Wo so viel zusammentrifft, läßt sich viel Verworrenheit erwenn eine so leichte Wend-
warten; es müßte wunderbar zugehen,
ung, eine einfache Selbstbesinnung, uns den unermeßlichen Verwick-
lungen entwinden könnte, die uns heute umfangen.
Vermutlich
ist
entweder die Hilfe ein bloßer Schein, oder der Gedanke der Persönlichkeit enthält
beilegt.
mehr und fordert mehr, als die übliche Fassung ihm
Sehen
wie es damit
wir,
y.
steht.
Untersuchung des Problems.
An dem Streit über Persönlichkeit ist ohne Zweifel viel Wortstreit;
so lange der eine
dem Ausdruck eine engere und niedrigere,
der andere eine weitere und höhere Bedeutung
ständigung ausgeschlossen.
um
das
Wort nur
Nicht deshalb
Aber wie oft, so
die Erscheinung
schätzen
ist
gibt,
ist
eine Ver-
auch hier der Streit
eines sachlichen Gegensatzes.
hervorragende Denker bis zur Gegenwart
»Persönlichkeit", weil ihnen das Wort in die Ohren klingt, sondern
weil
sie
zeichnet,
in
wenn auch unvollkommen,
ihm,
eine Tatsache
verzichten zu dürfen.
den Vorrang
brachte,
so
sie
Gedanken
be-
nicht glauben
Wie Person und Persönlichkeit von alters her
Menschen, des geistigen Wesens, zum Ausdruck
desist
einen
behauptet finden, auf die
es
eine
Grundüberzeugung vom Geistesleben, von
seinem Inhalt und seiner Bedeutung, die
in
ihm sich ein, wenn auch
unzulängliches Werkzeug schuf.
Wer im Zusammenhang einer Welt-
anschauung für
eintritt,
Persönlichkeit
Geistesleben keinen bloßen
Anhang
behauptet
damit,
daß das
der Natur, sondern eine eigen-
tümliche Art des Seins besagt; er behauptet, daß es nicht in einzelne
Persönlichkeit und Charakter.
349
Betätigungen und Vermögen aufgeht, sondern eine ihnen überlegene
und
umfassende Einheit enthält und damit zu einem Beisich-
sie
einem Selbstleben wird;
selbstsein,
Selbstleben
bloßer
kein
sondern aktiver Art
ist,
Sammelpunkt ihm
umwandelnde
eine
daß dies
behauptet endlich,
er
zugeführter
Elemente,
Kraft an allem
Empfan-
Nur
genen übt und das ganze Dasein auf eine höhere Stufe
hebt.
wenn
dieses
wesentlich
Neues
in
sie
von
alles
zutrifft,
bringt
Persönlichkeit
etwas
unser Dasein und rechtfertigt damit den Affekt, mit
dem
vielen ergriffen wird.
Nun
aber
das
ist
Ganze
Behauptungen nur im Recht,
jener
wenn im Gesamtbilde der Wirklichkeit und unserer Stellung in ihr
sich eingreifende Wandlungen vollziehen. An der besonderen Stelle
kann keine Wahrheit haben,
was nicht im Ganzen wahr und
Zusammenhängen begründet
seinen
in
wäre jene Bewegung zur
ist;
Persönlichkeit eine bloße Privatangelegenheit des Menschen, so wäre
sie
mit ihrem Weltbilde eine bloße Illusion, so würde
sie
ins Leere
Ein Durchdringen zur Wahrheit wird sie nur, wenn das Geistes-
fallen.
leben die Tiefe der Wirklichkeit bildet, in der sie ihr eignes Wesen
Nur auf Grund einer neuen Weltstufe und im Zusammen-
erreicht.
Wendung zur Persönlichkeit
hang mit
ihr
vollziehen
und die Menschheit persönliches Leben
kann der Einzelne die
entfalten.
Ja
es
muß jenes neue Leben, um den Menschen über die naturhafte Ordnung hinauszuheben,
sein
und
in
ihm wirken;
er
als
muß an
um der äußeren gewachsen
So handelt es sich bei Persönlichkeit um ein neues Grund-
inneren Unendlichkeit teilhaben,
einer
zu
umfängt und beherrscht,
die ihn zunächst
Ganzes seiner Seele zugegen
sein.
verhältnis zur Welt,
um eine neue Art des Lebens und Seins.
Steht aber die Sache so, so
keine
fertige
könnte,
ließe,
Tatsache,
die er
ist
Persönlichkeit für den Menschen
bequem und
kein sicherer Ausgangspunkt,
sondern
Aufgabe,
sie
sie
bedeutet
für
verlangt eine völlige
hurtig
sich
aneignen
der sich mühelos einnehmen
ihn
große
eine
und
schwere
Umwälzung der vorgefundenen
Lage.
Nicht
Selbst,
sondern der Gewinn eines neuen Selbst steht hier in Frage.
Die
die
Entfaltung
oder Ausschmückung des natürlichen
Wendung wird nur da den vollen Ernst und Nachdruck erwo sie auch ein entschiedenes Nein umfaßt, eine Ver-
langen,
neinung der naturhaften Selbsterhaltung, ein Hinausstreben über die
bloßmenschliche Form des Geisteslebens. Und es darf solches Nein
keinen
flüchtigen
Durchgangspunkt bilden,
es
muß
stets
zugegen
Zu den Problemen des Menschenlebens.
350
und energisch
bleiben
wenn nicht das Streben
festgehalten werden,
dem neuen Sein immer wieder zur
nach
zurücksinken
Lebensform
naturhaften
soll.
Ja auch innerhalb des Geisteslebens bildet Persönlichkeit einen
Anstieg und eine Konzentration,
die
Erfahrungen und
durch
erst
Entscheidungen des ganzen Menschen hindurch
Das
erreicht wird.
Leben durchläuft die drei Stufen einer grundlegenden, kämpfenden,
überwindenden Geistigkeit. Zuerst gilt es die Anerkennung einer
Aufgabe überhaupt, eine Erhebung des Lebens über die
geistigen
eine
Natur,
Entwicklung
geistiger
Größen und Güter
jenseit
Das ergibt die Scheidung eines
natürlichen Selbsterhaltung.
der
Idealis-
mus vom Naturalismus, der alles Geistesleben als eine Weiterführung
der
bloßen Natur
und damit
behandelt,
die
Scheidung der
erste
Aber der Boden des Idealismus erzeugt alsbald
Geister.
ein neues
Problem: das Reich d£r Erfahrung zeigt starke Widerstände gegen
die
vom
Idealismus geforderte Ordnung;
es
fragt
ob
sich,
Widerstände die Bewegung zum Stocken bringen, oder ob
Preisgebung der Sache; hier
muß irgendwelche Verstärkung, irgendDiese aber
welche Weiterbildung des Geisteslebens erfolgen.
der
in
von
Dort entsteht der Pessimismus mit seiner
ihr überwunden werden.
die
diese
sie
Wendung zum
Persönlichsein
behauptet
ist
wird.
es,
Das
Persönlichsein erscheint hier als der
Höhepunkt einer geistigen Be-
wegung, und zwar
der
indem
bindet,
als
ein solcher,
sie
zu einem Ganzen ver-
Momente fest-
er die früheren Stufen als beharrende
Denn das Leben bleibt unablässig in Fluß, immer neu ist der
Aufstieg von der Natur zum Geist zu vollziehen, immer neu der
hält.
Widerstand
gegen
die
Vergeistigung
des Daseins
zu
durchleben,
immer neu eine innere Überwindung zu suchen. So bleibt das
Ganze eine fortlaufende Tat, ein unablässiger Aufstieg; es läßt sich
erwarten, daß nicht der ganze Umfang des Daseins darin eingeht,
daß das Persönlichsein
hält;
es
Kraft,
Besitz,
ist
die
nicht
Seele
sondern
sein als ein
bei
uns
selbst einen
inneren Widerstand be-
sowohl unser ganzes Dasein
Augenscheinlich
der Seele.
das
höchste
Persönlichwerden.
Ziel,
Wie
nicht
als seine
ist
sowohl
es
so
ein
bewegende
nicht
ein
Persönlich-
dabei das Streben einem Ein-
strömen des naturhaften Ich unablässig zu widerstehen hat, so haben
die Begriffe immerfort ein Zurücksinken zur bloßmenschlichen Vorstellung abzuweisen,
lässig bedroht.
womit
eine Ermattung des
Denkens sie unab-
Persönlichkeit und Charakter.
35I
Wer solche Aufgaben und Verwicklungen im Persönlich>x'erden
der wird auch die Kämpfe zu würdigen wissen, die
anerkennt,
Problem
dies
Persönlichkeit
sich
knüpfen.
der Religion ward die Idee
In
zugunsten eines unpersönlichen Geisteslebens
mit jener
art
der
oft be-
das natürliche Ich des Menschen starr
kämpft,
weil
gelegt
und zugleich das höchste Wesen zu menschenartig gefaßt
schien;
die
Wendung zu einem
unpersönlichen Sein
derung einer völligen Auflösung des Einzelnen
dünkte
Unendlichkeit
nehmere Denkart.
demgegenüber eine
fest-
mit der For-
den Ozean der
in
größere,
reinere,
vor-
So die pantheistische Spekulation und die Mystik,
so die Höhe der indischen Religionen, so Spinoza mit seinem Worte,
wer Gott wahrhaft liebt,
daß,
wieder
Solche
liebe.
nicht verlangen könne,
Denkweise hat
ihr Recht
daß Gott ihn
der Abweisung
in
der kleinmenschlichen Daseinsform, aber mit dieser Verneinung,
Versinken
dem, Geistesleben keine neue,
wer
in
dem
den Abgrund der Ewigkeit, kann nur abschließen, wer
in
selbständige
zuerkennt,
Wirklichkeit
ihm wohl eine Befreiung von den Wirren und Mühen der
menschlichen Existenz,
von der Unruhe und Flucht der Zeit, von
der Enge und Starrheit des kleinen Ich erblickt, nicht aber das Auf-
und den Gewinn
steigen
Nein
kann
sich
neuen
eines
nur eine
Beruhigen beim
und vorwiegend passive
Lebens.
kontemplative
wo immer
Lebensführung, eine weichere, mattere, schlaffere Denkart;
das Geistesleben mehr Kraft und Zuversicht entwickelt, da wird es
das scheinbar Unmögliche wagen
einem Ja verlangen, da wird
der Persönlichkeit angibt.
Idee
und über das Nein hinaus nach
es die Bahnen einschlagen, welche die
Nun
von der Gefahr eines Zurücksinkens
begleitet sein;
in
aber wird dies Streben
stets
die natürliche Lebensform
in
der Tat pflegt bei den Religionen eine höhere und
eine niedere Art durcheinanderzulaufen: dort das Aufstreben zu einer
neuen Welt, einem neuen Leben und auch einem neuen Gedankenfür
reiche,
bole
liefert;
Daseins
das
unser
menschliches Dasein
hier dagegen
möglichst
Gegenstück der
gut
alten,
die
Neigung,
auszukommen,
mehr
nicht
innerhalb des
die
neue
Welt
als
Sym-
gegebenen
ein
bloßes
die höchsten Begriffe anthropomorph gestaltet,
in
dem allen weit mehr eine Festlegung des Kleinmenschlichen
ein
Aufklimmen zu neuen Höhen.
ihrer
Als
Verdunklung des unentbehrlichen
gegen das Persönlichsein
der weltgeschichtlichen
in
Abweisung
Nein
ist
als
dieser Art mit
der Widerstand
gutem Recht und gewiß dem Ganzen
Bewegung
unentbehrlich;
er
gerät aber ins
Zu den Problemen des Menschenlebens.
352
wenn er mit der niederen Art auch die höhere abweist und
Unrecht,
damit
An dem Gewinn einer solchen Gestaltung liegt schließlich
aufgibt.
auch
Hoffnung einer positiven Gestaltung des Geisteslebens
alle
Hoffnung einer gründlichen Überwindung des niederen
alle
Lebenstriebes.
Denn letzthin ist positivem Streben nur ein anderes
positive Streben gewachsen; alle Energie der Verneinung, alles Ver-
schwimmenwollen
die Unendlichkeit wird
positive
die
So
steht
des Nein
ist
hier
in
Frage,
Lebens- und Glücksdrang sich der bloßen Natur
entwinden und der Stufe des Geistes zuführen
Fall
Selbstsucht nicht
Bildung eines neuen geistigen Selbst
als die
und zwingenden Aufgaben.
mit großen
ob der
in
brechen
so gründlich
alle
läßt
oder
Im
nicht.
Mühe schließlich verloren.
unsägliche
Ahnliche Probleme wie die Religion zeigt an dieser Stelle auch
und
Künstlerische
die Kultur.
der
intellektuelle Weltbilder treffen in
Abneigung gegen eine leitende Stellung der Persönlichkeit zusammen.
Denn
zum
sehen darin ein Zurückziehen der geistigen Arbeit
sie
bloßen Menschen und eine widrige Störung ihres eignen Fortgangs
durch eine Einmischung seiner kleinen Sorgen.
Sich rein
ständig entfalten zu können scheint das Geistesleben nur,
wenn
es
vom Menschen und seinen Zwecken ledig-
unter völliger Ablösung
lich
und voll-
der eignen sachlichen Notwendigkeit folgt und sich im eignen
Kreise zu einem selbständigen Lebensganzen mit eigentümlichen Gesetzen zusammenschließt
Auch
hier aber
wird eine niedere und eine höhere Fassung
der Persönlichkeitsidee zusammengeworfen und mit dem, was herabzieht,
etwas aufgegeben, dessen die Kulturarbeit zur Erreichung ihrer
vollen
Höhe notwendig bedarf. Recht verstanden liegt das Sein und
die dem Streben nach Persönlichkeit vorschwebt, nicht
die Einheit,
neben der Arbeit, sondern innerhalb
zu bringen, das ein Selbstleben
in
ihrer.
Diese selbst
ist
dahin
ihr durchbricht, daß ein geistiges
dem
Denn es gibt schlechterdings keinen Inhalt ohne ein Selbst, das im Wirken und Geschehen
Sein sich in ihr erfaßt, die Erfahrungen in Erlebnisse umsetzt,
Geschehen
allererst einen
sich entfaltet.
Inhalt verleiht.
Nur ein solches innerhalb des Geisteslebens befindliches
Selbst gewährt diesem
eine
Seele
und einen
Halt,
behütet es vor
der Gefahr, ein leerer Mechanismus oder ein seelenloser Kulturprozeß
zu werden, gibt ihm die Kraft, seine eigne Arbeit zu bewältigen,
von
ihr
einem
bewältigt
und
erdrückt zu werden.
solchen Selbst das Leben
eine
statt
Auch kann nur mit
volle Wirklichkeit
gewinnen
Persönlichkeit und Charakter.
und
sich einer Wirklichkeit sicher
solchen Kern
ein schatten-
während
und traumhaftes Dasein
Empfangene verwandelt.
alles
fühlen,
353
es
ohne einen
führt
und darin
Die Inder zeigen uns solche Ver-
flüchtigung in klassischem Bilde. - Augenscheinlich handelt es sich hier
um eine eifrigere subjektive Aneignung einer gegebenen Wirksondern um eine reale Erhöhung und Umwandlung der
gesamten Wirklichkeit, es handelt sich um den letzten Abschluß der
Kultur, um die Möglichkeit eines neuen, wesenhafteren und seelennicht
lichkeit,
volleren Kulturideals.
Das Entscheidende dabei ist für Menschen
und Völker schließlich die Energie des Lebensaffekts, die kräftigere
oder mattere Erfassung des Lebens; die sachliche Entscheidung aber
Erwägungen, sondern
liegt
nicht bei begrifflichen
keit
der Entwicklung einer neuen Wirklichkeit
hier stehen wir bei den letzten
bei der Möglich-
Nirgends mehr als
Axiomen unserer geistigen Existenz.
Von solchen Überzeugungen aus, die für ein Persönlichwerden
das Aufsteigen einer neuen Welt und eine Umwälzung des natürlichen Seins verlangen, muß uns dje heutige Bewegung zum Persönlichsein als verworren, ja in mancher Hinsicht verfehlt erscheinen.
Die übliche Behandlung der Sache erschöpft sich
in
das Verlangen
einer kräftigeren Konzentration, einer Verstärkung des Subjekts, einer
größeren Selbständigkeit gegenüber der Umgebung.
das geschehen,
Welt
bleibt,
Aber wie soll
wenn der Mensch ein bloßes Stück der vorhandenen
nicht
von innen her an einer neuen Welt teil gewinnt?
Ohne eine Umkehrung des nächsten Anblicks der Wirklichkeit und
ohne den Gewinn eines neuen Lebensbodens könnte leicht die Sache
mehr schaden als nützen, indem sie in eine bloße Verbrämung des
Überspannung des Selbstgefühls, ein
und Fürsichzurechtlegen aller Dinge auslaufen
Und wenn gar die Bewegung zur Persönlichkeit als eine
natürlichen Lebenstriebes, eine
bloßes
müßte.
Genießen
Abwendung von den Weltfragen und eine Zurückziehung auf einen
Sonderkreis verstanden wird, so wäre das kaum etwas anderes als
eine Verherrlichung eines engen und öden Spießbürgertums.
Man
macht dadurch nichts Neues aus dem Menschen, daß man ihm die
Etiquette der Persönlichkeit anklebt. Ohne den Gewinn einer neuen
Welt und eine Erhöhung des eignen Lebens bleibt das Ganze eines
bequemen Surrogate, welche die Probleme des Menschenwesens
und den Ernst der Zeitlage verdecken.^
jener
Es wäre endlich wohl an der Zeit, das unablässige Zitieren des Qoethe„Höchstes Glück der Erdenkinder sei nur die Persönlichkeit«
Eucken, Qrundbegriffe. 4, Aufl.
23
^
schen Wortes:
Zu den Problemen des Menschenlebens.
354
Nach unserer Überzeugung muß das persönliche Leben einen
neuen Weltanblick entwickeln, aus seiner eignen Belebung, seinen
seinen Weiterbildungen
Erfahrungen,
Lebenswahrheiten erzeugen.
und
Reich von Grund-
ein
Mögen sich diese nicht in angemessene
Vorstellungsbilder umsetzen lassen, sie bleiben die Wahrheiten,
wie
schließlich
geistige
alles
die
Leben, so auch das Erkennen tragen,
zentralen Wahrheiten,
denen gegenüber alle anderen
Einsichten nur periphere bedeuten.
Nun behält unsere intellektuelle
sie
sind
die
und überhaupt unsere geistige Lage dadurch
eine starke Spannung,
daß ein Spalt zwischen den zentralen, persönlichen und den
peri-
pheren, unpersönlichen verbleibt, daß sich jene nicht unmittelbar in
Aber trotzdem dürfen wir die Wirklichkeit
diese überleiten lassen.
zwei endgültig getrennte Gebiete zerlegen und das Reich
nicht in
Denn
persönlichen Lebens gegen die große Welt starr abschließen.
das wäre nichts anderes,
und
als
das Leben zwischen leerer Subjektivität
seelenloser Arbeit zerteilen, es wäre ein Verzicht auf seine innere
und zugleich
Einheit
daher
nach
treibende
einer
Von
beiden Seiten
gilt,
das Ziel
seine volle Wahrheit.
Einigung zu streben,
es
ist
eine
als
und richtende Kraft tapfer festzuhalten, obschon wir keine
Aussicht haben, es gänzlich zu erreichen und die beiden Ausgangs-
punkte zu völliger Berührung zu bringen.
Bei solcher Überzeugung treten eine persönliche
jektive Gestaltung
Die subjektive Art
stellt
sich der Wirklichkeit
sobald
sie
über sich hinausgeht,
hinein;
die
persönliche möchte
dringen,
und
eine sub-
von Arbeit und Kultur uns deutlich auseinander.
nicht als zu etwas
ihre
gegenüber und
Besonderheit in
trägt,
Dinge
alle
zum eignen Leben der Dinge vor-
Fernem und Fremdem, sondern
als
zu
dem, worin das geistige Wesen sich selber, die Wahrheit seines eignen
Seins erreicht.
Mit der Verwandlung der Dinge in ein Selbstleben
vollzieht sich hier die Überwindung des Gegensatzes einer subjektivistischen
und
einer
solchen,
die
souverän
objektivistischen
oder
Behandlung zu gunsten einer
eigenständlich
heißen
die
Notwendigkeit der Sache
unmittelbar
ein
und wird
eigner Antrieb
Menschen, hier erst wird die volle Einigung mit der Sache
auf
Denn
könnte.
hier allein erlangt das Schaffen eine volle Selbständigkeit
des
erreicht,
Grund derer sie ihre eigne Natur rein und einfach aussprechen
einzustellen.
Die liebenswürdige und
anmutige Stelle, an der
es
sich
Westöstlichen Divan findet, will gar nicht so schwer genommen sein.
im
Persönlichkeit und Charakter,
Erst die persönliche, die souveräne Art
kann.
355
ist
über das hinaus,
was sich zwischen den Menschen und die Sache zu
Der Mensch kann
die Sache
darf vielfacher Mittel,
nicht
ohne weiteres
nischer Zurüstung,
Übung, Gelehrsamkeit u. s. w.
daß,
was nur
Ziele wird,
daß es den Menschen bei
Aber nun entsteht
und Weg ist, zum Zwecke und
Mittel
Kaum
Hauptsache ablenkt.
sich
festhält
und von der
dürfte ein Volk dieser Gefahr so aus-
wie das deutsche
sein
pflegt.
um an sie zu gelangen, er bedarf dazu tech-
die Gefahr,
gesetzt
stellen
ergreifen, er be-
mit der
Schwerfälligkeit seiner Art; besonders
Gründlichkeit,
aber auch
mühsam gelangt es zu voller
Überwindung der Technik durch das Schaffen, zu jenem Sichselbstin den Dingen, ohne das die Werke keine reinmenschliche
Größe und schlichte Einfalt erlangen können. So besteht eben heute
leben
in
unserem Leben ein arges Mißverhältnis zwischen dem Aufgebot
und
intellektueller
Schöpfungen, die
künstlerischer
Arbeit
und der Erzeugung von
zum ganzen Menschen sprechen und den ganzen
Wenn das Verlangen nach einer mehr persön-
Menschen fördern.
lichen Kultur dies heißen soll, daß
es einfache
Grundzüge aus der
Verwirrung herauszusehen und damit zum Ganzen des Menschenwesens zu wirken
dieses
gilt,
so
ist
dem nur freudig zuzustimmen.
Aber
keineswegs eine Sache raschen Entschlusses, sondern das
ist
Allerschwerste, etwas,
aller Kraft,
wozu es nicht nur der höchsten Anspannung
sondern auch vielfacher Gunst des Schicksals bedarf.
erfahren wir es auch heute:
wie wenig hat
alle
So
subjektive Beteuer-
ung des Wertes der Persönlichkeit uns innerlich weitergeführt, wie
wenig hat sie starke und selbstwüchsige Persönlichkeiten erzeugt!
b)
a.
Geschichtliches
Charakter,
zum Ausdruck und Begriff.
Charakter bedeutete bei den Griechen
sowohl das Werkzeug
zum Zeichnen und Prägen als das Gepräge selbst, das Merkzeichen;
es
schon
hat
im Altertum
geistige Gebiet gefunden.
als
in
die
den
Aristoteles,
spätere
Richtung.
Theophrasts,
tragen,
naheliegenden
Übergang auf das
Es geschah das aber sowohl
ästhetisch-literarischer
Namen
den
des
in
ethischer
Die ethischen Charaktere,
Schülers
und Nachfolgers des
sind freilich aller Wahrscheinlichkeit nach eine
Zusammenstellung aus größeren Werken des Mannes, aber
23*
Zu den Problemen des Menschenlebens.
356
Neigung zu genauer Beobachtung und scharf umrissener Zeichnung verschiedener menschlicher Typen geht auf Aristoteles, den
die
^
Kenner und
großen
Schule
seiner
Freund
eigentümlich.
zurück und blieb
Nach derselben Richtung wirkten die
Wirklichen,
alles
neuere Komödie und die Rhetoren;
war dem späteren Altertum
so
der Blick für die Eigentümlichkeit verschiedener menschlicher Arten
und Handlungen
geschärft.^
Zugleich
aber
bezeichnet
Charakter
auch die Eigentümlichkeit
schriftstellerischer
und künstlerischer Dar
Stellung, das individuelle
Gepräge
Im kirchlichen Sprach-
•gebrauch wurde es
Augustin der technische Ausdruck für ein
seit
w.
u. s.
der Seele durch gewisse Sakramente (im Mittelalter Taufe, Firmung,
Priesterweihe)
unverlierbar
eingeprägtes
geistiges
character sacramentalis, auch spiritualis genannt).
vereinzelt auch
So
(später
findet es sich
im Mittelhochdeutschen, ebenso vereinzelt ist hier die
Bedeutung Schriftzeichen (der karakter a b c).
Gegenwart
Zeichen
hineinreicht,
so
erinnert es
Wie das bis in die
auch an jenen älteren Ge-
wenn der amtliche Stil von der Beilegung eines »Charakters"
(Titels und Ranges) spricht.
Zu allgemeinerer Verwendung im psychologischen und ethischen
Sinn ist bei uns das Wort wohl von Theophrast her und durch die
brauch,
Vermittlung der Franzosen gelangt.^
La Bruyere:
ies
Typus und
^
caracteres
Im Jahre 1687 erschien von
de Theophraste, avec
les
caracteres
ou
typisch im jetzt üblichen Sinne als Bezeichnung gemein-
samer Seins- und Lebensformen dürfte aus der Medizin stammen. Es bemerkt
darüber Dilthey (Sitzungsberichte der K. Preuß. Akademie der Wissenschaften
1896, XIII, S. 18): „In diesem Sinne finden wir auch den Ausdruck zunächst
technisch gebraucht, wenn der Arzt Coelius (wahrscheinlich im 2. Jahrh. n. Chr.)
vom Typus des Wechselfiebers spricht und darunter die Regel seines Ablaufs
So sprechen wir überhaupt von einem typischen Verlauf."
über das Ganze Sauppe, Philodemi de vitiis 1. X, pag. 7: Peripatetici disciplinae suae principis et auctoris exemplum nulla in re magis secuti
sunt, quam ut omnia quae vel in natura rerum existerent vel in vita hominum
versteht.
*
et
S.
publica et privata usu venirent accuratissime observarent et observata sive
libris
singularibus explicarent sive ad sententias suas firmandas et illustran-
das adhiberent.
pag. 8:
neque vita ipsa tantum exempla suppeditabat, sed
maximam notationum copiam nova comoedia habebat. Quae ut eidem saeculi
ingenio originem debebat, atque aristoteleum illud Studium vitam quotidianam
moresque hominum observandi,
phrasti
libris
desumta
in
ita quaedam fortasse ex Aristotelis vel Theousum suum converterat, sed multa plura certe,
quam acceperat, deinde philosophis et rhetoribus suppeditavisse censenda est.
^ Wir besitzen darüber eine ebenso feinsinnige wie tiefdringende Untersuchung von R. Hildebrand: „Charakter
in
der Sprache des vorigen Jahr-
Persönlichkeit und Charakter.
les
moeurs de
viel
Beachtung
siede,
ce
ein
Buch, das auch
und Einfluß gewann.
357
Völkern
bei anderen
Sicherlich
stehen
damit im
Zusammenhang wie andere deutsche Werke zur Charakterschilderung,
so
auch
Qellerts
«Moralische Charaktere", eine Zugabe zu seinen
Hier wie sonst bedeutet Charakter soviel
moralischen Vorlesungen.
wie Bildnis, welches Wort bisweilen zu seiner Übersetzung dient,
Zeichnung, Porträt (so spricht
terisieren"
Rabener
B.
das fort
lebt
in
seinen Satiren von
dem Ausdruck „charakbis zur Gegenwart.
Von dem Bild überIn
der Ausdruck auf die Sache und wird zur Bezeichnung
sich
trägt
z.
zu meinen Charakteren").
w Originalen
der seelischen, namentlich der moralischen Art, der Orundbeschaffenheit des
Menschen.
diesem Sinne kann es eine Fülle verschiedener
In
Charaktere geben, guter und böser Art;
keinen
Charakter haben,
Woher der
Gabe der Natur oder als Werk
das heißt hier einer scharfen Ausprägung entbehren.
ob
Charakter
entstanden
freier Tat,
das bleibt dabei unentschieden.
sei,
Kant hebt den
Erst
wichtigen
die
Höhe,
ihn zu einer
die
These
moralischen Charakter.
Naturell
Begriff auf
und einem schweren Probleme macht.
scharfe Grenze zwischen einem physischen und einem
ethischen
Er zieht eine
als
Charakter schlechthin
und Temperament umfassend,
Menschen machen
läßt,
ist
zeigt an,
nur dieser;
jener,
was sich aus dem
der eigentliche Charakter hingegen, was er
aus sich selbst zu machen bereit
ist.
„Einen Charakter schlechthin
zu haben, bedeutet diejenige Eigenschaft des Willens, nach welcher
das Subjekt sich selbst an bestimmte
praktische Prinzipien
bindet,
die es sich durch seine eigne Vernunft unabänderlich vorgeschrieben
hat"
614).
(VII,
Natur aus
„Es
kommt
hierbei
auf das an,
nicht
was die
dem Menschen, sondern was dieser aus sich selbst macht."
„Die Gründung eines Charakters
Prinzipes des
ist
absolute Einheit des inneren
Lebenswandels überhaupt" (617).
In
diesem Sinne
wollte Kant nicht sagen, der Mensch habe diesen oder jenen Charakter,
sondern er habe überhaupt einen Charakter, „der nur ein einziger
oder gar keiner sein kann."
Diese kantische Fassung mit ihrer Erhebung des
die
Stufe
geistiger
Hunderts«
(Zeitschr.
Selbsttätigkeit
f.
d.
ist
rasch
deutschen Unterricht,
Lebens auf
durchgedrungen; ^
6.
Jahrgang,
7.
Heft.)
der
Ihr
folgt unsere Darstellung jener Zeit.
'
Wie
Charakter
(in
zeigt u. a. eine Abhandlung von
den Abh. d. K. Pr. Akad. d. Wiss. 1803).
rasch,
E. Biester
über den
Zu den Problemen des Menschenlebens.
358
hohe Ton,
dem
in
Wertschätzung, die
Folgezeit von
die
sie
Aber neben
Kant zurück.
und
Charakter spricht,
der Charakterbildung beilegt,
sie
die
weisen auf
seiner ethischen Fassung erhält sich auch
man nicht so viel
einem durch Anpassung und Gewöhnung er-
die ältere, empirisch-psychologische; sonst könnte
von einem
ererbten,
So laufen wiederum in einem
Wirkungen verschiedener Zeiten und Welt-
worbenen u. s. w. Charakter sprechen.
Wort
alltäglichen
die
anschauungen bunt durcheinander.
ß.
Die Lage der Zeit.
Der Begriff des Charakters im ethischen Sinne steht in engem
Zusammenhange mit dem der Persönlichkeit, nur hebt er die Selbstdes Menschen
tätigkeit
mit
dieser Begriff eine präzise
besonderer Stärke
hervor.
So gewiß
Bezeichnung erst spät gefunden
hat,
der
Gedanke einer durch eigene Willensenergie erreichbaren Selbständigkeit
und
Weltüberlegenheit
ihn
hielten
Zeiten,
wo
Zusammenhänge
licher
suchen
zwang.
Stoikern
gefunden,
die
weit
zurück;
hoch
besonders
Auflösung überkommener gesellschaftihren Halt in sich selbst zu
die Individuen
Seinen
ein
reicht
Ausdruck
klassischen
hat
das
bei
den
eigentümlicher Lebenstypus zieht sich von
dort durch die ganze Geschichte, er
ist
namentlich von hervorragen-
den Persönlichkeiten der Aufklärung aufs neue bekräftigt worden.
Auch Kants Erörterungen über den Charakter haben viel Verwandtschaft damit und benutzen mit Vorliebe Wendungen aus dem
stoischen
Die Entwicklung nach dieser Richtung
Gedankenkreise.
die
enthält
Gefahr einer schroffen Isolierung
genügsamkeit
des Individuums,
einer
und
stolzen
Selbst-
Verkennung der Bedingtheit
des Einzelnen, wenn nicht durch sichtbare,
so doch durch unsicht-
bare Zusammenhänge, aber bei solcher Gefahr blieb sie in beson-
deren Zeitlagen das einzige Mittel zur geistigen Selbsterhaltung des
Menschen.
Der Begriff des Charakters reicht aber über solche Zuspitzung
hinaus; als Bezeichnung des Selbstwertes und der Selbständigkeit
des Innenlebens gegenüber aller bloßen Außenwelt, als Bekenntnis
zur Überlegenheit
auch da
in
worfen wird.
dem
der
inneren
Ehren stehen,
Güter über
alle
äußeren kann er
wo jene Isolierung des Individuums ver-
Es berührt sich dann aber der Begriff so eng mit
der Persönlichkeit,
Erörterung bedarf.
So
daß er hier nicht noch einer besonderen
sei
nur mit einigen Worten darauf hinge-
Persönlichkeit und Charakter.
359
wie das Problem des Charakters und der Charakterbildung
wiesen,
vom Standort unserer eignen Zeit sich ausnimmt
Diese Zeit beschäftigt sich viel mit dem Problem des Charakters,
sie klagt zugleich
viel
und ausgeprägten Persönlichkeiten, sie fordert von der Kultur-
teren
mehr Fürsorge für die Bildung
namentlich von der Erziehung,
arbeit,
Aber
von Charakteren.
und Halbheit,
heit
über den heutigen Mangel an festen Charak-
in
dem allen erscheint wieder die Unklar-
die solchen Durchschnittsbestrebungen eigen ist
Nicht selten sieht es aus, als ob unversehens ein moralisches Sinken
und es nun nur einer eindringlichen Zuspräche oder ge-
erfolgt sei,
schickten Einrichtung
So einfach
bringen.
bedürfe,
um die Sache wieder in Stand zu
steht
doch
es
nicht.
Der Mangel an selbst-
wüchsigen und selbständigen Menschen, den wir heute schmerzlich
empfinden, hat sicherlich tiefere Gründe,
Immer mehr ist in den
Wandlungen der Jahrhunderte der Menschheit die mühsam erkämpfte
Innenwelt
erschüttert oder verdunkelt,
üben
kraft
daher
Güter,
ihre
immer weniger Anziehungs-
immer
leerer
werden zugleich die
Dazu die stürmische Okkupation des modernen Menschen
Seelen.
durch die Außenwelt,
die Streberei
nach sichtbaren Erfolgen,
der
wachsende Kampf ums Dasein und die unheimliche Beschleunigung
des Lebens, die Zerstückelung des Menschen durch die immer technischer, immer komplizierter gestaltete Arbeit, die abschleifende
Wirkung einer Durchschnitts- und Massenkultur. Kann ein solches
Getriebe einen Sinn und einen Platz für die Ausbildung selbstän-
diger Charaktere haben?
Wer im Vertrauen auf die innere Notwendigkeit der Sache jenes
Ziel
festhält,
der wird seine Erreichung sich keinenfalls so leicht
denken dürfen, wie manche Strömungen der Zeit, die ihre Schätzung
von Persönlichkeit und Charakter mit möglichstem Nachdruck beteuern, zugleich aber die Bedingungen zerstören, unter
für jene Größen ein Platz ist
ralismus,
Liebe
gültiges,
läßt,
indem
alles
er
bei
Dies tut
z.
allein
den Weltfragen mit besonderer Lust und
begrüßt, was den Menschen als klein
unselbständiges
denen
B. oft ein radikaler Libe-
Stück
und als ein gleich-
einer seelenlosen Natur
erscheinen
zugleich aber auf praktischem Gebiet für die Größe und Würde
des Menschen schwärmt, sich für Humanität begeistert und entrüstet
ist,
wenn er einen Mangel an selbständigen Charakteren,
wuchern der Streberei gewahrt
es
heute
in
den
ein
Über-
Ein solches Überwuchern, wie wir
mannigfachsten
Formen gewahren,
ist
schlimm
Zu den Problemen des Menschenlebens.
360
genug, aber wird ihm kräftig zu begegnen sein, wenn dem Menschen
alle
selbständige Innenwelt
höher
ein etwas
wenn er gar nichts anderes ist als
fehlt,
geartetes Tier
und daher keine anderen Ziele kennt
der natürlichen Lebenserhaltung?
als die
Wir werden an dieser Stelle in einen sicheren Fortgang nur
kommen, wenn die Probleme des inneren Menschen wieder zur
Hauptsache werden, und wenn sie sich zum Ganzen einer Lebensüberzeugung zusammenfassen, das mit aufrüttelnder, richtender,
Davon
höhender Macht die Gemüter zu ergreifen vermag.
Aber so sehr es abzulehnen
einstweilen noch weit entfernt.
die
Bildung von Persönlichkeit und Charakter
als
handelt werde, die sich so nebenbei abmachen
wirken dafür
läßt
unmittelbar auch
sich
erwägen
eignen Zeit;
wir
also
auf
Kürze,
in
ist,
daß
Sache be-
eine
läßt,^
er-
sind wir
einigermaßen
dem Boden
unserer
welchen Richtungen
in
vornehmlich das geschehen kann und geschehen muß.
Zunächst
ist
notwendig mehr Anerkennung und Hochhaltung
der bloßen
mehr Zurückstellen und Durchschauen
Aufmachung und des bloßen Scheins. Was uns daran
hindert,
namentlich der Epikureismus einer reifen,
der echten Werte des Lebens,
ist
überreifen
ja
Kultur, der jeden Einzelnen möglichst seinem individuellen Behagen
nachgehen, ihn ängstlich allen Zusammenstoß scheuen und willfährig
vor
sozialen
aller
alsdann
Wert,
Konvention sich ducken
Der Mensch stellt
läßt.
weniger auf sich selbst und gibt sich selbst seinen
sich
daß er das Gelingen seines Lebens an die Anerkennung
als
durch andere
setzt,
damit aber sich unvermeidlich zu ihrem Diener
Jedes Volk hat hier seine besonderen Gefahren, bei uns
erniedrigt.
Deutschen nehmen unverkennbar künstliche Abstufungen, Fragen des
Ranges, Dekorationen wie Titel und Orden,
werk des Lebens,
damit
einen
viel
das Aufsichselbststehen
zu
alles dieses
bloße Bei-
und schädigen
breiten
Platz
des Lebens
und
seine
Nebensachen
wie
Hauptsachen
Nebensachen
herabgesetzt
haftigkeit.
Unmöglich
lassen
behandeln,
ohne daß
Hauptsachen
sich
zu
ein
volle
Mann-
werden. Jeder Beruf und auch jeder Mensch hat ein Recht auf
Achtung und Anerkennung; er muß es sich erstreiten, wenn es ihm
versagt wird.
1
Es
sei
Aber er
es
„Wohl wachsen Schwämme
nur regnet, aber Menschenwürde,
wächst nicht
daß von außen her
hier an das derbe, aber nicht unberechtigte Wort Pestalozzis
erinnert (Wke. XII, 217):
wenn
erringt es nicht dadurch,
aus der Routine hervor,
wenn
ihr
leicht
aus
dem Mist,
und Charaktergröße
auch die Sonne scheint."
Geistestiefe
Persönlichkeit und Charakter.
ihm Rangklassen oder Dekorationen
bewilligt,
361
sondern dadurch, daß
seine Arbeit bei sich selbst gestärkt, unabhängig gestaltet, zu vollem
Ganze des Lebens gebracht wird.
Einfluß auf das
führt schon auf den zweiten
Das
Das
bildung bedarf.
ist
Wir
die eigne Verantwortung innerhalb unseres Lebenskreises.
und
über Vielregiererei und
pflegen
zu
durch
die
Bureaukratie
jener
die
Tendenz innewohnt,
Selbständigkeit zu
alle
Befugnis
hätte
die
Bureaukratie
von
als
den
Entfaltung
freier
gewiß insofern mit Recht,
zu
als
nur
eine
alles
andere von ihr abhängig zu
einzige
Stelle
voller
Aber
übertragen zu verstehen.
ihr
nimmer
langt, entspräche sie nicht einer
nicht
über Schädigung
klagen,
berufen und
machen^
es
Punkt, dessen die Charakter-
die Selbständigkeit, die freie Entscheidung
solche
eine
Macht
bei
uns
er-
uns innewohnenden Neigung, stäcke
uns eine Lust zu regulieren und zu schabionisieren, über
in
unsere Denkweise eigensinnig
anderen eine Polizei zu üben,
auch den anderen aufzudrängen. Es fehlt uns vielfach die Bereitschaft,
den anderen
lassen,
seiner Art gewähren,
in
wo er uns schroff widerspricht;
uns
erscheint
leicht
als
Verleugnen der eignen
dem Gedanken
eine
Überzeugung.
ein
frei
der eignen Gesinnung,
möglichst
als
erhalten
mög-
um ihnen vorzubeugen, drücken wir
von der Ausnahme
So
ein.
ein
Auch stehen uns leicht bei
den Gesamtstand des Lebens herab, gestalten
von der Regel
schalten zu
solches Gewährenlassen
der Freiheit vorwiegend die Gefahren eines
lichen Mißbrauclis vor Augen;
lieber
Mattheit
ihn auch da
wir
schnüren und engen es
her,
leicht
es nicht sowohl
konventionelle
Gestalten,
Menschen, Exemplare einer bloßen Gattung, während die
t)'pische
Ausbildung individueller Art unterdrückt wird und damit etwas verloren
geht,
dessen
dringend bedarf.
Aufrechterhaltung
die
Wie
vieles
wirkt
innerer
Selbständigkeit
im Leben der Gegenwart zu-
sammen,
um den Menschen abzuschleifen und ihn gleichförmig zu
gestalten,
wie sehr bedrohen Massenwirkungen die Entfaltung unserer
Individualität,
Massenwirkungen namentlich auch
der Individualität
mit besonderem
unsere Individualisten sind
oft
da,
wo das Recht
Nachdruck betont wird!
gar nichts anderes
als eine
Denn
besondere
Art von Gattungsmenschen mit durchaus uniformen Zügen.
Zur freien Entwicklung der Individualität bedarf es aber mehr
Muße,
mehr
innerer
Lebens zu gestatten
Sammlung,
pflegt.
nur zahllose Individuen,
als
die
Hast des gegenwärtigen
Die Überbürdung mit Arbeit, die nicht
sondern
ganze Stände bedrückt,
wird zu
Zu den Problemen des Menschenlebens.
362
Gefahr für eine innere Bildung,
ernstlichen
einer
ruhige
Selbstbesinnung,
alle
beharrende
sie
alle
zu-
Wir haben
sammenhängenden Aufbau des Lebens hemmt.
ausgezeichneten Lehrstand,
Vertreter
ist
ihm
Aber dieser Lehrstand
schwer überbürdet, zürn Teil mit mechanischen Geschäften,
ganz
abzunehmen
wohl
daß
wären;
frische
Menschen einer ganz anderen Wirkung fähig sind
abgestumpfte,
das
wenn erwogen,
wird
keine
nicht
und
als
Raum
für
Beisichselbstsein des Lebens schafft,
Wer
freudige
hier
es
findet,
wie auf
eine innere Bildung,
der
hilft
die
müde und
genügend erwogen, oder
durchgreifende Abhilfe.
anderen Gebieten mehr freien
ein
einen
den besten auf der Erde, den gebornen
und Wesenskultur.
Innen-
einer
indem
Konzentration, allen
für
auch zur Förder-
ung jener Ziele, die das Problem des Charakters vorhält. Wo so
Großes auf dem Spiele steht, da gewinnt auch das Bedeutung, was,
für sich selbst betrachtet, klein scheinen mag.
Freiheit des Willens.
6.
Einleitung.
a)
\/\ an kommt nie zu Ende, wenn man sich auf die Fragen der
"*^* Freiheit einläßt; jede Partei hat unbegrenzte Hilfsmittel",
so sagte der große Kritiker Bayle bei Erörterung des Willensproblems.*
Ein
angesehener
gegen die Akten
nismus für
die
letzten
deutscher
Jahrhunderte so
daß für uns
der Gegenwart
erklärt
da-
des Determinismus gegen den Indetermi-
in Sachen
M geschlossen ".2
aufgeklärt,
Gelehrter
Wer hat nun Recht?
viel
erledigt
völlig
Haben wirklich
Neues gebracht und die Sache so
ist,
was früher die Geister
Oder erscheint uns vielleicht nur deshalb
unversöhnlich entzweite?
die Sache als ausgemacht, weil wir sie unter dem Einfluß besonderer
Gedankenmassen nur von einer besonderen Seite zu betrachten pflegen?
Sehen
wir,
schon
als
wie es damit steht, und ob der Sieg des Determinismus
entschieden gelten darf.
Der Determinismus
seinem Grundgedanken nach
alt,
ver-
ändert ward nur das Nähere seiner Fassung und Begründung.
Bei
den
Stoikern,
dürfen, 3
ist
die
es der
als
ist
die
bewußten
Deterministen
gelten
Gedanke eines durchgängigen Kausalzusammen-
Oeuv. div. (Haag 1727)
^
ersten
III,
794a:
On ne finit point quand on s'en-
gage aux questions de la liberte, chaque parti a des ressources infinies.
'
S.
Meinong, Psychologisch-ethische UntersuchuTigen zur Werttheorie,
S. 209.
Es heißt dort: »Es ist aber nicht die Kontroverse des Determinismus,
die hier
aufgenommen werden soll
:
in ihr sind die Akten längst geschlossen
oder sollten es doch meines Erachtens sein, indem, wer an das Kausalgesetz
glaubt,
konsequenter Weise auch nicht Indeterminist sein kann."
meint bei Anführung dieser Stelle (Ethik,
bei
2.
Höffding
deutsche Aufl., 102), daß
Betrachtung der dänischen Literatur über
diese Frage
einen
man
anderen
Eindruck erhalte.
^
Daß Aristoteles keineswegs Indeterminist war, aber das Problem noch
nicht zu voller Klarheit brachte, hat in sorgfältigster Untersuchung R. Löning
gezeigt (in
dem Werk »Die Zurechnungslehre des Aristoteles", 1903).
Zu den Problemen des Menschenlebens.
364
hanges der Welt, der
es
alle
Willkür an den einzelnen Stellen aufhebt;
drängt hier weniger die psychologische Analyse
als
das Ganze
Nachdem
die
praktisch-
Weltansicht
der
zu
jener
moralische Richtung
des
Behauptung.
ältesten
Christentums
der Willensfreiheit
das entschiedene Übergewicht gegeben hatte, ohne sie aber wissenschaftlich irgend zu
begründen, verficht Augustins theozentrische Be-
trachtung der Wirklichkeit einen völligen Determinismus; jede eigne
Entscheidung des Menschen erscheint hier
Allmacht und der Allwissenheit Gottes.
als
Aufhebung der
eine
Die Milderung, welche die
Kirche und das Mittelalter daran vollzogen, wurde von der Reformation, namentlich ihren Anfängen, aufs schroffste abgewiesen und
der religiöse Determinismus wieder in voller Stärke aufgenommen.
Die Höhe der Aufklärung bringt einen kosmischen Determinismus
und
gibt
ihm
bei
Ein scheinbarer
Spinoza seine klassische Gestalt.
Gegner des Determinismus wie Leibniz bildet diesen in Wahrheit nur
feiner durch,
und
gibles Reich
gewährt unserm inmitten der Zeit befindlichen Leben
die kantische Rettung der Freiheit in ein intelli-
und Handeln keine genügende
Hilfe.^
So überwog schon vor dem
19. Jahrhundert
in
der geistigen
Arbeit entschieden der Determinismus, gerade an ihren Höhepunkten
gewann
er besondere Klarheit
und Kraft,
er scheint hier die Lebens-
energie nicht zu mindern, sondern zu steigern.
Paulus stand
dem
Determinismus näher als irgend ein anderer zu Beginn des Christen-
tums und durfte zugleich von
habe als die anderen
Natur und
von
alle;
sich
sagen,
daß er mehr gearbeitet
auch Augustin war eine unablässig tätige
gewaltigem Organisationsvermögen.
Und
in
den
Kämpfen der Reformationszeit war die Überzeugung, im Tun und
und gar, aber auch lediglich und allein von Gott,
Ergehen ganz
nicht von irgendwelcher menschlichen Macht, abhängig zu sein, die
Hauptquelle festen Vertrauens und unbeugsamer Willenskraft.
Zu diesem allen kommt die neueste Phase des Determinismus
im
19. Jahrhundert.
Ging er früher aus religiösen oder spekulativen
Überzeugungen hervor, so ist es jetzt die gründlichere Durchforsch-
ung der Erfahrung, welche von den verschiedensten Punkten her
zu ihm drängt und ihn sowohl anschaulicher als eindringlicher
macht
Immer dichter sehen wir jetzt das Netz der Kausalität den
*
Streng genommen, müßte jene intelligible Freiheit das ganze Leben
in
der Zeit
zur Untätigkeit
wegung rauben.
verdammen,
ihm
alle
Möglichkeit
eigner Be-
Freiheit des Willens.
Menschen umschlingen,
stock
seiner
Natur
Erfahrungen gelangen dabei durch die
alte
präzisere Fassung zu neuer
365
und
Wirkung.
gesteigerter
Den Grund-
der Mensch augenscheinlich ererbt,
hat
weitere Entwicklung hängt an der gesellschaftlichen
seine
Umgebung und
an der Erziehung; wenn er zu klarer Bewußtheit erwacht, findet er
im wesentlichen schon
sich
fertig,
das Schicksal,
nicht sein eigenes
daß
Wollen, hat ihn gebildet.
Die neuere Gesellschaftslehre
unser Handeln bis
Wurzeln hinein dem Einfluß der Gedie geschichtliche Betrachtung, daß wir
in
samtverhältnisse unterliegt,
ganz und gar Kinder der Zeit sind, selbst da,
Die moderne Psychologie aber
sprechen.
Gewebe des Innenlebens blicken,
verkettet
ein
und
zeigt,
seine
sie
zeigt
läßt
uns genauer
jede einzelne
bedingt, ja ringsumher festgelegt,
Geschehen unvermittelter Art.
wo wir ihr wider-
sie
in das
Handlung
duldet nirgends
Dabei aber hofft man der moral-
Man
ischen Seite des Lebens ihr volles Recht gewähren zu können.
sucht
darzutun,
antwortlichkeit,
daß,
was
ihr
notwendig
ist,
wie
z.
B.
Ver-
die
auch nach Aufhebung der Freiheit verbleibt;
ja
die
moralische Aufgabe scheint dadurch erheblich zu gewinnen, daß die
einzelne Handlung einem Ganzen des Lebens und dies Ganze wieder
geschichtlich-gesellschaftlichen
Denn wenn damit
die
Zusammenhängen fester eingefügt wird.
dieser Zusammenhänge die
Verbesserung
Hauptaufgabe des Handelns wird, so erhält es eine breitere Basis
sowie festere Angriffspunkte.
Zugleich entfaltet sich stärker das Ge-
lühl einer moralischen Solidarität
und
läßt die Verfehlung des
Indi-
viduums milder beurteilen. So zeigt namentlich das moderne Strafrecht bedeutende Bewegungen zu größerer Humanität unter dem Einfluß
des Determinismus.
Wenn in dieser Weise ^Ue Interessen zusammen-
streben und die engere Verflechtung mit dem unmittelbaren Dasein dem
Gedanken der Jahrtausende eine gesteigerte Wirkung gibt, so
verstummen zu müssen und an dem endsich nicht mehr zweifeln zu lassen. So
Determinismus
gültigen Siege des
alten
scheint aller Widerspruch
reicht
denn das Bekenntnis zum Determinismus weit über die Kreise
der Wissenschaft hinaus und gehört, wenigstens in Deutschland, zu
den Ausrüstungsstücken des Menschen, der sich »auf der Höhe der
Bildung" fühlt
Von hier aus glaubt der Adept moderner Weisheit
auf alle, die gegen den Determinismus noch irgend welche Bedenken
haben,
als
auf arg Zurückgebliebene
stolz
herabsehen zu dürfen.
Zu einiger Behutsamkeit demgegenüber könnte wohl die Wahrnehmung stimmen, daß es noch immer eine ganze Anzahl von
Zu den Problemen des Menschenlebens.
366
Forschern gibt,
tüchtigen
welche
dem Determinismus
widerstehen,
und daß sie eher zu wachsen als abzunehmen scheint, ^ sowie die
andere Wahrnehmung, daß bei anderen. Kulturvölkern der Determinismus keineswegs
so argem Verruf steht wie bei uns Deutschen.
in
Das gilt namentlich von Frankreich, wo die «Diskontinuitätsphilosophie"
bewußt und energisch den Determinismus ablehnt, wo
selbst die
in
lebensvolles Gemälde des Seelenlebens die Freiheit als wesent-
ein
lich
einschließt.
Das bezeugt doch
schon endgültig erledigt
deutschen Lebens
b)
Boutroux
ein
„Kontingenz" der Naturgesetze verteidigt,^ wo ein Bergson
ist
als
vielleicht,
unter
daß die Sache weniger
besonderen Strömungen des
erledigt erscheint.^
als
Erwägungen zur Behauptung des Determinismus.
Ein Problem,
das so
viel
Verwicklung
in
sich trägt,
und das
so sehr die Zeiten wie die Geister entzweit, in raschem Fluge nebenbei behandeln zu wollen, das
müßte als unbescheiden,
ja
vermessen
Aber wer sich mit den geistigen Strömungen der Gegenwart
gelten.
befaßt, der
muß irgend auch der Bewegungen gedenken, welche in
der Behandlung jenes Problems zutage treten.
Die Art, wie sich
heute der Determinismus in der Breite des Lebens ausnimmt, scheint
uns an einem starken Dogmatismus zu leiden und das alte Problem
viel
zu
aus
dem
Gesichtswinkel der besonderen Zeit zu
Wer auf
die
Geschichte
einseitig
behandeln.
blickt,
der
erhält
nicht den
Eindruck, daß der Determinismus sich zu seinem Gegner wie Höheres
zu Niederem
verhält,
und daß bei wachsender Aufklärung der Wider-
mehr und mehr verschwunden
stand
ist.
Denn schon vor Jahr-
tausenden lag die deterministische These deutlich vor Augen, es sind
dann immer wieder Gegenströmungen gekommen, und zwar nicht
nur in den Niederungen des Durchschnittslebens, sondern auch auf
^
Es seien
u. a.
von Werken der letzten Jahre erwähnt: von Rohland
»Die Willensfreiheit und ihre Gegner" (1905), Froehlich «Freiheit und Notwendigkeit" (1908), Joel „Der freie Wille" (1908).
"^
und
in
E.
Boutroux „Über den Begriff des Naturgesetzes in der Wissenschaft
der Philosophie
Boelitz „Die Lehre
der Gegenwart"
(deutsche Übers. 1907),
s.
auch
vom Zufall bei Emile Boutroux" (1907).
^ In
vortrefflicher Weise hat zur Klärung des Problems neuerdings
Windelband gewirkt („Über Willensfreiheit", 2. Aufl. 1905), indem er die
Notwendigkeit einer Scheidung verschiedener Formen zeigte, „die in dem
Worte ,Willensfreiheit' kritiklos zusammengefaßt zu werden pflegen" (s. S. 222).
Freiheit des Willens.
der
Höhe der
geistigen Arbeit,
den
leitenden
Deterministen
Durchführung
ja,
367
was am schwersten wiegt,
Zur
selbst.
vollen
bei
Konsequenz der
der Determinismus wohl an keiner Stelle gelangt.
ist
Wenn die Stoiker das All in ein
kausales
Gefüge verwandeln und
dieses auch das Schicksal des Menschen gänzlich bestimmen lassen, so
bleibt seiner eignen
Entscheidung vorbehalten, ob er den Weltlauf an-
erkennt und in die eigne Überzeugung aufnimmt, oder ob er sich
ihm fortschleppen läßt;
widerwillig von
die Möglichkeit einer solchen
Entscheidung, dieser Kern der stoischen Moral, durchbricht aber augen-
Augustin ist strenger Determinist nur
scheinlich den Determinismus.
so lange, als ihn gänzlich die theozentrische Betrachtung erfüllt;
so-
bald er das menschliche Handeln, im besondern das praktisch-kirchliche Leben, ins Auge faßt, gilt ihm der Mensch als zu selbstständiger
und zu eigner Entscheidung berufen. Auch Luther hat
die anfängliche Strenge seines Determinismus später beträchtlich ge-
Mitarbeit
mag den Menschen noch
Spinoza endlich
mildert.
so sehr einer
lückenlosen Weltverkettung angehörig zeigen, für ihre Anerkennung
ist
er
erst
gewinnen;
zu
Wendung des Lebens,
zu
mit
eine
einem
licher
Illusionen
nicht
endlich auch bei einer
minismus wie
ihr
ergibt
sich
aber
eine
völlige
Wendung von einem Gewebe menschMacht es
lauterer Wahrheit
mehr empirischen Fassung des Deter-
Reiche
der Gegenwart einen großen Unterschied, ob die
in
Verkettung uns gänzlich unbewußt wirkt, oder ob sie von uns durch-
und
schaut
in
unser Handeln aufgenommen wird?
Durchgängig
Ordnung nicht schon ihre
Anerkennung mit sich bringen, nach dem Ja oder Nein an dieser
Stelle aber das Leben sich grundverschieden gestalten. Ganz gleichwir die Tatsache einer kausalen
sehen
gültig scheint die Entscheidung des
Auch
die
Kants
Stellung
zu
Menschen also nicht zu sein.
unserem Problem könnte die
heutigen Deterministen vor einem allzu sicheren Auftreten warnen.
Kant pflegt auch ihnen
System
aber
ist
ein
entfernen heißt seinen
Man
großer Denker zu gelten und sein
philosophische Leistung der Neuzeit
Die
unentbehrlicher Eckstein dieses Systems,
ihn
als die gewaltigste
Freiheit
^
als ein
Gesamtbau
zerstören.^
Hat doch Kant die
sehe nur die Äußerungen Kants in der Vorrede zur Kritik der
praktischen Vernunft: »Der Begriff der Freiheit, sofern dessen Realität durch
apodiktisches Gesetz der praktischen Vernunft bewiesen ist, macht nun
den Schlußstein von dem ganzen Gebäude eines Systems der reinen, selbst
der spekulativen Vernunft aus« (V, 3, Hart.). Femer: „Freiheit ist aber auch
ein
Zu den Problemen des Menschenlebens.
368
Raum und Zeit und die Realität des Freiheitsbegriffes
um welche sich die Vernunftkritik drehe,
von
Idealität
die beiden Angeln genannt,
und ist auch seiner Erkenntnislehre der Freiheitsgedanke von Anfang
an gegenwärtig.
Mag man der besonderen Art, wie Kant das Frei-
heitsproblem löste,
noch so
kritisch
gegenüberstehen, es gibt doch
zu denken, daß dieser große Philosoph auf Freiheit glaubte unmögverzichten zu können.
lich
Was ist es wohl, das trotz aller Häufung anscheinend unwiderleglicher Gründe immer wieder über den
Determinismus hinaustreibt?
daß seine konsequente Durchführung alles zerstören
Dieses
wohl,
müßte,
was dem geistigen Leben des Menschen eigentümlich
ist.
Dem Determinismus ist, wie die Dinge draußen, so auch die Seele
aus
dem Zusammentreffen
ein gewisses Ergebnis mit
zwingender Notwendig-
Menschen eine gegebene
des
ihm
beider geht
hervor.
keit
Aber
läßt
sich
Größe;
im Grunde dann noch
von eignem
Handeln reden, und können wir uns innerlich irgend dafür verant-
Wird die Sache in ihren eignen Grenzen gehalten
überkommene Bild unseres Lebens
und Wesens verändert oder ergänzt, so müßten wir bloße Zuschauer
wortlich fühlen?
und
nicht unvermerkt durch das
dessen werden, was an uns vorgeht, an der Seele genau wie an dem
Körper; was
je
aus uns werden wird, das
liegt
mit sicherem Zuge
schon vorgebildet, wir spielen eine zugewiesene Rolle, wir müssen
eine vorgezeichnete Landstraße geduldig weiter
als völlige
Sklaven des Schicksals.
und weiter wandeln
Verschwinden würde damit
alle
wo es keinen Aufruf zur Entscheidung,
keine Spannung, kein ursprüngliches Handeln gibt, wo die Zukunft
wahrhaftige Gegenwart, denn
aus der Vergangenheit wie die Frucht aus
dem Samen
da gibt es nur den Schein einer Gegenwart.
zugleich aller innere Zusammenhang
und
alle
hervorgeht,
Auch verschwindet
beherrschende Einheit
des Lebens, denn eine solche läßt sich nicht überliefern,
nur aus eigner, ursprünglicher Tätigkeit hervorgehen,
sie
sie
kann
muß immer
von neuem geschaffen werden; so verwandelt sich unsere Seele dort
in
ein
Nebeneinander einzelner Elemente, das nach außen hin ein
Ganzes scheinen mag, das
entbehrt.
lichkeit,
die
In
in
Wahrheit
aller
inneren
Verbindung
Summa ist es namentlich die Leugnung aller Ursprüng-
welche jener Denkweise anhaftet, und dieser Verzicht auf
einzige
Möglichkeit
unter
a
allen
priori
Ideen
wissen,
der spekulativen Vernunft,
ohne
sie
wovon wir
doch einzusehen, weil
dingung des moralischen Gesetzes ist, welches wir wissen"
sie
(V, 4).
die
die Be-
Ursprünglichkeit,
dies
Freiheit des Willens.
35g
gänzliche Geschoben-
und Getriebenwerden
durch ein dunkles Geschick,
näher erwogen, etwas ganz Ent-
Das
Unerträgliches.
schlechterdings
setzliches,
ist,
Entsetzliche
Verstrickung in ein allgewaltiges, unentrinnbares Schicksal,
einer
das uns
nun und nimmer losläßt, haben namentlich die tieferen Geister unter
den Indern mit erschütternder Stärke empfunden; so ward ihnen die
Befreiung von solchem Schicksal, von dem Rad der Wiedergeburt,
zum dringendsten Anliegen, zur sehnlichsten Hoffnung.
Aber, so wird uns eingewandt, was
hilft
Sträuben gegen
alles
eine eherne Notwendigkeit? Ergebung und Resignation ist das einzige,
dem Menschen bleibt und
das
ziemt.
Erhält er nicht in Wahrheit
seine Natur als ein unabweisbares Erbe mitgeteilt, haben ihn zusammen
mit diesem nicht seine nähere und seine weitere Umgebung zu dem
Und ist es nicht das Schicksal, das den
gemacht, was er heute ist?
Fertigen
Motive,
wenn
oder dorthin
hieher
Bedarf
läßt?
ferner
nicht
stellt,
ihn
und würde nicht das Leben
oder jenes wirken
dieses
menschliche
das
Handeln
bestimmter
Chaos
in ein wirres
unter diesen Motiven willkürlich zu wählen wäre,
zerfallen,
wenn sich
ohne allen Zusammenhang mit dem bisherigen Tun der Gute zum
zum Guten entschließen könnte?
Bösen, der Böse
verlangen
Sicherlich
Würdigung; ob
sie
solche
Wahrheiten
gewissenhafte
eine
aber die Sache erschöpfen, ob
sie
das Eigen-
tümliche des Menschen mit seiner geistigen Art vollauf zur Geltung
bringen, das
ist
nicht so leicht zu sagen.
Es
ist
eine unbestreitbare
Tatsache, daß der Mensch, zunächst mit seinem Denken, nicht einfach
innerhalb der
sondern
daß
Verkettungen
des
Daseins verbleibt,
wie das Tier,
sich
ihnen
gegenüberzustellen und sie als Ganzes zu überschauen vermag.
Ohne
er
aus
diesen
Verkettungen
heraustritt,
das gäbe es kein Fragen nach Wahrheit, und schon daß eine solche
Frage überhaupt
des Lebens.
gestellt wird,
Steht es mit
enthält eine erhebliche Weiterbildung
dem Tun nicht ähnlich? Wir gehen nicht
auf in ein Nebeneinander einzelner Antriebe,
einer
eine neue Stufe des Lebens;
als
erheben uns zu
wir
überlegenen Einheit und gewinnen damit eine Selbsttätigkeit
heit überblicken
von
hier aus
können wir jene Viel-
und jedes Stück nach seinem Werte schätzen;
dieser
Wert ist nicht schlechthin gegeben und ausgemacht,
er gestaltet sich
von jener Einheit aus, und ihre Umgestaltung
muß auch ihn
erst
verändern.
Fragt man uns, wie jene Selbsttätigkeit,
wie jenes Hervor-
brechen ursprünglichen Geisteslebens im Menschen möglich
Eucken, Grundbegriffe.
4. Aufl.
24
sei,
wie
Zu den Problemen des Menschenlebens.
370
sie sich
aus
dem Ganzen des Alls erkläre,
so
bekennen wir offen
Unvermögen einer Antwort darauf. Aber wie arm
würden wir werden, wenn wir alles leugnen wollten, was wir nicht
zu erklären vermögen ? Wir sehen um uns in unerschöpflicher Fülle
bewußte und fühlende Wesen, eigentümliche Lebenseinheiten, immer
Ist uns das irgend erklärlich, könnte man
von neuem entstehen.
nicht, wenn es nicht eine unbestreitbare Tatsache wäre, es ebenso
und
ehrlich das
als
unmöglich
verwerfen
wie
Erwachen
ein
von
Selbsttätigkeit?
Denn eine Lebenseinheit scheint weder durch eine Zusammensetzung
von Leblosem noch durch eine Teilung von Lebendigem entstehen
zu
können.
kann es neue Lebenseinheiten
So müssen wir doch wohl unsere
leugnen.
keit
Also
in
keiner Weise
Aber sie quellen unablässig auf, wir können sie unmöglich
geben.
Begriffe
von Möglich-
der Wirklichkeit der Dinge unterordnen, nicht diese in ein von
unserm engen und kleinen Verstände gebildetes Maß hineinzwängen.
Im Grunde ist es der Intellektualismus, der dem Determinismus die
Wurzeln liefert
Dazu sei bei diesem Problem auch die besondere Lage des
Menschen mit ihrer Verwicklung nicht vergessen. Daß er einerseits
zur Natur gehört und andererseits den Beginn einer neuen Stufe
stärksten
der Wirklichkeit, eines Reiches des Beisichselbstseins,
wandelt
sein
bildet,
das ver-
ganzes Leben in eine Aufgabe, zu deren Lösung es
unbedingt seiner
eignen Entscheidung bedarf.
entgegengesetzte
Antriebe,
das
schlechterdings unvergleichlich.
gesellschaftliche Dasein
macht
die
Das stellt es unter
Motive
hie
und
da
Dort das« natürliche oder auch das
mit seiner Lust, hier die geistige
Ordnung
mit ihrem neuen Selbst und seiner Unendlichkeit,
Läßt sich direkt
miteinander vergleichen, und aneinander messen,
was ein Handeln
an
selbstischem
Glück,
und was
es
an Erhöhung des Wesens in
Erfüllung von Pflicht und Entwicklung von Liebe bringt?^
scheinlich
stehen
die Hauptrichtung
wohl um
*
das,
hier
einzelne Leistungen,
nicht
des Lebens in Frage,
was wir tun,
als
es
Augen-
sondern es steht
handelt
sich nicht so-
um das, was wir sind, oder vielmehr
So müssen wir auch die Vorstellung ablehnen,
als
ob die Motive
und gegebene Größen wären, die in der Seele wie Gewichte auf einer
Wagschale zusammenträfen und eine Entscheidung bewirkten. Muß denn
alles Handeln aus gegebenen Motiven erfolgen, können bei innerer Wandlung
feste
des Lebens nicht neue entstehen?
Und muß die Seele den Motiven ihren
Wert nicht immer von neuem erteilen?
Freiheit des Willens.
um
371
worin wir unser Wesen setzen.
Die Seele des Menschen
bildet nicht einen bloßen Schauplatz, an
dem zwei Stufen der Wirk-
das,
begegnen, sondern
lichkeit sich
wird selbst zur Mitwirkung auf-
sie
gerufen, nur durch selbsttätige Aneignung hindurch kann das Geistes-
leben
an dieser
darin
an
zur vollen Wirklichkeit
Stelle
Entscheidung
liegt,
das
kommen.
Was aber
ein
besonderer
nicht
vollzieht
Augenblick, sondern das vollzieht unser ganzes Leben, das will unablässig bejaht
und bekräftigt sein.
Geistiges Leben, so sahen wir,
verbleibt nicht, wie es einmal sich findet, es muß immer von neuem
So behält unser Leben
entstehen, soll es nicht aufs rascheste sinken.
stets eine Spannung, es wird, geistig angesehen, nie zu ruhigem Besitz.
Damit
erscheint
Wirken
freies
Wendung
Augenblicks, die
Sache
des
bloßen
nicht als ein plötzlicher Einfall.
Denn
nicht
eine
als
auch der Augenblick eine hervorragende Bedeutung ge-
so gewiß
winnen kann, er kann das nur in weiteren Zusammenhängen, nur als
der Höhepunkt eines durchgehenden Strebens.
An erster Stelle geht
die Frage auf das Ganze, auf die Hauptrichtung des Lebens,
nicht
auf einzelne Punkte.
Wie in unserem Leben Freiheit und Schicksal zusammenwirken,
wie das eine auf das andere angewiesen
ist,
das zeigt mit beson-
derer Anschaulichkeit die Bildung einer geistigen Individualität
bloßem Entschlüsse
Un-
möglich
kann
Schicksal
geht hier voran und weist uns die Richtung des Weges.
Aber sofern
erst
eine
dieses Individuelle geistiger Art
errungen,
geschieden,
Stärke
solche aus
muß
eigne
in
Tätigkeit
Mittelpunkt
als
erst
freigelegt
muß
aufgenommen,
anerkannt
und
ist,
werden.
der Seele der Seele, kann harte
des eignen Wesens,
von uns
es
von
das
Fremdem
Der Punkt unserer
sein.
ergriffen
entstehen,
Das Suchen
Mühen und
schwere Irrungen kosten, wir können uns weit verlaufen, ehe wir
Und wenn wir ihn gefunden haben,
Mühe und Arbeit, um ihn festzuhalten
zu jenem Punkte gelangen.
so
bedarf
es
weiterer
und durchzusetzen; so verwandelt der Verlauf des Lebens mehr
und mehr die Gabe des Schicksals in ein eignes Werk des
Menschen, so erhebt
sich
unser Leben
immer mehr zur
Selbst-
tätigkeit
Ähnliches
gilt
gebenem vorliegt,
von ganzen Völkern und
ist,
geistig angesehen,
Zeiten.
Was an Ge-
nur eine Möglichkeit,
zur Verwirklichung und zugleich zur konkreten Gestaltung
nur durch
unsere
eigne
Tat
gelangt.
die
immer
Wir können uns willenlos
24«
Zu den Problemen des Menschenlebens.
372
von dem, was wir empfangen und was uns umgibt,
treiben lassen
wir können ihm gegenüber eine Selbständigkeit gewinnen und von
aus erst ihm entringen, was an Geistigem in ihm angelegt
ihr
ist.
Die Geschichte baut sich ihrem Geistesgehalt nach nicht auf einer
und gesicherten Grundlage auf, sondern immer von
neuem mag über ihr Ganzes Zweifel entstehen, immer von neuem
gilt es, ihr eine Grundlage eret zu sichern, immer neu ist sie in ein
gegebenen
Ganzes zusammenzuschließen.
Solche Überzeugungen ergeben ein wesentlich anderes Bild der
Wirklichkeit, als es
dem Determinismus vorschwebt.
ihm
scheint bei
alles in einer einzigen
sein
ein
Fläche zu liegen oder doch aus
einem gegebenen Grundstock hervorzugehen,
Leben nicht so einfach und
Vor allem
Dem Determinismus
anderes Bild von unserem eignen Seelenleben.
Wahrheit
ist
unser
Inhalt nicht so gleichartig.
Ver-
in
schiedene Möglichkeiten und verschiedene Höhenlagen durchkreuzen
einander und ziehen bald hierher bald dorthin.
Eine von ihnen ist
durch den Verlauf des Lebens zur Herrschaft gelangt und
das Ganze
Lebens und Seins.
Aber
gilt leicht
es
brauchen nur
wesentlich neue Aufgaben, nur große Erschütterungen
und Umwälz-
als
unseres
ungen zu kommen, und
wartetes
uns
in
es
steigt
während
auf,
etwas
altes
völlig neues,
etwas uner-
und versinkt,
verblaßt
anderes in uns belebt, und es verändern sich
alle
es wird
Werte; es kann
uns unsäglich klein und nichtig erscheinen, was früher unsere Seele
erfüllte.
Das
an uns,
sondern durch unsere eigne Erregung und Bewegung hin-
alles
aber nicht
durch
daß
ein
was
mechanisches
Vorgehen
Ganze
durch.
Alsdann
dünkte,
nur eine gewisse Schicht, nur eine besondere Möglichkeit
war,
wird
deutlich,
uns
daß wir nur ein Stück unseres Wesens
schaftliche
Dasein und der
vorher
erlebten.
das
Das gesell-
Zwang der Lebenserhaltung drängen zu
einer Festlegung auf eine solche Besonderheit, hier gilt der Mensch
nur
als
Träger eines eigentümlichen Berufes, hier
darin aufgehen, hier wird
gezogene Linie
einhält.
ihm
alles verübelt,
soll
er möglichst
was nicht die dadurch
Wer sich aber ganz einer solchen Vereng-
nicht eine größere Weite mit offnen Mögdem muß es erstarren und verknöchern, der handelt
ung ergibt, seinem Leben
lichkeiten wahrt,
weniger
als
durch ihn gewirkt wird,
abgesteckte Chaussee,
verwandeln möchte.
in
die
der
geht
in
Wahrheit jene
der Determinismus unser aller Leben
Das ist der Segen des Leides und aller großen
Erschütterung, daß es den Menschen solcher Erstarrung zu entreißen
Freiheit des Willens.
373
Und darin hat
und ihm neue Lebensquellen zu eröffnen vermag.
auch die Kunst eine hervorragende Aufgabe, der üblichen Einpferch-
ung des Lebens den weiteren Kreis mit seinen offnen Möglichkeiten
entgegenzuhalten und dadurch der Bindung der Verhältnisse gegenüber zur Befreiung der Seele zu wirken. Immer ist dies der Scheidepunkt, ob unser Leben sich aus fertigen Daten zusammensetzt, oder
ob es noch mitten im Fluß
Was vom
Einzelnen,
ist.
das
von der Menschheit im
auch
gilt
Ganzen.
Wie jenen die Besonderheit seines Berufes und seines Ge-
schickes
festlegt
und
einspinnt,
so
ist
Menschheit eine
bei der
es
besondere Art der Kultur, auf welche die gewöhnliche Denkweise
sie
und
festlegt,
in
die
zur Verengung auch
man sie einspinnen möchte. Das aber bringt
Die Menschheit
eine Erstarrung des Lebens.
dann nur eine gewiesene Bahn zu verfolgen,
hätte
einer,
irgendwelcher
zu
bloßes Mittel
solchen Kultur fühlt, der
wer
Leistung;
mag glauben,
sich
würde ein
sie
Höhe
auf der
mit voller Sicherheit
nachweisen zu können, wie alles gekommen ist, wie alles so kommen
dem stellt sich die ganze Geschichte als eine Kette der Not-
mußte,
Aber auch Kulturen leben
wendigkeit dar.
altem; es wäre schlimm,
zum All
und
sie
von
dem
Alten
Altertum
ergreifen wie
die Gestaltung
welken und
durchlaufenen in sich
die
als
ausbilden
irgendwie
her
aus,
wenn die Menschheit in ihrem Verhältnis
nicht andere Möglichkeiten
trüge
sich
könnte.
ableitbar?
voraussehen,
irgend
Ist
nun dies Neue
Konnte das griechische
die
durch das
sich
Oder
Christentum und den Aufstieg neuer Völker vollzogen hat?
war vom
Neuzeit
Mittelalter
brachte?
modernen Kultur,
aus
Wendung
die
Und wenn
ein
wir
zu
jetzt
Greisenhaftwerden
welche
die
Schranken
der
erwarten,
innere
ihres
inneren
Bestandes
immer stärker empfinden, was anderes läßt uns freudig am Streben
und Wirken festhalten, als die Hoffnung, daß die Menschheit sich
auf den bisherigen Bahnen nicht schon ausgelebt hat, daß noch
Aber ohne unsere Selbsttätigoffne Möglichkeiten vor ihr liegen?
keit
werden solche Möglichkeiten
sich
schwerlich
beleben
lassen;
wir dürfen dabei nicht bloße Zuschauer sein, wir müssen Mitarbeiter
werden.
Sollte eine derartige
Denkweise mit ihrer Erweiterung und Be-
lebung des Wirklichkeitsbildes nicht auch für das Ganze des Weltalls
gelten?
Viel
zu
sehr neigen wir Modernen
und den Weltstand, der uns umfängt,
als die
dahin,
die
Welt
einzige Möglichkeit,
Zu den Problemen des Menschenlebens.
374
als
den Inbegriff
Wirklichen anzusehen.
alles
nur eine besondere
leicht
müssen?
ja bestehen
Ist
jene nicht
viel-
neben der andere bestehen können,
Art,
Als ein Anzeichen dessen könnten die mannig-
fachen Unfertigkeiten, Verwicklungen, Widersprüche gelten, in denen
Gemenge von Vernunft und UnverVon hier aus muß es als ein starrer und
unsere Welt sich befindet, das
das sie aufweist.
nunft,
einengender Dogmatismus erscheinen, alle Wirklichkeitsbildung an das
»Gegebene" zu binden.
und
licher
selbstverständlich gibt,
Ursprünglichkeit
ihrer
ein höchst unglück-
und da es alle Selbsttätigkeit
Eine
leugnet.
kaum,
empfindet heute
weise
ist
da es eine höchst problematische
irreleitender Begriff,
Behauptung als
mit
Dies «Gegebene"
eingeschüchterte
Herabsetzung
welche
der
Denk-
geistigen
»Der
Energie in der völligen Bindung an
die Gegebenheit
Geist geht auf alle Arten an die Kost
und schmiegt sich an das Ge-
gebene"
(J.
Wie
liegt.
Burckhardt).
sich
von unseren Überzeugungen aus das Problem der
und Gebundenheit des Handelns darstellt, das läßt sich
Freiheit
in
der hier gebotenen Kürze unmöglich erörtern, das hoffen wir bald
in einer Grundlegung der Ethik näher darlegen
es
zu können.
Hier galt
nur bemerklich zu machen, daß der Determinismus auf ganz be-
stimmten Voraussetzungen von der Wirklichkeit ruht, und daß die
Erkenntnis dessen seine Selbstverständlichkeit sofort zerstört.
handelt die Welt als gegeben
bloßes Stück dieser Welt;
als
ein
zu
einer
kaum
verständlichen
irgend zu zweifeln;
Er be-
und geschlossen und uns Menschen
hat er damit Recht,
Torheit,
an
seiner
so wird es
Wahrheit
noch
aber die Welt noch im Fluß und können
ist
wir selbst an vordringendem Schaffen teilnehmen,
so wird jemand,
der andere Wege versucht, nicht schon als ein intellektuell Verlorener
anzusehen
Schlimmsten Falles
sein.
freilich
müßte er sich dessen in
Gemeinschaft mit einem Plato und einem Kant vertrösten.
Die große Klärung und Förderung, welche der moderne Determinismus
dem Freiheitsprobleme gebracht hat, wird damit keines-
wegs verkannt, er hat das ganze Problem der Freiheit aufs wesentlichste
vertieft
und
Daß
viel
gemacht.
zum
guten Teil
nur das bleibt
auch
die
ihre
sammenstoß von
in
unserem Leben
das Schicksal es bereitet,
in Frage,
Freiheit
naive Bejahung schlechterdings unmöglich
Notwendigkeit
ein
Recht
Freiheit
ist
ob dieses das Ganze
behält,
und ob
waltet,
daß
nicht zu verkennen;
ist,
ob nicht zugleich
der Zu-
nicht eben
und Notwendigkeit unserem Leben seinen
Freiheit des Willens.
eigentümlichen
Sinne
erst
Charakter
möglich macht
verleiht,
ob
er
nicht
375
Leben
in
vollem
Wir stehen zu Schelling, wenn er sagt:
»Ohne den Widerspruch von Notwendigkeit und Freiheit würde nicht
Philosophie
allein,
Tod versinken."
sondern jedes höhere Wollen des Geistes
in
den
E.
Letzte Probleme.
1.
Der Wert des Lebens.
Einleitendes.
a)
D
ein
en Wert des Lebens nach der individuellen Empfindung mit
ihrer Zufälligkeit
und
ihren
Ding der Unmöglichkeit;
Schwankungen abzuschätzen,
hat die Frage
Pessimismus* keinen anderen Sinn, so
weisen.
Aber unmöglich
stellen,
schon deshalb
unwiderstehlicher
nicht,
sie
von vornherein abzuüber das Leben ein-
weil dies uns nicht mit schlichter
Tatsächlichkeit
ist
oder
Urteil
ist
läßt sich alles
des Optimismus
einnimmt,
sondern
weil
es
und
eine
Entscheidung unsererseits verlangt, diese aber entgegengesetzt auskann.
fallen
anschließen
aber wir
^
Denn entweder mögen wir uns seinem Strome freudig
und ihn möglichst durch unsere
oder
Kraft verstärken,
mögen uns ihm entgegenstemmen und
ihn bei uns
zum
Die Ausdrücke Optimismus und Pessimismus sind neueren Ursprungs,
zuerst zur Bezeichnung der Leibnizischen Lehre von der besten
Welt verwandt worden.
Brucker (IV, 2, S. 385) bemerkt darüber: Non
tacendum vero, ipsos Jesuitas Trivaltinos, magnos cetera Leibnizii admiratores,
jenes
ist
cum
recensione Theodiceae facta sententiam dicerent (wie eine
hinzufügt 1737, Febr.
art.
1),
Anmerkung
laudata ingenti lectionis et judicii copia,
tractationis ordine, accuratione et concinnitate systematica, fateri tamen,
errores
philosophum summum admisisse, maxime vero optimi mundi
tionem (optimismum vocant) non
nisi
larvatum materialismum
et
multos
asser-
et spiritualem
Spinozismum involvere, s. auch S. 415. Zu seiner Verbreitung hat wohl namentlich Voltaire mit seinem Candide ou l'optimisme beigetragen.
Bei Pessimismus
pflegen wir zuerst an Schopenhauer zu denken, aber dieser selbst gebraucht
das Wort nur spärlich; Caldwell in seinem vortrefflichen Werke über Schopenhauer bemerkt darüber (S. 522): He rarely uses the word pessimism
perhaps
three or four times at all — and than only about the philosophy of others,
and generally in the adjective form as opposed to an optimistic view of
—
—
things.
Der Wert des Lebens.
uns
stellen
bringen suchen.
Große
beide Antworten
deutlich
zu
Stillstand
377
geschichtliche Entwicklungen
Die Höhe des
vor Augen.
indischen Lebens war von der Überzeugung beseelt, daß das Leben
mit seiner endlosen Sorge,
Mühe und Not vornehmlich ein Leiden
und daß das Streben nach Befreiung von ihm oder doch nach
sei,
Herabsetzung
Summe
die
Stärke
seiner
Entgegen solcher Lebensverneinung
kreis
einer
Gut,
das
anderen Schätzung:
man
festhalten
eifrig
folgt
Lebensweisheit
unser westlicher Kultur-
zu erweisen.
dünkt
und
steigern möchte;
das
allumfassende
soweit das Problem
in
sie
Gerechtigkeit
und
sittliche
zu
Die griechischen
und Widersprüchen der Welt
erweisen;
beschäftigte,
Liebe
bemühen
diese als ein vollendetes Kunstwerk,
sie
Harmonie
der Wirklichkeit eine
hohes
ein
so
Die geschichtliche Bewegung nach
Dunkelheiten
überlegen zu werden, indem
eine
Leben
hier
dieser Richtung zerfällt aber in drei Hauptphasen.
Denker suchten den
bilde.
Bejahung zu begründen und die Wirk-
sich auch die Denker, jene
lichkeit als wertvoll
der
wie
die
z.
Ordnung,
voller
christlichen
Denker,
B. einen Augustin, sahen
von
die den Gegensatz
Ausgleichung
den
bringe;
modernen Denkern endlich ward die Welt ein vordringender Lebensstrom, ein unaufhörliches Anschwellen der Kraft; als ein Reiz und
Antrieb
als
dafür schien- sich auch
das zu rechtfertigen, was zunächst
Störung und Widerspruch erscheint
Diese Versuche der Denker sind oft hart angegriffen,
ja
arg ver-
spottet worden; sie möchten eine solche Behandlung verdienen,
wenn
nur spielende Reflexion sie hervorgebracht hätte, nicht tiefere Beweg-
ungen hinter ihnen stünden. Dieses aber war in der Tat der Fall. Denn
jene Versuche einer Rechtfertigung des Lebens wurzelten in einer tatsächlichen Gestaltung seiner, in einer Selbstkonzentration, die einen
Kern
vom übrigen Dasein schied und von ihm aus eine Weiterbildung des
So wären die griechischen Versuche, die Welt als
Kunstwerk darzustellen, ohne Saft und Kraft gewesen, hätte nicht
Ganzen erstrebte.
ein
die großartige künstlerisch-plastische Gestaltung des Lebens
ganzen Wirklichkeit, die wir an den Griechen bewundern,
tragen
und getrieben;
und der
sie
ge-
dies künstlerische Schaffen mit der ihm eignen
und Freude war es, was die griechische Welt gegen die Unvernunft des Daseins wappnete, die sie keineswegs gering anschlug, und
Kraft
was ihr die Zuversicht gab, dem Dunkel und Leid gewachsen zu sein.
Das Leben zerlegte sich damit in eine höhere und eine niedere Stufe, in
Gestalt
und Ungestalt; der Mensch aber vermochte sich auf die Seite
Letzte Probleme.
378
des
Höheren zu
und
stellen
seinem Bereiche dafür zu wirken.
in
An stärkster
Nicht anders stand es auf dem Boden des Christentums.
Empfindung des Bösen
wahrhaftig nicht,
hier
es
fehlte
aber das
Bewußtsein, ein unverlierbares Glied einer weltbeherrschenden
Ordnung zu
lichen
Zuversicht,
es
sein,
gab ihm
dem Menschen
verlieh
viel
zu tun,
es stählte ihn,
sitt-
Größe und
eine
den schweren
Kampf gegen die überwuchernde Unvernunft getrost zu beginnen.
Daß es sich in der Neuzeit ähnlich verhält, daß hinter dem Glauben
an das Vermögen der Entwicklung eine tatsächliche Steigerung des
Lebens,
eine
rastlose
Daseins
steht,
das haben wir
Arbeit
Verbesserung des
zur
menschlichen
vor Augen; ohne solche
alle deutlich
Erfahrung des Fortschritts hätte der Glaube an die Entwicklung die
Gemüter wohl nur flüchtig berührt.
So waren
waren
es durchgängig eigentümliche Lebensgestaltungen, es
Konzentrationen
des
Lebens bei
sich
selbst,
Tatsynthesen,
nicht bloße Begriffsklaubereien, Tatsynthesen, Lebensenergien, welche
dem Menschen die Überzeugung einflößten, den Gründen der Wirklichkeit
ihn
verbunden zu sein und aus ihnen Kraft zu empfangen, die
dem
aus
in
den
und ihn zugleich mit freudigem Mute
er-
Stande eines bloßen
der Aktivität versetzten
Übersichergehenlassens
Die Unvernunft verschwand damit keineswegs,
füllten.
mochte
sie
eher
noch gesteigert scheinen.
nicht
mehr einsam und wehrlos gegenüber, nun durfte er mitarbeiten
Aber nun
stand
der Mensch ihr
am Bau der Wirklichkeit, nun hatte sein Leben einen Sinn und zugleich einen Wert gewonnen.
Wer solche Zusammenhänge vor
Augen
hat,
schätzig
der
wird jene Versuche
ansehen,
wie
führung finden mag.
unzulänglich
Denker minder gering-
der
er
das
Nähere
ihrer
Beweis-
Aus Beweisen hat das Leben nie seine Kraft
gezogen.
b)
Die Verwicklung der Gegenwart.
Heute ergeht
es
uns bei diesem Problem wie bei vielen an-
deren; aus vermeintlichem Besitz sind wir wieder in ein tastendes
Suchen geworfen.
Alle jene
Lebenskonzentrationen sind
in
ihrem
Bestände und in ihrer Herrschaft aufs schwerste erschüttert: von der
künstlerischen
und der
ethischen
ist
es augenscheinlich,
der Fortschrittsgedanke hat nicht mehr die
Zauber; wir wissen nicht
recht,
alte Kraft
aber auch
und den alten
wohin uns der Fortschritt führt und
Der Wert des Lebens.
wem er zugute kommt,
tieferen
379
auch seine Tatsächlichkeit
ist
uns bei den
Lebensfragen stark ins Wanken geraten; ein hohles Phrasen-
tum umspinnt hier oft den Kern der Behauptung und zieht das
Ganze herab. Fehlt demnach dem heutigen Leben überhaupt ein
fester Zusammenschluß und mit ihm ein allbeherrschendes Ziel, so
zugleich
fehlt
dem Zustrom
die Kraft,
der Wirklichkeit gewachsen
zu werden, ihn innerlich zu bewältigen und in solcher Verwandlung
des
das Bewußtsein
Daseins in Aktivität
Dieser Mangel
wegung der Neuzeit
sie
läßt,
Größe zu
einer
finden.
muß um so empfindlicher werden, als eben die Bedie
Außenwelt aufs gewaltigste anschwellen
uns unvergleichlich größer vor Augen
und
stellt
stärker
Indem so immer mehr die
auf uns eindringen läßt als je zuvor.
Welt uns besiegt und uns zu ihrem bloßen
Anhange
herabsetzt,
verfallen wir bei aller Ausdehnung des Lebens einer inneren Schwäche
und Kleinmütigkeit; immer mehr
greift
die
Neigung
um sich, in
den mannigfachen Erfahrungen vor allem das zu sehen, was den
Menschen
als
klein
durchgängig scheint
ziehe,
als
Verhältnis
darstellt
es,
als
und
ihm
alle
Besonderheit
raubt;
ob sich mehr ein Schicksal an uns voll-
daß wir zu Herren der Dinge werden und ein inneres
zur Wirklichkeit
Außenwelt größer geworden
setzen haben,
ist
es,
ausbilden
ist,
als
können.
Weniger daß
die
daß wir ihr nichts entgegenzu-
was unseren Lebensmut niederdrückt und uns
am Inhalt der Wirklichkeit vor allem die Verneinung gewahren läßt
So erfahren wir
es zunächst
im Verhältnis zur großen Natur.
Auge fällt uns vor allem ihre Unermeßlichkeit, ihre UnendlichRaum und Zeit, ihre Grenzenlosigkeit im Großen wie auch
im Kleinen. Dieser Gedanke der Unendlichkeit hat frühere Zeiten
freudig ergriffen und innerlich erhöht, indem er ihnen ein AusIns
keit in
druck
der
unbegrenzten Lebensfülle
fanden
sie
darin einen besonderen
der Wirklichkeit war;i
auch
Vorzug des Menschen, daß sein
Denken ihn über alle Begrenzung hinaus zum Unendlichen und
Ewigen führe und ihn daran teilnehmen lasse. Heute aber denken
wir weniger an die innere Gegenwart der Unendlichkeit,
sie ausgebreitet in
als
daß
Raum und Zeit um uns liegt und zugleich unser
Zu voller
ganzes Dasein zu verschwindender Kleinheit herabsetzt
Gleichgültigkeit scheint
^
es
damit zu sinken.
Denn was
Die klassische Zeit des Altertums scheut das Unendliche
bedeutet.
als
etwas
Grenzenloses und damit einer künstlerischen Gestaltung Entzogenes, zu positiver
Schätzung hat den Begriff erst Plotin gebracht.
Letzte Probleme.
380
so hören wir,
was auf dieser winzigen Erde vorgeht, gegen-
alles,
über der unermeßlichen Fülle der Welten, die das geschärfte Auge
der Neuzeit uns wahrnehmen
das Maß,
ein anderes
Die äußere Größe gibt hier
läßt?
— Dazu
kennen wir nicht
Natur
bleibt die
uns innerlich fremd und verschlossen, sie scheint, indem die Forsch-
ung ihr immer mehr Boden abgewinnt, ihrem Grunde nach ferner
und ferner zu rücken. Denn es fehlt uns ein inneres Verhältnis
zu
wie es früheren Zeiten eine religiöse oder eine künstlerische
ihr,
Überzeugung gab.
Uns erfüHt ganz und gar der Gedanke der Be-
grenztheit des Menschen, wir scheinen einem besonderen Kreise an-
zugehören und ihn
wir
keiner Weise überschreiten zu können.
in
den
von
derart
Zusammenhängen
großen
Sind
abgeschnitten,
so
wird es zu einem vermessenen Unterfangen, zu einem eingebildeten
Anthropomorphismus,
Natur
die
deuten und verstehen zu wollen.
sie
ganz
ein
nis,
tätigen
irgendwie
tiefes
Geheimerzeugt
wir uns nur durch die Analogie eines
zweck-
Sie bleibt
Rätsel.
In
uns ein
Handelns irgend vorstellbar machen können, aber wenn sich
von Zwecken
überhaupt
einander
dem
die
Gestalten
unermeßlicher Fülle
unlösbares
Bildungen,
ihrem
mit
der Natur reden
widersprechen und sich
zu
nämlich
die
Natur
mit
so scheinen diese
läßt,
gegenseitig
aufzuheben.
In-
bewunderungswürdiger Sorgfalt eine
Art von Wesen bereitet, bereitet
sie
mit gleicher Sorgfalt eine andere,
was
die jene zu zerstören vermag, so scheint sie hier zu verneinen,
sie dort bejaht,
treibt sie
Individuen,
oft
Geschöpfe gegeneinander und
aufsteigender
mühsamen Umwegen,
auf
massenhaft werden
zierter
hetzt ihre eignen
sie
einen unerbittlichen Daseinskampf.
in
Zug
Inmitten alles
aufgeopfert.
sie
Massenhaft werden
gebildet,
des Lebens nicht zu
verkennen,
ebenso
aber
Kampfes ist
ein
immer kompli-
wird der Bau der Organismen, immer abgewogener die Diffe-
renzierung der Teile,
immer größer
die
seelische Leistung.
Aber
dem eignen Gebiet der Natur sehen wir keinen rechten Gewinn
dieses Forttriebs.
Denn wenn auch die höchste Stufe über die
Lebenserhaltung Im Kampf ums Dasein nicht hinauskommt, so wird
im Grunde nur dasselbe Ziel, was auch schon die niedere Stufe
auf
hat,
in
weit umständlicherer Weise erreicht;
Rückschritt als
ein
Fortschritt
zu nennen?
nach Leben und die völlige Leere des
bilden den härtesten Widerspruch.
viduen
an
das Dasein,
zu
seiner
ist
das nicht eher ein
Der ungeheure Drang
mühsam errungenen Lebens
Gierig klammern sich die IndiFesthaltung
bieten
sie
höchste
Der Wert des Lebens.
Kraft
und Leidenschaft auf.
wesen
selbst,
damit, welchen Sinn hat dies ganze
sie
zu walten, aber
sie scheint
unablässig
selbst
sobald
Irgendwelche Vernunft scheint
gehemmt und gebunden,
zu durchkreuzen,
sie
ja sie scheint
scheint in eine Breite
Dazu
die sich innerlich nicht zusammenfindet.
aufgelöst,
wir keine
weil
wir uns verwirrt und niedergeschlagen,
wir die Frage aufs Ganze richten.^
sich
und
darauf keine Antwort,
finden
fühlen
so
finden,
Aber was liefert dieses Dasein den Lebe-
was gewinnen
Wir
Getriebe?
381
zeigt die
Entwicklungslehre uns diesem dunklen Getriebe weit enger verkettet,
als
gegenwärtig war, immer enger
der Vorstellung früherer Zeiten
ziehen sich die Fäden zwischen uns und
nur
nicht
körperlich,
auch
dem was unter uns liegt;
scheinen
seelisch
ganz dieselben
uns
Kräfte zu beherrschen, die dort das Leben bestimmen; so wird das
Dunkel der Welt auch zum Dunkel für unser eignes Leben, eine
Notwendigkeit drängt und
nunft dient,
Nun
ist
treibt uns,
aber wie weit
sie einer
Ver-
nicht zu ersehen.
verbleibt
dem Menschen
die
Wendung von
der Natur
zur Kultur, er kann sich ein eignes Reich aufbauen und in ihm sich
eine Größe,
sowie seinem Leben einen Wert bereiten.
hier sieht heute,
als
wer die Sache
reinen Gewinn.
befriedigt
Aber auch
Ganze faßt, mehr Verwicklung
ins
Daß die Kultur den Einzelnen nicht unmittelbar
und glücklich macht, daran zweifeln wir heute
nicht;
so
muß sie ihm etwas über das Glück hinaus gewähren; was aber dies
sei,
das wissen wir nicht.
Gewiß wächst unaufhörlich unsere Macht
über die Umgebung, sicherlich verbessern sich unablässig die Be-
dingungen unseres Daseins, Schmerz und Not werden siegreich bekämpft, Genüsse
gewinnt
unser
eröffnen
Leben.
in
reichster Fülle,
alles
zusammen
sich
Aber
selbst
gibt
an Dauer
ihm als einem
Ganzen noch keinen Gehalt und Sinn; nach einem solchen zu fragen,
kann aber ein denkendes und überschauendes Wesen nicht
lassen.
Größe der Leistung jene Konzentration des Lebens bei sich selbst, die, wie wir sahen, dem Menschen
einen festen Halt und das Bewußtsein eines inneren Zusammenhanges mit dem Ganzen der Wirklichkeit gab, die ihm das Leben
Damit
zu einer großen und aussichtsreichen Aufgabe gestaltete.
Es fehlt unserer Kultur
bei aller
aber fehlt uns die Möglichkeit, uns über die Verwicklungen hinaus-
^
So mag sich wohl bei Erwägung des Ganzen das aristotelische Wort
aufdrängen: r «püm? SatjjLovia, aXX' ou
ö-£(«
(463b, 14).
Letzte Probleme.
382
zuheben und ihnen
Komplexe
zu begegnen, die jeder Kultur anhaften,
kräftig
moderne Kultur in besonderem Maße zeigt
die aber unsere
verbinden und verweben
die Kräfte
entstehen,
Große
mit
sich,
solchem Zusammenschluß befreit sich die Arbeit von der Zufälligkeit
der bloßen Individuen und gewinnt eine Selbständigkeit; kraft solcher
kann
Wege
eigne
sie
Aber zugleich
und glänzende Triumphe feiern.
der Einzelne immer mehr zu einem bloßen
sinkt
verfolgen
und Werkzeug, und
Mittel
mehr
je
er
das
tut,
mehr wird
desto
das Ganze der Kultur seiner Seele fremd, desto weniger kann
im Wirken
für
ein
sie
nach
Betriebsamkeit
größter
Selbst
geistiges
außen
behaupten.
mit
hin,
atemloser
Lebens eine innere Gleichgültigkeit verbunden sein und
die
und Freude
rechte Kraft
fehlen.
es
doch
Hast des
dem Leben
in lauter ein-
Erscheinungen aufgelöst und beinahe sich selbst entfremdet.
zelne
Auch das
schwer zu vermeiden,
ist
herrschende und bewegende Seele
alle
Wird
er
So kann mit
Kulturentwicklung
stark zur
daß,
fehlt,
wo der Kultur eine be-
das Kleinmenschliche, das
besonders aufwuchert und besonders
begleitet,
Empfindung kommt: die Verquickung
alles
Strebens mit
kleinmenschlichen Zwecken, die durchgängige Unwahrhaftigkeit des
gewöhnlichen Betriebes,
der
hohe
Ziele verkündet,
zugleich
aber
Wohl verfolgen lehrt;
dazu die vielfache Eitelkeit, die allen Erfolg zum Preise des kleinen
die
Handelnden
Ich
verkehrt;
und
es
erster
in
Linie
ihr
eignes
das braucht nur ins Ganze gefaßt zu werden,
alles
kann uns eine starke Unlust an diesem Ganzen befallen;
wir haben das Gefühl, hier Mächten gegenüberzustehen, denen wir
das Feld unmöglich überlassen dürfen, und denen gegenüber wir
doch nichts rechtes vermögen.
zeitweilig eine Hülfe
Auch der Fortschrittsgedanke, der
zu bieten schien, verblaßt uns bei diesen Pro-
blemen mehr und mehr; denn augenscheinlich reicht der Fortschritt
nicht
in
diese
elementaren Verhältnisse
hinein;
allem
Aufstreben
scheinen hier natürliche Triebe und Leidenschaften eine Grenze zu
setzen, die wir
hören
als
So
läßt
können.
Gegenwart
Lebens
doch
Schranke peinlich zu empfinden nicht aufsich
in der Kultur einen
findet,
nicht
sagen,
daß
der
Mensch der
genügenden Sinn und Wert seines
und daß ihn die Arbeit für sie über die Zweifel und
Nöte des Daseins sicher hinaushebt
Die
Strebens.
Kultur
In
bildet
nicht
die
äußerste
Grenze
menschlichen
kühnem Aufschwung kann der Mensch sich über ihr
ganzes Gebiet erheben, sich auf seine eigne Innerlichkeit stellen, in
Der Wert des Lebens.
383
Ausbildung weltumspannender Persönlichkeit jenem ganzen Gewirr
und Schein überlegen werden, von dort aus ein unmittelbares Verhältnis
Das haben mit bewußtem Streben zuerst die
zur Wirklichkeit suchen.
Stoiker unternommen, von ihnen zieht diese Denkweise sich als ein
bleibender Typus durch die weitere Geschichte,
in
der Aufklärungszeit
auf
stark
Weise sucht die Religion durch
nächsten
Daseins
ob
dabei
ziehung
die
in
Weg zu
fallen,
nicht
erörtern,
und Zurück-
ob uns heute dieser
wir fragen nur,
hat,
Jene Ablösung von der sichtbaren
ist
Welt und vom Kreise des Menschen
zu
hier
der Welt
mit ihrer Spaltung des Daseins nicht
Seele
möglich
betreten
anderer
Verwicklungen des
alle
Wir wollen
hinauszuheben.
eigne
In
die Entwicklung eines unmittelbaren
empfohlene Ablösung von
Gefahren und Schranken
hat besonders
sie
Seelen gewirkt.
Gott den Menschen über
Verhältnisses zu
die
die
um nicht ins Leere
verlangt,
den sicheren Besitz einer Innenwelt, und ein solcher kann
nur aus einem unmittelbaren Verhältnis zu einer überlegenen Macht,
möge sie als Gottheit, möge sie als Weltvemunft vorgestellt werden,
Dem modernen Menschen aber ist alle Wirklichkeit einer
hervorgehen.
solchen Macht unsicher geworden.
So hat bei ihm auch eine selb-
und
er
verliert
damit
die Möglichkeit eines Selbständigwerdens gegen
die
sichtbare
Welt
ständige
Innenwelt
keinen
festen
und das menschliche Getriebe.
keit
wird
mit
solchem
Grund,
Das Sichgroßfühlen der Persönlich-
Schwinden des Grundes zu einem
eitlen
Dünkel und zu einer leeren Phrase; was hat denn der Mensch ohne
ein
Reich selbstwüchsiger Innerlichkeit der Welt entgegenzusetzen,
die mit überwältigender Macht ihn umklammert?
Art der modernen Kultur steigert
Ja die besondere
noch das Gefühl der Abhängig-
Maße
immer mehr auf ein Wirken und
Schaffen in Reih und Glied verweist Wo aber das Vermögen des
Einzelnen als engbegrenzt und gering erscheint, da muß zugleich
keit,
indem
an seine
sie
der Antrieb
zu
ihn
eignem Beginnen sinken;
Seelenstande unsere eigne
als
in seiner Arbeit in wachsendem
den Menschen
Umgebung bindet,
Meinung uns
dazu
oft
bei
stellt
mattem
noch abhängiger
dar,
wir es in Wahrheit sind, wir neigen dahin, überall Anschluß an
andere zu suchen, unser Wollen dadurch bestätigen zu
lassen,
nur
das gibt uns ein Gefühl der Sicherheit; auch erwarten wir vieles
vom Vermögen gemeinsamer
bei
der
Einrichtungen,
Gesinnung des Einzelnen
liegt,
wo
kurz
Energie des Lebens ohne zwingende Not herab.
die
wir
Ist
Hauptsache
drücken
die
bei Überwiegen
Letzte Probleme.
384
solcher Gesinnung
hoffen ?i
So
sichtiges
ist
von
einem Aufruf zur Selbständigkeit
der Anblick des Ganzen wenig erfreulich.
viel
zu
Ein undurch-
Reich der Natur umschließt und beherrscht uns, ein Reich
der Kultur strebt darüber hinaus, aber es
seelenloser Arbeit
daher unser Glücksverlangen
Kraft möchte
unter den Gegensatz
fällt
und kleinmenschlicher Subjektivität und
nicht,
befriedigt
gesteigerte Anspannung geistiger
uns darüber erheben und uns einen sicheren Grund
im eignen Wesen geben, aber dem Wunsche entspricht nicht das
Vermögen, und der Versuch der Befreiung
läßt
noch
Mühe und
stärker
empfinden.
wandelt sich uns nicht
in
Sinnes zu entbehren; was
Die
unsägliche
einen
reinen
Ertrag,
unsere Bindung nur
Arbeit
ver-
scheint alles
ja sie
der Erfahrung des Lebens besonders
in
schmerzlich berührt: innerlich Überlegenes von Niederem abhängig
zu sehen, das scheint die Gesamtgestaltung unseres Daseins uns auf-
Denn wohl
zuerlegen.
regt
sich
in
ihm aufsteigende
Leben dringt vor und neue Ausblicke tun sich auf.
und Aufstrebende gelangt
nicht
das
Kraft,
Aber das Neue
zur Selbständigkeit,
es
bleibt
an
eben das gebunden, worüber es hinaus will, oft wird es dadurch zurück-
gezogen und
in
seinem Wirken gelähmt.
bei Erfahrung dessen,
lich
gilt,
ein trüber
eben den
tieferen
Pessimismus
Lebensbejahung reden,
*
Ist
Wunder, wenn
es ein
namentlich v/enn diese Lage als unveränderSeelen
der Lebensmut schwindet,
und
um sich greift? Wir hören ja heute viel von
Hymnen
ja
auf das Leben singen, aber es
Jenen Mangel an Selbstvertrauen, jenes Hoffen und Karren auf andere
schildert vorirefflich ein Artikel im Spectator über English Pessimism (11. Aug.
1906); es heißt dort S. 190:
we
try to correct
If
we were to
suggest the spirit which,
our pessimism, would be most efficacious;
increase in individual self-reliance.
it
when
would be an
We are not beaten in public affairs as
we imagine we are, and there is no necessity in carrying out our works of
philanthropy for relying so entirely upon associations. We establish far too
many societies. Everybody seems too feel that before he can do anything
he needs the protection of a crowd. He cannot even denounce or defend
motor-cars unless hundreds will join him to protect him from the consequences of thinking independently. The result is that every one who wants
to
do something good devotes to it some fraction of
Chip of his energy, and that the slrength which
his
mind, some little
derive from the
we would
We develop some new
is seldom or nfever present.
and small group, not a Loyola or a Wesley. This, always the danger of
democracy, is the danger also of the mental processes of our time, and deprives us first and foremost of all help from mdividual genius.
strong will of a leader
Der Wert des Lebens.
gehört das zu den vielen,
385
innerlich hohlen Schaustücken der Zeit-
oberfläche, es ist das eine erkünstelte Lebensbejahung, die dem Grunde
der Seele fremd
bleibt, es
ist
wie ein Rausch, der ein unbefriedigen-
des Dasein zeitweilig vergessen machen
soll.
Ein derartiges Schein-
wesen kann dem Pessimismus kaum etwas anhaben.
Aber jede eingehendere Erwägung des Pessimismus zeigt, daß
daß sich daher unmöglich
er in sich selbst einen Widerspruch trägt,
Ein wahrhaftiger Schmerz kann nur ent-
mit ihm abschließen läßt
wo es etwas wertvolles zu verlieren gibt; wo alles nichtig und
stehen,
gleichgültig wäre, da könnte auch die Versagung oder der Verlust in
keiner Weise
erregen.
Christentum
alte
Das ausgehende Altertum und mit ihm das
verfochten
die
wie
z.
sie
schöpften
B. erblinden
daß das Böse keine
Lehre,
ständige Wirklichkeit, sondern nur die
selb-
Wegnahme eines Guten sei,
nur könne, wer von Natur das Augenlicht besitzt;
daraus die
Überzeugung von einem sicheren Über-
freilich über die Klippe
ist nun
hinwegzukommen, das Böse bedeutet wohl mehr als einen
bloßen Mangel. Aber richtig ist, daß eine Empfindung, und gar
eine starke Empfindung des Bösen ohne irgendwelches Gegengewicht
wiegen
des Guten. ^
So einfach
nicht
schlechterdings undenkbar wäre.
»Wer anders findet sich
darüber, nicht König zu sein,
lich
als ein
meint Pascal gewiß mit Recht; würden
gänglichkeit der
z.
unglück-
entthronter König", so
B. die Menschen die Ver-
Dinge und den raschen Flug des Lebens so un-
ablässig beklagen, wenn sie bloße Eintagsfliegen wären, nicht in ihnen
irgend etwas wirkte, was in sich die Forderung ewiger Dauer trägt?
So
ist
inmitten aller Bedrängnisse der Zeit die tiefe Empfindung
dieser Bedrängnisse ein vollgültiges Zeugnis dafür,
in
jene Lage
enthält,
nicht ganz
und gar
was ihr Widerstand
Befreiung
von
leistet.
dem bloßen
aufgeht,
sein
Wesen etwas
Könnten wir so sehnlich eine
Kulturgetriebe
etwas an uns ihm überlegen wäre?
daß
daß der Mensch
erstreben,
Könnte uns
in
wenn
nicht
der Kultur der
Mangel innerer Zusammenhänge und reiner Sachlichkeit so sehr
schmerzen, wenn unsere Natur nicht solche verlangte?
Könnte das
tiefe
Dunkel der Welt unserem Leben zur Einengung werden, wenn
wir
nicht
müßten?
*
auf
irgendwelchem
Unmittelbar
inneren
Verhältnis
zu ihr bestehen
mag uns das alles an Positivem kaum etwas
Mit besonderem Nachdruck hat Augustin jene Lehre vertreten, nament-
dem enchiridion ad Laurentium de fide, spe et caritate. Nach seiner
Bezeichnung ist das Böse nicht causa efficiens, sondern nur causa deficiens.
Eucken, Grundbeg^riffe. 4. Aufl.
25
lich in
Letzte Probleme.
386
wohl aber überzeugt
liefern,
daß die
es uns,
glatte
Verneinung die
Sache nicht erschöpft, daß jenseit ihrer manches zu fragen
bleibt.
wenn wir
uns das Ganze des gegenwärtigen Lebens vor Augen stellen. Auch
Aber wir dürfen noch einen
wie es vorliegt, geht
auf in das Bild,
nicht
es
Schritt weiter
grenzenlosen Ausdehnung des modernen
Lebens
gehen,
das eine von der
streben,
enthält
herrscht.
Ein
mehr,
als
Ge-
überwältigte
sinnung von ihm entwirft; schon das, was wir haben, nicht
erst er-
den Blick und die Schätzung be-
dort
Leben aus weltumspannender Persönlichkeit
uns
ist
frommer und matter Wunsch; wohl können wir es
erreichen, aber wir bemühen uns eifrig
darum, wir suchen auf, was solches Streben verstärken könnte, wir
möchten im besondern die großen Persönlichkeiten, welche das geschichtliche Leben bietet, uns näher rücken und mit ihnen das eigne
Leben verknüpfen; mag das alles unvollkommen und unfertig sein,
mehr als
ein
Einem Schlage
nicht
mit
eine
Bewegung zu jenem Ziele ist zweifellos im Gange.
Noch deutlicher sind die Schranken der pessimistischen Be-
trachtung gegenüber der Kultur.
Es
ist
nicht richtig,
daß uns heute
nur der Mechanismus der Arbeit verbindet und uns ganz
zu
Stücken seines Räderwerkes macht;
Streites
und Werten
halten
wir besitzen inmitten
auch eine gemeinsame Gedankenwelt
wäre überhaupt nicht zu
— ohne
alles
eine solche
— uns umfängt mit geistigen
In-
gemeinsame Atmosphäre, wir sehen
bei
streiten
eine
und gar
,
näherer Betrachtung dessen eine innere Weiterbildung und Erhöhung
des Menschen durch die Kultur;
wir überzeugen uns, daß
eine neue Stufe der Wirklichkeit aufsteigt,
in
ihr
daß hier die Welt aus
einem bloßen Neben- und Gegeneinander ein innerer Zusammenhang
wird, und daß, was hier geschieht, über die Zwecke
Menschen weit
hinausreicht.
So
läßt
sich
des bloßen
auch die Leistung der
Gegenwart unmöglich verstehen vom kleinen
Ich des Menschen her.
Das gewaltige Vordringen der Wissenschaft und das unermüdliche
Gestalten
der Kunst
ist
nur begreiflich
als
das
Werk von inneren
Notwendigkeiten, die den Menschen packen und ins Schaffen treiben;
mag kleinmenschliche Art dies Schaffen noch so umspinnen, seine
Überlegenheit wird dadurch
es mit
dem
praktischen
nicht
gebrochen.
Wirken der Gegenwart.
Nicht anders steht
Eine
Zeit,
die an
Erweisung humaner Gesinnung alle früheren Zeiten weit überflügelt,
die zugleich das Recht jedes Einzelnen auf Entwicklung seiner geistigen
Kraft
und auf Teilnahme an den Lebensgütern bereitwillig anerkennt,
Der Wert des Lebens.
387
Macht über die Gemüter gfewährt,
die der sozialen Idee so viel
Wir sehen dieses Große nicht klar, weil wir
Egoismus beherrscht.
die
Erscheinungen
einzelnen
fassen,
aber es
nicht
genug
kräftig
Ganzes
ein
in
wir müssen es gewahren, sobald uns durch
da,
ist
ist
vom bloßen
keineswegs ganz und gar odec auch nur überwiegend
die trübe Oberfläche des Alltags hindurch die richtenden Grundlinien
hervorscheinen.
Die Anerkennung einer in der Menschheit aufsteigenden Geisteswelt verändert aber auch
den Gesamtanblick des Alls und unsere
Aufgabe ihm gegenüber.
Nun bildet die Natur nicht mehr das Ganze
Denn dar-
der Wirklichkeit, nun gewinnt diese eine größere Tiefe.
über kann kein Zweifel sein, daß, wenn eine solche
Wendung zur
Ganze von Haus aus mehr sein muß, als
es dem ersten Anblick erscheint. Auch die Entwicklung nimmt sich
Innerlichkeit statthat, das
anders
wenn
aus,
Natur ist, sondern
das
Geistesleben
Erzeugnis
kein
bloßen
der
aus der Natur nur hervorgehen kann, weil
es
diese eine tiefere Wirklichkeit hinter sich hat; die engere Verbindung
mit der Natur wird dann
des Menschen
nicht
sowohl ihn herab-
Wandlung der Grund-
als
die
Natur erhöhen.
anschauung
muß
auch der Arbeit neue Aufgaben
Mensch mit
seiner Geistigkeit
drücken
beschränktes Einzelwesen,
tritt
auch
Solche
kein
sondern wirkt ein Weltleben
Erkenntnisstreben
sein
stellen.
auf einen
bloßes,
günstigere
unter
Ist
der
Sonderkreis
in
ihm, so
Bedingungen.
Denn nun läßt sich fragen, ob sich bei ihm nicht zwischen Echtgeistigem und Kleinmenschlichem scheiden läßt, und ob sich von
jenem
aus
Brücke finden mag, die uns enger mit der
nicht eine
sie uns mehr zur Heimat macht.
Doch solche Gedankengänge lassen sich hier nicht weiter ver-
Welt verbindet und
folgen.
Hier
galt
es
nur
festzustellen,
daß
das
des
Weltbild
Pessimismus nicht das Ganze der Wirklichkeit aufnimmt, sondern
nur eine gewisse Durchsicht
lage
entspricht,
bindet.
Es
einer
Seelenlage,
Freilich
ist
die einer eigentümlichen Seelen-
die
uns
keineswegs
endgültig
mehr in unserer Wirklichkeit, als der Durch-
steckt weit
schnitt des Zeitlebens
bietet,
uns sehen
nicht
zu
läßt.
bestreiten,
daß
solches
sammengefaßt und vollauf angeeignet werden muß,
ständen gewachsen zu sein;
Mehr
um
erst
zu-
den Wider-
jenes aber kann nur geschehen,
wenn
es gelingt, wieder zu einer Selbstkonzentration des Lebens, damit zu
einem ausgeprägteren Charakter
und zugleich zu einem
25'
aktiveren
Letzte Probleme.
388
Verhalten zur Wirklichkeit zu gelangen.*
Mit
steckt in der Wirklichkeit;
wir können
Wendung
einer
bekannten Dürerschen Wortes könnten wir sagen:
des
«Die Vernunft
wer sie herausreißt, der hat sie."
Aber
sie nicht herausreißen, bevor sich uns selbst das Leben
zusammenschließt
und
sich
damit unsere
Organe
inneren
weiter-
bilden.
Es kann sich uns aber unmöglich das Positive
in
Leben und
Wirklichkeit zusammenschließen, ohne daß sich uns das Dasein zerlegt,
und Dunkel sich
Licht
des Menschen
wie der
schärfer scheiden, sich das ganze Leben
Menschheit
in
eine
durchgehende Aufgabe
So verschwindet mit dieser Wendung die Unvernunft
verwandelt.
keineswegs, wohl aber gewinnen wir die Möglichkeit,
zu
entrinnen.
Woher
der
ziehung des Höheren
gültigkeit
und
werden
überlegen
ihrem
damit
Widerstand
lähmenden
was
das,
woher
stamme,
zum Niedern, woher
des Weltlaufs gegen
ihr
innerlich
Druck
die
zu
Herab-
die scheinbare Gleich-
er
selbst
als Ziel
hervor-
zubringen scheint, das können wir Menschen unmöglich enträtseln;
die Philosophie wie die Religion haben das
Problem beim Versuch
einer Lösung nur immer weiter verwickelt.
So muß uns genügen und
kann uns genügen, daß bedeutendes
uns vorgeht, und daß wir
nicht
als
tatenlose
Zuschauer
lassen müssen, sondern
stellen
und zu
kommt
das
ihrer
das
bei
Weltgeschick über uns ergehen
daß wir uns auf die
Seite der Vernunft
Förderung zu wirken vermögen.
zu
Damit be-
Wort von Vauvenargues ein gewisses Recht: «Die
Welt ist, was sie für ein tätiges Wesen sein muß, voll von Hindernissen." Je mehr wir uns wieder einer vollen Lebenssynthese nähern,
desto mehr läßt sich auch der Lebensmut wiedergewinnen, desto mehr
wird uns das innere Gefüge des Lebens selbst einen sicheren Halt
gegen die Unvernunft des Daseins gewähren.
Wenn so unsere Zeit mit der Ergreifung tieferer Zusammenhänge der Wirklichkeit wieder zu
einer positiven
Schätzung
des
Daseins zurückkehren muß,
so bleibt dabei aller Optimismus, alle
Abschwächung des Dunkels
fern;
wir müssen im besondern
eine
nicht geringe Differenz zwischen der hier gewonnenen Lebensstimmung
und derjenigen anerkennen, welche
Literatur beherrschte.
^
die
Höhe
unserer klassischen
Dort schien die Welt ein Reich ungetrübter
Daß der von uns geforderte Aktivismus keineswegs mit der Wendung
zur praktischen Vernunft oder gar zur moralischen Betätigung zusammenfällt,
kam schon oben zur Sprache.
Der Wert des Lebens.
339
Vernunft zu bilden, der Mensch aber die Höhe seines Lebens in der
künstlerischen
Anschauung oder im denkenden Begreifen der Welt-
harmonie zu finden; die Hauptaufgabe der Menschheit war
voller Bewußtheit zu bringen,
allen Seiten
umfängt.
Uns Neueren haben sich
im menschlichen Leben die Probleme
daß wir so rasch
heraustreten dürften.
hier,
viel
in
der Natur wie
zu sehr verschärft,
mehr
als
wagen und aus dem Kampf
Aber wenn wir mit solcher Erschwerung des
einen
Abschluß
Daseins vieles verloren haben, wir haben eines gewonnen, das
Verluste
zu
was uns mit unbewußtem Wirken von
als aufwiegt:
wir
alle
dürfen selbst zur Förderung des
Ganzen wirken, weit mehr sind wir aus Zuschauern zu Mitarbeitern
am großen All geworden.
Das Problem der Religion.
2.
(Immanenz
— Transzendenz.)
Immanenz und Transzen-
|ie Behandlung des Gegensatzes von
I
*-^ denz könnte zur Aufrollung des gesamten religiösen Problemes
um so weniger, als wir uns über dies neuerdings
führen; sie soll es
wiederholt geäußert haben ;i so
tümliche Verhalten
hier
sei
nur das der Gegenwart eigen-
diesem Problem
zu
Die
jetzt
Auch
übliche Zusammenstellung von immanent und transBis dahin
zendent reicht nicht hinter Kant zurück.^
ander immanens
13.
(auch
sofern
sie
transeunt,
sofern
sie
das berühmte
^
S.
standen ein-
permanens) und transiens gegenüber;
Jahrhundert nannte
immanent,
ist
erörtert.
Zur Geschichte der Ausdrücke.
a)
dem
Kürze
in
mag das Wort uns zur Sache leiten.
man
seit
eine Handlung oder eine Ursache
des wirkenden Subjekts verbleibt,
innerhalb
darüber hinaus auf etwas anderes geht.^
So
Wort Spinozas zu verstehen, daß Gott die imma-
„Der Wahrheitsgehalt der Religion", 2. Aufl. 1905, »Hauptprobleme
der Religionsphilosophie der Gegenwart", 1907.
S. z. B. III, 245 (Hart.): „Wir wollen die Grundsätze, deren Anwendung sich ganz und gar in den Schranken möglicher Erfahrung hält, imma"^
nente, diejenigen aber, welche diese Grenzen überfliegen sollten, transzendente
Grundsätze nennen."
So unterscheidet z. B. Thomas v. Aquino eine actio manens und actio
s. Thomaslexikon von Schütz unter actio: duplex est actio, una
quae transit in exteriorem materiam, ut calefacere et secare, alia, quae manet
in agente, ut intelligere, sentire et velle. Das erstreckt sich bis in die Neuzeit.
Clauberg (op. omn. 1691, S. 322) gibt der Sache die Fassung: si ipsius rei,
quae dicitur agere, Status mutetur, est actio immanens, sin alterius, est actio
transiens.
Es stammt aber diese Unterscheidung, wie der gesamte Grundstock der scholastischen Terminologie, von Aristoteles. S. z. B. Met. 1050a, 24:
^
transiens;
TtTv [i£v eoyaTov
£T£pov
ot/.t'a
aKo T^;
napa
x;^v
r\
y^r'jii,
o'ieo)«;
oTov
epyov,
oiy.oo6[xri^v/.
oJ/ew?
aTi'Evi'wv
ri
opaat;,
8e
xa\
Yt^vexai
ouO'kv
Tt,
o!ov
yiYvexat
axo
xf?
jrapa
Tau"n)v
o?xo5o[itxf?
In soIcher Unterscheidung eines Tuns, das
gegen
Das Problem der Religion.
aber
nicht
nente,
die
daß Gott nicht aus
Dingen
wirkt,
Welt also
in
der Welt als
aller
sich selbst heraustritt,
sondern daß er dabei bei sich
Dinge
Es
sei.^
wenn er zu den
selbst verbleibt,
die
Demnach ist hier nicht sowohl Gott in
Neu .gegenüber der Scholastik ist
die Welt in Gott.
sich
nur die
dabei
Wirken
Ursache
transeunte
bedeutet,
39I
neben
trägt.
Ausschließiichkeit
der
Immanenz,
dem
hat
auch jene bereitwillig
transeunten
ein
immanentes
aner-
- Einen anderen Ausgangspunkt hat transzendent und trans-
kannt
zendental.
Transzendent
hießen
(transzendentia)
des Mittelalters die allgemeinsten
Hälfte
in
Eigenschaften
der
zweiten
der Dinge,
welche nach neuplatonischer Lehre jenseit der einzelnen Kategorien
liegen. 2
als
Von
aus ergab sich
hier
leicht
eine Beziehung auf Gott
das allen menschlichen Begriffen überlegene Sein; so wirkt der
Sprachgebrauch auch bis in die Neuzeit.^
Kant hat dann transzendent
und transzendental geschieden und unter Umprägung der Bedeutung
zu "V^erkzeugen seiner eigentümlichen Denkweise gemacht.*
sich selbst
und
richtet
begründet sich das deutliche Auseinandertreten von praktischer
ist,
eines anderen, das auf die
Hervorbringung eines Werkes ge-
und künstlerischer Wirksamkeit.
Für den Ausdruck Immanenz sei noch die
von Heman (Kantstudien VIII, 1, S. 58) angeführte Stelle des Augustin erwähnt (epist. 268 ad Nebr.): In se habeat haec tria et prae se gerat, primo
ut sit, deinde ut hoc vel illud sit, tertio, ut in eo quod est maneat, quantum
potest.
Primum illud causam ipsam naturae ostentat, ex qua sunt omnia.
Alterum speciem, per quam fabricantur et quodammodo formantur omnia.
Tertium manentiam quandam, ut ita dicam, in qua omnia sunt.
^
Ethic. pars
non vero transiens.
prop. XVIII: deus est
I,
omnium rerum causa immanens,
Begründung heißt es: om.nia quae sunt in deo
sunt et per deum concipi debent, adeoque deus rerum, quae in ipso sunt,
In der
est causa.
'
Grund der Schrift de causis zuerst die vier Beunum, verum, bonum, später außerdem res und aliquid; so sprach
Als solche gelten auf
griffe ens,
man von einer unitas, veritas transcendentalis u. s. w.
' S. z. B. Bayle oeuv div. (Haag 1727) III, 871a: Si l'Origeniste repond
que les vertus de Dieu sont transcendentelles, qu'elles ne peuvent point etre
dans la meme categorie que Celles de l'homme. Transzendental im älteren
Sinne verwenden noch Ch. Wolff und Lessing. Lambert nennt transzendente
Begriffe solche, die „das Allgemeine der Körper-
und Geisterwelt zusammen-
nehmen".
*
Über transzendent s. 0., über transzendental sagt er (Kritik d. r. V. III, 49)
»Ich nenne alle Erkenntnis transzendental, die sich nicht sowohl mit Gegen-
ständen,
sondern
mit
a priori möglich sein
unserer
soll,
Erkenntnis von Gegenständen
überhaupt beschäftigt."
sofern diese
Letzte Probleme,
392
Die
b)
Bewegung der Neuzeit zur Immanenz.
Der Gesamtverlauf der Neuzeit zeigt einen Zug zur Immanenz,
Eigentümlichkeit
dessen
durch
eine
Vergleichung
der
mit
Haupt-
bewegung der griechischen Kultur besonders deutlich erhellt. Das
Griechentum wurde durch die Erfahrungen seiner Arbeit immer
Von der Außenwelt,
weiter über die sinnliche Welt hinausgetrieben.
von der die Forschung begann,
schrittweise
religiösen
zurück
in
verlegte sich ihr der
Innenwelt,
die
bis
der
in
Gestaltung der Wirklichkeit bei Plotin die nächste Welt
zum bloßen Gleichnis einer unsichtbaren wurde.
religiösen
Überzeugung das
Jenseits
als
Die Neuzeit ver-
Galt im Mittelalter der
gerade entgegengesetzte Richtung.
folgt die
Schwerpunkt
abschließenden
das wahre Vaterland und
gab nur die Beziehung darauf dem Diesseits einen Wert, so beginnt
dem Verlangen,
Neuzeit mit
die
das Wirken des Göttlichen mehr
und
innerhalb der Welt aufzusuchen, ja diese als
einen Ausdruck
Abglanz des göttlichen Wesens zu verstehen.
Das ergibt zunächst
einen Panentheismus,
naissance.
das Bekenntnis der edelsten Geister der Re-
Bald aber verschiebt sich das weiter dahin, daß mehr
und mehr die Welt zur Hauptsache wird, daß die Gottesidee mehr
dahin wirkt, ihr eine größere Tiefe zu geben
lichkeit
So der Pantheismus
zu eröffnen.
und
eines Spinoza.
ratur
überwältigend angezogen, indem er
als eine
eines
neue Wirk-
Giordano Bruno
Er hat die klassische Zeit der deutschen
brücken, im besondern die weiteste
und
alle
Lite-
Gegensätze zu über-
freieste
Behandlung der
sichtbaren Welt mit aufrichtiger Anerkennung einer unsichtbaren zu
verbinden
versprach.
Solche pantheistische Denkweise
ist
auch im
wo dies seine eigentümda neigt es weit mehr, wenn nicht zum
19. Jahrhundert keineswegs erloschen; aber
Art vollauf
liche
entfaltet,
Atheismus, so doch zu einem Agnostizismus,^ einer Ablehnung
transzendenten
Fragen
als
schlechterdings
unlösbarer
aller
Probleme.
Über den Ursprung des Ausdrucks berichtet genau R. Flint in seinem
Der Schöpfer des Wortes agnostic,
aus dem dann bald agnosticism hervorging, ist Huxley. „According to Mr.
R. S. Hutton this latter word (d. h. agnostic) was suggested by Professor
*
hervorragenden Werke Agnosticism (1903).
Huxley, at a party held previous to the
at
in
now defunct Metaphysical Society,
James Knowles's house on Clapham Common, one evening in 1869,
my Hearing. He took it from St. Paul 's mention of the altar to ,the
iVlr.
unknown God'" (s. Hint, S. Iff.).
Das Problem der Religion.
Praktisch
ergibt das
Religion
aus
dem
wie
eine
eine Ausscheidung
das andere
der
War also das Göttliche zuerst unserem
Leben.
Dasein angenähert, dann
393
beseelende Kraft ihm aufs engste ver-
als
bunden, so verschwindet es schließlich gänzlich oder wird zu einer
unzugänglichen Ferne; so
aus
ist
die Religion
dem modernen Menschen
Macht zu
einer
allesbeherrschenden
der
Nebensache,
ja
zu
bloßen Illusion gesunken, und es hat die unmittelbar gegen-
einer
immer
wärtige Welt
Sinnen und Denken an
ausschließlicher alles
und
fehlt es
nicht an Widerständen
gegen eine solche Bewegung, schon deshalb
nicht, weil jede frühere
Phase sich gegen die späteren behauptet;
aufgehalten aber haben
Natürlich fehlte
sich gezogen.
sie jene nicht.
So tiefgehende Wandlungen bloß dem Unglauben und bösen
Willen der Individuen schuldgeben kann nur eine flache Denkweise;
sicherlich
hatte
die Sache tiefere Gründe in den allgemeinen Verhält-
Gründe, die unbefangen gewürdigt sein wollen. — Die ältere
nissen,
Art der Religion stieß zunächst mit einem wesentlich veränderten Lebensgefühle der Menschheit hart zusammen.
wo
Lebensmut,
aller
Glaube an
aller
Sie
entsprach einer Zeit,
irdische
eine
Zukunft ge-
wo man zur Religion seine Zuflucht nahm, um
Nun aber hatten
Frieden und Sicherheit zu finden.
brochen war, und
Ruhe,
dort
lange Jahrhunderte bei jugendfrischen Völkern einen neuen Lebens-
mut
erzeugt;
ein
solcher wollte nicht sowohl
nicht
sowohl
ein
sicheres Geborgensein
Ruhe als Betätigung,
Gefahr und
Wagnis,
als
Kampf; er konnte die Welt nicht verschmähen, ihn
in
sie
hinein,
Zu
steigern.
solcher
Wandlung der Grundstimmung kamen
entsprach
und zu
einer Arbeit,
wirkte.
Nach den verschiedensten Richtungen hin
tiefer
sich
die
ihr
die
ihrer Verstärkung
Erfolge
nächste Welt
trieb es mächtig
um seine Kraft an ihr zu erproben und durch sie zu
ist
die sinnlich-
dem Menschen bedeutender geworden, hat sie ihm sich
und zugleich einen engeren Zusammenhang bei
erschlossen
selbst
gefunden,
hat sie sein
größeren Leistungen geführt.
allgemeinen Gesetzen
und
in
Handeln
stärker
bewegt und zu
Die Wissenschaft zeigt die Natur unter
festen
Zusammenhängen,
sie entfernt
auch aus der Geschichte das Wunderbare und versteht sie aus ihren
eignen Verkettungen. Das gesellschaftliche Zusammensein der Menschheit
nimmt mehr
geistige
Aufgaben an
sich
und sucht mit Ein-
setzung gewaltiger Kraft aus unserm Dasein ein Reich der Vernunft
zu machen.
Alles
zusammen hat diese Welt mehr als je zuvor dem
Letzte Probleme.
394
Menschen auch
zur Heimat gemacht.
geistig
Sonderstellung
seine
aufs
ärgste
nämlich die Welt wird,
ständiger
und Walten eigne Gesetze
Zugleich aber wird
bedroht.
Je
mehr
sie
je
erweist, desto kleiner,
der wird ihr gegenüber der Mensch.
größer
durch
und
selb-
Wirken
alles
desto verschwinden-
Bei solcher Kleinheit können
unmöglich die ihm eigentümlichen Größen die Wirklichkeit fassen
und
seelisch
Annäherung
nahebringen.
innerlich
hältnis zu ihren
Rückt
Welt
die
bei
äußeren
aller
so entfällt alles innere Ver-
Gründen, so droht alle Religion ein bloßer Anthro-
pomorphismus zu werden
wo
so
Ferne,
weite
in
die Religion
und zur Mythologie zu sinken.
rückt sie leicht aus
behauptet,
sich
Auch
dem Zentrum
des Lebens in seine Peripherie, und wird sie aus einer natürlichen,
beinahe selbstverständlichen Überzeugung zu einer kühnen,
So
haltbaren Behauptung.
ist
kein
es
Wunder, daß
derer, die alles Überschreiten der Erfahrung verwerfen
«immanent" fassen wollen,
gaben
sich
immer
lauter
die
mühsam
Stimmen
und alle Auferheben und
wachsendes Echo finden; nie dürfte die Verneinung der Religion
ein
so in die Massen gedrungen sein,
gewaltet
haben,
mehr eine
wie
ist
Mag
der eine in ihr
Hemmung klarer Einsicht, der andere eine Lähmung der
Unterdrückung freudigen Lebensgefühls
Tatkraft, der dritte eine
blicken:
nie so viel Antipathie gegen sie
der Fall
heute
es
er-
dünkt sie ein verderblicher Wahn, der mit
hier wie dort
das der endgültige Abschluß des
allen
Kräften auszutreiben
alten
Problems, oder ist es eine bloße Woge der Zeit, die vorbeigehen
sei.
Ist
und vielleicht das gerade Gegenteil der eignen Absicht bewirken wird?
c)
Die Verwicklungen im Begriffe der Immanenz.
Ihre Stärke hat die
im Angriff; sobald
aus das Leben gestalten
wicklungen.
Bewegung gegen die Religion vornehmlich
eignes Vermögen zeigen und von sich
ihr
sie
soll,
erscheinen Verwicklungen
Erschreckend dürftig pflegt zu
sein,
über Ver-
was als Ersatz der
und selbst dies Dürftige ist zum guten Teil
fremdem Boden gewachsen und von dort aus zugeführt. Die
Religion geboten wird,
auf
«immanente" Lebensführung und Weltanschauung pflegt keineswegs
aus reiner Erfahrung zu schöpfen, sondern
sie
un-
idealisiert diese
vermerkt, sie mischt ihr etwas bei, was einer ganz anderen, nämlich
der pantheistischen Denkweise angehört; ein Pantheismus verblaßter
Gestalt
als
hat
sich
der
selbstverständlich.
einzelnen
Gebiete
bemächtigt und
Er pflegt dabei kein
gilt
dort
offenes Bekenntnis
zu
Das Problem der Religion.
395
wagen, sondern seine Erhöhung der Dinge eher zu verstecken, er ist
mit solcher Unklarheit gegenüber dem Pantheismus eines Spinoza und
eines
Goethe
und
ein unechter
schlechter Pantheismus.
Pantheismus erscheint
schlechter
in
sophie, welche unbedenklich die Natur beseelt
Wertbegriff behandelt;
erscheint
er
Ein solcher
monistischen Naturphilo-
einer
und als einen hohen
einer Geschichtsphilosophie,
in
welche bloße Massenbewegungen Vernunft erzeugen
und eine
läßt
Evolution zur Vernunft verkündet, obwohl dieser Begriff in ihrer Ge-
dankenwelt keinerlei Grundlage hat; er erscheint
groß behandeln.
politischsozialen
in
Bewegungen, die den Menschen wie er leibt und
und
lebt als edel
Überall ein verstecktes Idealisieren der Erfahrung,
zugleich aber ein Abschleifen der Gegensätze, ein
Verkümmern der
eigentümlich geistigen Art, ein Einschläfern aller Selbsttätigkeit.
Auch
nicht
so
wissenschaftlich angesehen
wie er sich zu geben
einfach
nächste Wirklichkeit, die uns ganz
wirklich an uns selbst?
einander,
wenn
Ist
völlig
er
Was
pflegt.
soll,
die
was
ist
Befund des Neben-
wird?
herausgestellt
denn
ist
und gar einnehmen
es der unmittelbare
rein
Immanenz
der Begriff der
ist
Dann müßten
wir uns in einen Haufen, ein Bündel einzelner Empfindungen auflösen;
das aber geht aus dem einfachen Grunde nicht, weil es freischwebende
Empfindungen überhaupt
ungen eines
Ich,
nicht gibt,
sondern immer nur Empfind-
meine und deine Empfindungen, nicht Empfind-
ungen an sich. So werden wir von der Empfindung immer wieder auf
eine
zusammenhaltende Einheit gewiesen,
erscheint
es
eignen Bereich ein Gegensatz, und es wird zur Frage,
des Lebens
erscheint
liegt.
bei
uns
Reicht
aber
selbst
eine
in
unserem
wo der Kern
das Problem so weit zurück,
Abstufung,
so
erhellt
deutlich,
und
wie
wenig mit dem Schlagwort der Immanenz gewonnen wird.
Beim
Recht der
religiösen
Problem im besondern geht gewiß mit gutem
Zug der Neuzeit gegen die mittelalterliche Transzendenz
mit ihrer Verdopplung der nächsten Welt, aber damit
ist
keineswegs
entschieden, daß unser ganzes Leben eine einzige Fläche bildet.
Es
notwendig werden,
Um-
könnten Abstufungen
kehrung dahin erfolgen
sichere
in
müssen,
daß,
es
ja
was
Boden unseres Lebens und Wirkens
könnte eine
als
der
selbst einen
Halt
uns zunächst
gilt,
einer tiefer gegründeten Welt erst zu suchen hätte.
die Wirklichkeit, die unser ganzes Leben
Was ist denn
und Streben umfassen soll?
Die Welt des unmittelbaren Sinneseindrucks dafür zu erklären, das
hieße den großen Vorkämpfern der Im.manenz,
einem Spinoza und
Letzte Probleme.
396
einem Goethe, schroff widersprechen, das hieße die seeHsche Tiefe
der gesamten modernen Kultur verkennen.
Die Anerkennung einer
von geistigem Leben getragenen WirkHchkeit aber erzeugt sofort die
Frage,
ob jenes den ganzen Umkreis unmittelbar an
es nicht
windung,
ja
Bekämpfung es erst nach
Hilfe weiterer Zusammenhänge
ob
sich zieht,
Hemmungen stößt, deren Über-
draußen und drinnen auf
weiterer Kräftigung
unternehmen kann.
und mit
Namentlich
ist
es
die Tatsache der vielfachen Unvernunft im Natur- und Menschenleben,
an der jedes System ausschließlich immanenter Vernunft mit seinem
Denn
hier
bleibt
die
Unvernunft
wegzudeuten
und
abzuschwächen,
aus
den Augen
zu
oder
sie
Pantheismus
scheitert.
rücken,
Wirklichkeit anzuerkennen
als
nur die Wahl,
ein
sie
entweder
möglichst
Grundelement
der
und damit für unangreifbar zu erklären.
Entweder also eine Tendenz zum Optimismus mit seiner verflachen-
den Art, oder zum Pessimismus mit seiner Verneinung und schließ-
demnach die Dinge nicht,
wie das Verlangen nach Immanenz es darstellt. Hüten wir uns, ein
Weltbild deshalb für wahr zu erklären, weil es sich unserer Vorstellung als das glatteste und bequemste empfiehlt. Denn was anderes
So einfach
lichen Verzweiflung.
liegen
wäre das als eine neue Art von Anthropomorphismus, die das menschliche
d)
Wollen und Wünschen zum Maßstab der Wirklichkeit macht?
Das Wiedererwachen des religiösen Problems.
So erscheinen arge Verwicklungen beim Versuch eines Aufbaues
des Lebens ohne alle Religion.
Aber das würde eine derartige Be-
wegung noch keineswegs hemmen; sehr viel Unklarheit und Widerspruch läßt sich ertragen, wenn der Zug des Lebens kräftig und
selbstbewußt
Problems
Nun aber ist ein Wiederaufsteigen des religiösen
ist.
inmitten
aller
leidenschaftlichen
Befehdung der Religion
heute nicht zu verkennen; das Vordringen der Verneinung in immer
breitere
Massen hindert nicht, daß auf der Höhe des Geisteslebens
die Religion wieder weit
schaften erregt; es
ist
mehr das Denken
einmal
so,
daß
beschäftigt
und Leiden-
in derselben Zeit verschiedene
Strömungen durch- und gegeneinandergehen können, und daß dabei
der Unterstrom
dem Zuge der Oberfläche direkt widersprechen mag.
Um aber der Tatsache eines Wiederaufsteigens der Religion gewiß
zu werden, brauchen wir nur unsere Zeit mit der unserer Klassiker
zu vergleichen; dort war die Religion mehr eine freundliche
säumung des Lebens, heute
ist
sie
in
Um-
seinen Mittelpunkt getreten,
Das Problem der Religion.
397
entzweit sie die Menschen bis zu härtestem Kampf, fließt sie in die
Behandlung
aller
gewaltige
eine
Angelegenheiten
ein,
übt
Denn auch
die
Verneinung
Kraft.
der Art, daß die Religion
als
heute nicht
ist
etwas welkes und greisenhaftes ruhig
geschoben würde, sondern die stürmische Leidenschaft des
bei Seite
genug als etwas noch sehr
Angriffs zeigt sie deutlich
und Wirksames.
tiges
im Ja wie im Nein
sie
Vielleicht besagt sogar die
Reales,
Kräf-
Verneinung selbst
oft weniger eine völlige Ablehnung der Religion, als sie ein
Verlangen
nach einer anderen, einfacheren, den Bedürfnissen der Gegenwart an-
gemesseneren Art der Religion bekundet. Jedenfalls erscheint danach
die Religion nicht wie ein mattes Licht, das langsam und still verglimmt.
Was mag
diese
es
Umschlag herbeigeführt hat?
das diesen
sein,
war er die
Schwerlich
von
Frucht
apologetischer
vorwiegend auf solche zu wirken,
pflegt
Denn
Arbelt.
welche schon ge-
wonnen sind; sie mag befestigen und zusammenhalten, vorzudringen
ist nicht ihre Art.
In Wahrheit ist es ein Rückschlag im modernen
Lebeh selbst, in dem die Bewegung wurzelt; eben indem jenes Leben
mit seiner Weltfreudigkeit sich frei entfalten und sein volles Vermögen einsetzen konnte, sind seine Schranken, ja ist seine Ohnmacht
gegenüber
den
erfahren
wir
welche
durch
Sichausleben
zwingend
Dasein
letzten
die
Verneinung
des Gegenteils,
und
geleistet
wurde,
dingungen
vertieft;
einmal
das
ungehemmte
einer
Behauptung
Die Richtung des Lebens auf das unmittelbare
Aberglauben ausgetrieben,
das
ist
in
viel
sonst
mannigfachster Weise
Aber was immer in dieser Richtung
weitergebildet.
vorwiegend peripherer Art, es hat die Be-
unseres Lebens
verbessert,
nicht
aber das Leben selbst
aus aller unsäglichen Arbeit
steigt
daher
erscheinen.
Zusammenhänge
muß
und damit
innere Leere auf,
länglich
wieder
Weltgeschichte,
durch
schlummernde Kraft erweckt, jenes Dasein
gefördert
der
Unerläßlichkeit
hindurch,
die
Wahn und
viel
geworden;
deutlich
Beweisführung
indirekte
erweist.
hat
Fragen
jene
Die Ablehnung
gestaltete
die
alles
jenes
aller
Kultur
schließlich
eine
Bemühen als unzu-
und jeder unsichtbaren
mehr und
mehr zu
einer
bloßen Menschenkultur; das mochte so lange keinen Anstoß erregen,
als
das Menschsein selbst
klärter Gestalt
^
So machte
umfassenden Ideal.
als
als
ein
gesehen wurde.^
Er selbst ist."
hoher Idealbegriff galt und in verDas aber geschah unter dem Ein-
vornehmster Weise Herder die „Humanität" zum all„Der Mensch hat kein edleres Wort für seine Bestimmung,
in
Letzte Probleme.
398
Denkweise, die
fluß derselben
lichkeit
abgelehnt
Verklärung
wird;
jetzt als
muß
so
der Mensch
fallen,
eine Verfälschung der Wirk-
mit ihrem Schwinden auch jene
seiner natürlichen
in
ohne Hülle erscheinen und zum einzigen Maß
Beschaffenheit
Wahren und
Nun hat aber eben das moderne Leben mit seiner
Guten werden.
Entfesselung
alles
so
Kräfte
aller
Niederes aufgewühlt, es
stellt
Unerquickliches,
viel
Trübes
und
uns das Kleine und Scheinhafte einer
vor Augen, daß die Hoffnung
immer geringer wird, von hier aus einen zusagenden Lebensstand
erreichen und dem menschlichen Dasein einen Sinn und Wert verleihen zu können. Immer mehr greift die Empfindung um sich, daß
im Menschen etwas steckt, was durch jenes immanente Kulturleben
nicht belebt wird, und daß jenes Verkümmerte etwas Unentbehr-
bloßen
liches,
Menschenkultur so
das Beste von allem
vielleicht
So erwächst
Menschen,
nach
festhält
und
einmal
wieder
sichtlich
von
von
Befreiung
einer
niederdrückt;
ist.
nach einer inneren Wandlung des
ein Verlangen
der
neue Zeit
eine
kleinen
Art,
gekommen,
ist
die
ihn.
es treibt
bloßen Menschenkultur zu einer wesen-
einer
erhöhenden und umbildenden Geisteskultur; das aber führt mit Not-
Forderung einer neuen Wirklichkeit und damit auf
wendigkeit zur
Weg der Religion.
den
Zunächst freilich ergibt es eine höchst verworrene Lage. Während
von innen her eine neue Art des Lebens und Seins verlangt wird,
hält
unser Verstand und unsere Arbeit uns beim unmittelbaren Da-
sein fest; wir
möchten etwas Höheres, aber wir finden keinen Weg
dahin und können doch
vom Ziele nicht lassen.
vom
hin-
einen
zum andern
So werden wir
und hergeworfen und widersprechen
unablässig uns selbst. Aber bei aller Unfertigkeit und allem Unbehagen
ist
wenigstens das eine gewonnen, daß wir aus
Besitz
wieder
gekommen
in
ein
Suchen,
sind,
daß
die
ein
alten
redliches
und ewigen
dem vermeintlichen
und
eifriges
Fragen
mit
Suchen
frischer
Kraft wieder aufsteigen.
Wie sich das freilich weitergestalten wird,
an
Bedingungen des Menschen und des Ge-
das
liegt
vielfachsten
schicks; darüber
e)
kann
erst die
Zukunft entscheiden.
Forderungen für die gegenwärtige Lage der
Religion.
Beim Überblick der gegenwärtigen Lage der Religion
bemerkenswert besonders die Tatsache
ins
fällt
als
Auge, daß zwischen der
Das Problem der Religion.
399
überkommenen kirchlichen Form der Religion und einer univerBewegung zur Religion aus dem eignen Streben der Zeit
eine schroffe Spannung besteht; es gibt heute viele, welche religiös,
saleren
aber durchaus nicht kirchlich sein wollen,
von der Kirche abgestoßen
Den
als
welche sich ebenso sehr
von der Religion angezogen fühlen.
Grund dessen mag man darin
nächsten
suchen, daß zwischen
der überkommenen Form des Christentums und dem Kulturleben der
Gegenwart eine weite Kluft besteht,
aufs
Äußerste
die ein gegenseitiges Verständnis
Das Weltbild
erschwert.
hat
wesentlich
sich
ver-
und Außermenschliche verschoben, die
ältere Gefühlslage erscheint dem modernen Menschen als zu weich
und zu matt, die Zeit stellt ihm neue praktische Aufgaben, die seine
ändert, namentlich ins Große
ganze Kraft verlangen; während das alte Christentum einer ermüdeten
und eingeschüchterten Menschheit neue
mut zuzuführen suchte,
und
starken Lebensdranges
dieses,
ist
das
Leben
in
Die Haupt-
tun.
dem Widerspruch
seine
Schärfe
daß
sich
der Zeit nicht mehr aus eigner Erfahrung
eine
Frage zusammenfaßt, deren Beantwortung
bildet:
Vor all dem Wirken und Schaffen
frischem
Menschheit verblaßt;
wo
sprünglichkeit
da
fehlt,
in
Kraftgefühl
der moralischen
Frage
die
in
Befreiung und
der inneren
vor jugendlich
frischen Lebens-
die
den Kern des Christentums
Rettung,
und
Wirkens zu
rastlosen
sache aber und was vornehmlich
gibt,
Kraft
hat jetzt die Religion mit einer Menschheit
Erneuerung der Menschheit.
die sichtbare Welt hinein und
ist
jene
Frage der modernen
aber der Frage die volle Kraft und Ur-
muß
auch
die Antwort die
Herzen gleich-
gültig finden, da verdunkelt sich das Recht und die Notwendigkeit der
Sache,
da wird hingegen
die Schätzung
voUkommnem enthält.
die
Abhängigkeit
gewährte Hilfe
der
viel
zu entfremden.
die
alles grell
was
beherrschen,
die
in
die
Augen fallen und
überlieferte
Endlich hat wohl auch bei
Kirche
vom
Staate
und
leicht
Fassung an Ununs Deutschen
die ihr
vom Staate
dazu beigetragen, jener die Gemüter innerlich
Denn
den anderen germanischen Völkern
bei
ist
Entfremdung anscheinend nicht so groß.
So
wegung
ist
es
ihre
älteren Art
begreiflich,
eignen
Bahnen
Vor allem
sucht.
religiöse
erstrebt
sie
Beder
gegenüber, die ihr zu eng und gebunden dünkt, eine
größere
Weite,
für
Weltumgebung;
die
daß die neu aufsteigende
eine
möglichste
sie
sorgt
Universalität,
sich
lungen im eignen Innern des Menschen
weniger
als
mehr
um
die
Offenheit
Verwick-
um sein Verhältnis zum
Letzte Probleme.
400
All;
dieses
keit ihn
möchte
ihm
sie
innerlich naherücken, seine Unendlich-
miterleben, seine Schönheit ihn genießen lassen; in solcher
Stimmung
künstlerischen
scheint
von
Befreiung
eine
sich
allem
Kleinmenschlichen zu vollziehen und die Seele im reinen Äther des
Alls sicher
und
zu schweben.
selig
Solches Streben gegen ein Sicheinspinnen des Menschen in die
bloße
Menschlichkeit,
solches
Verlangen
zum
der
Religion
und
in
wesentliche
Seite
deutende Wirkungen
hier aufstrebt,
die
hat
bedeutet
eine
Geschichte
be-
All
ihrer
Aber eine andere Frage
geübt.
ist,
ob,
was
Gesamtaufgabe der Religion zu lösen und damit
alle geschichtliche Gestalt zu
verdrängen und ersetzen vermag.
Wird
nämlich das Neue streng auf sein eignes Vermögen beschränkt und
nicht stillschweigend
durch das von der geschichtlichen Religion ge-
botene Leben in mannigfachster Weise ergänzt, so kann
und
Freiheit
Diese Art
und Leere
starke Vagheit
eine
dringt über feine
der Religion
nicht vor zu einer echten Tatsächlichkeit;
neue Welt zu eröffnen,
eine
stellt
sie
Weite
zarte
Stimmungen
dem Menschen eine
ihm nur die vorhandene Welt in
oder
Beleuchtung,
liebenswürdigere
und
statt
alle
wohl verdecken.
nicht
umsäumt
sein
sie
Leben
mit gefälligen Stimmungen, welche Mußestunden angenehm ausfüllen
mögen, die aber gegenüber dem Ernst des Lebens in kläglicher Weise
versagen.
Denn nun und nimmer wird damit
der
erreicht,
Seele
werden Kräfte
Schuld befreien, wird
Menschheit durch
sammengeführt.
entbunden,
eine Weiterbildung
die
von
Not und
dem Leben ein fester Halt geboten, wird die
eine
bei
Schöne
sich
Bilder,
selbst
befindliche
Innenwelt zu-
schöne Aussichten, aber Bilder, die
den bloßen Entwurf nicht überschreiten können
!
Die hier erwachsende
ästhetisch -pantheistische
Stimmung mag
Vorbereitungen
liefern,
der Hauptaufgabe der Religion
gewachsen; was
sie
an Wahrheit
und festerem verbinden,
enthält,
wertvolle
das
Anregungen und
ist
sie
nicht
muß sich mit anderem
um zur Förderung zu wirken.
Aber so wenig jene neue Art der Religion befriedigen kann,
gegen die kirchliche Form,
es bleibt die Tatsache des Widerspruchs
es bleibt ein Spalt
im
unabweisbar wird
damit
Überwindung
religiösen
die
arbeiten läßt.
wieder zusammenfinden, so
entgegenbringen, so
Leben und Streben der Gegenwart;
Frage,
ob und wie
Sollen sich die Zeit
sich
und
zu seiner
die Religion
muß die Zeit der Religion eine Frage
muß diese sie aber in einer Weise beantworten,
welche anzunehmen der Zeit nicht unmöglich
ist;
dazu aber wird
Das Problem der Religion.
beiden
auf
es
401
Wandlungen oder doch Weiter-
erheblicher
Seiten
Nur dann wird das Verlangen der Zeit nach
Religion wieder stark und überwältigend werden, wenn jene im
menschlichen Leben große innere Verwicklungen erkennt und zum
eignen Erlebnis macht, und wenn sie zugleich die Spitze dieser
bildungen bedürfen.
Verwicklungen im moralischen Probleme
Aber andererseits
findet.
muß auch die Religion das moralische Problem nicht nach der Enge
des
unmittelbaren
Eindrucks,
sondern
den Gipfel einer allum-
als
fassenden Bewegung verstehen und behandeln; damit wird sie selbst
Grundlage gewinnen und der
eine breitere
sonst unvermeidlich
die
ihr
die
Religion
einen
sicheren
der Zeit gewonnen, und
und
klar
ist
sicher geworden,
kommenen Bestand
Kontakt
sie
so
mit
dabei
kann
dem
ihrer
sie
innersten
Streben
eignen Grundtatsache
ohne Gefahr ihren über-
Prüfung daraufhin
einer
entgehen,
Partikularität
Hat an diesem Zentralpunkt
anhaftet.
was
unterziehen,
in
ihm wesentlich und unwandelbar, was aber nebensächlich und den
Wa'ndlungen der Zeit unterworfen ist. Vor allem muß die Religion
verschanzen.
nicht
Wesen besinnen und
auf ihr eignes
kräftig
sich
Ihrer
intellektuelle
letzten
Absicht nach
Aufklärungen
sich fest in
bringt sie
über die Welt,
ihm
dem Menschen
erweckt auch
sie
nicht bloß neue Gefühle oder stellt neue praktische Aufgaben, sondern
eröffnet aus
sie
Seins,
ja
zur
einem direkten Verhältnis zum
beherrschenden
neue Welt;
eine
erster Stelle
sie
Urkraft
führt den
des
tiefsten
Lebens ein
Grunde des
neues
Leben,
Beweis dieses neuen Lebens an
durch die Tatsächlichkeit seiner weltgeschichtlichen Ent-
wicklung, durch die Umgestaltung der Wirklichkeit, die unablässig
von ihm ausgeht; darauf freilich muß
botene Leben
als die
sie
beherrschende Seele
dringen, daß das hier gealles
Lebens, als die un-
erläßliche Bedingung aller Geistigkeit anerkannt wird.
dies
Leben
faltung auf
kann
seine Ent-
ist,
dem Boden der Menschheit steht unter den Bedingungen
der Zeit und Geschichte; es
es
Aber so sehr
seinem Grunde nach überzeitlicher Art
das,
ohne
sich
muß sich mit ihnen auseinandersetzen,
selbst
zu
verlieren,
nur,
wenn
bei
ihm
zwischen Substanz und Daseinsform deutlich geschieden wird; denn
dann wird eine Unwandelbarkeit der Substanz mit einer geschichtlichen
Bewegung der Daseinsform
Hier nun
liegt
eine
besonders wichtige und schwierige Aufgabe der Gegenwart, es
gilt
eine
der
weltgeschichtlichen
vereinbar.
Lage des Geisteslebens
bloßen Strömungen der Zeitoberfläche
Eucken, Grundbegriffe.
4. Aufl.
—
nicht
den
- entsprechende Daseinsform
26
Letzte Probleme.
402
der Religion zu gewinnen, ohne darüber ihre Substanz zu verlieren
oder auch nur irgend abzuschwächen.^
Diese Verständigung zwischen Christentum und Neuzeit
manchen
Vor allem
ist
aber
dazu
nötig,
Anerkennung und Würdigung
finde.
erscheint.
minder
daß der große Wandel der Zeiten mit allem, was in ihm an inneren
leicht,
als
Notwendigkeiten
In der
sie
liegt,
volle
gewöhnlichen Apologetik geschieht das nicht
ist
Sie faßlr nicht
die Sache ins Ganze, sondern sje verbleibt bei zerstreuten Punkten,
sie
sich
versetzt
nicht
in
den anderen hinein,
sondern behandelt
ihn gänzlich von außen her, sie operiert mit bloßen Möglichkeiten,
indem
sie
Wege
offen
zeigt,
daß die moderne Bewegung noch immer gewisse
lasse,
die
gutem Willen wohl
bei
mit
dem
kirch-
immer mehr ins
die
Gefahr
einer
inneren
Unwahrsie
kommt
in
hinein,
ja
Künstliche
lichen
Glauben vereinbar
haftigkeit;
sie
gerät damit
man mag hier des Hume'schen Wortes von der Vergeb-
lichkeit eines
mit
seien,
Unternehmens gedenken, einen Durchbruch des Ozeans
Strohwischen
verstopfen
zu
Auf diesem Wege wird
wollen.
nun und nimmer die Religion die erstrebte Stellung im Ganzen desLebens wiedererringen, nun und nimmer kann sie so die schlichte
Einfalt, die seelische Nähe, die sichere Überzeugungskraft erlangen, ohne
die
ihre
Aufgabe
sich
nicht
Religion in der Gegenwart
ist
lösen
läßt.
Ein
Unsicherwerden der
nicht zu verkennen
und sei nicht ver-
Der christliche Lebenstypus ist heute der Menschheit keineswegs mit solcher überzeugenden Nähe gegenwärtig, daß ihn sowohl
jeder einzelne unmittelbar ergreifen und erfahren, als von ihm die
dunkelt.
Gesamtgestaltung
des
Lebens
mächtige
Einflüsse
empfangen
könnte.
Wo die Religion so viel Fremdes und Veraltetes mit sich
schleppt,
wo vielfach der Schutt von Jahrtausenden die ewigen Wahr-
heiten
verdeckt,
da kann die Religion nicht ihre volle Kraft mit
siegesgewisser Zuversicht entfalten,
Gewißheit haben, da wird
da kann
sie selbst
sie
keine axiomatische
durch Angriffe von kläglicher
Weshalb uns das Christentum bei allem, was an ihm vergänglich ist,
und es daher keines Bruches mit
ihm zu bedürfen scheint, darüber haben wir uns in den religionsphilosophischen Werken näher geäußert. Hier sei nur noch gesagt, daß uns kaum
ein Unterfangen so töricht erscheint als das, eine Religion durch bewußte
In allen anderen Gebieten haben wir eine
Reflexion bereiten zu wollen.
derartige Denkweise der sich verflachenden Aufklärung glücklich überwunden, gerade das allerinnerlichste Gebiet aber, das sie am wenigsten verträgt, muß sich ihrer noch immer erwehren.
^
einen unvergänglichen Kern zu enthalten,
Das Problem der Religion.
403
Flachheit beunruhigt, die an einer gefestigten und selbsterlebten Über-
So bedarf
zeugung ohne weiteres abprallen müßten.
sie
dringend
durchgreifenden Revision, einer energischen Freilegung ihrer
einer
beherrschenden Grundlinien, einer Abstoßung alles dessen, was welk
und morsch
ward,
Eine
Interesse.
sie
bedarf dessen
solche Aufgabe aber
besonders in ihrem eignen
kann nur im Elemente voller
Freiheit gelingen.
Aber zugleich muß auch
die Substanz der Religion energisch
gewahrt, zu kräftiger Betätigung aufgerufen, zur Scheidung von Echtem
und Unechtem, von Wesen und Schein im Befunde der Zeit verwandt werden.
Bedeutendes
nichts
Religion
die
Ohne eine Selbständigkeit gegen die Zeitlage kann
schlechthin,
wohl aber eines
im besonderen
leisten;
Christentum mit seiner Verneinung nicht
letzten
freilich
steht
das
der nächsten Welt
Abschließens bei dieser Welt in
unversöhnlichem Gegensatze zu einer bloßen Daseins- und Menschenkultur, wie sie
ist "keinerlei
gewiß
den Hauptzug des modernen Lebens beherrscht. Hier
Ausgleichung möglich, sondern nur ein ehrlicher Kampf;
muß auch er einen schließlichen Frieden ins Auge fassen, aber
es macht für das Ergebnis einen gewaltigen Unterschied, ob der Gegen-
vor allem zunächst mit voller Klarheit herausgestellt, oder ob er
satz
von vornherein abgeschwächt wird.
Hier findet sich nun auf
dem
Boden des Protestantismus viel schwächliche Konnivenz und viel ängstliche Reverenz gegen die moderne Kultur, als sei sie im Ganzen ihres
Seins schon eine ausgemachte Wahrheit und nicht selbst voll schwerer
Probleme; man scheut alles kräftige Nein,
als
aus 'dem nicht ein Nein hervorspringt;
sein,
könne ein Ja von Wert
man denkt zu klein von
der eignen Sache und kommt daher natürlich nicht weiter. Bescheidenheit
mag den Menschen zieren, für die Sache kann sie zum Unrecht
Eine Religion von Gnaden der bloßen Kultur, eine Religion,
werden.
die gar allen Wandlungen der Zeitoberfläche folgeleistet,
liches
und
Freiheit
haltloses
binden; in der Sache
zu
erreichen.
Gegenwart
So
voller
ist
zeigt
muß
So
Ding.
und Ursprünglichkeit
sich
mit
das
ist
ein kläg-
nach
Verlangen
mehr
dem nach mehr Tiefe ver-
das möglich, es
liegt
nur am Menschen, es
auch
daß
freilich
sich
Verwicklungen
hier,
und Widersprüche
die Lage der
ist,
daß wir
aber diesen keineswegs wehrlos ausgeliefert sind, sondern bei Auf-
bietung geistiger Kraft über
sie
hinauskommen können.
26*
Schlußwort
\ V/ir haben verschiedene Gebiete durchwandert und verschiedene
^^ Probleme behandelt, soweit es im Plan unserer Arbeit lag.
Wir
überzeugten uns dabei von der überströmenden Fülle des Lebens,
das unsere Zeit durchdringt; eine greisenhafte Zeit
die
Arbeit
leistet.
namentlich
Stoffes
eine
deshalb,
geistigen Gehalt
und die Hauptist
das
dem unermeßlichen Zustrom
des
unfertige
des Zusammenfassens
die Kraft
nicht
Expansion des Lebens seine Konzentration
Aber wir sahen
zugleich,
es wahrlich
h-ervorragende
hat
durchaus
weil
ist
und so
Probleme
Aber es ist, auf den
angesehen,
richtung
aber
bedeutende
so
nicht,
Zeit;
sie
entspricht,
weil
die
weitem überwiegt.
bei
daß das nicht wie ein
starres Schicksal
hingenommen zu werden braucht, daß vielmehr das Leben der Gegenwart
auch
voller
Möglichkeiten
ist,
und fördern können; nur muß
ein
die
eine Synthese
vordringendes Schaffen diese
Wir sahen dabei weiter,
Möglichkeiten ergreifen und weiterführen.
daß das
nicht
«gegebenen"
unmittelbar aus der
vorbereiten
Lage heraus ge-
schehen kann, daß es vielmehr dazu eines Überlegenwerdens gegen
diese Lage, der Erreichung eines neuen Standorts des Lebens, einer
Umkehrung
des nächsten Daseins
bedarf;
eine solche
Umkehrung
aber geigte sich möglich und nicht anders möglich als durch ein Selbführten alle einzelnen Punkte
ständigwerden des Geisteslebens;
so
darauf zurück, und an jeder Stelle
kam zur Sprache, wie die Über-
zeugung von solcher Selbständigkeit die Aufgabe verändert und ihre
Lösung anbahnt.
Bei
solchem Gedankengange drängt die Betrachtung der Zeit
über den eignen Befund der Zeit hinaus
zum Ausblick in die Zu-
kunft; daß wir aber aus der Zerstreuung in vielfache Aufgaben zu einer
allumfassenden Aufgabe, von betrachtender und zerlegender Reflexion
zu mehr schöpferischer Synthese, von überwiegender Hingabe an
die
Außenwelt zu
mehr Leben
Selbständigkeit gelangen,
ist
bei
uns
selbst
und mehr innerer
dazu nach bestem Vermögen mitzuwirken
auch die Philosophie berufen.
Sachregister
Abstrakt, Geschichte des Begriffes
Macht abstrakter Begriffe in
55
der Gegenwart 55.
Agnostizismus, Ursprung des Aus;
—
drucks 392.
Aktivismus, sein Unterschied vom
Pragmatismus 51.
Aktuell, Geschichte des Wortes 220
Zurückweisung der landläufigen
—
Aktualität 220.
Beharrungslehre
Altertum
(im
und Mittelalter) 194 ff.
Beharrungs- und Bewegungslehre, ihr Kampf 216; — drei verschiedene Lebenstypen dabei 223 ff.
Bevölkerungszunahme, ihre Bedeutung
die
für
Auffassung der
Geschichte 214.
Allegorische Deutung, ihre Wirkung auf das logische Denken 58.
Allgemeinbegriffe, ihre eingewurzelte Überschätzung 54 ff.
Allgemeines Stimmrecht, wann
zuerst gefordert 290.
Altertum (s. Griechentum) sein Intellektualismus 39;
Ausgangspunkt der Erkenntnisarbeit, wo gesucht 64ff., 222.
— was an ihm
zu verschiedenen Zeiten geschätzt
Bildung, Geschichte des Begriffs
und Ausdrucks 232 ff.
Charakter, Geschichte des Aus-
— Erörterung aus
drucks 355 ff.;
der Lage der Zeit 358 ff.
Christentum,
seine Stellung
Intellektualismus
39 ff.;
—
zum
seine
Steigerung der Geschichte 197.
Demokratismus, seineEntwicklung
263 ff.
— a posteriori, Aus— modruck und Begriff 84
A priori
ff.;
derne Verflachung abgewiesen 86.
Arbeit, Eigentümlichkeit und Probleme der modernen Arbeit 18 ff.,
313ff.;
— seine Bedeutung und
seine Schranke 317 ff.
Denken, Verschiedenheit seiner geschichtlichen Stniktur 58.
Denknotwendigkeit,
Kritik ihres
Vermögens 94 ff.
73 ff.
Arbeitswelt (des Menschen) 113ff.
Ästhetizismus 336ff.
Aufklärung, ihr Intellektualismus
— ihre Scheidung von Natur
und Seele 173; — ihre Stellung zur
Geschichte 254; — Folgen ihrer
— ihre
Ungeschichtlichkeit 261
41;
ff.;
Fassung des Geisteslebens 271.
Determinismus,
seine
verschiede-
nen geschichtlichen Formen 363 ff.
— sein Wachstum im
Jahr— seine Voraushundert 364
setzungen 368; — sein Bild der
19.
ff.;
Wirklichkeit 372.
Diskontinuitätsphilosophie (in
Frankreich) 146.
Sachregister.
406
Dualismus, das Wort 170; — sein
Hervortreten
der Neuzeit 173;
in
— Dualismus innerhalb des Monismus der Gegenwart 190.
Empirismus, sein Erkenntnisbild
— sein Recht und sein Unrecht 117ff.; — sein Ungenügen
88, 90;
für die
Gegenwart
Energie,
Definition
141
;
124.
Ostwald
von
— geistige Energie = Lebens-
Entwicklung, Ausdruck 192ff.; —
ihr Aufkommen und Vordringen
— drei Hauptphasen der
Entwicklungslehre
(religiöse, künst-
empirische)
lerische,
—
— Schei-
198 ff.;
neuer Lebenstypus 204 ff.;
dung von Entwicklung und Evolutionismus 204 ff., 226;
— Gefahren
des Evolutionismus 210, 215.
Erfahrung, Ausdrücke 84;
— ihre
zwiefache Bedeutung für das Er-
kennen 114 ff.;
keit
auf
dem
— ihre SchwierigGebiet des Geistes-
lebens 120.
Denken,
irre-
leitender Sprachgebrauch 119.
Lebensprozesse 56 ff.
;
100 ff .; 1 1 1 ff.
— Kampf über seinen Ursprung
87; — Erkennen unterschieden vom
Kennen 99; — inwiefern immanenter Art 113; — inwiefern auf
Erfahrung angewiesen 114, — verschiedene Stufen 116; — wo die
Entscheidung darüber 121 ff.
Erkenntnislehre (moderne),
ihr
(s.
192ff.;
Ästhetik,
Entwicklung),
Aus-
— evolutionistische
Ethik, Rechtslehre 212 ff.
—
AufFortschritt, Ausdruck 200;
kommen und Durchdringen der
Idee 199ff.; — seine Verschieden-
heit in verschiedenen Lebensgebieten
— vager Fortschrittsglaube
213; — Schranken 382.
215;
111;
30ff.,
sein
Grundbegriff
— sein Verhältnis zum
geschichtlich -gesellschaftlichen
Da-
sein lOOff., 103ff.
Geistigkeit, ihre drei Stufen (grundlegende, kämpfende, überwindende)
350 ff.
Genie, zur Geschichte d^s Ausdrucks
307.
Geschichte, Zusammentreffen von
Zeitlichem und Zeitlosem in ihr 5,
103;
— ihre Bedeutung für die
Philosophie 65;
—
Bedingungen
der menschlichen Geschichte 21
ff.;
— Bedeutung und Schranke der
— Gründe der
Geschichte 253
—
Bewegung zur Geschichte 254
ff.;
ff ;
260ff.;
— Bedeutung der Geschichte
Geschichte
der
Philosophie,
Forderungen für ihre Behandlung
66 ff.
Geschichtliche Gesetze 166ff.
Geschichtsforschung und Geschichtsphilosophie 256.
Geschlechtliches, seine Behandlung im modernen Subjektivismus
in der modernen Kunst 342.
338;
—
Gesellschaft und Individuum,
Einfluß auf das Weltbild 187.
druck
Begriffes 374.
für das Geistesleben 265 ff.
Erkennen, seine Abhängigkeit vom
Evolution
auf das Gefühl 20 ff.
Gegebenes, das Bedenkliche dieses
der Bewegung gegen die Geschichte
Erfahrung und
letzte
Freiheit und Schicksal 371 ff.
Kritik des Zurückgehens
Gefühl,
Geistesleben,
konzentrationen 60, 62.
197 ff.;
Freiheit und Gleichheit, ihre Verbindung 290.
283 ff.;
— Art der gesellschaftlichen
Kultur 288.
Gesetz,
154 ff.;
Geschichte
und
Problem
— Kampf darum in den
verschiedenen Gebieten 159ff.
Gleichheitsidee, ihr Ursprung
und ihre Geschichte 289.
Gleichmacherei, ihre Gefahren in
der Gegenwart 299 ff.
Sachregister.
Gottesidee,
der
über
Streit
ihre
— zweierlei Fass-
Fassung 346ff.;
ungen innerhalb des Christentums
Griechische Denkweise (s. Altertum), ihr Verhältnis zum Wahrheits-
12; — ihr Intellektualismus 39; — Eigentümlichkeit ihres
logischen Verfahrens 58; — ihr Vor-
problem
Problem auf geistigem
Gebiet 31 Off.
große Persönihr
—
;
zur Zeitum-
lichkeit, ihr Verhältnis
gebung 297 ff.
Hierarchische Ordnung 285.
Historismus, seine Gefahr 258ff.;
— Abwägung von Historismus und
Rationalismus 271 ff.
Humanismus,
als moderner Terminus 48.
Höher, Ausdruck und Begriff 78.
Idealismus,
Kritik der
Ausdruck
68ff.;
—
überkommenen Formen
Idealismus
ungen
76 ff.;
für einen
— Forder-
neuen Idealismus
— kritischer und positiver
81 ff.;
Idealismus 124 ff.
Immanenz, Ausdruck 390; — Bewegung der Neuzeit zur Immanenz
— VePÄ'icklungen im Begriff
Immanenz 394 ff.
Individualkultur 302ff.
der
Ausdruck 283
— Begriff 284
— verschiedene geschichtliche Geff.;
ff .
Kultur, Geschichte des Ausdrucks
und Begriffs 228ff.;
unterschieden von Zivilisation 231 ff.;
ihr
Wesen und Wert 235 ff.;
ihre
—
—
Arten
das
von Mensch
Verhälthis
mikrokosmische) 284 ff.;
der
geistigen
— Bild-
Individualität
— Bedeutung der Individuen
— moGeschichte 297
in der
ff.;
derner Individualismus 306 ff.
Intellekt,
seiner
Geringschätzung
widersprochen 52 ff.
—
— seine
und
und
Kultur
— Teilkulturen
— Probleme
und Gesamtkultur 249;
gegenwärtigen
der
Kultur 250 ff.,
381 ff.
Kulturstaat, bei Fichte 231.
Kunst, ihr Verhältnis zur Moral
329 ff.; — ihre Stellung im modernen Leben 339;
Verhältnis
von Inhalt und Form 340;
Bedeutung der Kunst für den nordischen Menschen 341.
Künstlerischer
Individualis-
—
—
mus 306ff.
pour
L'art
Geschichte
l'art,
des
Ausdrucks 339 ff.
Leben,
wissenschaftliche
einer näheren
Versuche
Bestimmung 145 ff.
Lebensbejahung und Lebensverneinung 376 ff.; — gegen erkünstelte
Lebensbejahung 384 ff.
Lebensenergien 60.
Lebensprozeß als Ausgangspunkt
Stell-
ung zum Geistesproblem 342.
Logik, Überschätzung ihrer Form
—
54ff.
ihre Abhängigkeit
Lebensprozeß 58 ff.
Materialismus,
Ausdruck
vom
und
Lehre 175 ff.
Mechanismus,
zur Geschichte des
—
Ausdrucks 126;
zur Geschichte
des Problems 133 ff.;
unzulängr
—
lieh als philosophische Weltanschau-
ung
Intellektualismus, Wort und Sache
37 ff.;
Überflutung des modernen
Lebens durch ihn 53 ff.;
—
240 ff.;
Kultur
—
237 ff.;
geschichtlichen
;
staltungen(organische, hierarchische,
371;
Kant 41 ff.;
Lebenstypus 39.
der Untersuchung 31.
Literatur, ihre gegenwärtige
Individuum und Individualität,
ung
—
durch
Geistesleben 243 ff.;
anstellen des Beharrens 194 ff.
Größe,
392;
Brechung
intellektualistischer
—
347.
des
407
142.
Mechanismus und Teleologie,
ihre Ausgleichung bei Leibniz 134;
— bei Lotze 138.
Sachregister.
408
156 ff.;
Metaphysik, der Ausdruck 108;
— Metaphysik
das Problem 104
—
Metaphysik
108;
und Kultur
ff.;
und Moral 324 ff.
des Ausdrucks
Art
logischen
seines
— Hauptstätte der
— sein Ord— Bekämpfnungssystem 196
Verfahrens 58;
organischen Lehre 151
;
ff.;
ung dieses Begriffes 277 ff.
— Problem
— Bedeutung bei Kant 16,
Modern, Verschärfung des Begriffes
durch die Entwicklungslehre 205;
Ausdruck und Begriff 273 ff.;
—
— echtes und falsches Moderne
280ff.
Monismus, Wort 170ff.; — Lehre
_ Monismus der Gegen-
wart 183 ff.
Moral, gegenwärtiger Mangel einer
genügenden Begründung 322ff.;
überkommene Typen 323 ff.;
—
—
—
Moral und Metaphysik 324 ff.;
AbMoral und Kunst 329 ff.;
lehnung der ;neuen Ethik" 336ff.
Natur, der Ausdruck angegriffen
—
— Definition des
Aristoteles 172; — Naturbild der
von Boyle 127;
Gegenwart 380ff.
Naturgesetz, zur Geschichte 154 ff.;
Naturgesetz und Sittengesetz
163.
Naturkultus, seit Giordano Bruno
158.
Naturwissenschaft
Ge-
und
Neunzehntes jahrhundert, seine
Bewegung^
gegen
tualismus 43 ff.;
18ff.;
—
— seine
den
Intellek-
sein
Realismus
ihre
Goethe 28.
Ökonomismus 311ff., 318ff.
blem 376ff.
zwiefache
zum Geistesproblem 13ff.;
Organismus,
und
Organisch
Ausdrucks 128ff.,
Organische
des Problems 130 ff.;
des
Geschichte
—
Natur-
136 ff.;
Geschichtsauffassung
und
— Unterschied der alten
und der neuen Fassung
139;
—
organische Gesellschaftslehre 149 ff.
Panentheismus 392.
Pantheismus, der Neuzeit 392;
sein Verblassen
in der
—
Gegenwart
394 ff.
Persönlichkeit,
Geschichte
des
—
des Begriffes
Ausdrucks 343 ff.;
Problem der Persönlich345 ff.;
Motive des
keit Gottes 346;
—
—
Kampfes gegen die PersönlichkeitsPersönlichkeit und
idee 347 ff.;
—
Metaphysik 349 ff.;
— Kritik der
h^tigen Bewegung zur Persönlichkeit
353;
— persönliche Gestaltung
der Arbeit unterschieden von subBedeutung der
jektiver 354 ff.;
—
im geschichtlichen
Leben 297 ff.
Pessimismus, Ausdruck und
Pro-
blem 376ff.
Phantasie,
ihre
Unentbehriichkeit
für die Philosophie 98, 106 ff.
72 ff.
Neuzeit,
bei
Öffentliche Meinung 297.
Persönlichkeiten
schichte 168.
Bewegung
12 ff.;
Ontologie, Ausdruck 109.
Optimismus, Ausdruck und Pro-
Mittlerer Mensch 289.
innere
Normen 24, 163.
Objektiv, Wort lOff.;
289.
Mittelalter,
179ff.;
ff.;
ung des Individuums 151, 286 ff.
Noologische Betrachtung, unterschieden von psychologischer 33.
Mikrokosmos, Ausdruck 285.
Milieu, Geschichte
— ihr Eintreten für Ent— ihre Befrei197
wicklung
Stellung
— ihre
—
Behandlung des Denkens 41ff.;
ihr wissenschaftliches Verfahren 59,
Philosophie, ihre Aufgabe und Methode 63;
— das Recht einer selb-
ständigen Philosophie 63 ff.; 93 ff.;
der
ihr Zusammenhang mit
—
^9
Sachre gister.
Bewegung der Weltgeschichte
100 ff.;
—
philosophia
62,
perennis
278.
Pragmatismus,
und
Darstellung
Kritik 47.
— Geschichte der
— verschiedene
Aristoteles 37;
Begriffe
37 ff.;
Lebenstypen 39 ff.
Psychologismus, abgelehnt 19.
Rationalismus, sein Recht und
Unrecht
— sein Er-
11 7 ff.;
kenntnisbild 88;
—
sein Geschichts-
— im 19. Jahrhundert 72ff.;
— seine Schranken 74
69;
ff.
Reformation,
ihre
zum
Stellung
Intellektualismus
und zum Volun-
40 ff.;
— ihre Stellung
tarismus
Staatsidee, ihr stärkeres Hervortreten im 19. Jahrhundert 290, 301,
Stufen
und
des Lebens
kennens 102, 350.
des
—
Summierungder Vernunft 296
Verstand,
Ver-
reine
nunft) 86.
Religion,
ff.
System, Bedeutung und Grenze 67.
Teleologie 135ff.;
empirische
—
Teleologie 141.
tisch)
prak-
(s.
37 ff.
Transzendenz, Ausdruck 391.
Typus und typisch, Ursprung des
Ausdrucks 356,
Unendlichkeitsgedanke,
verschiedene
Schätzung
seine
und
Be-
handlung 379.
zur Kunst 332.
(reiner
Er-
Subjektiv, Wort 10;
Wendung
der Gegenwart zum Subjekt 19 ff.
Theoretische Vernunft
118.
Realismus (s. Idealismus) Ausdruck
Rein
Soziale Lebensführung 292ff.
318.
Praktische und theoretii^che
Vernunft, ihre Scheidung durch
bild
164.
Spiritualismus 178ff.
Politismus, 301 ff., 305 ff.
sein
Soziale Gesetze
Vitalismus 144ff.
Volltätigkeit 31.
Bewegung gegen
der Neuzeit 392 ff.;
aufsteigen 395ff.;
sie
in
— ihr Wieder— Forderungen
für ihre gegenwärtige
Lage 398 ff.
Renaissance, ihre Stellung
Kunst und zur Moral 331 ff.
zur
Romantik, ihre Schätzung des Or-
— ihre Entwicklungsidee 204, 215; — ihre Ge-
ganischen 136 ff.;
Voluntarismus, Wort und Problem
37 ff.;
— seine Entwicklung 43.
Von vornherein 86.
Wahrheitsbegriff und Wahrheitsproblem 12ff.; — nicht
bloß
intellektuell
64,
122;
ff.
Weltgeschichtlicher Stand der
Arbeit,
schichtsauffassung 255.
35ff.,
— unsere Fassung 35
als
Maßstab 267.
Schöne Seele 307.
Wert, Erörterung 22ff.
Scholastik, ihre Fassung der Wahr-
Wert des Lebens 376ff.
Wertschätzungen in der Ge-
heit
12 ff.;
— ihre Stellung zum
— Grenze
ihrer Leistung 57; — ihr Mangel
schichte 257.
Werturteile 20ff.
an disjunktiver Kraft 58ff.
Willensfreiheit,
Intellektualismus
40;
Souverän,
souveräne
Art
der
geistigen Arbeit 354 ff.
Sozialdemokratische Bewegung
313ff.
Erörterung
des
Problems 363 ff.
Selektionslehre 208ff.
Wirklichkeitsproblem 79ff.
Wirtschaftliche Frage,
ihre ge-
schichtliche Entwicklung 314 ff.
Wirtschaftliche
Gesetze
104ff.
Sachregister.
410
Zeitgeist, Aufkommen des Begriffs
und Problems 273.
Zivilisation,
drucks 229;
Kultur -231 ff.
Urspnmg
Zufall,
Grenze
seines
Vermögens
nach Aristoteles 132.
des
Aus-
— unterschieden von
Zweck,
sein
Recht
schauung 143 ff.
in
der Weltan-
VEIT & COMP, in Leipzig
Verlag von
GESCHICHTE DER MEÜEREH PHILOSOPHIE
VON NIKOLAUS VON KUES BIS ZUR GEGENWART.
Im Grundriß dargestellt
«
von
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an der Universität Erlangen.
o. Professor
Sechste, verbesserte und ergänzte Auflage,
gr. 8,
geh. 9 M., geb. in Ganzleinen 10 M.
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Großartig durch die
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Fülle des verarbeiteten Stoffes, da nicht allein alle Philosophen von Fach, sondern alle Männer,
deren Denkarbeit bestimmend auf das Geistesleben der modernen Völker eingewirkt hat, Berücksichtigung gefunden haben."
Durch die Gediegenheit des Inhaltes, die geschickte Anordnung und die Klarheit der Darstellung,
durch vorzügliche bibliographische Nachweise hat sich Falckenbergs Geschichte der neueren Philosophie allgemeine Anerkennung erworben, nicht nur in Deutschland, sondern auch im Auslande.
Den Schluß des Werkes bildet eine Erläuterung der wichtigsten philosophischen Kunstausdrücke.
—
GRIECHISCHE^ DENKER,
Eine Geschichte der antiken Philosophie.
Von
Theodor Gomperz.
Vollständig in drei Bänden.
»
Erster
und zweiter Band.
Zweite Auflage.
Lex.
8.
1903.
geh. 23 M, geb. in Halbfranz 28 M.
diesem einzig dastehenden, die Summe seiner durch zahllose Einzelforschungen ausgefüllten Lebensarbeit ziehenden Werke behandelt der große
Gelehrte nicht die Gedanken oder die Systeme oder die Geschichte der griechischen Philosophie, sondern die „Denker", die Individuen. Der Werdegang ihrer
Ideen wird auf dem Hintergrund ihrer Zeit geschildert, und die geistige, Jahrtausende
überspannende Brücke, die sie mit der Gegenwart verbindet, wird geschlagen.
Der dritte Band, dessen erste bis dritte Lieferung erschienen sind, wird
Aristoteles behandeln.
In
GESCHICHTE DES GELEHRTEN UNTERRICHTS
auf den deutschen Schulen
und Universitäten
vom Ausgang des Mittelalters bis zur Gegenv^^art.
Mit besonderer Rücksicht auf den klassischen Unterricht.
Von
Dr. Friedrich Paulsen,
o. ö.
Professor an der Universität Berlin.
Zweite, umgearbeitete und sehr erweiterte Auflage.
Zwei Bände.
1896 u. 1897.
gr. 8.
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:
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Der gelehrte Unterricht im Zeichen des alten Humaniimus. 1450—1740.
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nicht gänzlich fehlgeht,
so wäre hieraus für die Zukunft zu folgern, daß der gelehrte Unterricht bei den
modernen Völkern sich immer mehr einem Zustande annähern wird, in welchem
er aus den Mitteln der eigenen Erkenntnis und Bildung dieser Völker bestritten
wird."
VEIT & COMP, in Leipzig
Verlag von
JOHANN HEINRICH PESTALOZZI.
Seine Ideen in systematischer Würdigung.
Von
Dr. Hermann Leser,
Professor an der Universität Erlangen,
gr. 8.
geh. 3
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M 50 Pf.
VORLESünQEM ÜBER HRTÜRPHILOSOPHIE,
gehalten im
Sommer 1901 an der Universität Leipzig
von
Wilhelm Ostwaid.
Dritte,
Lex. 8.
1905.
vermehrte Auflage.
geh. 12 M, geb. in Halbfranz 14
M 50 Pf.
Die „Vorlesungen über Naturphilosophie" des berühmten Chemikers sind
eine der interessantesten neuzeitlichen Erscheinungen. Die „Vorlesungen" stellen
kein Lehrbuch oder System dar, sondern sind das Ergebnis umfassender Erfahrung bei Forschung und Unterricht, das durch die schöne Form, in der es geboten wird, eine außergewöhnliche Anziehungskraft auf den Leser ausübt.
PSYCHOLOGIE DER NATURVOLKER.
ENTWICKLUNGSPSVCHOLOOISCHE CHARAKTERISTIK
DES NATURMENSCHEN
IN INTELLEKTUELLER, AESTHETISCHER, ETHISCHER UND
RELIGIÖSER BEZIEHUNG.
Eine natürliche Schöpfungsgeschichte des menschl. Vorstellens, Wollens und Glaubens
von
Dr. Fritz Schultze,
ordentlicher Professor der Philosophie an der Technischen Hochschule zu Dresden,
gr. 8.
1900.
geh. 10
M.
WIRTSCHAFT UND RECHT
nach der materialistischen Geschichtsauffassung.
Eine sozialphilosophische Untersuchung
von
Dr. Rudolf Stammler,
o. 5. Professor
an der Universität Halle a. S.
Vitam impendere vero.
Zweite, verbesserte Auflage.
M
Lex. 8.
1906.
geh. 15 M, geb. in Halbfranz 17
50 Pf.
Die Erforschung der allgemein gültigen Gesetzmäßigkeit, unter der das
Der Versoziale Leben des Menschen steht, bildet den Inhalt dieses Werkes.
fasser gelangt zu dem Ergebnis, daß die allgemein gültige Richtschnur für das
dem sozialen Ideal entsprechende Recht nur die Herstellung einer Gemeinschaft von
Menschen sein kann, deren Wollen von bloß subjektivem Begehren frei ist, von
denen vielmehr jeder die objektiven Zwecke der anderen zu den seinigen macht.
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https://www.researchsquare.com/article/rs-3236892/latest.pdf
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Season and depth differences of soil moisture use for tree growth across wet and dry gradients in the Tibetan Plateau
|
Research Square (Research Square)
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Season and depth differences of soil moisture use for tree growth
across wet and dry gradients in the Tibetan Plateau Qian Li
Liang Jiao
(
jiaoliang@nwnu.edu.cn
)
Northwest Normal University
https://orcid.org/0000-0001-6066-2346
Ruhong Xue
Xichen Chen
Peng Zhang
Xuge Wang
Xin Yuan
Research Article
Keywords: Tree-ring oxygen isotopes, Wet and dry gradients, Water use dynamics, Spatial and temporal variation, Tibetan Plateau
Posted Date: August 23rd, 2023
DOI: https://doi.org/10.21203/rs.3.rs-3236892/v1
License:
This work is licensed under a Creative Commons Attribution 4.0 International
License. Read Full License Northwest Normal University
https://orcid.org/0000-0001-6066-2346 1. Introduction Global change affects tree adaptation in a variety of ways, leading to complex changes in forest ecosystem composition, structure and function,
especially in the colder northern regions (Kenneth et al. 2022; Wang et al. 2023). Global changes in tree growth are largely attributed to interactions
between temperature, precipitation and radiation balance (Seddon et al. 2016; Reichstein et al. 2017). In particular, the main factor influencing the
growth of trees in humid environments is temperature, while the growth of trees in arid areas is mainly limited by moisture (Amer et al. 2023). Thus, there are differences in the sensitivity of tree growth to climate change under different wet and dry conditions. Soil water is an important component of the water cycle that links the atmosphere, hydrosphere and biosphere, and is a major source of water for
plants (Eslaminejad et al. 2020), which plays an important role in maintaining vegetation growth and ecosystem productivity. Soil moisture
content varies with soil depth and season (Ma et al. 2020), and the water uptake mechanisms and adaptation strategies of trees to heterogeneous
environments vary especially under different hydrological changes (Bertrand et al. 2014). The soil moisture was found to be the most important
potential factors controlling the growth of vegetation on the Tibetan Plateau (Wang et al. 2018). The trees mainly use soil moisture during the
growing season (Zeng et al. 2020; Zhang et al. 2022). However, it is important to note that the measurement and estimation of soil moisture is very
complex and has great limitations for studying tree growth and water use strategies. Whereas tree rings can be used as a proxy for inversion of
soil moisture changes on long time scales due to their continuous record, accurate dating, sensitivity to climate change, and wide distribution (Gao
et al. 2018). For example, the oxygen isotopes in tree-ring have been used to reconstruct soil moisture in the western Himalayas over a period of
nearly 100 years (Bose et al. 2016). Due to the coupled processes of water uptake and transpiration by tree roots, interannual variation in tree-ring oxygen stable isotopes (δ18OTR) can
document seasonal changes in climate and the physiological-ecological response of trees to seasonal changes (Xu et al. 2020). 1. Introduction Variations in
δ18OTR are mainly influenced by δ18O uptake of source water by growing trees, evaporative enrichment of leaf water, biochemical fractionation
during photosynthesis, and xylem water exchange during the transport of carbohydrates to the trunk (Roden et al. 2000). This series of
fractionation processes is regulated by external climate change (Brinkmann et al. 2019; Xiao et al. 2020). The δ18OTR are derived from soil water,
which is ultimately derived from atmospheric water, mainly in relation to changes in precipitation, temperature and relative humidity (De et al. 2018). And changes in the atmospheric climate system are stored in soil moisture and are transferred to tree cellulose during the growing season
to affect tree-ring oxygen isotopes. In addition, δ18OTR has the potential to reflect the depth of water uptake (Gessler et al. 2014). Variations in soil
water δ18O at different depths are preserved in the tree rings, but the strength of the soil water isotope signal preserved in the tree rings depends on
the depth of water uptake by tree roots (Huang et al. 2019, Xu et al. 2020). In monsoon-controlled areas, precipitation δ18O has a seasonal pattern
and these signals are transmitted to various soil depths, causing soil water δ18O to vary in vertical gradients with seasonal strength and direction
(Lindh et al. 2014). Trees convert the isotopic composition of source water to the isotopic composition of cellulose through complex non-linear
physicochemical processes (Ballantyne et al. 2016). For example, it has been shown that trees in the East Asian monsoon zone significantly
correlated with soil moisture during the growing season, but the seasonal response was earlier in wet areas than in dry areas, and wet areas used
mainly shallow soil water while dry areas used deep soil water (Wang et al. 2021). However, other study also showed trees in the East Asian
monsoon region used deep water in the dry season and shallow soil water in the non-dry season (Yang et al. 2015). Therefore, there is uncertainty
in the study of water use strategies based on δ18OTR. Controlled by the Asian monsoon, the climate of the Tibetan Plateau has undergone some changes in recent years, mainly including increasing
surface temperatures (Sun et al. 2022), weakening monsoons (Wong et al. 2020) and reduced potential evaporation (Fang et al. 2019) since the
1960s. Research Article Page 1/20 Abstract Soil moisture has an important influence on tree growth in climate-sensitive regions. However, the seasonality of soil moisture utilization by trees
in the Tibetan Plateau, a typical climate-sensitive region, and the mechanisms of depth of uptake are still unclear. Therefore, the spatial and
temporal differences in the soil moisture use dynamics of tree growth were analyzed under wet and dry gradients on the Tibetan Plateau using
tree-ring δ18O (δ18OTR). The results showed that: 1) soil moisture during the growing season was the main influencing factor on tree growth under
different wet and dry gradients on the Tibetan Plateau. 2) The response of δ18OTR to soil moisture had a lag in arid areas than in wet areas. 3) In
wet areas, trees absorbed the soil moisture in surface, while absorbed the soil moisture in deep in arid areas. And meanwhile, trees could develop
more lateral and deep root systems using soil moisture from all soil layers to cope with climatic stress. In the future, more lateral and deep root
systems of trees will be better suited to survive in complex habitats on the Tibetan Plateau, and that monitoring of trees in single water use areas
should be enhanced. Further, it is of great significance to study soil water use strategies of tree growth under different wet and dry gradients for
predicting forest ecosystem changes in complex environments. 2.1 Study areas The six sites in the Tibetan Plateau were selected for the study such as Delingha region (DE), the Ganesh (GA) and Wache (WA) regions in the
south, the Manali (MA) and Jageshwar (JA) regions in the southwest, and the Karakorun (KA) region in the west (Fig. 1). Except for the KA region
which is affected by the westerly winds and the other regions are affected by the Asian summer winds. From the wettest JA region to the driest DE
region, the annual precipitation decreased by 1,311 mm (Fig.S1). The six regions are divided into humid and arid areas based on annual
precipitation. Specifically, the humid areas include the JA, WA and GA regions, while the arid areas include the MA, KA and DE regions. Please insert Fig. 1 about here. 1. Introduction Changes in summer wind activity affect monsoon precipitation by altering the spatial pattern of regional water vapor transport, ultimately
leading to interannual variability in δ18OTR (Ge et al. 2017; Tan et al. 2018). However, current studies on the Tibetan Plateau based on δ18OTR have
mainly focused on climate reconstruction (Grießinger et al. 2019; Xu et al. 2019) and physiological mechanisms (Zhang et al. 2018). Therefore,
this study aims to address three main questions: (1) whether there are differences in the seasonal dynamics of tree water use under different wet Page 2/20 and dry gradients on the Tibetan Plateau, (2) whether there are differences in the ability of tree growth to use water at different soil depths under
different wet and dry gradients, (3) and what are the combined adaptation mechanisms of trees based on the seasonal dynamics and soil depth of
tree water use under different wet and dry gradients? and dry gradients on the Tibetan Plateau, (2) whether there are differences in the ability of tree growth to use water at different soil depths under
different wet and dry gradients, (3) and what are the combined adaptation mechanisms of trees based on the seasonal dynamics and soil depth of
tree water use under different wet and dry gradients? 2.2 Tree-ring isotope data The δ18OTR data in this study were downloaded from NOAA Paleoclimatology Datasets (https://www.ncei.noaa.gov/pub/data/paleo/tree-
ring/isotope/asia). These δ18OTR chronologies are common to the years 1948–1998, and the specific locations, elevations, tree species, and
δ18OTR chronological lengths of the sampled sites were shown in Table 1. The sampling sites were largely located in high-elevation areas, and all
of the selected tree species were conifers. Tree cores were collected at breast height using increment borers. And alpha-cellulose extraction process
included chemical reactions with organic reagents, sodium chlorite and sodium hydroxide to remove extractions (ash, resin, etc.), lignin and hemi-
cellulose, respectively (Shi et al. 2015). Elevation gradients and age differences in the plateau had no significant effect on the climatic response of
δ18OTR records (Wernicke et al. 2015; Zeng et al. 2016). Please insert Table 1 about here. Table 1
Chronological information on tree rings
Sites
Location
Elevation
Species
period
Rbar
Date Source
JA
29°38′N,
79°51′E
1870m
Cedrus deodara
1621–2008
0.61–0.78
Xu et al (2018)
WA
27°59′N,
90°00′E
3500m
Larix griffithii
1743–2011
0.74–0.88
Sano et al (2013)
GA
28°10′N,
85°11′E
3500m
Abies spectabilis
1801–2000
0.56–0.78
Xu et al (2018)
MA
32°13′N,
77°13′E
2700m
Abies pindrow
1768–2008
0.71–0.92
Sano et al (2017)
KA
35°54′N,
74°56′E
2900m
Juniperus excelsa
1990–1998
0.51
Treydte et al (2006)
DE
37°48′N,
97°78′E
3500m
Juniperus przewalskii
-4680-2011
0.40–0.76
Yang et al (2021) 2.4 Soil moisture data Soil moisture data were obtained from the Global Land Data Assimilation System (http://Idas.gsfc.nasa.gov/gldas) with a temporal resolution of
1948–2015 and a spatial resolution of 0.25°×0.25°. This dataset has a longer temporal resolution and a higher resolution for soil moisture at
different depths compared to other datasets. We selected soil moisture data from 1948–1998 at three depths with 0-10cm, 10-40cm and 40-
100cm for our study. The soil moisture data we downloaded ((unit: kg/m2) had to be further converted to volumetric water content (VWC, unit:
m3/m3) for data analysis by the following conversion equation: (Eq. 4)
V WC = × × ×
kg
m2
m3
1000kg
1000mm
1m 1thicknessoflayerinmm 2.3 Meteorological data Our meteorological data included the maximum, mean and minimum temperatures, actual vapor pressure, and precipitation and the Palmer
drought severity index (PDSI) on monthly-scale for six regions (Fig.S1). The grid point data above were obtained from Climate Explorer
(http://climexp.knmi.nl/start.cgi). Since saturation water vapor pressure deficit (VPD) and relative humidity (RH) data for the region are difficult to
obtain, we calculated VPD (Murray 1966) and RH (New et al. 1999) based on the downloaded CRU TS4.04 (0.5◦×0.5◦) dataset, and the specific
calculation procedure as follows: Page 3/20 Page 3/20 (Eq. 1)
(Eq. 2)
(Eq. 3)
V PD = actualwatervapourpressure −saturatedwatervapourpressure = e0 × 10
( ) −actualwatervaporpressure
7.5T
T +237.3
RH = ( ) × 100
eesat
esat = 6.108 × exp(17.27 × T/(T + 273.3) RH = ( ) × 100
eesat In the above equation, e0 = 6.11hpa, T is the monthly mean temperature (unit: ℃). In the above equation, e = 6.11hpa, T is the monthly mean temperature (unit: ℃). 2.4 Soil moisture data 2.5 Data Analysis To investigate the elements of environmental factors that mainly affect δ18OTR, we used Pearson correlation as well as partial correlation analysis. In the meanwhile, One-way analysis of variance (ANOVA) was used to assess the variability of the δ18OTR chronology and the significance of
differences was tested using Tukey's multiple comparisons. Spatial correlation analysis was performed using Matlab 2023 and mapped using
ArcGIS 10.8. Please insert Fig. 4 about here. 3.1 Comparison of the spatial and temporal dynamics of δ18OTR under the wet and
dry gradients Please insert Fig. 2 about here. The spatial dynamics of δ18OTR showed an overall higher value in the north and lower value in the south (Fig. 3). The lowest value of δ18OTR
occurs in the southern part of the plateau (WA, 19.401‰) and the highest value is located in the northeastern part of the plateau (DE, 32.907‰). The value of δ18OTR decreased as the climate changed from arid to humid areas. Please insert Fig. 3 about here. g
3.3 Comparison of δ18OTR response to climatic and soil moisture factors under wet
and dry gradients 3.3 Comparison of δ18OTR response to climatic and soil moisture factors under wet
and dry gradients It is revealed that overall δ18OTR responded negatively to PDSI, precipitation and RH, and positively to temperature and VPD in different humid and
arid areas (Fig. 6). However, seasonal response differences were observed in humid and arid environments, showing δ18OTR more often correlated
with environmental factors in the middle of the growing season (June-August) in humid environments, and with environmental factors in the
middle and late growing season (July- September) in arid areas. Further, comparing the responses of δ18OTR to soil water content at different soil depths in humid and arid areas showed variability in depth as
well as time (Fig. 7). In general, the correlation was greater in wet areas with 0–10 cm and 10–40 cm in July (p < 0.01), while the arid environment
δ18OTR was significantly negatively correlated with soil moisture in 10–40 cm and 40–100 cm soil layers in September (p < 0.01). The seasonal
response of δ18OTR to soil moisture was lagged in the dry areas compared to the humid areas. Specifically, in humid areas, δ18OTR was
significantly negatively correlated with 0–40 cm soil moisture in July and August in JA and WA (r=-0.49 to -0.41, p < 0.05), while δ18OTR in GA
region was significantly negatively correlated with soil moisture at all three soil depths (r=-0.44 to -0.26, p < 0.05). However, in arid environments,
δ18OTR was significantly negatively correlated with soil moisture at all three soil depths from July to September in MA (r=-0.47 to -0.31, p < 0.05),
while δ18OTR in KA and DE regions were more significantly negatively correlated with soil moisture at 40–100 cm from July to September (r=-0.48
to -0.29, p < 0.05). Please insert Fig. 6 about here. 3.2 Comparison of the spatial and temporal dynamics of soil water content patterns at different depths under wet and dry gradients Soil water content at different gradients on the Tibetan Plateau showed inconsistent soil depth variation characteristics and temporal dynamics
(Fig. 4). Specifically, the soil water content in the six gradient regions showed the increasing trends with increasing soil depth from 0–10 cm, 10–
40 cm to 40–100 cm (Fig. 4a). At the same time, the soil moisture content in the wet areas was significantly greater than that in the arid areas,
accompanied by a greater variation in moisture content (VWCJA, WA, GA > VWCMA, KA, DE), especially in the DE area, where the moisture content of all
soil layers is lower than in the other areas. In terms of time, the seasonal dynamics of soil moisture content were more pronounced in the wetter areas than in the arid areas (Fig. 4b), where
the peak soil moisture content was mainly concentrated in June-September. The volumetric water content from July to September was 48.9% (JA),
49.1% (WA) and 47.8% (GA) of the year, respectively. However, the seasonal dynamics of soil moisture content in the arid areas varied more
between regions. In the MA region with the double-peaked model, the volumetric water content from June to September accounts for 32.8% of the
year. However, the high values of s volumetric water content in the KA region are concentrated in April-June, accounting for 32.3% of the year. But
no seasonal dynamic peaks in soil moisture content were formed in the driest DE region. Please insert Fig. 4 about here. Please insert Fig. 4 about here. Page 4/20 Page 4/20 The moisture content of each soil layer in the humid areas was significantly greater than in the arid areas and decreases significantly with the
decrease in regional precipitation (Fig. 5). Specifically, in the humid areas, the soil moisture in each soil layer in each regions tends to decrease and
the most significant decrease was in the 40–100 cm range. In the arid areas, soil moisture showed an overall increasing trend, but the MA area
shows a decreasing trend in all soil layers, which is closely related to the weakening of the monsoon in the region and the decrease in precipitation. Meanwhile, the linear fit of precipitation to soil moisture was stronger in the wetter regions than in the arid regions, with the fit becoming
progressively worse from the wettest JA region to the driest DE region (Fig. S2). Please insert Fig. 5 about here. Please insert Fig.7 about here. 3.5 Comparison of spatial characteristics of δ 18 O TR response to soil moisture content at different soil depths under wet and dry gradients 3.5 Comparison of spatial characteristics of δ 18 O TR response to soil moisture content at different soil depths under wet and dry gradients 3.5 Comparison of spatial characteristics of δ 18 O TR response to soil moisture content at different soil depths under wet and dry gradients The spatial correlation analysis revealed that the response of δ18OTR to soil moisture at all six sites in the study area was generally negatively
correlated. But the overall differences in the responses of δ18OTR to soil moisture content at 0–10 cm and 10–40 cm were not significant, and the
significant difference was found at 40–100 cm (Fig. 8). And the responses of δ18OTR to soil moisture in the southern and northeastern parts of the
plateau changed from negative to positive as the soil depth deepened (Fig. 8). Specifically, δ18OTR was negatively correlated with soil moisture at
0–10 cm of all sites. But the correlation showed a trend of JA > WA > GA > MA > KA > DLH. That is, the drier the climate, the lower the correlation
between δ18OTR and soil moisture at 0–10 cm and 10–40 cm. However, the positive responses of δ18OTR to soil moisture at the depth of 40–100
cm increases in DE of the northeastern part and in WA of the south. Please insert Fig.8 about here. 3.4 Correlation between δ 18 O TR and key environmental factors during the growing season from June to September under the wet and dry
gradients The VMC0-10 represents the volume moisture content of 0-10cm soil, the VMC10-40
represents the volume moisture content of 10-40cm soil, he VMC40-100 represents the volume moisture content of 40-100cm soil. Please insert Table 2 about here Please insert Table 2 about here 3.4 Correlation between δ 18 O TR and key environmental factors during the growing season from June to September under the wet and dry
gradients 3.4 Correlation between δ 18 O TR and key environmental factors during the growing season from June to September under the wet and dry
gradients 3.4 Correlation between δ 18 O TR and key environmental factors during the growing season from June to September under the wet and dry
gradients The correlation between δ18OTR and soil moisture during the growing season (June-September) was found to be significantly stronger than other
environmental variables through Pearson correlation analysis. Therefore, we further selected key environmental factors that strongly influenced
δ18OTR in different regions during the growing season for a partial correlation analysis (Table 2). The results showed that when VPD and RH were
controlled, δ18OTR was significantly negatively correlated with 0–10 cm soil moisture in the wet environment in JA (p < 0.05), while it was
significantly correlated with 0–10 cm and 10–40 cm soil moisture in WA and GA (p < 0.05). When VPD and RH were controlled, δ18OTR was highly
significantly negatively correlated with soil moisture at all soil depths in MA (p < 0.01), while δ18OTR was significantly negatively correlated with soil
moisture at 40–100 cm in KA and DE (p < 0.05). Please insert Fig.7 about here. Page 5/20 Page 5/20 Table 2 Partial correlation analyzes between δ18OTR and environmental factors from June-September. Control variables
Correlations with δ18OTR
JA
WA
GA
MA
KA
DE
VPD
VMC0 − 10
-0.396🞶🞶
-0.364🞶
-0.409🞶
-0.459🞶
-0.211
-0.145
VMC10 − 40
-0.265
-0.36🞶
-0.383🞶
-0.472🞶
-0.277
-0.286
VMC40 − 100
-0.21
-0.225
-0.241
-0.416🞶
-0.356🞶
-0.385🞶
PDSI
-0.161
-0.106
-0.255
-0.264
-0.278
-0.585🞶🞶
RH
0.27
0.179
0.423🞶
0.151
-0.064
-0.163
PDSI
VMC0 − 10
-0.251
-0.459🞶
-0.365🞶
-0.527🞶🞶
-0.004
0.173
VMC10 − 40
-0.213
-0.451🞶
-0.317
-0.542🞶🞶
-0.118
0.164
VMC40 − 100
-0.14
-0.156
-0.343
-0.5🞶🞶
-0.239
0.119
VPD
0.057
0.432
-0.141
-0.314
0.003
-0.021
RH
0.072
-0.396🞶
0.251
-0.028
-0.042
-0.022
RH
VMC0 − 10
-0.356🞶🞶
-0.377🞶
-0.392🞶
-0.53🞶🞶
-0.213
-0.13
VMC10 − 40
-0.289
-0.38🞶
-0.365🞶
-0.542🞶🞶
-0.276
-0.274
VMC40 − 100
-0.214
-0.225
-0.28
-0.499🞶
-0.352🞶
-0.365🞶
VPD
0.296
0.259
0.367🞶
0.159
-0.042
-0.008
PDSI
-0.208
-0.113
-0.242
-0.315
-0.277
-0.57🞶🞶
dicated p < 0.05 and p < 0.01, respectively. The VMC0-10 represents the volume moisture content of 0-10cm soil, the
olume moisture content of 10-40cm soil, he VMC40-100 represents the volume moisture content of 40-100cm soil. Note: * and ** indicated p < 0.05 and p < 0.01, respectively. 2 Difference strategies of season in soil moisture use dynamics for tree growth under different wet and d In climate-sensitive areas, soil moisture is critical to tree growth. A variety of factors can alter soil water uptake by trees, such as season (Rao et al. 2020), life history stage (Van et al. 2017), tree species (Schwendenmann et al. 2015), water availability and depth of plant access to water (Rao et
al., 2020), and the amount of recent rainfall (Xu et al. 2011). In our study, δ18OTR was found to be significantly correlated with soil moisture during
the growing season, but the seasonal response in arid and wet zones showed inconsistency, with tree-ring δ18O responding to soil moisture later in
arid zones than in wet zones (Fig. 7). Specifically, in the humid areas, we found that δ18OTR significantly negatively correlated with soil moisture from June to August ((Fig. 7). Because
the three humid regions are influenced by the Indian summer winds and receive more precipitation from June to August, which further affects
δ18OTR by influencing soil water 18O through infiltration. The JA and GA regions showed a significant negative correlation between δ18OTR and
July-September precipitation, Indian monsoon index, and monsoon circulation intensity in the region (Xu et al. 2018). And we also found a
significant negative correlation between δ18OTR and soil moisture during July-August in JA and GA regions. And meanwhile, δ18OTR values of
different tree species in the WA region were mainly influenced by summer precipitation since 1743 (Masaki et al. 2012). In our study, we also found
that δ18OTR in the WA region was significantly negatively correlated with soil moisture in July and August. We suggest that summer winds bring
abundant precipitation, and that plant roots take up water in the soil through a series of transpiration infiltration processes, finally retaining the
isotopic signal of precipitation in the xylem of trees expressed through δ18OTR in humid regions. We also found the response of δ18OTR in arid regions significantly correlated with soil moisture from July to September, especially in September
(Fig. 7). But the response of δ18OTR to soil moisture in arid regions was lagged compared to wet regions, which may be closely related to the
extended growing season of trees due to warming and humidification of the Tibetan Plateau in recent decades (zhang et al. 2021). However, the
specific response varied among the three arid regions. 4.1 The δ 18 O TR records soil water use by tree growth in different regions of the Tibetan Plateau 4.1 The δ 18 O TR records soil water use by tree growth in different regions of the Tibetan Plateau Since the 1960s, the surface temperature of the Tibetan Plateau has shown a continuous warming trend (Duan et al. 2015), and precipitation has
increased in the western and eastern parts of the plateau (Bao et al. 2019; Liu et al. 2020), but weakened in the southwestern part of the plateau
(Hu et al. 2021). In such a climatic change context, trees may adopt different water use strategies to maintain their growth. It has been shown that
δ18OTR could reflect the seasonality and the depth of plant water uptake (Gessler et al. 2014). This means that information on the variation of soil
water δ18O at different seasons and depths is stored and recorded in the tree-ring (Huang et al. 2019, Xu et al. 2020). Page 6/20 Page 6/20 The δ18OTR variations are mainly regulated by a combination of the oxygen isotope composition of the source water, and the external climate that
affects fractionation processes within the tree (Roden et al. 2000). In our results, the δ18OTR was more sensitive to moisture factors than to
temperature factors, and in particular δ18OTR showed the significant negative response to soil moisture during the growing season (Fig. 5). The
δ18OTR is derived from soil water, but that soil water δ18O varies with soil depth (Liu et al. 2017), which is consistent with our findings. In addition,
water enters the leaves from the soil through the xylem and then the lighter isotope (16O) evaporates more readily than the heavier isotope (18O),
leading to an enrichment of δ18O in leaf water (Farquhar et al. 1993). Trees take up soil water directly through the root system and retain the
isotopic signal in soil water in the xylem through a series of intra-tree fractionation processes. Therefore, the δ18OTR is a true reflection of soil water
use by trees based on the processes of water uptake, transport and dissipation in trees. At the same time, our study found that δ18OTR values were significantly greater in arid than in humid areas (Fig. 3), due to the fact that
transpiration affects leaf water enrichment by replenishing unenriched soil water to the leaves. 4.1 The δ 18 O TR records soil water use by tree growth in different regions of the Tibetan Plateau On the one hand, the δ18OTR is mainly negatively
correlated with stomatal conductance, and reduced soil moisture and humidity enhance evaporative enrichment of leaf and soil water through
effects on stomatal conductance and the ratio of internal leaf CO2 to atmospheric CO2 in arid environments (Mirfenderesgi et al. 2016). On the
other hand, soil water evaporation is enhanced in arid environments and the source water absorbed by tree roots becomes heavier, leading to an
enrichment of oxygen isotopes in leaf water (Roden et al. 2000). Sucrose produced by photosynthesis is transported through the bast to the xylem
of the trunk for cellulose synthesis, and during the conversion to cellulose, oxygen isotopes in the sugar are partially exchanged with oxygen
isotopes in the source water (Zhan et al. 2021), resulting in a mixed signal of both source water and transpiration-enriched oxygen isotopes in the
cellulose oxygen isotope ratio of the tree-ring (Giraldo et al, 2022). These two processes result in a higher δ18OTR in arid areas than in humid areas. Similarly, a moist atmosphere reduces transpiration and thus 18O water enrichment in needles, further leading to lower δ18OTR in humid areas than
in arid areas. And precipitation δ18O is inversely related to precipitation, more precipitation leads to lower precipitation δ18O values, resulting in
weaker δ18O enrichment of water from water sources absorbed by trees, both of which make δ18OTR in wet areas lower than in arid areas (Yang et
al. 2012). 2 Difference strategies of season in soil moisture use dynamics for tree growth under different wet and d The δ18OTR in the MA region was controlled by hydroclimatic variables such as summer
monsoon precipitation, RH and PDSI (Masaki et al. 2017). Those results suggesting that monsoon precipitation has a significant effect on tree
growth. Tree growth in the KA region is mainly influenced by winter snowfall, and although trees are still affected by precipitation δ18O during
summer precipitation, the melting snowfall provides a more stable water source, resulting in a much lower correlation between δ18OTR and summer
soil moisture in this region than in the other two regions (Kerstin et al. 2006). In the driest DE region, the δ18OTR were controlled mainly through soil
moisture and precipitation variability from May to September (Yang et al. 2006), and our study provided further evidence that δ18OTR in this region
was significantly negatively correlated with soil moisture in September. Page 7/20 Page 7/20 Page 7/20 In the meantime, it has been found that precipitation has a greater effect on soil moisture than temperature on the southeastern Tibetan Plateau. Soil moisture is significantly correlated with oxygen isotope growing season in the southeastern Tibetan Plateau trees (Wang et al. 2023).And the
δ18OTR were found to be significantly negatively correlated with growing season precipitation in the United Kingdom (Giles et al. 2015), southern
Mexico (Brienen et al. 2013), northwestern Tibetan Plateau (Zhang et al. 2021), and Ordos Plateau (Li et al. 2019), which was broadly in line with
our findings that summer rainfall to supplement plant water availability during the growing season (Hahm et al. 2019). We speculated that this
were two reasons for the lagging response of δ18OTR to soil moisture in arid areas relative to humid areas. On the one hand, it might be because
the Indian summer winds, which firstly reached the southern region, making precipitation events occur earlier in the southern region in their
northward push. Therefore, they affect the tree growth process through a series of fractionation as well as plant physiological processes, then
reflecting different moisture. On the other hand, this is closely related to the warming and humidification in the western and northern parts of the
plateau, where warmer temperatures and increased precipitation lead to an earlier and longer growing season for trees. 4.3 Difference strategies of depth in soil moisture use dynamics for tree growth under different wet and dry gradients on the Tibetan Plateau Precipitation on the Tibetan Plateau, where the natural environment is extremely fragile and sensitive, is highly spatially variable with annual
precipitation decreasing gradually with latitude (Li et al. 2022). Therefore, trees might maintain their growth by altering their water use strategies to
absorb soil moisture at different depths under different precipitation conditions. Most studies have shown that trees in arid environments tend to
absorb deeper water to resist drought (Hu et al. 2021), whereas trees in wet environments tended to use shallow soil water (Holdo et al. 2018). While this was supported in our study, we have also found that trees in complex climatic environments may develop complex root systems with
both lateral and deep roots, adjusting their water use strategies to maintain their growth in response to environmental changes. Our study also found that, the δ18OTR in the JA region was significantly negatively correlated with 0–10 cm soil moisture in a wet environment,
while that in the WA and GA regions were significantly negatively correlated with 0–10 cm and 10–40 cm soil moisture after controlling the VPD
and RH (Table 2), it suggesting that trees in humid areas tended to take up soil moisture in the surface and middle layers. Other studies also found
shallow soil water (35%) and precipitation (32.5%) were the main sources of water for trees in wet areas when annual precipitation was greater
than 800 mm (Zhou et al. 2011). Meanwhile, the topsoil in humid areas is the most active area for microbial activity, so that it provides nutrients to
the trees (Priyadarshini et al. 2016). For example, it has been shown that trees had a relatively large range of source water and a good water use
strategy in the shallow soil under humid climate conditions (Sun et al. 2019). In arid environments, the significant negative correlation between δ18OTR and 40–100 cm soil moisture in the KA and DE regions was revealed
after controlling VPD and RH (Table 2), it suggesting that trees in arid environments utilize more stable deep soil moisture. And trees in DE region
preferred to take up deep soil moisture through a study of cypress in the northeastern Tibetan Plateau (Grossiord et al. 2016), which validates our
results. 4.3 Difference strategies of depth in soil moisture use dynamics for tree growth under different wet and dry gradients on the Tibetan Plateau In arid environments, capillary water induces trees to form deeper root systems to absorb deep soil water and groundwater to meet higher
transpiration needs during the dry season, while deep root systems provide a more consistent and reliable source of water for trees (Lindh et al. 2014). But we also found the highly significant negative correlations between δ18OTR and soil moisture at all soil depths in MA, and there might be
two factors. On the one hand, soil moisture δ18O fractionation occurs when precipitation infiltrates into the soil (Oerter et al. 2014), creating an
isotopic gradient in the vertical profile while vigorous evaporation may mask some of the isotopic signal (Dai et al. 2015). On the other hand, it is
possible that the 'bimodal' precipitation pattern in this region encourages trees to develop lateral and deep root systems, using the surface root
system to obtain water from the upper soil layer when precipitation is adequate. The deep root system is used to obtain water from deep soil or
groundwater when precipitation is insufficient (zhang et al. 2017). This confirms our finding that trees adjust their water use strategies in time to
sustain self-sustaining growth (Zhao et al. 2018; Nolan et al. 2018). Trees under different environmental conditions have different coping strategies to climate extremes, and the resilience of trees to drought is closely
related to regional soil moisture, diurnal temperature differences, and plant physiological characteristics (Fang et al. 2018). It has been reported
that tree growth on the Tibetan Plateau is more vulnerable to challenges in the context of climate change (Zhang et al. 2021; Yao et al. 2023). Our
results suggest that trees in arid areas are better able to cope with drought stress due to their use of more stable deep soil water, whereas trees in
humid areas may be more vulnerable to drought stress in the context of climate variability due to their use of surface-middle soil water. As a result,
trees in wetter areas will face greater challenges to survival in the context of a weakening southwestern monsoon with reduced precipitation and
continued warming temperatures. Moreover, it is noteworthy that trees develop the water use strategy of absorbing moisture from different soil
layers with the stronger lateral and deep root systems as the environment changes, which can more be beneficial to trees in coping with complex
environment. 2 Difference strategies of season in soil moisture use dynamics for tree growth under different wet and d Moreover, precipitation in
early in the growing season was not replenished in time, while both precipitation and temperature at the end of the growing season promoted tree
growth, making the response of δ18OTR to soil moisture slightly later in the arid areas than in the humid areas (Guo et al. 2022). 5. Conclusion Page 8/20
The climate of the Tibetan Plateau has changed significantly in recent decades. It is of great significance to study the water use strategies of trees
in this region for monitoring its ecological environment. Therefore, the relationship between oxygen isotope data and environmental factors in tree Page 8/20 ring on the Tibetan Plateau were investigated. We found that δ18OTR recorded spatial and temporal differences in soil moisture use dynamics by
tree growth under different wet and dry gradients. Our results showed that δ18OTR responded more strongly to soil moisture than other
environmental factors during the growing season under different wet and dry gradients, but the response in the arid areas had a lag compared to
the humid areas. Meanwhile, trees in the humid southwestern region absorbed more shallow soil water, while trees in the arid western and
northeastern regions absorbed more deep soil water. We suggest that trees use shallow soil water alone increase the risk of drought stress in a
southwestern context of decreasing precipitation and increasing temperatures, whereas trees develop both lateral and deep root systems are better
able to survive in complex environments. As a water tower in Asia, the Tibetan Plateau has an important influence on the climate of the Northern
Hemisphere. We recommend the use of this method for larger-scale and more regional studies to research the response strategies of trees and
ecological changes in the context of climate change. ring on the Tibetan Plateau were investigated. We found that δ18OTR recorded spatial and temporal differences in soil moisture use dynamics by
tree growth under different wet and dry gradients. Our results showed that δ18OTR responded more strongly to soil moisture than other
environmental factors during the growing season under different wet and dry gradients, but the response in the arid areas had a lag compared to
the humid areas. Meanwhile, trees in the humid southwestern region absorbed more shallow soil water, while trees in the arid western and
northeastern regions absorbed more deep soil water. We suggest that trees use shallow soil water alone increase the risk of drought stress in a
southwestern context of decreasing precipitation and increasing temperatures, whereas trees develop both lateral and deep root systems are better
able to survive in complex environments. As a water tower in Asia, the Tibetan Plateau has an important influence on the climate of the Northern
Hemisphere. 5. Conclusion We recommend the use of this method for larger-scale and more regional studies to research the response strategies of trees and
ecological changes in the context of climate change. Data Availability Statemen All data sources analyzed in this study are included in this paper. Conflicts of Interest The authors declare no conflict of interest. The authors declare no conflict of interest. Funding This research was supported by CAS “Light of West China” Program (2020XBZG-XBQNXZ-A), Basic Research Innovation Group Project of Gansu
Province (No. 22JR5RA129), and the 2022 Major scientific Research Project Cultivation Plan of Northwest Normal University (WNU-LKZD2022-04). We also thank the anonymous referees for helpful comments on the manuscript. References 1. Amer A, Franceschi E, Hjazin A (2023). Structure and Ecosystem Services of Three Common Urban Tree Species in an Arid Climate City. Forests 14(4): 671 2. An W., Hou S, Zhang Q, Zhang W, Wu, S, Xu H, Liu Y (2017) Enhanced recent local moisture recycling on the northwestern Tibetan Plateau
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growing seasons lengthens in a cold and arid environment. Catena 196: 104936 Page 12/20 Page 12/20 88. Zhang J, Gou X, Manzanedo RD, Zhang F, Pederson N (2018) Cambial phenology and xylogenesis of Juniperus przewalskii over a climatic
gradient is influenced by both temperature and drought. Agricultural and forest meteorology 260: 165-175 88. Zhang J, Gou X, Manzanedo RD, Zhang F, Pederson N (2018) Cambial phenology and xylogenesis of Juniperus przewalskii over a climatic
gradient is influenced by both temperature and drought. Agricultural and forest meteorology 260: 165-175 89. Zhou Y, Chen S, Song W, Lu Q, Lin G (2011) Water-use strategies of two desert plants along a precipitation gradient in northwestern China. Chinese Journal of Plant Ecology 35(8): 789-80 89. Zhou Y, Chen S, Song W, Lu Q, Lin G (2011) Water-use strategies of two desert plants along a precipitation gradient in northwestern China. Chinese Journal of Plant Ecology 35(8): 789-80 Figures Figures
Figure 1
Geographical locations of study area and sampling sites. The green line in the figure represents the boundary line of the Tibetan Plateau, the green
dots are the tree wheel sampling points, the different shades of blue represent the different amounts of rainfall in the region, and the darker the
color means more precipitation. Figure 1 Figure 1 Geographical locations of study area and sampling sites. The green line in the figure represents the boundary line of the Tibetan Plateau, the green
dots are the tree wheel sampling points, the different shades of blue represent the different amounts of rainfall in the region, and the darker the
color means more precipitation. Page 13/20 Page 13/20 Page 13/20 Figure 2
The time series of the δ18OTR for each site. The red solid line indicates the trend for each chronology. Figure 2 The time series of the δ18OTR for each site. The red solid line indicates the trend for each chronology. Page 14/20 Page 14/20 Page 14/20 Figure 3
Significance test for study sites. The box in the figure represents the average value of δ18OTR for each point from 1948 to 1998. Inter-sample
differences in δ18OTR values were assessed using one-way ANOVA, with different letters indicating significant differences at the 0.05 confidence
level. And the red dashed line represents the mean value of δ18OTR in the arid areas and the blue dashed line represents the mean value of δ18OTR
in the humid areas. Figure 3 Significance test for study sites. The box in the figure represents the average value of δ18OTR for each point from 1948 to 1998. Inter-sample
differences in δ18OTR values were assessed using one-way ANOVA, with different letters indicating significant differences at the 0.05 confidence
level. And the red dashed line represents the mean value of δ18OTR in the arid areas and the blue dashed line represents the mean value of δ18OTR
in the humid areas. Page 15/20 Figure 4
Changes of soil moisture at different soil layers and annual scales. The squares in the box line diagram represent the median volumetric water
content, the black dots represent the outliers and the range of the boxes represents the 25%-75% data interval (a). The blue line in the graph
represents the 40-100 cm volumetric water content, the red line represents the 10-40 cm volumetric water content and the black line represents the
0-10 cm volumetric water content (b). Figure 4
Changes of soil moisture at different soil layers and annual scales. The squares in the box line diagra
content, the black dots represent the outliers and the range of the boxes represents the 25%-75% data
represents the 40-100 cm volumetric water content, the red line represents the 10-40 cm volumetric w
0-10 cm volumetric water content (b). Figure 1 Figure 4 Changes of soil moisture at different soil layers and annual scales. The squares in the box line diagram represent the median volumetric water
content, the black dots represent the outliers and the range of the boxes represents the 25%-75% data interval (a). The blue line in the graph
represents the 40-100 cm volumetric water content, the red line represents the 10-40 cm volumetric water content and the black line represents the
0-10 cm volumetric water content (b). Page 16/20 Figure 5
Interannual variation of soil moisture. The blue line represents 40-100cm volumetric water content, the red line represents 10-40cm volumetric
water content, the black line represents 0-10cm volumetric water content, and the grey dashed line represents the trend line. Figure 5
Interannual variation of soil moisture. The blue line represents 40-100cm volumetric water conte
water content, the black line represents 0-10cm volumetric water content, and the grey dashed li Figure 5 Interannual variation of soil moisture. The blue line represents 40-100cm volumetric water content, the red line represents 10-40cm volumetric
water content, the black line represents 0-10cm volumetric water content, and the grey dashed line represents the trend line. Page 17/20 Page 17/20 Page 17/20 Figure 6
Correlations of δ18OTR with monthly climate variables (temperature, PDSI, VPD, RH, PRCP). The “+” and “*” indicate significant correlations at the
95% and 99% level, respectively. The Tmax stands for maximum temperature, T stands for mean temperature, the Tmin stands for minimum
temperature, PDSI stands for Palmer Drought Index, VPD stands for vapor pressure deficit, RH stands for relative humidity, PRCP stands for
precipitation. Figure 6 Correlations of δ18OTR with monthly climate variables (temperature, PDSI, VPD, RH, PRCP). The “+” and “*” indicate significant correlations at the
95% and 99% level, respectively. The Tmax stands for maximum temperature, T stands for mean temperature, the Tmin stands for minimum
temperature, PDSI stands for Palmer Drought Index, VPD stands for vapor pressure deficit, RH stands for relative humidity, PRCP stands for
precipitation. Page 18/20 Figure 7
Correlations of δ18OTR with monthly soil moisture. The "+" represents p<0.05, "*" represents p<0.01, SM40-100 cm represents 40-100 cm soil
moisture, SM10-40 cm represents 10-40 cm soil moisture, SM0-10 cm represents 0-10 cm soil moisture. Figure 7 Figure 8 Figure 8 The spatial correlation between the June-September VWC in each soil layer and δ18OTR. The green point in the figure represents δ18OTR sampling
points and the black line is the Tibetan Plateau boundary line. Appendix.docx Figure 7 Correlations of δ18OTR with monthly soil moisture. The "+" represents p<0.05, "*" represents p<0.01, SM40-100 cm represents 40-100 cm soil
moisture, SM10-40 cm represents 10-40 cm soil moisture, SM0-10 cm represents 0-10 cm soil moisture. Page 19/20 Figure 8
The spatial correlation between the June-September VWC in each soil layer and δ18OTR. The green point in the figure represents δ18OTR sampling
points and the black line is the Tibetan Plateau boundary line. S
l
il Figure 8
The spatial correlation between the June-September VWC in each soil layer an
points and the black line is the Tibetan Plateau boundary line. Figure 8 The spatial correlation between the June-September VWC in each soil layer and δ18OTR. The green point in the figure represents δ18OTR sampling
points and the black line is the Tibetan Plateau boundary line. Supplementary Files This is a list of supplementary files associated with this preprint. Click to download. Page 20/20
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How to collect non-medical data in a pediatric trial: diaries or interviews
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To cite this version: Anaïs Le Jeannic, Hassani Maoulida, Sophie Guilmin-Crépon, Corinne Alberti, Nadia Tubiana-Rufi,
et al.. How to collect non-medical data in a pediatric trial: diaries or interviews. Trials, 2020, 21 (1),
pp.36. 10.1186/s13063-019-3997-9. inserm-02468132 How to collect non-medical data in a pediatric trial:
diaries or interviews Anaïs Le Jeannic, Hassani Maoulida, Sophie Guilmin-Crépon, Corinne
Alberti, Nadia Tubiana-Rufi, Isabelle Durand-Zaleski HAL Id: inserm-02468132
https://inserm.hal.science/inserm-02468132v1
Submitted on 5 Feb 2020 L’archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
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archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers. Le Jeannic et al. Trials (2020) 21:36
https://doi.org/10.1186/s13063-019-3997-9 Open Access Abstract Background: Non-medical data, such as the amount of time that patients and caregivers spend managing their
condition, may be relevant when assessing therapeutic strategies. For chronic pediatric conditions, the time that
patients and caregivers spend in seeking and providing care (which are the indirect costs in an economic evaluation)
can be significantly different depending on the treatment arm. To explore methods for collecting information on the
care burden for caregivers and patients, we investigated whether a patient diary provided additional information
compared to retrospective investigator-led interviews and whether a diary that was completed intermittently produced
more or less information than a diary completed continually. The main objective of this study was to identify which
type of data collection was most effective for measuring the time spent by caregivers and for estimating indirect
treatment costs over 9 months. Methods: Start-In! is a randomized controlled trial comparing the efficacy of three strategies of real-time continuous
glucose monitoring for 12 months in children and adolescents with type 1 diabetes. We designed an ancillary study to
assess methods of collecting information on the time spent by patients and caregivers in managing their condition
(indirect costs). Data were entered retrospectively in case report forms (CRFs) by investigators during quarterly follow-
up visits, which were supplemented with diaries completed prospectively by children or caregivers either continuously
or intermittently. Data about absences from school and work as well as the time that caregivers spent on diabetes care
were collected and the three collection methods were compared. Results: At the end of the 9-month study, 42% of the study participants failed to return their diary. For the diaries that
were received, less than 10% of expected data were collected versus 82% during investigators'interviews. Based on all
the information collected, we calculated that over 9 months, caregivers lost on average 3.9 days of working time (€786)
and 4 days of personal time, i.e. the equivalent of €526, and spent around 15 min of time on care per day, i.e. the
equivalent of €1700. Conclusions: The CRFs completed by investigators during quarterly visits cannot be replaced by a diary. Completing
the diaries appeared to represent an important additional burden to children and their caregivers, and the diaries
provided little additional information compared to investigators’ entries in the CRF. Trial registration: ClinicalTrials.gov, NCT00949221. Registered on 30 July 2009. How to collect non-medical data in a
pediatric trial: diaries or interviews Anaïs Le Jeannic1,2,3*
, Hassani Maoulida1, Sophie Guilmin-Crépon2,4,5,6,7, Corinne Alberti2,4,5,7,
Nadia Tubiana-Rufi5,6 and Isabelle Durand-Zaleski1,8,9 Anaïs Le Jeannic1,2,3*
, Hassani Maoulida1, Sophie Guilmin-Crépon2,4,5,6,7, Corinne Alberti2,4,5,7,
Nadia Tubiana-Rufi5,6 and Isabelle Durand-Zaleski1,8,9 Background The clinical study had two phases: all subjects wore glu-
cose monitors during the first 3 months, and thereafter
were randomized into three groups with different strat-
egies for glucose monitoring during the next 9 months. The ancillary study reported here concerns the 9-month
follow-up period of the patients (Fig. 1). The Start-In! economic evaluation required informa-
tion on consultations, absences from work or school due
to diabetes, and the time spent by the caregiver on aver-
age each day on providing diabetes care. This informa-
tion was entered for all patients by the investigators in
paper case report forms (CRF) at the end of each trimes-
ter and represented the reference strategy. To determine the best way to collect information on care
burden and time costs, we investigated whether or not a
patient-completed diary would provide additional informa-
tion compared to retrospective investigator-led interviews
and whether a diary completed intermittently would
produce more or less information than a continuous diary. We hypothesized first that a diary would enable investiga-
tors to shorten their interviews, which would be helped by
or even replace the diaries. We assumed that patients
would tend to underreport their time costs. Our second hy-
pothesis was that keeping a continuous diary would be an
onerous task and could lead to fatigue and poor data collec-
tion over time. However, the intermittent diary, while redu-
cing the data entry burden for patients [8], has the risk that
patients may fail to complete it if it was not a regular task. gy
To test whether diaries would provide additional informa-
tion, patients were randomized into two groups. In the con-
tinuous group, children and caregivers were asked to collect
data in a diary without interruption during one of the three
trimesters of the study (3 months continuously). In the
intermittent group, data were collected only during the last
month of each of the three trimesters. Randomization was
in a 3:1: ratio. In the continuous group, the diary was com-
pleted daily for one of the three trimesters of the study
follow-up period, allocated randomly in a 1:1:1 ratio. To
simplify the work for the investigators, the randomization
for the ancillary study was by center and not by patient
(Figs. 2 and 3). Background In! trial (NCT00949221) that compared the efficacy and
cost-effectiveness of three strategies of real-time continu-
ous glucose monitoring (RT-CGM) for children with type
1 diabetes [9]. This trial was well adapted to our study ob-
jective since insulin-dependent diabetes in children is a
chronic condition with major time costs to children and
their parents. In! trial (NCT00949221) that compared the efficacy and
cost-effectiveness of three strategies of real-time continu-
ous glucose monitoring (RT-CGM) for children with type
1 diabetes [9]. This trial was well adapted to our study ob-
jective since insulin-dependent diabetes in children is a
chronic condition with major time costs to children and
their parents. g
Non-medical data, such as the time spent by patients and
caregivers in managing their condition, may be relevant in
assessing the full burden and cost of a therapeutic strategy. The pertinence of including time costs for caregivers and
patients as indirect costs in economic evaluations of thera-
peutic strategies may be considered when writing the
protocol and mainly depends upon the perspective of the
evaluation and the type of illness being considered [1]. Productivity losses can represent a significant amount of re-
sources, especially for chronic pediatric conditions because
caregivers may need to seek and also provide care for chil-
dren. Information on absences from work or school is not
recorded in health insurance databases, so must be esti-
mated, either through retrospective interviews with an in-
vestigator [2] or from diaries completed by the parents or
patients [3, 4]. Investigator-led interviews require additional
work from the physicians, which could be reduced if the
children or their caregivers completed diaries prospectively
[5]. However, the use of paper or electronic diaries in clin-
ical trials requires logistical management and fastidiousness
from participants, especially for studies with long follow-up
periods [6]. This may be more problematic for pediatric tri-
als, since parental constraints may influence the acceptance
of additional methods of data collection [7]. The patients in Start-In! were aged from 2 to 17 years. They had been diagnosed with type 1 diabetes for more
than 1 year and were being treated by intensive insulin
therapy, achieving inadequate metabolic control. The
study was conducted in 11 pediatric diabetology units
with expertise in pump therapy and CGM. The primary
endpoint was long-term glycemic control at 12 months. Abstract Registry name: Study of Insulin Therapy
Augmented by Real Time Sensor in Type 1 Children and Adolescents (START-IN!). Keywords: patient diary, data collection, time costs, investigator-led interview © The Author(s). 2020 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver
(http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. * Correspondence: anais.lejeannic@urc-eco.fr
1AP-HP, Groupe hospitalier Hôtel-Dieu, URC Economie de la Santé Ile de
France, Paris, France
2Inserm, ECEVE UMR-S 1123, Paris, France
Full list of author information is available at the end of the article * Correspondence: anais.lejeannic@urc-eco.fr
1AP-HP, Groupe hospitalier Hôtel-Dieu, URC Economie de la Santé Ile de
France, Paris, France
2Inserm, ECEVE UMR-S 1123, Paris, France
Full list of author information is available at the end of the article © The Author(s). 2020 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver
(http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Le Jeannic et al. Trials (2020) 21:36 Le Jeannic et al. Trials (2020) 21:36 Page 2 of 7 Page 2 of 7 Background Data from the patient interviews and diaries
were entered into the CRFs by investigators following the
same schedule: one trimester for the continuous group and
the last month of each trimester for the intermittent group. The main objective of this study was to identify which
type of data collection method was most effective for
obtaining information on care burden and time costs over
the 9 months of a pediatric trial. The secondary objectives
were to determine the patient acceptance of an intermit-
tent versus a continuous diary and to check whether the
amount of data collected decreased over time. Endpoint The primary endpoint was the amount of information re-
corded in the diaries, compared to the amount of informa-
tion given to the investigator during the quarterly follow-up
visit. The Start-In! participants (patients and caregivers) were
asked to record in their diary: (1) for the child, the dates of
each period of absence from school or work, (2) for an adult
caregiver, the dates of each period of absence from work, (3)
the date of each consultation, (4) the number of caregivers
present at each consultation, and (5) an estimate of the time Methods
Study design Start-In! was a French multi-center trial with a random-
ized controlled prospective open parallel-group design,
comparing three therapeutic modalities in pediatric man-
agement [9]. We designed an ancillary study to the Start- Le Jeannic et al. Trials (2020) 21:36 Le Jeannic et al. Trials Page 3 of 7 Fig. 1 Design of the Start-In! trial. The discontinuous group used intermittent RT-CGM alternating with SMBG. RT-CGM real-time continuous
glucose monitoring, SMBG self-monitoring blood glucose spent on average each day by the caregivers to provide dia-
betes care such, as blood glucose monitoring and insulin ad-
ministration (Additional file 1). There was no minimum or
maximum for the number of absences or visits that could be
recorded. The same information was also collected quarterly
by the investigators and entered in the CRF with the help of
the diary, which was collected at the same time. entered in the diaries for each patient. If the amount of
time spent by the caregiver on average each day was not
completed, it was considered to be missing data rather
than zero. If the other data items were not completed at
all (dates of absences from school or work, and dates of
consultations), we considered that the data were missing
if the average time spent by the caregiver was also miss-
ing. The overall amount of information recorded was
compared between the intermittent and continuous
groups, and with the amount of information entered by
the investigators into the CRFs. Analysis y
The number of absences from work or school and the
number of consultations were calculated from the data Fig. 2 Groups for the Start-In! trial and the present study. Groups 1, 2, and 3 in the Start-In! trial used classic SMBG, 100% RT-CGM, and 40% RT-CGM,
respectively. CG continuous group, IG intermittent group, RT-CGM real-time continuous glucose monitoring, SMBG self-monitoring blood glucose Le Jeannic et al. Trials (2020) 21:36 Page 4 of 7 Fig. 3 Timeline of data collection for each randomization group. T0 marks the end of the third month of Start-In! Blue ovals represent the
continuous groups (CG1, CG2, and CG3) and orange circles represent the intermittent group (IG). CG, continuous group, CRF case report form, IG
intermittent group Fig. 3 Timeline of data collection for each randomization group. T0 marks the end of the third month of Start-In! Blue ovals represent the
continuous groups (CG1, CG2, and CG3) and orange circles represent the intermittent group (IG). CG, continuous group, CRF case report form, IG
intermittent group Intermittent group Of the 36 patients, just under half (n = 16) returned at
least one diary. Each patient should have submitted
three diaries, one for each trimester, but only 41 of the
108 diaries expected were received. Of those 41 diaries, Continuous group For the indirect costs in the economic evaluation, the
opportunity cost was based on the number of absences
from work or school for the patients and their families. For those who were in employment, it was based on the
average salary in France (€34 per hour) and for those
who were unemployed, on the average salary of a house
cleaner (€18.7 per hour). The average daily time spent
by a caregiver administering diabetes care to a patient
was valued based on the average salary of a home nurse
(€28.8 per hour). All costs are in euros (€) for 2017 (1
€ = 1.2US$) and were not discounted due to the short
time horizon. Of the 92 patients, only 53.3% (n = 49) returned their
diary. Of the 49 diaries returned, 44 were empty so data
were considered to be missing. Four diaries reported
that no absences had occurred and only one diary gave
the date of an absence from work or school. That is,
89.8% of the diaries received had missing data. Overall,
of the 92 diaries expected, 99.5% were either not re-
ceived or had missing data. Results Table 1 Frequency, mean age, and ratio of boys to girls at Table 1 Frequency, mean age, and ratio of boys to girls at
inclusion
Centre
Frequency
Mean age (years)
Sex ratio (boys/girls)
1
30
11.1
1.14
2
31
11.2
2.88
3
17
11.8
1.43
4
9
12.3
0.80
5
19
13.9
0.36
6
13
10.9
0.44
7
9
14.2
0.50
8
8
11.7
1.67
9
6
9.8
0.50
10
6
11.5
2.00
11
3
12.5
0.50
Overall
151
11.8
1.04 A total of 151 patients were randomized in the Start-In! trial, 52 into Group 1, 48 into Group 2, and 51 into
Group 3. Their characteristics were similar at inclusion
[9]. They had an average age of 12 years and there was
nearly a 50:50 ratio of boys to girls, although there was
large differences in age and sex between the centers, as
seen in Table 1. A total of 23 patients who left the study
were excluded from our ancillary analysis of the diary
data, as their diaries would, obviously, be empty. Thus,
diaries were expected from 128 patients for analysis, 92
in the continuous group and 36 in the intermittent
group. Information on care burden was additionally col-
lected on all 151 patients by the investigators during the
follow-up visits. However, almost half of the 128 patients
in this analysis did not return a single diary, so the infor-
mation they reported to the investigators and entered in
the CRF was based solely on their memory. Le Jeannic et al. Trials (2020) 21:36 Page 5 of 7 objective was to estimate the indirect medical costs in
each treatment arm and identify possible differences that
cannot be captured by calculating direct medical costs
based upon information systematically retrieved from
hospital insurance claims data. All investigators were
asked to record in the CRFs data on caregiver and pa-
tient burden (time), and they relied on information ob-
tained from both interviews and diaries. Our study
compared two prospective methods of documenting the
treatment burden for caregivers and children with type 1
diabetes during a clinical trial, which included an eco-
nomic evaluation. Our major finding was the magnitude
of the data missing in the diaries provided to caregivers
(around 99%). Results While the average cost of caregivers’ time
could be estimated from investigator-led interviews to
be around €3000 per patient over 9 months, the infor-
mation from diaries alone did not allow a reliable
estimate to be calculated. It would appear that asking
families to record non-medical information in a diary
during a pediatric trial of a severe chronic condition is
infeasible, and we do not recommend their use. 97.6% had missing data, as only one diary was complete. Overall, 99.5% of diaries from patients were either not
received or had missing data. Care burden and indirect costs Since the CRFs, which were based upon interviews and
diaries, held about 80% of the indirect cost data, we
calculated productivity losses and indirect costs. Over 9
months, parents lost on average 3.9 days of working time,
i.e. the equivalent of €786, and 4 days of unemployed time,
i.e. the equivalent of €526, and spent around 15 min of
time on care per day, i.e. the equivalent of €1700. Children
were absent from school for an average of 3 days over the
9 months. Many studies from different countries show a high
concordance between self-reported and medical records
of healthcare resource utilization in the general patient
population [11, 12]. We were interested in pediatric tri-
als, in which the diaries would be completed by care-
givers, who may be tired from caring for a chronically ill
child in addition to their jobs and may have other chil-
dren to take care of. In fact, family members of someone
with diabetes seems to be at higher risk of depression
[13] and parents of a child with type 1 diabetes not only
struggle with depression [14, 15] but also worry about
hypoglycemia [16], which can lead to parental emotional Investigator-completed CRF The same information was collected retrospectively from
the patients and caregivers by investigators, who entered
it into the CRFs at the trimestral follow-up visits. The
information was, potentially, supplemented by the data
entered prospectively in the diaries. Table 2 shows the numbers of CRFs completed for
each data item by the investigators for patients whose
diaries were missing or empty, by randomization group
(continuous or intermittent). For example, for the con-
tinuous group, regarding absences from work, among the
87 patients who either returned an empty diary or no
diary, this information was recorded for 73 in their CRF. For the consultations section, among the 35 patients
from the intermittent group with empty or no diaries,
this information was recorded for 34 in their CRF. We
calculated the percentage of data missing from the CRFs
in the same way we did for diaries. The continuous
group had more data missing in their CRFs than the
intermittent group (14% vs. 5%), which is far less than
the around 99% of data missing from the diaries. In con-
clusion, it appeared that for the diaries that were re-
ceived, less than 10% of expected data were collected
versus 82% during investigators'interviews. The amount of data entered by investigators for patients
in the intermittent group was slightly higher than for the
continuous group (95% vs. 86%). This may be because the
participants were asked to remember information for only
one month at the time of an interview instead of for three
[10]. In this trial, the CRFs always contained more data
than the diaries since the CRFs were based upon interviews
supplemented by the diaries (if the diaries had data). In
fact, diaries can be a reliable source of data to complement
usual reporting methods, as shown by Dunn et al. [5], but
we were not able to confirm this conclusion because of the
amount of missing diary data. References 1. Haute Autorité de Santé - Choix méthodologiques pour l’évaluation
économique à la HAS [Internet]. [cité 30 juill 2018]. Disponible sur: https://
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evaluation-economique-a-la-has 1. Haute Autorité de Santé - Choix méthodologiques pour l’évaluation
économique à la HAS [Internet]. [cité 30 juill 2018]. Disponible sur: https://
www.has-sante.fr/portail/jcms/r_1499251/fr/choix-methodologiques-pour-l-
evaluation-economique-a-la-has Additional file 1. Data collected by patients in the diaries and also in
the CRFs. 2. Laidi C, Prigent A, Plas A, Leboyer M, Fond G, Chevreul K, et al. Factors associated
with direct health care costs in schizophrenia: Results from the FACE-SZ French
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Not applicable. Consent for publication
Not applicable. Authors’ contributions ALJ wrote the manuscript. ALJ and HM carried out the analysis. IDZ
supervised the methodology and statistical analysis. ALC, IDZ, SGC, CA and
NTR contributed to the methodology. All authors revised the manuscript. All
authors read and approved the final manuscript. Acknowledgements The authors would like to thank Meryl Darlington for reviewing this work, as
well as Neil Jouanjean and Céline Quelen for their help in managing the
diary data. 3. Lin C-WC, March L, Crosbie J, Crawford R, Graves S, Naylor J, et al. Maximum
recovery after knee replacement--the MARKER study rationale and protocol. BMC Musculoskelet Disord. 2009;10:69. 4. Rösch M, Leidl R, Tirpitz C, Reinshagen M, Adler G, König HH. Cost
measurement based on a cost diary in patients with inflammatory bowel
disease. Z Gastroenterol. 2002;40(4):217–28. Competing interests The authors declare that they have no competing interests. Received: 29 August 2019 Accepted: 13 December 2019 Received: 29 August 2019 Accepted: 13 December 2019 Received: 29 August 2019 Accepted: 13 December 2019 Discussion Some trials require non-medical data that are not readily
available from patients’ charts or an insurance database. Investigators must either rely on patients’ recollection at
the time of a follow-up visit or provide diaries. Our Table 2 Frequency of CRFs completed for each data item for patients not returning diaries or returning empty diaries
Data item
Continuous group, n (%)
N = 87
Intermittent group, n (%)
N = 35
Absences from school
78 (90)
34 (97)
Absences from work
73 (84)
32 (91)
Consultations
81 (93)
34 (97)
Time spent on diabetes care
68 (78)
33 (94) Le Jeannic et al. Trials (2020) 21:36 Le Jeannic et al. Trials (2020) 21:36 Page 6 of 7 distress [17]. Difficulties are encountered with adoles-
cent patients, who are becoming more independent but
have troubled relationships with their parents [18] and
may have transition issues, like treatment adherence [19,
20]. In those circumstances, reporting non-medical data,
such as school absences, may not be a priority for the
family. Funding This study was supported by Direction générale de l’offre de soin and
sponsored by Assistance Publique, Hôpitaux de Paris (AP-HP). Unfortunately, the patients and their caregivers were
not asked to give their opinions about the different
methods of data collection, so we can only speculate
about why so much data were missing. This study has
other limitations. The amount of missing data prevented
us from performing statistical tests or evaluating our
secondary objectives of patient acceptance and fatigue. Moreover, with such little data, the quality of the infor-
mation provided in the diaries could not be assessed and
thus, only the quantity was explored in our trial. Supplementary information pp
y
Supplementary information accompanies this paper at https://doi.org/10. 1186/s13063-019-3997-9. y
Supplementary information accompanies this paper at https://doi.org/10. 1186/s13063-019-3997-9. Author details
1 1AP-HP, Groupe hospitalier Hôtel-Dieu, URC Economie de la Santé Ile de
France, Paris, France. 2Inserm, ECEVE UMR-S 1123, Paris, France. 3URC Eco IdF
(Paris health economics and health services research unit) and Inserm, ECEVE
UMR-S 1123, Paris, France. 4AP-HP, Hôpital Universitaire Robert Debré, Unité
d’Epidémiologie clinique, Paris, France. 5Université Paris Diderot, PRES
Sorbonne Paris Cité, Paris, France. 6AP-HP, Hôpital Robert Debré, Service
d’Endocrinologie-Diabétologie pédiatrique et Centre de référence des
Maladies Endocriniennes Rares de la Croissance, Paris, France. 7CIC-EC 1426,
Paris, France. 8AP-HP, Groupe hospitalier Albert Chenevier- Henri Mondor,
Service de Santé Publique, Créteil, France. 9Inserm METHODS CRESS UMR
1153, Paris, France. Conclusion Trials that require non-medical data may need to rely on
patients and families for information. In our study of a
pediatric chronic condition, investigator-led interviews
provided much more information than patient diaries. The completion of diaries appears to represent an import-
ant additional burden to children and their families, who
are already struggling with many competing issues. Notwithstanding, the large indirect costs associated with
the loss of parental productivity underlines the import-
ance of collecting these data, in one way or another. Availability of data and materials The datasets analyzed during the current study are available from the
corresponding author on reasonable request. Start-in! Study Group Nadia Tubiana-Rufi, Sophie Guilmin-Crépon, Jean-Bruno Lobut, Patricia Sierra,
Christine Delcroix (all CHU Robert Debré, AP-HP); Véronique Sulmont (CHU
Reims, CH Annemasse); Anne-Sophie Salmon, Pierre-François Souchon (both
CHU Reims); Claire Le Tallec (CHU Purpan, Toulouse); Régis Coutant, Natacha
Bouhours, Sylvie Dufresne, Frederique Gatelais (all CHU Angers); Fabienne
Dalla-Vale, Denis Morin (both CHU Arnaud de Villeneuve, Montpellier); Claire
Stuckens, Louis Montagne, Marie Dupre, Stéphanie Coopman (all CHU Lille);
Hélène Bony-Trifunovic, Karine Braun (both CHU Amiens); Hélène Crosnier,
Claire Personnier (both CH Intercommunal de Poissy); François Kurtz (CH
Saint-Avold); Florentia Kaguelidou, Laurence Corvez, Véronique Beruer, Eve-
lyne Jacqz-Aigrain (all Centre d’Investigation Clinique CIC 1426, Inserm, Rob-
ert Debré Hospital); Rolande Ducrocq (Laboratoire de Biochimie Métabolique,
CHU Robert Debré, AP-HP); Adyla Yacoubi, Tania Rilcy, Cécile Hoffart-
Jourdain, Akim Souag, Béatrice Andriss, Damir Mohamed, Isabelle Durand-
Zaleski (all Unité de Recherche Clinique Robert Debré, Unité de Recherche
Clinique Economie de la Santé and Direction de la Recherche Clinique et de
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patient diaries in conjunction with standard reporting methods: duplication
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cost data collection may be an efficient method in economic clinical trials. BMC Health Serv Res. 2012;12:318. 8. Seidl H, Meisinger C, Wende R, Holle R. Empirical analysis shows reduced
cost data collection may be an efficient method in economic clinical trials. BMC Health Serv Res. 2012;12:318. 9. Guilmin‐Crépon S, Carel J-C, Schroedt J, Sulmont V, Salmon A-S, Tallec CL,
et al. Is there an optimal strategy for real-time continuous glucose
monitoring in pediatrics? A 12-month French multi-center, prospective,
controlled randomized trial (Start-In!). Pediatric Diabetes [Internet]. [cité 21 9. Ethics approval and consent to participate Ethics approval and consent to participate Ethics approval and consent to participate
The AFSSAPS (National Agency for the Safety of Medicines and Health
Products) approved the protocol of the START-IN! trial (2007-AO1330-53). In-
formed consent was obtained from all study participants. Consent for publication
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0.1111/pedi.12820 févr 2019];0(0). Disponible sur: https://onlinelibrary.wiley.com/doi/abs/1
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with Type 1 Diabetes or Cystic Fibrosis. J Pediatr Psychol. 2010;35(8):814–22. 16. Aalders J, Hartman E, Nefs G, Nieuwesteeg A, Hendrieckx C, Aanstoot H-J,
et al. Mindfulness and fear of hypoglycaemia in parents of children with
Type 1 diabetes: results from Diabetes MILES Youth – The Netherlands. Diabet Med. 2018;35(5):650–7. 17. Haugstvedt A, Wentzel-Larsen T, Graue M, Søvik O, Rokne B. Fear of
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associated with poor glycaemic control and parental emotional distress: a
population-based study. Diabet Med. 2010;27(1):72–8. 18. Jaser SS, Grey M. A Pilot Study of Observed Parenting and Adjustment in
Adolescents with Type 1 Diabetes and their Mothers. J Pediatr Psychol. 2010;35(7):738–47. 19. Changes in Treatment Adherence and Glycemic Control During the Transition
to Adolescence in Type 1 Diabetes | Diabetes Care [Internet]. [cité 22 août
2019]. Disponible sur: https://care.diabetesjournals.org/content/35/6/1219 20. Lotstein DS, Seid M, Klingensmith G, Case D, Lawrence JM, Pihoker C, et al. Transition From Pediatric to Adult Care for Youth Diagnosed With Type 1
Diabetes in Adolescence. Pediatrics. 2013;131(4):e1062–70. Start-in! Study Group Guilmin‐Crépon S, Carel J-C, Schroedt J, Sulmont V, Salmon A-S, Tallec CL,
et al. Is there an optimal strategy for real-time continuous glucose
monitoring in pediatrics? A 12-month French multi-center, prospective,
controlled randomized trial (Start-In!). Pediatric Diabetes [Internet]. [cité 21 Page 7 of 7 Le Jeannic et al. Trials (2020) 21:36 Publisher’s Note Springer Nature remains neutral with regard to jurisdictional claims in
published maps and institutional affiliations. Springer Nature remains neutral with regard to jurisdictional claims in
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Effectiveness of Link Prediction for Face-to-Face Behavioral Networks
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Abstract Research on link prediction for social networks has been actively pursued. In link prediction for a given social network
obtained from time-windowed observation, new link formation in the network is predicted from the topology of the
obtained network. In contrast, recent advances in sensing technology have made it possible to obtain face-to-face
behavioral networks, which are social networks representing face-to-face interactions among people. However, the
effectiveness of link prediction techniques for face-to-face behavioral networks has not yet been explored in depth. To
clarify this point, here we investigate the accuracy of conventional link prediction techniques for networks obtained from
the history of face-to-face interactions among participants at an academic conference. Our findings were (1) that
conventional link prediction techniques predict new link formation with a precision of 0.30–0.45 and a recall of 0.10–0.20, (2)
that prolonged observation of social networks often degrades the prediction accuracy, (3) that the proposed decaying
weight method leads to higher prediction accuracy than can be achieved by observing all records of communication and
simply using them unmodified, and (4) that the prediction accuracy for face-to-face behavioral networks is relatively high
compared to that for non-social networks, but not as high as for other types of social networks. Citation: Tsugawa S, Ohsaki H (2013) Effectiveness of Link Prediction for Face-to-Face Behavioral Networks. PLoS ONE 8(12): e81727. doi:10.1371/
journal.pone.0081727 Editor: Peter Csermely, Semmelweis University, Hungary Received August 2, 2013; Accepted October 15, 2013; Published December 10, 2013 Received August 2, 2013; Accepted October 15, 2013; Published December 10, 2013 Copyright: 2013 Tsugawa, Ohsaki. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Copyright: 2013 Tsugawa, Ohsaki. This is an open-access article distributed under the terms of the Creative Commons Att
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This work was supported by JSPS KAKENHI Grant Number 25280030. The funders had no role in study design, data collection and analysis, decision to
publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: s-tugawa@cs.tsukuba.ac.jp Link prediction for face-to-face behavioral networks should be
useful for developing novel services and performing sociological
analyses. Abstract Link prediction is promising for predicting communica-
tions that are likely to occur; this can be viewed as potential
communication demands. Hence, link prediction in face-to-face
behavioral networks may contribute toward realizing novel
services such as friendship recommendation in real-world envi-
ronments, which is already common in online environments. Link
prediction techniques may also be useful for analyzing the
evolutionary dynamics of social networks in real-world environ-
ments, an important topic in social science. Sho Tsugawa1*, Hiroyuki Ohsaki2 ation and Systems, University of Tsukuba, Ibaraki, Japan, 2 School of Science and Technology, Kwansei Gakuin University, Hyogo, Japa of Engineering, Information and Systems, University of Tsukuba, Ibaraki, Japan, 2 School of Science and Technology, Kwansei Gakuin U Effectiveness of Link Prediction for Face-to-Face
Behavioral Networks Sho Tsugawa1*, Hiroyuki Ohsaki2 December 2013 | Volume 8 | Issue 12 | e81727 Link Prediction Techniques To clarify the effectiveness of link prediction techniques for face-
to-face behavioral networks, here we investigate the accuracy of
conventional link prediction techniques for networks obtained
from the SocioPatterns dataset [20]. We extend our preliminary
work [24] and investigate the prediction accuracy under various
conditions. Our results support the findings in [1], and provide
new findings, such as that incorporating temporal information
about communication
is
essential for
improving prediction
accuracy. Moreover, we discuss the effectiveness of link prediction
techniques for face-to-face behavioral networks compared with
other networks through experiments with the Enron email dataset
[26], which is one of the largest available email corpora. Among various link prediction techniques proposed in the
literature, the common neighbor (CN) [9], Adamic/Adar (AA)
[10], preferential attachment (PA) [9], Jaccard coefficient (JC)
[27], and resource allocation (RA) [11] are widely used and their
accuracies have been explored for several types of networks
[3,12,13,28]. For each node pair (i, j), a link prediction technique gives li, j,
an estimate of the likelihood of link formation between nodes i and
j. In other words, a link prediction technique predicts the
likelihood of future link formation as li, j. In the following paragraphs, we briefly introduce definitions of
the link prediction score li, j for the following types of link
prediction techniques for unweighted and weighted networks: CN
[9] and weighted CN (WCN) [4,28], AA [10] and weighted AA
(WAA) [4,28], PA [9] and weighted PA (WPA) [28], JC [27] and
weighted JC (WJC) [1], and RA [11] and weighted RA (WRA) [4]. In what follows, C(i) denotes the set of nodes adjacent to node i,
w(i, j) is a weight assigned to link (i, j), and a is a parameter for
controlling the contribution of link weights to the link prediction
score li, j. The main contributions of this study can be summarized as
follows. N We investigated the prediction accuracy of conventional link
prediction techniques for face-to-face behavioral networks by
using publicly available datasets. N We showed that in order to achieve high link prediction
accuracy for face-to-face behavioral networks, it is essential to
incorporate temporal information and to appropriately tune
the length of the training period, the period spanned by the
records analyzed; good tuning results in increased link
prediction accuracy. Common neighbor. Introduction Research on link prediction for social networks has been
actively pursued [1–8]. In link prediction for a given social
network obtained from time-window observation, new link
formation in the network is predicted from the topology of the
observed network. A social network is represented as a graph
where individuals are represented as nodes and social ties among
them are represented as links. In the literature, several link
prediction techniques have been proposed [4,5,9–11]. These
techniques can be used to predict new link formation by estimating
the likelihood of link formation between two nodes on the basis of
the observed network topology. Social ties can be defined in a
number of ways, and the accuracy of link prediction techniques
has been investigated for several types of social networks such as
coauthorship networks [3], email networks [12], and friendship
networks [13]. Link prediction techniques are expected to be
utilized for several applications such as recommendation [3],
anomaly detection [14], network modeling [15], missing link
detection [6], evaluation of network evolution mechanisms [16],
reconstruction of networks [17], and classification of partially
labeled networks [18,19]. However, the effectiveness of link prediction techniques for face-
to-face behavioral networks has not yet been fully explored. Our
research group has been investigating the effectiveness of link
prediction techniques for face-to-face behavioral networks using a
publicly available dataset called the SocioPatterns dataset [20],
which contains the history of face-to-face interactions among
participants at an academic conference. Our preliminary results
are presented in [24,25]. In addition, Scholz et al. [27] experi-
mentally investigated the accuracy of conventional link prediction
techniques for face-to-face behavioral networks by using their own
datasets. They also investigated the effectiveness of combining
face-to-face behavioral networks with other types of social
networks for link prediction [2]. To understand the effectiveness
of link prediction techniques when applied to real-world social
networks, however, more extensive investigation is necessary. For
instance, the effects of the observation period of social networks on Recent advances in sensing technology have made it possible to
obtain face-to-face behavioral networks, which are social networks
representing face-to-face interactions among people [20–23]. For
instance, in the SocioPatterns project, social networks representing
face-to-face interactions among participants at an academic
conference are constructed using badge-shaped sensing devices
[20]. Introduction December 2013 | Volume 8 | Issue 12 | e81727 1 PLOS ONE | www.plosone.org Link Prediction for Face-to-Face Networks Link Prediction for Face-to-Face Networks the prediction accuracy have not been discovered. The results in
[1] should be verified using other datasets. li, j ~ jC(i)\C(j)j
(in CN),
ð1Þ Overview of Experiments We evaluate the effectiveness of link prediction techniques
through experiments using the SocioPatterns dataset [20], which
contains the history of face-to-face communication among 110
participants over two and a half days at an academic conference
(ACM Hypertext 2009). At that conference, face-to-face commu-
nication was detected and stored on radiofrequency identification
(RFID) devices embedded in the conference badges worn by the
participants. Badges periodically broadcasted ultra-low power
radio packets that contained the participant’s ID. Each badge
collected packets every 20 s, and stored a list of IDs and
timestamps contained in the packets. Radio packet exchange
was possible only when two people were close (1.0–1.5 [m] apart)
and facing each other [20]. Face-to-face communication was
detected by the IDs stored in the badges. An interval of 20 s is
considered to be short enough to detect the occurrence of face-to-
face communication at social gatherings [20]. li, j ~
w(i,k)azw( j,k)a
log(1z
X
k0[C(k)
w(k0,k)a)
(in WAA):
ð4Þ ð4Þ Preferential attachment. The preferential attachment pre-
dicts new link formation from the idea that a high-degree node has
a higher chance of forming new links [9]. In PA [9] and WPA
[4,28], li,j is given by li, j ~ jC(i)j|jC( j)j
(in PA),
ð5Þ ð5Þ li, j ~
X
k[C(i)
w(i,k)a|
X
k[C( j)
w( j,k)a (in WPA):
ð6Þ ð6Þ For comparison purposes, we also use the Enron email dataset
[36] that contains 252,759 email messages with headers and body
texts exchanged between 151 employees of the Enron Corpora-
tion. Jaccard coefficient. Similarly to CN, the Jaccard coefficient
predicts new link formation from the number of common adjacent
nodes, but the link prediction score is normalized [27]. In JC [27]
and WJC [1], li, j is given by In our experiments, we divide the history of communication
into a training period and a testing period, and investigate how
accurately the occurrence of communications in the testing period
can be predicted using the history of communications in the
training period. Figure 1 shows an overview of the evaluation
method. li, j ~ jC(i)\C( j)j
jC(i)|C( j)j (in JC),
ð7Þ ð7Þ First, as an input for link prediction techniques, an undirected
graph G~(V, E), where each link weight is the number of
communications, is obtained from the history of communication
between times t0 and t1. The period between t0 and t1
corresponds to the training period. Link Prediction Techniques The common neighbor predicts new
link formation from the idea that the existence of many common
adjacent nodes between two nodes implies a high probability of
new link formation between those two nodes [10]. In CN [10] and
WCN [4], li, j, an estimate of the likelihood of link formation
between node i and j, is given by N We showed that the proposed decaying weight method can lead
to higher prediction accuracy than can be achieved by
observing all records of communication and simply using
them unmodified. The prediction accuracy of the decaying
weight method is comparable to the accuracy achievable with
an appropriately tuned training period. ð1Þ Figure 1. Overview of evaluation of link prediction techniques. New link formation is predicted from the topology of a social network
obtained from communications in the training period. We investigate the accuracy of link prediction techniques by comparing the predicted links
with actual new links created within the testing period. doi:10.1371/journal.pone.0081727.g001 Figure 1. Overview of evaluation of link prediction techniques. New link formation is predicted from the topology of a social network
obtained from communications in the training period. We investigate the accuracy of link prediction techniques by comparing the predicted links
with actual new links created within the testing period. doi:10.1371/journal.pone.0081727.g001 December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 2 Link Prediction for Face-to-Face Networks Table 1. Definitions of true positive (TP), true negative (TN), false positive (FP), and false negative (FN). ble 1. Definitions of true positive (TP), true negative (TN), false positive (FP), and false negative (FN). Number of communications is more
than or equal to S
Number of communications
is less than S
li, j is larger than or equal to T
TP
FP
li, j is smaller than T
FN
TN
TP, TN, FP, and FN are the numbers of node pairs (i,j) satisfying the conditions shown in this table. doi:10.1371/journal.pone.0081727.t001 Resource allocation. The resource allocation predicts new
link formation from an idea similar to that in AA [11]. In RA [11]
and WRA [4], li, j is given by ð2Þ li, j~
X
k[C(i)\C(j)
1
jC(k)j (in RA),
ð9Þ ð9Þ Adamic/Adar. The Adamic/Adar predicts new link forma-
tion from the idea that many common adjacent nodes with small
degree between two nodes implies a high probability of new link
formation between the nodes [10]. Link Prediction Techniques Similarly to CN, AA predicts
new link formation on the basis of the number of common
adjacent nodes, but assign a weight to li, j based on the degrees of
common adjacent nodes. In AA [10] and WAA [4], li, j is given by li, j~ w(i,k)azw( j,k)a
X
k0[C(k)
w(k0,k)a
(in WRA):
ð10Þ ð10Þ li, j ~
X
k[C(i)\C( j)
1
logjC(k)j (in AA),
ð3Þ ð3Þ December 2013 | Volume 8 | Issue 12 | e81727 Overview of Experiments In the graph G, a link (i, j)
represents the existence of a communication between individuals i
and j within the training period. The weight assigned to link (i, j) is
defined as the number of communications between individuals i
and j within the training period. li, j~
X
k[C(i)\C(j)
(w(i,k)azw( j,k)a)
X
k[C(i)
w(i,k)az
X
k[C(j)
w( j,k)a (in WJC):
ð8Þ ð8Þ PLOS ONE | www.plosone.org PLOS ONE | www.plosone.org December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 3 Link Prediction for Face-to-Face Networks Figure 2. The number of face-to-face communications recorded
in the SocioPatterns dataset over periods of one hour. The
onset of sensing face-to-face interactions is indicated by 0. doi:10.1371/journal.pone.0081727.g002 Precision evaluates the correctness, and recall evaluates the
completeness of link prediction Generally there is a tradeoff
Figure 2. The number of face-to-face communications recorded
in the SocioPatterns dataset over periods of one hour. The
onset of sensing face-to-face interactions is indicated by 0. doi:10.1371/journal.pone.0081727.g002
Figure 3. The number of email communications recorded in the
Enron email dataset over periods of one month. The onset
(indicated by 0) is April 1, 2000. doi:10.1371/journal.pone.0081727.g003 Figure 3. The number of email communications recorded in the
Enron email dataset over periods of one month. The onset
(indicated by 0) is April 1, 2000. doi:10.1371/journal.pone.0081727.g003 Figure 3. The number of email communications recorded in the
Enron email dataset over periods of one month. The onset
(indicated by 0) is April 1, 2000. doi:10.1371/journal.pone.0081727.g003 Figure 2. The number of face-to-face communications recorded
in the SocioPatterns dataset over periods of one hour. The
onset of sensing face-to-face interactions is indicated by 0. doi:10.1371/journal.pone.0081727.g002 Precision evaluates the correctness, and recall evaluates the
completeness of link prediction. Generally, there is a tradeoff
between precision and recall, whereby a larger threshold T
increases precision and decreases recall. AUC is defined as the
area under the ROC curve [32]. The ROC curve is obtained by
plotting the false positive rate (FP=(FNzTN)) versus recall by
changing the threshold T. AUC is 0.5 when the prediction
accuracy is equal to a random prediction, and is 1.0 with perfect
prediction accuracy [32]. Next, we calculate the link prediction score li, j for each node
pair (i, j) that is not associated with any communication before
time t1 by using one of the link prediction techniques introduced in
the previous section. Overview of Experiments Since li, j for CN, AA, PA, JC, and RA are
defined for an unweighted graph, we obtain li, j by simply ignoring
link weights in G. As a final step, we examine how accurately we can predict
which node pairs will have at least S communications between
times t2 and t3 despite not having any communication before time
t1. We perform this prediction by extracting node pairs whose link
prediction scores li, j meet or exceed a threshold T. The period
between times t2 and t3 corresponds to the testing period. It is
typical in studies of this kind to evaluate the accuracy of link
prediction techniques for S~1 only [4,28]. We introduce the
parameter S, and evaluate the effectiveness of link prediction
techniques not only for predicting new link formation but also for
predicting the formation of strong ties (i.e., links with large
weights). We perform experiments by changing S and a. Unless explicitly
stated we use S~1 and a~1. We first investigate the accuracy of
conventional link prediction techniques for face-to-face behavioral
networks using the SocioPatterns dataset, and then discuss the
effectiveness of link prediction techniques for face-to-face behav-
ioral networks compared to other networks through experiments
with the Enron email dataset. Figure 4. Degree distribution of a network obtained from the
entire history of face-to-face communications in the Socio-
Patterns dataset. In the network, each node represents a participant
at the conference, and each link between two nodes represents the
occurrence of face-to-face communication between participants during
the conference. doi:10.1371/journal.pone.0081727.g004 The link prediction problem can be formulated as ranking or
classification problems. Following prior work [1,6,13,29–31], we
use precision and recall, which are commonly used for evaluating
ranking problems, as well as area under the receiver operating
characteristic (ROC) curve (AUC) [32], which is commonly used
for evaluation of classification problems. Precision P and recall R
are defined as in Eqs. (11) and (12) using the concepts of true
positive (TP), true negative (TN), false positive (FP), and false
negative (FN). TP, TN, FP, and FN, which represent the
respective numbers of node pairs satisfying the corresponding
conditions in Table 1. P~
TP
TPzFP
ð11Þ
R~
TP
TPzFN
ð12Þ P~
TP
TPzFP
ð11Þ
R~
TP
TPzFN
ð12Þ ð11Þ Figure 4. Degree distribution of a network obtained from the
entire history of face-to-face communications in the Socio-
Patterns dataset. Comparison of Ten Link Prediction Techniques Comparison of Ten Link Prediction Techniques To investigate the accuracy of link prediction techniques for a
face-to-face behavioral network, we find the precision and recall of
ten link prediction techniques while varying the threshold T. Precision-recall curves for ten link prediction techniques are shown
in Fig. 6. The AUC scores of the ten link prediction techniques are
shown in Fig. 7. The training period is time 0–30 [h] (t0~0,
t1~30) and the testing period is time 30–60 [h] (t2~30, t3~60). Figure 6 shows that for instance, RA achieves a precision of
0.30–0.45 and a recall of 0.10–0.20 with an appropriate threshold
T for a face-to-face behavioral network obtained from the
SocioPatterns dataset. Unsurprisingly, this result suggests that link prediction tech-
niques cannot perfectly predict future link formation, but can
nevertheless
achieve
considerable
accuracy. It
is
therefore
expected that conventional link prediction techniques can be
applicable to services such as friendship recommendation that
require a moderate level of prediction accuracy. For instance, it is
reported in [29] that future friendship formation in an online
social networking service called LiveJournal can be predicted with
a precision of 0.18 and a recall of 0.18. Since networks in
LiveJournal and the SocioPatterns dataset are rather different in
terms of the number of nodes and links, direct comparison of the
results for those two networks is impossible. However, we expect
that a precision of 0.30–0.45 and a recall of 0.10–0.20 are
sufficient accuracy for a friendship recommendation service. Figure 5. Degree distribution of a network obtained from the
entire history of email communication in the Enron email
dataset. In the network, each node represents an employee of the
Enron Corporation, and each link between two nodes represents the
occurrence of email communication between the corresponding
employees from April 2000 to March 2002. doi:10.1371/journal.pone.0081727.g005 Results and Discussion y
p
Focusing on the differences among link prediction techniques,
we find that RA and PA achieve the highest accuracy. Figure 7
shows that the AUC scores of RA and PA are the highest. The
DeLong test [33] shows that the AUC scores of RA and PA are
significantly higher than those for other methods ( pv0:01), and
that there is no significant difference between RA and PA
( pw0:1). The high accuracy of RA is expected since the existence
of many common neighbors between two individuals intuitively
means a high probability of face-to-face communication. The
higher accuracy of RA compared to CN and AA, which are
similar to RA, is due to the existence of high-degree nodes (Fig. 4). RA successfully considers such high-degree nodes, which results in
high prediction accuracy. In [11], it is shown that the performance
of RA is higher than that of AA for networks containing nodes
with notably high degrees. The high accuracy of PA is also due to
the existence of high-degree nodes. Such nodes, which represent
eminent participants at the conference, tend to have many
communication events, which results in the high accuracy of PA. We can also see that the accuracies of techniques for the weighted Overview of Experiments In the network, each node represents a participant
at the conference, and each link between two nodes represents the
occurrence of face-to-face communication between participants during
the conference. doi:10.1371/journal.pone.0081727.g004 R~
TP
TPzFN
ð12Þ ð12Þ PLOS ONE | www.plosone.org
4
December 2013 | PLOS ONE | www.plosone.org PLOS ONE | www.plosone.org December 2013 | Volume 8 | Issue 12 | e81727 4 Link Prediction for Face-to-Face Networks Figure 5. Degree distribution of a network obtained from the
entire history of email communication in the Enron email
dataset. In the network, each node represents an employee of the
Enron Corporation, and each link between two nodes represents the
occurrence of email communication between the corresponding
employees from April 2000 to March 2002. doi:10.1371/journal.pone.0081727.g005 Characteristics of Datasets Before evaluating the effectiveness of link prediction techniques,
we analyze the characteristics of datasets used in this paper. Figure 2 shows the number of face-to-face communications
recorded in the SocioPatterns dataset over intervals of one hour,
and Fig. 3 shows the number of email communications recorded in
the Enron email dataset over intervals of one month. The degree
distributions of networks constructed from all communication logs
in the SocioPatterns dataset and the Enron email dataset are
shown in Figs. 4 and 5, respectively. Figure 2 shows that the number of face-to-face communications
fluctuates in the SocioPatterns dataset, and Fig. 3 shows that there
is no such tendency in email communications in the Enron email
dataset. Figures 4 and 5 show that the degree distribution is vastly
different in the two networks. For that reason, the effectiveness of
link prediction techniques might be different for the two networks. Figure 6. Relation between precision and recall for ten link prediction techniques. (training period: 0–30 (t0~0, t1~30), testing period:
30–60 (t2~30, t3~60), parameter for link prediction: a~1, dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g006 Figure 6. Relation between precision and recall for ten link prediction techniques. (training period: 0–30 (t0~0, t1~30), testing period:
30–60 (t2~30, t3~60), parameter for link prediction: a~1, dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g006 December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 5 Link Prediction for Face-to-Face Networks Figure 7. Comparison of the AUC scores of the ten link prediction techniques. (training period: 0–30 (t0~0, t1~30), testing period: 30–60
(t2~30, t3~60), parameter for link prediction: a~1, dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g007 Figure 7. Comparison of the AUC scores of the ten link prediction techniques. (training period: 0–30 (t0~0, t1~30), testing period: 30–60
(t2~30, t3~60), parameter for link prediction: a~1, dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g007 Figure 7. Comparison of the AUC scores of the ten link prediction techniques. (training period: 0–30 (t0~0, t1~30), testing period: 30–60
(t2~30, t3~60), parameter for link prediction: a~1, dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g007 networks (WCN, WAA, WPA, WJC, and WRA) are relatively
lower than techniques for the unweighted networks (CN, AA, PA,
JC, and RA). Lu¨ et al. [4] report the same phenomenon, which is analyzed in greater depth in the next section by varying the
parameter of the link prediction techniques. Effects of Link Prediction Parameters To investigate the effects of link prediction technique param-
eters on the prediction accuracy, we obtain the AUC scores of
WCN, WAA, WPA, WJC, and WRA while varying the parameter g
p
q
p
eters on the prediction accuracy, we obtain the AUC scores of
WCN, WAA, WPA, WJC, and WRA while varying the parameter
Figure 8. AUC scores of five link prediction techniques
obtained while varying the parameter a, which controls the
contribution of link weights to the link prediction score. (training period: 0–30 (t0~0, t1~30), testing period: 30–60 (t2~30,
t3~60), dataset: SocioPatterns dataset). a~0 is equivalent to ignoring
link weights, and a~1 is equivalent to simply using the number of face-
to-face communications as the link weight. doi:10.1371/journal.pone.0081727.g008
Figure 9. AUC scores of five link prediction techniques
obtained while varying the threshold S. (training period: 0–30
(t0~0, t1~30), testing period: 30–60 (t2~30, t3~60), dataset: Socio-
Patterns dataset). doi:10.1371/journal.pone.0081727.g009
PLOS ONE | www.plosone.org
6
December 2013 | Volume 8 | Issue 12 | e81727 Figure 8. AUC scores of five link prediction techniques
obtained while varying the parameter a, which controls the
contribution of link weights to the link prediction score. (training period: 0–30 (t0~0, t1~30), testing period: 30–60 (t2~30,
t3~60), dataset: SocioPatterns dataset). a~0 is equivalent to ignoring
link weights, and a~1 is equivalent to simply using the number of face-
to-face communications as the link weight. doi:10.1371/journal.pone.0081727.g008 Figure 9. AUC scores of five link prediction techniques
obtained while varying the threshold S. (training period: 0–30
(t0~0, t1~30), testing period: 30–60 (t2~30, t3~60), dataset: Socio-
Patterns dataset). doi:10.1371/journal.pone.0081727.g009
December 2013 | Volume 8 | Issue 12 | e81727 Figure 8. AUC scores of five link prediction techniques
obtained while varying the parameter a, which controls the
contribution of link weights to the link prediction score. (training period: 0–30 (t0~0, t1~30), testing period: 30–60 (t2~30,
t3~60), dataset: SocioPatterns dataset). a~0 is equivalent to ignoring
link weights, and a~1 is equivalent to simply using the number of face-
to-face communications as the link weight. doi:10.1371/journal.pone.0081727.g008 Figure 9. AUC scores of five link prediction techniques
obtained while varying the threshold S. (training period: 0–30
(t0~0, t1~30), testing period: 30–60 (t2~30, t3~60), dataset: Socio-
Patterns dataset). doi:10.1371/journal.pone.0081727.g009 December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 6 Link Prediction for Face-to-Face Networks Figure 10. AUC score of CN obtained while varying the training period length. Effects of Link Prediction Parameters (t1 and t2 are fixed to 12, 24, or 36 [h], and t0 is varied. The
testing period length is 3, 5, and 10 h. Dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g010 Figure 10. AUC score of CN obtained while varying the training period length. (t1 and t2 are fixed to 12, 24, or 36 [h], and t0 is varied
testing period length is 3, 5, and 10 h. Dataset: SocioPatterns dataset). d i 10 1371/j
l
0081727 010 Figure 10. AUC score of CN obtained while varying the training period length. (t1 and t2 are fixed to 12, 24, or 36 [h], and t0 is varied. The
testing period length is 3, 5, and 10 h. Dataset: SocioPatterns dataset). doi:10.1371/journal.pone.0081727.g010 a, which controls the contribution of link weights to the link
prediction score (Fig. 8). formation of strong ties, and a small S means that we predict the
formation of links including weak ties (links with small weights). We show the results for only CN, AA, PA, JC, and RA in this
investigation. This figure shows that appropriate link weights improve the
accuracy of link prediction. However, as reported in [4], the
prediction accuracy obtained when defining the link weight as the
number of face-to-face communications and simply using it
unmodified is even lower than that achieved when link weights
are ignored altogether. All five techniques achieve the highest
accuracy when a is approximately 0.3. This result suggests that the
accuracy of link prediction can be improved by increasing the
relative contribution of weak ties (links with small weights) to the
link prediction score li, j. More detailed analysis is, however,
required to determine the optimal value of a. This figure shows that the accuracy is higher when predicting
only the formation of strong ties than when predicting link
formation including weak ties. This suggests that link prediction
techniques are more effective at predicting the formation of strong
ties than weak ties. This result is considered natural since weak ties
may include noisy links, which are difficult to predict. Note that
this tendency is observed in different face-to-face behavioral
networks [1]. However, we should note that AUC scores do not
change significantly when S is six or higher. December 2013 | Volume 8 | Issue 12 | e81727 Effects of Training Period Length We next obtain the AUC score while varying the threshold S to
investigate the accuracy of each technique to predict the formation
of strong ties (links with large weights) (Fig. 9). In this investigation,
the training period and the testing period are time 0–30 [h] and
30–60 [h], respectively. A large S means that we predict only the In this section, we examine the effectiveness of using temporal
information about communication in addition to the network
topology for link prediction. By utilizing temporal information, we
can change the contribution of each communication event to the
link prediction score, which may result in more accurate December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 7 Link Prediction for Face-to-Face Networks Figure 11. Relation between the number of face-to-face contacts used for link prediction and AUC score of CN. t1 and t2 are fixed to
12, 24, or 36 [h], and t0 varies based on the number of contacts used for link prediction. 1000, 2000, and 5000 contacts are predicted immediately
following the training period, using the SocioPatterns dataset. doi:10.1371/journal.pone.0081727.g011 Figure 11 Relation between the number of face to face contacts used for link prediction and AUC score of CN t and t are f Figure 11. Relation between the number of face-to-face contacts used for link prediction and AUC score of CN. t1 and t2 are fixed to
12, 24, or 36 [h], and t0 varies based on the number of contacts used for link prediction. 1000, 2000, and 5000 contacts are predicted immediately
following the training period, using the SocioPatterns dataset. doi:10.1371/journal.pone.0081727.g011 Figure 10 shows that simply increasing the training period
length does not necessarily improve the prediction accuracy, and
can even degrade it. Although it is intuitively expected that using
more information should improve prediction accuracy, this result
suggests otherwise. prediction. A simple way to utilize temporal information is to
change the training period length. By changing the onset time of
the training period, we can evaluate the prediction accuracy when
the contribution of old communications are ignored. The effect of
the length of the training period on the prediction accuracy has
not been fully explored in existing work [1] or our preliminary
work [24,25]. When applying link prediction techniques to face-to-
face behavioral networks, it is important to know how to
determine the training period. December 2013 | Volume 8 | Issue 12 | e81727 Effects of Training Period Length Figure 10 also shows that the optimal training period length that
maximizes the AUC score depends on the starting time of the
testing period t2. Note that the optimal training period length does
not depend on the length of the testing period. We obtain the AUC scores of the link prediction techniques
while varying the training period length for several starting times
of the testing period t2 and lengths of the testing period. In this
investigation, the testing period immediately follows the training
period. Figure 10 shows the relation between the training period
length and AUC score for CN. Note that the results when using
other link prediction techniques are similar to the results shown in
Fig. 10. Since few communications take place at times 12–24 and
36–48 [h] (see Fig. 2), we use the data for periods 0–12, 24–36,
and 48–60 [h]. We next determine the training period length not by time, but
by the number of face-to-face contacts used for link prediction,
and perform a similar experiment to investigate the effect of
training period length. Since the number of face-to-face contacts
recorded in the dataset is different from hour to hour, this may
explain why the optimal training period length depends on the
starting time of the testing period. Figure 11 shows the relation
between the number of face-to-face contacts used for link
prediction and AUC score. In this investigation, the testing period
is also determined by the number of face-to-face contacts to
predict. PLOS ONE | www.plosone.org December 2013 | Volume 8 | Issue 12 | e81727 December 2013 | Volume 8 | Issue 12 | e81727 8 Link Prediction for Face-to-Face Networks Figure 12. Comparison of the AUC scores of WCN for four types of weighting methods. t1 and t2 are fixed to 12, 24, or 36 [h]. d~10{6. Parameter for link prediction: a~0:3, Dataset: SocioPatterns dataset. doi:10.1371/journal.pone.0081727.g012 Figure 12. Comparison of the AUC scores of WCN for four types of weighting methods. t1 and t2 are fixed to 12, 24, or 36 [h]. d~10{6. Parameter for link prediction: a~0:3, Dataset: SocioPatterns dataset. doi:10.1371/journal.pone.0081727.g012 Figure 12. Comparison of the AUC scores of WCN for four types of weighting methods. t1 and t2 are fixed to 12, 24, or 36 [h]. d~10{6. Parameter for link prediction: a~0:3, Dataset: SocioPatterns dataset. doi:10.1371/journal.pone.0081727.g012 prediction is increased. December 2013 | Volume 8 | Issue 12 | e81727 Effects of Training Period Length Hence, if we know that person A met person B, we can
infer that A will tend to communicate with persons near B and not
with persons located far away from B. However, as time elapses,
this information becomes irrelevant since persons A and B may
move. Therefore, recent information is more important than old
information, and the relevance of information about a particular
time segment decreases with time. cases, using approximately 3,000 contacts achieves the highest
accuracy. These results show that the decaying weight method achieves
higher accuracy than the flat weight method with t0~0. In
particular, the accuracy of the exponential decaying weight
method is comparable to that of the flat weight method with the
optimal training period length. The DeLong test [6] shows that
there are significant differences in AUC score between the
exponential decaying weight method and the flat weight method
with t0~0 when t1~t2~12 and t1~t2~24 ( pv0:01). The
DeLong test also shows that there are no significant differences in
AUC score between the exponential decaying weight method and
the flat weight method with the optimal training period length
( pw0:01). This result suggests that the decaying weight method
can be useful when the optimal training period length is unknown. Th
ff
i
f h
d
i
i h
h d i d
h These observations suggest that prolonged observation of face-
to-face contacts for link prediction often degrades the prediction
accuracy. We cannot draw general conclusions from our results
alone, but in this experiment, using approximately 3,000 contacts
(30 contacts per person) is optimal in many cases. Analysis using
other datasets is required in order to determine the optimal
training period length in the general case. We next investigate the prediction accuracy when using a
decaying weight method, which changes the contribution of each
contact on the link prediction score by weighting each face-to-face
contact. In the above experiments, all contacts used for link
prediction are weighted equally for link prediction score calcula-
tion. However, our experimental results suggest that simply using
all contacts often degrades the prediction accuracy. We therefore
change the contribution of each contact on the link prediction
score calculation by assigning a heavy weight to new information. In this experiment, we use two types of decaying weight methods,
linear and exponential. Let contacts between time t0 and time t1 be
c0, c1, c2,…, cn. Effects of Training Period Length In the linear decaying weight method, for a
contact ck a link weight in the graph G(V,E), which is used for
link prediction score calculation, is increased by k=n, and in the
exponential decaying weight method a link weight is increased by
eb(k{n). In exponential decaying weight method, b is determined
as e{bn~d, where d is a sufficiently small positive real number. The effectiveness of the decaying weight method is due to the
spatiotemporal
locality
of
face-to-face
communication. The
mobility of people is restricted by the space and their moving
speed. Hence, if we know that person A met person B, we can
infer that A will tend to communicate with persons near B and not
with persons located far away from B. However, as time elapses,
this information becomes irrelevant since persons A and B may
move. Therefore, recent information is more important than old
information, and the relevance of information about a particular
time segment decreases with time. Effects of Training Period Length We carefully investigated the results for
other starting times in the testing period, and found that prediction
accuracy increases as the number of contacts used for prediction
increases when the testing period includes the third day. In other These results show that in most cases, the prediction accuracy is
highest when using approximately 3,000 contacts. However, when
the starting time of the testing period is 36 [h], the prediction
accuracy increases together with the number of contacts used for Figure 13. Relation between precision and recall of ten link prediction techniques. (training period: April 2000–March 2001, testing period:
April 2001–March 2002, parameter for link prediction: a~1, dataset: Enron email dataset). doi:10.1371/journal.pone.0081727.g013 Figure 13. Relation between precision and recall of ten link prediction techniques. (training period: April 2000–March 2001, testing period:
April 2001–March 2002, parameter for link prediction: a~1, dataset: Enron email dataset). doi:10.1371/journal.pone.0081727.g013 December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 9 Link Prediction for Face-to-Face Networks Figure 14. Comparison of the AUC scores of the ten link prediction techniques. (training period: April 2000–March 2001, testing period:
April 2001–March 2002, parameter for link prediction: a~1, dataset: Enron email dataset). doi:10.1371/journal.pone.0081727.g014 Figure 14. Comparison of the AUC scores of the ten link prediction techniques. (training period: April 2000–March 2001, testing period:
April 2001–March 2002, parameter for link prediction: a~1, dataset: Enron email dataset). doi:10.1371/journal.pone.0081727.g014 These results show that the decaying weight method achieves
higher accuracy than the flat weight method with t0~0. In
particular, the accuracy of the exponential decaying weight
method is comparable to that of the flat weight method with the
optimal training period length. The DeLong test [6] shows that
there are significant differences in AUC score between the
exponential decaying weight method and the flat weight method
with t0~0 when t1~t2~12 and t1~t2~24 ( pv0:01). The
DeLong test also shows that there are no significant differences in
AUC score between the exponential decaying weight method and
the flat weight method with the optimal training period length
( pw0:01). This result suggests that the decaying weight method
can be useful when the optimal training period length is unknown. The effectiveness of the decaying weight method is due to the
spatiotemporal
locality
of
face-to-face
communication. The
mobility of people is restricted by the space and their moving
speed. Comparison with the Enron Email Dataset We next compare the results for the SocioPatterns dataset and
those for the Enron email dataset to investigate the characteristics
of link prediction techniques for face-to-face behavioral networks. Figure 13 shows the relation between precision and recall for each
link prediction technique for a network constructed from the
Enron email dataset. Figure 14 shows the AUC scores of the ten
link prediction techniques. In this investigation, the training period Figure 12 shows AUC scores of WCN when using flat weight
method, which equally weights all contacts, with the optimal
length of the training period, flat weight method with t0~0, linear
decaying weight method, and exponential decaying weight
method. PLOS ONE | www.plosone.org December 2013 | Volume 8 | Issue 12 | e81727 10 Link Prediction for Face-to-Face Networks Figure 15. Relation between the number of email messages used for link prediction and the AUC score of CN. (t1 and t2 are fixed to the
3rd month, 6th month, or 12th month, and t0 is changed based on the number of email messages used for link prediction. Predictions are made for
1000, 2000, and 5000 messages immediately after the training period. Dataset: Enron email dataset). doi:10.1371/journal.pone.0081727.g015 Figure 15. Relation between the number of email messages used for link prediction and the AUC score of CN. (t1 and t2 are fixed to the
3rd month, 6th month, or 12th month, and t0 is changed based on the number of email messages used for link prediction. Predictions are made for
1000, 2000, and 5000 messages immediately after the training period. Dataset: Enron email dataset). doi:10.1371/journal.pone.0081727.g015 and the testing period are April 2000 to March 2001 and April
2001 to March 2002, respectively. there are significant differences in AUC score between PA and
WPA and other techniques ( pw0:1). RA achieves the highest
accuracy for networks obtained from the SocioPatterns dataset,
but its accuracy is comparable to that of JC and AA for the
network obtained from the Enron email dataset. The DeLong test
shows that there are no significant differences in AUC score
among AA, JC, and RA ( pv0:01). In the network obtained from
the Enron email dataset, the number of high-degree nodes is
smaller than that in the network obtained from the SocioPatterns
dataset (Figs. 4 and 5). Comparison with the Enron Email Dataset Therefore, the link prediction scores of RA
tend to be similar to those of other similar techniques (i.e., AA and
JC), which determines the similarity in accuracy between these
techniques. From Fig. 13, we find that, for instance, JC achieves a precision
of 0.30–0.45 and a recall of 0.17–0.39 with an appropriate value of
the threshold T for an email network obtained from the Enron
email dataset. This result shows that conventional link prediction techniques
achieve higher precision and recall for email networks than for
face-to-face networks. Moreover, we should note that the number
of nodes in the network obtained from the Enron email dataset is
greater than that obtained from the SocioPatterns dataset. Hence,
link prediction techniques have higher performance for email
networks than for face-to-face networks. The topological structure
of a network may affect the accuracy of link prediction techniques
for the network. We next investigate the AUC score of CN for networks obtained
from the Enron email dataset by changing the number of email
messages used for prediction (Fig. 15). In this investigation, we
compare the effectiveness of tuning the training period length for a
face-to-face network and an email network. We also compare the
AUC scores of each weighting method (Fig. 16) and investigate the Moreover, Fig. 14 shows that the accuracy of link prediction for
the email network is significantly different among the ten link
prediction techniques. The accuracy of PA and WPA for the
network obtained from the Enron email dataset is significantly
lower than that for other techniques. The DeLong test shows that December 2013 | Volume 8 | Issue 12 | e81727 PLOS ONE | www.plosone.org 11 Link Prediction for Face-to-Face Networks Figure 16. Comparison of the AUC scores of WCN for four types of weighting methods. t1 and t2 are fixed to the 3rd, 6th, or 12th month. d~10{6. parameter for link prediction: a~0:3, Dataset: Enron email dataset. doi:10.1371/journal.pone.0081727.g016 gure 16. Comparison of the AUC scores of WCN for four types of weighting methods. t1 and t2 are fixed to the 3rd, 6th, or 12th mo Figure 16. Comparison of the AUC scores of WCN for four types of weighting methods. t1 and t2 are fixed to the 3rd, 6th, or 12th month. d~10{6. parameter for link prediction: a~0:3, Dataset: Enron email dataset. doi:10.1371/journal.pone.0081727.g016 effectiveness of incorporating temporal information with regard to
link prediction. behavioral network (see Fig. Acknowledgments The authors would like to thank Dr. Makoto Imase for valuable discussions
and Mr. Youhei Kobayashi for his kind support. The authors would like to thank Dr. Makoto Imase for valuable discussions
and Mr. Youhei Kobayashi for his kind support. Conclusions To clarify the effectiveness of link prediction techniques for face-
to-face behavioral networks, we have investigated the accuracy of
conventional link prediction techniques for such networks under
various conditions. We have performed several experiments
utilizing the SocioPatterns dataset containing a history of face-
to-face interactions among participants at an academic confer-
ence. Comparison with Other Networks We lastly compare the prediction accuracy for a face-to-face
behavioral network with that for other networks. Table 2 shows
the structural characteristics of several types of networks and the
AUC scores of CN for those networks. The AUC scores for
terrorist, and protein, and food web networks are taken from Fig. 3
in [6], and that for Facebook user networks is taken from Tables 3
and 5 in [13]. One possible method that can be used to avoid the need for
tuning the training period length in link prediction for face-to-face
behavioral networks is using the decaying weight method. Our
results showed that an exponential decaying weight method
achieves comparable accuracy with the method using the optimal
training period length, suggesting that the decaying weight method
can be useful when the optimal length of the training period is
unknown. Table 2 shows that CN achieves a higher AUC score for face-to-
face networks than protein networks, but achieves similar or lower
AUC scores compared to other networks. From this result, we
conclude that the accuracy of link prediction techniques for face-
to-face behavioral networks is higher than non-social networks, but
not so high among other types of social networks. We plan to
investigate the causes of this result in future research. We plan to investigate the causes of differences in accuracy
between different types of social networks. It is also important to
investigate the optimal training period length in other face-to-face
behavioral networks in future work. Link Prediction for Face-to-Face Networks sufficient for email networks. The DeLong test shows that there is
no statistically significant difference between the AUC scores of
the two methods ( pw0:1). This result also suggests that,
particularly for a face-to-face network, the training period should
be tuned carefully to improve the prediction accuracy. As
discussed in the previous subsection, in face-to-face networks,
using temporal information about communications is important
since face-to-face communication exhibits spatiotemporal locality. On the contrary, since email communication is not conducted in
real time and email messages are exchanged over the Internet, the
relevance of such locality for email networks is lower than that for
face-to-face networks. results for the SocioPatterns dataset and other datasets showed
that the accuracy of link prediction techniques for face-to-face
behavioral networks is relatively higher than that for non-social
networks, but not particularly high among other types of social
networks. These results suggest that conventional link prediction
techniques are applicable to services, such as friendship recom-
mendation, which require a moderate level of prediction accuracy. The experiments also reveal a unique characteristic of link
prediction for face-to-face behavioral networks whereby prolonged
observation of social networks often degrades the prediction
accuracy. For an email network, it is sufficient to use a naive
method that observes all records of email messages and simply uses
them unmodified. In contrast, for face-to-face behavioral net-
works, it is essential to determine an appropriate training period
length to ensure high prediction accuracy. In most cases in our
experiments, observing approximately 30 communications per
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techniques predict new link formation in a face-to-face behavioral
network with a precision of 0.30–0.45 and a recall of 0.10–0.20
through the use of an appropriate threshold. Comparison of the Conceived and designed the experiments: ST HO. Performed the
experiments: ST. Analyzed the data: ST HO. Contributed reagents/
materials/analysis tools: ST. Wrote the paper: ST HO. Conceived and designed the experiments: ST HO. Performed the
experiments: ST. Analyzed the data: ST HO. Contributed reagents/
materials/analysis tools: ST. Wrote the paper: ST HO. Comparison with the Enron Email Dataset 11), and suggests that the training
period must be determined carefully in order to improve the
prediction accuracy, particularly for a face-to-face behavioral
network. Figure 16 shows that the decaying weight method is not
effective, and that the naive method (i.e., the flat weight method) is Figure 15 shows that the prediction accuracy for an email
network is improved when the training period length is extended. This tendency is different from the results for a face-to-face Table 2. Structural characteristics of several types of networks and the AUC score of CN for those networks. network
number of nodes
average degree
clustering coefficient
average path length
AUC
face-to-face (SocioPatterns
dataset)
110
38.9
0.53
2.0
0.5–0.7
email (Enron email dataset)
151
9.3
0.70
6.0
0.6–0.85
terrorists [34]
–
4.9
0.36
2.6
0.5–0.8 [6]
Facebook [35]
63,731
24.3
–
–
0.8–0.94 [13]
protein [36]
–
4.8
0.06
3.7
0.5–0.58 [6]
food web [37]
–
3.0
0.17
3.3
0.5–0.7 [6]
doi:10.1371/journal.pone.0081727.t002
PLOS ONE | www plosone org
12
December 2013 | Volume 8 | Issue 12 | e81727 Table 2. Structural characteristics of several types of networks and the AUC score of CN for those netw 12 12 Link Prediction for Face-to-Face Networks g
y
(
) pp
6. Clauset A, Moore C, Newman MEJ (2008) Hierarchical structure and the
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English
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ChemicalToolBoX and its application on the study of the drug like and purchasable space
|
Journal of cheminformatics
| 2,014
|
cc-by
| 736
|
Lucas et al. Journal of Cheminformatics 2014, 6(Suppl 1):P51
http://www.jcheminf.com/content/6/S1/P51 Open Access ChemicalToolBoX and its application on the
study of the drug like and purchasable space
Xavier Lucas*, Björn A Grüning, Stefan Günther
From 9th German Conference on Chemoinformatics
Fulda, Germany. 10-12 November 2013 computed several physicochemical descriptors to dis-
cover general trends applying to each subset. The ever increasing amount of data and computational
capabilities in the cheminformatics field has led to a sce-
nario where efficient techniques for storage and proces-
sing in an integrated, modular, and easily accessible
platform are in vital demand. Here, we present Chemical-
ToolBoX, a compilation of more than 30 tools integrated
into a single computational chemistry and cheminfor-
matics platform based on the Galaxy workflow manage-
ment system [1,2]. We have recently designed a workflow
within the ChemicalToolBoX to generate a library of
compounds containing around 70 million unique com-
mercially available small molecules, i.e. the purchasable
space [3]. Subsequently, we have used filtering rules
based on structural patterns and chemical alarms to dis-
card problematic molecules, representing a very large
portion of the drug-like and purchasable space, along
with other drug discovery data sets including more than
2 million fragments (Figure 1). Furthermore, we have Published: 11 March 2014
References
1. [https://github.com/bgruening/galaxytools/tree/master/chemicaltoolbox]. 2. Grüning BA, et al:, submitted. 3. Lucas X, et al:, manuscript in preparation. doi:10.1186/1758-2946-6-S1-P51
Cite this article as: Lucas et al.: ChemicalToolBoX and its application on
the study of the drug like and purchasable space. Journal of
Cheminformatics 2014 6(Suppl 1):P51. Published: 11 March 2014 Published: 11 March 2014
References
1. [https://github.com/bgruening/galaxytools/tree/master/chemicaltoolbox]. 2. Grüning BA, et al:, submitted. 3. Lucas X, et al:, manuscript in preparation. doi:10.1186/1758-2946-6-S1-P51
Cite this article as: Lucas et al.: ChemicalToolBoX and its application on
the study of the drug like and purchasable space. Journal of
Cheminformatics 2014 6(Suppl 1):P51. References doi:10.1186/1758-2946-6-S1-P51
Cite this article as: Lucas et al.: ChemicalToolBoX and its application on
the study of the drug like and purchasable space. Journal of
Cheminformatics 2014 6(Suppl 1):P51. Open access provides opportunities to our
colleagues in other parts of the globe, by allowing
anyone to view the content free of charge. Publish with ChemistryCentral and every
scientist can read your work free of charge
W. Jeffery Hurst, The Hershey Company. available free of charge to the entire scientific community
peer reviewed and published immediately upon acceptance
cited in PubMed and archived on PubMed Central
yours you keep the copyright
Submit your manuscript here:
http://www.chemistrycentral.com/manuscript/
* Correspondence: xavier.lucas@pharmazie.uni-freiburg.de
Pharmaceutical Bioinformatics, Institute of Pharmaceutical Sciences, Albert-
Ludwigs-University, Freiburg, D-79104, Germany
Figure 1
© 2014 Lucas et al; licensee Chemistry Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons
Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in
any medium, provided the original work is properly cited. The Creative Commons Public Domain Dedication waiver (http://
creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. * Correspondence: xavier.lucas@pharmazie.uni-freiburg.de
Pharmaceutical Bioinformatics, Institute of Pharmaceutical Sciences, Albert-
Ludwigs-University, Freiburg, D-79104, Germany
Figure 1
© 2014 Lucas et al; licensee Chemistry Cent
Attribution License (http://creativecommo
any medium, provided the original work i
creativecommons.org/publicdomain/zero/ Figure 1 Open access provides opportunities to our
colleagues in other parts of the globe, by allowing
anyone to view the content free of charge. Publish with ChemistryCentral and every
scientist can read your work free of charge
W. Jeffery Hurst, The Hershey Company. available free of charge to the entire scientific community
peer reviewed and published immediately upon acceptance
cited in PubMed and archived on PubMed Central
yours you keep the copyright
Submit your manuscript here:
http://www.chemistrycentral.com/manuscript/ Publish with ChemistryCentral and every
scientist can read your work free of charge Publish with ChemistryCentral and every
scientist can read your work free of charge Open access provides opportunities to our
colleagues in other parts of the globe, by allowing
anyone to view the content free of charge. W. Jeffery Hurst, The Hershey Company. * Correspondence: xavier.lucas@pharmazie.uni-freiburg.de
Pharmaceutical Bioinformatics, Institute of Pharmaceutical S
Ludwigs-University, Freiburg, D-79104, Germany © 2014 Lucas et al; licensee Chemistry Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons
Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in
any medium, provided the original work is properly cited. The Creative Commons Public Domain Dedication waiver (http://
creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.
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https://openalex.org/W4313049783
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https://zenodo.org/record/7108648/files/11.%20Preparedness%20and%20Costing%20on%20Fire%20Safety.pdf
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English
| null |
Preparedness and Costing on Fire Safety Installation in Commercial Buildings
|
Social Science Research Network
| 2,022
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cc-by
| 9,178
|
ABSTRACT Purpose: The study was carried out to analyze the awareness among the management,
occupants and concerned authority of selected commercial buildings at Birtamode, jhapa and
prepare the tentative cost of additional tools and equipment’s needed. Design/Methodology/Approach: This research is focused on two commercial buildings;
Hanuman central and One stop mall (Birtamode Jhapa). These building are selected for
research as both of the buildings are considered as largest shopping malls and they also vary
in planning and construction technology. The fire safety preparedness and awareness of management, occupants were analyzed. After
then preparedness and awareness of concerned authority like fire fighters, Municipality and
Nepal police were analyzed. Data was collected from occupants, management and concerned
authority based on the questionnaires survey, site visits with check list and key informant
interview. Information regarding fire safety preparedness and awareness was presented in
tabular form, charts and Key informants interview and questionnaire. f
f
q
Findings/Result: Fire safety preparedness were lacking due to lack awareness on evacuation
procedure and equipment’s operation. Management seems to be more focused on financial
aspect rather than safety. Lack of proper firefighting equipment’s, equipment’s in poor
condition, unmanaged electrical wires is the existing problem in the buildings. Concerned
authority (Birtamode Municipality) had a fire Fighting team with a six fire men with a tank
and fire fighting vehicle. But the new technologies like GPS were not introduced to firefighter. There were many cases reported where due to communication gap between the fire fighter
and residents, the response was late which resulted more damage. Lack of fire hydrant near
the major commercial buildings created problem for fire fighter to refill their water tank. Narrow roads inside the municipality, poor implementation of fire safety code were major
problem for fire fighters. The tentative cost for the installation of fire safety tools and
equipment’s Rs 5, 15,000 and Rs 9, 50,000 for one stop mall and Hanuman central
respectively. Originality/Value: The study may be a guiding document for professionals, entrepreneurs
and occupants for improving awareness and preparedness by obtaining cost components
against fire safety. Paper Type: Research paper Keywords: Fire safety installation, Safety in commercial buildings, Preparedness and costing. 1 INTRODUCTION : Keywords: Fire safety installation, Safety in commercial buildings, Preparedness and costing. Keywords: Fire safety installation, Safety in commercial buildings, Preparedness and costing. 2 Professor, Institute of Management & Commerce, Srinivas University, Mangalore, India,
OrcidID: 0000-0002-4691-8736; E-mail:psaithal@gmail.com
Subject Area: Project Management.
Type of the Paper: Conceptual Research.
Type of Review: Peer Reviewed as per |C|O|P|E| guidance.
Indexed In: OpenAIRE.
DOI: https://doi.org/10.5281/zenodo.7108648
Google Scholar Citation: IJAEML
International Journal of Applied Engineering and Management Letters (IJAEML)
A Refereed International Journal of Srinivas University, India.
Crossref DOI: https://doi.org/10.47992/IJAEML.2581.7000.0152
Received on: 04/09/2022
Published on: 24/09/2022
© With Authors.
This work is licensed under a Creative Commons Attribution-Non-Commercial 4.0
International License subject to proper citation to the publication source of the work.
Disclaimer: The scholarly papers as reviewed and published by the Srinivas Publications
(S.P.), India are the views and opinions of their respective authors and are not the views or
opinions of the S.P. The S.P. disclaims of any harm or loss caused due to the published content
to any party.
How to Cite this Paper:
Mishra, A. K., & Aithal, P. S., (2022). Preparedness and Costing on Fire Safety Installation
in Commercial Buildings. International Journal of Applied Engineering and Management
Letters (IJAEML), 6(2), 182-196. DOI: https://doi.org/10.5281/zenodo.7108648 © With Authors. This work is licensed under a Creative Commons Attribution-Non-Commercial 4.0
International License subject to proper citation to the publication source of the work. Disclaimer: The scholarly papers as reviewed and published by the Srinivas Publications
(S.P.), India are the views and opinions of their respective authors and are not the views or
opinions of the S.P. The S.P. disclaims of any harm or loss caused due to the published content
to any party. This work is licensed under a Creative Commons Attribution-Non-Commercial 4.0
International License subject to proper citation to the publication source of the work. Disclaimer: The scholarly papers as reviewed and published by the Srinivas Publications
(S.P.), India are the views and opinions of their respective authors and are not the views or
opinions of the S.P. The S.P. disclaims of any harm or loss caused due to the published content
to any party. This work is licensed under a Creative Commons Attribution-Non-Commercial 4.0
International License subject to proper citation to the publication source of the work. Disclaimer: The scholarly papers as reviewed and published by the Srinivas Publications
(S.P.), India are the views and opinions of their respective authors and are not the views or
opinions of the S.P. The S.P. disclaims of any harm or loss caused due to the published content
to any party. PAGE 182 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
PUB International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION As the development works are rapidly increasing day by day fire safety should become a topic to be
considered. OHS should be given first priority by concerned authority and management. Although
firefighting is top priority for management in big metropolitan city like Kathmandu (capital city of
Nepal) and Pokhara, newly emerged city like Damak, Biratnagar, Birtamode are seems to be unaware
and negligent in this topic. Management, concerned authority and occupants are also seems to be
unaware on this topic and fire has been kept on the least priority by them. Firefighting preparedness
is not a priority for government compared to earthquake (UNDP, 2009) [4]. Fire wellbeing code was
formed in 1994 A.D. what's more, after then it has quite recently turned into a piece of report and
furthermore was never overhauled till now. Indeed, UNDP has expressed the requirement for present
day and state-of-the-art fire code. Since NBC 107:1994 [5] is definitely not a specialized yet in
addition an authoritative record, it should be refreshed in light of changing conditions to guarantee fire
safety. So the researcher has selected two commercial building and assessed fire safety in terms of
awareness and preparedness of Management, Concerned authority and Occupants and also calculated
the tentative cost of installation of fire safety tools and equipment’s in the buildings. 1. INTRODUCTION : Design is the mother of construction [1]. Design of Modern Malls and complex in Urban Areas seem
to consider the fire safety in some manner. But the available facility and measures are more like a
formality for owners. Date expired extinguisher, lack of emergency exit, poor design, more
commercial space than flowing space, owners motive of profit maximization, lack of awareness, lack
of electrical drawings, poor electrical connection, low quality electrical material, unmanaged use of
electrical appliances due to weather, less knowledge on fire safety, etc. are the major issues seen [2,
3]. PAGE 183 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
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PUBLICATION International Journal of Applied Engineering and Management
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PUBLICATION The study area is one stop mall and Hanuman Central located at Birtamode, Jhapa, Nepal. It can be
important document for all three parties for increasing their awareness and preparedness in case of
emergency. It can be helpful for the management about the cost of additional fire safety tools and
equipment’s that can be quickly installed. It can be helpful for new Designers and Developers about
the fire safety practice and importance of it in case of emergency. 3. OBJECTIVES : The Objective is to assess the status of awareness and preparedness of management, concerned
authority and occupants along with the cost of installation of fire safety tools and equipment’s for
Selected Commercial Buildings at Birtamode, Jhapa, Nepal. 4. LITERATURE REVIEW : 4.1 Fire Safety Preparedness: Readiness involves adjusting one's conduct to decrease the effect of catastrophes on people (Drabek,
1986) [3, 8]. It is a constant pattern of arranging, making due, sorting out, preparing, preparing,
working out, making, assessing, checking, and further developing exercises to guarantee powerful
coordination and the improvement of capacities of concerned associations in forestalling, safeguarding
against, answering, recuperating from, making assets, and moderating the impacts of catastrophic
events, demonstrations of psychological oppression, and other man-made calamities [3, 9]. During the
preparedness phase, emergency managers create strategies to manage and mitigate their risks, as well
as take steps to acquire the required capabilities to put those plans into effect. Common preparedness measures include: Common preparedness measures include: • Communication plans that use vocabulary and methods that are simple to understand. • Emergency services should be properly maintained and trained. • Emergency services should be properly maintained and trained. • The creation and testing of emergency population warning systems, as well as emergency shelters
and evacuation strategies. • Establish and maintain an emergency communication system that can assist in determining the
nature of an emergency and providing instructions as necessary (Ogajo, 2013) [3, 10]. 4.2 Fire Safety Management: 2. STATEMENT OF PROBLEMS : In July 27, 2019 Fire erupted in Hanuman Central at Birtamode which incurred loss of 20
million. The building caught a flame due to short circuit in an air conditioner machine of shoe shop
on ground floor of six storey building [6]. A few months back fire erupted in Sri Krishna market which
incurred loss of 15 million. Fire was started from an electronic shop go down around 1:00 – 2:00 am
and was taken under control only in 7 am in the morning. Fire fighters from Birtamode Municipality
and Mechinagar Municipality along with Nepali Army were able to control the fire. Fig. 1: Fire at Namaste Ply indsurty at chaitubari, Birtamode -2 (RSS) [7] Fig. 1: Fire at Namaste Ply indsurty at chaitubari, Birtamode -2 (RSS) [7] In Dec 4, 2020 A gigantic fire broke out at the Namaste Handle Industry at Chaitubari, Birtamod-2 in
Jhapa locale and destroyed property worth Rs 3 million. According to Representative Director of
Police of Region Police Office Jhapa, Rakesh Thapa, a short out prompted the fire.As police got the
data, a joint group comprising the Region Police Office Anarmani, the Territory 1 Debacle The board
Organization Chandragadhi and the Furnished Police Power Headquarters Sanischare went for the
rescue.Locals and security staff cooperated to drench the fire utilizing fire motors from Damak,
Bhadrapur and Birtamod. In case of Nepal, Safety seems to be of less priority either in design or in operational phase. Owner
doesn’t seem to be aware and prepared of unforeseen situation. Concerned authority seems weak in
implementing the fire safety awareness and fire safety code. NBC Fire code has become just a
document. This research mainly focuses on current fire safety condition existing commercial building
and awareness and preparedness of management, occupants and concerned authority and prepare a
tentative budget for installation of fire safety tools and equipment’s. PAGE 184 A. K. Mishra, et al. (2022); www.srinivaspublication.com 4.2 Fire Safety Management: In terms of fire prevention and safety, management is critical. A building's fire safety design consists
of a range of measures relating to the layout, structure, and other requirements, some of which are
activated in the event of a fire (Malhotra, 1993) [11]. Standard investigation, support, posting of notification and exit coordinated signs, normal fire works
out, departure methodologies, and the presence of fire superintendents are all essential for fire security
the executives (Ogajo, 2013) [10]. The essential objective of fire security the board is to ensure that all open fire wellbeing measures are
accessible for people to use in order to aid their escape (Baker, 2003) [12]. y
g
Exit Requirement: q
An entryway, hall, or path prompting an inside flight of stairs, an outside flight of stairs, a verandah
going to the road, the top of a structure, or the road is viewed as an exit. The egress could perhaps lead
to another structure in the area The exit should to another structure in the area. The exit should to another structure in the area. The exit should • Meet the minimum size criteria and enable for the evacuation of all people in a reasonable amount
of time. • Be devoid of any impediments and provide no resistance to movement. devoid of any impediments and provide no resistance to movement. • Be easily seen, especially with appropriate indicators. Be consistent and do not encroach on personal space. i • Be consistent and do not encroach on personal space. Stairs: • Be consistent and do not encroach on personal space. Stairs: • At least two fire escape staircases, one internal and the other external. • When a building's plinth area reaches 500 m2, more steps must be built in proportion to the
increased plinth space. • The minimum width is 1.5 meters. • The minimum width is 1.5 meters. • The distance between any point in a hallway and a building's stairway must not exceed 20 meters. Fire Escapes: The Building having 5 or more stories must provide a separate escape with of at least 75 cm width
Each riser on the fire escape must be no more than 19 cm high and 20 cm width The Building having 5 or more stories must provide a separate escape with of at least 75 cm width
Each riser on the fire escape must be no more than 19 cm high and 20 cm width. The Building having 5 or more stories must provide a separate escape with of at least 75 cm w
• Each riser on the fire escape must be no more than 19 cm high and 20 cm width. • Each riser on the fire escape must be no more than 19 cm high and 20 cm width. • The number of risers each flight must not exceed 15. • This type of fire escape should lead users to an open area. • This type of fire escape should lead users to an open area. s type of fire escape should lead users to an open area. Exit Doors: Leave entryways should open to a way or a hallway, however not to a way or a passageway, and they
should open outwards without impeding the development of individuals passing external the entryway. The greatest distance between any point in the course and such a leave entryway is 20 meters. The
width and level of the leave entryway should not be under 90 cm and 180 cm, separately. 4.5 Nepal National Building Code NBC 107: 1994: p
g
Nepal national building code NBC 107:1994 provides fundamental requirements for fire safety in
ordinary buildings (GoN, 1994). Summary of NBC 107: 1994 are listed below [2, 5]. Exit Requirement: Nepal national building code NBC 107:1994 provides fundamental requirements for fire safety in
ordinary buildings (GoN, 1994). Summary of NBC 107: 1994 are listed below [2, 5]. Exit Requirement: International Journal of Applied Engineering and Management
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PUBLICATION 4.4 Fire Safety Code: 4.3 Fire Safety Management Roles and Responsibilities: Fire Safety Management Roles and Responsibilities: re Safety Management Roles and Responsibilities: In a business facility, pre-planned processes for dealing with a fire emergency are required. The roles
and responsible of fire safety management are listed below. Roles and Responsibilities: In a business facility, pre-planned processes for dealing with a fire emergency are required. The roles
and responsible of fire safety management are listed below. Roles and Responsibilities: p
Select a fire security chief to guarantee that fire wellbeing conventions are followed and that upkeep
needs are met. Guarantee that staff knows about the wellbeing strategies and the tenants complete their
work in a protected way. An upkeep timetable ought to be produced for all fire recognition/caution frameworks, fire dousers,
hose reels, and other comparable things, and records of their review and fix ought to be safeguarded. Hold regular fire drills for representatives to guarantee that they know about the legitimate method for
get out, leave areas, and courses to take in case of a fire. Make arrangements for evacuation, fire suppression, and aid to the fire department when they arrive. The strategy should be addressed with the fire department, and their agreement on the planned
processes should be acquired [2, 3, 11]. PAGE 185 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
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PUBLICATION International Journal of Applied Engineering and Management
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PUBLICATION 4.4 Fire Safety Code: A building code is a rule or guideline that establishes the minimum standards for the design and
construction of structures and buildings. The goal of establishing those basic requirements is to
safeguard society's health and safety [13]. In new and existing buildings, structures, and processes, the fire safety code addresses fire prevention,
fire protection, life safety, and the proper storage and use of hazardous materials. They give a
comprehensive approach to hazard control in all structures and sites (International Code Council,
2009). The fire safety requirements for buildings were researched using NBC 107: 1994 [2, 5]. 5.1 Study Area: y
To assess the awareness and preparedness of Management, concerned authority and occupants
Buildings selected were one stop mall and Hanuman Central Birtamode. To assess the awareness and preparedness of Management, concerned authority and occupants
Buildings selected were one stop mall and Hanuman Central Birtamode. One Stop Mall: International Journal of Applied Engineering and Management
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PUBLICATION Hanuman central is also located at Birtamode, Jhapa touching national highway (Mahendra Highway). It is also known as Hanuman Complex. It was the first Commercial complex build in Birtamode. It
was opened in 2012. p
It is a Rectangular shaped building with atrium in the center. Vertical circulation is aided by elevators
and stairwells, whereas horizontal circulation is aided by walkways and lobby. There are 3 staircases
in the building with 3 elevators. 5.2. Research Philosophy: p y
The descriptive along with field based empirical research design was adopted in this research work. Figure 1 depicts the different stages applied in the research [14]. p y
The descriptive along with field based empirical research design was adopted in this research work. Figure 1 depicts the different stages applied in the research [14]. Figure 1 depicts the different stages applied in the research [14]. Fig. 1: Research Onion
The overall research philosophy is Pragmatism is in which only "solution" obtained from observation
with reliable assessment and objective interpretation. The study outcomes in these sorts of
investigations are frequently apparent and quantitative. Abduction approach for logical reasoning was followed through inductive and deductive ways. Regarding methodological choice, both qualitative as well as quantitative methods were followed. So
it’s a mixed method. The research strategy was survey. As the data were collected through the different
Key informant Interview, Survey and observation, this study was ex-post facto field based. The survey
was done on longitudinal time horizon. The data collection methods were observation method, survey,
and interview. Pragmatism
Abductive
Mixed
(Qualitative+Quantitative)
Survey
Longitudinal
Observation
& Survey Fig. 1: Research Onion g
The overall research philosophy is Pragmatism is in which only "solution" obtained from observation
with reliable assessment and objective interpretation. The study outcomes in these sorts of
investigations are frequently apparent and quantitative. Abduction approach for logical reasoning was followed through inductive and deductive ways. Regarding methodological choice, both qualitative as well as quantitative methods were followed. So
it’s a mixed method. The research strategy was survey. As the data were collected through the different
Key informant Interview, Survey and observation, this study was ex-post facto field based. The survey
was done on longitudinal time horizon. The data collection methods were observation method, survey,
and interview. g
One Stop Mall: One stop Mall is Located at Birtamode, Jhapa. It is located at North- west side from mukti chowk
(Central part of Birtamode). The building, which is a shopping complex with six stories above ground,
was finished in 2016. Three floors including the ground are used for shops and top three floors are
used for Cinema purpose. Building is U-shaped with a large steel structure at front connecting to
building. The structure has a variety of shops selling various goods and services, as well as leisure
venues such as a movie, a game center, and a pub. Underground Parking Facility is also available but
the building structural components don’t have any connection with the parking below. Vertical
circulation is assisted by escalators, elevators, and stairwells, while horizontal circulation is assisted
by walkways and lobby. C
l Hanuman Central: Hanuman Central: Hanuman Central: PAGE 186 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
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y
Based on the information from the literature review questionnaire was formulated. A survey of the
occupants of selected business buildings was done. Management of selected commercial buildings and
concerned authority. It was conducted to understand fire safety awareness and preparedness of all
three parties. Several references were takes into considerations [15, 16, 17, 18, 19, 20, and 21]. Key Informants Interview PAGE 187 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
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Key informants interview was done with Head of management of Hanuman central and One stop mall,
head of fire department( Birtamode Municipality) and Engineer of Birtamode Municipality( Building
Department) [15, 16, 17, 18, 19, 20, and 21]. International Journal of Applied Engineering and Management
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PUBLICATION International Journal of Applied Engineering and Management
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PUBLICATION Key informants interview was done with Head of management of Hanuman central and One stop mall,
head of fire department( Birtamode Municipality) and Engineer of Birtamode Municipality( Building
Department) [15, 16, 17, 18, 19, 20, and 21]. 5.4 Methodology in Brief: Table 1: Summary of Methodology
Objective
Activities
Analysis
1. To identify the level of awareness
among management, occupants
and concerned authority. 1. Key Informant
Interview
2. Questionnaire survey
1. Qualitative
Analysis
2. Quantitative
Analysis
2. To calculate the tentative cost of
installation of fire safety tools and
Equipment’s. 1. Rate collection and
calculation of cost. 1. Qualitative
Analysis Table 1: Summary of Methodology 6.1 Fire Safety Preparedness of Management, Occupants and Concerned Authority: 6.1 Fire Safety Preparedness of Management, Occupants and Concerned Authority: The data from the questionnaire was used to create the graphs below, which show the
inhabitants', management's, and concerned authority's level of fire safety readiness. 6.1.1 Awareness and Preparedness of Occupants on Fire Safety: 6.1.1 Awareness and Preparedness of Occupants on Fire Safety: The table 2 shows the level of fire safety readiness among the tenants of all chosen business
buildings, i.e., one Stop Mall and Hanuman Central Birtamode. Table 2: Awareness on Building Feature
Hanuman Central
One stop Mall
Awareness on building
feature
Yes
No
Can’t Say
Yes
No
Can’t Say
Building Design
10
12
18
20
19
1
Location of exit
28
11
1
27
13
0
Visibility of emergency
sign
4
22
14
26
10
4 ndents were asked to state whether or not they were aware of a certain building feature. Respondents were asked to state whether or not they were aware of a certain building feature. The respondent's response is shown in Table 4.21. Ten Hanuman Central respondents were
aware of the building design, 28 were aware of the position of the exit, and four respondents
had emergency signs visible. 12 respondents were unfamiliar of the building layout, 11 were
unaware of the position of the escape, and 22 respondents did not see an emergency sign. 18
respondents had no clue about the structure plan, 1 had no data about the position of the exit,
and 14 had no clue about the crisis sign. 6.1.3 Awareness of Emergency Egress: 20 respondents of Hanuman Central knew about crisis clearing, 12 respondents knew about get
together point. 18 respondents didn't know crisis clearing, 16 were uninformed about gathering point. Though, 2 respondents had no clue about crisis departure, 12 respondents had no clue about gathering
point. Table 3: Awareness of Emergency Egress
Hanuman Central (nos. of
occupants)
One stop mall (nos. of occupants)
Awareness on emergency
Egress
Yes
No
Can’t Say Yes
No
Can’t Say
Emergency
evacuation
20
18
2
22
7
11
Assembly point
12
16
12
14
20
6
52.5% of respondents were found aware, 31.25% of respondents were not aware and 16.25% can’t say
about emergency evacuation. 32.5% were aware, 45% were aware and 22.5% cants ay about Assembly
point. It shows that tenants needed mindfulness on crisis departure. A. K. Mishra, et al. (2022); www.srinivaspublication.com
PAGE 189
Table 3: Awareness of Emergency Egress
Hanuman Central (nos. of
occupants)
One stop mall (nos. of occupants)
Awareness on emergency
Egress
Yes
No
Can’t Say Yes
No
Can’t Say
Emergency
evacuation
20
18
2
22
7
11
Assembly point
12
16
12
14
20
6
52.5% of respondents were found aware, 31.25% of respondents were not aware and 16.25% can’t say
about emergency evacuation. 32.5% were aware, 45% were aware and 22.5% cants ay about Assembly
point. It shows that tenants needed mindfulness on crisis departure. Table 3: Awareness of Emergency Egress
Hanuman Central (nos. of
occupants)
One stop mall (nos. of occupants)
Awareness on emergency
Egress
Yes
No
Can’t Say Yes
No
Can’t Say
Emergency
evacuation
20
18
2
22
7
11
Assembly point
12
16
12
14
20
6 52.5% of respondents were found aware, 31.25% of respondents were not aware and 16.25% can’t say
about emergency evacuation. 32.5% were aware, 45% were aware and 22.5% cants ay about Assembly
point. It shows that tenants needed mindfulness on crisis departure. 52.5% of respondents were found aware, 31.25% of respondents were not aware and 16.25% can’t say
about emergency evacuation. 32.5% were aware, 45% were aware and 22.5% cants ay about Assembly
point. It shows that tenants needed mindfulness on crisis departure. 52.5% of respondents were found aware, 31.25% of respondents were not aware and 16.25% can’t say
about emergency evacuation. 32.5% were aware, 45% were aware and 22.5% cants ay about Assembly
point. It shows that tenants needed mindfulness on crisis departure. A. K. Mishra, et al. 6.1.2 Awareness of Building Feature: Twenty one-stop mall respondents were aware of the building design, 27 were aware of the position
of the exit, and 26 respondents reported that emergency signs were visible. The building design was PAGE 188 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
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unknown to 19 respondents, the location of the exit was unknown to 13 respondents, and the
emergency sign was not visible to 10 respondents. One individual had no clue about the structure
format, 0 had no clue about the position of the exit, and 4 had no clue about the crisis sign. Fig. 2: Awareness of Building Feature
Percentage of respondants
80
70
60
50
40
30
20
10
0
Building Plan
Location of Exit
Visibility of Emergency
Cannot Say
23.75
1.25
22.5 International Journal of Applied Engineering and Management
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unknown to 19 respondents, the location of the exit was unknown to 13 respondents, and the
i
i ibl
10
d
O
i di id
l h d
l
b
h International Journal of Applied Engineering and Management
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unknown to 19 respondents, the location of the exit was unknown to 13 respondents, and the
emergency sign was not visible to 10 respondents. One individual had no clue about the structure
format, 0 had no clue about the position of the exit, and 4 had no clue about the crisis sign. wn to 19 respondents, the location of the exit was unknown to 13 respondents, and the
ency sign was not visible to 10 respondents. One individual had no clue about the structure
, 0 had no clue about the position of the exit, and 4 had no clue about the crisis sign. emergency sign was not visible to 10 respondents. One individual had no clue about the structure
format, 0 had no clue about the position of the exit, and 4 had no clue about the crisis sign. Fig. 6.1.2 Awareness of Building Feature: 2: Awareness of Building Feature
Percentage of respondants
80
70
60
50
40
30
20
10
0
Building Plan
Location of Exit
Visibility of Emergency
Cannot Say
23.75
1.25
22.5 Fig. 2: Awareness of Building Feature 37.5% of respondents knew about building Plan, 38.75% of respondents don’t know about plan and
23.75 cannot say. 68.75% have knowledge about exit, 30% don’t know and 1.25 cannot say about it,
similarly 37.5% knows about emergency sign, 40% haven’t seen and 22.5% cannot say about that. 6.1.3 Awareness of Emergency Egress: International Journal of Applied Engineering and Management
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PUBLICATION Supervisory group ought to much of the time arrange normal fire drills to guarantee that inhabitants
stay mindful of the proper method for get out and where they will stand by after crisis departure. The
results are in similar trends to that of study conducted in Kathmandu [3], 22 respondents of LABIM
Shopping center knew about crisis clearing, 14 respondents knew about get together point. 7
respondents didn't know about crisis clearing, 20 were uninformed about gathering point. Though, 11
respondents had no clue about crisis departure, 6 respondents had no clue about gathering point. A. K. Mishra, et al. (2022); www.srinivaspublication.com 6.1.3 Awareness of Emergency Egress: (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
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Supervisory group ought to much of the time arrange normal fire drills to guarantee that inhabitants
stay mindful of the proper method for get out and where they will stand by after crisis departure. The
results are in similar trends to that of study conducted in Kathmandu [3], 22 respondents of LABIM
Shopping center knew about crisis clearing, 14 respondents knew about get together point. 7
respondents didn't know about crisis clearing, 20 were uninformed about gathering point. Though, 11
respondents had no clue about crisis departure, 6 respondents had no clue about gathering point. International Journal of Applied Engineering and Management
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bother with help to arrive at spot of wellbeing, 12 respondents accepted help won't be given during
fire crisis to arrive at spot of security. Though, 14 respondents couldn't say assuming they required
help, and 20 respondents had no clue in the event that help would be given during fire crisis. Table 4: Provision of Assistance
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of
occupants)
Provision of assistance
Yes
No
Can’t say
Yes
No
Can’t say
Need emergency assistance
15
10
15
5
21
14
Assistance
provided
7
17
16
8
12
20
5 respondents of One stop Shopping center required help to arrive at spot of security, 8 respondents
expected help will be given during fire crisis to arrive at spot of wellbeing. 21 respondents needn't
bother with help to arrive at spot of wellbeing, 12 respondents accepted help won't be given during
fire crisis to arrive at spot of security. Though, 14 respondents couldn't say assuming they required
help, and 20 respondents had no clue in the event that help would be given during fire crisis. 5 respondents of One stop Shopping center required help to arrive at spot of security, 8 respondents
expected help will be given during fire crisis to arrive at spot of wellbeing. 21 respondents needn't
bother with help to arrive at spot of wellbeing, 12 respondents accepted help won't be given during
fire crisis to arrive at spot of security. Though, 14 respondents couldn't say assuming they required
help, and 20 respondents had no clue in the event that help would be given during fire crisis. 5 respondents of One stop Shopping center required help to arrive at spot of security, 8 respondents
expected help will be given during fire crisis to arrive at spot of wellbeing. 21 respondents needn't
bother with help to arrive at spot of wellbeing, 12 respondents accepted help won't be given during
fire crisis to arrive at spot of security. Though, 14 respondents couldn't say assuming they required
help, and 20 respondents had no clue in the event that help would be given during fire crisis. p
p
p
g
g
Fig. 3: Awareness on Assistance
5
0
4
5
4
0
3
5
3
0
2
5
2
45
36
.2
Can't
Say
36
.2
38
.7
No
18
.7
25
Yes
Asistance Provided
Need Emergency Assistance
Percentage of Respondants A. K. 6.1.4 Provision of Assistance: 15 respondents of Hanuman Central assumed they need help to arrive at spot of security, 12
respondents expected help will be given during fire crisis to arrive at spot of wellbeing. 10 respondents
needn't bother with help to arrive at spot of wellbeing, 17 respondents expected help won't be given
during fire crisis to arrive at spot of security. Though, 15 respondents couldn't say assuming they
required help, and 16 respondents had no clue in the event that help would be given during fire crisis. Table 4: Provision of Assistance
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of
occupants)
Provision of assistance
Yes
No
Can’t say
Yes
No
Can’t say
Need emergency assistance
15
10
15
5
21
14
Assistance
provided
7
17
16
8
12
20
5 respondents of One stop Shopping center required help to arrive at spot of security, 8 respondents
expected help will be given during fire crisis to arrive at spot of wellbeing. 21 respondents needn't
bother with help to arrive at spot of wellbeing, 12 respondents accepted help won't be given during
fire crisis to arrive at spot of security. Though, 14 respondents couldn't say assuming they required
help, and 20 respondents had no clue in the event that help would be given during fire crisis. Fig. 3: Awareness on Assistance
5
0
4
5
4
0
3
5
3
0
2
5
2
45
36
.2
Can't
Say
36
.2
38
.7
No
18
.7
25
Yes
Asistance Provided
Need Emergency Assistance
Percentage of Respondants Table 4: Provision of Assistance
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of
occupants)
Provision of assistance
Yes
No
Can’t say
Yes
No
Can’t say
Need emergency assistance
15
10
15
5
21
14
Assistance
provided
7
17
16
8
12
20 Table 4: Provision of Assistance
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of
occupants)
Provision of assistance
Yes
No
Can’t say
Yes
No
Can’t say
Need emergency assistance
15
10
15
5
21
14
Assistance
provided
7
17
16
8
12
20
5 respondents of One stop Shopping center required help to arrive at spot of security, 8 respondents
expected help will be given during fire crisis to arrive at spot of wellbeing. 6.1.5 Awareness on Fire Safety Measures: Respondents of Hanuman Central were mindful about the presence of fire douser, 10 respondents
knew about the presence of water Hydrant in building. 10 respondents were uninformed about the
presence of fire quencher, 12 respondents were ignorant about the presence of water hydrant and 16
respondents didn't know about fire douser, 18 respondents didn't know about water hydrant. Table 5: Awareness on Fire Safety Measures
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of occupants)
Fire safety Measures
Exist
Do
not exist
Not sure
Exist
Do
not exist
Not sure
Fire
extinguisher
14
10
16
15
3
22
Water hose reel/water
hydrant
10
12
18
12
10
18 Table 5: Awareness on Fire Safety Measures Respondents of One stop knew about the presence of fire douser, 12 respondents knew about
the presence of water hose reel in building. 3 respondents were uninformed about the presence
of fire quencher, 10 respondents were ignorant about the presence of water hose reel and 22
respondents didn't know about fire douser, 18 respondents didn't know about water hose reel. International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION 25% of respondents expected help to lead them to place of wellbeing during fire crisis, 38.75% needn't
bother with any help and 36.25% tenants can't say regarding that. 18.75% accepted that help would be
given to during crisis, 36.25% of respondents imagine that help wouldn't be given during crisis and
45% can't say regarding that. 6.1.5 Awareness on Fire Safety Measures: A. K. Mishra, et al. (2022); www.srinivaspublication.com 6.1.4 Provision of Assistance: Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION
25% of respondents expected help to lead them to place of wellbeing during fire crisis, 38.75% needn't
bother with any help and 36.25% tenants can't say regarding that. 18.75% accepted that help would be
given to during crisis, 36.25% of respondents imagine that help wouldn't be given during crisis and
45% can't say regarding that. International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION 6.1.6 Ability to Operate Fire Safety Equipment: y
p
y
q
p
10 respondents of Hanuman Central think they had the option to work fire quencher, 5 respondents
expected they had the option to work water hose reel/water hydrant. 22 respondents couldn't work fire
douser, 25 respondents couldn't work water hose reel. While, 8 respondents were don't know whether
they could work fire quencher, 10 respondents were don't know whether they could work water hose
reel. A. K. Mishra, et al. (2022); www.srinivaspublication.com
PAGE 191
Table 6: Ability to Operate Fire Safety Equipment
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of
occupants)
Ability To Operate Fire
Equipment
Able to
Not able
to
Not sure
Able to
Not able
to
Not sure
Fire extinguisher
10
22
8
13
18
9
Water hose
reel/water hydrant
5
25
10
2
30
8
13 respondents of One stop Shopping center accepted they had the option to work fire douser, 2
respondents expected they had the option to work water hose reel/water hydrant. 18 respondents Table 6: Ability to Operate Fire Safety Equipment
Hanuman Central (nos. of
occupants)
One Stop Mall (nos. of
occupants)
Ability To Operate Fire
Equipment
Able to
Not able
to
Not sure
Able to
Not able
to
Not sure
Fire extinguisher
10
22
8
13
18
9
Water hose
reel/water hydrant
5
25
10
2
30
8
13 respondents of One stop Shopping center accepted they had the option to work fire douser, 2
respondents expected they had the option to work water hose reel/water hydrant. 18 respondents Table 6: Ability to Operate Fire Safety Equipment 13 respondents of One stop Shopping center accepted they had the option to work fire douser, 2
respondents expected they had the option to work water hose reel/water hydrant. 18 respondents 13 respondents of One stop Shopping center accepted they had the option to work fire douser, 2
respondents expected they had the option to work water hose reel/water hydrant. 18 respondents A. K. Mishra, et al. (2022); www.srinivaspublication.com PAGE 191 International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION SRINIVAS
PUBLICATION couldn't work fire douser, 30 respondents couldn't work water hose reel. While, 9 respondents were
don't know whether they could work fire quencher, 8 respondents were don't know whether they could
work water hose reel/water hydrant.28.75% of respondents had the option to work fire quencher, half
of respondent couldn't work and 21.25% were don't know. 8.75% had the option to work water hose
reel/water hydrant, 68.75 couldn't and 22.5% were don't know. Table 6 shows that of tenants needed
capacity to work the firefighting hardware's. Consequently, appropriate preparation on it will be given
to the inhabitants habitually so the misfortune during fire can be decreased. Equipment's with sensor
system could be also an effective way as in case of theft most of shopkeeper have found to install
sensor based siren which alerts the police on real time about its unauthorized opening. For this step,
direct interview was conducted with management. Management seems more focused on financial part
of building. Fire extinguisher was only tool for fire safety. There was smoke detector and alarm system
installed in One Stop Mall. Management of both the buildings have requested for installation of fire
hydrant to Municipality many times. Water hydrant installed in both the building weren’t in working
condition. Fire fighter of fire department of Birtamode had many problems regarding communication, training
and information about new technology. Concerned authority (Birtamode Municipality) seems weak in
implementation of code and law. They are not aware about the fire safety condition of existing
commercial buildings. No any seminar or awareness program has been conducted by municipality for
fire safety. Even the Municipality building didn’t have proper sign for safe evacuation. Fire fighter
seems always prepared but due to lack of direct communication they find hard to locate the site. Training for firefighter has been very often. Municipality seems just happy in collecting the fire tax
while registration of new house. A. K. Mishra, et al. (2022); www.srinivaspublication.com 6.2.1 Cost estimate For One Stop Mall: 6.2.1 Cost estimate For One Stop Mall:
Rates are collected from different website like (Daraz.com and sastodeal.com) and other rates are
collected from expert and experience of researcher. p
Rates are collected from different website like (Daraz.com and sastodeal.com) and other rates are
collected from expert and experience of researcher. Table 7: Cost estimate For One Stop Mall
S. No. Description
No
Unit
Rate
Quantity
Remarks
1
Exit Signs and boards(Digital)
10
per piece
2500
25000
2
Underground water tank for
water hydrant (maintenance)
1
L/s
100000
100000
for functioning of
existing water tank
3
Fire extinguisher (4.5 kg) CO2
5
per piece
12000
60000
4
Fire Blanket
5
per piece
5000
25000
for each floor
5
Foam fire extinguisher
5
per piece
5000
25000
for each floor
6
Smoke detector
40
per piece
2500
100000 10 for each floor up
to fourth
floor
7
Smoke detector Accessories and
installation charge
1
job
50000
50000
8
Fire drills
1
job
100000
100000
expert fee,
management,
accessories fee
9
Awareness program
1
job
25000
25000
expert fee,
management,
accessories fee
10
Coordination with concerned
authority
1
job
5000
5000
regular visit for
firefighter,
interaction International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION
Total
515000
Amount in words: - Five lakh fifteen thousand Only
Note: - The above estimates are done on current market rate which may subject to change anytime. As
mentioned it is a tentative estimate, so for exact estimate detail cost calculation should be done. Table 8: Cost estimate For Hanuman central
S. No. 6.2.1 Cost estimate For One Stop Mall: Description
No
Unit
Rate
Quantity
Remarks
1
Exit Signs and boards(Digital)
20
per piece
2500
50000
-
2
Overhead water tank for water
hydrant
1
L/s
250000
250000
for functioning of
existing water tank
on top floor
3
Fire Blanket
12
per piece
5000
60000
for 4 floor , at
staircase i:e, 3
staircase
4
Foam fire extinguisher
5
per piece
5000
25000
for each floor
5
Smoke detector
150 per piece
2500
375000
for each shop
6
Smoke detector Accessories and
installation charge
1
job
10000
10000
-
7
Fire drills
1
job
100000
100000
expert fee,
management,
accessories fee
8
Awareness program
1
job
25000
25000
expert fee,
management,
accessories fee
9
Coordination with concerned
authority
1
job
5000
5000
regular visit for
authority(Fire
brigade)
10
Emergency alarm system
1
job
50000
50000
Total
950000
Amount in words: - Nine lakh fifty thousand Only Amount in words: - Five lakh fifteen thousand Only Note: - The above estimates are done on current market rate which may subject to change anytime. As mentioned it is a tentative estimate, so for exact estimate detail cost calculation should be done. Rates are collected from different website like (Daraz.com and sastodeal.com) and other rates are
collected from expert and experience of researcher. The cost of installation is not meaningful in comparison to that of safety assured. This once again
verifies the standard theory of safety is investment with positive return, not expenditure as extra
burden. Assume chance of fire is very less but the impact once caught is very high so we must assure
fire safety system. For concerned authority: (1) Fire safety code NBC 107:1994 should be strictly implemented. (1) Fire safety code NBC 107:1994 should be strictly implemented. (2) Municipality should develop their own guideline according to the geography and literacy. (3) Municipality should organize site visit for firefighter to important commercial and public
buildings. (4) Municipality should strictly implement the fire safety code for newly registered buildings inside
its area. (5) The narrow road should be expanded as soon as possible. (6) New technology should be introduced to firefighter so that they can find exact location, should
organize regular training to the fire fighters. g
g
g
g
(7) Municipality should organize regular awareness program and seminar using different media. (8) Municipality should install fire hydrant in front of important commercial and public building (8) Municipality should install fire hydrant in front of important commercial a International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVA
PUBLICATION management in existing commercial buildings, no fire drills were held, and inhabitants received no
awareness or training. There was huge communication gap between all three parties. Management and
occupants don’t have direct communication (phone number) with fire brigade. In case of emergency
people use to call Nepal police and Nepal police then call the fire fighter. The gap creates the delay
for response. Also due to lack of new technology fire fighter finds hard to find the exact location. There was a problem for refilling the tank as there was no any fire hydrant installed by municipality. Fire fighters have to travel at least one kilometre to refill their tank. Due to unmanaged planning and
narrow road fire fighter find hard to reach the location. There was no any coordination among the
management, occupants and concerned authority. Lack of fire expert in the Birtamode area was also a major problem for management to perform fire
drills for occupants. The addition of fire safety tools and equipment’s to the existing buildings seems
less as compared to the loss that had occurred in the past due to fire in the buildings. Also the weakness
of Municipality in implementing fire safety codes and guidelines for the large commercial buildings
was a major problem seen during study. Fire extinguisher and Fire-fighter from municipality seems to
be only tool to fight in case of emergency. The following points can be recommended for the Management: The following points can be recommended for the Management: (1) The calculated cost of the addition of fire safety tools and equipment’s seems less as compared to
loss that might occur, so more fire safety tools and equipment’s should be added as soon as possible. (2) Fire drill and awareness program should be conducted frequently in coordination of concerned
authority with the help from expert. y
p
p
(3) The installed system (Water hydrant and water hose reels) should be made functional as soon as
possible in both buildings. (3) The installed system (Water hydrant and water hose reels) should be made functional as soon as
possible in both buildings. (4) Management should also focus on safety of occupants and put safety on top priority list. (5) Different signs and boards should be installed for awareness of occupants. For occupant (5) Different signs and boards should be installed for awareness of occupants. For occupants
(6) Occupants should always involve in safety program organized by authority and management (6) Occupants should always involve in safety program organized by authority and management
(7) Occupants also should be aware about building features, fire safety tools and equipment’s. ( )
p
y
y p
g
g
y
y
g
(7) Occupants also should be aware about building features, fire safety tools and equipmen ( )
p
g
y
q p
(8) Occupants should coordinate with the management for safe and healthy environment. Recommendation for further research work: Some issues were identified for future research: • Construction and specification of fire resistance • Problems based on human behavior. • Importance of fire safety for all the party. • Cost of fire safety before and after the incidence. 9. ACKNOWLEDGEMENT : The author is thankful to all the key informants and students of Madan Bhandari Memorial Academy
Nepal 1th Batch for helping in collecting data. The author is thankful to all the key informants and students of Madan Bhandari Memorial Academy
Nepal 1th Batch for helping in collecting data. REFERENCES : REFERENCES : 7. CONCLUSION : All three parties (occupants, management, and concerned authorities) lacked awareness and
preparedness. The residents of commercial buildings lacked fire safety knowledge and preparedness. They were uninformed about the building's general features, such as the floor plan and egress
placement. In the same way, the residents were unaware of the assembly point and emergency
evacuation method. They will also require aid in getting to a safe location. They were completely
unaware of the building's safety features and were unable to use them. There was no proper fire safety PAGE 193 A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION REFERENCES : A. K. Mishra, et al. (2022); www.srinivaspublication.com International Journal of Applied Engineering and Management
Letters (IJAEML), ISSN: 2581-7000, Vol. 6, No. 2, September 2022
SRINIVAS
PUBLICATION [1] Mishra, A. K., & Aithal, P. S., (2022). Causes and Effects of Design Change of Construction
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shpguidelines/hazard-prevention.html [Accesses 24 July 2017]. [20] OSHA
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shpguidelines/hazard-prevention.html [Accesses 24 July 2017]. [21] Yevgen Martyn et al, (2020). Software for Shelter’s Fire Safety and Comfort Levels Evaluation
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Data Stream Mining & Processing, Volume 1158. DOI: 10.1007/978-3-030-61656-4_31. PAGE 196
|
https://openalex.org/W4236951862
|
https://commons.clarku.edu/cgi/viewcontent.cgi?article=1920&context=faculty_geography
|
English
| null |
Response of carbon fluxes to water relations in a savanna ecosystem in South Africa
| null | 2,008
|
cc-by
| 12,271
|
Clark University
Clark University
Clark Digital Commons
Clark Digital Commons
Geography
Faculty Works by Department and/or School
1-1-2008
Response of carbon fluxes to water relations in a savanna
Response of carbon fluxes to water relations in a savanna
ecosystem in South Africa
ecosystem in South Africa
W. L. Kutsch
Max Planck Institute for Biogeochemistry
N. Hanan
Colorado State University
B. Scholes
The Council for Scientific and Industrial Research
I. McHugh
Monash University
W. Kubheka
Colorado State University
See next page for additional authors
Follow this and additional works at: https://commons.clarku.edu/faculty_geography
Part of the Geography Commons
Repository Citation
Repository Citation
Kutsch, W. L.; Hanan, N.; Scholes, B.; McHugh, I.; Kubheka, W.; Eckhardt, H.; and Williams, Christopher A.,
"Response of carbon fluxes to water relations in a savanna ecosystem in South Africa" (2008).
Geography. 921.
https://commons.clarku.edu/faculty_geography/921
This Article is brought to you for free and open access by the Faculty Works by Department and/or School at Clark
Digital Commons. It has been accepted for inclusion in Geography by an authorized administrator of Clark Digital
Commons For more information please contact larobinson@clarku edu cstebbins@clarku edu Clark University
Clark University
Clark Digital Commons
Clark Digital Commons
Geography
Faculty Works by Department and/or School
1-1-2008
Response of carbon fluxes to water relations in a savanna
Response of carbon fluxes to water relations in a savanna
ecosystem in South Africa
ecosystem in South Africa
W. L. Kutsch
Max Planck Institute for Biogeochemistry
N. Hanan
Colorado State University
B. Scholes
The Council for Scientific and Industrial Research
I. McHugh
Monash University
W. Kubheka
Colorado State University
See next page for additional authors
Follow this and additional works at: https://commons.clarku.edu/faculty_geography
Part of the Geography Commons
Repository Citation
Repository Citation
Kutsch, W. L.; Hanan, N.; Scholes, B.; McHugh, I.; Kubheka, W.; Eckhardt, H.; and Williams, Christopher A.,
"Response of carbon fluxes to water relations in a savanna ecosystem in South Africa" (2008). Geography. 921. https://commons.clarku.edu/faculty_geography/921
This Article is brought to you for free and open access by the Faculty Works by Department and/or School at Clark
Digital Commons. It has been accepted for inclusion in Geography by an authorized administrator of Clark Digital
Commons. For more information, please contact larobinson@clarku.edu, cstebbins@clarku.edu. Clark University
Clark University
Clark Digital Commons
Clark Digital Commons
Geography
1-1-2008
Response of carbon fluxes to wa
Response of carbon fluxes to wa
ecosystem in South Africa
ecosystem in South Africa
W. L. Kutsch
Max Planck Institute for Biogeochemistry
N. Hanan
Colorado State University
B. Scholes
The Council for Scientific and Industrial Research
I. McHugh
Monash University
W. Kubheka
Colorado State University
See next page for additional authors
Follow this and additional works at: https://common
Part of the Geography Commons
Repository Citation
Repository Citation
Kutsch, W. L.; Hanan, N.; Scholes, B.; McHugh, I.; Kub
"Response of carbon fluxes to water relations in a s
Geography. 921. https://commons.clarku.edu/faculty_geography/92
This Article is brought to you for free and open access by
Digital Commons. It has been accepted for inclusion in Ge
Commons. For more information, please contact larobins Authors
Authors
W. L. Kutsch, N. Hanan, B. Scholes, I. McHugh, W. Kubheka, H. Eckhardt, and Christopher A. Williams sch, N. Hanan, B. Scholes, I. McHugh, W. Kubheka, H. Eckhardt, and Christopher A. Williams This article is available at Clark Digital Commons: https://commons.clarku.edu/faculty_geography/92 This article is available at Clark Digital Commons: https://commons.clarku.edu/faculty_geography/921 1
Introduction Abstract. The principal mechanisms that connect carbon
fluxes with water relations in savanna ecosystems were stud-
ied by using eddy covariance method in a savanna ecosys-
tem at Kruger National Park, South Africa. Since the annual
drought and rewetting cycle is a major factor influencing the
function of savanna ecosystems, this work focused on the
close inter-connection between water relations and carbon
fluxes. Data from a nine-month measuring campaign lasting
from the early wet season to the late dry season were used. Savannas are defined as tropical and sub-tropical vegetation
types where woody plants and grasses co-dominate. They
are characterized by a seasonal change of water availability
with distinct rainy and dry seasons. Decades of research have
identified not only water, but also nutrients, herbivory, and
fire as major drivers of savanna vegetation dynamics (Walter,
1939, 1971; Scholes and Walker, 1993; Scholes and Archer,
1997; Sankaran et al., 2004; 2005; Beerling and Osborne,
2006). Total ecosystem respiration showed highest values at the
onset of the growing season, a slightly lower plateau during
the main part of the growing season and a continuous de-
crease during the transition towards the dry season. Despite large land surface coverage (∼20% of the global
land surface and 40% of the African surface), important bio-
diversity (∼8000 of 13 000 African plant species are savanna
endemic, White, 1980) and substantial land use change in
savannas, there have been relatively few projects focusing
on the patterns and processes controlling savanna carbon and
water vapour exchange (Levy, 1995; Verhoef et al., 1996;
Lloyd et al., 1997; Hanan et al., 1998; Scanlon and Albert-
son, 2004; Veenendaal et al., 2004; Williams and Albertson,
2004, 2005, for African savannas; Meir et al., 1996; Miranda
et al., 1997; Meinzer et al., 1999; da Rocha et al., 2002;
Santos et al. 2003, for South American savannas; Hutley
et al., 2000, 2001, 2005; Chen et al., 2002, 2003; Beringer et
al. 2003, for Australian savannas; Baldocchi et al., 2004 for
temperate Mediterranean savannas in California). The regulation of canopy conductance was changed in two
ways: changes due to phenology during the course of the
growing season and short-term acclimation to soil water con-
ditions. Clark University
Clark University Clark Digital Commons
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Kutsch, W. L.; Hanan, N.; Scholes, B.; McHugh, I.; Kubheka, W.; Eckhardt, H.; and Williams, Christopher A.,
"Response of carbon fluxes to water relations in a savanna ecosystem in South Africa" (2008). Geography. 921. https://commons.clarku.edu/faculty_geography/921 This Article is brought to you for free and open access by the Faculty Works by Department and/or School at Clark
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Commons. For more information, please contact larobinson@clarku.edu, cstebbins@clarku.edu. Biogeosciences, 5, 1797–1808, 2008
www.biogeosciences.net/5/1797/2008/ Biogeosciences © Author(s) 2008. This work is distributed under
the Creative Commons Attribution 3.0 License. Response of carbon fluxes to water relations in a savanna
ecosystem in South Africa W. L. Kutsch1, N. Hanan2, B. Scholes3, I. McHugh4, W. Kubheka2, H. Eckhardt5, and C. Williams6
1Max-Planck-Institute for Biogeochemistry, P.O. Box 10 01 64, 07701 Jena, Germany
2Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, USA
3Council for Scientific and Industrial Research (CSIR), Natural Resources and the Environment, P.O. Box 395,
0001 Pretoria, South Africa 4School of Geography and Environmental Science, Monash University, Clayton, Australia
5South African National Parks (SANParks), Scientific Services, Skukuza, South Africa
6Graduate School of Geography, Clark University Worcester, MA, USA 4School of Geography and Environmental Science, Monash University, Clayton, Australia
5South African National Parks (SANParks), Scientific Services, Skukuza, South Africa
6Graduate School of Geography, Clark University Worcester, MA, USA Received: 17 March 2008 – Published in Biogeosciences Discuss.: 27 May 2008
Revised: 23 September 2008 – Accepted: 6 October 2008 – Published: 23 December 2008 W. L. Kutsch et al.: Savanna water-carbon relations W. L. Kutsch et al.: Savanna water-carbon relations savannas in global carbon cycles, as well as informing poli-
cies related to the role of African countries in Kyoto Protocol. African savannas (Scholes and Archer, 1997). The site was
part of the SAFARI 2000 experiment to understand the in-
teractions between the atmosphere and the land surface in
southern Africa (Otter et al., 2002; Scholes et al., 2001;
Shugart et al., 2004). The location of the tower is about
13 km WS W of Skukuza in the N’waswitshaka catchment,
an ephemeral tributary of the Sabie River. The altitude of
the site is 365 m a.s.l. The underlying parent material is Ar-
chaean granite and gneiss. The landscape shows a typical un-
dulating form with drainage lines about 3 km apart and ridge
tops about 40 m above valley floors. This leads to a cate-
nal pattern of soils and vegetation, with nutrient-poor Com-
bretum savannas on the crests, and nutrient-enriched Acacia
savannas on the lower parts of the slope.The flux tower is lo-
cated at the boundary of the two plant communities. When
the wind blows from the SE (105◦–196◦), a fine-leaf Acacia
savanna is sampled, and when it blows from the N (270◦–
76◦), a broad-leaf Combretum savanna is sampled. Sectoral
analyses of eddy covariance measurements have been con-
ducted successfully in previous studies (e.g. Kutsch et al.,
2005). One of the few infrastructures enabling the study carbon
and water relations of African savannas is the flux tower in
the Kruger National Park, installed between Skukuza and
Pretoriuskop in 2000. In this study we investigate principal
mechanisms that connect carbon fluxes with water relations
in savanna ecosystems and differences between the two sa-
vanna types. Since the annual drought and rewetting cycle
is a major factor influencing the function of savanna ecosys-
tems, this work is focused on two questions concerning water
relations: (1) How do water availability and seasonality drive
ecosystem respiration?; and (2) how does the close inter-
connection between canopy conductance and photosynthesis,
influence the carbon uptake by the vegetation during times of
different water availability? Ecosystems respiration as measured by the eddy covari-
ance technique is the sum of soil, plant, and animal respira-
tion. In biomes with a pronounced rainfall seasonality and/or
water limitation, the temperature response may be secondary
to the response to water availability. W. L. Kutsch et al.: Savanna water-carbon relations 1 the monthly precip-
itation of the growing season 2002/2003 is compared to the
long-term mean. The 2002/2003 growing season was very
dry. Only half of the long-term amount of rain fell (276 mm). By combining the regulation of respiration on one hand
and stomatal conductance and photosynthesis on the other
we provide a principle analysis of the temporal dynamics in
the regulation of savanna carbon fluxes by water relations. We used data from a nine months period between January
and September 2003 for this study. For annual balances
of this site and improved gap-filling procedures we refer to
Archibald et al. (2008). W. L. Kutsch et al.: Savanna water-carbon relations Detailed soil and vegetation analyses of the two types are
given in Scholes et al. (2001). They show that the soil in the
Acacia savanna contains more clay, more nutrients such a ni-
trogen and phosphorous and also a higher cation exchange
capacity than in the Combretum savanna. Volumetric field
capacity (WCFC) is about 0.12 in the Combretum- and 0.2 in
the Acacia-savanna. Volumetric water content at −1500 kPa
(“wilting point”, WCWP) was 0.06 and 0.1, respectively. Rel-
ative plant available water (RPAW) was calculated from the
actual volumetric water content for the layers 0–10 cm and
0–30 cm by Ecosystem photosynthesis is likewise controlled by a com-
plex interaction of processes that connect carbon fluxes and
water relations. The control of CO2 and water vapour ex-
change between ecosystems and the atmosphere by the stom-
atal aperture of plant leaves is the hinge between the two cy-
cles – particularly notable in arid and semi-arid biomes. However, the same stomata do not always react to the
short-term (diurnal) variations of VPD in the same way. Leaf
stomatal sensitivity and maximum conductance can be mod-
ified within periods of several days to few weeks by phenol-
ogy and by acclimation to factors such as soil water status
(Halldin et al., 1984). Kutsch et al. (2001a) assessed the
influence of this mid-term stomatal acclimation on ecosys-
tem fluxes by calculating canopy water and carbon balances
for several years under different climates. In this context it
is important to consider that canopy photosynthetic capacity
(FP,sat) is strongly correlated to maximum surface conduc-
tance (Gs,sat; Schulze et al. (1994)). RPAW = (WCakt −WCWP)
(WCFC −WCWP) · 100
(1) (1) Rainfall at the site averages 547.1 mm but with significant
interannual variability. The first precipitation events of the
rainy season generally occur in late September or October,
with last rains in April or early May. However, much of the
interannual variability in total precipitation arises with vari-
ation in the dates of first and last rains. Figure 1 shows the
long-term monthly means of minimum and maximum tem-
perature and the mean precipitation at Skukuza (data from
1960–1999, after Scholes et al. (2001)). Precipitation is
shown as cumulative curve. In order to show a complete
growing season the data are shown from August to July. Most of the annual rainfall occurs during the summer (Octo-
ber to April). In the lower part of Fig. 1
Introduction The most constant parameter was water use efficiency that
was influenced by VPD during the day but the VPD response
curve of water usage did change only slightly during the
course of the growing season and decreased by about 30%
during the transition from wet to dry season. The regulation of canopy conductance and photosynthetic
capacity were closely related. This observation meets re-
cent leaf-level findings that stomatal closure triggers down-
regulation of Rubisco during drought. Our results may show
the effects of these processes on the ecosystem scale. Increasing our understanding of the physical and physio-
logical controls on savanna functioning using eddy covari-
ance measurements is of great importance both for funda-
mental ecological understanding as well as for improved
management and conservation of these important ecosys-
tems. Such measurements will, furthermore provide ba-
sic data for refining models of ecosystem carbon dynamics
and facilitate improved understanding of the role of tropical Correspondence to: W. L. Kutsch
(wkutsch@bgc-jena.mpg.de)
management an
tems. Such m
sic data for refi
and facilitate im
Published by Copernicus Publications on behalf of the European Geosciences Un Correspondence to: W. L. Kutsch
(wkutsch@bgc-jena.mpg.de) Published by Copernicus Publications on behalf of the European Geosciences Union. Published by Copernicus Publications on behalf of the European Geosciences Union. Published by Copernicus Publications on behalf of the European Geosciences Union. 1798 2.3
Supplemental measurements Additional weather measurements on the main tower in-
cluded measurements of incident and reflected shortwave
radiation (300–1100 nm, W m−2, CM14, Kipp and Zonen,
Delft, The Netherlands), incident and reflected near-infrared
(CM14 600–1100 nm, W m−2, Kipp and Zonen, Delft, The
Netherlands) and incoming and emitted longwave radiation
(CG2, >3.0 nm, W m−2, Kipp and Zonen, Delft, The Nether-
lands). Radiation measurements were made at 20 s inter-
vals and then recorded in the data-logger as 30 min aver-
ages. Precipitation events were recorded with a tipping-
bucket raingauge (Texas Instruments TE525, mm) totalled
for each 30 min period. In addition to the meteorological and flux measurements
on the main tower, we deployed supplemental microclimate
stations in the adjacent Acacia and Combretum savannas to
measure air temperature, air humidity, soil temperature and
volumetric soil moisture. In each savanna we deployed TDR
probes (Campbell Scientific CS615, cm3 water cm−3 soil)
and temperature probes (Campbell Scientific 108,◦C) in pro-
files with 4 or 5 measurement depths from near the soil sur-
face to just above bedrock. Fig. 1. Long-term monthly means of minimum and maximum tem-
perature (dotted lines) and the mean precipitation (solid line) at
Skukuza (data from 1960–1999, after Scholes et al. 2001). Pre-
cipitation is shown as cumulative curve. In order to show a com-
plete growing season the data are printed from August to July. In the lower part the monthly precipitation of the growing season
2002/2003 (hatched bars) is compared to the long-term mean (black
bars). Fig. 1. Long-term monthly means of minimum and maximum tem-
perature (dotted lines) and the mean precipitation (solid line) at
Skukuza (data from 1960–1999, after Scholes et al. 2001). Pre-
cipitation is shown as cumulative curve. In order to show a com-
plete growing season the data are printed from August to July. In the lower part the monthly precipitation of the growing season
2002/2003 (hatched bars) is compared to the long-term mean (black
bars). 2.4
Data treatment Post-processing of the raw high frequency (10 Hz) data to
obtain fluxes representative of thirty minute periods involved
the standard spike filtering, planar rotation of velocities, lag
correction for CO2 and water vapour concentrations, and cal-
culation of mass fluxes using conventional equations (Mon-
crieff et al., 1997; Aubinet et al., 2000). area average of approximately 1.0 (i.e. within canopy LAI
∼3.0) three or four weeks after the beginning of the rainy
season and then remaining at that level until several weeks
after the last rainfall. Thereafter, the trees in the Combretum
savanna dropped their leaves, whereas the Acacia species re-
mained leafy during the dry season. Herbaceous layer pro-
ductivity and LAI is more dynamic with inter-annual vari-
ability in total rainfall, with average LAI of the herbaceous
layer similar to the tree layer (i.e. 1.0), but in this case spread
out more or less evenly across the landscape rather than being
confined within the tree canopies. Carbon dioxide flux (FCO2) was also corrected to include
the canopy storage flux using the simple approach of assum-
ing that the CO2 is uniformly distributed within the air col-
umn between the soil surface and the inlet of the tube: F ′
CO2 = FCO2 + 1CO2 · h · 1000
η · t
(2) (2) (2) where F ′
CO2 is the corrected flux (µmol CO2 m−2 s−1), FCO2
is the flux measured by the eddy covariance system (µmol
CO2 m−2 s−1), 1CO2 is the difference in CO2 concentration
between t0 and t−1 (ppm), h is the height of the air column
(16 m), η is molar volume of an ideal gas (22.4 l mol−1), the
factor 1000 converts l into m3, and t is the time step of 1800 s. 2.1
Research site 2.1
Research site The vegetation of the study site is open woodland with ap-
proximately 30% tree canopy cover in both Acacia and Com-
bretum savanna types (Scholes et al., 2001). Leaf area index
of the trees in both savanna types is similar, increasing to an The Skukuza flux tower was established in early 2000 to
study the carbon, water and energy dynamics of semi-arid Biogeosciences, 5, 1797–1808, 2008 www.biogeosciences.net/5/1797/2008/ W. L. Kutsch et al.: Savanna water-carbon relations 1799 0
5
10
15
20
25
30
35
40
Temperature (°C)
0
150
300
450
600
Cumulative precipitation
(mm)
0
10
20
30
40
50
60
70
80
90
Monthly precipitation (mm)
A
S O N
D
J
F
M A
M J
J
Long- term mean
2002/2003
Fig. 1. Long-term monthly means of minimum and maximum tem-
perature (dotted lines) and the mean precipitation (solid line) at
Skukuza (data from 1960–1999, after Scholes et al. 2001). Pre-
cipitation is shown as cumulative curve. In order to show a com-
plete growing season the data are printed from August to July. In the lower part the monthly precipitation of the growing season
2002/2003 (hatched bars) is compared to the long-term mean (black
bars). 0
5
10
15
20
25
30
35
40
Temperature (°C)
0
150
300
450
600
Cumulative precipitation
(mm)
0
10
20
30
40
50
60
70
80
90
Monthly precipitation (mm)
A
S O N
D
J
F
M A
M J
J
Long- term mean
2002/2003 measure carbon dioxide (µmol mol−1) and water vapour
(mmol mol−1) concentrations. The gas samples were drawn
at 6 l min−1 through a 6 m teflon-coated tube to the gas ana-
lyzer enclosed in a weather-proof box. The sonic anemome-
ter and gas analyser measurements were recorded at 10 Hz. Cumulative precipitation
(
) 2.2
Eddy covariance system Ecosystem-level fluxes of water, heat and carbon diox-
ide were made using a closed-path eddy covariance sys-
tem mounted at 16 m. This consisted of a 3-dimensional
sonic anemometer (Gill Wind Master Pro, Gill Instruments,
Lymington, UK) to measure fluctuations in horizontal and
vertical wind speeds (m s−1) and temperature (K), and a gas
analyzer (LiCor 6262, LiCor, Lincoln, Nebraska, USA) to After calculation of the fluxes the eddy-covariance data
were combined with and fitted against microclimate data. Nighttime carbon dioxide flux (Fn), a surrogate for ecosys-
tem respiration (FR), was regressed against soil tempera-
ture at 7 cm depth (TS) and soil moisture (RPAW, Eq. (1). Biogeosciences, 5, 1797–1808, 2008 www.biogeosciences.net/5/1797/2008/ 1800 W. L. Kutsch et al.: Savanna water-carbon relations Only data with a coincident friction velocity (u∗) greater than
0.25 m s−1 were used for this analysis. leaf area index, and the friction velocity (Jensen and Hum-
melshøj, 1995). Finally, surface conductance was calculated
as the reciprocal of the residual resistance (Eq. 9): The data set was fitted to the ecosystem respiration model
described by Reichstein et al. (2002). This model pro-
vides response curves for soil moisture and temperature, and
changes the Q10-value of the temperature response curve
with changing soil moisture as observed in this study: GS =
VPD
E
−
u
u∗2 −rb
(9) (9) If VPD is given in g m−3, E in g m−2, u and u∗in
m s−1, and rb in s m−1, GS is obtained in m s−1. To con-
vert GS from m s−1 into the common plant physiological unit
of mmol m−2 s−1, a multiplication factor has to be applied
which is roughly 40 000. FR = FR,ref · f (Tsoil, RSWC) · g(RSWC)
(3)
f (Tsoil, RSWC) = eE0(RSWC)·(
1
Tref−T0 −
1
Tsoil−T0 )
(4) FR = FR,ref · f (Tsoil, RSWC) · g(RSWC)
(3) (3) FR = FR,ref · f (Tsoil, RSWC) · g(RSWC) f (Tsoil, RSWC) = eE0(RSWC)·(
1
Tref−T0 −
1
Tsoil−T0 )
(4) f (Tsoil, RSWC) = eE0(RSWC)·(
1
Tref−T0 −
1
Tsoil−T0 )
(4) (4) In order to obtain an indication of the sensitivity of the
regulation of canopy conductance to ambient humidity, GS
was plotted against the actual water vapour pressure deficit
of the air (VPD) for distinct periods of between two and four
weeks with different levels of soil moisture. 2.2
Eddy covariance system Each data set
was fitted separately to the following equation: g(RSWC) =
RSWC −RSWC0
(RSWC1/2 −RSWC0
) + (RSWC −RSWC0)
(5) (5) where E0 was not fixed but a linear function of relative water
content (RSWC) allowing for a RSWC effect on temperature
sensitivity. RWSC used as by Reichstein et al. (2002) was
derived from the RPAW for the 0–10 cm layer by: GS = GS,max
c · VPD2
(10) GS = GS,max
c · VPD2 (10) RWSC = RPAW
100
(6) where GS,max is a theoretical maximum conductance that
occurs when VPD (mbar) is zero (set constantly to
2000 mmol m−2 s−1 in that study), and c is an empirical co-
efficient that provides a measure of the sensitivity of the
regulation of GS. Envelope functions covering 90% of the
data were fitted to the data by quantile regression (Cade and
Noon, 2003). The seasonal course of the coefficient c in
Eq. (10) reveals how the canopy acclimates to changing con-
ditions. This function is simpler and more empirical than the
“Ball/Berry-Function” (Ball et al., 1987) that is commonly
used but c is defining the shape of the response curve in the
same way and reveals the same seasonal pattern as the co-
efficient D0 in the Ball/Berry approach does. We chose the
more empirical function to avoid deriving physiological pa-
rameters top-down which is inappropriate for a physiologi-
cally heterogeneous system like a savanna. (6) The modelled daytime ecosystem respiration (FR) was
then used to calculate canopy assimilation (FP ) from the
daytime fluxes: FP = F ′
CO2 −FR
(7) (7) Canopy assimilation was then fitted against global radia-
tion measured above the canopy by means of a tangens hy-
perbolicus (tan h) function (von Stamm, 1994): Canopy assimilation was then fitted against global radia-
tion measured above the canopy by means of a tangens hy-
perbolicus (tan h) function (von Stamm, 1994): FP = FP,sat · tan h( k · I
FP,sat
)
(8) (8) where k is an empirical coefficient describing the initial slope
of the light response curve and I is the incoming short-wave
radiation (W m−2). 3
Results Surface conductance, GS, defined as the surface conduc-
tance to water vapour transfer at a canopy scale, was cal-
culated from the eddy flux data as described by Jensen and
Hummelshøj (1995) and Herbst et al. (2002). First, the total
resistance, rs, of the system was calculated as the quotient
of vapour pressure deficit, VPD, of the air above the forest
and water vapour flux, E. Analogously, the resistance owing
to turbulent transport in the atmosphere, ra, (assumed to be
equal for momentum and water vapour) was determined as
the quotient of mean horizontal windspeed, u, and the square
of the friction velocity, u∗2. The resistance through the vis-
cous sublayers on the individual leaves, rb, was estimated as
a function of the kinematic viscosity of the air, the molec-
ular diffusivity of water vapour, the size of the leaves, the Biogeosciences, 5, 1797–1808, 2008 3.1
Climate and flux overview Since 2002/2003 was a dry year, periods with low rainfall
during the wet season resulted in periods of low soil moisture
even during the typical vegetation growth period (Fig. 2a, Ta-
ble 1). The dry periods can also be seen in decreases during
the annual course in the fAPAR (Fig. 2a). The net ecosystem
exchange (Fig. 2b) and its components (canopy photosynthe-
sis, FP , Fig. 2c, and ecosystem respiration FR, Fig. 2d) were
strongly influenced by the soil moisture dynamics. There-
fore, the observational data during the growing season were
divided in the following sub-sets for further eco(system)-
physiological analyses. Biogeosciences, 5, 1797–1808, 2008 www.biogeosciences.net/5/1797/2008/ W. L. Kutsch et al.: Savanna water-carbon relations 1801 J
F
M
A
M
J
J
A
S
1
3
2
4
5
6
7
8
Periods:
2
4
6
8
-5
0
5
10
15
20
-20
-15
-10
-5
0
5
FC (µmol m-2 s-1)
FP (µmol m-2 s-1)
FR (µmol m-2 s-1)
0
0.1
0.2
Soil moisture
0-10 cm (Vol %)
0.3
8
7
6
5
4
3
2
1
(a)
(b)
(c)
(d)
0
0.1
0.2
0.3
0.4
fAPAR
Fig. 2. Seasonal courses of volumetric soil water content for the
layers 0–10 cm (a) in the Acacia savanna (grey line) and the Com-
bretum savanna (black line) and of fAPAR of two Modis grid cells
around the tower (a), of net ecosystem exchange (NEE, (b), canopy
photosynthesis (FP , (c) and total ecosystem respiration (FR, (d). The periods defined in Table 1 are marked in the Fig. J
F
M
A
M
J
J
A
S
1
3
2
4
5
6
7
8
Periods:
2
4
6
8
-5
0
5
10
15
20
-20
-15
-10
-5
0
5
FC (µmol m-2 s-1)
FP (µmol m-2 s-1)
FR (µmol m-2 s-1)
0
0.1
0.2
Soil moisture
0-10 cm (Vol %)
0.3
8
7
6
5
4
3
2
1
(a)
(b)
(c)
(d)
0
0.1
0.2
0.3
0.4
fAPAR Table 1. Sub-sets for data evaluation in 2003. Dataset No. 3.1
Climate and flux overview Period
Characterization
Mean RPAW 0–10(1)
1
25 Jan–8 Feb
Wet soil,
early wet season
30.6
2
10–20 Feb
Dry soil,
early wet season
16.1
3
23 Feb–29 Mar
Wet soil,
mid wet season
28.2
4
1–25 Apr
Dry soil,
mid wet season
11.6
5
27 Apr–11 May
Wet soil,
transition period
39.1
6
14 May–5 Jun
Dry soil,
transition period
12.7
7
14 Jun–12 Jul
Wet soil,
dry season
28.2
8
28 Jul–3 Sep
Dry soil,
dry season
9.7
(1) Averages were build on the basis of only those datasets that con-
tained full information (climate and EC) after quality control and
filtering. They might slightly differ from the “real” averages. Kutsch et al. Fig 3 (1) Averages were build on the basis of only those datasets that con-
tained full information (climate and EC) after quality control and
filtering. They might slightly differ from the “real” averages. Kutsch et al. Fig 3 (1) Averages were build on the basis of only those datasets that con-
tained full information (climate and EC) after quality control and
filtering. They might slightly differ from the “real” averages. Kutsch et al. Fig 3 -20
-16
-12
-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
FP (µmol CO2 m-2 s-1)
Global radiation (W m-2)
Acacia-savanna
Combretum-savanna
FC (µmol CO2 m-2 s-1)
0
250
500
750
1000
1250
Fig. 3. Light response curves of net ecosystem exchange (FC, up-
per graph) and canopy photosynthesis (FP , lower graph) measured
during the peak of the growing season (Period 3 in March 2003). The open symbols show data from the Acacia-site, the filled sym-
bols from the Combretum-site. -20
-16
-12
-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
FP (µmol CO2 m-2 s-1)
Global radiation (W m-2)
Acacia-savanna
Combretum-savanna
FC (µmol CO2 m-2 s-1)
0
250
500
750
1000
1250 Fig. 2. Seasonal courses of volumetric soil water content for the
layers 0–10 cm (a) in the Acacia savanna (grey line) and the Com-
bretum savanna (black line) and of fAPAR of two Modis grid cells
around the tower (a), of net ecosystem exchange (NEE, (b), canopy
photosynthesis (FP , (c) and total ecosystem respiration (FR, (d). The periods defined in Table 1 are marked in the Fig. 3.4
Seasonal characteristics of canopy conductance and
photosynthesis 3.4
Seasonal characteristics of canopy conductance and
photosynthesis Fig. 4. Temperature response curves of nighttime respiration for dif-
ferent intervals of soil moisture. Filled cycles symbolize data from
the main growing season (January–March 2003), grey cycles from
the transition time (April–May 2003) and open symbols from the
dry period (June–September 2003). The values of the exponential
fits and the regression coefficient are related to the whole datasets
comprising data from all seasons. VPD response curves of GS were plotted for each of the de-
fined periods as envelope curves based on a 90% quantile
regression (Fig. 6). The response curves are paired. Each
pair shows data from a period with high soil water content
(left) and data from a dryer period thereafter (right). In or-
der to demonstrate the degree of stomatal acclimation the re-
sponse curve from the left dataset is also shown in the right
one as grey line. The graphs showed that stomatal sensitiv-
ity changed drastically within a few days when soil moisture
varied. Stomatal conductance was highest in January (6a)
and March (6c). The short period of drought during Febru-
ary (6b) slightly increased sensitivity to VPD, whereas dur-
ing April (6d) the drought led to much-reduced canopy con-
ductance over the whole range of VPD. After a large rain-
fall event in May (6e), plants reverted the regulation pat-
tern again but did not reveal as high conductance as in Jan-
uary or March, indicating the onset of the senescence of the
leaves that may be speeded up by the April drought period. Nevertheless, the following dry period during May (6f) re-
duced stomatal conductance again while the senescence of
the plants went on. Consequently, a rainfall event during late
June did not reverse the performance any more (6g) and the
during the dry season (August), canopy conductance was re-
duced to a minimum (6h). W. L. Kutsch et al.: Savanna water-carbon relations
Kutsch et al. Fig 5 W. L. Kutsch et al.: Savanna water-carbon relations
Kutsch et al. Fig 5 y = 0.3624e0.0903x
r2 = 0.433
0
2
4
6
8
10
y = 0.3437e0.0697x
r2 = 0.4945
0
2
4
6
8
y = 0.1763e0.0701x
r2 = 0.2301
0
2
4
6
8
Nighttime fluxes (µmol CO2 m-2 s-1)
Soil temperature at – 7 cm (°C)
Wet soil
Dry soil
Medium soil
10
20
30
40
Fig. 4. Temperature response curves of nighttime respiration for dif-
ferent intervals of soil moisture. Filled cycles symbolize data from
the main growing season (January–March 2003), grey cycles from
the transition time (April–May 2003) and open symbols from the
dry period (June–September 2003). The values of the exponential
fits and the regression coefficient are related to the whole datasets
comprising data from all seasons. y = 0.8734x + 0.3065
r2 = 0.6514
0
2
4
6
8
10
Modelled nighttime fluxes
Measured nighttime fluxes
(µmol CO2 m-2 s-1)
(µmol CO2 m-2 s-1)
0
2
4
6
8
10
Fig. 5. Comparison between measured and modelled data of night-
time respiration. Based on complete data set (January–September
2003). The solid line represents the regression (regression values
written at the top), the 1:1 line is dashed. y = 0.3624e0.0903x
r2 = 0.433
0
2
4
6
8
10
y = 0.3437e0.0697x
r2 = 0.4945
0
2
4
6
8
y = 0.1763e0.0701x
r2 = 0.2301
0
2
4
6
8
Nighttime fluxes (µmol CO2 m-2 s-1)
Soil temperature at – 7 cm (°C)
Wet soil
Dry soil
Medium soil
10
20
30
40 y = 0.8734x + 0.3065
r2 = 0.6514
0
2
4
6
8
10
Modelled nighttime fluxes
Measured nighttime fluxes
(µmol CO2 m-2 s-1)
(µmol CO2 m-2 s-1)
0
2
4
6
8
10 Fig. 5. Comparison between measured and modelled data of night-
time respiration. Based on complete data set (January–September
2003). The solid line represents the regression (regression values
written at the top), the 1:1 line is dashed. Figure 5 shows the modelled versus measured rates. Re-
sults indicate that this model approach explains much of the
variation in flux rates and confirm the hypothesis about an
indirect effect of water availability on ecosystem respiration
via changes of temperature sensitivity (Q10) for which Re-
ichstein et al. (2002) developed their model expression. 40 Soil temperature at – 7 cm (°C) Soil temperature at – 7 cm (°C) 3.2
Sectoral analysis and differences between savanna sites 3.2
Sectoral analysis and differences between savanna sites For a sectoral analysis data were sorted according to wind
direction in order to examine differences between the two sa-
vanna types. Data from the sector between 105◦–196◦were
taken for the fine-leaf Acacia savanna, from the sector be-
tween 270◦–76◦for the broad-leaf Combretum savanna. Data
from other wind directions were discarded. There were no
significant differences between the savanna types in terms of
fluxes. As an example, Fig, 3a and b show light response
curves of net ecosystem exchange (FC) and of canopy pho-
tosynthesis (FP ) for the two sectors during the peak of the
growing season (Period 3 as defined in Table 1, other pe-
riods showed the same picture). Consequently, data were
pooled again for further analysis. The following paragraphs
will show a principle analysis of carbon flux responses to
water relations in a savanna ecosystem without considering
possible small differences between the sub-types. Fig. 3. Light response curves of net ecosystem exchange (FC, up-
per graph) and canopy photosynthesis (FP , lower graph) measured
during the peak of the growing season (Period 3 in March 2003). The open symbols show data from the Acacia-site, the filled sym-
bols from the Combretum-site. Biogeosciences, 5, 1797–1808, 2008 www.biogeosciences.net/5/1797/2008/ 1802 W. L. Kutsch et al.: Savanna water-carbon relations 7. Conductance response curve of CO2-fluxes for the 8 periods
as defined in Table 1. Black dots symbolize the canopy gross photo-
synthetic flux, open circles the net ecosystem flux; data measured at
saturating light conditions with incoming shortwave radiation being
higher than 700 W m−2. Ecosystem net exchange (FC) and Canopy photosynthesis ( FP)
(µmol CO2 m-2 s-1) Canopy conductance (mmol m-2 s-1) Canopy conductance (mmol m-2 s-1) Canopy conductance (mmol m-2 s-1) Fig. 6. VPD-response curves of stomatal conductance for the 8
periods as defined in Table 1. Data measured at saturating light
conditions with incoming shortwave radiation being higher than
700 W m−2. The grey curves in the situations with dry soil (right
graphs) symbolize the curves from corresponding situations with
wet soils (left graphs). Fig. 7. Conductance response curve of CO2-fluxes for the 8 periods
as defined in Table 1. Black dots symbolize the canopy gross photo-
synthetic flux, open circles the net ecosystem flux; data measured at
saturating light conditions with incoming shortwave radiation being
higher than 700 W m−2. Each of these parameters showed a seasonal trend (dashed
grey line) that was overlaid by mid-term responses to the re-
spective drought periods, but the specific characteristics were
different. Each of these parameters showed a seasonal trend (dashed
grey line) that was overlaid by mid-term responses to the re-
spective drought periods, but the specific characteristics were
different. Light
saturated
canopy
photosynthesis
was
limited
strongly by canopy conductance below a threshold of
250 mmol m−2 s−1 during all of the periods under consider-
ation (Fig. 7a–h). During the periods with low soil mois-
ture (right figures) canopy conductance did almost never ex-
ceeded this threshold and CO2 uptake by the canopy was al-
most constantly reduced. In addition, the few data sets with a
canopy conductance around 250 mmol m−2 s−1 suggest a de-
crease of the photosynthetic capacity, which means that pho-
tosynthetic capacity was strongly coupled to stomatal sensi-
tivity. The seasonal trend of the coefficient c revealed a broad
minimum from January to March and constantly increased
during the transition period towards the dry season. C also
revealed mid-term responses to the drought periods: during
water stress, c was increased and showed the more conserva-
tive regulation of the stomata. W. L. Kutsch et al.: Savanna water-carbon relations Data measured at saturating light
conditions with incoming shortwave radiation being higher than PD (bar)
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
(b) Dry soil,
early wet season
(d) Dry soil,
mid wet season
(f) Dry soil,
transition period
(h) Dry soil,
dry season
0
10
20
30
40
50
0
10
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0
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50 -24
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-8
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-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
Ecosystem net exchange (FC) and Canopy photosynthesis ( FP)
(µmol CO2 m-2 s-1)
(a) Wet soil,
early wet season
(b) Dry soil,
early wet season
(c) Wet soil,
mid wet season
(d) Dry soil,
mid wet season
(e) Wet soil,
transition period
(f) Dry soil,
transition period
(g) Wet soil,
dry season
(h) Dry soil,
dry season
0
250
500
750
1000
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750
1000
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Canopy conductance (mmol m-2 s-1) -24
-20
-16
-12
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0
4
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-8
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0
4
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-8
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0
4
-24
-20
-16
-12
-8
-4
0
4
Ecosystem net exchange (FC) and Canopy photosynthesis ( FP)
(µmol CO2 m-2 s-1)
(a) Wet soil,
early wet season
(c) Wet soil,
mid wet season
(e) Wet soil,
transition period
(g) Wet soil,
dry season
0
250
500
750
1000
0
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500
750
1000
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500
750
1000
0
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Canopy conductan -24
-20
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-8
-4
0
4
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0
4
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-8
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0
4
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-8
-4
0
4
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-8
-4
0
4
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-20
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-8
-4
0
4
-24
-20
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-12
-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
Ecosystem net exchange (FC) and Canopy photosynthesis ( FP)
(µmol CO2 m-2 s-1)
(a) Wet soil,
early wet season
(b) Dry soil,
early wet season
(c) Wet soil,
mid wet season
(d) Dry soil,
mid wet season
(e) Wet soil,
transition period
(f) Dry soil,
transition period
(g) Wet soil,
dry season
(h) Dry soil,
dry season
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
Canopy conductance (mmol m-2 s-1)
Fig. W. L. Kutsch et al.: Savanna water-carbon relations 1803
t al. Fig 7 VPD (bar)
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
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1200
0
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0
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400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
Canopy conductance (mmol m-2 s-1)
(a) Wet soil,
early wet season
(b) Dry soil,
early wet season
(c) Wet soil,
mid wet season
(d) Dry soil,
mid wet season
(e) Wet soil,
transition period
(f) Dry soil,
transition period
(g) Wet soil,
dry season
(h) Dry soil,
dry season
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
Fig. 6. VPD-response curves of stomatal conductance for the 8
periods as defined in Table 1. Data measured at saturating light
conditions with incoming shortwave radiation being higher than
700 W m−2. The grey curves in the situations with dry soil (right
graphs) symbolize the curves from corresponding situations with
wet soils (left graphs). W. L. Kutsch et al.: Savanna water-carbon relations VP
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
Canopy conductance (mmol m-2 s-1)
(a) Wet soil,
early wet season
(c) Wet soil,
mid wet season
(e) Wet soil,
transition period
(g) Wet soil,
dry season
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50 VPD (bar)
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
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0
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0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
Canopy conductance (mmol m-2 s-1)
(a) Wet soil,
early wet season
(b) Dry soil,
early wet season
(c) Wet soil,
mid wet season
(d) Dry soil,
mid wet season
(e) Wet soil,
transition period
(f) Dry soil,
transition period
(g) Wet soil,
dry season
(h) Dry soil,
dry season
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
Fig. 6. VPD-response curves of stomatal conductance for the 8
periods as defined in Table 1. Data measured at saturating light
conditions with incoming shortwave radiation being higher than
700 W m−2. The grey curves in the situations with dry soil (right
graphs) symbolize the curves from corresponding situations with
wet soils (left graphs). W. L. Kutsch et al.: Savanna water-carbon relations -24
-20
-16
-12
-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
-24
-20
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-8
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0
4
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-8
-4
0
4
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-20
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-8
-4
0
4
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-20
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-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
-24
-20
-16
-12
-8
-4
0
4
Ecosystem net exchange (FC) and Canopy photosynthesis ( FP)
(µmol CO2 m-2 s-1)
(a) Wet soil,
early wet season
(b) Dry soil,
early wet season
(c) Wet soil,
mid wet season
(d) Dry soil,
mid wet season
(e) Wet soil,
transition period
(f) Dry soil,
transition period
(g) Wet soil,
dry season
(h) Dry soil,
dry season
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
0
250
500
750
1000
Canopy conductance (mmol m-2 s-1)
Fig. 7. Conductance response curve of CO2-fluxes for the 8 periods
as defined in Table 1. Black dots symbolize the canopy gross photo-
synthetic flux, open circles the net ecosystem flux; data measured at
saturating light conditions with incoming shortwave radiation being
higher than 700 W m−2. VPD (bar)
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
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400
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1200
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1200
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200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
0
200
400
600
800
1000
1200
Canopy conductance (mmol m-2 s-1)
(a) Wet soil,
early wet season
(b) Dry soil,
early wet season
(c) Wet soil,
mid wet season
(d) Dry soil,
mid wet season
(e) Wet soil,
transition period
(f) Dry soil,
transition period
(g) Wet soil,
dry season
(h) Dry soil,
dry season
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
0
10
20
30
40
50
Fig. 6. VPD-response curves of stomatal conductance for the 8
periods as defined in Table 1. 3.3
Night time respiration In a first approach, the night time data sets identified for
the analysis of the nocturnal fluxes after u∗filtering (1247
records) were divided into three classes of relative soil water
content. Within each of these classes an exponential temper-
ature response curves was generated. The analysis revealed
a clear exponential increase of ecosystem respiration with
increasing temperature and also a positive influence of soil
moisture (Fig. 4). In addition, the data showed that the tem-
perature response was modified by soil moisture: at low soil
moisture the Q10-value of the temperature function was re-
duced in comparison to high soil moisture. Thus, the data
set was fitted to the ecosystem respiration described by Re-
ichstein et al. (2002) which provides response curves for soil
moisture and temperature, and changes the Q10-value of the
temperature response curve with changing soil moisture as
observed in this study. www.biogeosciences.net/5/1797/2008/ Biogeosciences, 5, 1797–1808, 2008 W. L. Kutsch et al.: Savanna water-carbon relations www.biogeosciences.net/5/1797/2008/ Biogeosciences, 5, 1797–1808, 2008 W. L. Kutsch et al.: Savanna water-carbon relations Relations between photosynthetic capacity (FP,sat) and
canopy conductance at FP,sat (a) and between FP,sat and the co-
efficient c describing stomatal response sensitivity to VPD (b) for
the 8 periods under consideration. This might be a sampling issue, because the area is very
patchy (Archibald and Scholes, 2007). However, the two sa-
vanna types extend over at least 1 km in the respective foot-
print area. We suspect that the reasons for the functional
convergence of the savanna sub-types could only be found
by detailed studies on leaf-level eco-physiology. Kutsch (un-
published data) could show a spatial correlation between
soil properties and eco-physiological strategies of the grass
species. It was found that different species optimize their
fluxes by balancing stomatal conductance and photosynthetic
capacity between water availability (lower in the Acacia-
site soils) and nutrient availability (higher in the Acacia-site
soils). The resulting convergence allows to treat the data
as one dataset in the following principle analysis of water-
carbon interactions of fluxes in an African savanna ecosys-
tem. Fig. 8. Annual courses of the coefficient c describing stomatal
response sensitivity to VPD, of maximum canopy photosynthesis
(FP,sat) average nighttime respiration (Rav) and R15 as derived
from the model. Each point represents the center of a period de-
fined in Table 1. The dashed line shows a hypothetical seasonal
course without intermediate drought periods. during the drought periods, in particular in April. The ini-
tially high respiration rate is typical for seasonal ecosystems
facing drought or frost periods (Arneth et al., 2006; Schimel
and Mikan, 2005). Respiratory patterns showed a strong dependency on tem-
perature and soil moisture. In addition higher rates during
the early stages of the wet season and a decrease thereafter
even in periods of high soil moisture was detected. Sea-
sonal changes in ecosystem respiration rates due to changes
in biomass and activity were hard to detect because soil tem-
peratures differed between the phenological stages. In addi-
tion, changes in activity correlate with changes in tempera-
ture and rainfall patterns and are “masked” by the tempera-
ture and soil moisture response curves (Kutsch et al., 2001b). The relatively high Q10-value for wet conditions (∼2.5) in-
dicates a hidden seasonal change in activity. W. L. Kutsch et al.: Savanna water-carbon relations The seasonal trend of FP,sat was an increase until March
and a decrease thereafter – a typical performance for a plant
canopy during the course of the growing season. This trend
was overlaid by a reduction during drought. The seasonal trends of all important parameters are sum-
marized in Fig. 8a–c. The coefficient c (Eq. 10) is an in-
dicator for the sensitivity of surface conductance to VPD,
FP,sat for the photosynthetic capacity. Rav is the average
nighttime respiration for each period and R15 the normal-
ized respiration rate at 15◦C and optimum water conditions. Night time respiration seemed to be highest at the on-
set of the growing season, was stable on a slightly lower
level during the main part of the growing season and de-
creased thereafter. This trend was overlaid by depressions Biogeosciences, 5, 1797–1808, 2008 Biogeosciences, 5, 1797–1808, 2008 www.biogeosciences.net/5/1797/2008/ W. L. Kutsch et al.: Savanna water-carbon relations 1804 0
0
1
2
3
4
5
10
15
20
J
F
M
A
M
J
J
A
S
Coefficient c
0
0.02
0.04
0.06
0.08
0.1
FP,max (µmol CO2 m-2 s-1)
Rav and R15
(µmol CO2 m-2 s-1)
R15
Rav
Fig. 8. Annual courses of the coefficient c describing stomatal
response sensitivity to VPD, of maximum canopy photosynthesis
(FP,sat) average nighttime respiration (Rav) and R15 as derived
from the model. Each point represents the center of a period de-
fined in Table 1. The dashed line shows a hypothetical seasonal
course without intermediate drought periods. 0
0
1
2
3
4
5
10
15
20
J
F
M
A
M
J
J
A
S
Coefficient c
0
0.02
0.04
0.06
0.08
0.1
FP,max (µmol CO2 m-2 s-1)
Rav and R15
(µmol CO2 m-2 s-1)
R15
Rav 0
5
10
15
20
25
0
5
10
15
20
25
FP,max (µmol CO2 m-2 s-1)
FP,max (µmol CO2 m-2 s-1)
0
5
10
15
0
0.05
0.1
Coefficient c
GS at FP,max (mm s-1)
Fig. 9. Relations between photosynthetic capacity (FP,sat) and
canopy conductance at FP,sat (a) and between FP,sat and the co-
efficient c describing stomatal response sensitivity to VPD (b) for
the 8 periods under consideration. 0
5
10
15
20
25
0
5
10
15
20
25
FP,max (µmol CO2 m-2 s-1)
FP,max (µmol CO2 m-2 s-1)
0
5
10
15
0
0.05
0.1
Coefficient c
GS at FP,max (mm s-1) Fig. 9. W. L. Kutsch et al.: Savanna water-carbon relations This was con-
firmed by comparing mean night time fluxes throughout the
year: highest respiration rates were found at the onset of the
wet season in January and February, followed by constant de-
crease thereafter that was overlaid by a less severe influence
of water shortage during the growing period (Fig. 8c). Figure 9 summarizes the close relationship between stom-
atal regulation and photosynthetic capacity by the close re-
lationship between canopy photosynthetic capacity and sur-
face conductance (Fig. 9a) and, in addition, a strong correla-
tion between coefficient c and canopy photosynthetic capac-
ity (Fig. 9b) for the 8 periods under consideration. 4
Discussion The convergence of the two savanna types in terms of flux
properties may be the most surprising result of this study. Sectoral analysis of flux data has been conducted success-
fully in other studies, where distinct differences in fluxes be-
tween sectors could be related to differences in properties
such as species composition or LAI of the canopy (Kutsch
et al., 2005). At this site the fluxes were identical in spite
of different soil properties and plant species composition as
described by Scholes et al. (2001). It is assumed that this is due to the growth respiration of
the new biomass and the mineralization of easily decompos-
able organic matter that has been accumulated during the
dry season. Augustine and McNaughton (2005) showed re-
cently that the new growing plant biomass at the onset of the Biogeosciences, 5, 1797–1808, 2008 www.biogeosciences.net/5/1797/2008/ 1805 W. L. Kutsch et al.: Savanna water-carbon relations Kutsch et al. (2001a) showed by means of model calcu-
lations that stomatal acclimation allows the plant to opti-
mize for high carbon gain during periods of sufficient wa-
ter supply and to save water during drought. It is impor-
tant to note that decreases in stomatal conductance in many
plants are inter-related with decreases in photosynthetic ca-
pacity (Hall and Schulze, 1980; Schulze and Hall, 1982). Whereas those authors assumed an un-coordinated response
of stomatal conductance and photosynthetic capacity to soil
water stress (Schulze and Hall, 1982), Flexas et al. (2006a, b)
showed recently on the molecular level that stomatal closure
triggers down-regulation of Rubisco either directly through
its effects on chloroplast CO2 concentration or mediated by
some common signalling (e.g. a response to abscisic acid). Data from this study were obtained by eddy covariance at the
ecosystem scale and show the same pattern of inter-relation
between stomatal conductance and photosynthetic capacity
as leaf-level studies. growing season is also favoured by a large inorganic nitro-
gen pool that has accumulated during the dry season, dur-
ing which mineralization does not completely cease. Xu et
al. (2004) found a similar pattern oak/grass savanna ecosys-
tem in California. They also observed a “respiratory flush”
after a rainfall event, as did Levine et al. (1996) at Preto-
riuskop (30 km south of the Skukuza site) and Scholes et
al. (1997) in savannas at Nylsvley, South Africa. 4
Discussion Maximum canopy-scale carbon dioxide fluxes at light sat-
uration (FC,sat:−17 µmol CO2 m−2 s−1) and canopy-scale
photosynthetic capacity (FP,sat:−22 µmol CO2 m−2 s−1) re-
ported in this study are comparable with other eddy-
covariance studies of African savanna ecosystems, particu-
larly when bearing in mind that the 2002/2003 growing sea-
son at Skukuza was characterised by an extremely low pre-
cipitation. Scanlon and Albertson (2004) measured fluxes
from four savanna ecosystems along a precipitation transect
across the Kalahari sands. They report values of FP,sat be-
tween −15 and −20 µmol CO2 m−2 s−1 for the “drier end”
of the transect (mean annual rainfall 365 and 407 mm). A
similar range is also reported in Williams and Albertson
(2004) for a mixed Acacia-Combretum savanna in Botswana
receiving a mean annual rainfall of 400 mm, and by Veenen-
daal et al. (2004) for a Mopani-woodland in Botswana with
464 mm mean annual precipitation. It is important to note that the decrease in stomatal con-
ductance goes together with the down-regulation of photo-
synthetic capacity. As a consequence, Ci/Ca may remain rel-
atively stable and the isotopic fractionation will not be af-
fected by drought as severely as expected. Therefore, pre-
dicted values of isotopic fractionation from large scale mod-
eling may be wrong. The correlation between canopy photo-
synthetic capacity and maximum surface conductance seems
to be relevant for scaling processes from leaf to ecosystem
level and even to higher scales (e.g. Lloyd et al., 2008). Scanlon and Albertson (2004), who collected only short-
term data of a few days during the 2000 wet season, inter-
preted differences in the light response curves of FC and the
VPD response curve of water use efficiency between the sites
as a long-term adaptation of the different vegetation types
to the mean precipitation at each location. In our study we
found a highly dynamic ecosystem response to environmen-
tal factors that may also be part of this adaptation. Our
approach of defining different periods according to rainfall
characteristics and soil moisture provided valuable qualita-
tive insights. However, it is important to stress that theories developed
for individual plants or leaves cannot be scaled simply to
the vegetation level because different water use strategies
and life cycles may compete for the available water and in
this situation a conservative strategy may simply lead to a
higher water usage by the competing neighbour (Schulze et
al., 2005). 4
Discussion Moreover, different plant functional types have
different traits and life cycles that may influence the ecosys-
tem properties much more than leaf scale acclimation. De-
creases in the fAPAR from remote sensing as shown in Fig. 2a
may indicate that the LAI is reduced during the dry periods. However, this observation has to be interpreted very care-
fully, because the correlation between fAPAR and LAI is scat-
tering (Huemmrich et al., 2005) and influenced by changing
optical properties of leaves during drought stress. Therefore,
further studies including detailed process analyses and multi-
species canopy modelling are highly required to explain the
adaptive inter-relation of canopy conductance and photosyn-
thetic capacity which is only poorly represented in higher-
scale models. In particular, it showed that the response of canopy con-
ductance to VPD followed the soil moisture conditions very
dynamically and that conductance and photosynthetic capac-
ity were strongly coupled. Three findings are noteworthy: 1. The short-term regulation of canopy conductance was
modified by a general seasonal trend and some mid-term
acclimation to soil water conditions. This can be seen
in the variation of the coefficient c in Figs. 8 and 9. 2. The most constant parameter was water use efficiency
(data not shown) that was influenced by VPD during
the day but the VPD response curve of water usage did
change only slightly during the course of the growing
season and decreased by about 30% during the transi-
tion from wet to dry season. 5
Conclusion 3. The regulation of canopy conductance and photosyn-
thetic capacity were closely related, suggesting a com-
plex regulation pattern (Fig. 9). The eddy covariance measurements at a flux tower close to
Skukuza in the Kruger National Park, South Africa, revealed
the strong influence of water relations on the carbon fluxes www.biogeosciences.net/5/1797/2008/ Biogeosciences, 5, 1797–1808, 2008 W. L. Kutsch et al.: Savanna water-carbon relations 1806 in savanna ecosystems. While Merbold et al. (2008) showed
the close relationship between canopy conductance and Fpsat
being constant over space, this study gave useful insights
into the temporal dynamics of acclimation to changing con-
ditions. The mid-term ecophysiological response to climate
and/or soil moisture, respectively, is an important observa-
tion which is not included in any of the common models. change of forests: The euroflux methodology, Adv. Ecol. Res.,
30, 113–175, 2000. Augustine, D. J. and McNaughton, S. J.: Temporal asynchrony in
soil nutrient dynamics and plant production in a semiarid ecosys-
tem, Ecosystems, 7, 829–840, 2005. Baldocchi, D. D., Xu, L. K., and Kiang, N.: How plant functional-
type, weather, seasonal drought, and soil physical properties alter
water and energy fluxes of an oak-grass savanna and an annual
grassland, Agr. Forest Meteorol., 123, 13–39, 2004. Acknowledgements. Eddy covariance measurements in the Kruger
Park were supported by the US National Aeronautics and Space
Administration (NASA, Grant NAG5-8705 and TE/03-000-0008
to NPH), the National Science Foundation (EAR-0120630 to
NPH), NOAA (Grant NA17RJ1228 to NPH), by the South African
National Research Foundation (RJS) and Center for Scientific
and Industrial Research (RJS). WLK was funded by the German
Science Foundation (DFG, DFG Ku 1099/2-1) and the European
Commission (FP6, Project “CarboAfrica”, Nr. 037132 and strongly
supported by the Christian-Albrechts-University, Kiel. Construc-
tion of the micrometeorological tower near Skukuza was made
possible through NASA funding to Jeff Privette for participation in
EOS Validation and SAFARI-2000. The scientific collaboration of
Mary Scholes, Harold Annegarn, Lackson Marufu, Pierre Ngok and
numerous SAFARI-2000 personnel is gratefully acknowledged. The researchers and staff of Scientific Services in the Kruger
National Park provided scientific, technical and logistical support. Finally, the authors wish to thank the game-guards of the Kruger
National Park who kept us save during field work. Ball, J. T., Woodrow, I. E., and Berry, J. 5
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RPA and Pif1 cooperate to remove G-rich structures at both leading and lagging strand
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RPA and Pif1 cooperate to remove G-rich structures at
both leading and lagging strand
Laetitia Maestroni, Julien Audry, Pierre Luciano, Stephane Coulon, Vincent
Géli, Yves Corda a Maestroni, Julien Audry, Pierre Luciano, Stephane Coulon, Vincent
Géli, Yves Corda To cite this version: Laetitia Maestroni, Julien Audry, Pierre Luciano, Stephane Coulon, Vincent Géli, et al.. RPA and
Pif1 cooperate to remove G-rich structures at both leading and lagging strand. Cell Stress, 2020, 4,
pp.48 - 63. 10.15698/cst2020.03.214. hal-03012937 Research Article www.cell-stress.com HAL Id: hal-03012937
https://hal.science/hal-03012937v1
Submitted on 18 Nov 2020 L’archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
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lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers. RPA and Pif1 cooperate to remove G-rich structures at
both leading and lagging strand Laetitia Maestroni1, Julien Audry1, Pierre Luciano1, Stéphane Coulon1, Vincent Géli1,* and Yves Corda1,*
1 Aix-Marseille Univ, Inserm, CNRS, Institut Paoli-Calmettes, CRCM, Marseille, France. Equipe Labellisée par la Ligue Nationale contre
le Cancer. etitia Maestroni1, Julien Audry1, Pierre Luciano1, Stéphane Coulon1, Vincent Géli1,* and Yves C
x-Marseille Univ, Inserm, CNRS, Institut Paoli-Calmettes, CRCM, Marseille, France. Equipe Labellisée par la Ligue Nat
Cancer. 1 Aix-Marseille Univ, Inserm, CNRS, Institut Paoli-Calmettes, CRCM, Marseille, France. Equipe Labellisée par la Ligue
le Cancer. * Corresponding Authors: p
g
Vincent Géli, Aix-Marseille Univ, Inserm, CNRS, Institut Paoli-Calmettes, CRCM, Marseille, France. Equipe Labellisée par la Ligue
Nationale contre le Cancer ; E-mail: vincent.geli@inserm.fr; Yves Corda, Aix-Marseille Univ, Inserm, CNRS, Institut Paoli-Calmettes, CRCM, Marseille, France. Equipe Labellisée par la Ligue
Nationale contre le Cancer ; E-mail: yves.corda@inserm.fr doi: 10.15698/cst2020.03.214
Received originally: 25.09.2018;
in revised form: 03.01.2020,
Accepted 07.01.2020,
Published 17.01.2020. ABSTRACT In Saccharomyces cerevisiae, the absence of Pif1 helicase induces
the instability of G4-containing CEB1 minisatellite during leading strand but
not lagging strand replication. We report that RPA and Pif1 cooperate to
maintain CEB1 stability when the G4 forming strand is either on the leading or
lagging strand templates. At the leading strand, RPA acts in the same pathway
as Pif1 to maintain CEB1 stability. Consistent with this result, RPA co-
precipitates with Pif1. This association between Pif1 and RPA is affected by
the rfa1-D228Y mutation that lowers the affinity of RPA in particular for
G-rich single-stranded DNA. At the lagging strand, in contrast to pif1∆, the
rfa1-D228Y mutation strongly increases the frequency of CEB1 rearrange-
ments. We explain that Pif1 is dispensable at the lagging strand DNA by the
ability of RPA by itself to prevent formation of stable G-rich secondary struc-
tures during lagging strand synthesis. Remarkably, overexpression of Pif1 res-
cues the instability of CEB1 at the lagging strand in the rfa1-D228Y mutant
indicating that Pif1 can also act at the lagging strand. We show that the ef-
fects of the rfa1-D228Y (rpa1-D223Y in fission yeast) are conserved in Schiz-
osaccharomyces pombe. Finally, we report that RNase H1 interacts in a DNA-
dependent manner with RPA in budding yeast, however overexpression of
RNase H1 does not rescue CEB1 instability observed in pif1∆ and rfa1-D228Y
mutants. Collectively these results add new insights about the general role of
RPA in preventing formation of DNA secondary structures and in coordinating
the action of factors aimed at resolving them. RPA and Pif1 cooperate to remove G-rich structures at
both leading and lagging strand Keywords: RPA, Pif1, human
minisatellite CEB1, G-rich structures,
G-quadruplex. Keywords: RPA, Pif1, human
minisatellite CEB1, G-rich structures,
G-quadruplex. Abbreviations:
Co-IP – Co-immunoprecipitation,
RPA – replication protein A,
ssDNA – single-stranded DNA,
WT – wild type. INTRODUCTION INTRODUCTION fragments by governing the sequential action of Dna2 and
Fen1 [5]. During Pol δ synthesis most flaps generated on
the lagging strand, by strand displacement, are normally
cleaved by Fen1. However, a minor fraction escapes cleav-
age [6], and the 5' ssDNA flaps on Okazaki fragments get
extended by Pif1, a 5' to 3' helicase, to create substrates
for RPA binding that inhibits Fen1’s cleavage [5,7-11]. RPA
is next displaced by Dna2 which cleaves the long flap, gen-
erating a short flap structure that undergoes cleavage via
Fen1 [12-14]. Replication protein A (RPA) is the major eukaryotic single-
stranded DNA (ssDNA) binding protein that consists of 70,
32, and 14 kDa subunits [1]. RPA plays a key role in coordi-
nating DNA synthesis, repair, and DNA damage signalling
through binding to single-stranded DNA (ssDNA) interme-
diates generated during these processes [2]. RPA primarily
maintains ssDNA in an unfolded state through different
binding modes that are characterized by the length of the
interacting ssDNA [3]. Thus, RPA binds to ssDNA with high
affinity preventing the formation of DNA secondary struc-
tures and annealing of homologous sequences [4]. Among
its activities, RPA is involved in the maturation of Okazaki Interestingly, it has been reported in vitro, that RPA
binds to and unwinds G4 structures in a 5’ to 3’ direction
[15]. G4 are polymorphic and consist of four-stranded Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 48 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) mutant (rpa1-D223Y in fission yeast) also possesses a low-
er affinity for ssDNA and reduced ability in removing sec-
ondary structure from ssDNA [39, 40]. In this study, we
investigated the role of RPA in maintaining the stability of
CEB1 when the G-quadruplex-forming strand is either on
the leading or lagging strand template. Our results indicate
that both RPA and Pif1 cooperate at the leading strand to
maintain the stability of CEB1. Consistent with this hy-
pothesis, RPA co-precipitates with Pif1. In contrast to Pif1,
RPA is also required to stabilize CEB1 when the
G-quadruplex-forming strand is the lagging strand. How-
ever, under a situation that compromises RPA binding to
ssDNA, overexpression of Pif1 rescued lagging-CEB1 insta-
bility, suggesting that Pif1 can unwind G4 at the lagging
strand. INTRODUCTION Interestingly, Mms1 which binds to G-rich/G4 re-
gions and supports the binding of Pif1, is not required to
maintain the stability of CEB1. Based on these data we
propose a model in which RPA facilitates Pif1 action at the
leading strand DNA to unwind G4 while enriched RPA at
the lagging strand DNA prevents by itself formation of sta-
ble G4, explaining why Pif1 is dispensable. We extended
the role of RPA in preventing non-templated DNA single-
stranded structure by showing that RPA interacts with
RNAse H1 in a DNA-dependent manner in S. cerevisiae, as
previously reported in human cells [41]. However, we
found that overexpressing RNAse H1 did not restore CEB1
stability in both pif1∆ and rfa1-D228Y mutants, suggesting
that CEB1 instability is not due to R-Loop formation arising
during transcription. structures formed at specific G-rich motifs within DNA,
RNA, and into R loops, a RNA-DNA hybrid structure, that
can eventually lead to genome instability [16-19]. The core
of these structures is formed by a square arrangement of
four guanines held together by Hoogsteen hydrogen bonds
[20, 21]. Under specific conditions, G4 structures are rec-
ognized by specific factors and their formation is controlled
[22, 23]. However, it has been shown that highly stable G4
structures impede fork progression. Hence, their unwind-
ing by helicases is critical [17, 24, 25]. Many helicases are
able to unwind G4 structures in vitro such as the RecQ hel-
icases BLM, WRN and Sgs1 and other helicases such as Pif1,
FANCJ or RTEL1 [24,26-30]. G4 structures are also targeted
by additional proteins that protect them [23] or support
the function of an helicase at G4 [22]. In budding yeast,
unwinding of G4 is mainly performed by the Pif1 helicase
[26]. Indeed, a particular example is the 1.8 kb G4-forming
human minisatellite CEB1, a reporter of G4 formation and
processing [31, 32]. In cells lacking Pif1, CEB1 is unstable
when inserted into Saccharomyces cerevisiae genome,
near an early origin of replication (ARS305). Instability of
CEB1, which consists in 42 motifs of 39 nucleotides ar-
ranged as direct repeats, was correlated to the ability of
the CEB1 motif to form G4 [31, 33, 34]. Surprisingly, in
pif1∆ cells CEB1 was unstable only when the G-quadruplex-
forming strand was the leading strand template [33]. RESULTS We have previously shown in fission yeast that the
rpa1-D223Y mutation (rfa1-D228Y in budding yeast), that
exhibits a reduced affinity for ssDNA, impaired lagging
strand telomere replication and provoked accumulation of
secondary structures [38]. Consistently, expression of
ScPif1 rescued the phenotypes associated with the rpa1-
D223Y mutation [38]. These results suggested that rpa1-
D223Y cells accumulated G-rich structures at lagging
strand telomeres that were resolved by the heterologous
expression of ScPif1. In S. cerevisiae, RPA subunits are en-
coded by RFA1, RFA2, and RFA3 genes. The rfa1-D228Y INTRODUCTION This
result is apparently counterintuitive since ssDNA is mostly
formed during the discontinuous synthesis of the lagging
strand DNA [35] and because Pif1 has been reported to act
at the lagging strand [36] and binds G4 structures located
in the lagging strand [22, 37]. The rfa1-D228Y mutation affects both the leading-CEB1
and lagging-CEB1 As mentioned above, the G-rich minisatellite CEB1 can be
considered as a reporter of G4 formation and processing
[31, 32]. We used strains previously constructed in A. Nico-
las’ laboratory in which the 1.8 kb CEB1 is inserted in both
directions at 2.1 kb of ARS305 and 32.6 kb away from
ARS306 allowing to primarily replicate CEB1 only from the
proximal ARS305 origin (Figure 1). Depending on the orien-
tation of CEB1 insertion, the G4-forming strand will be rep-
licated by the leading or the lagging machinery [33]. We FIGURE 1: Map of the CEB1 insertion
within chromosome III at 2.1 kb
from the ARS305. In orientation I, the
G-quadruplex-forming strand is the
template of the leading polymerase
(leading CEB1). In orientation II, the
G-quadruplex-forming strand is the
template of the lagging polymerase
(lagging CEB1). FIGURE 1: Map of the CEB1 insertion
within chromosome III at 2.1 kb
from the ARS305. In orientation I, the OPEN ACCESS | www.cell-stress.com
49
Cell Stress | MARCH 2020 | Vol. 4 No. 3
FIGURE 1: Map of the CEB1 insertion
within chromosome III at 2.1 kb
from the ARS305. In orientation I, the
G-quadruplex-forming strand is the
template of the leading polymerase
(leading CEB1). In orientation II, the
G-quadruplex-forming strand is the
template of the lagging polymerase
(lagging CEB1). FIGURE 1: Map of the CEB1 insertion
within chromosome III at 2.1 kb
from the ARS305. In orientation I, the
G-quadruplex-forming strand is the
template of the leading polymerase
(leading CEB1). In orientation II, the
G-quadruplex-forming strand is the
template of the lagging polymerase
(lagging CEB1). G-quadruplex-forming strand is the
template of the leading polymerase
(leading CEB1). In orientation II, the
G-quadruplex-forming strand is the
template of the lagging polymerase
(lagging CEB1). Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 49 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) FIGURE 2: RPA is required to stabilize CEB1. (A) Experimental scheme. Yeast strains of interest are mated and the resulting diploid sporulated and dissec
After identification of spore-colonies of interest, the spore-colony is plated on media to obtain isolated colonies. Individual colonies are placed in liquid cult
until stationary growth phase. Genomic DNAs are extracted and analysed by Southern blot. (B) RPA is required to stabilize CEB1 when the G4-forming stran
replicated by the leading polymerase. The rfa1-D228Y mutation affects both the leading-CEB1
and lagging-CEB1 Genomic DNAs from yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI, and southern blotted. M
branes were hybridized with the CEB1-0.6 probe. (C) Genomic DNAs from yeast cells containing the leading-CEB1Gmut-1.7 were digested by ApaI and SacII,
southern blotted. The membranes were hybridized with the CEB1Gmut-1.7 probe. (D) In contrast to Pif1, RPA is required to stabilize CEB1 when the G4-form
strand is replicated by the lagging polymerase. Genomic DNAs from yeast cells bearing the lagging-CEB1 were digested by ApaI and NcoI, and southern blot
Membranes were hybridized with the CEB1-0.6 probe. M: ladder DNA serving as size standard (kbp). The number of colonies analysed per well, the percent
of rearrangement frequencies and the total numbers of colonies are indicated in Table 1 FIGURE 2: RPA is required to stabilize CEB1. (A) Experimental scheme. Yeast strains of interest are mated and the resulting diploid sporulated and disse
After identification of spore-colonies of interest, the spore-colony is plated on media to obtain isolated colonies. Individual colonies are placed in liquid cu
until stationary growth phase. Genomic DNAs are extracted and analysed by Southern blot. (B) RPA is required to stabilize CEB1 when the G4-forming stra
replicated by the leading polymerase. Genomic DNAs from yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI, and southern blotted. M
branes were hybridized with the CEB1-0.6 probe. (C) Genomic DNAs from yeast cells containing the leading-CEB1Gmut-1.7 were digested by ApaI and SacI GURE 2: RPA is required to stabilize CEB1. (A) Experimental scheme. Yeast strains of interest are mated and the resulting diploid sporulated and dis
ter identification of spore-colonies of interest, the spore-colony is plated on media to obtain isolated colonies. Individual colonies are placed in liquid
til stationary growth phase. Genomic DNAs are extracted and analysed by Southern blot. (B) RPA is required to stabilize CEB1 when the G4-forming s
plicated by the leading polymerase. Genomic DNAs from yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI, and southern blotted
anes were hybridized with the CEB1-0.6 probe. (C) Genomic DNAs from yeast cells containing the leading-CEB1Gmut-1.7 were digested by ApaI and Sa
uthern blotted. The membranes were hybridized with the CEB1Gmut-1.7 probe. (D) In contrast to Pif1, RPA is required to stabilize CEB1 when the G4-
and is replicated by the lagging polymerase. The rfa1-D228Y mutation affects both the leading-CEB1
and lagging-CEB1 Genomic DNAs from yeast cells bearing the lagging-CEB1 were digested by ApaI and NcoI, and southern
embranes were hybridized with the CEB1-0.6 probe. M: ladder DNA serving as size standard (kbp). The number of colonies analysed per well, the per
rearrangement frequencies, and the total numbers of colonies are indicated in Table 1. FIGURE 2: RPA is required to stabilize CEB1. (A) Experimental scheme. Yeast strains of interest are mated and the resulting diploid sporulated and dissected. After identification of spore-colonies of interest, the spore-colony is plated on media to obtain isolated colonies. Individual colonies are placed in liquid culture
until stationary growth phase. Genomic DNAs are extracted and analysed by Southern blot. (B) RPA is required to stabilize CEB1 when the G4-forming strand is
replicated by the leading polymerase. Genomic DNAs from yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI, and southern blotted. Mem-
branes were hybridized with the CEB1-0.6 probe. (C) Genomic DNAs from yeast cells containing the leading-CEB1Gmut-1.7 were digested by ApaI and SacII, and
southern blotted. The membranes were hybridized with the CEB1Gmut-1.7 probe. (D) In contrast to Pif1, RPA is required to stabilize CEB1 when the G4-forming
strand is replicated by the lagging polymerase. Genomic DNAs from yeast cells bearing the lagging-CEB1 were digested by ApaI and NcoI, and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. M: ladder DNA serving as size standard (kbp). The number of colonies analysed per well, the percentage
of rearrangement frequencies, and the total numbers of colonies are indicated in Table 1. Cell Stress | MARCH 2020 | Vol. 4 No. 3 50 OPEN ACCESS | www.cell-stress.com OPEN ACCESS | www.cell-stress.com A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) rangements of the lagging-CEB1 was clearly increased in
the rfa1-D228Y mutant (31%). These results suggest that
when G4-forming sequences are localized at the lagging
strand, RPA prevents the instability of CEB1. They are con-
sistent with the known enrichment of RPA at the lagging
strand [35]. Interestingly, despite the fact that pif1∆ did
not affect the stability of the lagging-CEB1, combining
pif1∆ with rfa1-D228Y, slightly aggravated the CEB1 insta-
bility phenotype of the single rfa1-D228Y mutant (41%). This result suggests that Pif1 can be active at the lagging-
CEB1. will name leading-CEB1 or lagging-CEB1 to indicate the
machinery that replicates the G4-forming strand. The rfa1-D228Y mutation affects both the leading-CEB1
and lagging-CEB1 Previous
results demonstrated that the helicase activity of Pif1 was
required to stabilize the leading-CEB1 but that Pif1 was
dispensable for the stability of the lagging-CEB1 suggesting
the existence of different mechanisms to resolve G4 when
the G4-forming strand is replicated by the leading or the
lagging machinery [33]. We thought to test the role of RPA in the CEB1 stability
by determining the effect of the rfa1-D228Y mutant whose
DNA-binding activity, in particular to G-rich ssDNA, is com-
promised [38, 39]. We used an experimental scheme (Fig-
ure 2A) adapted from Lopes et al. [33]. Briefly, tetrads ob-
tained from the sporulation of heterozygous diploids were
dissected. Spore colonies carrying the appropriated muta-
tion were then resuspended in water and streaked on YPD
plates in order to obtain about 200 isolated colonies. Colo-
nies were inoculated in YPD liquid cultures, grown to satu-
ration, and genomic DNAs were prepared from the liquid
cultures. The size of CEB1 was monitored by Southern blot. We calculated the frequency of instability by monitoring
CEB1 size variations (contractions and expansions). We
considered that CEB1 was unstable when the intensity of
the band(s) was superior to the one of the parental band. We therefore partially discriminated between early or late
events of CEB1 rearrangements. Overexpression of Pif1 in rfa1-D228Y cells rescues lagging-
CEB1 instability To further understand the functional interaction between
RPA and Pif1 in CEB1 instability, we overexpressed Pif1 in
rfa1-D228Y cells bearing leading-CEB1 or lagging-CEB1. Because strong Pif1 overexpression impairs cell viability
[42], we used a low-copy centromeric plasmid in which the
nuclear form of Pif1 is under the control of the GAL1 pro-
moter [43]. We first noticed that Pif1 overexpression was
slightly deleterious in rfa1-D228Y cells (Figure 3A). To es-
timate Pif1 overexpression we measured the expression of
PIF1 by reverse transcription followed by quantitative PCR. Our data indicate that PIF1 is overexpressed in a similar
level in WT and rfa1-D228Y cells expressing PIF1 under the
control of the GAL1 promoter (Figure 3B, left and 3C, left). Instability of both leading- and lagging-CEB1 was tested in
rfa1-D228Y cells grown in galactose (SGal) allowing the
overexpression of Pif1. We found that Pif1 overexpression
had only a modest effect on leading-CEB1 stability in rfa1-
D228Y cells while it had no effect in WT cells (Figure 3B,
right). Surprisingly, Pif1 overexpression completely rescued
the instability of lagging-CEB1 in rfa1-D228Y cells (Figure
3C, right). We concluded that when the G4-forming strand
was replicated by the lagging machinery in cells in which
the ssDNA binding activity of RPA is compromised, overex-
pressed Pif1 could act at the lagging strand to unwind G4. We first analysed the stability of the leading-CEB1 (ori-
entation I) in wild type (WT), pif1∆, rfa1-D228Y, and pif1∆
rfa1-D228Y cells (Figure 2B). As shown in representative
gels and as previously reported [31], the leading-CEB1 was
stable in the WT strain but extremely unstable in the pif1∆
mutant (53% of rearrangements) (Figure 2B, Table 1). In
most cases, we obtained contractions except in one case
(Figure 2B, second panel). In the rfa-1D228Y mutant, lead-
ing-CEB1 was also unstable (51% of rearranged colonies). Interestingly, the double mutant pif1∆ rfa1-D228Y exhibit-
ed a level of instability in a similar range of the two single
mutants (43%), suggesting that RPA and Pif1 could act in
similar pathways to unwind G4 and stabilize CEB1 when
the G4-forming strand is replicated by the leading poly-
merase (Figure 2B). The rfa1-D228Y mutation affects the interaction between
Pif1 and RPA Our results indicate that RPA and Pif1 could cooperate to
stabilize the leading-CEB1 and the lagging-CEB1. This
prompted us to test whether Pif1 interacts with RPA. Pif1-
myc was immunoprecipitated with anti-Myc antibody and
tested for the presence of RPA with a polyclonal antibody
directed against Rfa1 or against Rfa2 (Figure 4). Pif1-myc
was efficiently co-immunoprecipitated with both Rfa1 and
Rfa2 but to a lesser extend with rfa1-D228Y. To address
whether the robust Co-immunoprecipitations (Co-IP) of
Pif1-myc with Rfa1 and Rfa2 were dependent on the pres-
ence of DNA, DNase1 was added in the lysate. As shown in
Figure 4, DNA digestion affected the Co-IP suggesting that
the presence of DNA is required for the robust co-
precipitation of Pif1 and RPA. These results suggest that
the robust association of Pif1 with RPA relies on specific
structures on the DNA. We think that the interaction be-
tween Pif1 and RPA is not only due to unspecific interac-
tions mediated by DNA since the interaction between Pif1 We next analysed the effect of pif1∆, rfa1-D228Y, and
pif1∆ rfa1-D228Y on the stability of CEB1-Gmut, a version
of CEB1 mutated for its G-quadruplex-forming sequences
[31, 32]. The CEB1-Gmut was inserted at the same location
and in the same orientation as the leading-CEB1. As re-
ported, the CEB1-Gmut was stable in pif1∆ cells [33]. In the
rfa1-D228Y and pif1∆ rfa1-D228Y mutants, the % of CEB1-
Gmut rearrangements dropped to 22% and 20%, respec-
tively (Figure 2C). We concluded that the high leading-
CEB1 instability observed in the rfa1-D228Y mutant partly
relies on its ability to form G4. However, the residual CEB1-
Gmut instability suggests that the rfa1-D228Y mutation
affects the replication of the 42 repeated motifs of CEB1-
mut irrespectively of the presence of G4. We next examined the stability of the lagging-CEB1
(orientation II) in the same mutants as above. As reported
[33], deleting PIF1 had no effect on the stability of the lag-
ging-CEB1 (Figure 2D). In contrast, the frequency of rear- Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 51 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) L. Maestroni et al. (2020) L. Maestroni et al. (2020)
A critical role for RPA in CEB1 stability FIGURE 3: Pif1 overexpression rescues lagging-CEB1 instability in the rfa1-D228Y mutant. The rfa1-D228Y mutation affects the interaction between
Pif1 and RPA (A) Colonies containing the leading- or lagging-CEB1,
overexpressing Pif1 (Pif1 overexpression) or not (Empty vector) were plated on galactose medium and subsequently incubated at 30°C. (B) Left: c
was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was determined using qPCR. (-): Empty vector, (+): Pif1 overexp
sion. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (Pif1 overexpression) and containing the leading-CEB1 w
treated as in Figure 2. M: ladder DNA. (C) Left: cDNA was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was de
mined using qPCR. (-): Empty vector, (+): Pif1 overexpression. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (
overexpression) and containing the lagging-CEB1 were treated as in Figure 2. M: ladder DNA. Yeast strains were grown in medium containing galac
(2%). The number of colonies analysed per well, the percentage of rearrangement frequencies, and the total numbers of colonies are indicated in T
1. FIGURE 3: Pif1 overexpression rescues lagging-CEB1 instability in the rfa1-D228Y mutant. (A) Colonies containing the leading- or lagging-CEB1,
overexpressing Pif1 (Pif1 overexpression) or not (Empty vector) were plated on galactose medium and subsequently incubated at 30°C. (B) Left: cD
was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was determined using qPCR. (-): Empty vector, (+): Pif1 overexp
sion. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (Pif1 overexpression) and containing the leading-CEB1 w
treated as in Figure 2. M: ladder DNA. (C) Left: cDNA was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was de
mined using qPCR. (-): Empty vector, (+): Pif1 overexpression. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (
overexpression) and containing the lagging-CEB1 were treated as in Figure 2. M: ladder DNA. Yeast strains were grown in medium containing galact
(2%)
h
b
f
l
i
l
d
ll
h
f
f
i
d h
l
b
f
l
i
i di
d i FIGURE 3: Pif1 overexpression rescues lagging-CEB1 instability in the rfa1-D228Y mutant. (A) Colonies containing the leadin FIGURE 3: Pif1 overexpression rescues lagging-CEB1 instability in the rfa1-D228Y mutant. (A) Colonies containing the leading- or lagging-CEB1, and
overexpressing Pif1 (Pif1 overexpression) or not (Empty vector) were plated on galactose medium and subsequently incubated at 30°C. Mms1 is not required to maintain the leading-CEB1 and
the lagging-CEB1 stability Mms1 supports Pif1 helicase binding to G4 structures [22]. We thus analysed the importance of Mms1 in CEB1 stabil-
ity. We found that in mms1∆ cells, leading-CEB1 and lag-
ging-CEB1 were rather stable, showing 0% (0/116) and 3%
(3/116) of rearrangements, respectively (Figure 5A, Table
1). These results show that Mms1 is not required for the
stability of both leading-CEB1 and lagging-CEB1. They sug-
gest that CEB1 minisatellites are not targeted by Mms1. Moreover, they indicate that Mms1 does not support Pif1
function at CEB1 and, together with our previous results
showing that Pif1 and RPA interact, suggest that RPA could
contribute to the recruitment of Pif1 to CEB1, likely by di-
rectly recruiting Pif1. RNase H1 interacts with RPA but its overexpression
doesn't rescue CEB1 instability in the absence of Pif1 or
reduced levels of RPA Transcription by RNA polymerase can form a three-
stranded structure called R-loop [46], which can facilitate
or stabilize secondary structure formation in the exposed
ssDNA [16, 47, 48]. Thus, G4 structures may also result
from exposure of the G4 forming strand by formation on
the other strand of RNA:DNA hybrids. This may create a
complex structure involving G4 DNA on one strand and a
RNA:DNA hybrid on the other strand [49]. Interestingly,
Pif1 has been proposed to have a patrolling role that re-
moves any G4 or RNA/DNA structure [50]. Indeed, Pif1
regulates R-loop formation at specific genomic loci [51]
and potentially complements RNAse H for R-loop resolu-
tion [52]. On the other side, systematic analysis of protein
complexes in S. cerevisiae have shown that RPA interacts
with RNase H1 [53]. Finally, RPA was recently proposed to
act as a sensor of R-loop in human cells and to recruit and
stimulate RNase H1 to counteract R-loops [41]. To further
document the potential cooperation of RPA with Pif1, we
first tested whether RPA interacts with RNase H1 in a DNA-
dependent manner in budding yeast. Expression of HA-
Rnh1 was induced by the addition of galactose and its in-
teraction with Rfa1 was probed by Co-IP. The results
shown in Figure 6 indicate that Rnh1 interacts with RPA in
a way that is stimulated by the presence of DNA, thereby
extending the results of Gavin et al. [53] and Nguyen et al. [41]. Interestingly, we found that the rfa1-D228Y mutant
had a less efficient ability to bind Rnh1 (Figure 6). These
results suggest that the leading-CEB1 instability observed
on the leading strand in absence of Pif1 and by reducing
the association of RPA with ssDNA (rfa1-D228Y and rtt105∆
mutants) could be the consequence of the presence of R-
Loops. To determine if R-loops were responsible of CEB1
instability we analysed the stability of the leading-CEB1
and lagging-CEB1 in WT, pif1∆, rfa1-D228Y, and rtt105∆
cells overexpressing RNase H1, the enzyme responsible to
resolve R-loops. We found that RNase H1 overexpression
did not affect CEB1 stability in WT cells, and did not rescue
CEB1 stability in pif1∆, rfa1-D228Y, and rtt105∆ mutants Mms1 is not required to maintain the leading-CEB1 and
the lagging-CEB1 stability The rfa1-D228Y mutation affects the interaction between
Pif1 and RPA (B) Left: cDNA
was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was determined using qPCR. (-): Empty vector, (+): Pif1 overexpres-
sion. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (Pif1 overexpression) and containing the leading-CEB1 were
treated as in Figure 2. M: ladder DNA. (C) Left: cDNA was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was deter-
mined using qPCR. (-): Empty vector, (+): Pif1 overexpression. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (Pif1
overexpression) and containing the lagging-CEB1 were treated as in Figure 2. M: ladder DNA. Yeast strains were grown in medium containing galactose
(2%). The number of colonies analysed per well, the percentage of rearrangement frequencies, and the total numbers of colonies are indicated in Table
1. FIGURE 3: Pif1 overexpression rescues lagging-CEB1 instability in the rfa1-D228Y mutant. (A) Colonies containing the leading- or lagging-CEB1, and
overexpressing Pif1 (Pif1 overexpression) or not (Empty vector) were plated on galactose medium and subsequently incubated at 30°C. (B) Left: cDNA
was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was determined using qPCR. (-): Empty vector, (+): Pif1 overexpres-
sion. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (Pif1 overexpression) and containing the leading-CEB1 were
treated as in Figure 2. M: ladder DNA. (C) Left: cDNA was prepared from the indicated strains. The ration of PIF1 transcript to that of ACT1 was deter-
mined using qPCR. (-): Empty vector, (+): Pif1 overexpression. Right, genomic DNAs from wild-type (WT) or rfa1-D228Y cells overexpressing Pif1 (Pif1
overexpression) and containing the lagging-CEB1 were treated as in Figure 2. M: ladder DNA. Yeast strains were grown in medium containing galactose
(2%). The number of colonies analysed per well, the percentage of rearrangement frequencies, and the total numbers of colonies are indicated in Table
1. Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 52 L. Maestroni et al. (2020) A critical role for RPA in CEB1 stability and rfa1-228Y is lost despite the fact that the rfa1-D228Y
still binds to DNA, although with lower affinity [38]. RNase H1 interacts w
doesn't rescue CEB1 ins
reduced levels of RPA FIGURE 4: The rfa1-D228Y muta-
tion affects the interaction be-
tween
Pif1
and
RPA. The rfa1-D228Y mutation affects the interaction between
Pif1 and RPA Co-
immunoprecipitation experiments
were performed in triplicate. Pif1-
myc is immunoprecipitated with an
anti-Myc antibody (9E10). The
presence of RPA in the Pif1-myc IP
is monitored with an anti-Rfa1 or
an anti-Rfa2 antibody. (+): cell
extracts treated with benzonase. The asterisk (*) indicates a non-
specific band. MW: molecular
weight. FIGURE 4: The rfa1-D228Y muta-
tion affects the interaction be-
tween
Pif1
and
RPA. Co-
immunoprecipitation experiments
were performed in triplicate. Pif1-
myc is immunoprecipitated with an
anti-Myc antibody (9E10). The
presence of RPA in the Pif1-myc IP
is monitored with an anti-Rfa1 or
an anti-Rfa2 antibody. (+): cell
extracts treated with benzonase. The asterisk (*) indicates a non-
specific band. MW: molecular
weight. and rfa1-228Y is lost despite the fact that the rfa1-D228Y
still binds to DNA, although with lower affinity [38]. RNase H1 interacts with RPA but its overexpression
doesn't rescue CEB1 instability in the absence of Pif1 or
reduced levels of RPA Rtt105 is required to stabilize both the leading-CEB1 and
lagging-CEB1 Rtt105 functions as an RPA chaperone that escorts RPA to
the nucleus and facilitates RPA loading onto ssDNA [44]. Consequently, RTT105 inactivation reduces the association
of RPA with ssDNA generated during DNA transactions and
affects multiple RPA functions [44, 45]. We examined the
importance of Rtt105 on the stability of the leading-CEB1
and lagging-CEB1. We found that CEB1 is extremely unsta-
ble in rtt105∆ cells. Remarkably, we found that the fre-
quency of rearrangements reaches 100% in both leading-
and lagging-CEB1 (Figure 5B, Table 1). These results reveal
the importance of RTT105 in promoting the replication of
G4-forming CEB1 minisatellite during leading and lagging
strand synthesis and more generally, to maintain genome
stability. They confirm our previous results obtained with
rfa1-D228Y mutant indicating that RPA is crucial to remove
G-rich structures at both leading and lagging strand. OPEN ACCESS | www.cell-stress.com OPEN ACCESS | www.cell-stress.com Cell Stress | MARCH 2020 | Vol. 4 No. 3 53 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) FIGURE 5: Importance of Mms1 and Rtt105 on CEB1 stability. (A) Mms1 is not required to stabilize CEB1. Left, genomic DNAs from mms1∆ yeast
cells bearing the leading-CEB1 were digested by ApaI and XhoI and southern blotted. Right, genomic DNAs from mms1∆ yeast cells bearing the lag-
ging-CEB1 were digested by ApaI and NcoI and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. (B) Rtt105 is required to
stabilize both leading-CEB1 and lagging-CEB1. Left, genomic DNAs from rtt105∆ yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI
and southern blotted. Right, genomic DNAs from rtt105∆ yeast cells bearing the lagging-CEB1 were digested by ApaI and NcoI and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. WT: wild-type genomic DNA. The arrows show the position of stable leading-CEB1 (left) and
stable lagging-CEB1 (right). The horizontal dashed lines indicate the theoretical position of stable CEB1. M: ladder DNA serving as size standard (kbp). The number of colonies analysed per well, the percentage of rearrangement frequencies, and the total numbers of colonies are indicated in Table 1. FIGURE 5: Importance of Mms1 and Rtt105 on CEB1 stability. (A) Mms1 is not required to stabilize CEB1. Left, genomic DNAs from mms1∆ yeast
cells bearing the leading-CEB1 were digested by ApaI and XhoI and southern blotted. Rtt105 is required to stabilize both the leading-CEB1 and
lagging-CEB1 Right, genomic DNAs from mms1∆ yeast cells bearing the lag-
ging-CEB1 were digested by ApaI and NcoI and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. (B) Rtt105 is required to
stabilize both leading-CEB1 and lagging-CEB1. Left, genomic DNAs from rtt105∆ yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI
and southern blotted. Right, genomic DNAs from rtt105∆ yeast cells bearing the lagging-CEB1 were digested by ApaI and NcoI and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. WT: wild-type genomic DNA. The arrows show the position of stable leading-CEB1 (left) and
stable lagging-CEB1 (right). The horizontal dashed lines indicate the theoretical position of stable CEB1. M: ladder DNA serving as size standard (kbp). The number of colonies analysed per well, the percentage of rearrangement frequencies, and the total numbers of colonies are indicated in Table 1. FIGURE 5: Importance of Mms1 and Rtt105 on CEB1 stability. (A) Mms1 is not required to stabilize CEB1. Left, genomic DNAs from mms1∆ yeast
cells bearing the leading-CEB1 were digested by ApaI and XhoI and southern blotted. Right, genomic DNAs from mms1∆ yeast cells bearing the lag-
ging-CEB1 were digested by ApaI and NcoI and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. (B) Rtt105 is required to
stabilize both leading-CEB1 and lagging-CEB1. Left, genomic DNAs from rtt105∆ yeast cells bearing the leading-CEB1 were digested by ApaI and XhoI
and southern blotted. Right, genomic DNAs from rtt105∆ yeast cells bearing the lagging-CEB1 were digested by ApaI and NcoI and southern blotted. Membranes were hybridized with the CEB1-0.6 probe. WT: wild-type genomic DNA. The arrows show the position of stable leading-CEB1 (left) and
stable lagging-CEB1 (right). The horizontal dashed lines indicate the theoretical position of stable CEB1. M: ladder DNA serving as size standard (kbp). The number of colonies analysed per well, the percentage of rearrangement frequencies, and the total numbers of colonies are indicated in Table 1. Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 54 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) FIGURE 6: Rnh1 interacts with RPA. Co-immunoprecipitation experiments were performed in triplicate. HA-Rnh1 has been overexpressed by
galactose addition and immunoprecipitated with an anti-HA antibody. The presence of RPA in the HA-Rnh1 IP is monitored with an anti-Rfa1
antibody. Cell extracts treated with nuclease (Thermofisher) are indicated (+). Rtt105 is required to stabilize both the leading-CEB1 and
lagging-CEB1 Yeast strains were grown in medium containing 2% glucose (-)
or 2% galactose (+) when mentioned. Total proteins on the membrane were stained with Ponceau S as a loading control (Bottom). MW: mo-
lecular weight. FIGURE 6: Rnh1 interacts with RPA. Co-immunoprecipitation experiments were performed in triplicate. HA-Rnh1 has been overexpressed by
galactose addition and immunoprecipitated with an anti-HA antibody. The presence of RPA in the HA-Rnh1 IP is monitored with an anti-Rfa1
antibody. Cell extracts treated with nuclease (Thermofisher) are indicated (+). Yeast strains were grown in medium containing 2% glucose (-)
or 2% galactose (+) when mentioned. Total proteins on the membrane were stained with Ponceau S as a loading control (Bottom). MW: mo-
lecular weight. (Table 1). Taken together, our data suggest that the insta-
bility observed in absence of Pif1 and reduced levels of RPA
is not primarily related to R-loop formation. was calculated in both WT and rpa1-D223Y strains. The
level of instability of CEB25-L1T was 14% and 20% in the
WT (Figure 7B, left). In the rpa1-D223Y mutant, this level
increased up to 62% and 74%, respectively (Figure 7B,
right). Expectedly, the instability of CEB25-L1T-G12T that
cannot form G4 was reduced to 4% for leading replication
and 17% for lagging replication (Figure 7C). These results
clearly show that like in budding yeast the RPA complex
plays an important role in the replication and stability of
G4-forming sequences. Furthermore, the fact that fission
yeast lacks a functional homolog of Pif1 helicase, the Pfh1
helicase behaving more closely like the budding yeast Rrm3
helicase [56], may account for the prominent role of RPA
complex in G4 unwinding in fission yeast. The rpa1-D223Y mutation affects the stability of CEB25-
L1T in fission yeast Because the rpa1-D223Y mutation impaired replication of
the G-rich lagging strand telomere in Schizosaccharomyces
pombe, we investigated the stability of CEB in fission yeast. We took advantage of the minisatellite CEB25-L1T which
contains 14 repeats of a 44 nucleotides-sequence (0.62 kb)
that form a stable G4 [34, 54]. In this L1T version, the loop
of the repeated sequence of CEB25 has been reduced to
one thymine nucleotide. The shortening of the loop in-
creases its thermal stability in correlation with the in vivo
instability [54]. As a control the CEB25-L1T-G12T has been
used in which the guanine at the 12th position of the re-
peated sequence has been mutated into thymine, prevent-
ing the formation of G4. We introduced the CEB25-L1T and
CEB25-L1T-G12T in both orientations into the genome of
yeast cells at the leu1 locus at chromosome 2. The leu1
locus is located in between ARS-II-1964 and ARS-II-1983 at
a distance of 14 kb and 11 kb from CEB25, respectively
(Figure 7A). These two ARS have a relatively low firing effi-
ciency, 32% and 13%, respectively [55]. We monitored in
fission yeast the instability of CEB25 in both orientations
although this genomic context does not allow to clearly
distinguish whether the G4-forming sequence is replicated
by the leading or the lagging replication machinery. DISCUSSION We therefore propose that RPA cooperates
with Pif1 to resolved G-quadruplexes during leading strand
replication. Several interpretations can be invoked to explain the
difference in CEB1 stability observed in pif1∆ cells during
leading and lagging replication. It is possible that RPA re-
cruits Pif1. We think that the difference in Pif1 require-
ment at leading and lagging strands is related to RPA func-
tion. At the leading strand, both Pif1 helicase and RPA are
required. However, our results showing that mutated CEB1
(which are no longer able to form G4-strutures) and Pif1
overexpression both only partially rescue instability in rfa1-
D228Y cells reveal higher requirement of Pif1 compared to
RPA to maintain CEB1 stability at the leading strand. Inter-
estingly the roles are inverted between RPA and Pif1 at the
lagging strand. Because contrary to the leading strand the
lagging strand contains longer stretch of ssDNA between
elongating Okazaki fragments, RPA is present at a higher
concentration at the lagging strand [59]. Consequently, by
preventing formation/folding of G-quadruplex structures at
the exposed ssDNA, RPA is likely to directly contribute to
CEB1 stability independently of Pif1 at the lagging strand. Because rfa1-D228Y possesses a lower affinity for ssDNA
and a reduced ability in preventing and removing second-
ary structures from ssDNA, in rfa1-D228Y cells CEB1 be-
comes unstable. In that situation Pif1, which is not initially
required to remove G-quadruplex at the lagging strand, is
now required. Our results showing that the Pif1 overex-
pression almost fully rescues the stability of CEB1 in rfa1-
D228Y mutant at the lagging strand suggest that CEB1-G-
quadruplexes are formed in this mutant and can be pro- p
In contrast to Pif1, which is not required for CEB1 sta-
bility during lagging strand replication, we found that RPA
has a prominence in maintaining lagging CEB1 stability. This finding points out the differential behaviour of the
pif1∆ and rfa1-D228Y mutations according to the direction
of replication. However, our results indicate that Pif1 over-
expression drastically reduced the instability observed at
the lagging strand in the rfa1-D228Y mutant whose ssDNA
binding activity of RPA is compromised, in particular to
G-rich regions [38]. Finally, we report that Mms1, which
binds G4-structures and aids Pif1 binding to these struc-
tures is not required for CEB1 stability at both leading and
lagging strands. DISCUSSION In S. cerevisiae, Pif1 helicase processes G-rich secondary
structures thereby preventing deleterious events that may
lead to DNA breaks [33,38,57]. In vitro, Pif1 binds tightly to
G4 structures and unwinds them very efficiently [26,58]. Previous studies demonstrated that Pif1 prevents the
formation of G-quadruplex-dependent CEB1 internal rear-
rangements during leading strand, but not lagging strand
replication [33]. Here, we disclose that RPA cooperates
with Pif1 to remove G-quadruplex structures at both lead-
ing and lagging strand. We found that the rfa1-D228Y mu-
tation increases the frequency of CEB1 rearrangements
when the G-quadruplex forming strand is replicated by the
leading polymerase. The level of rearrangements is similar
to the level observed in pif1∆ cells. We report that mutat-
ing G-quadruplex-forming sequences strongly decreases As depicted in Figure 2A, the size of CEB25-L1T was
monitored by Southern blot and frequency of instability Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 55 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) TABLE 1. Rearrangement frequencies of CEB1 placed near ARS305 in both orientations in WT and mutant strains. Minisatellite
Genotype
Orientation I
Orientation II
CEB1-1.8
WT
1/154 (1%)
0/154 (0%)
pif1∆
57/108 (53%)
4/106 (4%)
rfa1-D228Y
50/98 (51%)
36/118 (31%)
pif1∆ rfa1-D228Y
43/101 (43%)
31/76 (41%)
WT GAL::PIF1
0/44 (0%)
0/43 (0%)
rfa1-D228Y GAL::PIF1
12/39 (31%)
2/43 (5%)
mms1∆
0/116 (0%)
3/116 (3%)
rtt105∆
116/116 (100%)
116/116 (100%)
WT GAL::RNH1
1/58 (2%)
0/58 (0%)
pif1∆ GAL::RNH1
30/58 (52%)
ND
rtt105∆ GAL::RNH1
58/58 (100%)
58/58 (100%)
rfa1-D228Y GAL::RNH1
39/87 (45%)
29/87 (33%)
CEB1-Gmut-1.7
WT
0/96 (0%)
ND
pif1∆
0/48 (0%)
ND
rfa1-D228Y
21/94 (22%)
ND
pif1∆ rfa1-D228Y
13/66 (20%)
ND
ND – not determined. TABLE 1. Rearrangement frequencies of CEB1 placed near ARS305 in both orientations in WT and mutant strains. RPA cooperate to remove G-quadruplex structures at both
leading and lagging strand (Figure 8). the instability of CEB1 in rfa1-D228Y while it totally abol-
ishes the instability of CEB1 in pif1∆ cells. Contrary to pif1∆
cells, in rfa1-D228Y cells the CEB1 instability does not com-
pletely rely on the G-quadruplex-forming sequence of CEB1,
indicating that the instability arising in this mutant partially
results from G-quadruplex-independent ssDNA-containing
secondary structures. We obtained similar results with the
double pif1∆ rfa1-D228Y mutant. Consistent with these
results, Pif1 overexpression suppresses the instability of
rfa1-D228Y mutant in the same proportion as G-mutated
CEB1 motif. DISCUSSION In contrast Rtt105 that promotes RPA nu-
clear import, and RPA-ssDNA complex formation at replica-
tion forks is required to stabilize CEB1 inserted in both ori-
entations. Collectively these results indicate that Pif1 and Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 56 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) estroni et al. (2020)
A critical role for RPA i
E 7: RPA is required to stabilize CEB25-L1T in fission yeast. (A) Map of the CEB25-L1T insertion within chromosome II at 14 kb
and at 11 kb from the ARS-II-1983. (B) Genomic DNA from cells containing CEB25-L1T in orientation 1 and 2 was digested by Pvu
d. Membranes were hybridized with CEB25 probe. (C) Genomic DNA from cells containing mutated CEB25-L1T-G12T in orientat
ed by PvuII and southern blotted. Membranes were hybridized with a CEB25-L1T probe. The number of colonies analysed per w
rearrangement frequencies, and the total number of colonies are indicated. Red stars mark unstable events. We calculated t
ility by monitoring the size and the intensity of the CEB25. We considered that CEB25 was unstable when the intensity of the sh
uperior to the one of the parental band or when the band disappears. The number of colonies analysed per well, the percentag
frequencies, and the total numbers of colonies are indicated in Table 1. FIGURE 7: RPA is required to stabilize CEB25-L1T in fission yeast. (A) Map of the CEB25-L1T insertion within chromosome II at 14 kb from the ARS-II-
1964 and at 11 kb from the ARS-II-1983. (B) Genomic DNA from cells containing CEB25-L1T in orientation 1 and 2 was digested by PvuII and southern
blotted. Membranes were hybridized with CEB25 probe. (C) Genomic DNA from cells containing mutated CEB25-L1T-G12T in orientation 1 and 2 was
digested by PvuII and southern blotted. Membranes were hybridized with a CEB25-L1T probe. The number of colonies analysed per well, the percent-
age of rearrangement frequencies, and the total number of colonies are indicated. Red stars mark unstable events. We calculated the frequency of
instability by monitoring the size and the intensity of the CEB25. We considered that CEB25 was unstable when the intensity of the short or high band
was superior to the one of the parental band or when the band disappears. DISCUSSION The number of colonies analysed per well, the percentage of rearrange-
ment frequencies, and the total numbers of colonies are indicated in Table 1. Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 57 L. Maestroni et al. (2020)
A critical role for RPA in CEB1 stab
FIGURE 8: Mechanistic model for the unfolding of CEB1-G-rich structures by RPA and Pif1. (A) In WT cells, when the G-rich strand (light grey)
leading strand (left part, CEB1 leading), stable G-quadruplexes structures are efficiently removed by Pif1, as proposed by Lopes and colleagues [33
the leading strand, RPA may cooperate with Pif1 either by preventing the refolding of the G4 structure or by recruiting Pif1 to the leading strand. W
the G-rich strand (light grey) is replicated by the lagging polymerase (right part, CEB1 lagging) binding of RPA to the G-rich strand prevents formati
stable G-rich secondary structures. In this context, Pif1 is dispensable. (B) In the rfa1-D228Y mutant, at the leading strand (left part), the reduced a
ty of RPA for G-rich ssDNA either reduce the ability of RPA to prevent the refolding of the G4 structure or decrease Pif1 recruitment. The decrea
RPA affects G4-unwinding and leads to the formation of CEB1 rearrangements. At the lagging strand (middle part), the decrease affinity of RPA f
rich ssDNA facilitates the formation of stable G-rich structures, affects their unwinding, and generates CEB1 instability. In such a case, overexpre
f Pif1 ( i ht
t)
ffi i
tl
i d G i h t
t
l
di
t CEB1 i
ti A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) L. Maestroni et al. (2020) (
) FIGURE 8: Mechanistic model for the unfolding of CEB1-G-rich structures by RPA and Pif1. (A) In WT cells, when the G-rich strand (light grey) is on
leading strand (left part, CEB1 leading), stable G-quadruplexes structures are efficiently removed by Pif1, as proposed by Lopes and colleagues [33]. At
the leading strand, RPA may cooperate with Pif1 either by preventing the refolding of the G4 structure or by recruiting Pif1 to the leading strand. When
the G-rich strand (light grey) is replicated by the lagging polymerase (right part, CEB1 lagging) binding of RPA to the G-rich strand prevents formation of
stable G-rich secondary structures. In this context, Pif1 is dispensable. DISCUSSION This may account for the even more prominent role of RPA
complex in G4 unwinding in fission yeast. This assumption
is also supported by the fact that the rpa1-D223Y mutation
has a stronger effect at telomeres in S. pombe than its
counterpart in S. cerevisiae [38,62]. Interestingly, we ob-
served expansions of the CEB25-L1T even in the WT strain,
especially when CEB25-L1T was preferentially replicated by
the lagging machinery. This result opens new avenues to
use fission yeast as a model organism to study expansions
of G-rich sequences that have been associated with neuro-
logical diseases. cessed by Pif1 indicating that RPA and Pif1 have redundant
G-quadruplex-processing activities at the lagging strand
and cooperate (Figure 8). Pif1 binds to G4 motifs with no preference for leading
and lagging strand templates [57] but Pif1 is essential only
for efficient replication through lagging strand G4s [60]. Consequently, the distinct behaviour of Pif1 at the leading
and the lagging strands may be due to different confor-
mations of the respective G-quadruplexes, affecting their
folding and/or their processing due to the occupancy rate
of RPA. Another possibility could be that the blocking
G-quadruplex structures are better tolerated and bypassed
by the lagging strand replication machinery due to its abil-
ity to prime DNA synthesis downstream of G-quadruplexes. However, this seems improbable because CEB1 instability
in rfa1-D228Y cells is fully suppressed by Pif1 overexpres-
sion suggesting that the blocking structures are present
and that the helicase activity of Pif1 is still required to re-
move G4-structures and rescue CEB1 stability. RPA interacts with RNase H1 and colocalizes with it at
R-loops raising the possibility that RPA could recruit and/or
stimulate RNase H1 [41,53]. We observed that the interac-
tion between RPA and RNase H1 is stimulated by the pres-
ence of DNA and is slightly affected in rfa1-D228Y cells. Furthermore, Pif1 regulates R-loop formation and poten-
tially complements RNase H for R-loop resolution [51, 52]. We found that RNase H1 overexpression did not rescue
CEB1 stability in pif1∆, rfa1-D228Y, and rtt105∆ mutants. These results suggest that R-loops are likely not responsi-
ble of CEB1 instability in these mutants, indicating that
CEB1 instability is not due to transcription. Collectively, our
results add new insights about the role of RPA as a general
sensor of secondary structures and regulator of genomic
stability. DISCUSSION (B) In the rfa1-D228Y mutant, at the leading strand (left part), the reduced affini-
ty of RPA for G-rich ssDNA either reduce the ability of RPA to prevent the refolding of the G4 structure or decrease Pif1 recruitment. The decrease in
RPA affects G4-unwinding and leads to the formation of CEB1 rearrangements. At the lagging strand (middle part), the decrease affinity of RPA for G-
rich ssDNA facilitates the formation of stable G-rich structures, affects their unwinding, and generates CEB1 instability. In such a case, overexpression
of Pif1 (right part) can efficiently unwind G-rich structures, leading to CEB1 size conservation. FIGURE 8: Mechanistic model for the unfolding of CEB1-G-rich structures by RPA and Pif1. (A) In WT cells, when the G-rich strand (light grey) is on
leading strand (left part, CEB1 leading), stable G-quadruplexes structures are efficiently removed by Pif1, as proposed by Lopes and colleagues [33]. At
the leading strand, RPA may cooperate with Pif1 either by preventing the refolding of the G4 structure or by recruiting Pif1 to the leading strand. When
the G-rich strand (light grey) is replicated by the lagging polymerase (right part, CEB1 lagging) binding of RPA to the G-rich strand prevents formation of
stable G-rich secondary structures. In this context, Pif1 is dispensable. (B) In the rfa1-D228Y mutant, at the leading strand (left part), the reduced affini-
ty of RPA for G-rich ssDNA either reduce the ability of RPA to prevent the refolding of the G4 structure or decrease Pif1 recruitment. The decrease in
RPA affects G4-unwinding and leads to the formation of CEB1 rearrangements. At the lagging strand (middle part), the decrease affinity of RPA for G-
rich ssDNA facilitates the formation of stable G-rich structures, affects their unwinding, and generates CEB1 instability. In such a case, overexpression
of Pif1 (right part) can efficiently unwind G-rich structures, leading to CEB1 size conservation. Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 58 A critical role for RPA in CEB1 stability L. Maestroni et al. (2020) the rpa1-D223Y mutant. The effects of the rpa1-D223Y
mutant were even more pronounced in S. pombe. This may
be due to the fact that fission yeast lacks a functional
ortholog of the Pif1 helicase. Indeed Pfh1 behaves more
closely to the budding yeast Rrm3 helicase than to Pif1 [56]. MATERIALS AND METHODS
Strains and growth conditions MATERIALS AND METHODS DISCUSSION RPA acts through direct interactions with proteins
acting at the level of specific secondary structures, which
are hotspots of genomic instability. This mode of RPA ac-
tion is conserved between yeasts and humans. How exactly Pif1 is recruited at the G-rich motifs and at
G-quadruplexes that form at CEB1 during replication? One
possibility is that Pif1 travels with the replication fork and
facilitates replication by processing G-quadruplex struc-
tures at the leading-CEB1. Indeed, Pif1 interacts with PCNA
and Cac1, the large CAF-1 subunit, which preferentially
assembles nucleosome onto replicating DNA [60, 61]. Thus,
Pif1 may be targeted to the replication fork by its ability to
interact with PCNA or the histone chaperone Cac1 and
preserves genome stability by acting at G-rich motifs at the
leading-CEB1. Alternatively, Pif1 could be directly recruited
to G-rich motifs and G-quadruplex structures. Mms1 is a
G4-DNA-binding protein that helps replication fork pro-
gression at G4 and Pif1 binding to specific G4 structures
[22, 37]. We show that Mms1 is not required to maintain
CEB1 stability at both leading- and lagging-CEB1 revealing
that G4 structures targeted by Mms1 are not deleterious
for CEB1 stability. Our results are in good agreement with
previous observations showing that Mms1 supports Pif1
function at G4 motifs only on the lagging strand [22, 37],
whereas Pif1 is not required for CEB1 stabilisation [33]. Here we show that RPA recruits Pif1 to CEB1. We report
that Pif1 associates with RPA and that this association is
affected in the rfa1-D228Y mutant and by DNA digestion,
suggesting that Pif1-RPA association relies on specific DNA
structures. In addition we show that rtt105∆ mutation,
which decreases the level of RPA associated with ssDNA (as
the rfa1-D228Y mutation) [44, 45], strongly affects CEB1
stability at both leading and lagging strands. These results
reveal the importance of Rtt105 in promoting the replica-
tion of G4-forming CEB1 minisatellite, during leading
strand and lagging strand synthesis, and confirm that the
level of RPA is crucial for CEB1 stability. RPA protects and
stabilizes ssDNA susceptible to secondary structure for-
mation. It is possible that the ability of RPA to directly bind
to Pif1 could be important to recruit Pif1 at specific G-rich
sequences/structures and/or to stimulate Pif1 activity at
G-rich motifs and G-quadruplex structures. OPEN ACCESS | www.cell-stress.com Strains and growth conditions LM361
LM411
diploïd CEB1-1.8 I rfa1-D228Y/RFA1 pif1::Kan/PIF1
diploïd CEB1-1.8 II rfa1-D228Y/RFA1 pif1::Kan/PIF1
This study
This study
LM349
diploïd CEB1-1.7 Gmut rfa1-D228Y/RFA1 pif1::Kan/PIF1
This study
LM396
diploïd CEB1-1.8 I rfa1-D228Y/RFA1 + pVS45 (GAL::PIF1)
This study
LM398
diploïd CEB1-1.8 II rfa1-D228Y/RFA1 + pVS45 (GAL::PIF1)
This study
LM401
diploïd CEB1-1.8 I rfa1-D228Y/RFA1 + pSH380 (empty vector)
This study
LM404
diploïd CEB1-1.8 II rfa1-D228Y/RFA1 + pSH380 (empty vector)
This study
YVC600
diploïd CEB1-1.8 I mms1::TRP1/MMS1
This study
YVC601
diploïd CEB1-1.8 II mms1::TRP1/MMS1
This study
YVC602
diploïd CEB1-1.8 I rtt105::Kan/RTT105
This study
YVC603
diploïd CEB1-1.8 II rtt105::Kan /RTT105
This study
YVC604
diploïd CEB1-1.8 I rtt105::Kan/RTT105 GAL::RNH1::natMX/RNH1
This study
YVC605
diploïd CEB1-1.8 II rtt105::Kan/RTT105 GAL::RNH1::natMX /RNH1
This study
YVC606
diploïd CEB1-1.8 I pif1::Kan/PIF1 GAL::RNH1::natMX /RNH1
This study
YVC607
diploïd CEB1-1.8 I rfa1-D228Y/RFA1 GAL::RNH1::natMX /RNH1
This study
YVC608
diploïd CEB1-1.8 II rfa1-D228Y/RFA1 GAL::RNH1::natMX /RNH1
This study
LM140
MATa rfa1-D228Y Pif1-myc::KanMX6
This study
W1042-7C
MATa can1-100,x SUP4-o::HIS3::pWJ317-CAN1-URA3 rfa1-D228Y
Rothstein R
LM301
pif1::KanMX6 rfa1-D228Y
This study
LM340
pif1::KanMX6
This study
YBL103
his3∆1; leu2∆0; ura3∆0; met15∆0; pGal-3HA-RNH1::NAT
Luke B. LM407
diploïd pGal-3HA-RNH1::natMX rfa1-D228Y/RFA1
This study
S. pombe strains
Genotype
Origin
JA961
h- leu1-32 ura4-D18 leu1+::pJK148-CEB25 orientation 1
This study
JA963
h- leu1-32 ura4-D18 leu1+::pJK148-CEB25-G12T orientation 1
This study
JA947
h- leu1-32 ura4-D18 leu1+::pJK148-CEB25 orientation 2
This study
JA948
h- leu1-32 ura4-D18 leu1+::pJK148-CEB25-G12T orientation 2
This study
SC387
h+ leu1-32 ura4-D18 ade6-M210 rad11-D223Y
Ueno M. All LM strains used are derivatives of W303-1B. All YVC strains used are derivatives of W303. Spore colonies were generated from the diploid strains. All LM strains used are derivatives of W303-1B. All YVC strains used are derivatives of W303. Spore colonies were generated from the diploid strains. gested DNA was resolved in 1% agarose gel and blotted onto
Hybond-XL membrane (GE Healthcare). After transfer, the
membrane was cross-linked with UV and hybridized with
CEB1-0.6 and CEB1 Gmut probes for CEB1-1.8 and CEB1Gmut -
1.7, respectively. 32P labelling of DNA probes was performed
by random priming using Klenow fragment exonuclease (New
England Biolabs), in presence of [α-32P]-CTP and hybridizations
were performed in Church buffer at 55°C. Radioactive signals
were detected using a BIORAD molecular imager FX. Correct and unique insertion of CEB25 at this locus generated
two fragments of 3 and 14.4 kb. To check the correct size of
CEB25-L1T and CEB25-L1T-G12T, a second digestion by PvuII
on genomic DNA was performed on selected clones. Strains and growth conditions Strains used in this study are listed in Table 2. Yeast strains
containing CEB1-1.8 I (orientation I), CEB1-1.8 II (orientation II),
or CEB1Gmut-1.7 were mated with pif1∆, rfa1-D228Y, pif1∆
rfa1-D228Y, mms1∆, or rtt105∆ mutants. After sporulation
and identification of the four resulting spores, the strains of
interest were plated on rich medium at 30°C to obtain isolated
colonies. Individual clones were grown in rich liquid culture at
30°C until stationary phase. For PIF1 overexpression experi-
ments, WT cells and rfa1-D228Y cells were transformed with
the centromeric plasmids pVS45 (expressing the nuclear form
of Pif1 under the control of the GAL1 promoter) and pSH380
(a pRS315-derived vector control) provided by Virginia Zakian
[43], then mated with leading-CEB1 (orientation I), or lagging-
CEB1 (orientation II) cells. Individual clones were grown in SD-
Leu (2% glucose), or SGal-Leu (2% galactose) media for repres-
sion or overexpression of PIF1, when mentioned. The 0.62 kb CEB25-L1T and CEB25-L1T-G12T were cloned
into pJK148 integrative plasmid in both orientations at Not1
site, from plasmid pPA84 and pPA84-G12T (A. Nicolas’s Labor-
atory), respectively. The corresponding plasmids were linear-
ized by Nde1 and transformed into S. pombe cells. To check
the insertion of CEB25 at the leu1 locus, genomic DNA was
digested by XbaI a nd southern blotted using a leu1 probe. Importantly, the results obtained in S. cerevisiae with
the rfa1-D228Y mutant were confirmed in S. pombe with Cell Stress | MARCH 2020 | Vol. 4 No. 3 OPEN ACCESS | www.cell-stress.com 59 L. Maestroni et al. (2020) A critical role for RPA in CEB1 stability Table2. Strains used in this study. S. cerevisiae strains
Genotype
Origin
ORT6119-4
MATa CEB1-1.8 I-ARS305
Nicolas A. ORT6135-36
MATa CEB1-1.8 II-ARS305
Nicolas A. ORT6157-1
MATa CEB1-1.7 Gmut-ARS305
Nicolas A. Co-Immunoprecipitation experiment Yeast cells were grown at 30°C in YPD to OD600=0.8. Extracts
were lysed with glass-beads in TMG-50 (10mM TrisHCl pH8.0,
1mM MnCl2, 10% (v/v) glycerol, 50 mM NaCl, 0.1 mM DTT)
containing a protease inhibitor cocktail (Calbiochem), MG132
(Sigma Aldrich), and 0.5% (v/v) Tween-20. Pif1-Myc and Rnh1-
HA proteins were immunoprecipitated with 9E10 monoclonal
anti-Myc antibody (Santa Cruz, CA) and 12CA5 anti-HA mono-
clonal antibody (Roche), respectively. The presence of Rfa1 Strains and growth conditions Southern
blot hybridized by a CEB25-L1T probe (Not1-fragment from
pPA84 plasmid), revealed a fragment of 1.1 kb. Fission yeast
strains containing CEB25-L1T and CEB25-L1T-G12T were then
mated with the rpa1-D223Y mutant. After sporulation and
identification of the resulting spores, the stability of the CEB25
was controlled a second time by Southern blot after digestion
of genomic DNA by PvuII. The strains of interest were plated
on rich medium to obtain isolated colonies and individual
clones were further grown in rich liquid culture at 32°C until
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h
h
d
l
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Apgar Score Is Related to Development of Atopic Dermatitis: Cotwin Control Study
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Hindawi Publishing Corporation
Journal of Allergy
Volume 2013, Article ID 712090, 6 pages
http://dx.doi.org/10.1155/2013/712090 Hindawi Publishing Corporation
Journal of Allergy
Volume 2013, Article ID 712090, 6 pages
http://dx.doi.org/10.1155/2013/712090 Hindawi Publishing Corporation
Journal of Allergy
Volume 2013, Article ID 712090, 6 pages
http://dx.doi.org/10.1155/2013/712090 Vibeke Naeser,1 Niklas Kahr,1 Lone Graff Stensballe,2 Kirsten Ohm Kyvik,3
Axel Skytthe,4 Vibeke Backer,1 Charlotte Giwercman Carson,5
and Simon Francis Thomsen6 1 Department of Respiratory Medicine, Bispebjerg Hospital, 2400 Copenhagen, Denmark
2 Danish Epidemiology Science Centre, Statens Serum Institut, 2300 Copenhagen, Denmark
3 Institute of Regional Health Services Research & Odense Patient Data Explorative Network,
University of Southern Denmark, 5000 Odense, Denmarkh y f
4The Danish Twin Registry, University of Southern Denmark, 5000 Odense, Denmark
5t The Danish Twin Registry, University of Southern Denmark, 5000 Odense, Denm
5 Danish Pediatric Asthma Center, Gentofte Hospital, 2900 Hellerup, Denmarkt h
5 Danish Pediatric Asthma Center, Gentofte Hospital, 2900 Hellerup, Denmarkt ft
6Department of Dermato-Allergology, Gentofte Hospital, 2900 Hellerup, Denmark Correspondence should be addressed to Simon Francis Thomsen; sft@city.dk Received 27 May 2013; Revised 20 September 2013; Accepted 21 September 2013 Received 27 May 2013; Revised 20 September 2013; Accepted 21 September 2013 Academic Editor: S. L. Johnston Copyright © 2013 Vibeke Naeser et al. This is an open access article distributed under the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Aim. To study the impact of birth characteristics on the risk of atopic dermatitis in a twin population. Methods. In a population-
based questionnaire study of 10,809 twins, 3–9 years of age, from the Danish Twin Registry, we identified 907 twin pairs discordant
for parent-reported atopic dermatitis. We cross-linked with data from the Danish National Birth Registry and performed cotwin
control analysis in order to test the impact of birth characteristics on the risk of atopic dermatitis. Results. Apgar score, OR (per
unit) = 1.23 (1.06–1.44), 𝑃= 0.008, and female sex, OR = 1.31 (1.06–1.61), 𝑃= 0.012, were risk factors for atopic dermatitis in
cotwin control analysis, whereas birth anthropometric factors were not significantly related to disease development. Risk estimates
in monozygotic and dizygotic twins were not significantly different for the identified risk factors. Conclusions. In this population-
based cotwin control study, high Apgar score was a risk factor for atopic dermatitis. This novel finding must be confirmed in
subsequent studies. Research Article
Apgar Score Is Related to Development of
Atopic Dermatitis: Cotwin Control Study Vibeke Naeser,1 Niklas Kahr,1 Lone Graff Stensballe,2 Kirsten Ohm Kyvik,3
Axel Skytthe,4 Vibeke Backer,1 Charlotte Giwercman Carson,5
and Simon Francis Thomsen6 1. Introduction No statistically
significant association was demonstrated between any of
the birth-related exposures and the four allergic conditions
studied, although some nonsignificant trends were noted for
those born by Caesarean section and those exposed to fetal
distress during labor.h Next, the risk factors that were unshared between the twins
within the pair were examined in a 1 : 1 matched conditional
logistic regression analysis (cotwin control analysis) in order
to determine the effect of these risk factors on atopic dermati-
tis. The factors entered into this analysis were birth weight,
birth length, Apgar score, and sex. The matching was done
with the affected (atopic dermatitis) twin in each pair being
the case and the unaffected twin being the control. DZ same-
sex and DZ opposite-sex twin pairs were considered together
in the analysis. The matching of the twins in the cotwin
control design allows an inherent adjustment for other factors
that would otherwise confound the association between
birth factors and atopic dermatitis such as genetic factors,
atopic predisposition, maternal smoking during pregnancy,
gestational age, mode of delivery, and current age, as well as
effects of the rearing environment. Therefore the estimated
risk of atopic dermatitis can be assumed to reflect more
precisely the influence of birth factors, for example, Apgar
score. The following predictions can be made for the cotwin
control analysis (Figure 1): if the association between birth
factors and atopic dermatitis is mediated via genetics, then we
do not expect to find any significant risk of atopic dermatitis
among discordant MZ twin pairs, because MZ twins are
concordant for early rearing environmental and genetic
factors. Alternatively, an increased risk of atopic dermatitis in
discordant DZ twin pairs would suggest that genetic factors
underlie the association, because DZ twins share the same
rearing environment, but only half their genetic variants. Finally, a direct (causal) effect of birth factors, for example,
Apgar score, on atopic dermatitis would be suggested by the
finding that both discordant MZ and DZ twin pairs are also
discordant on, for example, Apgar score, as the association
can neither be reflected by environmental nor genetic factors
[19]. Data were analyzed with the statistical package SPSS 18.0
(SPSS Inc., Chicago, IL). g
The cotwin control study design is a matched case-control
study that utilizes twin pairs discordant for an exposure or
an outcome. 2. Methods The study population comprised all live-born twins in Den-
mark between 1994 and 2000 included in the Danish Twin
Registry. In 2003 when the twins were 3–9 years of age
the twins’ parents responded to a two-page questionnaire
concerning atopic diseases, diabetes, febrile and epileptic
seizures, and mental health attributes. Atopic dermatitis was
diagnosed by the question “has your child ever had eczema in
the folds of the elbows or knees?” Zygosity was determined by
information regarding the twins’ similarity to family, friends,
teachers, and relations. This method determines zygosity cor-
rectly in more than 95% of the cases compared with genetic
marker information [18]. We cross-linked the information
from the questionnaire with data from the Danish National
Birth Registry where data on birth weight, birth length,
maternal smoking during pregnancy, gestational age, mode
of delivery (Caesarean section versus vaginal delivery), and
Apgar score were obtained. 1. Introduction Particularly, because twin pairs are inherently
matched on several confounding factors relating to early life
and upbringing, this type of study design is ideal to tease out
causative relationships in complex diseases. In the present
study we use the cotwin control design to identify birth-
associated risk factors for atopic dermatitis in twin pairs
discordant for the disease. 1. Introduction of atopic dermatitis has increased both in developed [4, 5]
and developing countries [6] in recent years, but the causes
for this increase are still poorly understood.h Atopic dermatitis is a chronic relapsing skin disease with
a lifetime prevalence of around 20%. The disease has an
early onset with 60% being affected during the first year of
life and 85% before the age of five [1]. Atopic dermatitis is
characterized by recurrent episodes of itching and eczema
occurring at typical sites such as the face and extensor aspects
of the arms and legs in early childhood, flexures in later
childhood, and head and neck as well as the hands in adult-
hood. Atopic dermatitis is highly heritable and individual
susceptibility to the disease is attributable particularly to
mutations in the filaggrin gene [2], although other genetic
variants [3] as well as environmental factors have been
implicated. Previous studies have shown that the prevalence The onset of atopic dermatitis early in life implies that
risk factors for the disease exert their effect already in utero
or in very early childhood. However, previous studies of
the association between perinatal factors and the risk of
atopic dermatitis are conflicting. With regard to alcohol
intake during pregnancy, results indicate an increased risk
for atopic dermatitis in children of mothers who consume
alcohol during pregnancy [7, 8]. Contrary to this, smoking
during pregnancy has been inversely related to development
of atopic dermatitis in the offspring [9, 10]. The associa-
tion between exclusive breastfeeding/increased duration of
breastfeeding and atopic dermatitis varies; while some studies 2 2 Journal of Allergy Journal of Allergy show a positive association [11] others show a negative [12]
or no [13] association. Some studies [9, 14] but not all [15]
have found that high birth weight increases the risk of atopic
dermatitis, whereas other studies have found that a higher
gestational age increases the risk of atopic dermatitis [16]. Mallen et al. investigated the association between six birth-
related exposures (birth weight, mode of delivery, artificial
commencement of labor, prematurity, neonatal intensive care
unit admission, and fetal distress) and asthma, allergic rhini-
tis, eczema, and hay fever in adulthood [17]. 3. Results The proportions of girls and boys in the population were
48.9% and 51.1%, respectively. The mean age was 5.9 years
with no difference between girls and boys. The overall
prevalence of atopic dermatitis was 14.9%, 14.0% in boys and
16.0% in girls, 𝑃= 0.003. Increasing birth length (𝑃= 0.038),
nonsmoking of the mother during pregnancy (𝑃= 0.002),
and monozygosity (0.012) were also significant predictors for
atopic dermatitis in the whole sample when ignoring the
familial relationship between the twins (Table 1). A total of 10,809 twin subjects participated in the ques-
tionnaire study (participation rate 68%). Of these, complete
information on atopic dermatitis was available for 5399 intact
twin pairs: 850 monozygotic (MZ) twin pairs, 2,279 dizygotic
(DZ) twin pairs of the same sex, 1,999 DZ twin pairs of
opposite sex, and 271 twin pairs of unknown zygosity. There
were 907 twin pairs discordant for atopic dermatitis, and of
these 58 and 827 were monozygotic (MZ) and dizygotic (DZ)
pairs, respectively. Within twin pairs discordant for atopic dermatitis, the
affected twin had, on average, a higher Apgar score and was
smaller at birth compared with the unaffected twin (Table 2). In multiple conditional logistic regression comparing the
affected with the unaffected twin, Apgar score, OR (per unit)
= 1.23 (1.06–1.44), 𝑃= 0.008, and female sex, OR = 1.31 (1.06–
1.61), 𝑃= 0.012, were significantly predisposing factors for
atopic dermatitis, whereas birth anthropometric factors were
not significantly related to development of atopic dermatitis. Risk estimates in discordant MZ and DZ twins were not
significantly different (Table 3). 2.1. Statistical Analysis. First, the relationship between birth-
associated risk factors and atopic dermatitis was examined in
the whole population using unpaired 𝑡-tests and chi-square
tests, ignoring the familial relationship between the twins. 2.1. Statistical Analysis. First, the relationship between birth-
associated risk factors and atopic dermatitis was examined in
the whole population using unpaired 𝑡-tests and chi-square
tests, ignoring the familial relationship between the twins. 3 Journal of Allergy 3 Table 1: Characteristics of subjects with and without atopic dermatitis in a sample of Danish twin pairs, 3–9 years of age. Table 1: Characteristics of subjects with and without atopic dermatitis in a sample of Danish twin pairs, 3–9 years of age. 3. Results Atopic dermatitis
OR (95% CI)
𝑃value
Yes
𝑁= 1614
No
𝑁= 9195
Birth weight, g
2593
2571
22 (−9–52)
0.167
Birth length, cm
48.24
48.05
0.19 (0.01–0.38)
0.038
Apgar score
9.76
9.73
0.03 (−0.02–0.09)
0.172
Gestational age, days
254.9
254.6
0.32 (−0.61–1.26)
0.496
Caesarean section
14.4
15.3
0.93 (0.84–1.04)
0.220
Maternal smoking
12.9
15.5
0.80 (0.70–0.92)
0.002
Sex
Boys
14.0
86.0
1.00
Girls
16.0
84.0
1.17 (1.05–1.30)
0.003
Age, years
5.92
5.86
0.06 (−0.05–0.17)
0.257
Zygosity
DZ
14.4
85.6
1.00
MZ
16.8
83.2
1.20 (1.04–1.38)
0.012
Means and proportions are calculated from available data for the individual variables. OR is mean difference for birth weight, birth length, Apgar score, gestational age, and age. ristics of subjects with and without atopic dermatitis in a sample of Danish twin pairs, 3–9 years of age. Table 2: Distribution of perinatal risk factors for atopic dermatitis in 907 twin pairs, 3–9 years of age, discordant for atopic dermatitis. All twin pairs (𝑛= 907)
MZ twin pairs (𝑛= 58)
DZ twin pairs (𝑛= 827)
Affected twins
Unaffected twins
Affected twins
Unaffected twins
Affected twins
Unaffected twins
Birth weight, g
2627
2649
2624
2625
2632
2650
Birth length, cm
48.37
48.41
48.60
48.17
48.40
48.44
Apgar score
9.81
9.68
9.80
9.75
9.80
9.67
Female sex
52.3
47.4
53.4
53.4
53.1
47.8
Genetic factors
Environmental factors
Apgar score
Atopic dermatitis
Genetic factors
Environmental factors
Apgar score
Atopic dermatitis
Genetic factors
Environmental factors
Apgar score
Atopic dermatitis
Larger effect size in DZ twins than in MZ twins
discordant for atopic dermatitis
Larger effect size in MZ twins than in DZ twins
discordant for atopic dermatitis
Equal effect sizes in MZ and DZ twins discordant
for atopic dermatitis
(3) Causal relationship
(2) Environmental confounding
(1) Genetic confounding
Figure 1: Possible association between Apgar score and atopic dermatitis predicted by the cotwin control method. al risk factors for atopic dermatitis in 907 twin pairs, 3–9 years of age, discordant for atopic dermatitis. Table 2: Distribution of perinatal risk factors for atopic dermatitis in 907 twin pairs, 3–9 years of age, discordant for atopic dermatitis. 4. Discussion However, previous
studies have found a harmful effect on atopic dermatitis
of other birth-related factors linked to high Apgar score
such as increased fetal growth [9], high socioeconomic
status (SES) [26, 27], and nonsmoking of the mother [9,
10]. Particularly, Lundholm et al. examined the association
between fetal growth and atopic diseases and found that the
rate of atopic dermatitis increased with higher levels of the
mother’s education and furthermore that children of mothers
who did not smoke during pregnancy also had slightly higher
rates of atopic dermatitis compared with children of mothers
who smoked during pregnancy [9]. Furthermore, a Dutch
study showed that smoking during pregnancy resulted in
decreased birth weight and that women of low SES give
birth to lighter babies [28]. These findings are inline with
the assumption that atopic dermatitis is a disease affecting
apparently robust children from high SES families. Because
the affected (atopic) and the unaffected twin were matched on
exposure status in our paired analysis, the observed positive
association between atopic dermatitis and Apgar score is
adjusted for confounding due to SES, parental smoking, and
gestational age. Taken together, multiple factors are involved
in the development of atopic dermatitis, particularly, genetics,
fetal programming, and environmental effects, which makes We found an overall higher prevalence of atopic dermati-
tis among the entire group of MZ twins than among the group
of DZ twins. This finding could be due to confounding from
factors in the MZ group not present in the DZ group, for
example, a larger susceptibility gene pool among MZ twins
or a more unfavorable intrauterine environment in MZ than
in DZ twins. Also, these could be factors such as differences
in the MZ and DZ families in terms of smoking and SES, but
evidence in support of this theory is lacking. The large population size with identification of 907 twin
pairs discordant for atopic dermatitis and the high response
rate are strengths of this study, although the number of
discordant MZ pairs was quite small (58 in total). We had
complete ascertainment of all live-born twins in Denmark
between 1993 and 2000 and cross-linked these data to data
from the Danish Birth Registry combining information from
two independent sources. Data on perinatal risk factors were
obtained by professional midwifes’ records, which increased
the validity of the data. 4. Discussion it difficult to deduce which risk factors represent direct effects
and which represent mediatory effects. The equal effect sizes
in MZ and DZ twins for Apgar score in cotwin control
analysis suggest a direct effect of one or more of the factors
attributable to the Apgar score in relation to atopic dermatitis. In this nationwide cohort of Danish twin pairs discordant
for atopic dermatitis, a high Apgar score and female sex
were significant determinants of atopic dermatitis, whereas
birth weight and birth length were not significantly related to
disease development. The prevalence of atopic dermatitis was higher in girls
than in boys in the present study. Other studies contradict
this, concluding that boys have a higher prevalence of atopic
dermatitis and yet other studies find no significant difference
between the prevalence of atopic dermatitis in boys and girls. Notably, a study from Taiwan found that the prevalence of
atopic dermatitis in females is lower than in males before the
age of 8 years but subsequently becomes higher [29], whereas
studies from Japan and Italy show no significant difference
in the prevalence of atopic dermatitis in boys and girls [30,
31]. A recent study of another (comparable) Scandinavian
population from Sweden found a female preponderance of
atopic dermatitis (14.5% in girls versus 9.4% in boys) [32]. p
Apgar score is a clinical test performed on a newborn
one and five minutes after birth. It is a composite measure
of breathing effort, heart rate, muscle tone, reflexes, and
skin colour and is an indicator of the newborn’s need for
medical attention shortly after birth. A detrimental effect
of high Apgar score on the risk of atopic dermatitis is, to
our knowledge, a novel finding that needs to be confirmed
in subsequent studies. Among Finnish children low Apgar
score increased the risk of asthma [20, 21]. Contrary to this,
Apgar score was not related to asthma [22] or hay fever
[23] in Finnish twin children and did not interact with in
vitro fertilization on the risk of asthma in children born after
assisted reproduction [24]. Moreover, Gilman et al. found
no significant association between Apgar score and maternal
smoking [25]. It is somewhat counterintuitive that it is the
twin with the highest Apgar score that has the highest risk
of later development of atopic dermatitis. 3. Results Genetic factors
Environmental factors
Apgar score
Atopic dermatitis
Genetic factors
Environmental factors
Apgar score
Atopic dermatitis
Genetic factors
Environmental factors
Apgar score
Atopic dermatitis
Larger effect size in DZ twins than in MZ twins
discordant for atopic dermatitis
Larger effect size in MZ twins than in DZ twins
discordant for atopic dermatitis
Equal effect sizes in MZ and DZ twins discordant
for atopic dermatitis
(3) Causal relationship
(2) Environmental confounding
(1) Genetic confounding
Figure 1: Possible association between Apgar score and atopic dermatitis predicted by the cotwin control method. Environmental factors Atopic dermatitis Environmental factors Equal effect sizes in MZ and DZ twins discordant
for atopic dermatitis
(3) Causal relationship Atopic dermatitis Figure 1: Possible association between Apgar score and atopic dermatitis predicted by the cotwin control method. Journal of Allergy 4 Table 3: Significant risk factors for atopic dermatitis in 907 twin pairs, 3–9 years of age, discordant for atopic dermatitis. All twin pairs
MZ twin pairs
DZ twin pairs
ORMZ = ORDZ
OR (95% CI)
P value
OR (95% CI)
P value
OR (95% CI)
P value
P value1
Birth weight
1.02 (0.81–1.28)
0.884
0.96 (0.12–7.86)
0.972
0.98 (0.77–1.24)
0.880
NS
Birth length
1.00 (0.08–13.03)
1.00
8.31 (N/A)
0.854
1.62 (0.11–22.01)
0.758
NS
Apgar score
1.23 (1.06–1.44)
0.008
1.19 (0.60–2.34)
0.615
1.21 (1.04–1.42)
0.015
NS
Female sex
1.31 (1.06–1.61)
0.012
1.31 (1.07–1.61)
0.011
NS
Only statistically significant predictors for atopic dermatitis are presented in the table (adjusted for birth weight, birth length). 1Test for heterogeneity between odds ratios for MZ and DZ twin pairs. NS: nonsignificant. ors for atopic dermatitis in 907 twin pairs, 3–9 years of age, discordant for atopic dermatitis. References [17] C. D. Mallen, S. Mottram, G. Wynne-Jones, and E. Thomas,
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The Lancet, vol. 368, no. Journal of Allergy Journal of Allergy 5 The fact that we studied twins may have had implications
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environment, and mode of delivery, could also be expected to
influence the variation in Apgar score. However, in a matched
case-control study of 112 infant twins and singletons from the
United States 5-minute Apgar score was not different in twins
compared with singletons when matched for gestational
age and mode of delivery [33], suggesting that results in
relation to Apgar score can be extrapolated to the general
singleton population. Still, we acknowledge that there may be
unmeasured factors in our sample that could have interfered
with our conclusions. [10] A. Linneberg, J. B. Simonsen, J. Petersen, L. G. Stensballe, and
C. S. Benn, “Differential effects of risk factors on infant wheeze
and atopic dermatitis emphasize a different etiology,” Journal
of Allergy and Clinical Immunology, vol. 117, no. 1, pp. 184–189,
2006. [11] D. J. Purvis, J. M. D. Thompson, P. M. Clark et al., “Risk factors
for atopic dermatitis in New Zealand children at 3.5 year of age,”
British Journal of Dermatology, vol. 152, no. 4, pp. 742–749, 2005. [12] C. S. Benn, J. Wohlfahrt, P. Aaby et al., “Breastfeeding and risk
of atopic dermatitis, by parental history of allergy, during the
first 18 months of life,” American Journal of Epidemiology, vol. 160, no. 3, pp. 217–223, 2004. [13] J. F. Ludvigsson, M. Mostrom, J. Ludvigsson, and K. Duchen,
“Exclusive breastfeeding and risk of atopic dermatitis in some
8300 infants,” Pediatric Allergy and Immunology, vol. 16, no. 3,
pp. 201–208, 2005. [14] P. Leadbitter, N. Pearce, S. Cheng et al., “Relationship between
fetal growth and the development of asthma and atopy in
childhood,” Thorax, vol. 54, no. 10, pp. 905–910, 1999. We conclude that a high Apgar score may be linked
to development of atopic dermatitis. Although this finding
agrees with the theory that robust babies of high SES are at
increased risk of atopic dermatitis, this has not been previ-
ously documented and awaits support from future studies. [15] A. Sevelsted and H. Journal of Allergy Bisgaard, “Neonatal size in term children is
associated with asthma at age 7, but not with atopic dermatitis or
allergic sensitization,” Allergy, vol. 67, no. 5, pp. 670–675, 2012. [16] M. M. Moore, S. L. Rifas-Shiman, J. W. Rich-Edwards et al.,
“Perinatal predictors of atopic dermatitis occurring in the first
six months of life,” Pediatrics, vol. 113, no. 3, pp. 468–474, 2004.h 4. Discussion Contrary to this, the diagnosis of
atopic dermatitis was self-reported by the parents of the
twins, which could have induced recall and reporting bias. Moreover, the question used to diagnose atopic dermatitis
has not been systematically validated. Another limitation
alludes to the differing age at ascertainment of the eczema
diagnosis, which may have influenced the results. However,
age was automatically eliminated as possible confounder in
the cotwin control analysis model since it was the same for
the two twins within the pair. References 9537, pp. 733–743, 2006. [23] M. R¨as¨anen, J. Kaprio, T. Laitinen, T. Winter, M. Koskenvuo,
and L. A. Laitinen, “Perinatal risk factors for hay fever—a study
among 2550 Finnish twin families,” Twin Research, vol. 4, no. 5,
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Olausson, “Asthma in Swedish children conceived by in vitro
fertilisation,” Archives of Disease in Childhood, 2012. h
[7] C. G. Carson, L. B. Halkjaer, S. M. Jensen, and H. Bisgaard,
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dermatitis. The COPSAC prospective birth cohort study of a
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during pregnancy and children’s cognitive and physical devel-
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during pregnancy and atopic dermatitis in the offspring,”
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factors of atopic diseases in German children and adolescents,”
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dermatitis, allergic rhinitis and asthma in Taiwan: a national
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matitis in Japanese elementary schoolchildren,” British Journal
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matitis in Italian schoolchildren: factors affecting its variation,”
Acta Dermato-Venereologica, vol. 89, pp. 122–125, 2009. References [32] N. Ballardini, I. Kull, C. S¨oderh¨all, G. Lilja, M. Wickman, and C. F. Wahlgren, “Eczema severity in preadolescent children and its
relation to sex, filaggrin mutations, asthma, rhinitis, aggravating
factors and topical treatment: a report from the BAMSE birth
cohort,” British Journal of Dermatology, vol. 168, pp. 588–594,
2013. [33] S. A. Friedman, E. Schiff, L. Kao et al., “Do twins mature
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pp. 1193–1199, 1997.
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Reverse Wedge Effect Following Intramedullary Nailing of a Basicervical Trochanteric Fracture Variant Combined with a Mechanically Compromised Greater Trochanter
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Reverse Wedge Effect Following Intramedullary
Nailing of a Basicervical Trochanteric Fracture
Variant Combined with a Mechanically
Compromised Greater Trochanter
Yu Zhang
Jiangsu Province Hospital and Nanjing Medical University First Affiliated Hospital
https://orcid.org/0000-0003-2274-429X
Jun Hu
Jiangsu Province Hospital
Xiaodong Qin
(
qinxiaodongnjmu@163.com
)
Xiang Li
Jiangsu Province Hospital
Guqi Hong
Jiangsu Province Hospital
Research article
Keywords: trochanteric fracture, intramedullary nailing, intraoperative complication
Posted Date: January 20th, 2020
DOI: https://doi.org/10.21203/rs.2.14211/v2
License:
This work is licensed under a Creative Commons Attribution 4.0 International
License.
Read Full License
Version of Record: A version of this preprint was published at BMC Musculoskeletal Disorders on March
28th, 2020. See the published version at https://doi.org/10.1186/s12891-020-03212-6. Reverse Wedge Effect Following Intramedullary
Nailing of a Basicervical Trochanteric Fracture
Variant Combined with a Mechanically
Compromised Greater Trochanter
Yu Zhang
Jiangsu Province Hospital and Nanjing Medical University First Affiliated Hospital
https://orcid.org/0000-0003-2274-429X
Jun Hu
Jiangsu Province Hospital
Xiaodong Qin
(
qinxiaodongnjmu@163.com
)
Xiang Li
Jiangsu Province Hospital
Guqi Hong
Jiangsu Province Hospital
Research article
Keywords: trochanteric fracture, intramedullary nailing, intraoperative complication
Posted Date: January 20th, 2020
DOI: https://doi.org/10.21203/rs.2.14211/v2
License:
This work is licensed under a Creative Commons Attribution 4.0 International
License. Read Full License
Version of Record: A version of this preprint was published at BMC Musculoskeletal Disorders on March
28th, 2020. See the published version at https://doi.org/10.1186/s12891-020-03212-6. Research article License:
This work is licensed under a Creative Commons Attribution 4.0 International
License. License:
This work is licensed under a Creative Commons Attribution 4.0 International
License. Read Full License Version of Record: A version of this preprint was published at BMC Musculoskeletal Disorders on March
28th, 2020. See the published version at https://doi.org/10.1186/s12891-020-03212-6. Page 1/18 Page 1/18 Page 1/18 Abstract Background: To introduce an unreported intraoperative complication in intramedullary nailing (IN) of an
anatomically reduced trochanteric fracture variant characterized by a basicervical fracture line and
coronally disrupted greater trochanter (GT). Materials and Methods: A total of 414 trochanteric fractures (TF) treated with intramedullary nails from
2013 to 2017 were included in this study. After analysis of intraoperative fluoroscopy data, 33 cases,
including 21 females and 12 males, with a mean age of 72.5 years (33 to 96 years) were identified for
internal rotation of the cephalocervical fragment and inferior opening at the basicervical fracture line
caused by nailing a satisfactorily reduced TF. The morphological features of this group of patients were
analyzed on computed tomography (CT) scan. On radiograph, the magnitude of the displacement and
final femoral neck-shaft angle (NSA) were measured. Results: CT analysis demonstrated that the basicervical fracture line and the posterolateral fragment
(PLF) detached from the GT were the two dominant features of this cohort. They were classified
according to the number of main fragments: a 3-fragmentary subgroup containing three consistent
fragments (cephalocervical fragment, PLF and distal femoral shaft) and a 4-fragmentary subgroup
embracing one additional fragment (lesser trochanter). The four subtypes were as follows: the 3-
fragmentary S indicating a small PLF (6 cases), the 3-fragmentary M presenting a moderate PLF (3
cases), the 3-fragmentary L standing for the PLF involving whole lesser trochanter (LT) (4 cases) and the
4-fragmentary GL incorporating separated PLF and LT fragments (20 cases). Geological analysis
demonstrated that the majority of the basicervical fracture lines (81.8%) just crossed the center of the
piriformis fossa, while the others marginally involved the medial wall of the GT. Postoperatively, the mean
width of the inferior opening at the basicervical region was 9.2±4.6 mm. The mean NSA was 135.2±7.8
degrees. The comparison between the 3- and 4-fragmentary subgroups revealed no significant
differences in magnitude of displacement and NSA. Conclusion: This unreported intraoperative complication predominantly occurred in the intramedullary
nailed basicervical trochanteric fracture variant combined with a PLF from the GT. The magnitude of the
secondary displacement was substantial and resulted in a relative valgus reduction. This secondary
displacement was caused by an impingement of the reamer with the superolateral cortex of the
cephalocervical fragment and should be addressed during the operation. Level of Evidence: Therapy IV Level of Evidence: Therapy IV Introduction The increase in the aging population worldwide has resulted in a growing amount of TF among geriatric
people, which presents challenges to the public health system and to orthopedic trauma surgeons [1]. Surgical intervention has been the preferred treatment for TF patients because of the benefits of effective
pain control and the opportunity for early weight bearing. Compared to dynamic hip screw plates, INs are Page 2/18 gaining more popularity and could provide superior stability to unstable TF patterns that are
characterized by posteromedial comminution, reverse oblique configuration, lateral wall disruption, GT
disruption and basicervical variants [2, 3]. At the same time, much attention has been paid to the increasing postoperative complications associated
with the IN application. Those complications include sliding screw cut-out or cut-through [4], extensive
fracture collapse and reduction loss, thigh pain associated with nail tip collision, and varus malunion. However, the intraoperative complications are less noticeable. Recently, Hak reported an uncommon
intraoperative complication when IN was performed to stabilize a group of TFs: a varus malreduction
presented as a secondary fracture displacement caused by inserting a cephalated IN from the tip of the
GT [5]. It was featured by a lateral displacement of the femoral shaft and an opening of the superior part
of the primary fracture line (trochanteric region), which was later named the “wedge effect” by O’Malley [6]
(Figure 1). Interestingly, we also noticed another pattern of intraoperative complications closely associated with
inserting a trochanter tip starting point IN in a cohort of TFs (Figure 2). In contrast to the aforementioned
“wedge effect”, the reaming/IN insertion generated internal rotation of the cephalocervical fragment and
an inferiorly oriented gap at the primary fracture line (basicervical region). This pattern was named the
“reverse wedge effect”. Further studies revealed that this intraoperative complication completely occurred
in an uncommon TF pattern distinguished by the primary fracture line at the basicervical region
(basicervical TF variant) and a detached PLF from the GT. The purpose of the present study was to
investigate the incidence of the “reverse wedge effect” at a level-one tertiary trauma center, summarize its
morphological features and analyze its cause and potential influence on treatment outcomes. Materials And Methods We obtained Institutional Review Board approval for this study. A total of 414 TF cases that underwent IN
fixation at our hospital between January 2013 and January 2017 were included in this study. The
inclusion criteria were skeletally mature, unilateral TF with complete radiographs, intraoperative
fluoroscopy and preoperative CT scan which improved our understanding of the fracture characteristics
and the degree of instability. The exclusion criteria were as follows: (1) patients younger than 18 years;
(2) high-energy injury or multiple injuries such as car accidents or high-altitude fall injuries; and (3)
patients with pathological fractures and previous histories of hip malformation or surgery of either hip. By observing intraoperative fluoroscopy, the TF cases demonstrating the “reverse wedge effect” during
reaming or nail insertion were identified. We obtained Institutional Review Board approval for this study. A total
fixation at our hospital between January 2013 and January 2017 were
inclusion criteria were skeletally mature, unilateral TF with complete ra
fluoroscopy and preoperative CT scan which improved our understand
and the degree of instability. The exclusion criteria were as follows: (1)
(2) high-energy injury or multiple injuries such as car accidents or high
patients with pathological fractures and previous histories of hip malfo
By observing intraoperative fluoroscopy, the TF cases demonstrating th
reaming or nail insertion were identified. The demographic information of this cohort was extracted from patient medical records. The
radiographic data were available in the Picture Archiving and Communication System (PACS), and the
measurements were carried out with the built-in gauge tools. One of the senior surgeons at the hospital
classified the fractures according to the AO/OTA classification on plain radiographs (H.J.). Page 3/18 Page 3/18 To eliminate the negative influences of initial displacement on the fracture morphology analysis, three-
dimensional CTs (3D-CT) of both sides of the proximal femurs were reconstructed simultaneously. The
fracture line at the basicervical region was designated the primary fracture line, and the coronally
propagated fracture line affecting the GT was designated the secondary fracture line. The analysis of the
reconstructed 3D images revealed that there were three consistent fragments: cephalocervical fragment,
femoral shaft and posterolateral fragment from GT. A variable fragment (LT) was also frequently
identified. Therefore, each case could be defined as either a 3-fragmentary or 4-fragmentary pattern. Then, the rationale of Shoda’s 3D-CT classification of the proximal femur was borrowed to further
classify it into one of four subgroups [7]. Materials And Methods The 3-fragmentary S was the first subgroup, where “S” referred to
a small PLF fragment combined with cephalocervical and shaft fragments (Figure 3A). The second
subgroup was the 3-fragmentary M, in which a moderate PLF (indicated by “M”) coexisted with other two
consistent fragments. Compared to the 3-fragmentary S, the secondary fracture line in the 3-fragmentary
M propagated more medially and was in the vicinity of the lateral border of the intact LT (Figure 3B). In
the third subgroup, one large banana-like fragment consisting of the PLF and the LT as one unit occurred
in the 3-fragmentary L (Figure 3C). When the LT was also concomitantly fractured, there were four large
fragments; thus, it was defined as the 4-fragmentary GL subtype (Figure 3D). On the 3D-CT, the morphological feature of the primary fracture line was specifically studied [7]. As the GT
of the injured limb was routinely comminuted and the normal spatial relationship between the main
fragments was altered, we utilized the mirrored image of the contralateral uninjured proximal femur as a
template for the injured hip to precisely evaluate the distribution primary fracture line. After the DICOM
data were entered into Materialise 3-matic (Materialise NV Inc., Leuven, Belgium), the cephalocervical
fragment of the TF was maximally matched to the mirrored 3D image of the contralateral femur. On the
fused images, the relationship between the primary fracture line and the GT could be analyzed (Figure 4). Thirty-one patients were stabilized by proximal femoral nail anti-rotation (PFNA, Synthes Inc., Oberderf,
Switzerland) and two younger patients by a TRIGEN INTERTAN nail (Smith–Nephew, Memphis, USA). The
postoperative radiography was assessed by another fellow training surgeon who did not take part in the
operation (H.G.Q). The width of the secondary displacement at the inferior part of the primary fracture
line and the NSA were measured on the anteroposterior (AP) view (Figure 5). All statistical analyses were performed using Statistical Package for the Social Sciences (SPSS) (ver
23.0, International Business Machines, Inc. Armonk, New York, USA). Quantitative data were expressed as
median and compared among groups by Student’s t-test or Mann-Whitney U test. All P-values were two-
sided and a P-value < <lue < 0.05 was considered to be statistically sign Surgical technique Under general anesthesia, the patient was positioned on the operation table in supination with consistent
distraction of the injured limb, and the manual reduction of the fracture was performed. By persistent
traction and an internal rotation maneuver, the limb length and rotational alignment were restored first. Page 4/18 After the medial cortical continuity and acceptable NSA were confirmed on AP view of the fluoroscopic
image, the anterior cortex reduction was checked on lateral view. If there was negative reduction of the
anterior cortex, a small elevator was percutaneously inserted to reduce it. No attempt was made to reduce
an isolated LT fragment or PLF from the GT. When the reduction was satisfactory on both views, a PFNA/TRIGEN INTERTAN nail was inserted
according to the manufacturer’s operation manual. After making a 5-cm incision proximal to the tip of the
GT, the fasciae and muscle fibers of the gluteus medius were split. As the standard manipulation, the
entry point of the nail was slightly lateral to the tip of the GT, and guide wire placement was followed by
opening the femur, reaming the medullary canal and inserting a PFNA/TRIGEN INTERTAN nail. Frequently,
“reverse wedge effect” occurred either during canal reaming or nail insertion. Before 2017, we resorted to enlarging the entrance of the femoral canal or overdistracting the injured limb
to overcome this secondary deformity. However, the design of the reamer made the over-reaming
technique extremely difficult, and the overdistraction method generally failed. Later, we chose to secure
the acquired initial reduction by inserting a 3.5-mm Kirschner wire (K-wire) before reaming and
PFNA/TRIGEN INTERTAN nail insertion (Figure 6E). Under image intensifier control, the K-wire was
inserted along the axis of the femoral neck and closely underneath the anterior cortex. Then, the femoral
canal was reamed, and the nail was inserted. A bone hook should sometimes be added to augment the
maintenance of the primarily achieved reduction (Figure 6F). We found that those methods were
extremely effective and dramatically shortened the operation time if the risk of the “reverse wedge effect”
could be predicted preoperatively. Results A total of 33 patients met the inclusion criteria in the final analysis so that the overall incidence of the
“reverse wedge effect” was 7.9%. The preoperative plain radiograph analysis revealed that all the 33
patients were basicervical TF variants while there were another 12 cases of this special fracture type did
not present similar intraoperative complication. The patients included 12 males and 21 females, with a
mean age of 72.5 years (33 to 96 years). The internal rotation of the cephalocervical fragment caused by
reamer/IN impingement was the main underlying cause of this intraoperative complication. On the
postoperative radiograph, this secondary displacement demonstrated an inferior gap at the primary
fracture line and increased NSA (Figure 2B). The AO/OTA classification scheme failed to properly
categorize this special fracture type properly [8]: on the plain radiograph, the coronally oriented secondary
fracture line made the measurement of the lateral wall thickness unreliable and generated great
confusion in differentiating AO-31 A1 GT from A2. In contrast, the classification system we devised in this
study depicted each case briefly and concisely. The incidence of the four-fragment TF (4-fragmentary GL) was the highest (Table 1). In all four
subgroups, the majority of the primary fracture lines transversed the center of the piriformis fossa Page 5/18 Page 5/18 Page 5/18 (83.8%) (Figure 4A). The remaining fracture lines marginally involved a small portion of the GT medial
wall (Figure 4B). Therefore, all the primary fracture lines originated extracapsularly. A Mann-Whitney U test was run to determine if there were differences in magnitude of displacement and
the NSA between 3-fragmentary and 4-fragmentary subgroups. Distributions of the secondary
displacement and NSA for 3-fragmentary and 4-fragmentary subgroups were similar, as assessed by
visual inspection. Median displacement for 3-fragmentary (8.20 mm) and 4-fragmentary (9.20 mm) was
not statistically significantly different, U = 80.5, z = -1.825, p = 0.068, using an exact sampling distribution
for U. While, the median NSA for 3-fragmentary (136.0 degrees) and 4-fragmentary (133.0 degrees) was
not statistically significantly different, U = 116.0, z = -0.518, p = 0.624. Discussion As
Cho’s study found that the incidence of the PLF in TF was as high as 88.4% [15], we observed a similar prevalence among all the 414 patients, and every “reverse wedge effect” case had a PLF. We believe that
further classification and comparison would facilitate understanding and exploring the influences of
those two fragments on “reverse wedge effect” formation and treatment/strategy design. Although Van Embden stated that AO/OTA classification was more comprehensive and reliable
compared to the Jensen-Evans classification [16], we found that it failed to cover this special fracture
pattern. According to AO/OTA classification (2018 version), AO-31A1 and A2 were designated as
pertrochanteric fractures in which the main fracture line propagated through the trochanters. Saarenpaa,
Watson and other authors described basicervical TF variant as fracture at the base of femoral neck that
is medial to the intertrochanteric line [9, 17, 18]. So basicervical TF variant could not be classified into
pertrochanteric (AO - 31A1 and A2) or intertrochanteric fracture(AO - 31A3). In contrast, in 1949, even
before the clinical application of computed tomography, Evans had proposed 3- and 4-fragmentary
fracture patterns in his classification scheme [19]. Considering the complexity of the fracture morphology,
a three-dimensional classification system originating from Evans’ would be more pragmatic [16]. The
principle from Babhulkar and Shoda’s classifications of GT fracture [2, 7] could be incorporated to further
classify basicervical TF variants according to the morphology of the PLF. Although the sample size was not large enough to make statistical comparisons between the four
subgroups, the measured displacements of “reverse wedge effect” and ultimate NSA were similar among
them. Therefore, we could infer that the volume of the PLF minimally influenced the extent of the
secondary displacement in this cohort of patients. From a mechanical view, it is the existence of the PLF
contributes to the development of the “reverse wedge effect”. When the GT region is intact, the guide wire could be constrained around the tip of the GT, and the
trajectory of the reamer/IN was centralized into the femoral canal. However, an incompetent GT made the
guide wire and reamer float at the start site. Under an image intensifier, we observed that the reamer was
prone to skew medially, especially when attempting to over-ream the superolateral corner of the
cephalocervical fragment (Figure 6D). As a result, impaction between the cephalocervical fragment and
the reamer/nail occurred. Discussion The purpose of the present study was to investigate the incidence, characteristics and causes of the
“reverse wedge effect” during application of the IN in TF. In contrast to the previously reported “wedge
effect”, which was defined as an intraoperative secondary displacement at the superior part of the
pertrochanteric fracture line [6], we observed another IN-related intraoperative complication, which was
demonstrated as an inferior opening at the already reduced primary fracture line. We also identified that
all the cases occurred in basicervical TF variants combined with a disrupted GT. By reviewing
intraoperative fluoroscopy, we noticed an impingement between the reamer/IN and the superolateral edge
of the cephalocervical fragment, which caused this secondary displacement (Figure 6). Because all the
414 reviewed cases were treated by PFNA/TRIGEN INTERTAN nailing but the incidence of the “reverse
wedge effect” was nearly 7.97%, we believe that anatomic factors other than implantation should play a
major role. The basicervical fracture line is the most significant feature of this cohort of patients. Strictly, a
basicervical fracture is defined as a 2-part fracture where the fracture line originates from the base of the
femoral neck and exits above the lesser trochanter [9]. Despite the concomitant PLF or LT fracture, the
primary fracture lines in this cohort of patients met this description exactly. There is little controversy
about the fact that a basicervical fracture is an unstable fracture and prone to varus deformity [10]. Sliding
screws were initially proposed as a treatment for these fractures [11, 12], but the recent trend favors IN
fixation [9, 13]. Anatomically, the piriformis fossa is the transitional region between the hard cortex of the
femoral neck and the relatively weak cancellous bone of the GT [14]. Our study demonstrated that the
primary fracture lines just crossed the piriformis fossa center. This indicated that the superolateral corner
of the cephalocervical fragment was primarily made from the hard cortex, which is hard enough to resist
reaming and colliding with the reamer and PFNA/TRIGEN INTERTAN nail insertion. Eventually, the reamer
or EFNA/TRIGEN INTERTAN nail would push the cephalocervical fragment, internally rotating and
inferiorly displacing, resulting in the “reverse wedge effect”. Page 6/18
Notably, in this cohort, the PLF and LT fragment were the other two prominent anatomical features. Discussion Although mild valgus reduction is more preferred than varus reduction for allowing interfragment
compression and reducing bone-implant stresses, Ciufo et al. recently showed that residual basicervical
gapping was closely associated with fixation cutout [21]. Considering that the magnitude of the secondary
displacement exceeded 4 mm, which was proposed by Baumgaertner as the lowest threshold for
malreduction [22], we believe that this newly reported intraoperative complication is worth further
investigation. There were some limitations to the present study. First, this study was a retrospective analysis. Therefore,
the robustness of the analysis is undetermined. Second, a comparison between the different trochanteric
fracture patterns in the incidence rate of the “reverse wedge effect” was not carried out. However, we
rigorously reviewed all the intraoperative fluoroscopy findings during the study period and did not find
any “reverse wedge effect” in other fracture patterns. Thus, we believe that this complication is closely
correlated with the nailing basicervical TF variant combined with the PLF. Third, the treatment outcomes
of this cohort of patients were not evaluated for several reasons. The primary reason is that this cohort of
patients was highly heterogeneous, and some very young patients were included, which made building an
age- and sex-matched control group difficult. Conclusions Our study reveals traumatic anatomic factors associated with the “reverse wedge effect” in
cephalomedullary nailing in a group of trochanteric fractures. In contrast to the previously proposed
“wedge effect”, this “reverse wedge effect” is characterized by an inferior opening at the primary fracture
line in the trochanteric region and a mild valgus malreduction. This secondary displacement exclusively
occurred during the reaming/nailing of an anatomically reduced trochanteric fracture featuring a
basicervical variant type and a disrupted GT. Further analysis demonstrated that the basicervical fracture
line is the major contributor to this intraoperative complication and that the impingement between the
reamer/nail and the superolateral cortex tip of the cephalocervical fragment was the direct cause. Thus,
the magnitude of the displacement is significant, and early postoperative weight bearing might be
compromised. We advocate that this special intraoperative complication should be prevented in advance
of reaming and nail insertion to optimize the final reduction quality. Discussion Since the volume of the PLF plays a relatively minor role, this fragment could be
excluded during the management of the “reverse wedge effect”. In this cohort of patients, the incidence of PLF was 100% and the LT disruption was 60.6%. To investigate
the role of LT disruption in this operative complication, comparisons between 3- and 4-fragmentary
subgroups in the magnitude of displacement and postoperative NSA were made and revealed no
significant differences. We could conclude that the appearance of LT fragment contributed minimally to
the occurence and degree of the “reverse wedge effect”. When preventing the reverse wedge effect in
basicervical TF variant, the lesser trochanter fragment did not have to be reduced and temporarily
stabilized. It is unclear how this intraoperative complication negatively impacts the treatment outcome. The
postoperative measurement demonstrated that the magnitude of the secondary displacement in the Page 7/18 “reverse wedge effect” was significant. Zhang et al. advocated the “medial positive reduction” concept to
validate a stable medial cortex mismatch, but they did not quantify the threshold of the acceptable
diastasis at the medial cortex [20]. It is reasonable that an obvious opening at the primary fracture line
represents instability in the fracture. Thereafter, postoperative excessive collapse would occur because of
failure to restore cortex interdigitation. Early postoperative weight bearing had to be postponed. The
postoperative radiograph demonstrated a mild tendency of valgus reduction in the majority of patients. Although mild valgus reduction is more preferred than varus reduction for allowing interfragment
compression and reducing bone-implant stresses, Ciufo et al. recently showed that residual basicervical
gapping was closely associated with fixation cutout [21]. Considering that the magnitude of the secondary
displacement exceeded 4 mm, which was proposed by Baumgaertner as the lowest threshold for
malreduction [22], we believe that this newly reported intraoperative complication is worth further
investigation. “reverse wedge effect” was significant. Zhang et al. advocated the “medial positive reduction” concept to
validate a stable medial cortex mismatch, but they did not quantify the threshold of the acceptable
diastasis at the medial cortex [20]. It is reasonable that an obvious opening at the primary fracture line
represents instability in the fracture. Thereafter, postoperative excessive collapse would occur because of
failure to restore cortex interdigitation. Early postoperative weight bearing had to be postponed. The
postoperative radiograph demonstrated a mild tendency of valgus reduction in the majority of patients. Authors’ contributions QXD and ZY performed the study. HGJ and HJ participated in the operation and data
collection/interpretation for the study. ZY performed the statistical analysis. QXD conceived of the study
and participated in its design. LX was responsible for coordination, data collection/interpretation and
proofreading the final manuscript. All authors read and approved the final manuscript. Yu Zhang will be contact to provide data and materials. Yu Zhang will be contact to provide data and materials. Consent for publication Not applicable. Competing interests The authors declare that they have no competing interests. The authors declare that they have no competing interests. We gratefully acknowledge the support of all the doctors in our departmen No funding was obtained for this study Ethics approval and consent to participate Approved by the Institutional Review Board (IRB)/Independent Ethics Committee (IEC) of Jiangsu
Province Hospital (The First Affiliated Hospital of Nanjing Medical University) at 300 Guangzhou Road,
Nanjing 210029, China. All subjects provided informed consent in written to take part in the study. Authors’ Affiliations Department of Trauma, the First Affiliated Hospital of Nanjing Medical University and Jiangsu Province
Hospital Acknowledgements Acknowledgements Page 8/18 Abbreviations IN: intramedullary nailing
GT: greater trochanter
TF: trochanteric fractures
CT: computed tomography IN: intramedullary nailing Page 9/18 AP: anteroposterior K-wire: Kirschner wire K-wire: Kirschner wire References 1. Cummings SR, Rubin SM, Black D. The future of hip fractures in the United States: numbers, costs,
and potential effects of postmenopausal estrogen. Clin Orthop 1990,252:163–166 1. Cummings SR, Rubin SM, Black D. The future of hip fractures in the United States: numbers, costs,
and potential effects of postmenopausal estrogen. Clin Orthop 1990,252:163–166 2. Babhulkar S. Unstable trochanteric fractures: Issues and avoiding pitfalls. Injury. 2017;48(4):803-818. 3. Sharma G, Kumar G N K, Yadav S, et al. Pertrochanteric fractures (AO/OTA 31-A1 and A2) not
amenable to closed reduction: causes of irreducibility. 2014;45(12):1950-1957. 4. Kashigar A, Vincent A, Gunton MJ, et al. Predictors of failure for cephalomedullary nailing of
proximal femoral fractures. Bone Joint J. 2014;96-B(8):1029-1034. 5. Hak DJ, Bilat C. Avoiding varus malreduction during cephalomedullary nailing of intertrochanteric hip
fractures. Arch Orthop Trauma Surg. 2011;131(5):709-710. 6. O'Malley MJ, Kang KK, Azer E, et al. Wedge effect following intramedullary hip screw fixation of
intertrochanteric proximal femur fracture. Arch Orthop Trauma Surg. 2015;135(10):1343-1347. 6. O'Malley MJ, Kang KK, Azer E, et al. Wedge effect following intramedullary hip screw fixation of
intertrochanteric proximal femur fracture. Arch Orthop Trauma Surg. 2015;135(10):1343-1347. 7. Shoda E, Kitada S, Sasaki Y, et al. Proposal of new classification of femoral trochanteric fracture by
three-dimensional computed tomography and relationship to usual plain X-ray classification. J
Orthop Surg (Hong Kong). 2017;25(1):1-5. 8. Meinberg EG, Agel J, Roberts CS, et al. Fracture and Dislocation Classification Compendium-2018 J
Orthop Trauma. 2018;32 Suppl 1:S1-S170. 9. Watson ST, Schaller TM, Tanner SL, et al. Outcomes of Low-Energy Basicervical Proximal Femoral
Fractures Treated with Cephalomedullary Fixation. J Bone Joint Surg Am. 2016;98(13):1097-1102. 10. Gill JM, Johnson GR, Sher JL, et al. Biomechanical aspects of the repair of intertrochanteric
fractures. J Biomed Eng. May 1989; 11 (3):235-239. 11. Baumgaertner MR, Higgins TF. Femoral neck fractures. In:Rockwood CA, Bucholz RW, Heckman JD,
Green DP, eds. Rockwood and Green’s Fractures in Adults. Vol 2. 5th ed. Philadelphia, Pa:Lippincott
Williams and Wilkins; 2001:1588. Page 10/18 12. Su BW, Heyworth BE, Protopsaltis TS, et al. Basicervical versus intertrochanteric fractures: an
analysis of radiographic and functional outcomes. Orthopedics. 2006;29(10):919-925. 13. Hu SJ, Yu GR, Zhang SM. Surgical treatment of basicervical intertrochanteric fractures of the
proximal femur with cephalomeduallary hip nails. Orthop Surg. 2013;5(2):124-129. 14. Reece GM, Poole KE, Gee AH. Imaging the femoral cortex: thickness, density and mass from clinical
CT. Med Image Anal. 2012;16(5):952-65. 15. References Cho JW, Kent WT, Yoon YC, et al. Fracture morphology of AO/OTA 31-A trochanteric fractures: A 3D
CT study with an emphasis on coronal fragments. Injury. 2017\;48(2):277-284. 16. Van Embden, S. Meylaerts, S. Rhemrev, et al. The Comparison of Two Classifications for Trochanteric
Femur Fracture: The AO/ASIF Classification and the Jensen Classification. Injury. 2010;41(4):377-81. 17. Saarenpaa I, Partanen J, Jalovaara P. Basicervical fracture-a rare type of hip fracture. Arch Orthop
Trauma Surg. 2002;122(2):69-72. 18. Okano I, Sawada T, Kushima N, et al. Treatment With Helical Blade Cephalomedullary Nail for Two-
Part Basicervical Proximal Femoral Fracture in Elderly Patients: A Retrospective Observational Study. Geriatr Orthop Surg Rehabil. 2017;8(4):244–251. 19. Evans EM. The treatment of trochanteric fractures of the femur. J Bone Joint Surg Br. 1949;31
(2):190-203. 19. Evans EM. The treatment of trochanteric fractures of the femur. J Bone Joint Surg Br. 1949;31
(2):190-203. 20. Chang SM, Zhang YQ, Ma Z, et al. Fracture reduction with positive medial cortical support: a key
element in stability reconstruction for the unstable pertrochanteric hip fractures. Arch Orthop Trauma
Surg. 2015 Jun;135(6):811-818. 21. Ciufo DJ, Zaruta DA, Lipof JS, et al. Risk Factors Associated With Cephalomedullary Nail Cutout in
the Treatment of Trochanteric Hip Fractures. J Orthop Trauma. 2017 ;31(11):583-588. 22. Baumgaertner MR, Curtin SL, Lindskog DM, et al. The value of the tip-apex distance in predicting
failure of fixation of peritrochanteric fractures of the hip. J Bone Joint Surg Am 1995;77(7):1058–
1064. Table subgroups
3-fragmentary
4-fragmentary
p value
Frequency
13
20
-
Median Displacement (mm)
8.2 (6.90, 8.85)
9.2 (8.45, 10.95)
0.068
Median NSA (degree)
136.0 (130.5, 138.5)
133.0 (132.0, 138.8)
0.624
Table 1. The 3-fragmentary and 4-fragmentary subgroups were compared with Mann-
Whitney U tests
Figure
s Page 11/18 Page 11/18 Figure 1
The simulated diagram depicting the “wedge effect”, where the intramedullary nail insertion causes
lateralization of the femoral shaft and varus malalignment. Figure 1 The simulated diagram depicting the “wedge effect”, where the intramedullary nail insertion causes
lateralization of the femoral shaft and varus malalignment. Page 12/18 Page 12/18 Figure 2
(A) The preoperative anteroposterior (AP) radiograph showing an apparently simple AO 31.A 2.2
pertrochanteric fracture. (B) Following fixation with PFNA, there was an obvious inferior gap at the
basicervical region (white arrow) and moderate valgus deformity. The lucency around helix indicates a
micromotion and instability of the fracture-implantation complex. Figure 2 (A) The preoperative anteroposterior (AP) radiograph showing an apparently simple AO 31.A 2.2
pertrochanteric fracture. (B) Following fixation with PFNA, there was an obvious inferior gap at the
basicervical region (white arrow) and moderate valgus deformity. The lucency around helix indicates a
micromotion and instability of the fracture-implantation complex. (A) The preoperative anteroposterior (AP) radiograph showing an apparently simple AO 31.A 2.2
pertrochanteric fracture. (B) Following fixation with PFNA, there was an obvious inferior gap at the
basicervical region (white arrow) and moderate valgus deformity. The lucency around helix indicates a
micromotion and instability of the fracture-implantation complex. Page 13/18 Page 13/18 Figure 3 Figure 3 Figure 3 Figure 3 Page 14/18 (A) The cephalocervical fragment was fused on the mirror image of the contralateral proximal femur. The
primary fracture line of the injured side just crossed the piriform fossa. (B) The primary fracture line
involved a small piece of the posteromedial wall of the GT. Figure 4 (A) The cephalocervical fragment was fused on the mirror image of the contralateral proximal femur. The
primary fracture line of the injured side just crossed the piriform fossa. (B) The primary fracture line
involved a small piece of the posteromedial wall of the GT. (A) The cephalocervical fragment was fused on the mirror image of the contralateral proximal femur. The
primary fracture line of the injured side just crossed the piriform fossa. (B) The primary fracture line
involved a small piece of the posteromedial wall of the GT. Page 15/18 Figure 5 Figure 5 The tangential lines to the medial cortexes of the cephalocervical and shaft fragments are drawn. The
distance between the two lines represents the width of the basicervical gap. Page 16/18 e 6 B) Preoperative 3D image demonstrating a subtype 4-fragmentary GL
us deformity. (C and D) The wide arrow indicates the impact between
ateral cortex of the cephalocervical fragment. The resultant basicervic Figure 6
(A and B) Preoperative 3D image demonstrating a subtype 4-fragmentary GL pertrochanteric fracture with
mild varus deformity. (C and D) The wide arrow indicates the impact between the reamer and the
superolateral cortex of the cephalocervical fragment. The resultant basicervical gap is designated by a
narrow arrow. (E) When the risk of reverse wedge effect was high, a 3.5-mm Kirschner wire (K-wire) was inserted to secure the reduction. (F) Occasionally, a single K-wire was not strong enough, and a bone hook
was inserted to prevent inferior opening at the fracture line. Figure 6 (A and B) Preoperative 3D image demonstrating a subtype 4-fragmentary GL pertrochanteric fracture with
mild varus deformity. (C and D) The wide arrow indicates the impact between the reamer and the
superolateral cortex of the cephalocervical fragment. The resultant basicervical gap is designated by a
narrow arrow. (E) When the risk of reverse wedge effect was high, a 3.5-mm Kirschner wire (K-wire) was Page 17/18 Page 17/18 Page 17/18 inserted to secure the reduction. (F) Occasionally, a single K-wire was not strong enough, and a bone hook
was inserted to prevent inferior opening at the fracture line. Page 18/18 Page 18/18
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First Live-Experience Session with PET/CT Specimen Imager: A Pilot Analysis in Prostate Cancer and Neuroendocrine Tumor
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Disclaimer/Publisher’s Note: The statements, opinions, and data contained in all publications are solely those of the individual author(s) and
contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting
from any ideas, methods, instructions, or products referred to in the content.
Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Disclaimer/Publisher’s Note: The statements, opinions, and data contained in all publications are solely those of the individual author(s) and
contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting
from any ideas, methods, instructions, or products referred to in the content. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 A pilot analysis in Prostate Cancer and Neuroendocrine Tumor. Lorenzo Muraglia1, Francesco Mattana1, Laura Lavinia Travaini1, Gennaro Musi2,3, Emilio Bertani4,
Giuseppe Renne5, Eleonora Pisa5, Mahila Esmeralda Ferrari6, Uberto Fumagalli Romario4, Ottavio De
Cobelli2,3, Nicola Fusco3,5, Francesco Ceci1,3* 1 Division of Nuclear Medicine, IEO European Institute of Oncology IRCCS, Milan, Italy
2 Division of Urology, IEO European Institute of Oncology IRCCS, Milan, Italy
3 Department of Oncology and Hemato-Oncology, University of Milan, Milan, Italy
4 Division of Digestive Surgery, IEO European Institute of Oncology IRCCS, Milan, Italy
5 Division of Pathology, IEO European Institute of Oncology IRCCS, Milan, Italy
6 Unit of Medical Physics, IEO European Institute of Oncology IRCCS, Milan, Italy ABSTRACT Objective: to evaluate the feasibility of the intra-operative application of a specimen PET/CT imager
in a clinical setting. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Materials and methods: this is a pilot analysis performed in three patients who received an intra-
operative administration of 68Ga-PSMA-11 (n=2) and 68Ga-DOTA-TOC (n=1), respectively. Patients
were administrated with PET radiopharmaceuticals to perform radio-guided surgery with a beta-
probe detector during radical prostatectomy for prostate cancer (PCa) and salvage
lymphadenectomy for recurrent neuroendocrine tumor (NET) of the ileum, respectively. All
procedures have been performed within two ongoing clinical trials in our Institute (NCT05596851
and NCT05448157). Pathologic assessment with immunohistochemistry (PSMA-staining and SSA
immunoreactivity) was considered as standard of truth. Specimen images were compared with
baseline PET/CT images and histopathological analysis. Results: Patients received 1 MBq/Kg of 68Ga-PSMA-11 (PCa) or 1.2 MBq/Kg of 68Ga-DOTA-TOC (NET)
prior to surgery. Specimens were collected, positioned in the dedicated specimen container, and
scanned to obtain high resolution PET/CT images. In all cases a perfect match was observed between
the findings detected by the specimen imager and histopathology. Overall, the PET spatial resolution
was sensibly higher for the specimen images compared to the baseline whole-body PET/CT images. Furthermore, the use of the PET/CT specimen imager did not significantly interfere with any
procedures, and the overall length of the surgery was not affected using the PET/CT specimen
imager. Finally, the radiation exposure of the operating theater staff was lower than 40 µSv per
procedure (range 26 – 40 Sv). Conclusion: the image acquisition of specimens obtained by patients who received intra-surgery
injection of 68Ga-PSMA-11 and 68Ga-DOTA-TOC was feasible and reliable also in a live-experience
session and has been easily adapted to surgery daily-practice. The high sensitivity, together with the
evaluation of intra-lesion tumor heterogeneity, were the most relevant results since the data
derived from specimen PET/CT imaging matched perfectly with the histopathological analysis. Keywords: PET/CT; PET; PET/CT specimen imager; radio-guided surgery Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444 Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444 doi:10.20944/preprints202212.0444.v2 The aim of this pilot analysis was to assess the feasibility of a PET/CT specimen imager in a live-
experience session in patients who received an intraoperative injection of 68Ga-PSMA-11 and 68Ga-
DOTA-TOC. The aim of this pilot analysis was to assess the feasibility of a PET/CT specimen imager in a live-
experience session in patients who received an intraoperative injection of 68Ga-PSMA-11 and 68Ga-
DOTA-TOC. Patient Population This pilot analysis was performed as a secondary objective of two trials currently ongoing in our
Institute in Pca (NCT05596851) and NET (NCT05448157) designed to evaluate the safety and the
efficacy of a positron probe detector (DROP-IN β-probe) for live intra-operative radio-guided
surgery. Three patients enrolled in these trials have been considered eligible for this supplemental
analysis, two affected by Pca and one affected by NET of the ileum. In all cases, patients received a
baseline whole-body PET/CT to assess the burden of the disease (68Ga-PSMA-11 PET/CT and 68Ga-
DOTA-TOC PET/CT respectively). The scans were performed in the Division of Nuclear Medicine of
our Institute (GE Discovery MIDR) according to international procedural guidelines, 4-6 weeks prior
to surgery. All patients signed informed consent form prior to enrollment and studies have been
approved by local ethical committee and scientific review board. INTRODUCTION Positron Emission Tomography (PET) imaging is considered one of the leading imaging techniques
to investigate oncological patients and it is considered the standard-of-care in many solid tumors
and hematological malignancies (1). Recently, several technological improvements appeared in the
field of PET imaging, ranging from digital detectors (e.g. silicon photomultipliers) to long axial field-
of-view scanners (2). However, the attention moved also to smaller PET/computed tomography (CT)
tomographs that might be implemented in the daily clinical practice of surgical oncology. Recently,
a compact and mobile PET/CT specimen imager was developed to obtain full 3D high resolution
images, assisting surgeons and pathologists during the surgical procedure directly in the operating
theater. This specimen imager combines the picomolar sensitivity of PET (at sub-millimeter
resolution) with anatomical information obtained from a high-resolution micro-CT. The first
research application was in breast cancer using 18F-FDG as a PET radiopharmaceutical, where a
multicenter clinical trial is currently on-going (NCT04999917). Intra-operative high-resolution
PET/CT imaging has shown promising results for margin assessment in breast malignancies, where
a respective sensitivity and specificity of 84% and 96% was reached for the identification of positive
margins of the invasive component of invasive ductal carcinoma (3). Explorative studies in
pancreatic adenocarcinoma and in head and neck cancers have also demonstrated the feasibility of
intra-operatively imaging radiotracer uptake in high resolution specimen PET/CT images (4,5). Recently, novel radiopharmaceuticals emerged as game changers in prostate cancer (PCa) with
prostate specific membrane antigen (PSMA) PET and in neuroendocrine tumor (NET) with
somatostatin receptor (SSR) based PET. PSMA-PET showed its superiority compared to conventional
imaging or other PET tracers in randomized clinical trials (6,7) or registry trials (8-10) to stage Pca or
in case of disease recurrence, while SSR-agonist PET, is currently the standard-of-care to investigate
gastro-entero-pancreatic NET (GEP-NET), especially in case of low-grade disease (G1-G2) (11). Furthermore, both PSMA inhibitors and SSR agonists are theranostic agents and can be labeled with
isotopes for PET imaging (e.g. Gallium-68) or high-energy beta or alpha emitters (e.g. Lutetium-177
or Actinium-225) for radio-ligand therapy. Therefore, there is rising and increasing interest to
explore the potential of PET/CT specimen imagers beyond 18F-FDG, applying this high-resolution
imaging in patients undergoing surgery for Pca or NET. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 through a dedicated viewer interface (red square). As such the viewer does provide a 3D image of
the specimen. Additionally, a 3D render is present, which can be rotated by the user. through a dedicated viewer interface (red square). As such the viewer does provide a 3D image of
the specimen. Additionally, a 3D render is present, which can be rotated by the user. Specimen PET/CT imager PET/CT images were obtained using the AURA10® Specimen PET/CT imager (Xeos Medical NV,
Belgium). Subsequently, optical, micro-CT and micro-PET images were acquired. Specimens were
scanned in a dedicated specimen container (10 cm diameter and 4 cm height) (Figure 1). CT images
were reconstructed using the Image Space Reconstruction Algorithm at 100µm voxel size. PET
images were reconstructed using 20 iterations of the ordered subset expectation maximization
algorithm at 400µm voxel size. The total duration between specimen insertion and visualization of
the images on the device was 10 minutes. Figure 1: Overview of the PET/CT specimen imager. On the right is the device with its main
constitutes: the medical display (red triangle), the specimen box container (red circle), the viewer
interface (red square) and the hardware that allows a two-step acquisition (CT and PET – red star). The viewer of the PET/CT specimen imager shows the images in three different planes (coronal,
sagittal and transversal) as shown in the left figure. The user can scroll through each of the planes Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 In all cases, surgical specimens were collected in the operating theater and placed in dedicated
containers by the pathologist, who oriented the specimens in the same position adopted for
histopathological analysis. Specimen containers were placed in the PET/CT specimen imager and
one scan was performed for each specimen. PET/CT acquisitions were performed 2.5 to 4.5 hours
post-injection. Such delayed acquisition times can be made due to the high detection sensitivity of
the Imager. Of some specimens, several acquisitions were performed, to explore the potential of
administering less activity. High-resolution fused 3D images were obtained and reviewed in a
multidisciplinary setting involving nuclear medicine physicians, surgeons, and pathologists. Pathologic assessment with immunohistochemistry (PSMA-staining and SSA immunoreactivity) was
considered as standard of truth. Specimen images were subsequently compared with baseline
PET/CT images and histopathological analysis. Baseline whole-body PET/CT images were performed according to EANM guidelines after an
intravenous injection of 2 MBq/kg of 68Ga-PSMA11 and 2,5 MBq/kg of 68Ga-DOTATOC. Baseline
scans were performed within six weeks before the day of the surgery (patient-1: 4 weeks; patient-
2; 6 weeks; patient-3: 4 weeks). Description of the surgery procedure This analysis was a secondary objective of two ongoing phase II trials at our institution
(NCT05596851; NCT05448157), involving the intra-operative administration of 68Ga-PSMA-11 and
68Ga-DOTATOC for radio-guided surgery with a dedicated beta-probe. Therefore, the activity used
is that defined by the protocol of the trials. In this pilot study we demonstrated that high-quality
images can be made with the specimen PET/CT imager, using a low activity of 68Ga-PSMA-11 and
68Ga-DOTATOC. The PCa patients received 1 MBq/kg of68Ga-PSMA-11 and the NET patient
received 1.2 MBq/kg of 68Ga-DOTA-TOC. PET radiopharmaceutical administration was performed
directly in the surgery theater at the beginning of the procedure. For Pca patients a robotic-assisted
laparoscopic approach was adopted. Extended pelvic lymph node dissection (ePLND) was
performed according to a standard template, including internal/external iliac nodes, obturatory
nodes and common iliac ones. Radical prostatectomy (RP) was performed after ePLND, according
to the study protocol (NCT05596851). For the ileum GEP-NET patient, open surgery was chosen as
surgical approach, according to study protocol (NCT05448157). Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 the areas of PSMA-uptake in the micro-PET image corresponded to prostate cancer lesions on
histopathological analysis. Extra-prostatic extension, as suspected from the micro-PET images, was
also confirmed. Lymph nodes that showed PSMA uptake on micro-PET were confirmed to contain
metastatic tumor cells on histopathological analysis. All positive lymph nodes detected in
preoperative whole-body PET/CT were resected and imaged. Patient n.2 was referred to PSMA-PET (Figure 3: A,E,I) for high-risk Pca (Gleason Score= 4+5; fusion
biopsy, 21/21 positive cores bilateral, initial PSA= 8 ng/mL). PSMA-PET showed pathological and
heterogeneous uptake in the whole prostate, with suspicion of extra-capsular extension, two PSMA-
positive pelvic lymph nodes and one PSMA-positive extra-pelvic lymph node. According to E-PSMA
reporting criteria (12) the patient was considered miT3N1M1a. The day of surgery, 90 MBq of 68Ga-
PSMA-11 was administered intravenously. The prostate gland and lymph nodes were resected
(Figure 2: B,F,J) and all specimens were scanned in the PET/CT specimen imager. Scans were
performed sequentially, once the specimen was positioned in the container and properly oriented
by the pathologist, at 206 min (prostate) and 265 min (lymph nodes) post-injection. Images (Figure
3: C,G,K) showed a high spatial resolution, and margin involvement was strongly suspected (Figure
3: C,D), in accordance with baseline PSMA-PET. PET/CT specimen images offered a high-resolution
view of the distribution of PSMA inside the prostate lesion, highlighting high heterogeneity of PSMA
expression in the prostate. As in the first case, areas of PSMA-uptake in the micro-PET image
corresponded to prostate cancer lesions on histopathological analysis and a positive margin – as
suspected from the microPET-CT images – was confirmed. Lymph nodes that showed PSMA uptake
on micro-PET were confirmed to contain metastatic tumor cells on histopathological analysis. Patient n.3 underwent 68Ga-DOTA-TOC-PET (Figure 4: A) for recurrent ileum GEP-NET (G2; Ki-67=
5%). The PET scan showed the presence of a peri-pancreatic lymph node suspected of disease
location. The day of surgery 70 MBq of 68Ga-DOTA-TOC was administered intravenously. The PET-
positive lymph node was resected and placed in the PET/CT specimen container. The specimen was
scanned in the PET/CT specimen imager (Figure 4: B,C) at 150 minutes post-injection properly
showing a high-resolution view of homogeneous 68Ga-DOTA-TOC biodistribution within the lymph
node. The specimen was analyzed for histopathology and immune-histochemistry assessment with
SSR immune-reactivity (Figure 4: D). RESULTS Patient n.1 was referred to PSMA-PET (Figure 2: A,E) for high-risk Pca (Gleason Score= 4+4; biopsy,
6/6 positive cores bilateral, initial PSA= 10 ng/mL). PSMA-PET showed pathological and
heterogeneous uptake diffusely comprising the left part of the prostate, without suspicion of extra-
capsular extension. Additionally, five PSMA-positive pelvic lymph nodes were detected. According
to E-PSMA reporting criteria (12) the patient was considered miT2N1M0. The day of surgery, 80
MBq of 68Ga-PSMA-11 was administered intravenously. The prostate gland and pelvic lymph nodes
were resected and all specimens were scanned in the PET/CT specimen imager. Scans were
performed sequentially, once the specimens were positioned in the container and properly oriented
by the pathologist, at 192 min (prostate) and 273 min (lymph nodes) post-injection. High-resolution
PET/CT images (Figure 2: C,G) showed higher spatial resolution compared to conventional PET
images. This indicates a potential for intraoperative margin assessment, complementary to
pathological analysis. Left basal margin’s involvement was strongly suspected on the prostate gland
image (Figure 2: C,D). All resected lymph nodes were scanned. Histopathology analysis was
performed and compared with the specimen PET/CT images (Figure 2: D,H). For the prostate gland, Patient n.1 was referred to PSMA-PET (Figure 2: A,E) for high-risk Pca (Gleason Score= 4+4; biopsy,
6/6 positive cores bilateral, initial PSA= 10 ng/mL). PSMA-PET showed pathological and
heterogeneous uptake diffusely comprising the left part of the prostate, without suspicion of extra-
capsular extension. Additionally, five PSMA-positive pelvic lymph nodes were detected. According
to E-PSMA reporting criteria (12) the patient was considered miT2N1M0. The day of surgery, 80
MBq of 68Ga-PSMA-11 was administered intravenously. The prostate gland and pelvic lymph nodes
were resected and all specimens were scanned in the PET/CT specimen imager. Scans were
performed sequentially, once the specimens were positioned in the container and properly oriented
by the pathologist, at 192 min (prostate) and 273 min (lymph nodes) post-injection. High-resolution
PET/CT images (Figure 2: C,G) showed higher spatial resolution compared to conventional PET
images. This indicates a potential for intraoperative margin assessment, complementary to
pathological analysis. Left basal margin’s involvement was strongly suspected on the prostate gland
image (Figure 2: C,D). All resected lymph nodes were scanned. Histopathology analysis was
performed and compared with the specimen PET/CT images (Figure 2: D,H). For the prostate gland, Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 A specipin was positioned in a region with high radiotracer
uptake in the specimen as seen on the high-resolution PET/CT images. The positioning of this pin
guided the pathologist to slice the specimen in the region of interest. This allowed an accurate
comparison between the 68Ga-DOTA-TOC uptake and the histopathological SSR immune-reactivity. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 All three methodologies (baseline PET/CT, specimen PET/CT and histopathology) were fully
concordant, and the lymph node was considered as a true positive finding. Table 1 and Table 2 summarize all semi-quantitative data about radiopharmaceuticals uptake,
administered activity and acquisition time both in whole-body PET/CT and high-resolution PET/CT
specimen imaging. Table 1 and Table 2 summarize all semi-quantitative data about radiopharmaceuticals uptake,
administered activity and acquisition time both in whole-body PET/CT and high-resolution PET/CT
specimen imaging. Finally, in all procedures the radiation exposure of the operating theater staff located near the
surgical table was measured using an Electronic Personal Dosimeters (Thermo Scientific™ EPD
Mk2™) and the absorbed dose was below 40 Sv (range 26 – 40 Sv) for each procedure. Figure 2 Patient-1 (prostate cancer): 68Ga-PSMA-11 whole-body PET/CT image showed pathological
PSMA uptake (high expression, E-PSMA Visual Score=3) in the prostate and in five pelvic lymph
nodes. In this figure are represented also high-resolution specimen PET/CT images and
histopathological analysis in the specimens were immunohistochemical analysis (PSMA staining)
was performed. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Figure 3 Patient-2 (prostate cancer): 68Ga-PSMA-11 whole-body PET/CT images showed pathological PSMA
uptake (high expression, E-PSMA Visual Score=3) in the prostate and in three pelvic lymph nodes. In
this figure are represented also high-resolution specimen PET/CT images and histopathological
analysis in the specimens were immunohistochemical analysis (PSMA staining) was performed. Patient-2 (prostate cancer): 68Ga-PSMA-11 whole-body PET/CT images showed pathological PSMA
uptake (high expression, E-PSMA Visual Score=3) in the prostate and in three pelvic lymph nodes. In
this figure are represented also high-resolution specimen PET/CT images and histopathological
analysis in the specimens were immunohistochemical analysis (PSMA staining) was performed. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Figure 4 Figure 4 Patient 3 (GEP-NET): 68Ga-DOTA-TOC whole-body PET/CT images showed pathological DOTA-TOC
uptake in one single peripancreatic lymph node. High-resolution specimen PET/CT images
confirmed the positivity and the presence metastasis from GEP-NET was confirmed by
histopathological analysis of the specimen. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Table 1 Semi-quantitative data measured both in pre-operative whole-body PET/CT and high-resolution
specimen PET/CT images. PET/CT SUVmax
Specimen Imager SUVmax
Pt-1 Prostate
26.5
89.2
Pt-1 Left Obturatory Nodes
11.4
51.5
Pt-2 Prostate
36.2
57.4
Pt-2 Right Common Iliac Node
11.8
23.2
Pt-2 Left Obturatory Nodes
25.9
36.4
Pt-3 Peri-pancreatic Node
52.1
86.6
SUVmax: Maximum Standardized Uptake Value DISCUSSION DISCUSSION DISCUSSION In this pilot analysis, first we assessed the feasibility of integrating this new generation imaging
approach in a live-surgery session. The collection and the scan of the specimens did not significantly
interfere with any procedure, and the overall length of the surgery was not affected using the
PET/CT specimen imager. Second, the radiation exposure generally measured for radio-guided
surgery using gamma emitting isotopes (e.g., 99mTc or 18F) is low and this technique is extensively
used in clinical practice. With 99mTc radiation doses per procedure to the surgeon’s chest is
comprised between 0.2–0.8 µSv per MBq of activity present at time of surgery, while for 18F
radiation exposures to the surgeon’s chest is comprised between 10 to 217 µSv for procedures. The
radiation exposure for the operating theater staff measured in our pilot analysis was low as well
(less than 40 Sv; range 26 – 40 Sv), confirming the feasibility of injecting positron emitting
isotopes (e.g., 68Ga or 18F) directly in the surgery theater. Finally, the specimen PET images were
reliable, and in full concordance with the information derived by the histopathological analysis. Moreover, they had a higher spatial resolution compared to conventional PET images. The
extraction of semiquantitative parameters (e.g., SUVmax or metabolic tumor volume) was also
feasible. At present, the implementation in clinical practice of this device has not been broadly explored. However, the development of PET/CT specimen imagers has a promising perspective and embraces
the integration of new generation imaging with the most updated surgery techniques. This new
approach might improve surgeon’s confidence in ensuring complete removal of the primary tumor. At present, the implementation in clinical practice of this device has not been broadly explored. However, the development of PET/CT specimen imagers has a promising perspective and embraces
the integration of new generation imaging with the most updated surgery techniques. This new
approach might improve surgeon’s confidence in ensuring complete removal of the primary tumor. Moreover, the precise correspondence of receptor-target radiopharmaceuticals uptake and
immuno-histochemistry staining confirmed the precision of this imaging procedure to evaluate
intra-lesion heterogeneity. Thus, the high spatial resolution of the PET specimen imager offers a
deeper understanding of the bio-distribution pattern of the radiotracers inside cancer lesions. In
this scenario, the use of a specimen imager might be an adjunct tool for the pathologist to better
optimize resources and for a “live” assessment of specific tumor biomarkers directly during surgery. Table 2 Administered activity and acquisition time differences between standard whole body PET/CT scans
and PET/CT specimen imager scans in each patient. Administered
activity
for
baseline PET
Acquisition time
Baseline PET
Administered
activity
for
PET/CT Specimen
Imager
Acquisition time
PET/CT Specimen
Imager
Patient-1
Pt. 1: 160 MBq
60 min
80 MBq
192-273 min
Patient-2
Pt. 2: 180 MBq
60 min
90 MBq
206-275 min
Patient-3
Pt. 3: 175 MBq
60 min
70 MBq
150 min Administered activity and acquisition time differences between standard whole body PET/CT scans
and PET/CT specimen imager scans in each patient. Administered activity and acquisition time differences between standard whole body PET/CT scans
and PET/CT specimen imager scans in each patient. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 the PET/CT specimen imager might detect micro-metastasis. Furthermore, in patient 1 extra-
capsular extension was not suspected in baseline PSMA PET/CT. However, extra-capsular extension
was observed with the specimen imager (and subsequently confirmed by histology), thus hinting at
a potential superior sensitivity compared to the whole-body PET images used in clinical practice. Since surgical margin status is an important prognostic factor in PCa, this is a topic of particular
interest. At present, even more deeply studied techniques for surgical margins assessment such as
intraoperative frozen section (IFS) for the detection of positive surgical margins in PCa, did not show
a significant advantage in terms of oncological outcomes and have still sub-optimal accuracy (13). In clinical practice, new generation molecular imaging has already proven its crucial role in the
decision-making process of several malignancies, including PCa and NET, and the transition from
pre-clinical imaging to clinical practice has been quick and clinically significant. However, new
radiopharmaceuticals are emerging beyond PSMA and SSR based PET, like inhibitors of the fibroblast
activating protein (FAPi PET) or ligand of the chemokine receptor 4 CXCR4 (Pentixafor PET). All these
new radiopharmaceuticals will increase the number of pathways that can be explored with in vivo
molecular imaging, and it will increase the understanding of tumor heterogeneity both in primary
tumor and in metastatic locations. In this scenario, intra-operative PET/CT specimen imaging offers
a link between molecular biology, histopathology assessment, molecular imaging and radio-guided
surgery. DISCUSSION Another potential advantage might be related to the high negative predictive value (NPV) of this
imaging procedure. In patient 1 and 2, histopathology confirmed the lymph node showing high
PSMA expression as true positive findings. However, also PSMA negative nodes were included in the
specimen container. The histopathological analysis confirmed the lack of malignant cells in these
lymph nodes, thus resulting as true negative findings. The future challenge will be understanding if LIMITATIONS AND FUTURE PERSPECTIVES This is a pure explorative analysis, as this was the first experience with a PET/CT specimen imager
used in prostate cancer and a neuroendocrine tumor in a live-surgery setting. Due to the limited
number of patients (n=3), this analysis only provides only a first indication of the possibilities of
intraoperative specimen PET/CT imaging for prostate cancer and neuroendocrine tumors. Extensive
future research is needed to further investigate the clinical value of this technique. In this
perspective, a deeper understanding of radiotracers’ uptake pattern inside surgical specimens
through a PET/CT specimen imager could open newer research pathways that range from imaging
(e.g detection of surgical margins’ involvement) to pathology (e.g. correlation with
immunohistochemistry), to clinical oncology (e.g. changes in uptake patterns after treatment). Finally, nor comparative analysis with other technique (e.g. mp-MRI) or an evaluation of the impact
on clinical management of the patients cannot be performed at this stage. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 CONCLUSION The use of the PET/CT specimen imager in a live-experience session was feasible and its application
has been easily adapted to surgery daily-practice. In this pilot analysis, the image acquisition of
specimens obtained by patients who received an intra-surgical injection of 68Ga-PSMA-11 and
68Ga-DOTA-TOC was feasible and reliable, and imaging findings matched perfectly with
histopathological analysis. The high sensitivity, together with the evaluation of intra-lesion tumor
heterogeneity, were the most relevant results derived from this pilot analysis. Even if far from
definitive evidence, this pilot analysis showed promising results that need to be confirmed by
further prospective studies in larger cohorts of oncological patients. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444.v2 REFERENCES 1) Kapoor V, McCook BM, Torok FS. An introduction to PET-CT imaging. Radiographics. 2004 Mar-
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Calais J. Diagnostic Accuracy of 68Ga-PSMA-11 PET for Pelvic Nodal Metastasis Detection
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Smith R, Small EJ, Slavik R, Carroll PR, Herrmann K, Czernin J, Hope TA. Assessment of 68Ga-PSMA-
11 PET Accuracy in Localizing Recurrent Prostate Cancer: A Prospective Single-Arm Clinical Trial. JAMA Oncol. 2019 Jun 1;5(6):856-863. doi: 10.1001/jamaoncol.2019.0096. PMID: 30920593; PMCID:
PMC6567829. 11) Bozkurt MF, Virgolini I, Balogova S, Beheshti M, Rubello D, Decristoforo C, Ambrosini V, Kjaer
A, Delgado-Bolton R, Kunikowska J, Oyen WJG, Chiti A, Giammarile F, Sundin A, Fanti S. Guideline for PET/CT imaging of neuroendocrine neoplasms with 68Ga-DOTA-conjugated
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33604691; PMCID: PMC8113168. 12) Ceci F, Oprea-Lager DE, Emmett L, Adam JA, Bomanji J, Czernin J, Eiber M, Haberkorn U,
Hofman MS, Hope TA, Kumar R, Rowe SP, Schwarzenboeck SM, Fanti S, Herrmann K. E-PSMA:
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33604691; PMCID: PMC8113168. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 doi:10.20944/preprints202212.0444 Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 CONCLUSION 406 doi:10.1016/S0140-6736(20)30314-7 407; 7) Emmett L, Buteau J, Papa N, Moon D, Thompson J, Roberts MJ, et al. The Additive Diagnostic
408 Value of Prostate-specific Membrane Antigen Positron Emission Tomography Computed
409 Tomography to Multiparametric Magnetic Resonance Imaging Triage in the Diagnosis of
Prostate 410 Cancer (PRIMARY): A Prospective Multicentre Study. European Urology. 2021;80(6):682-9. doi: 411 10.1016/j.eururo.2021.08.002; 8) Calais J, Ceci F, Eiber M, Hope TA, Hofman MS, Rischpler C, Bach-Gansmo T, Nanni C, Savir-Baruch B,
Elashoff D, Grogan T, Dahlbom M, Slavik R, Gartmann J, Nguyen K, Lok V, Jadvar H, Kishan AU, Rettig
MB, Reiter RE, Fendler WP, Czernin J. 18F-fluciclovine PET-CT and 68Ga-PSMA-11 PET-CT in patients
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comparative imaging trial. Lancet Oncol. 2019 Sep;20(9):1286-1294. doi: 10.1016/S1470- Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 2 February 2023 13) Dinneen EP, Van Der Slot M, Adasonla K, Tan J, Grierson J, Haider A, Freeman A, Oakley N, Shaw G. Intraoperative Frozen Section for Margin Evaluation During Radical Prostatectomy: A Systematic
Review. Eur Urol Focus. 2020 Jul 15;6(4):664-673. doi: 10.1016/j.euf.2019.11.009. Epub 2019 Nov 29. PMID: 31787570. 13) Dinneen EP, Van Der Slot M, Adasonla K, Tan J, Grierson J, Haider A, Freeman A, Oakley N, Shaw G. Intraoperative Frozen Section for Margin Evaluation During Radical Prostatectomy: A Systematic
Review. Eur Urol Focus. 2020 Jul 15;6(4):664-673. doi: 10.1016/j.euf.2019.11.009. Epub 2019 Nov 29. PMID: 31787570.
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Study on <i>r</i>-truncated degenerate Stirling numbers of the second kind
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* Corresponding author: Taekyun Kim, Department of Mathematics, Kwangwoon University, Seoul 139-701, Republic of Korea,
e-mail: tkkim@kw.ac.kr
Dae San Kim: Department of Mathematics, Sogang University, Seoul 121-742, Republic of Korea, e-mail: dskim@sogang.ac.kr
Hyekyung Kim: Department of Mathematics Education, Daegu Catholic University, Gyeongsan 38430, Republic of Korea,
e-mail: hkkim@cu.ac.kr Taekyun Kim*, Dae San Kim, and Hyekyung Kim
Study on r-truncated degenerate Stirling
numbers of the second kind Taekyun Kim*, Dae San Kim, and Hyekyung Kim https://doi.org/10.1515/math-2022-0535
received June 30, 2022; accepted November 24, 2022 https://doi.org/10.1515/math-2022-0535
received June 30, 2022; accepted November 24, 2022 Abstract: The degenerate Stirling numbers of the second kind and of the first kind, which are, respectively,
degenerate versions of the Stirling numbers of the second kind and of the first kind, appear frequently when
we study various degenerate versions of some special numbers and polynomials. The aim of this article is to
consider the r-truncated degenerate Stirling numbers of the second kind, which reduce to the degenerate
Stirling numbers of the second for
=
r
1, and to investigate their explicit expressions, some properties and
related identities, in connection with several other degenerate special numbers and polynomials. Keywords: r-truncated degenerate Stirling numbers of the second kind, r-truncated degenerate Bernoulli
polynomials of order alpha MSC 2020: 11B73, 11B83 Open Access. © 2022 the author(s), published by De Gruyter.
This work is licensed under the Creative Commons Attribution 4.0
International License. Open Mathematics 2022; 20: 1685–1695 is obtained by truncating the first r terms of the degenerate exponential function (see (8), (10), (35)). In the
same way, the r-truncated degenerate Bernoulli polynomials of order α are defined by truncating the first r
terms of the degenerate exponential function in their generating function (see (17)). In some works related
to truncated degenerate special numbers and polynomials, we let the reader refer to [4,6,7,15] and refer-
ences therein. The aim of this article is by using generating functions to study their explicit expressions, some proper-
ties and related identities on the r-truncated degenerate Stirling numbers of the second kind, in connection
with the r-truncated degenerate Bernoulli polynomials of order α, the degenerate Bernoulli numbers,
the degenerate Stirling numbers of the second kind and the degenerate Bernoulli numbers of order α. The outline of this article is as follows. In Section 1, we recall the degenerate exponentials and loga-
rithms. We remind the reader of the degenerate Stirling numbers of both kinds. Furthermore, we recall the
degenerate Bernoulli polynomials of order α. Section 2 gives the main results of this article. We consider the
r-truncated degenerate Stirling numbers of the second kind and find three explicit expressions for those
numbers in Theorems 1–3. Then we introduce the r-truncated degenerate Bernoulli polynomials of order α
and obtain some results in connection with the r-truncated degenerate Stirling numbers of the second kind. In Theorem 4, the degenerate Bernoulli numbers are expressed in terms of the Stirling numbers of the
second kind. In Theorems 5 and 8, some identities involving the r-truncated degenerate Stirling numbers of
the second kind (for r = 2 and r = 3, respectively), the degenerate Bernoulli numbers and the degenerate
Stirling numbers of the second are obtained. In Theorem 6, we find an identity connecting the 2-truncated
degenerate Stirling numbers of the second kind and the degenerate Bernoulli numbers. In Theorem 7, we
obtain an identity relating the 2-truncated degenerate Stirling numbers of the second kind and the degen-
erate Bernoulli numbers of order α. In the rest of this section, we recall the facts that are needed throughout
this article. For any nonzero
∈
λ
, the degenerate exponentials are defined by he degenerate exponentials are defined by For any nonzero
∈
λ
, the degenerate exponentials are defined by ( )
( )
(
)
(
)
∑
=
! 1 Introduction Carlitz [1,2] initiated to study of the degenerate Bernoulli and Euler polynomials and numbers, which are
degenerate versions of the Bernoulli and Euler polynomials and numbers. In recent years, studying degen-
erate versions of some special numbers and polynomials has regained interests of some mathematicians
and yielded quite a few interesting results (see [3–11] and references therein). It is noteworthy that studying
degenerate versions is not only limited to polynomials but also extended to transcendental functions, like
gamma functions (see [12,13]). It is also remarkable that the degenerate umbral calculus is introduced as a
degenerate version of the classical umbral calculus (see [14]). Degenerate versions of special numbers and
polynomials have been explored by various methods, including combinatorial methods, generating func-
tions, umbral calculus techniques, p-adic analysis, differential equations, special functions, probability
theory and analytic number theory. The Stirling number of the second
(
)
S n k
,
2
is the number of ways to partition a set of n objects into k
nonempty subsets. The (signed) Stirling number of the first kind
(
)
S n k
,
1
is defined so that the number of
permutations of n elements having exactly k cycles is the nonnegative integer (
)
(
)
∣(
)∣
−
=
−S n k
S n k
1
,
,
n k
1
1
. The degenerate Stirling numbers of the second kind
(
)
S
n k
,
λ
2,
(see (5), (8)) and of the first kind
(
)
S
n k
,
λ
1,
(see
(4), (7)) were introduced as degenerate versions of the Stirling numbers of the second and of the first kind,
respectively. These degenerate Stirling numbers of both kinds appear frequently when we study degenerate
versions of some special numbers and polynomials. Here we consider the r-truncated degenerate Stirling numbers of the second which reduce to the
degenerate Stirling numbers of the second for
=
r
1. We note that the generating function of these numbers 1686
Taekyun Kim et al. 1686 λ
α
λ
x
n
n λ
α
n
0
,
(3) (3) When
=
x
0,
( )
( )
( )
=
β
β
0
n λ
α
n λ
α
,
,
are called the degenerate Bernoulli numbers of order α. In particular, for
=
α
1,
( )
( )
( )
=
β
x
β
x
n λ
n λ
,
,
1
are called the degenerate Bernoulli polynomials. For
≥
n
0, the degenerate Stirling numbers of the first kind are defined by When
=
x
0,
( )
( )
( )
=
β
β
0
n λ
α
n λ
α
,
,
are called the degenerate Bernoulli numbers of order α. In particular, for
=
α
1,
( )
( )
( )
=
β
x
β
x
n λ
n λ
,
,
1
are called the degenerate Bernoulli polynomials. For
≥
n
0, the degenerate Stirling numbers of the first kind are defined by When
=
x
0,
( )
( )
( )
=
β
β
0
n λ
α
n λ
α
,
,
are called the degenerate Bernoulli numbers of order α. In particular, for
=
α
1,
( )
( )
( )
=
β
x
β
x
n λ
n λ
,
,
1
are called the degenerate Bernoulli polynomials. For
≥
n
0, the degenerate Stirling numbers of the first kind are defined by ( )
(
)( )
(
[ ])
∑
=
=
x
S
n k x
,
see 3 ,
n
k
n
λ
k λ
0
1,
,
(4) (4) where ( )
( )
(
)
(
)
(
)
(
[
])
=
=
−
⋯
−
+
≥
x
x
x x
x
n
n
1,
1
1 ,
1
see 1–11, 14–22 . n
0 formula of (4), the degenerate Stirling numbers of the second kind are defined by As the inversion formula of (4), the degenerate Stirling numbers of the second kind ar ( )
(
)( )
(
)
(
[ ])
∑
=
≥
=
x
S
n k x
n
,
,
0
see 3 . =
+
[
]
=
∞
e
t
x
k
t
λt
1
see 3–7
,
λ
x
k
k λ
k
0
,
x
λ
(1)
( )
( )
(
)(
)
(
(
) )
(
)
=
=
−
−
⋯
−
−
≥
x
x
x x
λ x
λ
x
n
λ
n
1,
2
1
,
1 . λ
n λ
0,
,
(2) ( )
( )
(
)
(
)
∑
=
! =
+
[
]
=
∞
e
t
x
k
t
λt
1
see 3–7
,
λ
x
k
k λ
k
0
,
x
λ
(1)
( )
(
)(
)
(
(
) )
(
)
≥
x
x x
λ x
λ
x
n
λ
n
2
1
1
(2) ( )
( )
(
)
(
)
∑
=
! =
+
[
]
=
∞
e
t
x
k
t
λt
1
see 3–7
,
λ
x
k
k λ
k
0
,
x
λ
(1) (1) ( )
( )
(
)(
)
(
(
) )
(
)
=
=
−
−
⋯
−
−
≥
x
x
x x
λ x
λ
x
n
λ
n
1,
2
1
,
1 . λ
n λ
0,
,
(2) (2) 1,
( )
( )
(
)
=
=
+
e t
e t
λt
1
λ
λ
1
λ
1 (see [8,15]). When
=
x
1,
( )
( )
(
)
=
=
+
e t
e t
λt
1
λ
λ
1
λ
1 (see [8,15]). When
=
x
1,
( )
( )
(
)
=
=
+
e t
e t
λt
1
λ
λ
1
λ
1 (see [8,15]). In [1,2], Carlitz considered the degenerate Bernoulli polynomials of order α given by ( )
( )
(
)
λ
λ
(
[
])
In [1,2], Carlitz considered the degenerate Bernoulli polynomials of order α given by ⎜
⎟
⎛
⎝
( )
⎞
⎠
( )
( )
( )
∑
−
=
! =
∞
t
e t
e
t
β
x t
n
1
. 2 r-Truncated degenerate Stirling numbers of the second kind ! ⋯
! + +⋯+
=
≥
S
n kr
k
n
l l
l
,
1
1
1
1
. λ
r
l
l
l
n
l
r
l λ
l λ
l λ
k
2,
,
,
,
1
2
k
i
k
1
2
1
2 Theorem 1. For
≥
n k
,
0 with
≥
n
kr, we have Theorem 1. For
≥
n k
,
0 with
≥
n
kr, we have Theorem 1. For
≥
n k
,
0 with
≥
n
kr, we have (
)
( )
( )
( )
[ ]
∑
=
! ! ⋯
! ! ⋯
! + +⋯+
=
≥
S
n kr
k
n
l l
l
,
1
1
1
1
. λ
r
l
l
l
n
l
r
l λ
l λ
l λ
k
2,
,
,
,
1
2
k
i
k
1
2
1
2 From the binomial expansion, we note that ⎜
⎟
⎜
⎟
⎛
⎝
( )
( )
⎞
⎠
⎛
⎝
⎞
⎠
( )(
) ⎛
⎝
( )
⎞
⎠
⎛
⎝
⎞
⎠
(
)
(
)
( )
( )
( )
⎛
⎝
⎜
⎛
⎝
⎞
⎠
(
)
( )
( )
( )
(
)
(
)
⎞
⎠
⎟
∑
∑
∑
∑
∑
∑
∑
∑
∑
! −
! =
! −
! =
! −
! −
⋯
! ! ⋯
! =
! −
! ⋯
−
! ! ⋯
! −
−⋯−
! ! =
−
=
−
=
−
=
=
∞
…
=
−
+⋯+
=
∞
=
…
=
−
−−−⋯−
k
e t
l
t
k
k
m e
t
l
t
k
k
m
k
m
t
j
l l
l
t
k
k
m
n
k
m
l l
l
n
l
l
t
n
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
. n
n
(9) Let
( )
[[ ]]
= ∑
∈
=
∞
f t
a t
t
k
k k
0
. For
≥
n
0, the operator [ ]
tn is defined by
[ ] ( )
(
)
(
[
])
=
≥
t
f t
a
n
,
0
see
22 . n
n
(9) Let
( )
[[ ]]
= ∑
∈
=
∞
f t
a t
t
k
k k
0
. For
≥
n
0, the operator [ ]
tn is defined by
[ ] ( )
(
)
(
[
])
=
≥
t
f t
a
n
,
0
see
22 . n
n
(9) (9) n λ
k
n
λ
k
,
0
2,
(5) (5) The degenerate logarithm
( )t
logλ
is the compositional inverse of
( )
e t
λ
satisfying
(
( ))
(
( ))
=
e
t
e t
log
log
λ
λ
λ
λ
. Then we have The degenerate logarithm
( )t
logλ
is the compositional inverse of
( )
e t
λ
satisfying
(
( ))
(
( ))
=
e
t
e t
log
log
λ
λ
λ
λ
. Then we have Study on r-truncated degenerate Stirling numbers of the second kind
1687 Study on r-truncated degenerate Stirling numbers of the second kind
1687 (
)
((
)
)
( )
(
[ ])
∑
+
=
+
−
=
! =
∞
−
t
λ
t
λ
n
t
log 1
1
1
1
1
see 3 . λ
λ
n
n
n
n
1
1
, λ
1
(6) (6) From (4) and (5), we note that From (4) and (5), we note that From (4) and (5), we note that
(
(
))
(
)
(
)
∑
! +
=
! ≥
=
∞
k
t
S
n k t
n
k
1
log 1
,
,
0
λ
k
n k
λ
n
1,
(7) (
(
))
(
)
(
)
∑
! +
=
! ≥
=
∞
k
t
S
n k t
n
k
1
log 1
,
,
0
λ
k
n k
λ
n
1,
(7) (7) ! ! =
k
n
n k
and
(
( )
)
(
)
(
[ ])
∑
! −
=
! =
∞
k
e t
S
n k t
n
1
1
,
see
3 . λ
k
n k
λ
n
2,
(8) and (
( )
)
(
)
(
[ ])
∑
! −
=
! =
∞
k
e t
S
n k t
n
1
1
,
see
3 . λ
k
n k
λ
n
2,
(8) (8) Let
( )
[[ ]]
= ∑
∈
=
∞
f t
a t
t
k
k k
0
. For
≥
n
0, the operator [ ]
tn is defined by
[ ] ( )
(
)
(
[
])
=
≥
t
f t
a
n
,
0
see
22 . 2 r-Truncated degenerate Stirling numbers of the second kind For
∈
r
, we consider the r-truncated degenerate Stirling numbers of the second kind given by ⎜
⎟
⎛
⎝
( )
( )
⎞
⎠
(
)
(
)
[ ]
∑
∑
! −
! =
! ≥
=
−
=
∞
k
e t
l
t
S
n kr t
n
k
1
1
,
,
0 . λ
l
r
l λ
l
k
n kr
λ
r
n
0
1
,
2,
(10) (10) (10) We agree that
(
)
[ ]
=
S
n kr
,
0
λ
r
2,
, for
≤
<
n
kr
0
. Note that
(
)
(
)
(
)
[ ]
=
≥
S
n k
S
n k
n k
,
,
,
,
0
λ
λ
2,
1
2,
. From (10), we note that We agree that
(
)
[ ]
=
S
n kr
,
0
λ
r
2,
, for
≤
<
n
kr
0
. Note that
(
)
(
)
(
)
[ ]
=
≥
S
n k
S
n k
n k
,
,
,
,
0
λ
λ
2,
1
2,
. From (10), we note that ⎜
⎟
⎜
⎟
⎛
⎝
( )
( )
⎞
⎠
⎛
⎝
( )
⎞
⎠
⎛
⎝
⎜
⎜⎜
( )
( )
( )
⎞
⎠
⎟
⎟⎟
∑
∑
∑
∑
! −
! =
! ! =
! ! ⋯
! ! ⋯
! ! =
−
=
∞
=
∞
+ +⋯+
=
≥
k
e t
l
t
k
l
t
k
n
l l
l
t
n
1
1
1
1
1
1
1
1
. λ
l
r
l λ
l
k
l r
k λ
l
k
n kr
l
l
l
n
l
r
l λ
l λ
l λ
k
n
0
1
,
,
,
,
,
1
2
k
i
k
1
2
1
2
(11) (11) ⎝
⎠
≥
l
r
i
Thus, by (9) and (11), we obtain the following theorem. Thus, by (9) and (11), we obtain the following theorem. Thus, by (9) and (11), we obtain the following theorem. heorem 1. For
≥
n k
,
0 with
≥
n
kr, we have
(
)
( )
( )
( )
[ ]
∑
=
! ! ⋯
! Therefore, by comparing the coefficients on both sides of (12), we obtain the following theorem. Theorem 2. For
≥
n k
,
0, we have Theorem 2. For
≥
n k
,
0, we have Theorem 2. For
≥
n k
,
0, we have Theorem 2. For
≥
n k
,
0, we have ⎛
⎝
⎞
⎠
(
)
( )
( )
( )
(
)
(
)
⎧
⎨⎩
(
)
[ ]
∑
∑
! −
! ⋯
−
! ! ⋯
! −
−⋯−
! =
≥
≤
<
=
…
=
−
−−−⋯−
k
k
m
n
k
m
l l
l
n
l
l
S
n kr
if n
kr
if
n
kr
1
1
1
1
1
,
,
,
0,
0
. m
k
m
l l
l
r
l λ
l λ
l
λ
n l
l
l
m
m
λ
r
0
, ,
,
0
1
,
,
,
1
2
1
2,
m
m
m λ
1 2
1
2
1
2
, When
=
k
1 in (10), we have ⎜
⎟
( )
⎛
⎝
( )
( )
⎞
⎠
(
)
(
)(
)
(
) (
)
[ ]
[ ]
[ ]
∑
∑
∑
∑
∑
! =
−
! =
! =
+
+
! =
+
! +
! ! =
∞
=
−
=
∞
=
∞
+
=
∞
l
t
e t
l
t
S
n r t
n
S
n
r r
t
n
r
t
S
n
r r
n
n
r
t
n
1
1
,
,
,
. l r
l λ
l
λ
l
r
l λ
l
n r
λ
r
n
n
λ
r
n r
r
n
λ
r
n
,
1
1
,
2,
0
2,
0
2,
(13) (13) For
=
k
2, we have For
=
k
2, we have For
=
k
2, we have For
=
k
2, we have ⎜
⎟
⎜
⎟
⎛
⎝
( )
⎞
⎠
⎛
⎝
⎜
(
)(
)
⎞
⎠
⎟
⎛
⎝
(
)(
)
⎞
⎠
⎛
⎝
⎜
(
)
(
) (
) (
)
⎞
⎠
⎟
[ ]
[ ]
[ ]
[ ]
∑
∑
∑
∑∑
! =
+
+
! +
+
! 2 r-Truncated degenerate Stirling numbers of the second kind λ
l
r
l λ
l
k
m
k
λ
k m
m
l
r
l λ
l
m
m
k
j
j λ
j
m
l l
l
r
l λ
l λ
l
λ
m
l
l
n
m
k
m
l l
l
r
l λ
l λ
l
λ
n l
l
l
m
m
n
0
1
,
0
0
1
,
0
0
,
, ,
,
0
1
,
,
,
1
2
0
0
, ,
,
0
1
,
,
,
1
2
1
m
m
m
m
m
m λ
1 2
1
2
1
1 2
1
2
1
2
,
(12) (12) Taekyun Kim et al. 1688
Taekyun Kim et al. 1688
Taekyun Kim et al. 1688 =
+
−
+
! −
+
! +
! ! =
∞
=
∞
=
∞
=
∞
=
l
t
t
S
j
r r
t
j
r
S
l
r r
t
l
r
t
S
j
r r S
n
j
r r
n
n
j
r
j
r
t
n
1
,
,
,
,
. l r
l λ
l
r
j
λ
r
j
l
λ
r
l
r
n
j
n
λ
r
λ
r
n
,
2
2
0
2,
0
2,
2
0
0
2,
2,
(14) ⎜
⎟
⎜
⎟
⎛
⎝
( )
⎞
⎠
⎛
⎝
⎜
(
)(
)
⎞
⎠
⎟
⎛
⎝
(
)(
)
⎞
⎠
[ ]
[ ]
∑
∑
∑
! =
+
+
! +
+
! =
∞
=
∞
=
∞
l
t
t
S
j
r r
t
j
r
S
l
r r
t
l
r
1
,
,
l r
l λ
l
r
j
λ
r
j
l
λ
r
l
,
2
2
0
2,
0
2, (14) Continuing this process, we have Continuing this process, we have ⎜
⎟
⎛
⎝
( )
⎞
⎠
⎛
⎝
⎜
(
)
(
)
(
)
(
) (
)
(
)
⎞
⎠
⎟
[ ]
[ ]
[ ]
∑
∑
∑
! =
! +
+
⋯
+
+
! +
! ⋯
+
! ! =
∞
=
∞
+ +⋯+
=
l
t
t
n S
j
r r S
j
r r
S
j
r r
j
r
j
r
j
r
t
n
1
,
,
,
. l r
l λ
l
k
kr
n
j
j
j
n
λ
r
λ
r
λ
r
k
k
n
,
0
2,
1
2,
2
2,
1
2
k
1
2
(15) ⎜
⎟
⎛
⎝
( )
⎞
⎠
⎛
⎝
⎜
(
)
(
)
(
)
(
) (
)
(
)
⎞
⎠
⎟
[ ]
[ ]
[ ]
∑
∑
∑
! =
! +
+
⋯
+
+
! +
! ⋯
+
! ! =
∞
=
∞
+ +⋯+
=
l
t
t
n S
j
r r S
j
r r
S
j
r r
j
r
j
r
j
r
t
n
1
,
,
,
. l r
l λ
l
k
kr
n
j
j
j
n
λ
r
λ
r
λ
r
k
k
n
,
0
2,
1
2,
2
2,
1
2
k
1
2
(15) ⎜
⎟
⎛
⎝
( )
⎞
⎠
⎛
⎝
⎜
(
)
(
)
(
)
(
) (
)
(
)
⎞
⎠
⎟
[ ]
[ ]
[ ]
∑
∑
∑
! =
! +
+
⋯
+
+
! +
! ⋯
+
! ! =
∞
=
∞
+ +⋯+
=
l
t
t
n S
j
r r S
j
r r
S
j
r r
j
r
j
r
j
r
t
n
1
,
,
,
. l r
l λ
l
k
kr
n
j
j
j
n
λ
r
λ
r
λ
r
k
k
n
,
0
2,
1
2,
2
2,
1
2
k
1
2
(15) (15) On the other hand, by (10), we obtain On the other hand, by (10), we obtain On the other hand, by (10), we obtain ⎜
⎟
⎜
⎟
⎛
⎝
( ) ⎞
⎠
⎛
⎝
( )
( )
⎞
⎠
(
)
(
)(
)
(
)
(
)
[ ]
[ ]
[ ]
∑
∑
∑
∑
∑
! =
−
! =
! ! =
! +
+
! =
! +
+
! =
∞
=
−
=
∞
=
∞
+
=
∞
t
l
e t
l
t
k
S
n kr t
n
k
S
n
kr kr
t
n
kr
t
k S
n
kr kr
n
kr
t
1
1
,
,
,
. l r
l
l λ
k
λ
l
r
l λ
l
k
n kr
λ
r
n
n
λ
r
n kr
kr
n
λ
r
n
,
1
1
,
2,
0
2,
0
2,
(16) (16) Therefore, by (15) and (16), we obtain the following theorem. Therefore, by (15) and (16), we obtain the following theorem. Therefore, by (15) and (16), we obtain the following theorem. Theorem 3. For
≥
n k
,
0, we have
(
)
(
)
(
)
(
)
[ ]
[ ]
[ ]
[ ]
∑
! +
=
+
+
⋯
+
k
S
n
kr kr
S
j
r r S
j
r r
S
j
r r
,
,
,
r
λ
r
λ
r
λ
r
k
2,
1
2,
2
2, (
)
(
)
(
)
(
)
(
)
(
) (
)
(
)
[ ]
[ ]
[ ]
[ ]
∑
! +
! +
=
+
+
⋯
+
+
! +
! ⋯
+
! + +⋯+
=
k
n
kr
S
n
kr kr
S
j
r r S
j
r r
S
j
r r
j
r
j
r
j
r
,
,
,
,
. λ
r
j
j
j
n
λ
r
λ
r
λ
r
k
k
2,
2,
1
2,
2
2,
1
2
k
1
2 Let us consider the r-truncated degenerate Bernoulli polynomials of order α given by (
( )
( ) )
( )
( )
[
]
∑
−∑
=
! =
−
! =
∞
−
t
e t
e
t
β
x t
n
1
. αr
λ
l
r
t
l
l λ α
λ
x
n
n λ
r
α
n
0
1
,
0
,
1,
l
(17) (17) When
=
x
0,
( )
[
]
[
]
=
−
−
β
β
0
n λ
r
α
n λ
r
α
,
1,
,
1,
are called the r-truncated degenerate Bernoulli numbers of order α. Study on r-truncated degenerate Stirling numbers of the second kind
1689 ⎜
⎟
⎜
⎟
⎛
⎝
( )
( ) ⎞
⎠
(
)
⎛
⎝
⎜
(
)
(
)
⎞
⎠
⎟
⎛
⎝
⎛
⎝
⎞
⎠
(
)
⎞
⎠
[
]
[ ]
[
]
[ ]
[
]
[ ]
[
]
∑
∑
∑
∑
∑
∑
∑
∑
=
−
! ! =
! ! ! =
! −
! −
! ! ! =
! −
! =
−
=
∞
−
=
∞
=
∞
−
=
∞
=
−
−
=
∞
=
−
−
t
e t
t
l
β
t
l
α
S
j αr t
j
β
t
l
α
S
n
l αr n
n
l l
β
t
n
α
n
l S
n
l αr β
t
n
1
,
,
,
,
αr
λ
l
r
l
l λ
α
l
l λ
r
α
l
j αr
λ
r
j
l
l λ
r
α
l
n αr
l
n αr
λ
r
l λ
r
α
n
n αr
l
n αr
λ
r
l λ
r
α
n
0
1
,
0
,
1,
2,
0
,
1,
0
2,
,
1,
0
2,
,
1,
(18) ⎜
⎟
⎜
⎟
⎛
⎝
( )
( ) ⎞
⎠
(
)
⎛
⎝
⎜
(
)
(
)
⎞
⎠
⎟
⎛
⎝
⎛
⎝
⎞
⎠
(
)
⎞
⎠
[
]
[ ]
[
]
[ ]
[
]
[ ]
[
]
∑
∑
∑
∑
∑
∑
∑
∑
=
−
! ! =
! ! ! =
! −
! −
! ! ! =
! −
! =
−
=
∞
−
=
∞
=
∞
−
=
∞
=
−
−
=
∞
=
−
−
t
e t
t
l
β
t
l
α
S
j αr t
j
β
t
l
α
S
n
l αr n
n
l l
β
t
n
α
n
l S
n
l αr β
t
n
1
,
,
,
,
αr
λ
l
r
l
l λ
α
l
l λ
r
α
l
j αr
λ
r
j
l
l λ
r
α
l
n αr
l
n αr
λ
r
l λ
r
α
n
n αr
l
n αr
λ
r
l λ
r
α
n
0
1
,
0
,
1,
2,
0
,
1,
0
2,
,
1,
0
2,
,
1, (18) where α is a positive integer. Thus, by (18), we obtain ⎛
⎝
⎞
⎠
(
)
⎧
⎨
⎩
(
)
[ ]
[
]
∑
−
=
! ! =
>
=
−
−
n
l S
n
l αr β
αr
α
n
αr
n
αr
,
,
if
,
0,
if
. n
k
n k
n k
n k
1
2
1
1
2
1
1
2
1 {(
)∣
(
)
}
=
…
+
+ ⋯+
=
+
+ ⋯+
−
+
=
−+
−+
−+
k k
k
k
k
k
k
k
k
n
k
k
n
Λ
,
,
,
,
2
1
. n
k
n k
n k
n k
1
2
1
1
2
1
1
2
1
h {(
)∣
(
)
}
=
…
+
+ ⋯+
=
+
+ ⋯+
−
+
=
−+
−+
−+
k k
k
k
k
k
k
k
k
n
k
k
n
Λ
,
,
,
,
2
1
. n
k
n k
n k
n k
1
2
1
1
2
1
1
2
1
h We observe that ⎜
⎟
⎜
⎟
( )
( )
( )
(
)
⎛
⎝
( )
⎞
⎠
(
)
( ( )
( )
( )
)
⎛
⎝
(
)
( ( )
( )
( )
)⎞
⎠
∑
∑
∑
∑
∑∑
+
+
+ ⋯+
+ ⋯
=
−
! ! ! =
+
−
! …
! =
+
−
! …
! ! ! ! =
∞
=
∞
=
∞
=
∞
−+
−+
=
∞
=
−+
t
t
t
k k
x
l
t
k
B
x
x
x
t
n
k B
x
x
x
t
n
1
1
1
1
1
1
1
1
1
1
1
,
1
,
,
1
1
1
1
,
1
,
,
. x
λ
x
λ
x
k
k λ k
k
k
l
l
l λ
l
k
k
k
n k
n k
λ
λ
n k
n k
n
n
k
n
k
n k
λ
λ
n k
λ
n
1
1,
2
2,
2
,
0
1
,
1
,
1
1,
2
2,
1
1
1
1
,
1
1,
2
2,
1,
k
1
2
(19) ⎜
⎟
⎜
⎟
(
)
⎝
⎠
(
)
( ( )
( )
( )
)
⎛
⎝
(
)
( ( )
( )
( )
)⎞
⎠
∑
∑
∑
∑
∑∑
=
−
! ! ! =
+
−
! …
! =
+
−
! …
! l
n αr
λ
r
l λ
r
α
0
2,
,
1, It is well known that the partial Bell polynomials are defined by It is well known that the partial Bell polynomials are defined by ⎜
⎟
⎛
⎝
⎞
⎠
(
)
∑
∑
! ! =
…
! =
∞
=
∞
−+
k
x t
l
B
x x
x
t
n
1
,
,
,
,
l
l
l
k
n k
n k
n k
n
1
,
1
2
1 where k is a nonnegative integer. Thus, we note that Thus, we note that ⎜
⎟
⎜
⎟
(
)
⎛
⎝
⎞
⎠
⎛
⎝
⎞
⎠
⎛
⎝(
)
⎞
⎠
⎛
⎝
⎞
⎠
(
)
∑
∑∏
…
=
! ! ! ⋯
! ! ! ⋯
−
+
! =
! ! ! −+
+ +⋯+
=
+
+⋯+
−+
=
−+
−+
=
−+
−+
−+
−+
B
x x
x
n
l l
l
x
x
x
n
k
n
k
x
j
,
,
,
1
2
1
1
,
n k
n k
l
l
l
k
l
l
n k
l
n
n k
l
l
n k
l
j
n k
j
j
k
,
1
2
1
2
1
1 2
1
1
2
1
Λ
1
1
n k
n k
n k
n
k
j
1
2
1
1
2
1
1
2
1 ⎜
⎟
⎛
⎝
⎞
⎠
∑∏
=
! ! ! =
−+
n
k
x
j
1
,
j
n k
j
j
k
Λ
1
1
n
k
j where {(
)∣
(
)
}
=
…
+
+ ⋯+
=
+
+ ⋯+
−
+
=
−+
−+
−+
k k
k
k
k
k
k
k
k
n
k
k
n
Λ
,
,
,
,
2
1
. n
k
n k
n k
n k
1
2
1
1
2
1
1
2
1 {(
)∣
(
)
}
=
…
+
+ ⋯+
=
+
+ ⋯+
−
+
=
−+
−+
−+
k k
k
k
k
k
k
k
k
n
k
k
n
Λ
,
,
,
,
2
1
. 1690 1690
Taekyun Kim et al. Then, by (19) and (20), we obtain Then, by (19) and (20), we obtain Then, by (19) and (20), we obtain (
)
(
)
( ( )
( )
( )
)
(
)
∑
… =
−
! …
≥
=
−+
−+
K
x x
k B
x
x
x
n
,
,
1
1
,
1
,
,
1
,
1 ,
n λ
k
n
k
n k
λ
λ
n k
n k
λ
,
1
2
1
,
1
1,
2
2,
1
1, (
)
(
)
( ( )
( )
( )
)
(
)
(
)
∑
… =
−
! …
≥
… =
=
−+
−+
K
x x
k B
x
x
x
n
K
x x
,
,
1
1
,
1
,
,
1
,
1 ,
,
,
1. n λ
k
n
k
n k
λ
λ
n k
n k
λ
λ
,
1
2
1
,
1
1,
2
2,
1
1,
0,
1
2 (
)
… =
K
x x
,
,
1. λ
0,
1
2 Note that, from (19) and (20), we have Note that, from (19) and (20), we have (
)
(
)
(( )
( )
( )
)
(
)
(
)
∑
∑
… =
−
! …
=
−
! =
−+
=
K
k B
k S
n k
1, 1,
1
1
, 1
,
, 1
1
,
. n λ
k
n
k
n k
λ
λ
n k
λ
k
n
k
λ
,
0
,
1,
2,
1,
0
2, Taking
=
α
1 in (17), we have Taking
=
α
1 in (17), we have ( )
( )
( )
( )
[
]
∑
−∑
=
! =
−
! =
∞
−
t
e t
t
e
t
β
x t
n . r
λ
l
r
l
l
λ
x
n
n λ
r
n
0
1 1
0
,
1,1
l λ
,
(21) ( )
( )
( )
( )
[
]
∑
−∑
=
! =
−
! =
∞
−
t
e t
t
e
t
β
x t
n . =
=
=
∞
=
∞
−+
−+
=
∞
=
−+
k k
l
t
k
B
x
x
x
t
n
k B
x
x
x
t
n
1
1
1
1
,
1
,
,
1
1
1
1
,
1
,
,
. k
k
l
l
k
k
n k
n k
λ
λ
n k
n k
n
n
k
n
k
n k
λ
λ
n k
λ
n
0
1
1
,
1
1,
2
2,
1
1
1
1
,
1
1,
2
2,
1,
(19) (19) ⎜
⎟
⎝
⎠
(
)
( ( )
( )
( )
)
⎛
⎝
(
)
( ( )
( )
( )
)⎞
⎠
∑
∑
∑∑
! ! =
+
−
! …
! =
+
−
! …
! =
=
=
∞
=
∞
−+
−+
=
∞
=
−+
k
l
k
B
x
x
x
t
n
k B
x
x
x
t
n
1
1
1
,
1
,
,
1
1
1
1
,
1
,
,
. k
l
k
k
n k
n k
λ
λ
n k
n k
n
n
k
n
k
n k
λ
λ
n k
λ
n
0
1
1
,
1
1,
2
2,
1
1
1
1
,
1
1,
2
2,
1,
(19)
We denote (19) by We denote (19) by (
)
( )
∑
∑
=
…
! =
∞
! =
∞
x t
K
x x
t
n
1
,
,
. n
n
n n
n
n λ
n
0
1
0
,
1
2
n λ
,
(20) (
)
( )
∑
∑
=
…
! =
∞
! =
∞
x t
K
x x
t
n
1
,
,
. n
n
n n
n
n λ
n
0
1
0
,
1
2
n λ
,
(20) (20) λ
x
r
λ
l
r
l λ
l
l
l λ
r
l
r
j r
j λ
j
l
l λ
r
l
j
j r λ
j
l
l λ
r
l
n
j
n
j r λ
n j λ
r
n
1
1
,
0
,
1,1
,
0
,
1,1
0
,
0
,
1,1
0
0
,
,
1,1 (22) By (1) and (22), we obtain By (1) and (22), we obtain ⎜
⎟
( )
⎛
⎝
⎞
⎠(
) ( )
( )
(
)
[
]
∑
=
! +
! ≥
=
+
−
−
x
n
j
j
j
r
β
x
n
1
,
0 . n λ
j
n
j r λ n j λ
r
,
0
,
,
1,1
(23) ⎜
⎟
( )
⎛
⎝
⎞
⎠(
) ( )
( )
(
)
[
]
∑
=
! +
! ≥
=
+
−
−
x
n
j
j
j
r
β
x
n
1
,
0 . n λ
j
n
j r λ n j λ
r
,
0
,
,
1,1
(23) (23) In particular, for
=
n
0,
[
]
( )
=
−
! β
λ
r
r
0,
1,1
1 r λ
, . By (23), we obtain In particular, for
=
n
0,
[
]
( )
=
−
! β
λ
r
r
0,
1,1
1 r λ
, . By (23), we obtain n particular, for
=
n
0,
[
]
( )
=
−
! β
λ
r
r
0,
1,1
1 r λ
, . By (23), we obtain
( )
⎛
⎝
⎞
⎠
( )
⎧
⎨⎩
(
)
(
)
(
)
(
)
(
)(
) (
)(
(
) )⎫
⎬⎭
[
]
[
]
=
! −
−
+
=
! −
+
−
+
−
−
+
+
+
+
−
−
+
−
−
β
r
x
rλ
r
β
r
x x
λ
x
rλ
r
rλ
r
r
r
rλ
r
λ
1
1
1
,
1
2
1
1
2 1
1
2
1
2
1
1
1
. λ
r
r λ
λ
r
r λ
1,
1,1
,
2,
1,1
,
2
2 In particular, for
=
n
0,
[
]
( )
=
−
! β
λ
r
r
0,
1,1
1 r λ
, . By (23), we obtain By (23), we obtain ( )
⎛
⎝
⎞
⎠
( )
⎧
⎨⎩
(
)
(
)
(
)
(
)
(
)(
) (
)(
(
) )⎫
⎬⎭
[
]
[
]
=
! −
−
+
=
! −
+
−
+
−
−
+
+
+
+
−
−
+
−
−
β
r
x
rλ
r
β
r
x x
λ
x
rλ
r
rλ
r
r
r
rλ
r
λ
1
1
1
,
1
2
1
1
2 1
1
2
1
2
1
1
1
. λ
r
r λ
λ
r
r λ
1,
1,1
,
2,
1,1
,
2
2 ( )
⎛
⎝
⎞
⎠
[
] =
! −
−
+
−
β
r
x
rλ
r
1
1
1
,
λ
r
r λ
1,
1,1
, From (6), we note that (
)
( )
( )
(
)
∑
∑
+
=
! =
+
! =
∞
−
=
∞
+
t
t
t
λ
n
t
λ
n
t
1 log 1
1
1
1
1
. λ
n
n
n
n
n
n
n
n
1
1
,
0
1,
λ
λ
1
1
(24) (24) Replacing t by
( ) −
e t
1
λ
in (24), we obtain Replacing t by
( ) −
e t
1
λ
in (24), we obtain
( )
⎛
( )
⎞
λ
λ
1
1
k
n
k Replacing t by
( ) −
e t
1
λ
in (24), we obtain
( )
( )
(
)
(
( )
)
⎛
⎝
⎜
( )
(
)⎞
⎠
⎟
∑
∑
∑∑
! =
=
+
! −
=
+
! ∞
∞
+
∞
+
β
t
n
t
e t
λ
k
k
e t
λ
k
S
n k
t
n
1
1
1
1
1
1
1
,
. n λ
n
k
k
λ
k
n
k
k
λ
n
,
1,
1,
2,
λ
λ
1
1
(25) ( )
( )
(
)
(
( )
)
⎛
⎝
⎜
( )
(
)⎞
⎠
⎟
∑
∑
∑∑
! =
−
=
+
! −
=
+
! r
λ
l
r
l
l
λ
x
n
n λ
r
n
0
1 1
0
,
1,1
l λ
, (21) When
=
r
1,
( )
( )
[
]
=
β
x
β
x
n λ
n λ
,
0,1
,
are the degenerate Bernoulli polynomials. From (21), we note that When
=
r
1,
( )
( )
[
]
=
β
x
β
x
n λ
n λ
,
0,1
,
are the degenerate Bernoulli polynomials. From (21), we note that When
=
r
1,
( )
( )
[
]
=
β
x
β
x
n λ
n λ
,
0,1
,
are the degenerate Bernoulli polynomials. From (21), we note that When
=
r
1,
( )
( )
[
]
=
β
x
β
x
n λ
n λ
,
0,1
,
are the degenerate Bernoulli polynomials. From (21), we note that When
=
r
1,
( )
( )
[
]
=
β
x
β
x
n λ
n λ
,
0,1
,
are the degenerate Bernoulli polynomials. From (21), we note that ⎜
⎟
⎜
⎟
( )
⎛
⎝
( )
( )
⎞
⎠
( )
( )
( )
( )
(
)
( )
⎛
⎝
⎜
⎛
⎝
⎞
⎠
( )
(
)
( )⎞
⎠
⎟
[
]
[
]
[
]
[
]
∑
∑
∑
∑
∑
∑
∑∑
=
−
! ! =
! ! =
+
! ! =
! +
! ! =
−
=
∞
−
=
∞
=
∞
−
=
∞
+
=
∞
−
=
∞
=
+
−
−
e
t
t
e t
l
t
β
x t
l
t
j
t
β
x t
l
j
r
t
β
x t
l
n
j
j
j
r
β
x
t
n
1
1
1
1
1
1
. =
∞
=
∞
+
=
∞
=
+
β
t
n
t
e t
λ
k
k
e t
λ
k
S
n k
t
n
1
1
1
1
1
1
1
,
. n
n λ
n
λ
k
k
k
λ
k
n
k
n
k
k
λ
n
0
,
0
1,
0
0
1,
2,
λ
λ
1
1
(25) ( )
( )
(
)
(
( )
)
⎛
⎝
⎜
( )
(
)⎞
⎠
⎟
∑
∑
∑∑
! =
−
=
+
! −
=
+
! =
∞
=
∞
+
=
∞
=
+
β
t
n
t
e t
λ
k
k
e t
λ
k
S
n k
t
n
1
1
1
1
1
1
1
,
. n
n λ
n
λ
k
k
k
λ
k
n
k
n
k
k
λ
n
0
,
0
1,
0
0
1,
2,
λ
λ
1
1
(25) (25) Comparing the coefficients on both sides of (25), we have Comparing the coefficients on both sides of (25), we have Comparing the coefficients on both sides of (25), we have ( )
(
)
(
)
∑
=
+
≥
=
+
β
λ
k
S
n k
n
1
1
,
,
0 . n λ
k
n
k
k
λ
,
0
1,
2,
λ
1
(26) (26) Study on r-truncated degenerate Stirling numbers of the second kind
1691 Study on r-truncated degenerate Stirling numbers of the second kind
1691 j
n
λ
j λ
j
n
λ
j λ
j
n
n j k
λ
k
j
λ
j
n
n j
λ
k
k
j
λ
n
λ
k
k
λ
n
j
k
k j
λ
j k j
k
λ
k
n
λ
j
k
k j
n
λ
j
j
k
n λ
j
k
k j
n
λ
j
j
k
n λ
j
k
k j
n
l
λ
l
k
n λ
j
k
k j
λ
0
2,
2
,
0
2,
2
,
0
0
0
1
1
1
,
1
1
1
,
1
0
2,
,
1
2,
(29) ⎜
⎟
(
)⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
[
](
( )
) [ ]
( )
(
)
[
](
( )
) [ ]
( )
(
( )
)
[ ]
[ ]
∑
∑
∑
∑
−
+
+
=
+
! −
+
−
+
! ! =
+
! −
−
! −
=
+
! −
−
! −
=
=
=
−+
=
−
S
n
j
k
k
n
k
j
β
n
k
S
n
j
k
k
n
j
k
β
j
n
k
t
e t
t
k
t
t
e t
n
k
t
e t
t
k t
t
t
e t
t
t
t
, 2
, 2
1
1
1
1
1
j
n
λ
j λ
j
n
λ
j λ
j
n
n j k
λ
k
j
λ
j
n
n j
λ
k
k
j
λ
k
0
2,
2
,
0
2,
2
,
0
0 (29) ⎜
⎟
(
) [ ]
⎛
⎝
⎞
⎠
(
)
(
( )
)
(
( )
)
(
) (
) [ ]
( )
(
)
(
)
[ ] (
( )
)
(
)
(
) (
)
(
)
(
)
(
) [ ](
( )
)
(
) ⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
[ ]
(
)
(
)
(
) ⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
∑
∑
∑
∑
∑
∑
=
+
! −
−
! −
=
−
+
! ! −
+
+
! −
+
+
! −
−
! −
! =
−
+
! ! ! +
+
! −
! −
! −
=
−
+
+
+
! −
−
! ! −
! +
−
−
+
−
! =
−
+
+
+
! −
−
! +
−
−
+
−
! =
=
=
−+
=
−
=
−
−
=
−
−
−
=
−
−
−
=
−
=
∞
=
−
S
n
j
k
k
n
k
j
β
n
k
S
n
j
k
k
n
j
k
β
j
n
k
t
e t
t
k
t
t
e t
n
k
t
e t
t
k t
t
t
e t
n
k
t
e t
t
t
k t
e t
n
k
t
k
j
e t
t
t
k t
e t
n
k
k
t
t
e t
n
k
t
e t
j
k
j
t
n
k
k
β
n
n
k
j
k
j
t
e t
t
n
k
k
β
n
k
j
j k
j
t
S
l
j
j
l
j
t
n
k
k
β
n
k
j k
j
S
n
j
j
n
j
, 2
, 2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1,
1
1
1
1
1,
1
1
. Therefore, by (29), we obtain the following theorem. Replacing t by
(
)
+ t
log 1
λ
in (3), with
=
=
α
x
1,
0, we have Replacing t by
(
)
+ t
log 1
λ
in (3), with
=
=
α
x
1,
0, we have Replacing t by
(
)
+ t
log 1
λ
in (3), with
=
=
α
x
1,
0, we have ⎜
⎟
( )
(
)
(
)
(
(
))
(
)
⎛
⎝
(
)⎞
⎠
∑
∑
∑
∑
∑∑
+
! =
+
=
! +
=
! =
! =
∞
+
=
∞
=
∞
=
∞
=
∞
=
λ
n
t
t
t
β
k
t
β
S
n k t
n
β
S
n k
t
n
1
1
1 log 1
1
log 1
,
,
. n
n
n
n
λ
k
k λ
λ
k
k
k λ
n k
λ
n
n
k
n
k λ
λ
n
0
1,
0
,
0
,
1,
0
0
,
1,
λ
1
(27) (27) By comparing the coefficients on both sides of (27), we obtain By comparing the coefficients on both sides of (27), we obtain By comparing the coefficients on both sides of (27), we obtain (
)
( )
(
)
∑
=
+
≥
=
+
β
S
n k
λ
n
n
,
1
1
,
0 . k
n
k λ
λ
n
n
0
,
1,
1, λ
1 (
)
( )
(
)
∑
=
+
≥
=
+
β
S
n k
λ
n
n
,
1
1
,
0 . k
n
k λ
λ
n
n
0
,
1,
1, λ
1
(28) (
)
( )
(
)
∑
=
+
≥
=
+
β
S
n k
λ
n
n
,
1
1
,
0 . k
n
k λ
λ
n
n
0
,
1,
1, λ
1 (28) Therefore, by (26) and (28), we obtain the following theorem. Therefore, by (26) and (28), we obtain the following theorem. Therefore, by (26) and (28), we obtain the following theorem. Theorem 4. For
≥
n
0, we have Theorem 4. For
≥
n
0, we have ( )
(
)
∑
=
+
=
+
β
λ
k
S
n k
1
1
,
n λ
k
n
k
k
λ
,
0
1,
2,
λ
1
and
( )
(
)
∑
+
=
+
=
n
λ
β
S
n k
1
1
,
. n
n
k
n
k λ
λ
1,
0
,
1,
λ
1 ( )
(
)
∑
=
+
=
+
β
λ
k
S
n k
1
1
,
n λ
k
n
k
k
λ
,
0
1,
2,
λ
1 ( )
(
)
∑
=
+
=
+
β
λ
k
S
n k
1
1
,
n λ
k
n
k
k
λ
,
0
1,
2,
λ
1 ( )
(
)
∑
=
+
=
+
β
λ
k
S
n k
1
1
,
n λ
k
n
k
k
λ
,
0
1,
2,
λ
1 and ( )
(
)
∑
+
=
+
=
n
λ
β
S
n k
1
1
,
. n
n
k
n
k λ
λ
1,
0
,
1,
λ
1 From Theorem 4, we note that ⎜
⎟
⎜
⎟
(
)⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
[
](
( )
) [ ]
( )
(
)
[
](
( )
) [ ]
( )
(
) [ ](
( )
)
(
( )
)
(
) [ ]
⎛
⎝
⎞
⎠
(
)
(
( )
)
(
( )
)
(
) (
) [ ]
( )
(
)
(
)
[ ] (
( )
)
(
)
(
) (
)
(
)
(
)
(
) [ ](
( )
)
(
) ⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
[ ]
(
)
(
)
(
) ⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
[ ]
[ ]
∑
∑
∑
∑
∑
∑
∑
∑
∑
∑
−
+
+
=
+
! −
+
−
+
! ! =
+
! −
−
! −
=
+
! −
−
! −
=
+
! −
−
! −
=
+
! −
−
! −
=
−
+
! ! Therefore, by (29), we obtain the following theorem. −
−
! −
! =
−
+
! ! ! +
+
! −
! −
! −
=
−
+
+
+
! −
−
! ! −
! +
−
−
+
−
! =
−
+
+
+
! −
−
! +
−
−
+
−
! =
−
−
=
−
−
−
=
−
−
−
=
−
=
∞
=
−
n
k
t
k
j
e t
t
t
k t
e t
n
k
k
t
t
e t
n
k
t
e t
j
k
j
t
n
k
k
β
n
n
k
j
k
j
t
e t
t
n
k
k
β
n
k
j
j k
j
t
S
l
j
j
l
j
t
n
k
k
β
n
k
j k
j
S
n
j
j
n
j
1
1
1
1
1
1
1
1
1
1
1
1
1
1,
1
1
1
1
1,
1
1
. n
j
k
k j
λ
j k j
k
λ
k
n
λ
j
k
k j
n
λ
j
j
k
n λ
j
k
k j
n
λ
j
j
k
n λ
j
k
k j
n
l
λ
l
k
n λ
j
k
k j
λ
0
1
1
1
,
1
1
1
,
1
0
2,
,
1
2, Therefore, by (29), we obtain the following theorem. Therefore, by (29), we obtain the following theorem. 1692
Taekyun Kim et al. 1692
Taekyun Kim et al. 1692
Taekyun Kim et al. Theorem 5. For
≥
n k
,
0, we have Theorem 5. For
≥
n k
,
0, we have Theorem 5. For
≥
n k
,
0, we have (
)⎛
⎝
⎞
⎠
(
) ⎛
⎝
⎞
⎠
(
)
(
)
(
) (
)
(
)
[ ]
∑
∑
−
+
+
= −
+
+
+
! −
−
! +
−
! +
−
−
=
=
−
S
n
j
k
k
n
j
k
β
n
k
k
β
n
k
j k
j
n
j
S
n
j
j
, 2
1
1
1
1,
1 . j
n
λ
j λ
n
n λ
j
k
k j
λ
0
2,
2
,
,
1
2, Observe that, for any formal power series
( ) = ∑=
∞
f t
a t
i
i i
0
, we have [ ] ( )
(
)
(
)[
] ( )
′
=
+
=
+
+
+
t
f t
n
a
n
t
f t
1
1
. n
n
n
1
1
(30) (30) [ ] ( )
(
)
(
)[
] ( )
′
=
+
=
+
+
+
t
f t
n
a
n
t
f t
1
1
. 1692
Taekyun Kim et al. n
n
n
1
1 By making use of (30), we have By making use of (30), we have ⎜
⎟
⎛
⎝
⎞
⎠
(
)
[ ]
∑
+
−
−
+
=
n
k
j
S
n
j
k
k β
1
, 2
j
n
λ
j λ
0
2,
2
, ⎜
⎟
⎜
⎟
⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
(
)[
](
( )
) [ ]
( )
(
)
[
]
⎛
⎝
(
( )
) ⎞
⎠
[ ]
( )
(
)
[
] (
( )
)
(
( )
)
(
)
[ ]
( )
(
)
[
] (
( )
)
(
( )
)
(
) (
)
[ ]
( )
(
)
[
] (
( )
)
(
( )
)
(
) (
)
[ ]
( )
(
) [ ] (
( )
)
(
) (
)
(
)
[ ] (
( )
)
(
) (
)
( )
(
) [ ]⎧
⎨⎩
(
)
(
)
(
)
(
)
(
)
(
)
(
)
⎫
⎬⎭
(
)
(
)
(
)
(
)
(
)
(
)
(
)
(
) (
)
[ ]
[ ]
[ ]
[ ]
[ ]
[ ]
∑
∑
∑
∑
∑
∑
∑
∑∑
∑∑∑
∑
∑∑
+
−
−
+
=
+
−
! −
+
−
+
−
! ! =
+
−
! −
+
−
−
! −
=
+
−
! −
−
! −
=
+
−
! −
−
−
−
! −
=
+
−
! −
−
−
−
−
! +
−
=
+
−
! −
−
−
−
−
! +
−
=
+
−
! −
−
−
! +
−
+
−
! −
−
−
! +
−
=
+
−
! +
−
−
+
−
! −
−
+
−
−
+
−
! −
−
! =
+
−
! +
−
−
+
−
! −
−
+
−
! +
−
−
−
+
−
! −
! 1692
Taekyun Kim et al. =
=
=
−+
=
−+ −
=
−+ −
−
−
=
−+ −
−
=
−
−
−
−
−
−
−
=
∞
=
−
=
∞
=
=
−
−
=
−
=
=
−
−
n
k
j
S
n
j
k
k β
n
k
S
n
j
k
k
n
j
k
β
j
n
k
n
j
k t
e t
t
k
t
t
e t
n
k
t
d
dt
e t
t
k
t
t
e t
n
k
t
e t
t
e
t
k
t
t
e t
n
k
t
e t
t
e t
λt
k
λt
t
t
e t
n
k
t
e t
t
e t
λt
t
k
λt
t
t
e t
n
k
t
e t
t
t
k
λt
n
k
λ t
e t
t
t
k
λt
t
e t
n
k
t
S
l
k
k
l
k
λ
t
λ
S
l
k
k
l
k
λ
β
m
j
t
n
k
S
l
k
k
l
k
λ
λ n
k
S
l
k
k
β
λ
l
k
n
j
1
, 2
1
, 2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2, 2
2
2
2, 2
2
2
1
2, 2
2
2
1
2, 2
2
2
. 1692
Taekyun Kim et al. =
−
−
−
−
=
∞
=
−
=
∞
=
=
−
−
=
−
=
=
−
−
t
k
λt
e t
n
k
t
e t
t
t
k
λt
n
k
λ t
e t
t
t
k
λt
t
e t
n
k
t
S
l
k
k
l
k
λ
t
λ
S
l
k
k
l
k
λ
β
m
j
t
n
k
S
l
k
k
l
k
λ
λ n
k
S
l
k
k
β
λ
l
k
n
j
1
1
1
1
1
1
1
1
1
1
1
1
1
2, 2
2
2
2, 2
2
2
1
2, 2
2
2
1
2, 2
2
2
. j
λ
n
λ
k
k
n
λ
k
k
λ
n
m k l k
m
λ
m l m
m k j k
m
l k
j
λ
j l
m j λ
m
l k
n
λ
n l
j k
n
l k
j
λ
n j λ
j l
0
1
2
1
2
2,
2
2,
2
,
2,
2
2,
2
, Therefore, by (31), we obtain the following theorem. Therefore, by (31), we obtain the following theorem. Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have 1692
Taekyun Kim et al. j
n
λ
j λ
j
n
λ
j λ
j
n
n j k
λ
k
j
λ
j
n
n j k
λ
k
j
λ
j
n
n j k
λ
k
λ
λ
j
λ
j
n
n j k
λ
k
λ
j
λ
j
n
n j
λ
k
λ
k
j
λ
n
λ
k
k
n
λ
k
k
λ
n
m k l k
m
λ
m l m
m k j k
m
l k
j
λ
j l
m j λ
m
l k
n
λ
n l
j k
n
l k
j
λ
n j λ
j l
0
2,
2
,
0
2,
2
,
0
0
1
0
1
1
1
0
1
1
0
1
1
1
2
1
2
2,
2
2,
2
,
2,
2
2,
2
, (
)
(
)
(
)
[ ]
∑
=
+
−
! −
+
−
+
−
! ! =
n
k
S
n
j
k
k
n
j
k
β
j
1
, 2
1
j
n
λ
j λ
0
2,
2
, (31) (
) (
)
( )
(
) [ ] (
( )
)
(
) (
)
(
)
[ ] (
( )
)
(
) (
)
( )
(
) [ ]⎧
⎨⎩
(
)
(
)
(
)
(
)
(
)
(
)
(
)
⎫
⎬⎭
(
)
(
)
(
)
(
)
(
)
(
)
(
)
(
) (
)
[ ]
[ ]
[ ]
[ ]
∑∑
∑∑∑
∑
∑∑
! +
=
+
−
! −
−
−
! +
−
+
−
! −
−
−
! +
−
=
+
−
! +
−
−
+
−
! −
−
+
−
−
+
−
! −
−
! =
+
−
! +
−
−
+
−
! −
−
+
−
! +
−
−
−
+
−
! −
! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have ⎜
⎟
⎛
⎝
⎞
⎠
(
)
(
) ⎧
⎨⎩
(
)
(
)
(
)
(
)
(
)
(
) (
)
⎫
⎬⎭
[ ]
[ ]
[ ]
∑
∑
∑∑
+
−
−
+
=
+
−
! +
−
−
+
−
! −
+
+
−
−
−
+
−
! −
! =
=
−
=
=
−
−+
n
k
j
S
n
j
k
k β
n
k
S
l
k
k
l
k
λ
S
l
k
k
β
λ
l
k
n
j
1
, 2
1
2, 2
2
2
2, 2
2
2
. j
n
λ
j λ
l k
n
λ
n l
j k
n
l k
j
λ
n j λ
j l
0
2,
2
,
2,
2
2,
2
,
1 Study on r-truncated degenerate Stirling numbers of the second kind
1693 Now, we observe that Now, we observe that Now, we observe that ⎜
⎟
⎜
⎟
⎜
⎟
⎜
⎟
⎜
⎟
⎜
⎟
⎜
⎟
(
)⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
[
](
( )
) [ ]⎛
⎝
( )
⎞
⎠
(
)
[
](
( )
) [ ]⎛
⎝
( )
⎞
⎠
(
) [ ]⎛
⎝
( )
⎞
⎠
(
)
⎛
⎝
⎞
⎠
(
)
[ ]⎛
⎝
( )
⎞
⎠
(
)
⎛
⎝
⎞
⎠
(
)
[ ]
( )
[ ]
( )
(
)
∑
∑
∑
∑
∑
∑
−
+
+
=
+
! −
+
−
+
! ! =
+
! −
−
! −
=
+
! ! −
−
−
=
+
! ! −
−
=
+
! ! −
−
=
+
! ! −
! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
−
−
−
=
−
−
−
−
=
∞
=
−
=
−
−
=
−
−+
n
k
t
λ
t
k
e t
n
k
k
t
k
j
e t
t
λ t
n
k
t
k
j
λ
β
m
j k
t
n
k
k
t
k
j
e t
k
j
l
λ t
1
1
1
1
1
1
1
1
2
1
1
2
1
1
1
2
n
k
t
k
λ
n
j
k
k j
λ
j
j
k j
n
m
k
j
m
j
m j λ
j
m
n
j
k
k j
λ
j
l
k j
l
l j
l
2
1
1 1
0
0
,
1
1
0
1 ⎜
⎟
⎜
⎟
⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
(
)
[
](
( )
( )
) [
]
( )
(
)
[ ](
( )
( )
) [
]⎛
⎝
( )
⎞
⎠
[ ]
[ ]
∑
∑
∑
∑
+
+
+
=
+
! +
+
! −
! =
+
! −
−
−
∕
! −
=
+
! −
−
−
∕
! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
=
−
=
−
=
+
−
=
−
n
k
j
k S
j
k
k β
n
k
S
j
k
k
j
k
β
n
j
n
k
t
e t
t
t
k
t
t
e t
n
k
t
e t
t
t
t k
t
t
e t
, 3
, 3
1
1
2
1
1
1
2
1
j
n
λ
n j λ
j
n
λ
n j λ
j
n
j k
λ
λ
k
n j
λ
j
n
j
λ
λ
k
k
n j
λ
0
2,
3
,
0
2,
3
,
0
2,
2
0
2,
2
(33) ⎜
⎟
⎜
⎟
⎜
⎟
⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
(
)
[
](
( )
( )
) [
]
( )
(
)
[ ](
( )
( )
) [
]⎛
⎝
( )
⎞
⎠
(
) [ ](
( )
( )
)
(
( )
)
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
) (
(
) )
(
( )
)
[ ]
[ ]
∑
∑
∑
∑
∑
+
+
+
=
+
! +
+
! −
! =
+
! −
−
−
∕
! −
=
+
! −
−
−
∕
! −
=
+
! −
−
−
∕
! −
=
+
! −
−
+
−
! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
−
−
−
=
−
−
−
−
=
∞
=
−
=
−
−
=
−
−+
n
k
t
λ
t
k
e t
n
k
k
t
k
j
e t
t
λ t
n
k
t
k
j
λ
β
m
j k
t
n
k
k
t
k
j
e t
k
j
l
λ t
1
1
1
1
1
1
1
1
2
1
1
2
1
1
1
2
n
k
t
k
λ
n
j
k
k j
λ
j
j
k j
n
m
k
j
m
j
m j λ
j
m
n
j
k
k j
λ
j
l
k j
l
l j
l
2
1
1 1
0
0
,
1
1
0
1 (33) ⎜
⎟
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
) (
(
) )
(
( )
)
∑
=
+
! −
−
+
−
! −
=
−
−
n
k
t
k
j
e t
t
λ
t
k
e t
t
1
1
1
1
n
j
k
k j
λ
j
t
k j
λ
k
0
2
2 ⎜
⎟
⎜
⎟
⎜
⎟
(
) [ ]
(
) (
(
) )
(
( )
)
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
)
⎛
⎝
(
) ⎞
⎠
(
) [ ]
⎛
⎝
⎜(
)
⎛
⎝
⎞
⎠
(
)
(
)
⎞
⎠
⎟
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
)
⎛
⎝
⎞
⎠
(
)
∑
∑
∑
∑
∑
=
+
! −
+
−
! −
+
+
! ! ×
−
−
+
−
=
+
! −
−
−
! ! +
+
! ! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have =
=
=
−+
=
−
=
−
−
=
−
−
S
n
j
k
k
n
k
j
β
n
k
S
n
j
k
k
n
j
k
β
j
n
k
t
e t
t
k
t
t
e t
n
k
k
t
e t
t
t
t
t
e t
n
k
k
t
t
e t
n
k
k
k
j
t
t
e t
n
k
k
k
j
β
n
, 2
, 2
1
1
1
1
1
1
1
1
1
. j
n
λ
j λ
k
j
n
λ
j λ
k
j
n
n j k
λ
k
j
λ
k
j
n
n j
λ
k
k
j
λ
k
n
λ
k
j
k
k j
n
λ
k j
j
k
k j
n λ
k j
0
2,
2
,
0
2,
2
,
0
0
0
0
,
(32) (32) Therefore, by (32), we obtain the following theorem. Therefore, by (32), we obtain the following theorem. Therefore, by (32), we obtain the following theorem. Theorem 7. For
≥
n k
,
0, we have Theorem 7. For
≥
n k
,
0, we have ⎜
⎟
⎜
⎟
(
)⎛
⎝
⎞
⎠
⎛
⎝
⎞
⎠
⎛
⎝
⎞
⎠
(
)
[ ]
( )
(
)
∑
∑
−
+
+
=
+
−
=
=
−
−
S
n
j
k
k
n
k
j
β
n
k
k
k
j
β
, 2
1
. j
n
λ
j λ
k
j
k
k j
n λ
k j
0
2,
2
,
0
, From Theorems 4 and 5, we have From Theorems 4 and 5, we have ⎜
⎟
⎜
⎟
⎜
⎟
⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
(
)
[
](
( )
( )
) [
]
( )
(
)
[ ](
( )
( )
) [
]⎛
⎝
( )
⎞
⎠
(
) [ ](
( )
( )
)
(
( )
)
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
) (
(
) )
(
( )
)
[ ]
[ ]
∑
∑
∑
∑
∑
+
+
+
=
+
! +
+
! −
! =
+
! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
−
−
∕
! −
=
+
! −
−
−
∕
! −
=
+
! −
−
−
∕
! −
=
+
! −
−
+
−
! −
=
−
=
−
=
+
−
=
−
=
−
−
n
k
j
k S
j
k
k β
n
k
S
j
k
k
j
k
β
n
j
n
k
t
e t
t
t
k
t
t
e t
n
k
t
e t
t
t
t k
t
t
e t
n
k
t
e t
t
t
t
t k
e t
n
k
t
k
j
e t
t
λ
t
k
e t
t
, 3
, 3
1
1
2
1
1
1
2
1
1
1
2
1
1
1
1
1
j
n
λ
n j λ
j
n
λ
n j λ
j
n
j k
λ
λ
k
n j
λ
j
n
j
λ
λ
k
k
n j
λ
n
λ
λ
k
k
λ
n
j
k
k j
λ
j
t
k j
λ
k
0
2,
3
,
0
2,
3
,
0
2,
2
0
2,
2
2,
2
0
2
2
(33)
⎜
⎟
⎜
⎟
⎜
⎟
(
) [ ]
(
) (
(
) )
(
( )
)
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
)
⎛
⎝
(
) ⎞
⎠
(
) [ ]
⎛
⎝
⎜(
)
⎛
⎝
⎞
⎠
(
)
(
)
⎞
⎠
⎟
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
)
⎛
⎝
⎞
⎠
(
)
∑
∑
∑
∑
∑
=
+
! −
+
−
! −
+
+
! ! ×
−
−
+
−
=
+
! −
−
−
! ! +
+
! ! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
=
−
=
−
=
+
−
=
−
=
−
−
n
k
j
k S
j
k
k β
n
k
S
j
k
k
j
k
β
n
j
n
k
t
e t
t
t
k
t
t
e t
n
k
t
e t
t
t
t k
t
t
e t
n
k
t
e t
t
t
t
t k
e t
n
k
t
k
j
e t
t
λ
t
k
e t
t
, 3
, 3
1
1
2
1
1
1
2
1
1
1
2
1
1
1
1
1
j
n
λ
n j λ
j
n
λ
n j λ
j
n
j k
λ
λ
k
n j
λ
j
n
j
λ
λ
k
k
n j
λ
n
λ
λ
k
k
λ
n
j
k
k j
λ
j
t
k j
λ
k
0
2,
3
,
0
2,
3
,
0
2,
2
0
2,
2
2,
2
0
2
2
(33) ⎜
⎟
⎜
⎟
(
)
(
)
(
)
(
)
(
)
[
](
( )
( )
) [
]
( )
(
)
[ ](
( )
( )
) [
]⎛
⎝
( )
⎞
⎠
(
) [ ](
( )
( )
)
(
( )
)
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
) (
(
) )
(
( )
)
[ ]
∑
∑
∑
∑
=
+
! +
+
! −
! =
+
! −
−
−
∕
! −
=
+
! −
−
−
∕
! −
=
+
! −
−
−
∕
! −
=
+
! −
−
+
−
! Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
−
=
−
=
+
−
=
−
=
−
−
n
k
S
j
k
k
j
k
β
n
j
n
k
t
e t
t
t
k
t
t
e t
n
k
t
e t
t
t
t k
t
t
e t
n
k
t
e t
t
t
t
t k
e t
n
k
t
k
j
e t
t
λ
t
k
e t
t
, 3
1
1
2
1
1
1
2
1
1
1
2
1
1
1
1
1
j λ
j
n
λ
n j λ
j
n
j k
λ
λ
k
n j
λ
j
n
j
λ
λ
k
k
n j
λ
n
λ
λ
k
k
λ
n
j
k
k j
λ
j
t
k j
λ
k
,
0
2,
3
,
0
2,
2
0
2,
2
2,
2
0
2
2
(33)
⎜
⎟
⎜
⎟
⎜
⎟
(
) [ ]
(
) (
(
) )
(
( )
)
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
)
⎛
⎝
(
) ⎞
⎠
(
) [ ]
⎛
⎝
⎜(
)
⎛
⎝
⎞
⎠
(
)
(
)
⎞
⎠
⎟
(
) [ ]
(
)
⎛
⎝
⎞
⎠
(
( )
)
⎛
⎝
⎞
⎠
(
)
∑
∑
∑
∑
∑
=
+
! −
+
−
! −
+
+
! ! ×
−
−
+
−
=
+
! −
−
−
! ! +
+
! ! Theorem 8. For
≥
n k
,
0, we have ⎜
⎟
⎜
⎟
⎛
⎝
⎞
⎠
(
)
(
) (
)
⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
⎛
⎝
⎞
⎠
(
)(
)
(
)
[ ]
∑
∑
∑
∑
+
+
+
= −
+
! −
−
! ! +
+
! −
−
! −
−
+
−
−
−
−
+
−
! =
−
=
−
=
−
=
n
k
j
k S
j
k
k β
n
k
k
j
λ
n
j k β
n
k
j k
j
k
j
l
S
n
l
j
j
λ
n
l
j
, 3
1
1
1
2
1
1,
1 1
2
1
. j
n
λ
n j λ
k
j
n
j
j
n j λ
j
k
k j
l
n
λ
l
l
0
2,
3
,
0
,
1
1
2,
(34) (34) Remark 9. As the counterpart of (10), we may consider the r-truncated degenerate Stirling numbers of the
first kind given by Remark 9. As the counterpart of (10), we may consider the r-truncated degenerate Stirling numbers of the
first kind given by ⎛
⎝
⎜
(
)
( )
⎞
⎠
⎟
(
)
[ ]
∑
∑
! +
−
! =
! =
−
−
=
∞
k
t
λ
l
t
S
n kr t
n
1
log 1
1
,
,
λ
l
r
l
l
l
k
n kr
λ
r
n
1
1
,
1
1,
λ
1
(35) (35) where r is a positive integer. These numbers will be investigated in a forthcoming article. Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have Therefore, by (33), we obtain the following theorem. Therefore, by (33), we obtain the following theorem. Therefore, by (33), we obtain the following theorem. Theorem 8. For
≥
n k
,
0, we have Theorem 8. For
≥
n k
,
0, we have Theorem 6. For
≥
n k
,
1, with
≥
n
k, we have −
−
−
−
=
−
−
−
−
=
∞
=
−
=
−
−
=
−
−+
n
k
t
λ
t
k
e t
n
k
k
t
k
j
e t
t
λ t
n
k
t
k
j
λ
β
m
j k
t
n
k
k
t
k
j
e t
k
j
l
λ t
1
1
1
1
1
1
1
1
2
1
1
2
1
1
1
2
n
k
t
k
λ
n
j
k
k j
λ
j
j
k j
n
m
k
j
m
j
m j λ
j
m
n
j
k
k j
λ
j
l
k j
l
l j
l
2
1
1 1
0
0
,
1
1
0
1 Taekyun Kim et al. 1694
Taekyun Kim et al. ⎜
⎟
⎜
⎟
(
) (
)
⎛
⎝
⎞
⎠
(
)
(
)
(
) [ ]
(
)
(
)
(
)
⎛
⎝
⎞
⎠
(
)
(
) (
)
⎛
⎝
⎞
⎠
(
)
(
)
(
)
(
)
(
)
⎛
⎝
⎞
⎠
(
)(
)
(
)
∑
∑
∑
∑
∑
∑
∑
= −
+
! −
−
! ! +
+
! −
−
! −
! −
−
= −
+
! −
−
! ! +
+
! −
−
! −
−
+
−
−
−
−
+
−
! =
−
=
−
= −
∞
=
−
−+
=
−
=
−
=
n
k
k
j
λ
n
j k β
n
k
t
j k
j
S
l j
t
l
k
j
l
λ t
n
k
k
j
λ
n
j k β
n
k
j k
j
k
j
l
S
n
l
j
j
λ
n
l
j
1
1
1
2
1
,
1
1
2
1
1
1
2
1
1,
1 1
2
1
. k
j
n
j
j
n j λ
n
j
k
k j
l
j
λ
l
l
k j
l
l
l j
k
j
n
j
j
n j λ
j
k
k j
l
n
λ
l
l
0
,
1
1
2,
1
1
1
1
0
,
1
1
2,
1
1 Therefore, by (33), we obtain the following theorem. References [1]
L. Carlitz, Degenerate Stirling, Bernoulli and Eulerian numbers, Utilitas Math. 15 (1979), 51–88. [2]
L. Carlitz, A degenerate Staudt-Clausen theorem, Arch. Math. (Basel) 7 (1956), 28–33. [3]
D. S. Kim and T. Kim, A note on a new type of degenerate Bernoulli numbers, Russ. J. Math. Phys. 27 (2020), no. 2,
227–235. [4]
H. K. Kim, H. Baek, and D. S. Lee, A note on truncated degenerate exponential polynomials, Proc. Jangjeon Math. Soc. 24 (2021), no. 1, 63–76. 24 (2021), no. 1, 63–76. [5]
T. Kim and D. S. Kim, Degenerate Whitney numbers of first and second Kind of Dowling lattices, Russ. J. Math. Phys. (
),
,
[5]
T. Kim and D. S. Kim, Degenerate Whitney numbers of first and second Kind of Dowling lattices, Russ. J. Math. Phys. 29 (2022), no. 3, 358–377. [6]
T. Kim and D. S. Kim, Degenerate zero-truncated Poisson random variables, Russ. J. Math. Phys. 28 (2021), no. 1, 66–72. [6]
T. Kim and D. S. Kim, Degenerate zero-truncated Poisson random variables, Russ. J. Math. Phys. 28 (2021), no. 1, 66–72. [7]
T Kim and D S Kim Some identities on truncated polynomials associated with degenerate Bell polynomials Russ J [7]
T. Kim and D. S. Kim, Some identities on truncated polynomials associated with degenerate Bell polynomials, Russ. J. Math. Phys. 28 (2021), no. 3, 342–355. [8]
T. Kim, D. S. Kim, L.-C. Jang, H. Lee, and H. Kim, Representations of degenerate Hermite polynomials, Adv. A
139 (2022), 102359. [9]
T. Kim, D. S. Kim, H. K. Kim, and H. Lee, Some properties on degenerate Fubini polynomials, Appl. Math. Sci. Eng. 30 (2022), no. 1, 235–248. (
)
[10] T. Kim, D. S. Kim, H. Lee, S. Park, and J. Kwon, New properties on degenerate Bell polynomials, Complexity 2021 (2021)
7648994. [11]
S. K. Sharma, W. A. Khan, S. Araci, and S. S. Ahmed, New type of degenerate Daehee polynomials of the second kind, Adv. Difference Equations 2020 (2020), 428. [12] T. Kim and D. S. Kim, Degenerate Laplace transform and degenerate gamma function, Russ. J. Math. Phys. 24 (2017), no. 2,
241–248. T. Kim and D. S. Kim, Note on the degenerate gamma function, Russ. J. Math. Phys. 27 (2020), no. 3, 352–358. [14] D. S. Kim and T. Kim, Degenerate Sheffer sequences and λ-Sheffer sequences, J. Math. Anal. Appl. 493 (2021), no. 1,
124521. [15] U. Duran and M. 3 Conclusion In recent years, we have witnessed that many degenerate versions of quite a few special numbers and
polynomials were investigated and some nice results were obtained by adopting various tools. In this article, we considered the r-truncated degenerate Stirling numbers of the second, which reduce
to the degenerate Stirling numbers of the second for
=
r
1, and studied by using generating functions their
explicit expressions, some properties and related identities on those numbers, in connection with several
other degenerate special numbers and polynomials. As one of our future projects, we would like to continue this line of research, namely, to explore various
degenerate versions of some special numbers and polynomials, and to find their applications to physics,
science and engineering. Funding information: This work was supported by the Basic Science Research Program, the National
Research Foundation of Korea (NRF-2021R1F1A1050151). Author contributions: All authors contributed equally to the writing of this article. All authors read and
approved the final manuscript. Conflict of interest: The authors state no conflict of interest. 1695 Study on r-truncated degenerate Stirling numbers of the second kind Disclosure: An earlier version of the article has been presented as a preprint in the following link: https://
arxiv.org/abs/2206.04402. Data availability statement: No data were used to support this study. References Acikgoz, On degenerate truncated special polynomials, Mathematics 8 (2020), no. 1, 144. [16] S. Araci, A new class of Bernoulli polynomials attached to polyexponential functions and related identities, Adv. Stud. Contemp. Math. 31 (2021), no. 2, 195–204. [17] A. Bayad and J. Chikhi, Apostol-Euler polynomials and asymptotics for negative binomial reciprocals, Adv. Stud. Contemp. Math. (Kyungshang) 24 (2014), no. 1, 33–37. [18] L. Comtet, Advanced Combinatorics, D. Reidel Publishing Co., Dordrecht-Holland, 1974. [19] R. Dere and Y. Simsek, Applications of umbral algebra to some special polynomials, Adv. Stud. Contemp
(Kyungshang) 22 (2012), no. 3, 433–438. ] Y. Simsek, On Boole-type combinatorial numbers and p [20] Y. Simsek, On Boole-type combinatorial numbers and polynomials, Filomat 34 (2020), no. 2, 559–565. [21] R. F. Tate and R. L. Goen, Minimum variance unbiased estimation for the truncated Poisson distribution, Ann. Math. Statist. 29 (1958), 755–765. (
)
[22] S. Roman, The umbral calculus, Pure and Applied Mathematics, Vol. 111, Academic Press, Inc. [Harcourt Brace Jovanovich
Publishers], New York, 1984.
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ASPECTS OF LEGAL PROTECTION FOR CHILDREN FROM CYBERCRIME
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Jurnal Pembaharuan Hukum/Jurnal pembaharuan hukum
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1 Article 1 paragraph 3 of the 1945 Constitution of the Republic of Indonesia. ASPECTS OF LEGAL PROTECTION FOR CHILDREN FROM CYBERCRIME Ida Musofiana
Sultan Agung Islamic University
ida.musofiana@unissula.ac.id Ida Musofiana
Sultan Agung Islamic University
ida.musofiana@unissula.ac.id Aji Sudarmaji
Sultan Agung Islamic University
aji.s@unissula.ac.id Aji Sudarmaji
Sultan Agung Islamic University
aji.s@unissula.ac.id Ira Alia Maerani
Sultan Agung Islamic University
Ira.alia@unissula.ac.id Abstract The 21st century has progressed further. As in Indonesia, currently, the
development of technology is increasingly rapid. This certainly affects aspects
of social life in society and its security. Today's threats are not only enemies in
the military, but also non-military. Among them are the threats faced by
internet users, the penetration rate shows a high cumulation, opening up
opportunities for the emergence of cyber-based crimes. The development of
infrastructure is in line with the development of technology. Increasingly
sophisticated technology is not only used by the community for positive things. However, not a few are taking advantage of technological sophistication to reap
benefits that result in losses for technology users themselves. This study aims
to research, examine how legal protection for children from cybercrime. In this
case, children become vulnerable to cybercrime in digital era. Why is that? Considering that the control for internet access, as well as legal products in
Indonesia, cannot yet fully guarantee protection for victims of cybercrime. The
method used in this research is to use the descriptive analysis approach. Assessing and analyzing legal protection for children from cybercrime based on
legal theories and applicable laws and regulations. Besides, it also reveals the
phenomena of the current situation in the community. urnal Pembaharuan Hukum urnal Pembaharuan Hukum JPH: Jurnal Pembaharuan Hukum
Volume 7, Number 3, December 2020 A. INTRODUCTION A. INTRODUCTION
The Republic of Indonesia as a state law.1 Adhere to the principles of
protection law, protection of human rights for citizens country, and
democratic principles, including protection of children, so it is absolute for
state and government provide protection law and human rights to children
as part of the nation's children who are expected to become continuation of
the ideals of the struggle of the Indonesian nation. ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 201 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Precise in 2001, Indonesia stated its commitment to the Declaration of
a World Worthy for Children. This is following the provisions The Convention
on the Rights of the Child ratified by the government Indonesia through
Presidential Decree Number 36 of 1990 and later also poured in Law
Number 4 of 1979 concerning Child Welfare. Apart from that, the Law
Number 23 of 2002 concerning Child Protection as amended by Law Number
35 of 2014 concerning Amendments to the Law Number 23 of 2002
concerning Protection Children and Law Number 11 of 20122 all about the
Juvenile Criminal Justice System both put forward the principles general
protection of children, namely non-discrimination, best interests of children,
survival, and growth and respect for participation child. Entering the 21st century, any country has its challenges. The threats
faced are not only in the military field but also non-military. Several threats
to national security and defense are carried out by non-state actors such as
insurgency, terrorism, human trafficking, cybercrime, narcotics, piracy at
sea, including violations of human rights. In a conflict asymmetric warfare,
carried out by non-state actors as weak parties against the government. The use of information technology, media, and communication in
society, especially young students, has changed both the behavior of society
and human civilization globally. The development of information and
communication technology has also caused world relations to become
borderless and caused significant social, economic, and cultural changes to
take place so rapidly. Information technology is currently a double-edged
sword because, in addition to contributing to the improvement of welfare,
human progress, and civilization, it is also an effective means of acting
against the law.3 g
The more sophisticated the technology is not a few who use it for
positive activities, also not a few who use it for negative activities. 2 Ridwan Mansur's opinion is quoted in Nevey Varida Ariani, Implementation of Law Number 11
of 2012 concerning the System Juvenile Justice to Protect Children's Interests, Media Hukum
Journal, Volume 21, Nomor 1, 2014, page 108-109. 3 Ivan Fauzi Raharja, Wise to Use Social Media Among Students According to Law Number 19
of 2016 Concerning Electronic Information and Transactions, Journal Selat, Volume 6,
Number 2, May 2019, page 236. 2 Ridwan Mansur's opinion is quoted in Nevey Varida Ariani, Implementation of Law Number 11
of 2012 concerning the System Juvenile Justice to Protect Children's Interests, Media Hukum
Journal, Volume 21, Nomor 1, 2014, page 108-109.
3 Ivan Fauzi Raharja, Wise to Use Social Media Among Students According to Law Number 19
of 2016 Concerning Electronic Information and Transactions, Journal Selat, Volume 6, 202
ASPECTS OF LEGAL PROTECTION FOR Jurnal Pembaharuan Hukum
CHILDREN FROM CYBERCRIME Volume 7 No.3 Sept-Dec 2020
Ida Musofiana Aji Sudarmaji Ira Alia Maerani
2 Ridwan Mansur's opinion is quoted in Nevey Varida Ariani, Implementation of Law Number 11
of 2012 concerning the System Juvenile Justice to Protect Children's Interests, Media Hukum
Journal, Volume 21, Nomor 1, 2014, page 108-109.
3 Ivan Fauzi Raharja, Wise to Use Social Media Among Students According to Law Number 19
of 2016 Concerning Electronic Information and Transactions, Journal Selat, Volume 6,
Number 2, May 2019, page 236. A. INTRODUCTION In this
research technology in terms of the use of cyberspace. Along with the
development of the times, the use of technology cannot be separated from
people's lives. This makes technology vulnerable to crime. Children at the
age categorized as children following Law Number 11 of 2012 concerning
the Juvenile Criminal Justice System, namely under the age of 18 (eighteen)
years. The internet is currently attached to the lives of today's children,
even toddlers (babies under five years) can use gadgets to watch YouTube. According to their age, children fall into the category of groups
vulnerable to cybercrime. Children can easily access the internet anywhere
and anytime. Things like this open the opportunity to become victims of
cybercrime more easily. It was 2001 when the crime of a pornographic
website in Texas was exposed. The two perpetrators who became the
perpetrators were Indonesians. In 2006 the Polda Metro Jaya arrested Peter 202
ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani W. Smith (an Australian citizen) who molested 50 (fifty) children from
Indonesia, India, and Vietnam.4 W. Smith (an Australian citizen) who molested 50 (fifty) children from
Indonesia, India, and Vietnam.4 The benefits of the virtual world so far provide social changes in
people's lives, especially for children. Children are like a white paper, what
we write down, that's where it will be drawn or formed because children
have imitation qualities. Deviant behavior can occur in children. Data from
the Indonesian Child Protection Commission shows that in 2016 there were
496 cases of pornography and cybercrime.5 Specifics regarding legal protection towards children, the state pays
attention bypassing the Law Number 35 of 2014 which is the change of Law
Number 23 of 2002 regarding Child protection. These changes are to
emphasize the importance of weighting criminal sanctions and also fines for
criminals against children to provide a deterrent effect, as well to encourage
concrete steps for recover physically, psychologically, and also social
children as victims and/or children as criminals as an anticipatory step so
children as victims or not as perpetrators be the perpetrator of the same
crime later day. Currently, a new legal regime has been born, known as cyberlaw or
telematics law. 4 Data sourced from the Indonesian Child Protection Commission in 2011 2016.
5 Hardianto Djanggih, The conception of Legal Protection for Children as Victims of Cyber
Crimes Through Penal and Non Penal Approaches, Mimbar Hukum Journal, Volume 30 Nomor
2, June 2018, page 318.
6 Ivan Fauzi, Op Cit, Journal Selat, page 237.
7 Ibid
page 238 4 Data sourced from the Indonesian Child Protection Commission in 2011 – 2016. 5 Hardianto Djanggih, The conception of Legal Protection for Children as Victims of Cybe
Crimes Through Penal and Non Penal Approaches, Mimbar Hukum Journal, Volume 30 Nom
2, June 2018, page 318. 6 Ivan Fauzi, Op Cit, Journal Selat, page 237.
7 Ibid., page 238 A. INTRODUCTION Cyberlaw is internationally used for legal terms related to the
use of information and communication technology. Likewise, telematics law
is the embodiment of the convergence of telecommunications law, media
law, and informatics law. Other terms that are also used are the law of
information technology, virtual world law, and cyberspace law. These terms
were born considering activities carried out through computer systems
networks and communication systems both locally and globally (internet) by
utilizing computer system-based information technology which is an
electronic system can be seen virtuallay.6 The emergence of this virtual version of social networks affects the
relationship between people, generally teenagers. The community site
above was created to fulfill the desire of individuals to communicate without
any time and space restrictions. Not infrequently social networks often have
the potential to influence a person's thinking patterns and shape individual
personalities. A large number of social media users among junior and senior
high school students at this time did not escape its spread to all corners of
the country. However, these social media users have very little information
about the existence of Law Number 19 of 2016 concerning Electronic
Information and Transactions, which is a legal product that limits the
freedom of social media users to be wiser in its use, especially among
teenagers.7 As we know, teenagers are the largest community in Indonesian
society who use social media regularly. The initial reason they are very ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 203 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani active in using social media is to seek attention, ask for opinions, and foster
an image, but over time they become dependent. Although social media has
a positive impact on adolescents, when it is difficult for them to escape from
activities related to social media, it will have a less positive impact. Several
studies show that due to the excessive use of social media, adolescents are
found to experience incongruence in their self-concept.8 Indonesian Internet User Services Association revealed the results of
its released survey 2016 on children of internet users. 8 Pamela Felita and Friends, Use of Social Media and Self Concept in Teens, Psikologi MANASA
Journal, Volume 5 Nomor 1, March 2016, page 31
9 Ministry of Communication and Information of the Republic of Indonesia, Indonesian
Internet User Penetration and Behavior, https://aptika.kominfo.
go.id/index.php/berita/274-rilis-hasil-survei-penetrasi-dan-perilaku-pengguna-internet-
indonesia-tahun-2018, acceded on 22 September 2020, 09.55 o’clock
10 Setiono, Understanding of Legal Research Methodology Postgraduate Law studies of Sebelas
Maret University, 2010, page 20 A. INTRODUCTION The result of a survey
showed 76.4% were not safe on the internet for the child.9 This is because
of the internet or information technology is an instrument potential criminal
instrument, and the second indicates the need for immediate Fixing the
legal sector in the field of child protection as victims of cybercrime in
Indonesia. The researcher in this case formulates several problem
formulations including How is legal protection for children from cybercrime? Then, Providing Legal Protection for Child Victims from Cybercrime. 204
ASPECTS OF LEGAL PROTECTION FOR Jurnal Pembaharuan Hukum
CHILDREN FROM CYBERCRIME Volume 7 No.3 Sept-Dec 2020
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
8 Pamela Felita and Friends, Use of Social Media and Self Concept in Teens, Psikologi MANASA
Journal, Volume 5 Nomor 1, March 2016, page 31
9 Ministry of Communication and Information of the Republic of Indonesia, Indonesian
Internet User Penetration and Behavior, https://aptika.kominfo.
go.id/index.php/berita/274-rilis-hasil-survei-penetrasi-dan-perilaku-pengguna-internet-
indonesia-tahun-2018, acceded on 22 September 2020, 09.55 o’clock
10 Setiono, Understanding of Legal Research Methodology Postgraduate Law studies of Sebelas
Maret University, 2010, page 20 11 Andi Hamzah, Indonesian Criminal and Criminal System from Retribution to Reform, Printed
for 2, Pardnya Paramita, Jakarta, page 16.
12 Faizin Sulitio, etc, Alternative Model of Crime of Cyber Pornography Crime, Arena Hukum
Journal, Volume 9, number 3, 2016, page 359.
13 Ibid., C. RESULT AND DISCUSSION RESULT AND DISCUSSION
1. The Concept of Child Protection as Victims of Cyber Crime p
y
According to Phillips in his book A First Book English Law, as
quoted by Andi Hamzah, which is seen as a goal the current
punishment is a variation from deterrent forms, good addressed to
violations of the law itself as well as those who have potential become
criminals, protection to society from evil deeds, improvement or reform
to criminals. The latter is the most modern and popular today, not only
for purposes improve imprisonment conditions but also looking for
other alternatives that are not criminal in fostering lawbreakers.11 Frequently asked questions criminal law experts in various
seminars are the question of how to criminal law should or should at
this time and future. This question is related to the field of criminal law
politics (penal policy), namely how better criminal law should be it is
created, structured, and used to organize or controlling human behavior
in Public. especially to cope with crimes committed by members society
as well as by the authorities.12 Crime is a deviant behavior that will always exist and cling to
every form of society. Crime is the oldest social problem therefore must
be overcome. Judging from the consequences of crime can disturb or
destroy and hinder the achievement of national goals and also prevent
optimal use of national sources Sudarto stated that the use of means
penal in tackling crime cannot be released from its connection with
legal policy criminal or political criminal law (criminal law policy). In this
connection, then Soedarto argued that implementing criminal law
politics meant making an assessment and elections to achieve statutory
results criminal at best, in a sense meet the requirements of justice and
efficiency. Efforts to find alternatives penalties for deprivation of liberty in
the short term were also supported by the Union Nations. In the 2nd
UN Congress regarding "The Prevention of Crime and Treatment of
Offenders ”in 1960 London recommended that the practice is thorough
removal short term imprisonment is not maybe. only realistic solutions
can be achieved by reducing the number of uses.13 That gradual
reduction by increasing substitute forms or alternative to conditional
punishment, supervision/probation, fines, work outside the agency. and
actions others that do not contain deprivation independence. B. RESEARCH METHODS Legal research is closely related to the concept of law used that
according to Soetondyo Wigyosoebroto of the five legal concepts, these
legal concepts are as follows: 1. Law is the principle of truth and justice that applies naturally and
applies universally; 2. Law is positive norms in the statutory system; 3. Law is what is decided by the judge (in concreto) and is systemized as
a judge-made law; 4. Laws are institutionalized social behavior patterns that exist as empirical
social variables; and 5. Law is a manifestation of symbolic meanings of social behavior as seen
in their interactions.10 5. Law is a manifestation of symbolic meanings of social behavior as seen
in their interactions.10 This study uses a descriptive-analytical literature method. Analytical
descriptive research can be done without using a basic supposition or
proposition, but it has begun with an outline of the thoughts and problems
to be studied. The literature method is one of the data collection methods
used in social research methodology to trace event record data ”. The
literature used in this research is sourced from journals, books, and other
online literature contains the concept of legal protection for child victims of
social media crimes in general as data in this paper. 204
ASPECTS OF LEGAL PROTECTION FOR Jurnal Pembaharuan Hukum
CHILDREN FROM CYBERCRIME Volume 7 No.3 Sept-Dec 2020
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
8 Pamela Felita and Friends, Use of Social Media and Self Concept in Teens, Psikologi MANASA
Journal, Volume 5 Nomor 1, March 2016, page 31
9 Ministry of Communication and Information of the Republic of Indonesia, Indonesian
Internet User Penetration and Behavior, https://aptika.kominfo. go.id/index.php/berita/274-rilis-hasil-survei-penetrasi-dan-perilaku-pengguna-internet-
indonesia-tahun-2018, acceded on 22 September 2020, 09.55 o’clock
10 Setiono, Understanding of Legal Research Methodology Postgraduate Law studies of Sebelas
Maret University, 2010, page 20 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 204 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani C. RESULT AND DISCUSSION Sudarto had a very good reason relation to cybercrime
phenomenon this one of which resulted in the child being victim, so to
overcome it must with the formulation and use of legislation precisely
so that the purpose of that law can be achieved. In Indonesia, child ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 205 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani protection from harmful effects of cyber exploitation can be found in
various laws, namely: protection from harmful effects of cyber exploitation can be found in
various laws, namely: a. Article 38 paragraph (3) of the Law Number 32 of 2002 concerning
The broadcast stated that “Fill broadcast must provide protection
and empowerment to the public specifically, namely children and
adolescents.14 b. Article 15 Law Number 44 2008 about the Pornography states that
“Everyone obliged to protect children from pornography influence
and prevent children's access to information pornography.15 c. Article 43 of Law Number 33 of 2009 concerning Film emphasized
that "Business actors movies are prohibited from doing dubbing
import movie sound into language Indonesia, except for imported
films educational interests and/or research.16 d. Article 27 of Law Number 19 of 2016 concerning Change Law
Number 11 of 2008 concerning Information & Transactions
Electronic confirms prohibition of distribution and/or transmits
and/or creates Electronic Information accessible and/or Electronic
Documents has a charge that breaks decency and gambling.17 d. Article 27 of Law Number 19 of 2016 concerning Change Law
Number 11 of 2008 concerning Information & Transactions
Electronic confirms prohibition of distribution and/or transmits
and/or creates Electronic Information accessible and/or Electronic
Documents has a charge that breaks decency and gambling.17 g
y
g
g
e. Article 59 of Law Number 35 of 2014 concerning Change Law
Number 23 of 2002 on Child Protection states that "the Government
and other state institutions are obliged and responsible for provide
special protection to children in emergencies, children in conflict
with the law, children from minority groups and isolated exploited
children economic and/or sexual, child who is trafficked, the child
who is fall victim to abuse narcotics, alcohol, psychotropic drugs,
and other addictive substances (drugs), children victims of
kidnapping, sales and trafficking, victim's child.18 2. 14 Law Number 32 of 2002 concerning The broadcast. 14 Law Number 32 of 2002 concerning The broadcast.
15 Law Number 44 of 2008 about the Pornography.
16 Law Number 33 of 2009 concerning Film emphasized.
17 Law Number 19 of 2016 concerning Change Law Number 11 of 2008 concerning Information
& Transactions Electronic.
18 Law Number 23 of 2002 on Child Protection.
19 Indah Setyowati, Ida Musofiana, Juridical Studies Against Diversion In Criminal Justice
System, Jurnal Pembaharuan Hukum, Volume 6 Number 2, 2019, page 241. g
15 Law Number 44 of 2008 about the Pornography. 206
ASPECTS OF LEGAL PROTECTION FOR Jurnal Pembaharuan Hukum
CHILDREN FROM CYBERCRIME Volume 7 No.3 Sept-Dec 202
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
14 Law Number 32 of 2002 concerning The broadcast.
15 Law Number 44 of 2008 about the Pornography.
16 Law Number 33 of 2009 concerning Film emphasized.
17 Law Number 19 of 2016 concerning Change Law Number 11 of 2008 concerning Information
& Transactions Electronic.
18 Law Number 23 of 2002 on Child Protection.
19 Indah Setyowati, Ida Musofiana, Juridical Studies Against Diversion In Criminal Justice
System, Jurnal Pembaharuan Hukum, Volume 6 Number 2, 2019, page 241. C. RESULT AND DISCUSSION Legal Protection for Child Victims from Cybercrime
Indonesia itself as a country that has ratified the Convention on
the Rights of the Child through Presidential Decree Number 36 of 1990
dated August 25, 1990, has further integrated children's rights into
national law, namely into Law Number 23 of 2002 concerning Child
Protection19 which has been amended by-law number 35 of 2014. Frequently asked questions criminal law experts in various
seminars are the question of how to criminal law should or should at
this time and future. Law Number 19 of 2016 Although this law is not
specific regulates pornography, but regulations on that matter are also
regulated in this law. One of them is regulated in Article 27 paragraph Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 206 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani (1), namely: "Every people knowingly and without rights distribute
and/or transmit and/or make accessible Electronic Information and/or
Electronic Documents has a charge that violates decency. "The meaning
of decency regulated in the article includes related to pornography. The internet further supports the child sex tourism industry by
providing patrons of the detours a means by which to easily connect
with other like-minded steps throughput the world.20 Constitution in
protecting children from cyber exploitation, according to the author has
not been able to reach there is a cybercrime development resulting in
children as victims. Author's reason is that there are still forms of
cybercrimes that need to be criminalized and necessary also, formulate
the appropriate form of punishment for cybercriminals who resulted in
children as a victim. Protection efforts for victims of a criminal act is an effort to
recover losses that have been obtained by the victim. This will be more
interpreted if victims are directly involved in the process of settlement
of criminal cases. Enforcement law is a development effort which is
continuous aiming realizing national life and a safe, orderly, and
peaceful state dynamic environment in the world independent.21 The purpose of providing compensation is not others to develop
justice and the welfare of the victim as community members by
benchmarks in practice, victims are given rights and obligation to be
developed as people, children are also people. 20 Ahmad Taufan Damanik, The Efforts of ASEAN-ACWC in Addressing the Trend of Sexual
Crimes Against Children Online”, Makalah, Conference on Sexual Crime against Children
Online: Law Enforcement and Regional Cooperation, Jakarta, 29 – 30 October 2012.
21 Thantawi, et al., Protection of Victims of Crime Cyber Crime in the Indonesian Criminal Law
System, Journal Ilmu Hukum, Vol. .2, No. 1, 2014, page. 39.
22 Hardianto Djanggih, The conception of Legal Protection for Children as Victims of Cyber
Crimes Through Penal and Non Penal Approaches, Mimbar Hukum Journal, Volume 30
Number 2, June 2018, page 323. y
p g
22 Hardianto Djanggih, The conception of Legal Protection for Children as Victims of Cyber
Crimes Through Penal and Non Penal Approaches, Mimbar Hukum Journal, Volume 30
Number 2, June 2018, page 323. Jurnal Pembaharuan Hukum ASPECTS OF LEGAL PROTECTION FOR
Volume 7 No.3 Sept-Dec 2020 CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
207
20 Ahmad Taufan Damanik, The Efforts of ASEAN-ACWC in Addressing the Trend of Sexual
Crimes Against Children Online”, Makalah, Conference on Sexual Crime against Children
Online: Law Enforcement and Regional Cooperation, Jakarta, 29 – 30 October 2012.
21 Thantawi, et al., Protection of Victims of Crime Cyber Crime in the Indonesian Criminal Law
System, Journal Ilmu Hukum, Vol. .2, No. 1, 2014, page. 39.
22 Hardianto Djanggih, The conception of Legal Protection for Children as Victims of Cyber
Crimes Through Penal and Non Penal Approaches, Mimbar Hukum Journal, Volume 30
Number 2, June 2018, page 323. C. RESULT AND DISCUSSION Therefore, required
strict, simple regulations, and easy to understand so it can avoid
discrimination in the application by law enforcement and intimidation
from certain parties which exacerbates it the condition of the victim in
its suffering prolonged. g p
g
In the development of laws and regulations in Indonesia,
penalties for compensation disadvantages are not only the domain of
civil law but has also entered into criminal law. Development this
happens because it is increasing world community attention to access
victims of criminal acts in the justice system criminal.22 The penalty for damages is a crime that obliges someone who
has acted detrimentally to others to pay some money or goods to the
injured person. The existence of the Criminal Code which is Dutch heritage that
was formed in 19th century did not recognize the punishment of
change loss. This is because the paradigm at the time of its formation
more to retribution (retributive). However, Article 14c of the Criminal ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 207 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Code has provisions conditional criminal stating that judge in imposing
conditional punishment against the perpetrator of a criminal act other
than determining general conditions can also impose special conditions
in the form of replacement losses that must be met before the term the
experiment ended. Apart from the Criminal Code, it is deep Another
positive criminal law provision about indemnity is also known. Corruption Criminal Act in Article 34 regulates additional penalties in the
form of payment of replacement money in the amount as much as
possible as wealth objects obtained from corruption. In this case, the compensation is given to the state because the
state
is
the
victim
(collective
victim). In
the
context
of
pornographycyber then this compensation sanction can also be in the
form of collective victims represented by the state and their use for
mental rehabilitation victims of pornography. Also, compensation can
be given to people who directly fall victim to pornography cyber. Based on the results of interviews with Police investigators, a
special criminal investigation Unit 1 in the section of cybercrime,
explained that during the pandemic era that had started in Indonesia
last March 2020. 23 Nanung, Reksrimsus Investigator At The Central Java Police, Cybercrime Unit 1, 26 October
2020 C. RESULT AND DISCUSSION The results of the police report in Unit 1, only one
case of a child victim of cybercrime in the Central Java Regional
Police.23 in the implementation of legal protection for victims by the
articles violated by the perpetrator. since the perpetrator is a child, then
diversion is done. Sanctions for damages as referred to has stated above is the
sanction intended to protect victims of action criminal. Apart from that,
there are still some other benefits that provide added value on
compensation sanctions if included in the new KUHP. 208
ASPECTS OF LEGAL PROTECTION FOR Jurnal Pembaharuan Hukum
CHILDREN FROM CYBERCRIME Volume 7 No.3 Sept-Dec 2020
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
23 Nanung, Reksrimsus Investigator At The Central Java Police, Cybercrime Unit 1, 26 October
2020 Book: Ahmad Taufan Damanik, 2012, The Efforts of ASEAN-ACWC in Addressing the
Trend of Sexual Crimes Against Children Online, Paper Conference on
Sexual Crime against Children Online: Law Enforcement and Regional
Cooperation, Jakarta; Ahmad Taufan Damanik, 2012, The Efforts of ASEAN-ACWC in Addressing the
Trend of Sexual Crimes Against Children Online, Paper Conference on
Sexual Crime against Children Online: Law Enforcement and Regional
Cooperation, Jakarta; D. CONCLUSION D. CONCLUSION
The conclusion in this article is that it is preferable to use social media
following applicable regulations and subject to the Law of the Republic of
Indonesia Number 19 of 2016 concerning Amendments to Law of the
Republic of Indonesia Number 11 of 2008 concerning Electronic Information
and Transactions. As well as the need for an active role from the community
and parents in supervising children in using social media so that they are
not entangled with criminal sanctions and/or compensation. 208
ASPECTS OF LEGAL PROTECTION FOR Jurnal Pembaharuan Hukum
CHILDREN FROM CYBERCRIME Volume 7 No.3 Sept-Dec 2020
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
23 Nanung, Reksrimsus Investigator At The Central Java Police, Cybercrime Unit 1, 26 October
2020 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Journal: Hardianto Djanggih, The conception of Legal Protection for Children as Victims
of Cyber Crimes Through Penal and Non Penal Approaches, Mimbar
Hukum Journal, Volume 30 No. 2, June 2018; Hardianto Djanggih, The conception of Legal Protection for Children as Victims
of Cyber Crimes Through Penal and Non Penal Approaches, Mimbar
Hukum Journal, Volume 30 No. 2, June 2018; Ivan Fauzi Raharja, Wise to Use Social Media Among Students According to Law
Number 19 of 2016 Concerning Electronic Information and
Transactions, Journal Selat, Volume 6, Number 2, May 2019; Nevey Varida Ariani, Implementation of Law Number 11 of 2012 concerning the
System Juvenile Justice to Protect Children's Interests, Media Hukum
Journal, Volume 21, Nomor 1, 2014; Pamela Felita and Friends, Use of Social Media and Self Concept in Teens,
Psikologi MANASA Journal, Volume 5 Nomor 1, March 2016; Rochmawati, Weny, Internet Utilization of Behavior (Descriptive Study of Using
the Internet for Entertainment and Academic Interests among
Children in the City of Surabaya), Journal Libri-Net, Vol. 1, No. 1,
2012; S., Kandi Aryani, Youth Acceptance of Pornography Discourse on Sex Sites in
Online Media, Journal Penelitian Dinamika Sosial, Vol. 7, 2008; Setiono, Understanding of Legal Research Methodology Postgraduate Law
studies of Sebelas Maret University, 2010; Sulistio, Faizin, et al., Alternative Model of Crime of Cyber Pornography Crime,
Journal Arena Hukum, Vol. 9, No. 3, 2016; Jurnal Pembaharuan Hukum ASPECTS OF LEGAL PROTECTION FOR
Volume 7 No.3 Sept-Dec 2020 CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
209
Sulistio, Faizin, et al., Alternative Model of Crime of Cyber Pornography Crime,
Journal Arena Hukum, Vol. 9, No. 3, 2016;
Thantawi, et al., Protection of Victims of Crime Cyber Crime in the Indonesian
Criminal Law System, Journal Ilmu Hukum, Vol. .2, No. 1, 2014;
Ulinnuha, Masyari, Protecting Children from Internet Negative Content: A Study
of Children's Web Browsing, Jurnal Sawwa, Vol. 8, No. 3, 2013;
Legislation:
KUHP;
Law Number 32 of 2002 concerning The broadcast;
Law Number 44 of 2008 about the Pornography;
Law Number 33 of 2009 concerning Film emphasized; Thantawi, et al., Protection of Victims of Crime Cyber Crime in the Indonesian
Criminal Law System, Journal Ilmu Hukum, Vol. .2, No. 1, 2014; Ulinnuha, Masyari, Protecting Children from Internet Negative Content: A Study
of Children's Web Browsing, Jurnal Sawwa, Vol. 8, No. Journal: 3, 2013; Jurnal Pembaharuan Hukum ASPECTS OF LEGAL PROTECTION FOR
Volume 7 No.3 Sept-Dec 2020 CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
209
Legislation:
KUHP;
Law Number 32 of 2002 concerning The broadcast;
Law Number 44 of 2008 about the Pornography;
Law Number 33 of 2009 concerning Film emphasized; Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 Legislation: ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
209 Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 209 Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani
Law Number 35 of 2014 Change Law Number 23 of 2002 on Child Protection;
Law number 19 of 2016 change Law 11 of 2008 Concerning Electronic
Information and Transactions;
Website:
Ministry of Communication and Information of the Republic of Indonesia,
Indonesian
Internet
User
Penetration
and
Behavior,
https://aptika.kominfo.go.id/index.php/berita/274-rilis-hasil-survei-
penetrasi-dan-perilaku-pengguna-internet-indonesia-tahun-2018;
The Indonesian Child Protection Commission in 2011 – 2016. www.kpai.go.id; Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Law Number 35 of 2014 Change Law Number 23 of 2002 on Child Protection;
Law number 19 of 2016 change Law 11 of 2008 Concerning Electronic
Information and Transactions;
Website:
Ministry of Communication and Information of the Republic of Indonesia,
Indonesian
Internet
User
Penetration
and
Behavior,
https://aptika.kominfo.go.id/index.php/berita/274-rilis-hasil-survei-
penetrasi-dan-perilaku-pengguna-internet-indonesia-tahun-2018;
The Indonesian Child Protection Commission in 2011 – 2016. www.kpai.go.id; Law Number 35 of 2014 Change Law Number 23 of 2002 on Child Protection;
Law number 19 of 2016 change Law 11 of 2008 Concerning Electronic
Information and Transactions; Law Number 35 of 2014 Change Law Number 23 of 2002 on Child Protection;
Law number 19 of 2016 change Law 11 of 2008 Concerning Electronic
Information and Transactions; Website: Ministry of Communication and Information of the Republic of Indonesia,
Indonesian
Internet
User
Penetration
and
Behavior,
https://aptika.kominfo.go.id/index.php/berita/274-rilis-hasil-survei-
penetrasi-dan-perilaku-pengguna-internet-indonesia-tahun-2018;
The Indonesian Child Protection Commission in 2011 – 2016. www.kpai.go.id; 210
ASPECTS OF LEGAL PROTECTION FOR
CHILDREN FROM CYBERCRIME
Ida Musofiana, Aji Sudarmaji, Ira Alia Maerani Jurnal Pembaharuan Hukum
Volume 7 No.3 Sept-Dec 2020 210
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https://openalex.org/W2986808514
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https://www.cb.cityu.edu.hk/ef/doc/GRU/WPS/GRU%232019-030%20Benjamin.pdf
|
English
| null |
Predicting mid-life capital formation with pre-school delay of gratification and life-course measures of self-regulation
|
Journal of economic behavior & organization
| 2,020
|
cc-by
| 16,267
|
a b s t r a c t How well do pre-school delay of gratification and life-course measures of self-regulation
predict mid-life capital formation? We surveyed 113 participants of the 1967–1973 Bing
pre-school studies on delay of gratification when they were in their late 40’s. They re-
ported 11 mid-life capital formation outcomes, including net worth, permanent income,
absence of high-interest debt, forward-looking behaviors, and educational attainment. To
address multiple hypothesis testing and our small sample, we pre-registered an analysis
plan of well–powered tests. As predicted, a newly constructed and pre-registered mea-
sure derived from preschool delay of gratification does not predict the 11 capital formation
variables (i.e., the sign-adjusted average correlation was 0.02). A pre-registered composite
self-regulation index, combining preschool delay of gratification with survey measures of
self-regulation collected at ages 17, 27, and 37, does predict 10 of the 11 capital formation
variables in the expected direction, with an average correlation of 0.19. The inclusion of
the preschool delay of gratification measure in this composite index does not affect the
index’s predictive power. We tested several hypothesized reasons that preschool delay of
gratification does not have predictive power for our mid-life capital formation variables. © 2019 The Authors. Published by Elsevier B.V. y
This is an open access article under the CC BY license. ( http://creativecommons.org/licenses/by/4.0/ ) his is an open access article under the CC BY license. ( http://creativecommons.org/licenses/by/4.0/ ) ✩ Authors are listed in alphabetical order. Sadly, our extraordinary friend and coauthor, Walter Mischel, who led the Bing study starting with the first
experiments in 1967, passed away during the editorial process. The Bing Longitudinal Project was supported by a number of grants from NIMH and NSF
to Walter Mischel and Yuichi Shoda, of which the most recent are National Institutes of Health Grant MH39349 and National Science Foundation Grant
BCS-0624305 . Research reported in this publication was also supported by the National Institute on Aging of the National Institutes of Health under
Award Numbers P30AG034532 and P01AG005842 and the Pershing Square Fund for Research on the Foundations of Human Behavior. The content is the
sole responsibility of the authors and does not necessarily represent the official views of any of these grant agencies. For her initial work preparing the
economics survey, the authors thank Dawn DeGere. Journal of Economic Behavior and Organization journal homepage: www.elsevier.com/locate/jebo Predicting mid-life capital formation with pre-school delay of
gratification and life-course measures of self-regulation ✩ Daniel J. Benjamin a , b , David Laibson b , c , Walter Mischel d , Philip K. Peake e ,
Yuichi Shoda f , Alexandra Steiny Wellsjo g , ∗, Nicole L. Wilson h a Center for Economic and Social Research and Department of Economics, University of Southern California, Los Angeles, CA 900
United States b National Bureau of Economic Research, Cambridge, MA 02138, United States
c Department of Economics, Harvard University, Cambridge, MA 02138, United States
d Department of Psychology, Columbia University, New York, NY 10027, United States
e Department of Psychology, Smith College, Northampton, MA 01063, United States
f Department of Psychology, University of Washington, Seattle, WA 98195, United States
g Department of Economics, University of California, Berkeley, CA 94720, United States
h Department of Management, Lundquist College of Business, University of Oregon, Eugene, OR 97403, United States a r t i c l e
i n f o
Article history:
Received 9 May 2018
Revised 28 May 2019
Accepted 17 August 2019
JEL Classifications:
D910
D140
I310
I210
I120
Keywords:
Self-regulation
Delay of gratification
Mid-life capital formation Predicting mid-life capital formation with pre-
school delay of gratification and life-course
measures of self-regulation Predicting mid-life capital formation with pre-
school delay of gratification and life-course
measures of self-regulation Dan Benjamin, University of Southern California/NBER
David Laibson, Harvard University/NBER
Walter Mischel, Columbia University
Philip K. Peake, Smith College
Yuichi Shoda, University of Washington
Alexandra Steiny Wellsjo, University of California,
Berkeley
Nicole L. Wilson, University of Oregon © 2019 by Benjamin, Laibson, Mischel, Peake, Shoda, Wellsjo & Wilson. All rights
reserved. Short sections of text, not to exceed two paragraphs, may be quoted
without explicit permission provided that full credit, including © notice, is given to the
source. © 2019 by Benjamin, Laibson, Mischel, Peake, Shoda, Wellsjo & Wilson. All rights
reserved. Short sections of text, not to exceed two paragraphs, may be quoted
without explicit permission provided that full credit, including © notice, is given to the
source. ARTICLE IN PRESS JID: JEBO Journal of Economic Behavior and Organization xxx (xxxx) xxx Journal of Economic Behavior and Organization xxx (xxxx) xxx E-mail address: alexsteiny@berkeley.edu (A.S. Wellsjo). 1. Introduction The capacity to self-regulate matters for a wide array of life outcomes. A critical component of self-regulation is the
ability to delay gratification. Working at Stanford’s Bing Nursery School beginning in the late 1960’s, Mischel and colleagues
conducted numerous experimental variations of the now classic “self-imposed delay of gratification” paradigm to examine
the cognitive and contextual mechanisms that affect preschoolers’ ability to wait to obtain a more desired outcome ( Mischel
et al., 1989; Mischel, 2014 ). Follow-up investigations beginning in the 1980’s reported that preschool waiting time predicted
better self-regulation as teenagers, especially among children who participated in certain experimental variations of the
preschool delay task ( Shoda et al., 1990 ). These “diagnostic” experimental conditions were challenging in that they com-
bined the physical presence of the tempting outcomes with requiring children to rely on their own spontaneously generated
strategies for coping with the frustration of waiting ( Shoda et al., 1990 ). Longitudinal research with the Bing cohort has
evolved into a multi-disciplinary exploration. Research on the Bing cohort now spans roughly 50 years, making it the
longest study of the delay of gratification paradigm. 1 Outcomes that have been studied include coping, social, and academic
competence ( Mischel et al., 1988; Mischel et al., 1989; Shoda et al., 1990 ), SAT scores ( Shoda et al., 1990 ), substance use
( Ayduk et al., 20 0 0 ), borderline personality features ( Ayduk et al., 2008 ), BMI ( Schlam et al., 2013 ), executive functioning,
and neural activation patterns ( Berman et al., 2013; Casey et al., 2011 ). The predictive power of the delay of gratification
task has also been documented in other samples with childhood and adolescent outcomes such as BMI ( Connell and
Francis, 2014; Francis and Susman, 2009; Seeyave et al., 2009 ), ADHD symptoms ( Campbell and von Stauffenberg, 2009 ),
academic competence ( Duckworth et al., 2013 ; Watts et al., 2018 ), and social competence ( Yang and Wang, 2007 ). As one
would expect, partial regression coefficients between preschool delay and subsequent life outcomes are lower when early
childhood, delay-related variables such as family background and early cognitive development are added to the model as
predictors ( Watts et al., 2018 ). 1. Introduction In previous research in the Dunedin cohort, other survey and observer-reported measures
of early life self-control have been found to be predictive of indices of adult outcomes, including socioeconomic status,
financial planfulness, and income ( Moffitt et al., 2011 ). The network of documented longitudinal relations of preschool delay of gratification are extensive. Nonetheless, both
media and academic accounts of the research commonly exaggerate the scope of the actual findings ( Watts et al., 2018 ). In
these accounts, preschool delay is cast as prognostic of almost all adult life milestones. In light of this common miscasting, it
is important that ongoing research empirically examine whether and how self-control actually relates to later life outcomes. The current research explores one such connection: the relation to mid-life capital formation. Economists conceptualize capital formation to include any costly activity that accumulates a resource (i.e., “capital”) that
generates future benefits. This definition implies that capital formation includes investments in human capital (e.g., working
hard at school or on the job to obtain knowledge/experience that will pay off later), investments in social capital (e.g.,
developing a network of professional contacts that will increase long-run professional success), and investments in financial
capital (e.g., saving now so that the household can spend more later). In the current paper, we revisit 113 individuals from the original Bing cohort, roughly 45 years after they participated
in the original experiments. Within this sample, we examine associations between measures of self-regulation based on
multiple assessments during the first four decades of life (including preschool delay) and a comprehensive array of mid-life
measures of capital formation. In addition, we also study preschool waiting time on its own as a predictor of mid-life capital
formation. To our knowledge, ours is the first attempt to look at these relationships because no cohort for which delay of
gratification was measured in childhood has become old enough to survey in mid-life until now, with the exception of the
Dunedin cohort, which is roughly ten years younger than the Bing cohort which we study. In the conclusion of this paper,
we compare our results to the small set of partially overlapping results from the Dunedin study and find a remarkable
degree of concordance despite the starkly different socioeconomic contexts of the two studies (and the somewhat younger
age of the Dunedin study participants). The rest of the paper proceeds as follows. D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 2 1. Introduction In Section 2 we discuss our sample. In Section 3 we describe our key mea-
sures and our pre-registered analysis plan. In Section 4 we state our pre-registered hypotheses and report our main results. In Section 5 we present additional results. In Section 6 we conclude and discuss the generalizability of our findings. The
Online Appendix (OA) contains further details about sample recruitment, survey design, and analyses, as well as a number
of additional and robustness analyses that we conducted. a b s t r a c t For helpful comments, the authors thank Constança Esteves-Sorenson and Charlie Sprenger; participants
at the Behavioral Economics Annual Meeting, the Drawing Links between Happiness, Health, and Self-regulation conference, the LABEL-IEPR Conference on
Children’s Decision Making, the NBER Summer Institute, and the RAND Summer Institute; and seminar participants at the University of California Berkeley,
the University of California San Diego, and the University of Southern California. ∗Corresponding author. E-mail address: alexsteiny@berkeley.edu (A.S. Wellsjo). https://doi.org/10.1016/j.jebo.2019.08.016
0167-2681/© 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license. ( http://creativecommons.org/licenses/by/4.0/ ) https://doi.org/10.1016/j.jebo.2019.08.016
0167-2681/© 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license. ( http://creativecommons.org/licenses/by/4.0/ ) https://doi.org/10.1016/j.jebo.2019.08.016
0167-2681/© 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license. ( http://creativecommons.org/licenses/by/4.0/ ) Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 JID: JEBO D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 ARTICLE IN PRESS ARTICLE IN PRESS JID: JEBO D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
3
[m3Gsc; November 16, 2019;10:34 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
[ D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
Table 1
Means, medians, and standard deviations of survey measures (prior to rank-normalizations). Survey Sample ( N = 113)
Full Sample ( N = 543)
Mean
Median
SD
Mean
Median
SD
p -value
Preschool measures
Male
37%
48%
0.01
Age at wait task (months)
51.6
52.0
5.6
52.1
52
5.7
0.36
Raw wait time (seconds)
509.7
556.0
369.1
470.1
440
367.7
0.20
Deviation from predicted preschool wait time (ln seconds)
0.08
0.51
2.37
0.00
0.27
2.30
0.66
Demographics at economics survey
Age
46.0
46.0
2.0
Married
79%
Has children (among those with information)
89%
Missing information on children
20%
White
94%
Asian
4%
Capital formation measures
Net worth ($ millions)
1.8
0.9
3.4
Permanent income ($ thousands)
131
78
147
Wealth-income ratio
13.5
10.1
14.6
High interest-rate debt ($ thousands)
1.2
0.0
2.6
Credit card misuse
–
–
–
Delay choice (% later choices)
85%
89%
14%
Savings rate
11.0%
10.0%
9.3%
Financial health
–
–
–
Educational attainment (years)
19.1
19.0
2.1
Forward-looking behaviors
–
–
–
Social Status
7.6
8.0
1.4
Notes : Entries of “-” are for measures with no natural scale. p -value is from an OLS regression of each variable on a constant and an
indicator variable for being in the survey sample. See OA Section II for more detailed information on sample size for each outcome. Table 1
Means, medians, and standard deviations of survey measures (prior to rank-normalizations). During 2012–2013, we contacted the 156 Bing cohort participants for whom we had current contact information (28% of
the 550 total), and we invited them to participate in an online survey. 2 The survey is reproduced in the OA. 113 participants
completed it (72% of those we contacted). Table 1 shows the demographics of our survey sample, as well as summary statistics for the measures we discuss in
Section 3 . ARTICLE IN PRESS Survey respondents averaged 46 years of age at the time they took the survey, 37% are male, and they are on
average white, married with children, well educated, and wealthy. Table 1 also compares our survey sample with the full
Bing sample on several variables we have for the full sample: sex, age during the delay of gratification task, and wait time. While our survey sample has fewer males (37% versus 48%), it does not differ substantially from the full Bing cohort on the
other variables. 3.1. Capital formation measures 3.1. Capital formation measures From the survey responses, we constructed the following 11 capital formation measures, which we use as dependent
variables in our study: • Net worth is the sum of assets minus the sum of debts. • Permanent income is defined as total household income (the average of reported income in the calendar year preceding
the survey and the calendar year when the respondent was age 35) divided by the reported number of adults living in
the household. Income at age 35 is inflation-adjusted to dollars in the year before the survey. • Permanent income is defined as total household income (the average of reported income in the calendar year preceding
the survey and the calendar year when the respondent was age 35) divided by the reported number of adults living in
the household. Income at age 35 is inflation-adjusted to dollars in the year before the survey. W
l h i
i
i
h di id d b
i • Wealth-income ratio is net worth divided by permanent income. • High interest-rate debt (reverse scaled) is the annual amount of interest above 6% paid on high-interest rate debt. For each
debt category, respondents selected an interest rate bucket for each type of debt. We assumed the interest rate is the
midpoint of the selected category. For each high interest-rate debt category we calculate the amount of interest above
6% as the dollar amount of debt multiplied by the interest rate above 6%. • Credit card misuse (reverse scaled) is an index with four components: high interest-rate debt, amount of carried credit
card debt (i.e., not paid off each month), a binary measure for having been denied a credit card in the last year, and
number of late payments on a credit card bill in the last year. • Credit card misuse (reverse scaled) is an index with four components: high interest-rate debt, amount of carried credit
card debt (i.e., not paid off each month), a binary measure for having been denied a credit card in the last year, and
number of late payments on a credit card bill in the last year. 2 The survey was approved by the institutional review boards at Harvard University and the University of Washington. All participants provided informed
consent. Many of the questions used to form the economic outcome variables were taken from the Cognitive Economics Project ( http://ebp-projects.isr. umich.edu/CogEcon/ ). 2. The Bing cohort and our survey sample The original Bing experiments were conducted during 1967–1973 (first described in Mischel and Ebbesen, 1970 ). They
included a total of 550 students from Stanford’s Bing Nursery School, aged about 4 years old (ranging from 2 to 6). Many of
the participants are children of Stanford faculty and staff. 1 Other long-running longitudinal studies with measures of self-regulation include the Dunedin Multidisciplinary Health and Development Study and the
NICHD Study of Early Child Care and Youth Development (SECCYD). The Dunedin cohort was first assessed at age 3 in the mid-1970’s and most recently
assessed at age 38 (with an ongoing assessment at age 45). While the Dunedin study did include parent and teacher survey measures and observer reports
of self-regulation in preschool, it did not include the Mischel delay of gratification task. The NICHD-SECCYD began collecting data in the early 1990’s when
children were aged 0-3 and collected the final wave of data in the mid-20 0 0’s when the children were 15 years old. The NICHD-SECCYD contains a version
of the Mischel delay of gratification task as well as parent and teacher survey measures of self-regulation from birth through early adolescence. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx • Delay choice is the percent of questions for which the respondent selected the “money later” instead of the “money
sooner” option in 40 tradeoff questions (20 questions with a choice between money today and thirty days from today
and 20 questions with a choice between money thirty and sixty days from today). • Savings rate is the self-reported percentage of income saved over the past few years, including retirement accounts and
all other savings. • Savings rate is the self-reported percentage of income saved over the past few years, including retirement accounts and
all other savings. • Financial health is an index comprised of three components: self-ratings about having enough money to meet needs,
difficulty in paying monthly bills, and current financial situation. • Financial health is an index comprised of three components: self-ratings about having enough money to meet needs,
difficulty in paying monthly bills, and current financial situation. y
p y
g
y
• Educational attainment (years of education) is calculated from reported degrees received. • Educational attainment (years of education) is calculated from reported degrees received. • Forward-looking behaviors is an index comprised of 5 equal-weighted components: measures of diet, exercise habits, and
BMI; smoking and alcohol behavior; preventative health and dental care; procrastination; and consideration of future
financial consequences. • Forward-looking behaviors is an index comprised of 5 equal-weighted components: measures of diet, exercise habits, and
BMI; smoking and alcohol behavior; preventative health and dental care; procrastination; and consideration of future
financial consequences. • Social status is self-reported placement in society on a 10-rung ladder. The ladder is described as representing where
people stand in the United States, with the top of the ladder being those who have the most money, the most education,
and the most respected job. • Social status is self-reported placement in society on a 10-rung ladder. The ladder is described as representing where
people stand in the United States, with the top of the ladder being those who have the most money, the most education,
and the most respected job. Summary statistics for these measures are in Table 1 . D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx We applied a rank-order inverse normal transformation to all final variables – including both the capital formation vari-
ables described above, as well as the self-regulation variables described below – because our statistical procedures have
better small-sample properties when the variables are normally distributed (for discussion of this transformation, see, e.g.,
Bishara and Hittner, 2012 ). In general, the raw data for our capital formation and self-regulation variables do not appear
to be normally distributed (see OA Sections I and II for distributions of the raw and transformed variables). The rank-
normalization approach has the advantage that we can apply it uniformly, rather than making different parametric assump-
tions for each variable (e.g., that wealth is log-normally distributed). To implement the transformation, we first calculated
the rank order of the subject within the sample and then used the inverse normal cdf to fit the ranks to a standard nor-
mal distribution. The transformation preserves the rank ordering of the original variables but alters their scaling and may
therefore affect the correlations with other variables. However, our transformation ensures that variables are approximately
normal by construction and minimizes the effects of outliers. Because the transformation relies on rank order, the variables
in our analyses represent a normalized measure of relative standing. 3 In prior work, wait times were adjusted to account for experimental condition by taking the difference from the mean wait time in a condition
(equivalent to a fixed-effects specification). In the current work, we instead treat the experimental conditions as random effects (as described below)
because mean wait time in a condition is estimated imprecisely when the per-condition sample size is small. Essentially, the random effects model predicts
the effect of an experimental condition as a weighted average of the observed mean wait time in the condition (conditional on age and sex) with the mean
wait time across experimental conditions. The larger the sample size for an experimental condition, the larger the weight assigned to the observed mean
wait time in that condition. In addition to the treatment of experimental conditions, RND also differs from wait time used in prior work because it accounts
for the child’s age and sex, it adjusts for the fact that observed wait times are capped at 15 minutes, and we measure wait time in log-seconds rather
than seconds. Finally, we note that our new rank-normalized measure may or may not be an improvement on the old absolute-deviation-from-the-mean
measure. See OA for more discussion of the methodological differences. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 ARTICLE IN PRESS
JID: JEBO D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 3.2. Self-regulation measures We study two measures of self-regulation. The first measure, rank-normalized delay (RND) , is derived from children’s
waiting times observed while participating in experimental variants of the self-imposed delay of gratification paradigm
when the children were in preschool. The second measure, rank-normalized self-regulatory index (RNSRI) , consists of four
equal-weighted components, each measuring self-regulation at a different age. One of the components of RNSRI is RND. We
will describe RND followed by RNSRI. RND is a new measure derived from preschool waiting that takes into account the fact that there were diverse treatment
conditions in the wait-task experiments differing in a number of ways that affected how long children on average waited
in each condition (including both “diagnostic” and “non-diagnostic” conditions, as we discuss below). 3 In addition, RND
also addressed the fact that wait times are censored (the criterion for successfully waiting was typically set to 15 min). Specifically, we constructed the RND measure in three steps: (a) we generated an expected log wait time for each treatment
condition using a tobit random-effects model, controlling for the child’s sex and age at the time of the experiment; (b) we
calculated the difference between each child’s actual log wait time and this expected log wait time; (c) we rank-normalized
these differences. We now explain steps (a) and (b) in more detail. For step (a), we used wait-time data from nearly all the Bing participants: a total of 543 children for whom data on
wait time, age at which they did the wait-time task, and sex is available. The sample size per experimental condition is 3
to 165 participants. Our survey data, which is from a subset of 113 of these participants across 20 of the 21 experimental
conditions, contains 1 to 34 participants per condition. For children who participated in multiple studies at the Bing School,
we use only the wait time from their first delay of gratification study (because the psychological meaning of the waiting
experiences in subsequent studies may be different from their first time). p
q
y
)
A histogram of the 543 raw wait times is shown in Fig. 1 A. The distribution has many observations at just a few seconds,
as well as a long right tail (censored at 900 s). For our model of expected wait times, we therefore adopted a log-normal Please cite this article as: D.J. Benjamin, D. Laibson and W. 3.1. Capital formation measures Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 3.2. Self-regulation measures Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
5
ARTICLE IN PRESS
JID: JEBO
[m3Gsc; November 16, 2019;10:34 ] [m3Gsc; November 16, 2019;10:34 ] JID: JEBO JID: JEBO D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
5
[m3Gsc; November 16, 2019;10:34 ] D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
istribution of wait times across Bing cohort participants and experimental conditions
stribution of wait times from the 543 children who participated in the original Bing experiments and for whom data on wait time, ag
the wait-time task, and sex is available. Figure 1A displays the distribution of individual wait times. Figure 1B displays the distributio
es across the 21 experimental conditions. ution. Fig. 1 B shows a histogram of mean raw wait times in the 21 experimental conditions. This distribution 5 5 Fig. 1. Distribution of wait times across Bing cohort participants and experimental conditions Fig. 1. Distribution of wait times across Bing cohort participants and experimental conditions
Notes : Distribution of wait times from the 543 children who participated in the original Bing experiments and for whom data on wait time, age at which
they did the wait-time task, and sex is available. Figure 1A displays the distribution of individual wait times. Figure 1B displays the distribution of mean
wait times across the 21 experimental conditions. Fig. 1. Distribution of wait times across Bing cohort participants and experimental conditions
Notes : Distribution of wait times from the 543 children who participated in the original Bing experiments and for whom data on wait time, age at which
they did the wait-time task, and sex is available. Figure 1A displays the distribution of individual wait times. Figure 1B displays the distribution of mean
wait times across the 21 experimental conditions. Notes : Distribution of wait times from the 543 children who participated in the original Bing experiments and for w
they did the wait-time task, and sex is available. Figure 1A displays the distribution of individual wait times. Figur
wait times across the 21 experimental conditions. distribution. Fig. ARTICLE IN PRESS We calculated an empirical-Bayes estimate of the expected ln(wait time) for each participant as the expected value
of the posterior distribution of the experimental condition’s random effect, treating the model’s parameter estimates as the
true parameter values. Table 2 shows the parameter estimates. The age coefficient of 0.08 (SE = 0.02) implies that, on average, every additional
month of age when participating in the wait-time task is associated with waiting roughly 8% longer. This is consistent with
prior work in the Bing and other samples on the development of delay of gratification strategies ( Mischel and Mischel, 1983 )
and directionally consistent with prior results on the correlation between age and delay (e.g., Carlson et al. (2018) find
a marginally significant positive correlation between wait time and age in the Bing and two other samples). The male
coefficient of −0.67 (SE = 0.23) implies that, controlling for age, males on average wait less than 50% as long as females—a
remarkably large difference between the sexes. Previous analyses of the Bing cohort have found mixed results with respect
to gender. Mischel and Underwood (1974) do find that males on average wait less than females when the data is aggregated
across experimental arms. However, other analyses of subsets of the Bing cohort have not found statistically different waiting
times between boys and girls ( Mischel and Moore, 1973, 1980; Mischel et al., 1988; Ayduk et al., 20 0 0 ), perhaps due to the
smaller sample sizes in these analyses. A recent exception is Carlson et al. (2018) , who find that girls waited significantly
longer than boys in the diagnostic subsample of Bing participants. In a different sample of 135 preschoolers in the 1980’s,
Carlson et al. (2018) find a similar, though statistically insignificant, sex difference. In a third sample of 540 preschoolers in
the 20 0 0’s, they find no sex difference. Turning to step (b), for each participant we calculate ln(subject’s actual wait time) – expected ln(wait time). If the par-
ticipant’s wait time was at the boundary of 900 s, we used the estimated model to calculate the expected value of the
uncensored wait time (conditional on the participant waiting at least 900 s, as well as on the participant’s age and sex) and
used this expected value in place of the participant’s actual wait time. ARTICLE IN PRESS D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
[m3Gsc; November 16, 2019;10:34 ] D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
[ D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
[ D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 6 Table 2
Parameter estimates from tobit random effects model of log preschool wait time, con-
trolling for age, sex, and experimental condition. Wait time in log-seconds
β1 : Age at delay of gratification task (months)
0.081 ∗∗∗
(0.020)
β2 : Male
−0.673 ∗∗∗
(0.227)
α: Constant
2.289 ∗∗
(1.071)
σε : SD of Individual-Specific Error
2.469 ∗∗∗
(0.101)
ση : SD of Random Effect (Experimental Condition)
0.786 ∗∗∗
(0.218)
N
543
Notes: Parameter estimates from a tobit random effects regression of preschool wait
time (measured in log-seconds) on age at delay of gratification task, sex, and a con-
stant. ση is the estimated standard deviation of the experimental condition effect, and
σε is the estimated standard deviation of the individual-specific error. Standard errors
are in parentheses. ∗p < 0.1, ∗∗p < 0.05, ∗∗∗p < 0.01. Table 2
Parameter estimates from tobit random effects model of log preschool wait time, con-
trolling for age, sex, and experimental condition. Notes: Parameter estimates from a tobit random effects regression of preschool wait
time (measured in log-seconds) on age at delay of gratification task, sex, and a con-
stant. ση is the estimated standard deviation of the experimental condition effect, and
σε is the estimated standard deviation of the individual-specific error. Standard errors
are in parentheses. ∗p < 0.1, ∗∗p < 0.05, ∗∗∗p < 0.01. month older β1 , the effect of being male β2 , the variance of the experimental-condition random effects σ 2
η , and the variance
of the individual-specific error σ 2
ε . month older β1 , the effect of being male β2 , the variance of the experimental-condition random effects σ 2
η , and the variance
of the individual-specific error σ 2
ε . p
ε
We estimated the tobit random-effects model by maximum likelihood (using the GLLAMM procedure in Stata version
14.0). Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 4 As we report in Section 5 , contrary to what we expected, the predictive power of the self-regulation measures does not increase as the age of mea-
surement gets closer to the age at which we measure the capital formation variables. 3.2. Self-regulation measures 1 B shows a histogram of mean raw wait times in the 21 experimental conditions. This distribution appears
to have a long right tail, so we also assumed that the condition means are log-normally distributed. We estimated the following tobit model of expected wait time: y ∗
gi = α + β1 × ag e gi + β2 × mal e gi + ηg + ε i , where i indexes individuals; g indexes experimental conditions; y ∗
gi is the underlying, uncensored, log wait time; ag e gi is age
in months at time of the delay task ; mal e gi is a binary indicator of sex; ηg is the group random effect ∼N( 0 , σ 2
η ) , and ε gi is
an individual-specific error ∼N( 0 , σ 2
ε ) . We modeled the effect of experimental condition as a random effect to deal with
the very small sample sizes in some conditions. We used a tobit model to account for the maximum wait time of 900 s
(or 15 min) imposed in the original experiments. The model has five parameters: a constant term α, the effect of being one Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 3.3. Statistical approach With the data we had, we faced two main statistical challenges. First, our sample size is small: 113 survey respondents. Given the small magnitude of associations we anticipated, we expected to have low statistical power for detecting any
particular association between a self-regulation measure and a capital formation measure. Moreover, due to low power,
any association we identified as statistically significant would be likely to have an exaggerated effect-size estimate (e.g.,
Gelman and Carlin, 2014 ), a bias sometimes called the “statistical winner’s curse” (e.g., Ioannidis, 2008 ). Second, because
the self-regulation and capital formation variables we examine could yield more than 600 different hypothesis tests, we
faced a potential multiple-testing problem and its associated “data-mining” problems. Consequently, there would be a high
rate of false positives at a p -value threshold of 0.05. Moreover, if we adopted a family-wise significance threshold such as a
Bonferroni correction, we would exacerbate the problem of low power. To address the potential multiple-testing problem, we pre-registered and time-stamped on Open Science Framework
( https://osf.io/u39hg/ ) our coding of the variables and our planned series of analyses before calculating any correlations
between independent variables and dependent variables. 5 Moreover, we distinguished between four types of analyses: (i)
tests of primary hypotheses, (ii) tests of secondary hypotheses, (iii) ex-post analyses, and (iv) robustness checks. Our pre-
registration plan specified our tests of primary and secondary hypotheses and our robustness checks, and we committed
to identifying any ex-post analyses as such in the paper. We further committed to treat tests of secondary hypotheses
as exploratory. We pre-registered only two primary hypotheses, which we describe in Section 4 . We report some of our
secondary and ex-post analyses in Section 5 , and we report all of them in the OA Section V. Our planned series of analyses of our primary hypotheses were designed to address the challenge of our small sample
size. We planned four analyses. First, with each of our capital formation measures as a dependent variable, we run an OLS
regression on each measure of self-regulation, controlling for sex. (Regressions with the savings rate as the outcome also
control for permanent income.) While we report the regression results because they are simple and serve as inputs to our
other three tests, we do not use a p -value threshold to judge statistical significance due to the multiple-testing problem. D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 7 “general cognitive ability” (with the items selected based on their face validity for these concepts). We then factor-analyzed
the remaining 25 items within the Smith sample using principal axis factoring and oblique rotation. We labeled the four
oblique factors resulting from this analysis “attention,” “coping,” “goal pursuit,” and “concern for others”; see OA Section I-C
for a list of the items included in each of the six subscales. For each participant at each age, we created rank-normalized
subscale scores, giving equal weight to each item in the subscale. We then calculated a rank-normalized CCQ score for each
respondent at each age, giving equal weight to each of the 6 subscales and then rank-normalizing the resulting variable. We computed RNSRI by taking an equal-weighted mean of RND and rank-normalized CCQ score at ages 17, 27, and 37,
and then rank-normalizing the resulting variable. 5 In a slight abuse of terminology, we will often refer to “correlations” between our independent and dependent variables. As described below, what we
will actually report are coefficients from OLS regressions of our dependent variables on measures of self-regulation, controlling for sex and other variables.
Because both the independent and dependent variables are normalized, the regression coefficients can be interpreted as partial correlation coefficients.
6 For example, for an average correlation of 0.15 between our dependent variables and an independent variable, our simulations yielded on average 3.9
of 11 dependent variables associated with a p -value less than 0.05, 3.4 associated at a false discovery rate threshold of 0.1, and in 97% of simulations, we
Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 ARTICLE IN PRESS We then factor-analyzed
the remaining 25 items within the Smith sample using principal axis factoring and oblique rotation. We labeled the four
oblique factors resulting from this analysis “attention,” “coping,” “goal pursuit,” and “concern for others”; see OA Section I-C
for a list of the items included in each of the six subscales. For each participant at each age, we created rank-normalized
subscale scores, giving equal weight to each item in the subscale. We then calculated a rank-normalized CCQ score for each
respondent at each age, giving equal weight to each of the 6 subscales and then rank-normalizing the resulting variable. We computed RNSRI by taking an equal-weighted mean of RND and rank-normalized CCQ score at ages 17, 27, and 37, D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
7
ARTICLE IN PRESS
JID: JEBO
[m3Gsc; November 16, 2019;10:34 ] JID: JEBO JID: JEBO 5 In a slight abuse of terminology, we will often refer to “correlations” between our independent and dependent variables. As described below, what we
will actually report are coefficients from OLS regressions of our dependent variables on measures of self-regulation, controlling for sex and other variables.
Because both the independent and dependent variables are normalized, the regression coefficients can be interpreted as partial correlation coefficients.
6 For example, for an average correlation of 0.15 between our dependent variables and an independent variable, our simulations yielded on average 3.9
of 11 dependent variables associated with a p -value less than 0.05, 3.4 associated at a false discovery rate threshold of 0.1, and in 97% of simulations, we
Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre- ARTICLE IN PRESS We calculated this expected value using simulation,
using 10,0 0 0 draws from the model under the assumption that the parameter estimates in Table 2 are the true parameter
values. Our second measure, RNSRI, has four components: RND and survey measures of self-regulation measured at ages 17,
27, and 37. The latter three components are derived from the California Child Q-Set (CCQ), a set of personality descrip-
tors ( Block and Block, 1980 ). Versions of the CCQ were completed by participants’ parents with respect to their children’s
personality around age 17 and by the participants themselves around ages 27 and 37. 4 For each of the three ages, we constructed a measure of overall self-regulation from subsets of CCQ items, with items and
weights determined by the following procedure. First, a preliminary set of 37 CCQ items relating to self-control was selected
based on ratings by three of us (Peake, Shoda, and Wilson). These items were administered to an independent sample of 191
Smith College students. We conducted a principal components analysis, which found that all but one of these items loaded
positively on the first unrotated principal component. In our Bing survey sample, using ratings obtained at ages 17, 27, and
37, we then conducted a confirmatory principal component analysis on the remaining 36 items. We eliminated five items
from the analysis due to failure to demonstrate consistent positive loadings on the first unrotated principal component. From among the remaining 31 items, we formed two subscales, each consisting of 3 items, for “delay of gratification” and 4 As we report in Section 5 , contrary to what we expected, the predictive power of the self-regulation measures does not increase as the age of mea-
surement gets closer to the age at which we measure the capital formation variables. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
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JID: JEBO
[m3Gsc; November 16, 2019;10:34 ]
“general cognitive ability” (with the items selected based on their face validity for these concepts). In this Section, we describe the two primary hypotheses that represent the core of our study Hypothesis 1. We hypothesize that RNSRI will have a positive, but relatively modest, correlation with the measures of mid-
life capital formation. Although we predicted that self-regulation plays a significant role in development, given that many other factors (peers,
career choice, marital outcomes, family wealth, inheritance, chance, etc.) influence capital formation, we predicted that the
correlation between self-regulation and capital formation is modest. Specifically, our pre-registered prior distribution for the
correlation of RNSRI with each measure of mid-life capital formation had a mean of 0.15 (and standard deviation of 0.2). These predicted correlations are consistent with past results for these types of outcomes (e.g., Moffitt et al., 2011; Richard
et al., 2003 ). Tests of Hypothesis 1 are reported in the top panel of Table 3 . Of the 11 capital formation measures, 10 are associated
with the predicted sign. Six variables are significant at our FDR threshold of 0.1 (and the same six have p -values < 0.05):
net worth, credit card misuse, financial health, forward-looking behaviors, educational attainment, and permanent income. In an F -test, the (joint) null that all 11 capital formation measures have no association is rejected (empirical Wald test
p -value = 0.0 0 05). OA Section V-A and V-D contains details of all these analyses, Bayesian posterior means and standard
deviations for the correlation with each outcome, as well as pre-registered robustness analyses. The average observed correlation across all 11 variables, ¯r = 0.19 (SE = 0.04), is close to our pre-registered prior mean of
0.15. The average of our Bayesian posterior means is a correlation of 0.18. Hypothesis 2. On its own, RND (measured around age four) will have only a very small correlation with the measures of
mid-life capital formation. As specified in our pre-registration, we predicted very small correlations between our measure of preschool waiting
and measures of mid-life capital formation for three reasons (which we discuss further in Section 5 ). First, most of the
respondents to the capital formation survey were tested in experimental conditions that have previously been designated as
“non-diagnostic” (e.g., the experimenter suggested a specific self-regulation strategy to the child). Theoretical considerations
and earlier findings ( Shoda et al., 1990 ) led us to anticipate that variation within non-diagnostic conditions would generate
less predictive power than variation within diagnostic conditions. D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 8 Note that we originally hypothesized that the relationship between preschool waiting and mid-life capital formation
would be stronger in the “diagnostic” conditions of the original Bing Study (for example, see Shoda et al., 1990 ; and p. 5 of
our pre-registered analysis plan). Nevertheless, in order to maximize sample size, for tests of our primary hypotheses, we
pooled respondents who participated in diagnostic and non-diagnostic experimental conditions, as we did in some of the
prior work ( Mischel et al., 1988; Ayduk et al., 20 0 0 , 20 08; Schlam et al., 2013 ). Our pre-registered priors and corresponding
power calculations take into account this pooling across conditions. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 reject the (joint) null that all 11 variables each have no effect with an empirical Wald test p -value less than 0.05. Instead, for an average correlation of 0.05,
our simulations yielded on average 0.9 of 11 dependent variables associated with a p -value less than 0.05, 0.3 associated at a false discovery rate threshold
of 0.1, and in 18% of simulations, we reject the (joint) null that all 11 variables each have no effect with an empirical Wald test p -value less than 0.05. 8
D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
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[m3Gsc; November 16, 2019;10:34 ] 3.3. Statistical approach Second, to identify “significance” of associations between a self-regulation measure and a specific capital formation mea-
sure, we use a false discovery rate (FDR) threshold of 0.1. Compared with a family-wise error rate threshold, an FDR thresh-
old has greater statistical power, albeit with a different interpretation: rather than controlling the error rate of any particular
association, an FDR threshold of 0.1 means that among the set of “significant” associations, at most 10% are false positives. We compute the FDR of each association using the Benjamini and Hochberg (1995) algorithm, applied separately to the OLS
p -values for each of the two self-regulation measures. Third, for each self-regulation measure, we test for the joint significance of associations with our 11 capital formation
measures. We use a Wald test with a p -value threshold of 0.05. Because the standard Wald test p -value is biased downward
in small samples, we calculate an empirical p -value by comparing the observed Wald test statistic to a distribution of Wald
test statistics under the null, which we generate by permuting our data. Specifically, we randomly permute the values of the
dependent variable within the sample 10 0 0 times, and we re-calculate the Wald test statistic each time. We calculate our
empirical p -value as equal to the percentile where our observed Wald test statistic falls in the distribution of test statistics
from the permuted samples. Finally, to obtain an estimate of the magnitude of the association between each self-regulation measure and each cap-
ital formation measure, we report our Bayesian posterior mean and standard deviation for each coefficient. For each self-
regulation measure, we pre-registered our prior about the mean and standard deviation of the OLS coefficients for the 11
capital formation measures. We calculate the posterior mean and standard deviation for each coefficient assuming normal
distributions for both the prior and the observed OLS coefficient. Due to Bayesian “shrinkage” toward the prior, the Bayesian
posterior means do not suffer from the “statistical winner’s curse” bias ( Ioannidis, 2008 ) and can thus serve as our “best
guess” effect-size estimates. g
Before pre-registering our analysis, we used Monte Carlo (i.e., simulation) methods to confirm that given our priors, our
planned FDR and Wald tests were well-powered. 6 In Section 4 , we compare our pre-specified priors to what we found. In this Section, we describe the two primary hypotheses that represent the core of our study Second, as noted for Hypothesis 1, self-control is only one
of the many factors that influence mid-life capital formation. Third, the assessment of RND is separated in time from the
capital formation variables by over 40 years. Research and theory suggest that while many people are stable in self-control
over the life-course, many exhibit change, either towards increasing or decreasing self-control ( Berman et al., 2013; Casey
et al., 2011 ). In contrast, the other components of RNSRI average a separation of around 20 years from the capital formation
variables and represent an average of many indicators of self-regulation. The predictive power of an index consisting of
many items typically exceeds the predictive power of each of its components. Since RND is only one component of the 86-
item RNSRI, we expected it to have less predictive power. Specifically, our pre-specified prior distribution for the correlation
of RND with each measure of mid-life capital formation had a mean of 0.05. Tests of Hypothesis 2 are reported in the bottom panel of Table 3 . Of the 11 capital formation measures, 6 are positively
correlated with RND, but none are significantly associated at our FDR threshold of 0.1 (and all have p -values > 0.05). In an
F -test, the joint null of no effect for all 11 capital formation variables is not rejected (empirical Wald test p -value = 0.45). None of our pre-registered robustness analyses, including alternative measures of childhood wait time, substantively affects
the conclusions from the bottom panel of Table 3 (see OA Section V-D). The average observed correlation across all 11 variables, ¯r = 0.02 (SE = 0.05), is close to our pre-registered prior mean of
0.05. The average of our Bayesian posterior means is a correlation of 0.03. ation m
e contro ation m
e contro 5.1. Mechanisms As explained above, three mechanisms motivated our pre-registered hypotheses that the index of self-regulatory mea-
sures would be moderately predictive, while the preschool delay of gratification task would be very weakly predictive. These
mechanisms flow from standard statistical properties and our understanding of the psychological processes underlying the
cumulative findings of past empirical studies: 1. The index of self-regulatory measures is comprised of 86 responses per participant, whereas the preschool delay of
gratification task is a single behavioral task. An index of similar measures tends to have a higher signal-to-noise ratio
than its components. 1. The index of self-regulatory measures is comprised of 86 responses per participant, whereas the preschool delay of
gratification task is a single behavioral task. An index of similar measures tends to have a higher signal-to-noise ratio
than its components. p
2. The preschool delay of gratification task is measured using a diagnostic variant of the task for 34 of our 113 participants;
the remaining 79 participants experienced a non-diagnostic variant of the pre-school delay of gratification task. Pooling
across diagnostic and non-diagnostic conditions weakens the correlation with outcome variables. 2. The preschool delay of gratification task is measured using a diagnostic variant of the task for 34 of our 113 participants;
the remaining 79 participants experienced a non-diagnostic variant of the pre-school delay of gratification task. Pooling
across diagnostic and non-diagnostic conditions weakens the correlation with outcome variables. g
g
3. The index of self-regulatory measures is comprised of questions that are measured throughout the life course up to
age 37 (specifically, ages 4, 17, 27, and 37), whereas the preschool delay of gratification task is measured at age 4. Self-
regulation measured closer in time to the observed outcomes will be more strongly related to them. 3. The index of self-regulatory measures is comprised of questions that are measured throughout the life course up to
age 37 (specifically, ages 4, 17, 27, and 37), whereas the preschool delay of gratification task is measured at age 4. Self-
regulation measured closer in time to the observed outcomes will be more strongly related to them. We now discuss (pre-registered) secondary and (non-pre-registered) ex-post analyses that shed some light on these
mechanisms. We find support for the first, suggestive evidence in favor of the second, and no support for the third. Table 4 Table 4
Mean of coefficients from regressions of the 11 capital formation measures on RNSRI, RND, and individual CCQ items. Measure(s) of Self-Regulation
Type of Analysis
Mean of Coefficients Across Measures
of Capital Formation
RNSRI (includes RND and Age 17, 27, and 37 CCQ indices)
Pre-registered primary
0.19 (0.04)
RND
Pre-registered primary
0.02 (0.05)
RND (diagnostic condition)
Pre-registered secondary
0.07 (0.09)
RND (non-diagnostic condition)
Pre-registered secondary
−0.01 (0.06)
All CCQ items (avg over 85 items)
Ex-post
0.07 (0.03)
Age 17 CCQ items (avg over 23 items)
Ex-post
0.08 (0.05)
Age 27 CCQ items (avg over 31 items)
Ex-post
0.07 (0.02)
Age 37 CCQ items (avg over 31 items)
Ex-post
0.08 (0.02)
Notes : All regressions control for sex, and regressions with savings rate as the dependent variable also include control for permanent
income (coefficients on control variables not reported). For the regressions on CCQ items, the sign of each coefficient is adjusted to be
positive before the coefficients are averaged. Bootstrapped standard errors are in parentheses. e 4
n of coefficients from regressions of the 11 capital formation measures on RNSRI, RND, and individual CCQ items. from regressions of the 11 capital formation measures on RNSRI, RND, and individual CCQ items. Notes : All regressions control for sex, and regressions with savings rate as the dependent variable also include control for permanent
income (coefficients on control variables not reported). For the regressions on CCQ items, the sign of each coefficient is adjusted to be
positive before the coefficients are averaged. Bootstrapped standard errors are in parentheses. 7 Some of the outcome variables, such as the total years of education and income at age 35, precede the age 37 self-regulation measurement. In ex-post
analyses, we found that this does not drive our main results; excluding educational attainment, permanent income, and wealth-to-income ratio from the
primary outcomes, the average correlation between RNSRI and the remaining outcomes is 0.18. This is consistent with our finding that self-regulation
measured at age 37 is not a better predictor of our primary outcomes than measurements at age 17 and 27. Focusing on the three potentially problematic
outcomes, while educational attainment does appear to be most strongly predicted by self-regulation at age 37, permanent income and wealth-to-income
ratio are not. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 ARTICLE IN PRESS 0
D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
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[m3Gsc; November 16, 2019;10 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
[ D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
[ 10 mation measures, controlling for sex. (7)
(8)
(9)
Savings
rate
Financial
health
Education
years
−0.01
0.24 ∗∗
0.24 ∗∗∗
(0.09)
(0.09)
(0.09)
103
110
110
−0.06
−0.07
0.13
(0.09)
(0.09)
(0.09)
106
113
113
de control for permanent income (coefficients n ation
s
∗∗∗
9)
9)
ents n 5.1. Mechanisms To test the first mechanism, we evaluate the predictive power of each of the 85 components of the CCQ index compared
to the predictive power of the RNSRI index taken as a whole. In this ex-post analysis, we regressed each of the capital
formation measures on each of the 85 items in the RNCCQ indices (23 at age 17, 31 at age 27, and 31 at age 37) in separate
regressions (i.e., 85 CCQ items x 11 outcomes = 935 regressions). In Table 4 , we report the average coefficient across all 935
regressions of a capital formation measure on a rank-normalized CCQ item. This average correlation is 0.07 (SE = 0.03), which
is smaller than 0.19 (SE = 0.04), the average correlation reported above between the RNSRI index and each of the 11 capital
formation variables. In a related ex-post analysis, we found stronger correlations between self-regulation measures and the
capital formation measures when we use an index of capital formation variables rather than examining them separately. We
constructed the capital formation index by averaging the 11 capital formation measures for the 102 subjects with no missing
outcome variables and then rank-normalizing this average. The correlation between RNSRI and the capital formation index
is 0.43 (SE = 0.09), and 0.13 (SE = 0.10) between RND and the capital formation index; see OA. 7 The second mechanism is a potential explanation for the weak relationships we find with RND. The preschool delay of
gratification task was run in many different ways and only some of those trials were theorized to be diagnostic ( Shoda et al.,
1990 ). For example, in some trials the four-year-old participants were asked to think about the rewards in a particular way
that was hypothesized to facilitate delay (e.g., think of marshmallows as clouds). In other trials, the participants were not
given any suggested cognitive strategies. These two methods are referenced in the literature as suggested ideation and spon-
taneous ideation . In the original Bing studies, this ideation manipulation was crossed with a reward-exposure manipulation:
in some cases the rewards were exposed (elevating appetitive responses and thereby making delay relatively more difficult),
and in other cases the rewards were covered or removed. Experimental settings with spontaneous ideation and exposed Please cite this article as: D.J. Benjamin, D. Laibson and W. 5.1. Mechanisms Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 ARTICLE IN PRESS [m3Gsc; November 16, 2019;10:34 JID: JEBO JID: JEBO D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 11 rewards were theorized to be the most likely to be “diagnostic” of self-regulatory competencies. Indeed, it was only these
settings that produced the significant correlation between preschool wait time and subsequent adolescent outcomes, includ-
ing parental CCQ item ratings of adolescents employed in the current research and SAT scores ( Shoda et al., 1990 ). rewards were theorized to be the most likely to be “diagnostic” of self-regulatory competencies. Indeed, it was only these
settings that produced the significant correlation between preschool wait time and subsequent adolescent outcomes, includ-
ing parental CCQ item ratings of adolescents employed in the current research and SAT scores ( Shoda et al., 1990 ). Specifically, in our current data, only 34 participants were assessed in the diagnostic version of the preschool delay task. We pre-registered an analysis to examine the predictive power of RND separately for participants in the diagnostic and
non-diagnostic version of the preschool delay task, but we classified it as a secondary analysis because we anticipated large
standard errors due to the smaller sample sizes. Based on past results, we hypothesized that RND would have more pre-
dictive power among the diagnostic participants. As shown in Table 4 , in the diagnostic condition the average correlation
between RND and the 11 human capital measures is 0.07 (SE = 0.09), which, incidentally, is equal to the average correlation
between the individual components of the CCQ index and the 11 human capital measures. In contrast, the average correla-
tion between preschool wait time in the non -diagnostic conditions and the 11 human capital measures is −0.01 (SE = 0.06). Hence, our evidence provides suggestive support for the earlier finding that the diagnostic conditions are the relevant ones
for assessing individual differences in self-regulatory competence—albeit with two caveats, both related to the large stan-
dard errors: (i) the coefficients in both conditions are statistically indistinguishable from zero, and (ii) the diagnostic and
non-diagnostic correlations are not statistically distinguishable from each other. 5.1. Mechanisms If we take the point estimates at face value,
they suggest that once we level the playing field, by comparing single questions from the self-regulatory index to the diag-
nostic variant of the preschool delay task, the predictive power of preschool waiting time is identical to the typical reported
CCQ item assessing self-regulation later in the life-course. However, re-analysis of prior evidence raises questions about the
consistency of the differences between diagnostic and non-diagnostic conditions. In an ex-post analysis suggested in the
editorial process, we re-analyzed the correlation between BMI and preschool wait time in Schlam et al. (2013) separately
for the diagnostic and non-diagnostic subsamples. Of the 164 subjects in the BMI study, 37 were in a diagnostic condition
(25 of whom are also in the current survey sample). Contrary to our findings, the relationship between preschool delay
(measured as the deviation from the condition mean) and BMI is weaker in the diagnostic group ( r = −0.03, SE = 0.14) than
in the non-diagnostic group ( r = −0.24, SE = 0.08), though the difference is not statistically distinguishable. The correlations
between BMI and delay deviation are similar if we minimize the effect of outliers by winsorizing the data. See OA Section
V-C-8 for more detail. It is also worth noting that two studies using data from the NICHD-SECCYD find the expected negative
relationship between wait time in the diagnostic version of the preschool task and adolescent BMI ( Seeyave et al. (2009)
and Francis and Susman (2009) ). In light of all these results, evidence for the importance of the diagnostic methodology is
mixed. To test the third mechanism, in an ex-post analysis we examined the relationship between the date of measurement
and predictive power. Surprisingly to us, we found no relationship between the age at which the self-regulation questions
from the CCQ were administered and the predictive power of those questions for mid-life capital formation. Specifically,
the mean correlation with the capital formation measures for the questions administered at age 17 is 0.08 (SE = 0.05), the
mean correlation from the questions administered at age 27 is 0.07 (SE = 0.02), and the mean correlation for the questions
administered at age 37 is 0.08 (SE = 0.02) ( Table 4 ). 8 8 Similarly, there is limited evidence for an age trend in the predictive power of the indices of CCQ questions. In ex-post analysis, we found that the
average correlations between the mid-life capital formation measures and RNCCQ at ages 17, 27, and 37 are 0.15 (SE = 0.09), 0.17 (SE = 0.06), and 0.19
(SE = 0.05), respectively (see OA). 5.2. Excluding RND from the self-regulation measure 5.2. Excluding RND from the self-regulation measure 5.2. Excluding RND from the self-regulation measure We formulated our primary hypotheses in terms of RND and RNSRI, where the former self-regulation measure is a com-
ponent of the latter. Among our pre-registered secondary analyses, we also examined an RNCCQ index, which is otherwise
the same as RNSRI (i.e., it is composed of the CCQ indices from ages 17, 27, and 37) but excludes RND. This RNCCQ index
has correlation r = 0.16 (SE = 0.09) with RND in the survey sample. 9 Considering the predictive power of the RNCCQ index
on its own, we find that its mean correlation with the capital formation measures is 0.20 (SE = 0.04) whether we control for
RND (pre-registered secondary analysis) or not (ex-post analysis); see OA Sections V-B-2 and V-C-14 for more details. Thus,
the inclusion of RND in the RNSRI index does not change its predictive power for the capital formation measures. 5.3. Money now vs. money later as a predictor average correlations between the mid-life capital formation measures and RNCCQ at ages 17, 27, and 37 are 0.15 (SE = 0.09), 0.17 (SE = 0.06), and 0.19
(SE = 0.05), respectively (see OA).
9 The overall correlation between RND and RNCCQ is calculated by combining RNCCQ data across ages 17, 27, and 37. The correlations for each respective
age bucket are r = 0.06 (SE = 0.10), r = 0.23 (SE = 0.09), and r = 0.15 (SE = 0.10). Readers interested in other subcomponents of RNCCQ (including the factor-
analytic subscales) can find the associated correlations with RND in the OA. 10 For a review and methodological critique of money earlier vs. later questions, see Cohen et al (forthcoming) . For example, in principle choices in the
money earlier vs. later paradigm should reveal market interest rates, transactions costs, and trustworthiness of the experimenter, and should not measure
true time preferences (because money is temporally fungible). Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 8 Similarly, there is limited evidence for an age trend in the predictive power of the indices of CCQ questions. In ex-post analysis, we found tha
average correlations between the mid-life capital formation measures and RNCCQ at ages 17, 27, and 37 are 0.15 (SE = 0.09), 0.17 (SE = 0.06), and
(SE = 0.05), respectively (see OA).
9 The overall correlation between RND and RNCCQ is calculated by combining RNCCQ data across ages 17, 27, and 37. The correlations for each respe
age bucket are r = 0.06 (SE = 0.10), r = 0.23 (SE = 0.09), and r = 0.15 (SE = 0.10). Readers interested in other subcomponents of RNCCQ (including the fa
analytic subscales) can find the associated correlations with RND in the OA.
10 For a review and methodological critique of money earlier vs. later questions, see Cohen et al (forthcoming) . For example, in principle choices i
money earlier vs. later paradigm should reveal market interest rates, transactions costs, and trustworthiness of the experimenter, and should not me
true time preferences (because money is temporally fungible).
Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with p 9 The overall correlation between RND and RNCCQ is calculated by combining RNCCQ data across ages 17, 27, and 37. The correlations for each respective
age bucket are r = 0.06 (SE = 0.10), r = 0.23 (SE = 0.09), and r = 0.15 (SE = 0.10). Readers interested in other subcomponents of RNCCQ (including the factor-
analytic subscales) can find the associated correlations with RND in the OA. 6. Concluding remarks We have reported pre-registered analysis of the latest survey wave of the Bing pre-school study on delay of gratification. Respondents were in their late 40’s at the time of this latest wave and were asked to report 11 mid-life capital formation
outcomes (e.g., net worth and permanent income). Our analysis plan both described our methods and predicted what we
would find. As predicted, a newly constructed measure derived from preschool delay of gratification does not predict the
11 capital formation variables (i.e., the sign-adjusted average correlation was 0.02). By contrast a composite self-regulation
index, combining preschool delay of gratification with survey measures of self-regulation collected at ages 17, 27, and 37,
does predict 10 of the 11 capital formation variables in the expected direction, with an average correlation of 0.19. The inclu-
sion of the preschool delay of gratification measure in this composite index does not affect the index’s predictive power for
two reasons. Most importantly, the index of self-regulatory measures is comprised of 86 responses per participant, whereas
the preschool delay of gratification task is a single behavioral task. In addition, the preschool delay of gratification task is
measured using a diagnostic variant of the task for 34 of our 113 participants; the remaining 79 participants experienced a
non-diagnostic variant of the pre-school delay of gratification task. The data we have analyzed is unique because the Bing cohort is the only sample where preschool delay of gratification
has been studied long enough to examine relationships with mid-life outcomes. While the tests of our primary hypotheses
were well powered, we caution that our sample is relatively small and not representative of the overall population—e.g., 97%
of our sample has a four-year college degree (the exceptions are one participant who has a two-year college degree and two
who have some college but no degree)—limiting the generalizability of the results. However, we can compare our results to the small set of overlapping analyses that have been conducted using the
Dunedin cohort, which began collecting data in 1972–1973 and has childhood self-regulation measures but no preschool
delay of gratification measure. 11 The cohort is from a small town in New Zealand with much lower levels of educational
attainment ( Moffitt et al., 2011 ). Specifically, 29% of the Dunedin sample is college educated ( Belsky et al., 2016 ). 11 Specifically, the measure of self-regulation used in Moffitt et al. (2011) is an index of examiner ratings of lack of control at ages 3 and 5; parent and
teacher ratings of impulse aggression and hyperactivity at ages 5, 7, 9, and 11; parent and teacher ratings of lack of persistence and impulsivity at ages 9
and 11; and self-reported symptoms of hyperactivity, inattention, and impulsivity at age 11. This measure does not include performance on the delay of
gratification task that we use to construct our RND measure. 5.4. Other secondary, robustness, and ex-post analyses 5.4. Other secondary, robustness, and ex-post analyses In OA, we report the results from all pre-registered secondary and robustness analyses as well as all ex-post analyses we
conducted. In our secondary analyses, we examine additional outcome variables, secondary sets of independent variables
(e.g., breaking out the RNSRI into its components), and comparisons of diagnostic and non-diagnostic conditions. In our ex-post analyses, we report correlations between our key variables, results using individual questions from the
RNCCQ as independent variables, an analysis of the characteristics of the Bing sample over time, an in-depth analysis of
BMI as an outcome variable, results using delay choice as an independent variable, robustness of the primary analyses
to controlling for age, results using an index of capital formation measures as the dependent variable, and an analysis of
the effects of rank-normalization. We also report results from two ex-post analyses resulting from the editorial process:
analyses of RNCCQ as an independent variable on its own, and a comparison of the BMI correlations in the diagnostic and
non-diagnostic subsamples using BMI data collected in previous surveys. In our robustness analyses, we test the robustness of our primary analyses to measuring wait time in seconds rather
than log-seconds, winsorizing RNSRI and RND, estimating a non-linear effect of RND, using alternative imputation methods
for the RNCCQ indices, using the delay deviation measure from prior work, and dropping survey respondents for whom we
had to make assumptions in order to assign values to any of their capital formation measures. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 12 paradigm ( Mischel and Ebbesen, 1970 ; see Peake, 2017 for discussion). Nonetheless, monetary delay choice measures (e.g.,
money earlier vs. later) have been popular among economists and psychologists as a tool for assessing time preferences and
hence, as a putative index of self-control and impulsive behavior (e.g. Thaler, 1981 ; see Stevens, 2017 for examples). We con-
sider (monetary) delay choice to be a dependent variable in our primary analysis, but in an ex-post analysis, we examined
it as a predictor of the remaining capital formation variables (as in Meier and Sprenger (2010), Meier and Sprenger (2013) ,
and Falk et al. (2018) ). Delay choice is a rank-normalized variable measuring the percent of times a respondent selected the later response in
a series of incentivized money earlier vs. later questions. The questions included 20 of the form “Which would you prefer? (A) getting $X today (B) getting $Y thirty days from today” and 20 of the form “Which would you prefer? (A) getting $X
thirty days from today (B) getting $Y sixty days from today,” with values ranging from $10 to $29. y
y
y ( ) g
g
y
y
y
g
g
We regressed the remaining 10 capital formation measures on delay choice, controlling for sex. Although all 10 correla-
tions have the predicted sign, none is significant at the 5% level. The average correlation with the capital formation measures
is 0.11 (SE = 0.06). See OA Section V-C-9 for more detailed information. 12
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[m3Gsc; November 16, 2019;10:34 ] JID: JEBO 5.3. Money now vs. money later as a predictor One of our capital formation variables, delay choice, is based on responses to money earlier vs. later questions. 10 Early
research on self-control in children relied heavily on delay choice data ( Mischel, 1958, 1961a, 1961b; Mischel and Gilli-
gan, 1964; Bandura and Mischel, 1965; Mischel and Staub, 1965; Mischel and Grusec, 1967 ). Recognition of the conceptual
limitations of delay choice measures was a primary motivation for the development of the preschool delay of gratification 8 Similarly, there is limited evidence for an age trend in the predictive power of the indices of CCQ questions. In ex-post analysis, we found that the
average correlations between the mid-life capital formation measures and RNCCQ at ages 17, 27, and 37 are 0.15 (SE = 0.09), 0.17 (SE = 0.06), and 0.19
(SE = 0.05), respectively (see OA). (
)
p
y (
)
9 The overall correlation between RND and RNCCQ is calculated by combining RNCCQ data across ages 17, 27, and 37. The correlations for each respective
age bucket are r = 0.06 (SE = 0.10), r = 0.23 (SE = 0.09), and r = 0.15 (SE = 0.10). Readers interested in other subcomponents of RNCCQ (including the factor-
analytic subscales) can find the associated correlations with RND in the OA. 10 For a review and methodological critique of money earlier vs. later questions, see Cohen et al (forthcoming) . For example, in principle choices in the
money earlier vs. later paradigm should reveal market interest rates, transactions costs, and trustworthiness of the experimenter, and should not measure
true time preferences (because money is temporally fungible). 10 For a review and methodological critique of money earlier vs. later questions, see Cohen et al (forthcoming) . For example, in principle choices in the
money earlier vs. later paradigm should reveal market interest rates, transactions costs, and trustworthiness of the experimenter, and should not measure
true time preferences (because money is temporally fungible). Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 The authors declare no conflicts of interest. The authors declare no conflicts of interest. Code availability Code for data analysis is available in the Online Appendix. D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx 13 to the self-regulation measures in the Bing sample. For example, Moffitt et al. (2011) found that a 1 SD increase in child-
hood self-control as measured in the Dunedin study predicts a 0.24 SD increase in income at age 32. In the Bing sample,
a 1 SD increase in RNSRI predicts a 0.32 SD increase in rank-normalized permanent income. Similarly, a 1 SD increase in
the Dunedin self-control measure predicts a 0.12 SD decrease in credit card problems, while a 1 SD increase in the Bing
RNSRI predicts a 0.18 SD decrease in rank-normalized credit card misuse. A 1 SD increase in the Dunedin self-control mea-
sure predicts a 0.14 SD decrease in money management difficulties, while a 1 SD increase in the Bing RNSRI predicts a 0.24
SD increase in financial health. 12 Despite these intriguing similarities across the two samples, the issue of generalizability
remains an important question to be addressed in future research as mid-life data becomes available in more childhood
longitudinal cohorts. to the self-regulation measures in the Bing sample. For example, Moffitt et al. (2011) found that a 1 SD increase in child-
hood self-control as measured in the Dunedin study predicts a 0.24 SD increase in income at age 32. In the Bing sample,
a 1 SD increase in RNSRI predicts a 0.32 SD increase in rank-normalized permanent income. Similarly, a 1 SD increase in
the Dunedin self-control measure predicts a 0.12 SD decrease in credit card problems, while a 1 SD increase in the Bing
RNSRI predicts a 0.18 SD decrease in rank-normalized credit card misuse. A 1 SD increase in the Dunedin self-control mea-
sure predicts a 0.14 SD decrease in money management difficulties, while a 1 SD increase in the Bing RNSRI predicts a 0.24
SD increase in financial health. 12 Despite these intriguing similarities across the two samples, the issue of generalizability
remains an important question to be addressed in future research as mid-life data becomes available in more childhood
longitudinal cohorts. Supplementary materials Supplementary material associated with this article can be found, in the online version, at doi: 10.1016/j.jebo.2019.08.016 . Data availability Data collection for the Bing Longitudinal Study now spans nearly 50 years. Due to a host of historical issues connected to
the consenting procedures in this work and promises made to participants to protect both their privacy and the confiden-
tiality of their responses, it has been a long-standing policy and IRB requirement that we not share the data connected with
this work with outside parties or agencies. We believe it is imperative that we honor the promises made to our participants
who have generously provided input to the project over the years. References b ll
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teacher ratings of impulse aggression and hyperactivity at ages 5, 7, 9, and 11; parent and teacher ratings of lack of persistence and impulsivity at ages 9
and 11; and self-reported symptoms of hyperactivity, inattention, and impulsivity at age 11. This measure does not include performance on the delay of
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school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 D.J. Benjamin, D. Laibson and W. Mischel et al. / Journal of Economic Behavior and Organization xxx (xxxx) xxx
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Mischel, W. , Ebbesen, E.B. , 1970. Attention in delay of gratification. J. Pers. Soc. Psychol. 16, 329–337 . 12 Other outcomes studied in Moffitt et al. (2011) at age 32 include measures of physical health, depression, substance dependence, socioeconomic status,
single-parent child rearing, financial planfulness, and criminal conviction. 12 Other outcomes studied in Moffitt et al. (2011) at age 32 include measures of physical health, depression, substance dependence, socioeconomic status,
single-parent child rearing, financial planfulness, and criminal conviction. Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 ARTICLE IN PRESS Ability to delay gratification at age 4
years and risk of overweight at age 11 years. Arch. Pediatr. Adolesc. Med. 163, 303–308 . Shoda, Y. , Mischel, W. , Peake, P.K. , 1990. Predicting adolescent cognitive and self-regulatory competencies from preschool delay of gratification: identifying
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age 9. Acta Psycholog. Sin. 39, 668–678 . Please cite this article as: D.J. Benjamin, D. Laibson and W. Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016 Please cite this article as: D.J. Benjamin, D. Laibson and W. y
ng, J. , 2007. A follow-up study of self-imposed delay of gratification at age r as a predictor of children’s school-based socia
ta Psycholog. Sin. 39, 668–678 . ARTICLE IN PRESS Mischel et al., Predicting mid-life capital formation with pre-
school delay of gratification and life-course measures of self-regulation ✰ , Journal of Economic Behavior and Organization,
https://doi.org/10.1016/j.jebo.2019.08.016
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IDEALISMO À ITALIANA: FILOSOFIA NA ITÁLIA E INFLUÊNCIA GERMÂNICA NOS SÉCULOS XIX E XX: O CASO DA INFLUÊNCIA E RECEPÇÃO DE FEUERBACH, KIERKEGAARD E NIETZSCHE NAS FILOSOFIAS DE BENEDETTO CROCE E GIOVANNI GENTILE
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IDEALISMO À ITALIANA: FILOSOFIA NA ITÁLIA E
INFLUÊNCIA GERMÂNICA NOS SÉCULOS XIX E XX:
O CASO DA INFLUÊNCIA E RECEPÇÃO DE FEUERBACH,
KIERKEGAARD E NIETZSCHE NAS FILOSOFIAS
DE BENEDETTO CROCE E GIOVANNI GENTILE Márcio Gimenes
Universidade de Brasília Resumo: O objetivo do presente trabalho é investigar o panorama da filosofia italiana dos séculos XIX e
XX e a respectiva influência germânica por ela sofrida, avaliando ainda a articulação do idealismo
italiano e a crítica ao cristianismo produzida na Itália nesse período. Para tanto, dividiremos nosso artigo
em duas partes principais: na primeira, investigaremos o idealismo italiano e a crítica à religião na Itália
entre os séculos XIX e XX, dando especial ênfase a dois dos seus principais autores: Benedetto Croce e
Giovanni Gentile. Já na segunda parte, investigaremos a influência de três pensadores do idealismo
germânico na Itália desse período, a saber, Feuerbach, Kierkegaard e Nietzsche, bem com a sua
recepção em contexto italiano e sua importância para o idealismo local. Palavras-chave: Filosofia italiana; filosofia contemporânea; idealismo; cristianismo. Abstract: The objective of this study is to investigate the panorama of Italian philosophy of the
nineteenth and twentieth centuries and its Germanic influence for her suffering, still evaluating the joint
Italian idealism and criticism of Christianity produced in Italy during that period. In this way, we will divide
our article into two main parts: first, we will investigate Italian idealism and criticism of religion in Italy
between the nineteenth and twentieth centuries, with special emphasis on two of its main authors:
Benedetto Croce and Giovanni Gentile. Already in the second part, we will investigate the influence of
three thinkers of Germanic idealism in Italy of that period, namely Feuerbach, Kierkegaard and
Nietzsche, as well as their reception in the Italian context and their importance for local idealism. Keywords: Italian Philosophy; Contemporary Philosophy; Idealism; Christianity. Introdução A despeito de o Brasil possuir uma imensa afinidade cultural com a
Itália, não apenas por partilhar a herança linguística latina, mas também por
haver recebido, especialmente a partir do final do século XIX e início do
século XX, inúmeros imigrantes italianos, a filosofia italiana, talvez excluindo-
se aqui autores clássicos como Maquiavel, Galileu, Vico e alguns outros, ainda
é, infelizmente, pouco conhecida e estudada no Brasil e, segundo avaliamos, o
mesmo ocorre em Portugal e em em outros países de língua portuguesa. Obteve maior êxito entre os brasileiros – e cremos que entre os falantes de
português em geral – o estudo dos autores franceses, alemães e ingleses, o que © Dissertatio [45] 184-2082017 Márcio Gimenes avaliamos como importante, mas não excludente. Afinal, a própria filosofia
italiana dos séculos XIX e XX não é feita, a rigor, como filosofia italiana pura,
isto é, como um produto ligado a um modo de pensar ou a uma posição
geográfica, tal como aponta Roberto Esposito em sua obra O pensamento vivo –
a origem e a atualidade da filosofia italiana (ESPOSITO, 2013). A filosofia, por
definição, teria imensa dificuldade em ser tomada como pertencente a esse ou
aquele país, uma vez que ela é, por princípio, universal. Desse modo, dissertar
sobre filosofia italiana, francesa, alemã, inglesa, brasileira, latino-americana,
europeia ou africana sempre comportaria, em si mesmo, um problema e uma
impossibilidade. Contudo, pensamos que aqui é que se insere efetivamente o
interesse do nosso trabalho, isto é, em saber como ocorre aquilo que se
convencionou denominar como filosofia italiana dos séculos XIX e XX. Para
tanto, daremos especial destaque ao idealismo e à crítica da religião na Itália e a
analisaremos a partir do pensamento de autores como Benedetto Croce,
Giovanni Gentile, Antonio Gramsci, Maurizio Viroli, Carlo Augusto Viano,
Eugenio Garin e Franco Lombardi, dentre outros. Assim almejamos
compreender, ao menos em linhas gerais, um pouco do panorama da filosofia
italiana do período e do contexto religioso italiano da época. Julgamos tal
passo, inclusive, importante para uma adequada compreensão da maneira
como os italianos interpretaram a crítica religiosa do cristianismo operada
pelos autores germânicos do século XIX como Feuerbach e Nietzsche (que
eram alemães) e bem como Kierkegaard (que era dinamarquês), uma vez que
tais autores foram centrais para o idealismo italiano e influenciaram
diretamente na sua crítica ao cristianismo e na articulação intelectual do seu
tipo de idealismo. Introdução Assim, o objetivo desse trabalho consiste na avaliação do problema
religioso italiano no século XIX e na sua problematização, ou seja, como,
também a partir dele, se pode compreender o idealismo italiano desse século e
do século seguinte. Com efeito, nosso intuito é avaliar o hegelianismo e o pós-
hegelianismo na Itália, para tanto nos debruçaremos especialmente sobre
algumas reflexões delimitadas dos autores já citados. Avaliamos que aqui, com
as delimitações adequadas, conseguiremos apresentar um quadro significativo
de diferentes matizes do hegelianismo na cultura italiana dos séculos XIX e
XX. Feito tal percurso, apresentaremos nossa conclusão crítica que, segundo
avaliamos, pode apresentar importantes desafios para uma compreensão mais
ampla da história da filosofia e para uma valorização do pensamento italiano. 185 Dissertatio [45] 184-2082017 1. O contexto do idealismo italiano e da crítica da religião na Itália entre
os séculos XIX e XX: com especial atenção para Benedetto Croce e
Giovanni Gentile 1. O contexto do idealismo italiano e da crítica da religião na Itália entre
os séculos XIX e XX: com especial atenção para Benedetto Croce e
Giovanni Gentile 1. O contexto do idealismo italiano e da crítica da religião na Itália entre
os séculos XIX e XX: com especial atenção para Benedetto Croce e
Giovanni Gentile Carlos Augusto Viano, em sua obra La filosofia italiana del Novecento
(VIANO, 2006), apresenta, a partir do debate do idealismo italiano,
notadamente a partir da polêmica das teses de Benedetto Croce e Giovanni
Gentile, um inspirador retrato da filosofia italiana no início do século XX. Tal
retrato é feito em forma panorâmica e, segundo avaliamos, apesar dos seus
limites, pode ser importante para uma primeira compreensão de aspectos
significativos da filosofia italiana do século XX. Sua apresentação, de uma
forma geral, nos fornece a interpretação crociana de Hegel, Marx e também
um pouco de aspectos da posição liberal do pensador napolitano. De igual
modo, avalia e apresenta os principais movimentos de renovação da cultura
italiana do período dando, especial ênfase, às revistas e outras produções. O
autor igualmente investiga a polêmica intelectual e pessoal entre Croce e
Gentile, o fascismo, a religião ensinada para a massa na escola pública, bem
como a tese do Estado laico como um ideal tanto do fascismo quanto da
escola pública. Afirmo, portanto, a minha profissão de fé agrade ou não agrade a quem está a
me ouvir: eu sou cristão. Sou cristão porque creio na religião do espírito, mas
quero rapidamente acrescentar a fim de evitar equívocos: eu sou católico. E não
de hoje, fique isso bem claro. Católico, a rigor, sou desde junho de 1875, a saber, Introdução Por isso, não fortuitamente, o tema da laicidade, tão caro à filosofia
moderna, foi um pilar central da reforma que Gentile operou enquanto
ministro da instrução pública e acabou, na verdade, por se constituir no seu
calcanhar de Aquiles nas suas polêmicas contra a Igreja. Desde o primeiro
momento, Gentile, como ministro e como educador, teve uma difícil relação
com o pensamento católico. A tese por ele defendida do Estado laico e da
religião como educação básica para as massas incultas nunca pareceu muito
aceitável aos olhos das instituições religiosas. Sua posição apontava para a
Igreja como uma espécie de iniciadora dos espíritos na cultura, mas, ao mesmo
tempo, afirmava, ainda indiretamente, que ela, de fato, não fazia ciência. Como
era de se esperar, notadamente num contexto como o italiano, os católicos
reagem e não aceitam nem as teses de Croce (da religião da liberdade), nem a
posição de Gentile, nem o humanismo modernista. Contudo, também recusam
o antigo tomismo. Assim é que se delinea a tumultuada relação da Igreja com o
fascismo. A crise da Igreja Católica em perder o seu lugar de posto cultural,
especialmente na Itália tem uma forte repercussão. É verdade que Gentile se
proclamava católico e advogava a tese de uma religião estatal e racional, tal
como se pode notar na sua declaração La mia religione: 186 Márcio Gimenes desde quando estou no mundo. E sou, por isso, desolado de não poder-vos
anunciar também eu, uma crise, uma tempestade da alma, uma súbita conversão,
um golpe fulminante. Estou, prosaicamente, desde o dia do meu nascimento,
percorrendo a estrada de Damasco (GENTILE, 1965, p.406). desde quando estou no mundo. E sou, por isso, desolado de não poder-vos
anunciar também eu, uma crise, uma tempestade da alma, uma súbita conversão,
um golpe fulminante. Estou, prosaicamente, desde o dia do meu nascimento,
percorrendo a estrada de Damasco (GENTILE, 1965, p.406). Contudo, tal posição não atenua os fatos de que, aos olhos de muitos
católicos – e da estrutura eclesiástica – o máximo de sua religião alcançava um
mero tipo de religião da razão ao estilo hegeliano. O mesmo ocorre com Croce
ao advogar a tese da religião da liberdade. g
g
Na filosofia italiana do século XX, a politica sempre foi uma tônica e
um ponto central. As discussões acaloradas entre liberais e socialistas foram
marcantes em todo o período. Introdução Croce era, inclusive,
sobrinho de Bertrando e de Silvio Spaventa, introdutores do pensamento
hegeliano na Itália. Gentile foi um grande admirador e aluno de Bertrando. Ambos partem, portanto, do idealismo filosófico hegeliano. Aliás, como bem
salienta Garin, o hegelianismo italiano deve ser compreendido num
“quadrado” que inclui Spaventa, De Sanctis, Croce e Gentile. Com efeito,
ambos renovam a vida cultural italiana, ampliam os debates, produzem
inúmeras revistas, livros e retiram a Itália do seu gueto intelectual, ampliando seu
escopo de discussão com autores alemães e também ingleses e franceses. p
g
O mesmo Garin (GARIN, 1966, p.1225-1243) aponta que os irmãos
Spaventa e a chamada escola de Napóles foram extremamente importantes
para a inserção do pensamento hegeliano na Itália, salientando ainda que
Firenze e Palermo foram dois importantes centros da filosofia italiana no
período. Os antecedentes de tal pensamento podem ser percebidos em dois
pensadores: Domenico Mazzoni e Giambattista Passerini. Ambos foram
sacerdotes católicos e os primeiros italianos a estudarem, ainda que de modo
mais rudimentar, o pensamento de Hegel. Por isso, na concepção de Garin, o
primeiro hegeliano com estudo um pouco mais aprofundado foi, na verdade,
Augusto Vera. Tal autor escreveu, em Paris, no ano de 1855, uma Introduction à
la philosophie de Hegel. Sua obra foi basicamente construída dentro do ambiente
acadêmico francês e foi a primeira a apresentar uma certa robustez filosófica. O mesmo Garin (GARIN, 1966, p.1225-1243) aponta que os irmãos
Spaventa e a chamada escola de Napóles foram extremamente importantes
para a inserção do pensamento hegeliano na Itália, salientando ainda que
Firenze e Palermo foram dois importantes centros da filosofia italiana no
período. Os antecedentes de tal pensamento podem ser percebidos em dois
pensadores: Domenico Mazzoni e Giambattista Passerini. Ambos foram
sacerdotes católicos e os primeiros italianos a estudarem, ainda que de modo
mais rudimentar, o pensamento de Hegel. Por isso, na concepção de Garin, o
primeiro hegeliano com estudo um pouco mais aprofundado foi, na verdade,
Augusto Vera. Tal autor escreveu, em Paris, no ano de 1855, uma Introduction à
la philosophie de Hegel. Sua obra foi basicamente construída dentro do ambiente
acadêmico francês e foi a primeira a apresentar uma certa robustez filosófica. Contudo, para Spaventa, Vera seria um hegeliano mais hegeliano do que o
próprio Hegel. Introdução Nesse contexto também se pode compreender
o tema da renovação moral nos debates entre Croce e Gentile. Muito se ouve
falar, no contexto posterior a Segunda Guerra, da tese fascista de Gentile, da
sua atuação como ministro, como político, de sua enorme produção intelectual
e do desfecho trágico de sua vida e do seu assasinato em 1944 pelas forças
opositoras ao regime. Contudo, avaliamos que, primeiro, cabe entender que a
sua posição e a sua atuação política tem uma estreita relação com a sua posição
filosófica. Em outras palavras, sua interpretação, como filósofo, do estado
ético e moral hegeliano, isto é, o seu hegelianismo é quem efetivamente o
coloca no centro dos acontecimentos políticos do século XX. Assim, a imagem
do filosófo que é legada à posteridade, ao menos nos primeiros anos da sua
morte, isto é, aquela imagem do filósofo aliado ao fascismo é verdadeira, mas
não faz total justiça ao pensador, visto que, sem entrarmos no âmbito do
julgamento moral ou político, Gentile foi um intelectual de fôlego, com uma
imensa produção intelectual e um dos primeiros ministros a construir um
plano de instrução pública para toda a Itália, além de levar adiante o
empreendimento da Enciclopédia italiana, que era um desdobramento, a notar
pelo seu próprio nome, da sua posição idealista em filosofia, tal como é
igualmente perceptível nesse pequeno trecho da declaração La mia religione: “A
religião cresce, se espande, se consolida e vive, dentro da filosofia, que elabora
incessantemente o conteúdo imediato da religião e o introduz na vida da
história” (GENTILE, 1965, p.424). Em outras palavras, a religião para Gentile
está inserida dentro do todo do sistema hegeliano. g
De igual modo, o filósofo italiano talvez mais conhecido no contexto
após a Segunda Guerra é Benedetto Croce. O filósofo napolitano, antigo
amigo de Gentile, com quem rompe após discordâncias políticas, é uma figura
de destaque por sua forte oposição ao fascismo e por sua clara posição
contrária aos intelectuais que o defenderam. Tal posição lhe confere destaque
inclusive no exterior, tendo também o mérito de levar a filosofia italiana além
dos seus confins. Entretanto, tal como muito bem relembra Garin, citando 187 Dissertatio [45] 184-2082017 Gramsci: “Croce não pode ser compreendido sem Gentile” (GRAMSCI apud
GARIN, 1997, p.172, n.2). Assim, Gentile e Croce possuem alguns traços
comuns: ambos são herdeiros da tradição hegeliana. Introdução Desse modo, ao analisar criticamente sua obra, aponta para a
falta de um filosofar italiano e também percebe nas suas reflexões uma falta da
compreensão de Hegel dentro de um contexto de maior liberdade e
secularização. O tema da falta de liberdade para a reflexão filosófica é um
aspecto central a ser percebido aqui. Para Spaventa, o fato de a Itália ser a sede
do poder católico já a impederia, por natureza, a tal reflexão. Desse modo, sua
posição aponta, com extrema argúcia, para uma direção que pode ser
explorada conceitualmente: a filosofia italiana clássica foi, no seu entender,
emigrada para autores não italianos como Espinosa, Leibniz e Kant. Esses
seriam italianos sem serem italianos por nascimento. Em outras palavras,
trariam em sua filosofia todo o tema clássico da filosofia italiana da liberdade
de reflexão. Desse modo, Espinosa teria um equivalente com Giordano Bruno
e Kant com Vico. Tal tese, que já aparece em Rodolfo Mondolfo
(MONDOLFO, 1924, p.2ss), ressoa mesmo em autores como Gentile e
Croce. O primeiro, por exemplo, escreve um importante trabalho sobre a
filosofia de Giordano Bruno, o que não parece desprovido de significado 188 Márcio Gimenes (GENTILE, 1920). O segundo desenvolve, na obra Storia d´Europa nel secolo
decimonono, o importante conceito de religião da liberdade. Com efeito, o
hegelianismo italiano possui um arco que começa por Vera, os irmãos
Spaventa e alcança autores como Gentile e Croce, incluindo ainda, nesse
horizonte conceitual, Gramsci. Garin destaca ainda aqui a importância de
autores como Antônio Tari, que, juntamente, com Spaventa e Vera teria
formado a “santíssima trindade hegeliana na Itália”, incluindo ainda os nomes
de Labriola e Francesco de Sanctis. Não se poderia fazer justiça à filosofia italiana do século XX sem
mencionarmos o surgimento da Crítica em 1903, o famoso periódico dirigido
por Croce. Nessa época, o pensador napolitano afirma-se como um autor mais
maduro. Começam a se delinear aqui os seus interesses mais claros que, em
geral, acompanham aqueles que abandonaram a religião, ou, ao menos, não
dão mais a ela uma importância central. Em seu lugar, surgem a arte, a moral, o
direito, a metodologia histórica. Introdução Em outras palavras, Croce configura-se aqui
como um hegeliano típico, como aponta Garin: “Negada a filosofia da história
como reencarnação da teologia, a filosofia já se fazia para Croce, compreensão
da aparência da história humana ou, se se quiser, do ritmo profundo da
humanidade no seu processo” (GARIN, 1997, p.184). Junto com Garin
podemos observar, inclusive, diversos Croces e Gentiles, autores com diversas
fases. Por influência de Labriola, ambos se tornam leitores de Hegel e também
de Marx. Croce passa de tal fase para a Estética, realizando, mais ou menos,
um sistema ideal no decorrer de sua obra. No fundo, sua filosofia fica na
fronteira e cumpre o drama da consciência italiana: “[...] o antigo drama da
consciência italiana próxima do interesse mundano, da alegria da vida, do amor
das formas e dos arroubos místicos de uma outra cidade” (GARIN, 1997,
p.266). Assim, avaliamos que notadamente para compreendermos melhor o
tema da religião da liberdade em Croce, cumpre fazer uma avaliação daquilo
que Galasso (GALASSO, 2002) denomina com a sua fase histórica. q
(
)
Um dos marcos do pensamento italiano do século XX foi a
publicação, em 1932, da já citada Storia d´Europa nel secolo decimono de Benedetto
Croce. Ali o filósofo napolitano desenvolve um tema que já existe em outros
autores que o antecedem, a saber, o tema da religião da liberdade. Contudo,
sua abordagem é diferenciada. Na sua concepção, a era posterior a Napoleão
possui, em todo o mundo, um anseio pela liberdade. Essa é a nova palavra de
ordem. Segundo Croce, tal aspiração é oriunda de nossa herança greco-
romana, tendo um acréscimo, depois da Revolução Francesa do século XVIII,
dos temas da igualdade e da liberdade. Na verdade, a liberdade pode ser dita de
muitos modos e possui, sem dúvida alguma, muitas facetas: liberdade civil,
liberdade política, liberdade do indivíduo singular, liberdade dos indivíduos 189 Dissertatio [45] 184-2082017 coletivos. Desse modo, Croce promove nessa obra uma espécie de inventário
da liberdade, discorrendo sobre ela nos gregos, nos romanos, nos filósofos, na
Renascença, na Reforma Protestante, na Revolução Francesa e etc. Sua tese é,
no fundo, de matriz hegeliana, a saber, a história deve ser vista como a história
da liberdade. A partir dessa visão apresenta-se também a sua perspectiva acerca
da moral. E, ao abordar o tema da moral, surgem outras indagações: o que é a
liberdade? Para que ela serve? A quem ela pertence? Introdução Um dos erros mais grosseiros do catolicismo foi a defesa da
monarquia absolutista e a manutenção de uma ideia limitada sobre o poder do 190 Márcio Gimenes governante que, aliás, ainda se pode perceber também em Lutero. A tese
crociana é que, na verdade, o pensamento moderno se caracteriza por uma
passagem do naturalismo e do racionalismo para uma progressiva caminhada
em direção à dialética e ao idealismo. Por isso, segundo o pensador de
Nápoles, o liberalismo tem forte relação com o espiritualismo e o comunismo
com o materialismo mais estrito. Tal concepção forneceria ao liberalismo um
pouco mais de sofisticação intelectual. Nesse contexto, pode-se entender também um importante fator
constituinte dos séculos XIX e XX, isto é, a presença do romantismo que,
segundo Croce, pode ser divido em dois tipos: o primeiro seria teórico e
especulativo; o segundo seria prático e, por isso, mais sentimental e moral. O
fato concreto é que no romantismo encontra-se uma reação ao academicismo
e nele se tem, por exemplo, a geração da Estética e de suas preocupações
filosóficas. Por sua vez, o romantismo de matriz religiosa e mais moral, além
de ser protestante é profundamente prático. Tal romantismo sonha com um
mundo capaz de conciliar a natureza e o espírito, ao contrário da estética, que
privilegiará a fantasia e o sonho e também um espírito de cunho mais
especulativo, que, de algum modo, sempre percorrerá uma trilha atrás de uma
síntese. A partir de tal diversidade de espécies de romantismo temos, no
entender de Croce, até mesmo um neopaganismo, visto que ele vai em busca,
inclusive, das suas origens primitivas germânicas. Entretanto, também existe
nesse romantismo um lado ético e político, que talvez possa ser melhor
explorado. Seu erro básico foi, na verdade, dar uma forte ênfase no mundo da
vida prática, sem compreender a importância da teoria e da especulação. Assim, a religião da liberdade crociana ultrapassaria o romantismo mas, ao
mesmo tempo, se nutriria de suas principais ideais e as reelaboraria,
desenvolvendo uma guerra do espírito, uma dialética diferenciada, tal como se
poderá notar na resistência e na oposição ao predomínio absolutista
promovido pelo liberalismo. A tese central de Croce é que ocorre uma vitória
do ideal liberal em todo o mundo. O absolutismo vai pouco a pouco perdendo
seu terreno. Introdução Croce assiste, no seu tempo, os diversos nacionalismos se
apropriarem do conceito de liberdade e tentarem circunscrevê-lo, aos seus
domínios territoriais, concretizando-o a partir do solo onde se vive. Entretanto, a liberdade é também uma ruptura com vínculos tradicionais,
podendo ainda ser vista como liberdade econômica. O pensador napolitano vê,
na figura do intelectual do século XX, a afirmação da liberdade por excelência. No seu entender, o intelectual seria um tipo de novo missionário em uma nova
cruzada, a saber, a cruzada da liberdade. Desse modo, o pensador avalia que a
liberdade é uma religião. Ela possui uma ética própria e aponta para um tipo de
religião do futuro. Figuras como Sócrates e Jesus e eventos como o paganismo,
o cristianismo, o agostinianismo, o calvinismo mostrariam claramente tal tese. A era da liberdade seria, no fundo, a era do espírito preconizada por Joaquim
de Fiore no século II. O problema para Croce é que tal religião da liberdade
não existe na Igreja Católica. No seu entender, o catolicismo é a negação da
ideia liberal. Seu grande intuito é a pregação de uma religião ultramundana,
ainda que, em alguns momentos, defenda ou propicie o aparecimento de
defensores da religião da liberdade: “a história, que é a história da liberdade, se
mostra mais forte do que a sua doutrina e seu programa, e o derrota e o força a
se contradizer no campo dos fatos” (CROCE, 1965, p.23). Nesse mesmo
escopo é que se pode compreender os desafios do Renascimento e da Reforma
Protestante diante de uma Igreja Católica corroída. No entender do pensador,
a Contra-Reforma salvou o corpo político da Igreja, mas não foi capaz de
salvar a sua alma. Assim, seu domínio ficou restrito ao mundano e perdeu a
sua vocação religiosa. Em decorrência de tal situação, a Igreja Católica assiste a fuga dos
seus pensadores, bem como dos seus cientistas, restando nela basicamente o
construto da Escolástica. Em outras palavras, ela não alcança a modernidade:
não chega a autores como Lutero, Calvino, Rousseau e Voltaire. Tal instituição
se configura como inimiga do liberalismo. Contudo, Croce é muito claro ao
afirmar não um anti-cristianismo, mas um anti-clericalismo: “[...] a aversão era
não ao catolicismo, mas ao ‘clericalismo’, ao ‘negro clericalismo’” (CROCE,
1965, p.27). Introdução Os ideais liberais de desenvolvimento, progresso, educação,
civilidade, ganham corpo e se ampliam em escala mundial, ainda que isso
ocorra de modo diverso, a depender do contexto. Para além do domínio
político e econômico, afirma-se aqui uma tese bastante instigante: os séculos
XIX e XX revelam uma vida intelectual bastante ativa. Aqui se pode
claramente perceber a ênfase cultural dos ideais liberais e a reação dos ideais
católicos. A curiosa posição de Croce é que na Itália (e mesmo na França) a
renovação do pensamento se dá por meio dos jovens pensadores alemães:
Kant, Hegel e autores pós-hegelianos. No seu entender, acontece na Igreja
Católica uma queda cultural e, diante disso, o que resta a ela é o exercício do 191 Dissertatio [45] 184-2082017 poder político: “[...] a Igreja Católica não podia seguir os seus adversários na
alta esfera em que se moviam e, mais e mais, se reduzia a potência
prevalentemente política” (CROCE, 1965, p.84). Assim, os padres são cada
vez mais vistos na companhia dos reis e dos seus ministros. Por isso, Croce
percebe em tudo isso uma mistura dos interesses da sacristia e dos ideais
policiais. O fato é que, no seu entender, o liberalismo consegue colher nessa
época, inclusive na Igreja, os seus melhores pensadores e por isso, não
despropositadamente, muitos dos pensadores dos séculos XIX e XX são
oriundos da estrutura eclesiástica. A Revolução Francesa foi, sem dúvida alguma, um ponto central para
a posição liberal desde o século XVIII. Croce aponta que seus ecos e
repercussão podem ser percebidos na Itália em autores como Giuseppe
Mazzini. Afirmam-se aqui dois curiosos tipos de catolicismo italianos: o
primeiro é o tipo do católico liberal; o segundo o do anti-clerical. Ambos
acontecem num período marcado por debates intensos no campo social e
entre as teses de socialistas, comunistas e liberais. Percebe-se aqui, segundo
Croce, a importância decisiva dos pensadores pós-hegelianos pois esses,
nutridos também por autores franceses e pelo vento da Revolução Francesa
promovem, em seu campo, uma reviravolta no idealismo alemão. Tal
configuração torna-se decisiva para a nova compreensão do idealismo não
apenas na Alemanha, mas no restante do mundo. Além disso, Croce destaca a
contribuição de autores como Tocqueville, autor da célebre análise da
Democracia na América, que, no seu entender, afirma-se como um profeta dos
novos tempos do século XIX e da sua passagem para o século XX. Ainda naqueles primeiros meses um jovem filósofo de inspiração hegeliana,
Silvio Spaventa, tematizava pensar o impensável e dizia que o infinito abstrato
da Igreja e aquele vivo na nacionalidade e no Estado, o infinito da religião e o
infinito da sociedade, por obra e fato de “um homem tido por infalível”, eram
reconhecidos na sua unidade, na unidade de Deus, “que reina no intelecto e no
coração como no próprio céu se nutre da sua luz” (CROCE, 1965, p.151). Introdução Os ideais liberais tem diante de si, especialmente na Itália, um inimigo
decisivo: o papado. Como Croce bem aponta, seguindo a trilha de Silvio
Spaventa, seu tio, tal guerra existe desde os dias de Maquiavel e persiste até o
século XX: Ainda naqueles primeiros meses um jovem filósofo de inspiração hegeliana,
Silvio Spaventa, tematizava pensar o impensável e dizia que o infinito abstrato
da Igreja e aquele vivo na nacionalidade e no Estado, o infinito da religião e o
infinito da sociedade, por obra e fato de “um homem tido por infalível”, eram
reconhecidos na sua unidade, na unidade de Deus, “que reina no intelecto e no
coração como no próprio céu se nutre da sua luz” (CROCE, 1965, p.151). Assim, a Igreja parece não conviver bem com os ideais liberais. Quando ela se encontra fragilizada, clama por maior liberdade. Contudo,
quando ela é forte e poderosa, tolhe a liberdade, mostrando efetivamente o seu
intuito. Tal será sempre a tônica da difícil relação entre o liberalismo e a Igreja,
que piora substancialmente quando há um problema efetivo a se responder, a
saber: o que fazer com o poder temporal do papa? Ou melhor, o que fazer 192 Márcio Gimenes quando tal poder, por força do avanço dos ideais liberais, acaba. Como
justificá-lo? A resposta à pergunta não é simples, mas Croce ensaia uma
possível: [...] porque a Igreja, sociedade perfeita, inclui com o espiritual o temporal, e a
seu tempo estendia bastante largo o seu poder, e investia e coroava os príncipes
da terra e os excomungava e os depunha, e se ora se via reduzida a dominar
somente um pedaço da Itália, nem por isso tinha renunciado a um direito ao
qual não lhe era dado renunciar sem contradizer a própria doutrina e natureza
(CROCE, 1965, p.199). O fim do poder temporal eclesiástico não deixa de ser o fim de um
dado tipo de teocratismo e por isso repercute de forma positiva na Itália e em
toda a Europa. Tal posição ajuda, inclusive, na redescoberta de valores cristãos
mais sinceros e honestos deixando mais claro, por exemplo, um debate entre
as teses da Igreja triunfante (aliada ao poder) e a Igreja militante (a Igreja dos
mártires, dos que sofrem). Por isso, de forma muito curiosa, mas não sem
propósito, Croce nas palavras finais do seu livro usa, uma vez mais, o termo
“religião da liberdade” (CROCE, 1965, p.316). Introdução Em outras palavras, completa o
seu inventário da liberdade. Segundo Galasso, o período compreendido após 1870 é uma época
fundamental da história europeia. Por isso, não fortuitamente, duas das obras
crocianas abordam claramente tal período, a saber, Storia d´Europa nel secolo
decimono e Storia d´Italia dal 1871 al 1915. Aqui, no horizonte posterior à guerra
franco-prussiana afirmam-se a discussão de temas como o nacionalismo, o
socialismo, o republicanismo, os valores liberais, o positivismo, bem como o
hegelianismo e a sua herança. Assim, o próprio título da obra de Galasso (Croce
e lo spirito del suo tempo) parece claramente demonstrar as intenções do autor. Por isso, no seu entender, Croce seria, na verdade, um autor do diagnóstico da
crise do final do século XIX e daquilo que se sucederia no século XX. q
q
Com efeito, a obra do autor napolitano, ao seu modo, não deixa de
espelhar um debate entre o romantismo e a ciência, notadamente o debate
entre o positivismo e a ciência. Assim, há também aqui uma proximidade entre
os temas crocianos e os temas desenvolvidos por Comte e por Hegel. Por isso,
há também na obra do pensador, apesar dele nunca ter sido um positivista,
fortes traços do laicismo e do anticlericalismo típicos do positivismo. Curiosamente, tal positivismo mostra, uma vez mais, que a cultura italiana é
ora europeia e ora parece distante dos temas filosóficos da Europa, a depender
do contexto e da perspectiva avaliada. No entender de Galasso, o positivismo
foi, na Itália, muito mais um clima cultural do que uma filosofia: “O
positivismo foi, com efeito, um clima cultural e moral, antes e mais do que
uma filosofia” (GALASSO, 2002, p.93). 193 Dissertatio [45] 184-2082017 Do mesmo modo que o positivismo possui uma recepção particular
na Itália, o mesmo ocorre com o marxismo. Aqui, seguindo a esteira dos
introdutores de Hegel na Itália como Spaventa e Labriola, Croce e Gentile, por
exemplo, aproximam-se dos temas da filosofia marxista. Aqui compreendemos
o que se poderá chamar como idealismo italiano. Nesse contexto, tanto Croce,
como também Gentile, se inserem como filósofos interessados nos temas da
filosofia hegeliana e como críticos do positivismo. Assim, o Croce desse
período é um autor hegeliano, estudioso da filosofia do espírito, alguém que
busca recuperar os temas da filosofia da história e, nesse sentido, um crítico do
positivismo. Introdução Aliás, não há distinção entre eles. Por isso, sua
pista de liberdade pode nos livrar do autoritarismo tanto da religião como dos
totalitários políticos de esquerda ou de direita e, nesse sentido, ele é inspirador
mesmo para seus opositores como Gramsci (1949a, 1949b). Além disso, possui
o mérito de inserir a filosofia italiana em um debate filosófico internacional na
medida em que suas teses foram discutidas por importantes autores como
Löwith e Cassirer, por exemplo. Entretanto, como bem parece salientar Viano,
o idealismo crociano (e gentiliano) sempre conviveram também numa certa
dubiedade: ao mesmo tempo em que pareciam integrar a filosofia italiana ao
mundo, pareciam também fechá-la ao mundo, não sendo tal fechamento
unicamente culpa do fascismo: Do ponto de vista filosófico-cultural em geral o predomínio da filosofia crociana,
juntamente com o domínio daquela gentiliana, criou um clima de prevalência
idealística: esse tinha garantido à Itália a uniformidade cultural, mas a manteve
alheia aos movimentos que da Primeira Guerra Mundial até metade do século
tinham caracterizado a cultura dos outros grandes países europeus e dos Estados
Unidos. A segunda parte do problema de fronte ao qual se encontrava a filosofia
italiana depois da guerra se configurava assim como a abolição do fechamento cultural
italiano, um fechamento que o fascismo tinha reforçado e institucionalizado com
a própria política cultural, mas que já antes o neoidealismo havia praticado
(VIANO, 1985, p.49). Assim, abre-se aqui também um importante espaço para que se possa
pensar a filosofia italiana do século XX com Croce (e Gentile), mas também
contra ambos, o que sempre parece mais rico e significativo. Tal exercício é
efetivado, segundo uma perspectiva gramsciana, por Piccioni (PICCIONI,
1983) que, na sua obra Ideologia e filosofia del neoidealismo italiano, aponta
claramente que tanto Croce como Gentile são autores pós-positivistas e, ao
mesmo tempo, autores conservadores e anti-marxistas. Como vimos, para
Gentile – mas também para Croce – a religião é um dado tipo de moral que é,
inclusive, necessária ao povo. Assim, o que se afirma aqui é uma clássica tese
conservadora. O idealismo desenvolvido por ambos, apesar de todos os
méritos, não deixa de flertar, segundo Piccioni, com um elitismo filosófico já
presente na filosofia hegeliana e, a rigor, presente no próprio platonismo, que
foi incorporado pela posição idealista hegeliana. Introdução Para Galasso, Croce era “idealista porque se inseria no sulco da
tradição do pensamento que de Kant o tinha levado até Hegel” (GALASSO,
2002, p.153). Contudo, o Croce leitor de Hegel, mesmo da Lógica e do
Sistema, nunca deixou de ter em seu horizonte o tema hegeliano por
excelência: a racionalidade dentro da história. A conciliação entre a história e a
filosofia. Aqui, no entender de Galasso, talvez exista uma divergência entre
Croce e Gentile. É certo que ambos foram leitores de Hegel, de Marx e deram
importância ao tema da história. Entretanto, para Galasso, Gentile não teria
conseguido se desapegar do tema da Lógica. Assim, mais do que afirmar que
Croce e Gentile romperam em função do fascismo, Galasso parece construir
um argumento teórico para a primeira separação entre ambos os pensadores, o
que seria mais significativo filosoficamente. Com muita propriedade, Galasso percebe na obra de Croce um
momento historiográfico. Inserem-se nesse horizonte, muitas de suas obras,
mas há um especial destaque para seu trabalho sobre filosofia da história
denominado Teoria e storia della storiografia. Há aqui a afirmação do pensador da
crise da consciência europeia, o que aproxima Croce da sua tradição e de
autores como Burckhardt, Weber, Horkheimer. Desse modo, o contexto
cultural italiano dos anos do fascismo e do pós-fascismo apontam para a
importância de autores como Croce, Gentile e dos pensadores católicos. Aqui
notamos uma questão que mereceria uma investigação mais aprofundada: por
que o idealismo de Croce e de Gentile sempre se reporta aos temas da religião
e da religiosidade? No entender de Galasso, tal coisa ocorre pelo fato da
religião ser, de algum modo, a base do idealismo italiano, como igualmente
suspeitava Viroli na sua obra Come se Dio ci fosse – Religione e libertà nella storia
d´Italia (VIROLI, 2009). Assim, “de resto, fora do idealismo e do reflorescente
catolicismo filosófico, o panorama filosófico italiano não oferecia muito”
(GALASSO, 2002, p.322). Com efeito, nesse clima, percebemos a afirmação
do liberalismo crociano, que fica mais evidente na obra Storia d´Europa nel secolo
decimono, tal como constatamos. Aqui Croce, em seu momento historiográfico,
afirma o tema do liberalismo e da liberdade em oposição ao catolicismo e ao 194 Márcio Gimenes fechamento político italiano dos anos do fascismo e de acordos entre o Estado
e a Igreja. Devemos notar que o ciclo histórico crociano não é maior e nem
menor que sua obra filosófica. Introdução Tal tipo de posição serve tanto
para dar autoridade ao filósofo como aquele que pensa melhor, e por isso
governa e lidera, como também pode fornecer ao mesmo filósofo as bases
para sua crítica. Por isso, segundo Piccioni, Croce como filósofo idealista, está
na base tanto do pré-fascismo como do anti-fascismo, sendo um conservador
de raiz. Assim, seu apreço pelo tema da filosofia da história é um apreço 195 Dissertatio [45] 184-2082017 hegeliano por excelência, um ideal de progresso e superação estão contidos no
seu filosofar: hegeliano por excelência, um ideal de progresso e superação estão contidos no
seu filosofar: A defesa dos valores de cultura preservados daqueles empíricos – na realidade
do processo histórico real portador de novos agentes da história – vem a
constituir a essência do pensamento teórico e político crociano sob um plano de
fechado conservadorismo e a garantir a continuidade mesma do pré-fascismo ao
anti-fascismo (PICCIONI, 1983, p.60). Paolo Rossi chama a atenção para o fato de que os intelectuais
italianos, formados nos anos do fascismo, foram, ainda que depois viessem a
se opor a ele, inegavelmente formados dentro do seu ambiente cultural. Por
isso, segundo ele, há uma reação, de certo modo doentia, ao tentar renegá-lo: Muitos italianos – certo a maioria – foram muito hábeis na arte de fazer
desaparecer o fascismo da sua história pessoal. Intelectuais de várias
proveniências e natureza não se comportaram de modo muito diferente dos seus
compatriotas e tem ao longo aplicado essa mesma técnica à história nacional
(ROSSI, 2012, p.13). Por isso, como já percebeu como argúcia Gramsci, Croce e Gentile,
por exemplo, não podem ser vistos separadamente, visto que fazem parte do
mesmo escopo cultural. Assim, Rossi faz uma curiosa citação, a partir de um
exemplo particular, do quanto tais filósofos eram vistos conjuntamente tanto
antes, como até mesmo depois do fascismo: Meu pai, um estimado e assaz culto professor de Liceu, tinha uma religião laica
de trabalho intelectual e da função da escola clássica, se inspirava principalmente
em Croce nos seus trabalhos de crítica dantesca, mas admirava também Gentile
e lia e apostilava os seus escritos. Era muito longe (por temperamento e por
formação cultural) de cada forma de intolerância e fanatismo, mas julgo que
compartilhava plenamente a imagem gentiliana da história contemporânea da
Itália e o diagnóstico gentiliano sobre o significado e a função história do
fascismo (ROSSI, 2012, p.26). Introdução Uma vez apresentado esse muito breve panorama da filosofia italiana
dos séculos XIX e XX através de dois dos seus principais expoentes, avaliamos
que cabe agora investigar um dos aspectos daquilo que podemos chamar de
idealismo germânico à italiana, a saber, a recepção das obras de Feuerbach,
Kierkegaard e Nietzsche em solo italiano. 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália O filósofo Feuerbach possui uma curiosa recepção ao seu trabalho:
ora sua obra é vista e interpretada entre os hegelianos, ora entre os marxistas e
ora se afirma nos debates teológicos e religiosos. Perone (PERONE, 1972)
aponta que, para além de tais interpretações, parece importante ressaltar que 196 Márcio Gimenes seu legado, de fato, alcança um patamar mais elevado na medida em que tais
posições são superadas e sua crítica começa a ser feita a partir de uma
interpretação científica e historiográfica. Como um exemplo de tal avanço na
pesquisa feuerbachiana notamos, em contexto italiano, a obra do pensador
italiano Franco Lombardi, que ocorre na sequência de outras interpretações
que haviam explorado a relação de Feuerbach com o idealismo: O L. Feuerbach, La Nuova Italia, Firenze 1935, de F. Lombardi, de certo modo,
aproveita-se do precedente, visto que, movendo-se a partir do idealismo, intenta
a sua própria superação através de Feuerbach, e está mais interessado na
contribuição filosófica de Feuerbach do que na sua crítica religiosa. Por outro
lado, isso se afirma sob o fundo da contemporânea situação cultural italiana,
dominada pelo neoidealismo; nela o interesse por Feuerbach assume
imediatamente uma veia polêmica e um significado teorético (PERONE, 1972,
p.183). O L. Feuerbach, La Nuova Italia, Firenze 1935, de F. Lombardi, de certo modo,
aproveita-se do precedente, visto que, movendo-se a partir do idealismo, intenta
a sua própria superação através de Feuerbach, e está mais interessado na
contribuição filosófica de Feuerbach do que na sua crítica religiosa. Por outro
lado, isso se afirma sob o fundo da contemporânea situação cultural italiana,
dominada pelo neoidealismo; nela o interesse por Feuerbach assume
imediatamente uma veia polêmica e um significado teorético (PERONE, 1972,
p.183). Segundo Perone, destacam-se ainda, na primeira metade do século
XX, na crítica italiana de Feuerbach, alguns importantes trabalhos: dois de
Pareyson (PAREYSON, 1950;1971) e as clássicas interpretações de Mondolfo
(MONDOLFO, 1923; 1924), além, evidentemente, da já citada obra pioneira
de Franco Lombardi. Na segunda metade do século XX temos como destaque
dois importantes trabalhos de Claudio Cesa (CESA, 1963; 2003), igualmente
conhecido por suas célebres traduções de obras do pensador alemão. Especialmente nessa última obra, de modo mais elaborado, começa a se
desenhar um estudo de Feuerbach e da importância da temática da consciência
como um ponto importante na reflexão filosófica do autor. 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália Entre os anos
sessenta e setenta do século XX, destacam-se ainda as obras de Rambaldi
(RAMBALDI, 1970; 1966a; 1966b), Perone (PERONE, 1972), Severino
(SEVERINO, 1972), Alessi (ALESSI, 1975), bem como o importante artigo de
Caracciolo (CARACCIOLO, 1978). Nos anos setenta, e já adentrando nos
anos oitenta, ganham destaque na recepção italiana de Feuerbach os estudos
sobre os textos inéditos e póstumos do pensador, os chamados Nachlaβ. Assim, afirmam-se as importantes obras de Casini (CASINI, 1979; 1974) e
Tomasoni (TOMASONI, 1982), estudioso dos inéditos do pensador alemão e
tradutor de algumas obras importantes de Feuerbach, que também publicará,
em 2011, a mais completa biográfica filosófica do pensador alemão disponível
em língua italiana até então (TOMASONI, 2011). Já a recepção kierkegaardiana na Itália, padece daquilo que Cornélio
Fabro, grande especialista e maior tradutor italiano da obra do autor
dinamarquês, denominou como “o provincianismo laico-clerical da cultura
italiana” (PIZZUTI, 1995, p.226). Há, contudo, no seu entender, uma honrosa
exceção: “Se excetua a monografia de F. Lombardi, Kierkegaard (FIRENZE,
1936, republicada integralmente em 1967)” (PIZZUTI, 1995, p.226). Tal 197 Dissertatio [45] 184-2082017 trabalho de Lombardi pode ser melhor compreendido dentro do contexto
intelectual que antecedia à Segunda Guerra Mundial e está especialmente
localizado em torno dos debates culturais promovidos em Florença, tal como
bem acentua Garin e, nesse sentido, trata-se de um Kierkegaard bastante
específico e com a grande virtude de possuir uma abrangência intelectual:
“Como não recordar, apenas para um exemplo, que Kierkegaard, a quem hoje
cabe o amargo destino de servir de tema a tantas dissertações e aulas
universitárias, era apresentado aos italianos propriamente no ‘Leonardo’1 e
traduzido na papiniana ‘Cultura dell´anima’” (GARIN, 1997, p.22). Na esteira dessa interpretação de Lombardi, podemos ainda elencar
outros trabalhos de autores italianos: Abbagnano (La struttura dell´esistenza),
Paci (Principi di una filosofia dell´essere), Carlini (Il mito del realismo), Battaglia (Il
problema morale nell´esistenzialismo), Stefanini (Esistenzialismo ateo ed esistenzialismo
teistico), Cantoni (La coscienza inquieta), tal como acentua Pizzuti (PIZZUTTI,
1995, p.226-228) e Basso (2009, p.82-85). Entretanto, a despeito dos avanços e
méritos de todos esses trabalhos, eles parecem ainda estar englobados naquilo
que se convencionou denominar como a Kierkegaard Renassaince alemã. Desse
modo, Pizzuti avalia que a interpretação de Kierkegaard tem um efetivo
avanço em solo italiano com a obra e com as traduções de Cornélio Fabro, já
na metade do século XX. 1 O Leonardo era uma revista literária. Em 1906, na sua edição de Abril-Junho, ela publicou um ensaio
de Hoffding, célebre comentarista dinamarquês de Kierkegaard. Em 1907, publicou uma tradução de In
Vino Veritas feita por Knud Ferlov. Se pode afirmar, em síntese, que Fabro na Itália realizou o mesmo programa que
Thulstrup promoveu na Dinamarca, a diferença do estudioso dinamarquês, que
ignorou quase totalmente o contributo dos estudiosos católicos, Fabro
conduzido pela mesma lógica da sua linha hermenêutica sublinhou, com uma
insistência que por vezes parecia (mas não era) uma afirmação confessional, a
objetiva convergência de alguns pontos kierkegaardianos com a posição da 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália Fabro, que era religioso e um importante conhecedor de Santo
Tomás e da tradição escolástica, faz uma espécie de ligação de Kierkegaard
com o mundo católico, tal coisa também se pode notar, por exemplo, na
Espanha, na recepção de Unamuno. Contudo, ainda que Fabro tenha tal
posição, ele não compreende, em momento algum do seu trabalho,
Kierkegaard como uma espécie de católico. Para Pizzutti seu trabalho, a
despeito de algumas limitações, é um avanço, por exemplo, em relação a um
outro importante estudioso dinamarquês de Kierkegaard do mesmo período,
Niels Thulstrup, que não coloca nenhuma possibilidade de diálogo entre o
autor dinamarquês e o mundo católico, circunscrevendo-o aos domínios
protestantes e nórdicos: Se pode afirmar, em síntese, que Fabro na Itália realizou o mesmo programa que
Thulstrup promoveu na Dinamarca, a diferença do estudioso dinamarquês, que
ignorou quase totalmente o contributo dos estudiosos católicos, Fabro
conduzido pela mesma lógica da sua linha hermenêutica sublinhou, com uma
insistência que por vezes parecia (mas não era) uma afirmação confessional, a
objetiva convergência de alguns pontos kierkegaardianos com a posição da 198 Márcio Gimenes doutrina católica, e absolutamente tomista, não sem, todavia, abafar as razões
precisas pela qual Kierkegaard não pode ser tomado nem por católico e nem por
cripto-católico (PIZZUTI, 1995, p. 228). doutrina católica, e absolutamente tomista, não sem, todavia, abafar as razões
precisas pela qual Kierkegaard não pode ser tomado nem por católico e nem por
cripto-católico (PIZZUTI, 1995, p. 228). Karl Löwith, pensador que teve curta passagem pela Itália no período
que antece à Segunda Guerra Mundial, é figura importante para
compreendermos a recepção de Feuerbach, Kierkegaard e Nietzsche em solo
italiano. O autor do clássico De Hegel a Nietzsche possui uma relação acadêmica
com três pensadores italianos bastante importantes da época: Giovanni
Gentile, Benedetto Croce e Franco Lombardi. O primeiro deles, autor com
quem o pensador colaborou nas suas publicações, era, na sua concepção, um
homem bastante culto e gentil, embora tivesse ligações com o fascismo. Löwith narra que, numa dada ocasião social, espantou-se com o modo como
ele, a fim de defender a política externa italiana colocou “na mesma panela
fascista, Maquiavel, Mazzini e Mussolini” (LÖWITH, 1988, p.117), o segundo
é merecedor do seu mais alto elogio. 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália Na curta experiência de um passeio em
sua companhia em Nápoles, Löwith encontrou-se com Benedetto Croce, a
quem tomava por um dos poucos espíritos livres em atividade numa Europa
sitiada pelo nazi-fascismo, tal como se pode perceber na elogiosa menção: Eu tive uma vez a sorte de encontrar-me com Croce na sua casa junto com
alguns de seus amigos. Ele depois passeou conosco, pelas ruas de Nápoles, até a
meia-noite e mesmo se nós mais jovens nem sempre pudessemos concordar
com sua avaliação das mutações que ocorriam em nossa época, uma coisa é
certa: na Europa ele é um dos poucos espíritos que permaneceu livre e em posse
de um saber e de uma cultura que fazia envergonhar todos os mais jovens
(LÖWITH, 1988, p.118). Já o terceiro pensador italiano da sua relação o acompanha até o
momento do seu embarque para o exílio no Japão, como podemos atestar por
esse emotivo trecho da sua autobiografia: Em Nápoles passamos ainda um belo entardecer junto a Franco Lombardi, que
no dia seguinte – era 11 de outubro – nos acompanhou ao navio japonês Suwa
Maru. A separação da Itália foi para mim mais dolorosa porque era a separação
de uma terra que tinha negado até mesmo o direito de hospitalidade a um
estrangeiro (LÖWITH, 1988, p.146-147). Tratava-se de Franco Lombardi, autor das obras já citadas sobre
Kierkegaard e Feuerbach. Na visão de Bobbio, suas teses sobre tais os dois
autores germânicos colocam ambos os pensadores numa chave de leitura
secularizada que estabelecerá um novo patamar na interpretação italiana: Com uma singular intuição antecipadora, Franco Lombardi escreve em 1935 um
livro sobre Feuerbach, em 1936 um outro sobre Kierkegaard: dois filósofos que
permaneceram à margem da filosofia acadêmica, reencontrarão à liberação sobre
uma via mestra, um do renascimento marxista, o outro do existencialismo
(BOBBIO, 1986, p.149). 199 Dissertatio [45] 184-2082017 Lombardi escreve também uma importante história da filosofia
italiana compreendida nos últimos cem anos, isto é, entre 1848 e 1948. Em La
filosofia italiana negli ultimi cento anni (s/data), mesmo com a forma um pouco
panorâmica, pode-se perceber a importante contribuição do pensador italiano
em seu inventário. É notória sua tentativa de compreender o pensamento
italiano dentro de um contexto europeu e, por isso, são inúmeras as citações
aos eventos e temas europeus por excelência. 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália Vico, por exemplo, filósofo
moderno italiano, será sempre a fonte de inspiração, coisa igualmente
perceptivel na obra de Croce e Gentile. Assim, não é fortuita que, em 1848,
mesmo período em que Marx e Engels redigiam o seu Manifesto do Partido
Comunista, o autor situe, como ponto importante do contexto filosófico
italiano, o trabalho dos jovens hegelianos de Nápoles: De Sanctis, Bertrando
Spaventa, Salvatore Tomassi e Antonio Labriola. Contudo, Lombardi, que é
ele próprio um leitor atento de Kierkegaard e Feuerbach, acredita, como bom
pós-hegeliano, que é preciso superar a filosofia hegeliana sistemática: “a
libertação da filosofia ‘do sistema’ ou a posição de uma filosofia do homem. Esta constituia, doravante, o programa que seria posto para o pensamento
filosófico italiano” (LOMBARDI, s/data, p.33). Assim, para o pensador, todos
os pensadores antisistemáticos são, de algum modo, irmãos de Kierkegaard. Tal
como ele assim defende ao analisar a filosofia de Labriola: Não diferentemente de Kierkegaard (que de todos foi irmão na sua reação
contra a filosofia “sistemática”, outra que contra a filosofia “do sistema”, isto é,
contra a ideologia hegeliana), esses pensadores podiam dizer de si mesmos: “Vi
um pássaro do qual se diz que prenuncia a chuva – e um tal pássaro sou eu. Se
em uma geração começam a reunir-se as nuvens prenunciadoras de um
temporal... como eu” (LOMBARDI, s/data, p.50-51). Se Vico foi inspirador dos filósofos italianos modernos, Lombardi
sublinha que Galileu será, por sua vez, o inspirador renascentista dos autores
do positivismo italiano, chegando tal influência até mesmo a autores do
idealismo italiano como Croce e Gentile. Por isso, de forma muito espirituosa,
ele compara Croce, o filósofo campeão da liberdade em contexto italiano com
a figura do bispo Mynster como “testemunha da verdade” nos tempos de
Kierkegaard (LOMBARDI, s/data, p.108, nota 1). Não menos espirituosa será
a comparação feita entre Croce e os devotos napolitanos de São Gennaro: “o
culto pátrio se repartiu igualmente em Nápoles entre ‘os parentes’ de São
Gennaro e os parentes de Don Benedetto” (LOMBARDI, s/data, p.119). 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália Kierkegaard, autor a quem Lombardi dedicou uma boa parte dos seus
estudos, é o inspirador de uma contudente crítica de sua parte ao catolicismo
italiano e ao modo como seus compatriotas o vivem enquanto identidade
nacional: “Aqui se poderia perguntar em qual modo se pode falar de 200 Márcio Gimenes ‘convertidos ao catolicismo’ em um país como a Itália: não somos talvez nós
todos se podemos perguntar tal como Kierkegaard nascidos na Itália? E não é
a Itália, por assim dizer, não é talvez a Itália – um reino católico apostólico
cristão? E assim não seremos todos cristãos?” (LOMBARDI, s/data, p.142). O
autor dinamarquês, tomado ainda como pai do existencialismo produz em solo
italiano, alguns bons comentadores e estudiosos no entender de Lombardi. Pareyson, por exemplo, seria um deles. Contudo, a despeito de reconhecer a
importância do trabalho do estudioso de Kierkegaard que foi Cornélio Fabro,
Lombardi é contundente em sua crítica a ele, o que o coloca numa posição
direta de crítica aos kierkegaardianos e faz sua interpretação do pensador
dinamarquês mais próxima dos pós-hegelianos: “Padre Cornélio Fabro que
dos primeiríssimos estudos honestos e severos sobre a psicologia passou a
ocupar-se ativamente do existencialismo kierkegaardiano, com um tentativa de
relacioná-lo com o existencialismo kierkegaardiano, que na verdade repugna a
isso, sob o velho tronco da doutrinas tomistas” (LOMBARDI, s/data, p.180). Nesse mesmo sentido é que podemos compreender a continuidade
daquilo que se convecionará denominar posteriormente de existencialismo
italiano. Aqui, segundo Viano, teremos destaque para os nomes de Luigi
Pareyson, Bobbio, Abbagnano. O foco de tais autores se dará
preferencialmente nos temas da condição humana, da finitude e, com menor
ênfase no tema do irracional. Como se pode notar, são todos temas marcantes
do existencialismo pós-guerra. Feita tal visão panorâmica da recepção italiana de Feuerbach e
Kierkegaard, falta ainda a compreensão da obra nietzschiana na Itália. Segundo
Penzo (PENZO, 1990, p.199-204), a recepção da obra nietzschiana na Itália
possui, no seu primeiro momento, uma configuração literária, tal como ocorre
também na Alemanha. O autor é visto como o pensador anárquico, aquele que
destruiu Deus, a moral, os valores tradicionais. Assim, tal imagem oscila entre
um negativo e um positivo, funcionando como um mito. A democracia se reduz então a uma luta de egoísmos vaidosos, que se
desenvolvem sob o abaixamento sistemático da superioridade legítima e
adquirida. É o triunfo do burguês, do filisteu, da trufa, do asno presunçoso, do pedante que faz o sábio, do idiota que se crê igual ao homem de engenho, de
todas as mediocridades e todas as baixezas (D´ANNUNZIO, 1994, p.14). pedante que faz o sábio, do idiota que se crê igual ao homem de engenho, de
todas as mediocridades e todas as baixezas (D´ANNUNZIO, 1994, p.14). Para o intérprete italiano de Nietzsche, a desigualdade existira na
natureza humana e não reconhecê-la seria ilógico, contrário àquilo que é
natural. Nesse sentido, ele justificaria um direito do mais forte, uma vez que
isso é obtido pela natureza e assim justificaria, inclusive, a colonização africana
(e de outras partes do mundo) pelas nações europeias. O problema mais grave
para D´Annunzio residiria no fato da decadência europeia, já bem percebida
por Nietzsche em sua obra: “Segundo a doutrina de Frederico Nietzsche uma
entre as razões da geral decadência está nisso: que a Europa inteira recebeu a
sua definitiva marca da noção de bem e de mal tomada no sentido moral dos
escravos” (D´ANNUNZIO, 1994, p.14). Segundo Michelini, a interpretação
apresentada por D´Annunzio é, na verdade, não apenas a primeira aparição do
autor em contexto italiano, mas também prova ser a primeira interpretação
feita mais num sentido cultural, isto é, ainda se trata de um Nietzsche muito
geral e menos elaborado de modo acadêmico ou investigativo, tal como se
poderá notar nos anos posteriores: É sobretudo o Nietzsche de D´Annunzio, isto é, o Nietzsche filtrado pelo
esteticismo decadente e provincial e morbidamente sensual do poeta italiano,
aquele destinado a ter maior sucesso e divulgação, próprio porque era mais fácil,
menos profundo e consciente – ao contrário do original – do tormento que
atravessa a inteira civilização ocidental (MICHELINI, 1978, p.18-19). É sobretudo o Nietzsche de D´Annunzio, isto é, o Nietzsche filtrado pelo
esteticismo decadente e provincial e morbidamente sensual do poeta italiano,
aquele destinado a ter maior sucesso e divulgação, próprio porque era mais fácil,
menos profundo e consciente – ao contrário do original – do tormento que
atravessa a inteira civilização ocidental (MICHELINI, 1978, p.18-19). Em 1898, surge em contexto italiano uma tradução, feita por
Edmondo Weisel, de Para além de bem e mal. Note-se, conforme aponta
Michelini, que tal coisa é bastante prematura, uma vez que até mesmo na
Alemanha, pátria do filósofo, algumas obras ainda não haviam aparecido na
integra. Aqui pode haver um aspecto singular na relação do filósofo com a
Itália, também para compreensão da sua recepção entre os italianos. Nietzsche,
como é conhecido, teve diversas estadias na Itália, começando-as em 1876. 2. O idealismo germânico à italiana: a recepção de Feuerbach,
Kierkegaard e Nietzsche na Itália Desse modo, é
significativo que a primeira configuração nietzschiana na Itália se afirme na
interpretação de Gabriele D´Annunzio: “De resto, é significativo que a fortuna
de Nietzsche na Itália seja ligada ao nome de D´Annunzio, que lê Nietzsche
entre 1892-1894 em um senso estético-heróico” (PENZO, 1990, p.199). A interpretação feita por D´Annunzio é ainda importante pela
apresentação de um Nietzsche crítico da democracia, tal como se pode
perceber num texto seu de 25 de Setembro de 1892 e publicado com o
instigante título de A besta eletiva: 201 Dissertatio [45] 184-2082017 pedante que faz o sábio, do idiota que se crê igual ao homem de engenho, de
todas as mediocridades e todas as baixezas (D´ANNUNZIO, 1994, p.14). Como aponta Michelini: “Na Itália, Nietzsche havia encontrado a atmosfera
ideal para escrever a sua obra filosófica. A longa série de estadias no nosso país
foi iniciada treze anos antes, em 1876, e terminava, tragicamente, na cidade
italiana que tinha mais amado, Torino” (MICHELINI, 1978, p.29). Também
Buddensieg, comentador alemão da obra de Nietzsche, corrobora com a tese
da importância das estadias italianas na obra de Nietzsche. Ele, na sua obra A
Itália de Nietzsche, analisa de modo minucioso os nove locais por onde o
filósofo esteve: Sorrento e Napoli, Genova, Veneza, Roma, Florença, Nice
(hoje França), Torino. Tal como também aponta Michelini, que afirmava que
Nietzsche sempre procurou viver na Itália como uma pessoa comum, sem
fazer contatos acadêmicos ou de trabalho, Buddensieg aponta para o quanto o 202 Márcio Gimenes pensador sentia-se à vontade em território italiano: “Contudo, na Itália, ele
vive e escreve e se sente mais como um cidadão das localidades onde habita”
(BUDDENSIEG, 2006, p.19). Por isso, com extrema propriedade, D´Iorio,
estudioso contemporâneo da obra nietzschiana, aponta uma curiosa ligação
afetiva de Nietzsche com o sul da Itália, notadamente por ocasião da sua
passagem por Sorrento: “Da varanda de seu quarto diante de Sorrento, ele
avista a ilha de Ischia, ilha vulcânica, lugar real e imaginário que servirá ao
filósofo de modelo para as ‘ilhas bem-aventuradas’, as ilhas dos discípulos de
Zaratustra, as ilhas bem-aventuradas são as do futuro, da esperança, da
juventude” (D´IORIO, 2014, p.14). Depois dessa primeira interpretação de cunho literário livre e também
de um Nietzsche mais politizado ao estilo fascista, afirma-se, notadamente na
primeira metade do século XX, uma interpretação com um viés existencial,
notadamente presente nas obras de Banfi, Paci e alguns outros, tal como
aponta Penzo (PENZO, 1990, p.199-204). Um importante apontamento sobre
Nietzsche é feito, ainda em 1904, numa resenha feita pelo filósofo italiano
Giovanni Papini a um livro de Francesco Orestano intitulado As ideias
fundamentais de F. Nietzsche no seu progressivo desenvolvimento. Para ele, que era um
crítico de Nietzsche, era preciso que se fosse além da concepção rasa de um
“filósofo de manicômio” para um real compreensão de Nietzsche: A desventura maior de Nietzsche como filósofo foi, depois a loucura, a sua
fortuna. Por alguns anos ele tornou-se o pensador da moda, o oráculo dos
literatos, o fornecedor de frases de efeitos para cenários estéticos. pedante que faz o sábio, do idiota que se crê igual ao homem de engenho, de
todas as mediocridades e todas as baixezas (D´ANNUNZIO, 1994, p.14). Houve uma
grande quantidade de gente que acreditava entender o filósofo porque amava o
artista, e essa admiração, dizemos assim ‘literária’, o tornavam suspeitos aos
filósofos puros. Nem um e nem outro estavam totalmente errados (PAPINI,
1904, p.63). Contudo, conforme Michelini também destaca uma importante
interpretação de Nietzsche em contexto italiano será aquela fornecida por
Benedetto Croce, apresentada em 1907 nas páginas de A Crítica. O pensador
napolitano, mesmo não sendo um comentador de estrito senso do autor
alemão, e ainda que apresentando pontuais discordâncias com o seu
pensamento, aponta, pela primeira vez, para um significado filosófico da obra
nietzschiana, fato fundamental para uma mudança de rumos: Mas já Croce tinha providenciado, por ocasião da tradução italiana do
‘Nascimento da tragédia’, por ele patrocinada, a enquadrar na justa medida o
valor do modo nietzschiano de fazer filologia. Naquele artigo, publicado na
‘Critica’, Croce coloca, como veremos, em relevo a fecundidade daquele método
no estudo dos acontecimentos antigos [...] (MICHELINI, 1978, p.19). Cabe ressaltar que o momento da primeira chegada de Nietzsche na
Itália, ainda nos círculos de D´Annunzio, é marcada por forte influência do 203 Dissertatio [45] 184-2082017 positivismo, o que pode ter forçado o filósofo a ser tomado pelos círculos
literários ou estéticos. Fato que, conforme podemos ver em Croce, é
recuperado de um modo mais preciso. De igual modo, tal como aponta
Michelini, talvez esse seja um contexto adequado para compreendermos o
tema nietzschiano do Super-homem na exagerada interpretação de D´Annunzio:
“D´Annunzio encontrou na moral do Super-homem o meio mais adequado e
próximo da sua personalidade mórbida e sensual para entrar no ciclo da
literatura europeia” (MICHELINI, 1978, p.221). Uma interpretação, que se afirma na segunda metade do século XX,
aproxima Nietzsche de Heidegger e pode ser percebida notadamente na
intepretação de Gianni Vattimo em La fine della modernita (VATTIMO, 2014). Nesse mesmo registro, percebe-se a relação entre Nietzsche e Freud, bem
como entre Nietzsche e Marx, sendo seu articulador o já citado Karl Löwith. Depois da Segunda Guerra Mundial, afirma-se o trabalho interpretativo e de
tradução de G. Colli e M. Montinari, como bem aponta Marton: Ponto de inflexão maior, porém, constitui a edição crítica das obras completas
do filósofo. E isto porque seus efeitos não se limitam à Itália, mas se fazem
sentir nos estudos nietzschianos de maneira geral. pedante que faz o sábio, do idiota que se crê igual ao homem de engenho, de
todas as mediocridades e todas as baixezas (D´ANNUNZIO, 1994, p.14). Não há dúvida de que os
esforços de Giorgio Colli e Mazzino Montinari serão cruciais para trazer a
pesquisa internacional sobre a filosofia de Nietzsche ao ponto em que se
encontra hoje (2007, p.49). Por isso, de modo não fortuito, Garin, estudioso da história da
filosofia italiana, avalia que Nietzsche ainda é um desafio para a filosofia do
século XX: E assim avança o anúncio de Nietzsche, que próprio colocava aquele início de
século no abismo de uma loucura que parece mais que uma crise patológica, a
lúcida consequência de uma posição coerente: Deus está morto, todos os deuses
estão mortos, e também o homem está morto, e a natureza com suas leis
legítimas é o mundo conhecido (1997, p.31). Tal tese não parece equivocada e, na realidade, retoma o problema da
morte de todos os deuses e de toda a metafísica, fato talvez já ocorrido em
Kant. Contudo, o problema é retomado por outro viés interpretativo e, por
isso, adquire significado filosófico e relevância ainda nos dias atuais. Considerações finais A dúvida que surge, logo após o final da Segunda Guerra, é, num
dado grau, a mesma tanto na Alemanha quanto na Itália: um regime filosófico
pode resultar no nazismo ou no fascismo? Nesse sentido, a crítica alemã
operada por Adorno e Horkheimer em A dialética do esclarecimento teria estreita
relação com a crítica italiana feita a posteriori, ou seja, Giovanni Gentile seria, no 204 Márcio Gimenes contexto italiano, o equivalente daquilo que teria sido Heidegger? Em ambos,
pode-se perceber que a tomada de posição política é uma consequência do seu
pensamento filosófico. O mesmo também ocorre na posição política de Croce,
liberal oposto ao regime e na posição politica socialista de Franco Lombardi. Em outras palavras, a oposição ao regime possui uma raiz muito próxima do
mesmo idealismo que ajudou na sustentação do regime. Assim, aquilo que chamamos como filosofia italiana entre os séculos
XIX e XX é melhor percebida, talvez, com uma investigação da influência
alemã. Um dado curioso é que talvez passe desapercebido é que Croce e
Gentile, como bem pontua Verucci (2006) foram condenados ao mesmo
tempo pela Igreja Católica, isto é, em 1932. Ambos foram tomados como
autores que pouco nacionalistas e que acentuavam demais a chamada filosofia
estrangeira isto é, a filosofia idealista alemã. Desse modo, “a filosofia de Croce
nascia da ‘fonte teutônica’, isto é, hegeliana, e gerou uma filosofia similar e
talvez também, se fosse possível, pior que a sua, isto é, o atualismo”
(VERUCCI, 2006. p.145). Desse modo, se os italianos possuem uma dada recepção de Hegel,
eles também possuirão uma interpretação muito peculiar dos filósofos da vida,
isto é, dos pós-hegelianos que se lançam contra o mestre Hegel. Aqui
podemos, incluir autores como Nietzsche, Schopenhauer, Feuerbach,
Kierkegaard e inúmeros outros. Para Viano, as críticas de Croce como
antiacadêmico podem ser compreendidas dentro desse espírito. D´Annunzio,
precursor de uma dada interpretação nietzschiana na Itália, estaria na mesma
linha, com um pouco menos de ênfase. Assim, Viano ressalta aqui um
isolamento da filosofia italiana, que se perdia internamente num debate entre
idealistas e positivistas (VIANO, 2006), crítica igualmente realizada por Garin
(GARIN, 1997). Passadas as guerras e querelas mais marcantes, talvez seja essa
a época de uma observação menos contaminada e mais ampla do que foi – e
do segue sendo – a filosofia italiana e de qual seria a sua relavância no contexto
atual dos estudos filosóficos. Considerações finais Curiosamente, talvez Adorno e Horkheimer, de
algum modo, espelham um pouco da crítica do idealismo italiano e da sua
crítica ao cristianismo: “A velhinha italiana que, em sua piedosa simplicidade,
consagra uma vela a San Gennaro em favor do neto que partiu para a guerra,
pode estar mais próxima da verdade do que os papas e bispos que, imunes à
idolatria, abençoam as armas contra as quais San Gennaro é impotente. Mas,
para a simplicidade, a própria religião torna-se um sucedâneo da religião”
(ADORNO e HORKHEIMER, 1985, p.167). Logo, talvez existam mais pontes
entre a Itália e a Alemanha do que se possa imaginar num primeiro exame. Referências Referências 205 Dissertatio [45] 184-2082017 ADORNO, T. W. e HORKHEIMER, M. Dialética do Esclarecimento. Rio de
Janeiro: Jorge Zahar Editor, 1985. ALESSI, A. L´ateismo di Feuerbach. Roma: LAS, 1975. BASSO, I. “Italy: From a Literary curiosity to a Philosophical Comprehension”. In: Kierkegaard´s International Reception, v. 8, Tome II, Southern, Central and
Eastern Europe, edited by Jon Stewart, Farnham, Ashgate, 2009 BOBBIO, N. Profilo ideológico del novecento italiano. Torino: Einaudi, 1986. CARACCIOLO, A. “Dio e spazio religioso in Feuerbach”. In: AA.VV.,
Dialettica e Religione, vol. I, Benucci, Perugia, 1978, p. 203-269. BUDDENSIEG, T. L´Italia di Nietzsche. Milano: Libri Scheiwiller, 2006. CASINI, L. Feuerbach postumo. Il panteismo della Lezioni di Erlangen. Firenze:
Sansoni, 1979. ______. Storia e umanesimo in Feuerbach. Bologna: Il Mulino, 1974. ______. Cesa, C. Il giovane Feuerbach. Bari: Laterza, 1963. ______. Introduzione a Feuerbach. Roma: Laterza, 2003. CROCE, B. Storia d´Europa nel secolo decimono. Roma: Laterza, 1965. ______. Storia d´Italia dal 1871 al 1915. Roma: Laterza, 1977. ______. Teoria e storia della storiografia. Milano: Adelphi, 2001. D´ANNUNZIO, G. Su Nietzsche. Catania: De Martinis & C. Editori, 1994. D´IORIO. Nietzsche na Itália – a viagem que mudou os rumos da filosofia. Rio de
Janeiro: Zahar, 2014. ESPOSITO, R. Pensamento vivo – origem e atualidade da filosofia italiana. Belo
Horizonte: Editora UFMG, 2013. GALASSO, G. Croce e lo spirito del suo tempo. Roma: Laterza, 2002. GARIN, E. Cronache di filosofia italiana 1900-1960 v.1. Roma: Laterza, 1997. ______. Storia della filosofia italiana – volume terzo. Torino: Einaudi, 1966. GENTILE, G. Giordano Bruno e il pensiero del Rinascimento. Firenze: Vallecchi
Editore, 1920. ______. La religione. Firenze: Sansoni, 1965. GRAMSCI, A. Gli intellettuali e l´organizzazione della cultura. Torino: Einaudi,
1949a. 206 Márcio Gimenes ______. Il materialismo storico e la filosofia di Benedetto Croce. Torino: Einaudi,
1949b. LOMBARDI, F. Feuerbach. Firenze: La Nuova Italia, 1935. ______. La filosofia italiana negli ultimi cento anni. Asti: Casa Editrice Arethusa,
s/data. ______. Kierkegaard. Firenze: La Nuova Italia, 1936. ____. Kierkegaard. Firenze: La Nuova Italia, 1936. LOSURDO, Domenico. Dai Fratelli Spaventa a Gramsci – per una storia politico-
sociale dela fortuna di Hegel in Italia. Napoli: La Città del Sole, 1987. LÖWITH, K. La mia vita in Germania: prima e dopo il 1933. Milano: Il Saggiatore,
1988. MARTON, S. (org.). Nietzsche pensador mediterrâneo: a recepção italiana. São Paulo:
Discurso, 2007. MICHELINI, G. Nietzsche nell´Italia di D´Annunzio. Palermo: S. F. Flaccovio
Editore, 1978. MONDOLFO, R. La Filosofia política in Italia nel secolo XIX (conferenze sulla Storia
d´Italia del secolo XIX svolte nell´anno accademico 1922-1923. Padova: La Litotipo
Editrice Universitaria, 1924. ______. Sulle orme di Marx. Bologna: Cappelli, 1923-1924. PAPINI, G. “Francesco Orestano – le idee fondamentali di Fed. Niezsche nel
loro svolgimento”. In: La Critica. Rivista di Letteratura, Storia e Filosofia diretta da
Benedetto Croce, 2, 1904, p. 63-66. PAREYSON, L. “Due possibilità: Kierkegaard e Feuerbach”. In: Esistenza e
persona, Taylor, Torino, 1950. ______. Studi sull´esistenzialismo. Firenze: Sansoni, 1971. PENZO, G. Invito al pensiero di Nietzsche. Milano: Mursia, 1990. PERONE, U. Teologia ed esperienza religiosa in Feuerbach. Milano: Mursia, 1972. PICCIONI, L. Ideologia e filosofia del neoidealismo italiano. Urbino: Università degli
Studi di Urbino, 1983. PIZZUTI, G.M. Invito al pensiero di Kierkegaard. Milano: Mursia, 1995. RAMBALDI, E. Feuerbach 1842 ‒ Necessità di un cambiamento. Firenze: Sansoni,
1970. ______. La critica antispeculativa di L. Feuerbach. Firenze: La Nuova Italia, 1966a. 207 Dissertatio [45] 184-2082017 ______. Le origini della sinistra hegeliana. Firenze: La Nuova Italia, 1966b. ROSSI, P. Un breve viaggio e altre storie – le guerre , gli uomini, la memoria. Milano:
Raffaello Cortina Editore, 2012. ______. Le origini della sinistra hegeliana. Firenze: La Nuova Italia, 1966b. ROSSI, P. Un breve viaggio e altre storie – le guerre , gli uomini, la memoria. Milano:
Raffaello Cortina Editore, 2012. SEVERINO, G. Origine e figure del processo teogonico in Feuerbach. Milano: Mursia,
1972. TOMASONI, F. Feuerbach e la dialettica dell´essere. Firenze: La Nuova Italia,
1982. ______. Ludwig Feuerbach – Biografia intellettuale. Brescia: Morcelliana, 2011. VATTIMO, G. La fine della modernità. Milano: Garzanti, 2014. VERUCCI, G. Idealisti all´Indice – Croce, Gentile e la condanna del´Sant´Uffizio. Roma: Laterza, 2006. VIANO, C. . La filosofia italiana del Novecento. Bologna: Il Mulino, 2006. VIROLI, M. Come se Dio ci fosse – Religione e libertà nella storia d´Italia. Torino:
Einaudi, 2009. E-mail: marciogimenes@unb.br E-mail: marciogimenes@unb.br Recebido: 31/05/2016
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UND FRIEDENSFORSCHUNG
| Digol, Debate in Russia on Conflict in Georgia
NATO-Mitglieder bzw. ihrer Gesellschaften vor politischem
Zwang zum Kernbereich des Artikels 5 gehöre. Daher führe der
Kampf gegen eine terroristische Bedrohung die Allianz – wenn
auch unter veränderten Vorzeichen – wieder an ihren Beginn
zurück. Eine solche strategische Besinnung sei nicht nur unter
sicherheitspolitischen Aspekten dringend geboten, sondern ge
währleiste auch die notwendige Legitimation der NATO in den
Mitgliedstaaten, die durch die Vielzahl von Auslandseinsätzen
in den vergangenen Jahren nicht (mehr) hergestellt werden
konnte. Unter dem Überbegriff »Transatlantic Homeland Defense« plädieren diese Beobachter daher für eine multidimensio
nale Strategie der Allianz, die Elemente diplomatischer, finan
zieller, wirtschaftlicher, geheimdienstlicher und polizeilicher
Zusammenarbeit umfassen soll. Konkret soll die Zivilverteidi
gung in die Allianzdoktrin mit dem Ziel aufgenommen wer
den, das Territorium der NATO vor terroristischen Angriffen
zu schützen.25 Ob diese Vorschläge jedoch geeignet sind, die
skizzierten strukturellen Beschränkungen zu überwinden, darf
bezweifelt werden.
25 Vgl. Amas, Neyla u.a., Transatlantic Homeland Defense, Center for Techno
logy and National Security Policy – National Defense University (Mai 2006),
Special Report (<www.ndu.edu/inss/press/CTNSP-INSS/spl-rpt.pdf>); sowie
Hamilton, Daniel S., NATO Summit I: In Area, or in Trouble, in: International
Herald Tribune, 27.11.2006. Am stärksten wird diese Position von dem ehema
ligen spanischen Ministerpräsidenten Aznar vertreten. Vgl. Aznar, Jose-Maria,
NATO – An Alliance for Freedom, in: The RUSI Journal, 151 (2006) 4, S. 38-40.
Right or wrong: Debate in Russia on Conflict in Georgia
Diana Digol*
Abstract: The article presents a comprehensive overview of opinions in Russia on the August 2008 military conflict with Georgia.
The positions presented include not only those of the political elite and political opinion-makers but also those of the cultural
and social elite and of representatives of various ethnic groups involved in the conflict. The overview concludes by presenting
results of public opinion polls. The period covered is between the commencement of hostilities on 8 August 2008 and the end of
that year. The article is limited to Russia and does not cover the debates of neighbouring countries.
Keyworlds: Russia, Georgia, conflict, opinion
Russland, Georgien, Konflikt, Meinung
1. In����������
troduction
F
rom the start of the conflict between Georgia and Russia
the leading Western news agencies dedicated several re
ports daily to this conflict. The desire to understand the
events was overwhelming. Experts on Russia and the Caucasus
were in high demand by local and national news agencies. Ini
tial assessments rapidly found their way into major European
and American news channels and newspapers. The majority of
the media blamed Russia for the developments and were look
ing for details.
What was happening in Russia? How were these events shown,
interpreted, received and discussed within Russia, by Russians,
Georgians, Ossetians and others? From the outset of the con
flict, an increasingly intense and comprehensive debate took
place on TV, radio and in print media. The mass media provid
ed a platform to numerous persons of various political inclina
tions as well as apolitical individuals, intellectuals, academics
and artist.
This article presents a comprehensive overview of the opinions
in Russia on the August 2008 military conflict between Georgia
and Russia. The premise is that analysing opinions of a variety
of people would help to create a picture of these events from a
Russian point of view. Telling this story, I speak with the voices
of the people allowing myself to use summaries of their argu
112
| S+F (27 . Jg.) 2/2009
ments. I conclude the overview by presenting the results of
public opinion polls.
The opinions of the two highest ranking leaders of Russia,
Dmitry Medvedev and Vladimir Putin are presented to deline
ate the official position of the political elite. The opinion of
the official opposition (parties, other than the ruling one, rep
resented in the parliament) is presented through the voices of
Gennady Zyuganov, leader of the Communist Party of the Rus
sian Federation (KPRF), Vladimir Zhirinovsky, the leader of the
Liberal Democratic Party of Russia (LDPR) and Sergei Mironov,
a leader of the party “A Just Russia”. The position of other par
ties (unofficial opposition) is presented through the voices of
Mikhail Kasyanov, leader of People’s Democratic Union (PDU),
Sergei Mitrokhin, a leader of the “Yabloko” party and Garry
Kasparov, a leader of “United Civic Front”.
The views of Natalia Narochnitskaya, from the Institute for
Democracy and Cooperation, Stanislav Belkovskii, Director of
the Institute for National Strategy and Alexandr Konovalov,
President of the Institute of Strategic Evaluation are presented
to delineate the position of Russian political scientists and ana
lysts. The views of Russian intellectuals are presented by the
journalist Vladimir Pozner and the writer Boris Strugatskii. In
*
Dr. Diana Digol is researcher at the Center for OSCE Research (CORE) at the
Institute for Peace Research and Security Policy (IFSH), Germany.
In the current State Duma (October 2008), only these three parties, in addi
tion to the governing party “United Russia”, are represented.
https://doi.org/10.5771/0175-274x-2009-2-112
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Digol, Debate in Russia on Conflict in Georgia
addition, the reactions of a group of intellectuals from St. Pe
tersburg and of the Imperial House of Russia are presented. The
Georgian elite is represented by the TV star Tina Kandelaki, the
journalist Otar Kushanashvili, the singer Vakhtang Kikabidze,
Nikolai Tsiskaridze, a principal dancer of the Bolshoi Ballet,
and a number of others. The position of the Ossetian elite is
represented by opera director Valery Gergiev and the actors
Bulat Budaev and Soslan Fidarov. To conclude, the opinion of
the general public is summarised, based on results of the two
prominent opinion polls in Russia: the public opinion research
centre VCIOM and the non-governmental organization “Le
vada Analytical Centre” (“Levada-Center”).
This article is different from other articles on similar topics
in presenting not only the position of the political elite and
political opinion-makers, but also that of members of the cul
tural and social elite as well as representatives of various ethnic
groups involved in the conflict. These views are then contrasted
with results of the public opinion polls. The article is limited
to the period between the 8 of August 2008 and the end of
December 2008, when research for the article was finalised. It is
confined to Russia and does not include debates of neighbour
ing countries.
2. Political elite
2.1 Official position
The official position of Russia has been framed and expressed
by Medvedev and Putin in political addresses, speeches and
interviews (Medvedev, 2008a, 2008b, 2008c, 2008d, 2008e,
2008f, 2008g, 2008h), (Putin, 2008a, 2008b, 2008c, 2008d).
It has been broadcasted on national TV and published in na
tional newspapers. Hence, the official position is that Georgia
attacked not only the small territory of South Ossetia, but also
the lightly armed peacekeepers, causing deaths mainly among
Russian peacekeepers. This fact served as additional evidence
for the official position that Georgia planned in advance and
prepared an attack on South Ossetia and was, potentially, plann
ing a similar one on Abkhazia. As a response, Russia was forced
to intervene with the goal of preventing a humanitarian cri
sis and a new genocide. It was argued that this decision was
deeply rooted in the long term historical ties existing between
Russia and the people of South Ossetia and the geographical
proximity of Russia to South Ossetia, making Russia the only
possible state able to intervene in a timely fashion. Thus, Russia
had no choice but to intervene promptly to protect the civil
ian population; in this respect a parallel has been drawn to the
massacres in Srebrenica in July 1995, when Dutch peacekeepers
preferred to retreat rather than intervene, which literally led to
the deadly consequences (Putin, 2008c).
It has been further argued that Russia was against any new form
of confrontation and wished to avoid any new edition of the
“Cold War” and global isolation of Russia. Russia expected to
Arguably, Georgian peacekeepers were informed about the imminent attacks
and left their positions beforehand (“Peace enforcement: war chronicle,”
2008).
| BEITRÄGE AUS SICHERHEITSPOLITIK
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find understanding in the West (Medvedev, 2008h), (Putin,
2008b).
As far as the economic consequences are concerned, Russia is
aware that the West can try to use leverage against Russia in
the economic sphere, in particular with respect to its attempts
to become a WTO member. However, it was stressed again and
again that when a decision had to be taken to intervene and
protect the people of South Ossetia economic considerations
were of secondary importance (Medvedev, 2008g), (Putin,
2008a).
Regarding the recognition of independence of South Osse
tia and Abkhazia and related accusations of Russia applying
double standards in the case of South Ossetia versus the case
of Chechnya, Medvedev and Putin, drew a parallel between
South Ossetia and Kosovo rather than between South Ossetia
and Chechnya. The argument is that as much as Kosovo was
a casus sui generis, South Ossetia and Abkhazia were also such
cases (Medvedev, 2008d), (Putin, 2008c). Moreover this recog
nition was the implementation of the UN principle of the right
of self-determination of nations.
The Georgian attack is believed by Russian officials to have been
staged by Americans and with Americans; their physical pres
ence in the midst of military action has been suspected and is
being investigated. In the words of Medvedev, the Americans
must be feeling sad that their “virtual project ’Free Georgia’ had
failed” (Medvedev, 2008h). A possible link between the elec
toral campaign in the USA and the need to support one of the
presidential candidates was also mentioned (Putin, 2008c).
2.2 Official opposition
The position of the official opposition is presented through the
party positions of the only three opposition parties in the cur
rent State Duma: the Communist Party of the Russian Federa
tion (KPRF), the Liberal Democratic Party of Russia (LDPR) and
“A Just Russia” Party.
The position of the Communist Party of the Russian Federation
(KPRF) was in line with the official position. Its leader Gennady
Zyuganov argued that Europe should have been grateful to Rus
sia for stopping a big Caucasian war. There was no doubt, he
continued, that Georgia initiated military actions by assaulting
and killing the Russian peacekeepers (Zyuganov, 2008b). It was
Russia’s military duty to get involved in the conflict (Zyuga
nov, 2008a). The KPRF organised a protest meeting in front of
the Georgian Embassy, waving banners with slogans such as:
“Freedom to South Ossetia”, “Saakashvili – to the Hague”, etc.
(Vladov, 2008). Moreover, the KRPF authored an appeal to the
Russian government asking them to stop the colonial regime
of Saakashvili and to immediately recognise the independ
ence of South Ossetia and Abkhazia (Zyuganov, 2008a, 2008b,
2008c).
One might recall, however, that in the case of Serbia-Kosovo, Russia was sup
porting Serbia’s position arguing for the precedence of territorial integrity
over the right of self determination. The change of the position is explained
in the interview of Putin to the German channel ARD (Putin, 2008c).
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| Digol, Debate in Russia on Conflict in Georgia
The position of the Liberal Democratic Party of Russia (LDPR)
was also in line with the official position, but was more radical
and far-reaching in its interpretations and suggestions (Zhiri
novsky, 2008b). Thus, the party’s leader, Vladimir Zhirinovsky,
argued that a pre-emptive attack of Russia would have been a
much better scenario (Zhirinovsky, 2008c). He stated that in
such a case, Tskhinvali and the lives of thousands of Ossetians
would have been saved. The recognition of South Ossetia was
justified: never again would Ossetians be living together with
Georgians (Zhirinovsky, 2008a).
The party “A Just Russia” was equally united in its opinion and
supportive of official Russia’s position. In the opinion of its
leader, Sergei Mironov, Russia had no other choice but to inter
vene since Georgia was systematically physically eliminating
the local Ossetian and Abkhazian population. He described the
actions of Saakashvili as “a strategic mistake”, further adding
that “He showed the world his intentions towards his people.
He is killing them” (Mironov, 2008a). Mironov believes that
with his actions “Saakashvili has signed a death sentence for
himself” politically (Mironov, 2008b). The party views the
worsening of the relations between Russia and the EU as tem
porary and foresees no serious economic consequences for Rus
sia (Ponomarev, 2008).
2.3 Non-official opposition
By contrast to the official positions, the former prime minister
of Russia, and currently a leader of the opposition party, The
People’s Democratic Union (PDU), Mikhail Kasyanov said that
Russia had continuously and incessantly provoked Georgia. It
was regrettable that Georgia attacked militarily; however, Rus
sia should have stayed within its peacekeepers’ mandate. In
stead it launched a full-scale war. He thoroughly criticised the
actions of Russia’s government and argued that these actions
were unprofessional and disproportionate. By these actions, he
continued, Russia turned itself from a peacekeeper and media
tor in the region into a party to the conflict, which might lead
to international isolation. He was one of the first to suggest
that, as a consequence, Russia might be deprived of hosting
the Olympic Games in 2014 (Stack, 2008).
The “Yabloko” party expressed concerns about the situation in
South Ossetia. Its leader, Sergey Mitrokhin, expressed his wish
that Russia should not turn from a “saviour of lives in South
Ossetia” into an aggressor and should stop the war as soon as
possible (Mitrokhin, 2008b). At the same time, he appealed to
the NATO countries to take the issue of Georgia’s membership
in NATO off its agenda (Mitrokhin, 2008a).
The Russian chess grandmaster, former World Chess Champi
on and nowadays writer and political activist as well as leader
of the political movement “United Civic Front” (UCF), Garry
Kasparov, partially echoed the official position. In particular,
he did not doubt that it was Saakhasvili who had provoked the
war (Kasparov, 2008c). However, he maintained that the postwar developments were against the interests of Russia and its
citizens, that they contradict the international treaties Russia
has signed and ratified and conflict with the norms of inter
national cooperation (Kasparov, 2008a). Moreover, he blamed
114
| S+F (27 . Jg.) 2/2009
Russia for acting against the agreed-upon Sarkozy-Medvedev
plan for the peaceful regulation of the conflict in South Ossetia
and Abkhazia (Kasparov, 2008a). As a consequence, he envi
sioned a growing instability in the Caucasus and the prolifera
tion of separatist movements, the complete political isolation
of Russia, the acceleration of negative tendencies in the Russian
economy and the weakening of its international position. By
recognising South Ossetia and Abkhazia, Russia has prepared a
long-term “headache” for itself (Kasparov, 2008b). In short, he
viewed this conflict and Russia’s response as a military political
adventure with catastrophic political consequences.
3. Social and cultural elite
A variety of divergent opinions of Russia’s intellectuals, includ
ing those of ethnic Georgians and Ossetians are presented be
low.
3.1 Political analysts
Natalia Narochnitskaya, head of the Paris office of the Institute
for Democracy and Cooperation, a Russian non-profit founda
tion, defined the operation in South Ossetia as Russia’s victory.
For the first time in the last 15 years Russia acted as suggested
by its “national pride, historical obligations and own interests”
(Ovchinnikov, 2008). Russia demonstrated that the opinion of
its own people was more important than the opinion of the
West. According to her, the whole Western rhetoric about the
Georgia-Ossetian conflict was actually not because of Georgia,
but because of the new rising power and will of Russia. She was
sure that if Georgia had been located somewhere else geograph
ically, for example in Siberia, no one would have cared about it.
She argued that the “anti-Russian policy of Georgia is necessary
for the West” (Ovchinnikov, 2008). Georgia was supposed to
serve the purposes of “squeezing Russia out” as a main military
player in the Black Sea area (Ovchinnikov, 2008).
Stanislav Belkovskii, director of the Institute for National Strat
egy, argued that Saakashvili had achieved two of his three goals.
First, the total destruction of the infrastructure of South Ossetia
and its capital, Tskhinvali, was achieved as well as “cleansing”
the territory of Ossetians by making them flee. Second, Russia
was deprived of the exclusive status of a peacekeeper in the re
gion, which was his long-term goal. He failed only in his third
goal, which was to take full control of the territory of South
Ossetia. Belkovskii further argued that accepting the SarkozyMedvedev plan would mean a full defeat for Russia. Recognis
ing South Ossetia and Abkhazia was the only winning strategy
which would allow Russia to legitimise its economic and mili
tary presence in the region. What seemed probable in the im
mediate aftermath of the crisis – depriving Russia of the right
to hold the Olympic Games in Sochi – is no longer relevant.
The reasoning is that the Olympic Games are at first instance
a commercial project and involve many partners, including
foreign ones.
Alexandr Konovalov, president of the Institute of Strategic
Evaluation, considered that the events might have sped up the
accession of Georgia into NATO, but without the two regions
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Digol, Debate in Russia on Conflict in Georgia
of South Ossetia and Abkhazia (Mekhtiev, 2008). He doubted
that a change of leadership in Georgia was one of Russia’s aims
in this conflict and argued that in Georgia “there is no strong
political leader who is also pro-Russian”(Mekhtiev, 2008).
Moreover, the conflict influenced the whole of Georgian antiRussian sentiments. The main outcome for Russia, according to
him, is that the USA and Western Europe understood that they
“should take into account the position of Russia, in particular
in matters related to the Caucasus” (Mekhtiev, 2008). He also
said that Russia stopped at the right moment, not crossing the
line after which a new “Cold War” would have been unavoid
able (Mekhtiev, 2008).
3.2 Russian intellectuals
Vladimir Posner, a Russian journalist and president of the
Russian Academy of TV (until October 2008), might be known
to a Western reader from the first talk shows and TV bridges
between the Soviet Union and USA, which he initiated in the
1980s. He was the one who suggested in 2007 TV-bridges be
tween Russia and Georgia to talk about the conflict among all
parties and look together for ways to solve it. According to him,
“every person who sees a bit further than his own nose should
have understood that after the recognition of Kosovo, Russia
would, inevitably – with 98 per cent probability – recognise
South Ossetia and Abkhazia” (Posner, 2008). According to Pos
ner, Putin, at that time still president, was talking about it very
clearly. To a Russian, the situation in Kosovo is not different
from the one in Abkhazia and South Ossetia. It was clear that if
Georgia would not change its policy and attitude toward these
entities (South Ossetia and Abkhazia), then this would be an
outcome. He continued by arguing that it is difficult now to im
agine that South Ossetia and Abkhazia would ever again wish
to become a part of Georgia. The Georgians have humiliated
and mistreated Abkhazians since Soviet times. This attitude was
exacerbated during the rule of former Georgian president Zviad
Gamsakhurdia and has continued during the rule of Saakash
vili. Although not easy, recognition of these entities was the
right thing to do from the point of view of Russian state interest
(Posner, 2008).
With respect to consequences for Russia, these events indeed
placed Russia in a position of international isolation. There
is, however, mutual interdependence between Russia and the
West, thus Posner believed that Russia would not be excluded
from the “G8”, no new war was imminent and no return to the
Iron Curtain was possible. However, the image of Russia was
certainly damaged. Posner suspected American involvement
as well; he made particular reference to partisan coverage of
the crisis on American TV by which he felt deeply disgusted
(Posner, 2008).
A famous Soviet and Russian science fiction writer, Boris Stru
gatskii mentioned philosophically that he “cannot think of
any politician, any political power which is not guilty either
through its action or through its inaction” (Strugatskii, 2008).
This might have created the impression that he approved of the
He had his own political TV-show “Vremena” which, once a week, on Sunday
evening, replaced the official evening news.
| BEITRÄGE AUS SICHERHEITSPOLITIK
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actions taken by Russia. However, he continued by describing
Russia as a totalitarian regime which thrives on short victorious
wars. He further stated that the Caucasus has been balancing
between peace and war since the collapse of the USSR. One of
the reasons is that Russia and Russian elites will never agree to
let Georgia out of its sphere of interest. In the dilemma between
the “right to self determination” and “territorial integrity”,
“power” wins. And only in those cases where “power” is not
interested, can the UN be useful somehow (Strugatskii, 2008).
Strugatskii compared South Ossetia to Chechnya, and argued
that Georgia did not do in South Ossetia anything Russia had
not done in Chechnya, even using the same justification of
“bringing constitutional order to separatist regions” (Vish
nevskyi, 2008a). He further blamed Russia for a lack of consist
ency in its positions. He argued Russia had negatively assessed
the actions of the USA in Serbia against the Milosevic regime,
but then, in his view, took similar actions in Georgia (Vish
nevskyi, 2008a).
A group of intellectuals from St. Petersburg, including poets,
writers, artists, professors, political scientists, and sociologists,
wrote an open letter to their colleagues in Georgia. This letter
addressed to the Shota Rustaveli Institute of Georgian Litera
ture in Tbilisi was in essence dedicated to all Georgian intel
lectuals. The letter containing the words “this is not our war
and [these are] not our tanks” (Vishnevskyi, 2008b) and stated
that the politicians of two or maybe three states put cultural
and human relations between Russia and Georgia at risk. It also
recalled that WWII influenced the attitude of Russians towards
the Germans. Thus, the authors of the letter were concerned
about preventing a similar change in attitude between Rus
sians and Georgians. The answer from the Georgian side, little
expected from the Russian side mainly because of the broken
lines of communications, arrived promptly. The letter was full
of gratitude for a public expression of opinion and hope that
the links between intellectuals would remain untouched and
unchanged (Petlyanova, 2008). This was one of the first signs
that human relations might be above the politics.
The Imperial House of Russia echoed the official position of
Russian government. The Head of the House of Romanoff,
H.I.H. Grand Duchess Maria Vladimirovna is convinced that
“the present conflict in South Ossetia has its origins abroad.”
She expressed her profound anguish and “is troubled by the ac
tions of political leaders, who are only fulfilling the will of for
eign masters and, because of this, have turned the territory of
their own country into a bombing range for foreign weapons,
and who have sacrificed their own people for foreign interests”
(Imperial House of Russia, 2008).
3.3 Famous Georgians who made a career in the
USSR or in Russia
This section features various famous persons of Georgian ori
gin (not necessarily with Georgian citizenship) who are or were
active in Russian intellectual or cultural life. Thus, Tina Kan
delaki, TV star and one of the top television presenters, nick
named in the Russian press the “Main Georgian of Moscow”
(Antonova, 2008), argued in her blog that the Georgian people
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| Digol, Debate in Russia on Conflict in Georgia
were against the Saakashvili regime and were against the war
(Kandelaki, 2008). She described Saakashvili as a person with
weak nerves but high ambitions and narcissism who compares
himself to David the Builder. She argued, “Saakashvili is bound
to go down into history of Georgia as Mikheil the Destroyer”
(Walker, 2008). She also suspected American involvement in
this conflict and suggested that the main interest of Americans
in Georgia were not people but strategy: “We are only 40 min
utes away from Iran” (Walker, 2008). She blamed Saakashvili,
however, she believed that Russian-Georgian relations would
shortly resume on a normal path again since their cultures have
been intertwined for centuries.
Nikolai Svanidze, a famous TV journalist, working at the progovernment channel “Russia”, author of Medvedev’s biogra
phy and creator of the “Historical Chronicles” was one of the
most outspoken persons during the days of conflict and im
mediately afterwards (Svanidze, 2008). He argued that Russia
would have scored higher politically if it had not crossed the
border between South Ossetia into Georgia. Additionally, he
saw the recognition of South Ossetia and Abkhazia in the af
termath of the military actions as inevitable (Svanidze, 2008).
He further argued that these events showed that Russia is not
surrounded by a circle of friends (referring to the nearby states),
and there are few friends further away (referring to the rest of
the world). He is concerned with an intensification of Russia’s
isolation. In consequence, he sees that it would be a difficult
task to increase Russia’s influence on the international scene
(Svanidze, 2008).
Otar Kushanashvili, a charismatic journalist and TV presenter,
openly named Saakashvili a “recruitee” of the USA and believed
that he should be judged by an international criminal tribunal
for these events (Russia.Ru, 2008). He further argued that no
single life is worth to be lost to enhance the self-esteem of a
politician and called for politicians on both sides to overcome
their ambitions and start negotiations immediately (Baranets,
2008). Kushanashvili stated that Georgia had no basis for its
souvereignty. In his opinion, the only real partner of Georgia
is Russia. Thus, Georgia should search for ways to restore its
friendship with Russia as soon as possible. As for Saakashvili,
the only good move he could still make for Georgia was to vol
untarily give up his post and ask the Georgians for forgiveness
(Russia.Ru, 2008).
Vakhtang Kikabidze, a Georgian and Soviet singer, screenwriter,
producer, composer and actor who had played in an iconic So
viet movie “Mimino”, was truly exceptional in his scale of criti
cism of Russia. He accused Russia of staging a long-planned
military and political attack and then occupying Georgia, as
sessing it as act against Saakashvili and his rule. As a sign of
protest against Russian aggression in Georgia he cancelled a
scheduled concert in September in St. Petersburg and refused
an Order of Honour decoration, offered by the President of
Russia, to honour his 70th birthday (Pleshakova, 2008). The
translation of his critical interview in the Georgian newspaper
“Kvela Siakhle” was widely discussed in Russian mass media.
His words concerning the actions Georgia should take against
Russia were initially translated from Georgian into Russian,
as Georgia “should bring this country [Russia] to an end” or
“should put an end to this country [Russia]” (Vorsobin, 2008).
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| S+F (27 . Jg.) 2/2009
After a wave of disapproval in the Russian mass media of a vari
ety of persuasions and public calls to ban him indefinitely from
entering Russia, he offered his interpretation of his words with
the new version sounding like Georgia “should distance itself
from Russia” (Sapozhnikova, 2008). He gave further assurances
that Georgians are still treating Russians as brothers. Russia is
good, its policy is bad would seem a perfect short version of
his opinion.
Another stand was taken by the principal dancer of the Bolshoi
theatre, Nikolai Tsiskaridze, who is Russia’s People’s artist. He
was born in Tbilisi and holds Georgian and Russian artistic ti
tles. He mentioned that while he is, indeed, of Georgian origin,
he did not feel any different from those who were bombed and
killed in South Ossetia. He continued by suggesting that artists
should stay out of politics, and he as artist and Russian artist
of Georgian origin would like to do so. For the same reason he
disapproved the position and involvement of Kikabidze (Tsiska
ridze, 2008).
The Russian actor and citizen with a Georgian family name,
Oleg Bashilashvili, born in 1942, is personally attached to Geor
gia, including South Ossetia, because this is the place where he
spent the early years of his life during the WWII evacuation.
He abstained from giving his opinion on the events because
he doubted the objectivity of the coverage by Russian media.
Nevertheless, he pointed to two issues. First, that Abkhazia and
South Ossetia are de iure Georgian territory. Until the interna
tional community accepts South Ossetia and Abkhazia as inde
pendent states they should be considered as a part of Georgia.
Thus, he continued, “the entrance of our [Russia’s] military is
an occupation”(Basilashvili, 2008). Second, he disapproved of
Russia’s policy of giving Russian citizenship to the applicants
from South Ossetia and Abkhazia (Basilashvili, 2008).
Boris Akunin (a pen name of Grigory Chkhartishvili), a phi
lologist, critic, essayist born in Georgia, was named Russian
Writer of the Year in 2000. He defines Russia as not (yet) being
a democratic country and strongly disapproves of Putin’s au
thoritarianism. He argued that, for most Europeans, the Rus
sian invasion into Georgia beyond the borders of South Ossetia
immediately revived the memory of the events in Budapest and
Prague several decades ago and this might explain the negative
reaction of a number of European countries. With respect to
the interstate developments between Russia and Georgia, he
believed that relations between Russia and Georgia might only
get worse unless the leadership in one country or the other
were to change. Further, he believed that in the aftermath of
the conflict the efforts of intellectuals alone would not suffice
to restore and maintain good relations between Russians and
Georgians. This is a moment, he argued, for the political elites
to assess once again the actions of Russia and to open a public
discussion of the political steps taken by both countries. As far
as the consequences are concerned, he saw that Russia found
itself isolated, was risking being expelled from the G-8, endan
gered the Sochi Olympic games and dashed hopes of becoming
a member of WTO in the near future. Thus, it was “a small mili
The People’s Artist of the Union of Soviet Socialist Republics / Russia, also
sometimes translated as National Artist of the USSR, is an honorary title
granted to citizens of the Soviet Union / Russian Federation.
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Digol, Debate in Russia on Conflict in Georgia
tary victory followed by a big political defeat” (Akunin, 2008).
Even if it were a part of the big American plan, Russia’s inter
vention did not alter it significantly (Akunin, 2008).
Soso Pavliashvili, a famous pop singer, said in interviews that
he felt equally hurt by the sufferings of Georgians and Osse
tians in this senseless war. He views Ossetians as ethnically the
closest to the Georgian nation. He has advocated for Russia to
remain an important player in the Caucasus. Hee advised his
son to study at the Russian military school in Russia and not at
the American one in Georgia. He believed that Georgia was not
a place for the American military. “If Russia flourishes, Georgia
will flourish”, he concluded (Veligzhanina, 2008).
Among the opinions and reactions of yet other famous Geor
gians are that of Georgi Danelia, a Georgian film director, who
condemned the military actions and regretted that he had lived
to see a day of war between Georgia and Russia (Veligzhanina,
2008). Tamara Gverdtsiteli, a variety singer and People’s art
ist of Georgia born in Tbilisi, found these events shocking and
could not understand how such horror could happen. Stanis
lav Sadalskii, an actor and citizen of Georgia, argued that he
is “sure that in this case it is not Georgia but America fighting
against Russia” (Baranets, 2008). The cancelled performance by
Nani Bregvadze, a Georgian People’s artist and famous singer
of Russian romances, can be interpreted as a protest against
Russia. She postponed her jubilee concert, which was to be
performed at the Academic Cappella in St. Petersburg on 26
September 2008 (Bezmenova, 2008). The views of some other
famous Georgians were difficult to understand and interpret, as
for example the opinion of the President of the Russian Acad
emy of Arts, Zurab Tsereteli, a controversial Russian-Georgian
painter, sculptor and architect. Commenting on these events
he said: “Sometimes you need to take a break from love, and
that’s what’s happening now. But tomorrow the romance will
start again, and it will be passionate!” (Walker, 2008) Some re
fused to comment on the situation, for example, the director
of a singer Valerii Meladze said that the artists would feel hurt
to see his name next to any mention of war (Guru Ken Show,
2008).
3.4 Famous Ossetians who made a career in the
USSR or in Russia
Only a few famous Ossetians could be identified, which is no
doubt yet more evidence of how small the Ossetian nation is.
A world famous opera director, Valery Gergiev, Ossetian by na
tionality was full of grief, commenting on the events stressing
that for such a small nation as Ossetian to lose 1,5-2 thousand
people was a big loss (Beroeva, 2008). The “world most famous
Ossetian”, as named by the Russian press (Rozhaeva, 2008),
deeply regretted the almost complete destruction of Tskhin
vali, comparing the ruins of this city to the ruins of Stalingrad
Georgi Danelia originated a series of “sad comedies” as he styles them.
He might be known to a Western reader as the creator of the sculpture “Tear
of Grief” officially donated by the Russian government as a memorial to those
that died in the September 11th terrorist attacks.
Valery Gergiev is a general director and artistic director of the Mariinsky
Theatre, principal conductor of the London Symphony Orchestra and the
Rotterdam Philharmonic Orchestra and principal guest conductor of the Met
ropolitan Opera.
| BEITRÄGE AUS SICHERHEITSPOLITIK
UND FRIEDENSFORSCHUNG
destroyed during WWII (Kireeva, 2008). He strongly believed
that it was a campaign carefully planed in advance and organ
ised by Saakashvili. He promised to carry the message about the
initiators of this tragedy on every possible occasion to Europe,
the USA or elsewhere. In an interview with the BBC Russian
Service, he expressed the gratitude to the Russian army cred
iting their timely action and support with saving the lives of
another 2-3 thousand Ossetians (Beroeva, 2008). Gergiev gave a
concert on the ruins of Tskhinvali. He was accompanied by two
other famous artists, Dmitri Hvorostovsky and Anna Netrebko
(Rozhaeva, 2008).
An actor, Bulat Budaev, believed that this was not a war between
people, but rather a war provoked by Saakashvili: “I do not
know why he is so indebted to the USA” (Veligzhanina, 2008).
Another actor, Soslan Fidarov, mentioned in grief that this war
took away from him his best friend who died in Tskhinvali.
These are his people, he continued, and thus he was suffering
with them. The irony of the life, as he put it, was that in his last
movie about WWII he, an Ossetian, played a heroic Georgian,
while in real life a Georgian took away the life of his best friend
(Veligzhanina, 2008).
4. Opinion polls
In this section I present the results of opinion polls carried out
by two of the most prominent institutions in Russia. The polls
included here were conducted in the period from August to Oc
tober 2008 by the governmental public opinion research centre
VCIOM and the non-governmental organization “Levada Ana
lytical Center” (“Levada-Center”).
4.1 Public Opinion Research Center – VCIOM
In the public opinion polls conducted by VCIOM (WCIOM,
2008a, 2008b, 2008c, 2008d), the majority of Russians ap
proved of the policy of the Russian government and fully sup
ported Russia’s recognition of the independence of South Os
setia and Abkhazia. More than 70 per cent of the respondents
believed that this conflict touched directly on the interests of
Russia; more than 80 per cent of the respondents defined it as a
full scale military conflict or war. Three out of four respondents
(75%) blamed Georgia and the USA for the conflict. Russia’s
involvement in the conflict was supported by more than 80
per cent of respondents; more than 70 per cent believed that
Russia should keep its peacekeepers in South Ossetia. Those op
posing the presence of Russian peacekeepers in South Ossetia
argued against new casualties among the Russian population
and believed that Georgians and Ossetians should resolve the
situation by themselves. Those who supported the continua
tion of Russia’s military presence in the area argued that this
was a way to keep peace, protect the South Ossetian civilian
population and prevent further attacks, chaos and genocide.
Russians believed that only Russia would be able to protect the
civilian population of South Ossetia and Abkhazia (WCIOM,
2008b). Every second inhabitant of Russia became less friend
ly to Georgians believing that there were more divisive than
common features between these two nations (51%) (WCIOM,
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| Digol, Debate in Russia on Conflict in Georgia
2008d). As regards the future of Russia’s relations with Western
countries, Russians are against “cold relations and a hot war”
with the West (WCIOM, 2008c). Although expecting a pull
back in the relations with Western countries in the immedi
ate aftermath of the conflict, Russians saw it as a temporary
phenomenon. Every third respondent saw Germany, the UK,
Japan and the USA as partners of Russia and considered Russia
a European country (WCIOM, 2008a).
4.2 “Levada Analytical Center”
Public opinion surveys carried out by the “Levada-Center”
clearly showed that an ever-increasing share of respondents
(70%-87%) considered that Russia’s leadership did everything
possible to avoid an escalation of the conflict and bloodshed
(Levada-Center, 2008a, 2008c, 2008e). As seen by the Russian
population, the top three reasons for Russia’s military interven
tion were maintaining peace, protecting the Ossetian popula
tion and defending Russian peacekeepers. The majority of Rus
sians took the side of South Ossetia in this conflict and was in
favour of governmental humanitarian assistance to this region
(81 % of respondents). With respect to military assistance, half
of the respondents were in favour while more that one third
was clearly against (Levada-Center, 2008b).
Notwithstanding this, the overwhelming majority of the popu
lation (80%) justified the recognition by Russia of South Osse
tian and Abkhazian independence. Concerning the future sta
tus of South Ossetia, 34% were in favour of the independence of
South Ossetia from Georgia and 46% for a unification of South
Ossetia with Russia (Levada-Center, 2008c). Equal numbers
of respondents saw it as a timely act (34%) or overdue (30%),
only 12% as a premature act or as wrong (12%) (Levada-Center,
2008d). Among those who disapproved of the act of recogni
tion, 70 per cent believed that it would increase tensions in the
Caucasus, speed up the separatist movements inside Russia and
possibly lead to conflict with third countries. Furthermore,
the majority of Russians believed that Russia should keep its
peacekeepers in the region, encourage the return of Georgian
refugees into South Ossetia and Abkhazia, protect them, and
finance the rebuilding of the urban and rural areas in South
Ossetia (Levada-Center, 2008b).
While the overwhelming majority of Russians (74%) believe
that the Georgian people have fallen hostage to the geopoliti
cal ambitions of the USA, they disagree (52%) that the Ossetian
people have fallen victim to the conflicting ambitions of Russia
and Georgia (Levada-Center, 2008c).
The roots of the conflict were seen in the US policy of expand
ing its influence on Russia’s neighbouring countries (49%) and
in the discriminatory policy towards Ossetians and Abkhazians
by the Georgian government (32%). It was also seen in Saakash
vili opting for military action to resolve the territorial conflict
in order to facilitate the path towards NATO membership and
strengthen his own position so he could remain the country’s
president (81%) (Levada-Center, 2008a).
Russians also believe that Western countries were supporting
Georgia in this conflict because they aim at weakening Russia
118
| S+F (27 . Jg.) 2/2009
and decreasing its role in the Caucasus to the minimum (66%).
Nevertheless, almost the same number of respondents (60%)
argued in favour of an international resolution of the con
flict with the participation of the UN and EU (Levada-Center,
2008e).
There was some uncertainty concerning future relations of
Russia with Western countries. Restoring the status quo in re
lations was expected by 48%, while another 35% believed in
a “new edition” of the Cold War and yet other 18% being in
doubt. Every fifth Russian feared Russia’s isolation, every fifth
believed in the possibility of a third World War. The majority,
however, were of the opinion that the tension would ease in the
near future (Levada-Center, 2008d).
Every fifth person believed that the USA was the winner in this
conflict and almost every third person believed that Russia was
the winner. One fourth saw no winners at all (Levada-Center,
2008d).
As the results of these polls demonstrate, Russians have over
whelmingly supported the course of action taken by their gov
ernment, including the recognition of the independence of
South Ossetia and Abkhazia. They have also been concerned
about the potential consequences of the crisis for Russia. Rus
sians strongly believe in a third party involvement in the con
flict. The majority approved of the continuation of Russia’s
involvement in the Caucasus region.
5. Conclusion
The objective of this article was to present the debate that
took place in Russia in the aftermath of the Russian-Georgian
military conflict in August 2008. The article comprises diverse
opinions of people ranging from official politicians to political
analysts to the public responding to opinion polls.
Those who approved the course of the Russian government
were united on several issues. They believed that Georgia was
the initiator of military action and justified Russia’s military
intervention to save the lives of South Ossetians. They also jus
tified the intervention of the Russian army beyond the borders
of South Ossetia on military tactical grounds. They further ap
proved of the recognition by Russia of South Ossetia and Abkhazia in the aftermath of these events, arguing that never
again would Ossetians and Abkhazians be willing to live to
gether with Georgians in the same state. Moreover, they ap
proved of Russia acting similarly to the way the international
community (in particular, the USA and EU) acted in Kosovo.
They suspected the involvement of the USA and they agreed
that this conflict has damaged the image of Russia abroad.
Those who disapproved of Russia’s actions did so based on one
or more points. Some suggested that Georgia responded to
constant military provocations from the side of Russia. Others
accused Russia of double standards comparing this conflict to
the situation in Chechnya, or in imitating the USA, compar
ing this conflict to the situation in Kosovo. Still others blamed
Russia for becoming a party to the conflict long ago, in par
ticular by awarding Russian citizenship to the South Ossetians
and Abkhazians. Yet others were concerned with the negative
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Digol, Debate in Russia on Conflict in Georgia
consequences for Russia in the aftermath of the events: the in
stability of Russia’s borders, political and economical isolation,
damaged image, etc.
In this analysis those who approved of the actions taken by Rus
sia outnumbered those who disapproved of these actions. In
both camps, there were people of Russian and Georgian origins.
No Ossetian or Abkhazian was identified who condemned the
actions taken by Russia in South Ossetia. Those who approved
mainly focused on the steps taken immediately after the start
of the military campaign: intervention in South Ossetia and be
yond its borders, recognition of the statehood of South Ossetia
and Abkhazia. Those who disapproved based their arguments
on the potential future negative consequences for Russia: yet
more casualties among Russians, Russia’s international image,
political isolation and economic consequences.
It is possible – even probable – that not all the variations of the
opinions of diverse actors were presented and certainly not all
those actors directly or indirectly involved were given a voice in
this article. I believe, nevertheless, that mainstream opinions
of various persuasions were presented here and might serve as
a point of reference for an informed and interested reader.
| BEITRÄGE AUS SICHERHEITSPOLITIK
UND FRIEDENSFORSCHUNG
Kasparov, G. (2008b, 10 September). Self-sufficiency of econo
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Levada-Center. (2008b). 12.08.2008. Russia –Georgia conflict:
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Medvedev, D. (2008b, 26 August 2008). Interview with Al
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Vladov, P. (2008, 14 August). Protests in Moscow at the embas
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|
https://openalex.org/W3022059855
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https://scindeks-clanci.ceon.rs/data/pdf/1451-2092/2020/1451-20922001075I.pdf
|
Kirghiz, Kyrgyz
| null |
Assessment of negative influence of manganese in welding fumes on welder's health and ways to reduce it
|
FME Transactions
| 2,020
|
cc-by
| 5,508
|
1.
INTRODUCTION component), and the disperse medium is formed by gas
mixture (welding fumes gaseous component). Welding
fumes solid-state component is especially hazardous not
only due to its toxic level, but also due to microscopic
dust particles with low sedimentation rate (no more than
0.08m/s), and are equally distributed in the airspace at
the breathing level in a radius of 5 m, inevitably getting
into the body of the welder or any human nearby [6, 7]. Welding works are known to be dangerous for human
health as far as a certain amount of welding fumes is
emitted into the working area alongside with noise and
light irradiation. Apart from gases contained in the
welding fumes, a great amount of welding dust resulted
from oxidation of the evaporated metal as well as
shielding gas, alloyed metals and flux reach environ-
ment. Chemical composition of the solid part in the
welding fumes primarily depends on the type of filling
materials. When the welding arc contacts the metal,
evaporations emitted into the air above the working area
are gradually oxidized producing fine dust [1-3]. Excessive amount of welding fumes metals in a
body causes negative health conditions [8, 9]. Penetration of nano- and micro-dust particles of
iron, manganese and silicon oxides into a human body
triggers natural protective mechanisms, which in turn
can not only activate macrophages, but also lead to
inflammatory processes and consequently develop a
number of diseases [10-12]. In 2013 American Conference of Governmental and
Industrial Hygienists (ACGIH) reconsidered its recom-
mendations on quantitative composition of welding
fumes and suggested reducing limiting concentrations
[4]. This might become a regulation, and make welding
materials manufactures follow new standards. These
changes are caused by the problem of harmful influence
of welding fumes on people’s health. In many countries,
particularly in the USA, there are lawsuits between
welders who suffer from occupational diseases and
manufactures of welding materials [5]. Particles up to 10 μm and more are retained on the
surface of the bronchial tree mucosa, while small solid
particles (up to 5 μm) penetrate much deeper. While
passing through the alveoli membranes, small-sized
particles can get into blood which is of the utmost
danger [13-16]. When passing from one medium to
another, heavy metals can change their chemical
structure and cause deficiency of essential elements
replacing them in metal-containing protein structures. D.A. Chinakhov D.A. Chinakhov
Director,
Yurga Institute of Technology National
Research Tomsk Polytechnic University
Russia The article presents the results of a theoretical study (literature review of
publications), which allowed to establish the negative impact of welding
aerosol (manganese and other elements) on the human body: you can use
special welder protection equipment (ventilation and individual welder
protection); reduce the quantitative and qualitative content of manganese
welding aerosols (welding technology, power sources, modern welding
materials); reduce the content of manganese in the human body, removing
it with medicines. E.D. Chinakhova
Researcher,
Medical Faculty,
Department of Anatomy,
Siberian State Medical University
Russia E.D. Chinakhova
Researcher,
Medical Faculty,
Department of Anatomy,
Siberian State Medical University
Russia Experimental studies have shown that the use of an inverter power source,
compared with a diode rectifier, contributes: to ensuring the drop-by-drop
transfer of electrode metal to reduce the time of their formation by 46%
and the transition by 28%; ensures the transition of alloying elements from
welding materials to the weld metal by 6% and reduces its losses from the
fusion line by 6% and HAZ by 3%; to reduce the intensity of education (g /
min) SA and their components by 23%; to reduce the specific allocation of
CA and their components by 23%. K.Yu. Kirichenko
Researcher,
SEC Nanotechnology FEFU
School of Engineering,
Far Eastern Federal University
Russia E.V. Verkhoturova
Researcher,
Research Department,
Irkutsk National Research
Technical University
Russia Keywords: welding, welding fumes, welder’s health, welding process
optimization, inverter power supplies. © Faculty of Mechanical Engineering, Belgrade. All rights reserved
Received: May 2019, Accepted: July 2019
Correspondence to: Dmitry P. Il’yaschenko, Yurga
Institute of Technology National Research Tomsk
Polytechnic University, Russia,
E-mail: mita8@rambler.ru
doi doi:10.5937/fmet2001075I D.P. Il’yaschenko
Associate Professor,
Industrial Technology Department,
Yurga Institute of Technology National
Research Tomsk Polytechnic University,
Russia
D.A. Chinakhov
Director,
Yurga Institute of Technology National
Research Tomsk Polytechnic University
Russia
E.D. Chinakhova
Researcher,
Medical Faculty,
Department of Anatomy,
Siberian State Medical University
Russia
K.Yu. Kirichenko
Researcher,
SEC Nanotechnology FEFU
School of Engineering,
Far Eastern Federal University
Russia
E.V. Verkhoturova
Researcher,
Research Department,
Irkutsk National Research
Technical University
Russia D.P. Il’yaschenko Assessment of Negative Influence of
Manganese in Welding Fumes on
Welder’s Health and Ways to Reduce it Associate Professor,
Industrial Technology Department,
Yurga Institute of Technology National
Research Tomsk Polytechnic University,
Russia Keywords: welding, welding fumes, welder’s health, welding process
optimization, inverter power supplies. Keywords: welding, welding fumes, welder’s health, welding process
optimization, inverter power supplies. 1.
INTRODUCTION It
increases glutomate content in blood and the content of γ-
aminobutyric acid which is the main inhibitory mediator of
the brain decreases accordingly [49, 50]. In addition, another brake mediator – dopamine falls
under the excessive influence of manganese. Accele-
rating dopamine oxidation (while reducing its con-
centration), manganese stimulates accumulation of a ne-
urotoxic compound which destruct a specific type of
receptor on the surface of nerve cells [23, 49, 51-53]. Manganese is essential for synthesis and exchange of
neurotransmitters [23, 29]: it increases excitability of
adrenoreactive systems and sensitivity of chemore-
ceptors, enhances protective inhibition in the cortex of the
cerebral hemispheres. Manganese is also a cofactor for
some enzymes (hydrolases, transferase, lyases, glutamine
synthetases, etc.). In oxidation states (II) or (III),
manganese enters the active site of a types of superoxide
dismutase and catalase, which are the enzymes necessary
for neutralization of a active oxygen intermediate. Researches by a number of authors [19, 54]
demonstrate that divalent metallic manganese can alter
stability of prion proteins suggesting that stabilization of
prion protein caused by manganese can play a role in
the pathogenesis of prion diseases. Manganese stimulates synthesis of cholesterol, fatty
acids, vitamin C and insulin. It is believed [27, 30] that
manganese competes with iron when interacting with
proteins and enzymes that include iron in the active site
(for example, mitochondrial complex I and aconitase)
and if in excess it replaces magnesium in cellular
enzymatic reactions. Thus, [30, 55, 56], bioconcentrarion of manganese
can lead to a wide range of occupational diseases in
latent and active forms due to disturbance in oxidation-
antioxidant and metabolic processes, development of
central nervous system specific disorders and other
negative influences on workers' health. Based on a literature review we found out that the
following can reduce negative influence of manganese
and other elements on human body: There is evidence [15, 19, 31] that nano compounds
of elements oxides containing manganese found in
welders’ lungs induce development of certain types of
fibrosis as well as interstitial inflammation. This
explains the potential risk to respiratory health caused
by repeated exposure to welding fumes. 1. Using special means of protection of respira-
tory system of the welder (ventilation and personal
protective equipment). 2. Reducing an amount of manganese and its
compounds in welding fumes (welding technology, po-
wer sources, modern welding materials). 1.
INTRODUCTION They can not only enter metabolic cycles but also
accumulate in human the body and hair [17-21]. By origin, welding fumes fall into the category of
condensation aerosols. They can be described as a
disperse system in which the disperse phase is formed
by small solid particles (welding fumes solid-state Some metals [22, 23] released during welding are
able to associate with functional groups of proteins (SH,
NH-, NH2-, COO-), change a configuration of the
enzymes active centers thereby violating protein
functions and suppressing proteins activity, which also
causes health disorders. © Faculty of Mechanical Engineering, Belgrade. All rights reserved
Received: May 2019, Accepted: July 2019
Correspondence to: Dmitry P. Il’yaschenko, Yurga
Institute of Technology National Research Tomsk
Polytechnic University, Russia,
E-mail: mita8@rambler.ru
doi doi:10.5937/fmet2001075I Manganese oxides are the most harmful chemicals
[23-26] released during welding (40% of the total dust). FME Transactions (2019) 48, 75-81 75 They can cause organic damage of human nervous
system, disrupt lungs, liver and blood circulatory
system. Silicon compounds inhaled by humans also dis-
rupt body function and cause silicosis. Chromium com-
pounds, although released in smaller quantities, accu-
mulate in the body causing headaches, diseases of dig-
estive system and anemia, while titanium oxide can
cause lung disease. This fact [37, 41, 47-49] can be explained by a high
rate of oxidative metabolism in nervous tissue and less
active antioxidant protection than in other tissues. Manganese easily penetrates to neurons through the
blood-brain barrier inhibiting activity of mitochondrial
enzymes and accumulating in mitochondria. This again
increases lipid peroxidation and causes damage to neu-
ron membrane due to a number of biochemical
processes. An influence of manganese (Mn) [27, 28] is worth
special attention since it accounts for a large proportion
of the entire toxic content of welding fumes solid
component. Manganese is an essential microelement
that greatly contributes to normal body functioning. It
takes part in regulating various types of metabolism in
cells and body´s tissues, blood clotting, growing of
bones and connective tissue. It also regulates function-
ing of reproductive organs and affects spermatogenesis. It is believed that astrocytes regulate the manganese
content of in the brain. Being damaged first, these nervous
tissue cells may lose some of their functions, in particular,
capture of glutamate (which excites amino acid). 1.
INTRODUCTION Manganese in oxidation states (+4, +6, +7) is most
dangerous for humans [23] because it causes oxidative
stress due to oxidation of dopamine and other catec-
holamines. There is evidence that the content of iron ca-
rriers in the body such as transferrin and ferritin incre-
ases significantly with excessive intake of manganese,
while the content of transferrin receptor decreases. This
can be a signal of iron deficiency [23, 32-34]. 3. Reducing bioconcentration of manganese in
human body by removing it with medicines. The research objective is to assess reduction in
qualitative and quantitative content of manganese in
welding fumes of manual metal arc welding (MMA)
with modern inverter power supplies. The research objective is to assess reduction in
qualitative and quantitative content of manganese in
welding fumes of manual metal arc welding (MMA)
with modern inverter power supplies. Changes in iron metabolism can decrease hemoglobin
contents and stimulate irritation of the erythrocyte lineage
of the bone marrow, which can lead to some changes in
red blood cells (eg, microcytosis) [32, 35, 36]. RESULTS AND DISCUSSION 3. e results are shown in Table 1 and in Figure 1. ure 1. Oscillograms of droplet transfer current and
tage (3mm in diameter E19.9NbB20 electrodes): - d
e; b – Inverter type
An analysis of the oscillograms in Fig. 1 (Tab
ows that the droplet transfer time is reduced an The results are shown in Table 1 and in Figure 1. Figure 2. Images obtained by micro-X-ray analysis with
INCA (Oxford Instruments)
Figure 3. Micro-X-ray spectral analysis graph: 1 - weld
metal; 2 – weld fusion line; 3 – eat affected zone Figure 2. Images obtained by micro-X-ray analysis with
INCA (Oxford Instruments) Figure 2. Images obtained by micro-X-ray analysis with
INCA (Oxford Instruments) Figure 3. Micro-X-ray spectral analysis graph: 1 - weld
metal; 2 – weld fusion line; 3 – eat affected zone An analysis of the data in Table 2 shows that the
content of alloying elements increases in all areas of the
weld when an inverter is used, which can be explained
by fine-droplet transfer (Table 1). by fine droplet transfer (Table 1). Samples of the welding fumes for hygienic
assessment of electrodes were taken during bead
welding of steel plates with X12CrNiTi18-9 electrodes. Diode and inverter rectifiers were used as welding arc
power supplies. The welding was performed with
constant reverse polarity current in the modes shown in
Table 1. Welding fumes were captured by a special
cover isolating the welding zone. At the same time,
according to technological guidelines (MU 1927-78
1980; MU 4945-88 1990), welding fumes were deposi-
ted on filters made from FPP-15-1.5 tissue to evaluate
the level of fuming and on AFA-HA-18 filters for the
following chemical analysis of the welding fumes
samples. The emitted fume and their components were
weighed. For this purpose, at least five welding fumes
samples were taken. The results of the tests (Tables 4-6)
were subjected to statistical processing [11]. Figure 1. Oscillograms of droplet transfer current and
voltage (3mm in diameter E19.9NbB20 electrodes): - diode
type; b – Inverter type An analysis of the oscillograms in Fig. 1 (Table 1)
shows that the droplet transfer time is reduced and the
number of short circuits increases when using a high
frequency energy conversion power supply (inverter). In
this case the transferred electrode metal droplets are
smaller in size which influences the weld metal
composition [58]. To confirm this, a microradiography
analysis of the composition (Fig. RESULTS AND DISCUSSION 2) of sections 1,2,3
shown in Fig. 3 and Table 2 was carried out. In MMA with E19.9NbB20 coated electrodes,
deoxidation (alloying) of weld metal occurs in a
combined way – through the electrode coating. VOL. 48, No 1, 2020 ▪ 77 2.
EXPERIMENTAL PART Welding stability was assessed by means of statistic
analysis of oscillograms of welding current and voltage
generated by power supplies with different dynamic
characteristics. Manganese [37-40] intensifies the process of cell
membranes lipid peroxidation by generating free radicals
as a result of binding glutathione sulfhydryl groups and
blood (tissues) proteins. It breaks the antioxidant defence
of the body which maintains homeostasis. We analyzed the following power supplies: welding
rectifier of VD-306E type and inverter power supply of
Nebula 315 type. To study their dynamic characteristics
we compared current and voltage oscillograms obtained
with the help of a digital "AKIP-4122 / 1V" storage
oscilloscope, "Pintek Electronics" DP-50 " differential
probe, "current probe PR 1030"; and "OWON_
Oscilloscope_2.0.8.26" software. Subsequently, lipid peroxidation products accumu-
late leading to disruption of functions and integrity of
cell membranes and intracellular metabolism in general. According to many authors [41-46], the main
negative effect of the excess intake of manganese is
central nervous system damage. 76 ▪ VOL. 48, No 1, 2020 FME Transactions FME Transactions Figure 2. Images obtained by micro-X-ray analysis with
INCA (Oxford Instruments)
Figure 3. Micro-X-ray spectral analysis graph: 1 - weld
metal; 2 – weld fusion line; 3 – eat affected zone FME Transactions Power supply
- rectifier
Electrode
Average welding mode
parameters
Number of
short
circuits
Arc gap short
circuit time
ts.c., ms
Short circuit
time
Ts.c., ms
Imax, А
Imin, А
Diode type
12
12±3.76
249±76.3
132.1 ±4.7
54.9 ± 4.75
Inverter type
E19.9Nb
B20
Current 86±2,7А
Voltage 24.5±0,6V
Welding speed
0.27 m/min
24
8.7±3.52
140±45.8
123 ± 1.63
78 ± 1.06 Table1 . Welding modes with TsL-11 electrodes from various power supplies and static data of coated electrod
droplets transfer [57]. 78 ▪ VOL. 48, No 1, 2020
FME Transactions
Diode type
12
12±3.76
249±76.3
132.1 ±4.7
54.9 ± 4.75
Inverter type
E19.9Nb
B20
Current 86±2,7А
Voltage 24.5±0,6V
Welding speed
0.27 m/min
24
8.7±3.52
140±45.8
123 ± 1.63
78 ± 1.06
Table 2. Chemical composition of weld metal of X12CrNiTi18-9 steel produced by E19.9NbB20 electrodes. Power Supply
Element content
Diode rectifier
Inverter
Weight %
Atomic %
Weight %
Atomic %
Weld metal (1)
Si K
0.36
0.71
0.38
0.75
Ti K
1.44
1.45
1.90
1.91
Cr K
18.54
19.66
19.21
20.37
Fe K
67.71
66.85
69.45
68.57
Ni K
10.19
9.57
10.81
10.15
Weld fusion line (2)
Si K
0.50
0.98
0.56
1.10
Ti K
0.60
0.69
0.65
0.75
Cr K
16.45
17.45
17.40
18.42
Fe K
70.52
69.53
71.44
70.77
Ni K
10.87
10.11
11.01
10.20
Heat affected zone (3)
Si K
0.50
0.97
0.53
1.05
Ti K
0.61
0.70
0.76
0.88
Cr K
17.19
18.21
17.41
18.45
Fe K
70.96
70.00
70.70
69.73
Ni K
10.63
9.98
10.71
10.05
Table 3. Emission (g/min) of welding fumes and their components. Electrode type
(diameter, mm)
Power supply -
rectifier
Welding
fumes
CrO3
Cr2O3
Mn
Ni
F
Soluble
F
Insoluble
Diode
0.257
0.012
0.006
0.007
0.006
0.012
0.013
E19.9NbB20 (3)
Inverter
0.2
0.09
0.004
0.005
0.0045
0.01
0.01
Table 4. Specific emissions (g / kg) of welding fumes and their components. Electrode type
(diameter, mm)
Power supply -
rectifier
CA
CrO3
Cr2O3
Mn
Ni
F
Soluble
F
Insoluble
Diode
10.48
0.48
0.26
0.31
0,26
0.50
0.51
E19.9NbB20
Inverter
8.07
0.37
0.20
0.24
020
0.42
0.42
Table 5. Toxic level of welding fumes (TLVf) and air exchange (NHL). Electrode type
(diameter, mm)
Power supply - rectifier
TLVf , mg / m3
NHL, m3 / h
Diode
0.15
1480
E19.9NbB20 (3)
Inverter
0.1
1000 Table 2. Chemical composition of weld metal of X12CrNiTi18-9 steel produced by E19.9NbB20 electrodes. FME Transactions FME Transactions The results (Table 4-5) of the research showed that
inverter rectifiers provide better health conditions for a
MMA welder compared with diode rectifiers as far as
the amount of emitted hazardous chemicals is reduced
(welding fumes from 0.257 to 0.2 (19%), manganese from 0.007 to 0.005 (23%), chromium oxide from 0.012
to 0.09 (24%)) Thus, the risk of toxic poisoning and
inflammation of the airways mucous membrane of
welders and indirect workers is lower. (
g
(
),
g
Table1 . Welding modes with TsL-11 electrodes from various power supplies and static data of coated electrodes metal
droplets transfer [57]. Power supply
- rectifier
Electrode
Average welding mode
parameters
Number of
short
circuits
Arc gap short
circuit time
ts.c., ms
Short circuit
time
Ts.c., ms
Imax, А
Imin, А
Diode type
12
12±3.76
249±76.3
132.1 ±4.7
54.9 ± 4.75
Inverter type
E19.9Nb
B20
Current 86±2,7А
Voltage 24.5±0,6V
Welding speed
0.27 m/min
24
8.7±3.52
140±45.8
123 ± 1.63
78 ± 1.06
Table 2. Chemical composition of weld metal of X12CrNiTi18-9 steel produced by E19.9NbB20 electrodes. Power Supply
Element content
Diode rectifier
Inverter
Weight %
Atomic %
Weight %
Atomic %
Weld metal (1)
Si K
0.36
0.71
0.38
0.75
Ti K
1.44
1.45
1.90
1.91
Cr K
18.54
19.66
19.21
20.37
Fe K
67.71
66.85
69.45
68.57
Ni K
10.19
9.57
10.81
10.15
Weld fusion line (2)
Si K
0.50
0.98
0.56
1.10
Ti K
0.60
0.69
0.65
0.75
Cr K
16.45
17.45
17.40
18.42
Fe K
70.52
69.53
71.44
70.77
Ni K
10.87
10.11
11.01
10.20
Heat affected zone (3)
Si K
0.50
0.97
0.53
1.05
Ti K
0.61
0.70
0.76
0.88
Cr K
17.19
18.21
17.41
18.45
Fe K
70.96
70.00
70.70
69.73
Ni K
10.63
9.98
10.71
10.05 with TsL-11 electrodes from various power supplies and static data of coated electrodes metal Table1 . Welding modes with TsL-11 electrodes from various power supplies and static data of coated electrodes metal
droplets transfer [57]. N
b
f A
h
t
Sh
t i
it lding modes with TsL-11 electrodes from various power supplies and static data of coated electrodes metal
nsfer [57]. Table1 . Welding modes with TsL-11 electrodes from various power supplies and static data of coated electrodes metal
droplets transfer [57]. REFERENCES [12] Kirichenko, K.Yu: Estimation of ecological harm
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grant. FME Transactions Power Supply
Element content
Diode rectifier
Inverter
Weight %
Atomic %
Weight %
Atomic %
Weld metal (1)
Si K
0.36
0.71
0.38
0.75
Ti K
1.44
1.45
1.90
1.91
Cr K
18.54
19.66
19.21
20.37
Fe K
67.71
66.85
69.45
68.57
Ni K
10.19
9.57
10.81
10.15
Weld fusion line (2)
Si K
0.50
0.98
0.56
1.10
Ti K
0.60
0.69
0.65
0.75
Cr K
16.45
17.45
17.40
18.42
Fe K
70.52
69.53
71.44
70.77
Ni K
10.87
10.11
11.01
10.20
Heat affected zone (3)
Si K
0.50
0.97
0.53
1.05
Ti K
0.61
0.70
0.76
0.88
Cr K
17.19
18.21
17.41
18.45
Fe K
70.96
70.00
70.70
69.73
Ni K
10.63
9.98
10.71
10.05 ble 2. Chemical composition of weld metal of X12CrNiTi18-9 steel produced by E19.9NbB20 electrodes. Table 3. Emission (g/min) of welding fumes and their components. Electrode type
(diameter, mm)
Power supply -
rectifier
Welding
fumes
CrO3
Cr2O3
Mn
Ni
F
Soluble
F
Insoluble
Diode
0.257
0.012
0.006
0.007
0.006
0.012
0.013
E19.9NbB20 (3)
Inverter
0.2
0.09
0.004
0.005
0.0045
0.01
0.01 Table 3. Emission (g/min) of welding fumes and their components. Table 4. Specific emissions (g / kg) of welding fumes and their components. Table 5. Toxic level of welding fumes (TLVf) and air exchange (NHL). Electrode type
(diameter, mm)
Power supply - rectifier
TLVf , mg / m3
NHL, m3 / h
Diode
0.15
1480
E19.9NbB20 (3)
Inverter
0.1
1000 Table 5. Toxic level of welding fumes (TLVf) and air exchange (NHL). Electrode type
(diameter, mm)
Power supply - rectifier
TLVf , mg / m3
NHL, m3 / h Table 5. Toxic level of welding fumes (TLVf) and air exchange (NHL). 78 ▪ VOL. 48, No 1, 2020 FME Transactions FME Transactions 4.
CONCLUSION [8] Filova, V.A. et al.: Hazardous substances in the en-
vironment. Elements of V-VH groups of the perio-
dic system and their inorganic compounds: ref. ed. /Ed.St.Petersburg: NGO "Professional",2007;452 p. The research showed that inverter power supplies
compared with diode rectifiers: 1. ensure electrode metal fine-droplet transfer, reduce
time of droplet generation by 46% and time of droplet
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ЦЕНТРАЦИЈЕ МАНГАНА У ГАСОВИМА ОД
ЗАВАРИВАЊА НА ЗДРАВЉЕ ЗАВАРИВАЧА И
КАКО ДА СЕ ТАЈ УТИЦАЈ СМАЊИ [53] Erikson K.M., Aschner M. Manganese Neuro-
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E.M., Richt J.A., Kanthasamy A., Kanthasamy
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Proteolysis: Relevance to Role of Metals in Patho-
genesis of Prion Disease. Toxicological Sciences. 2010;115(2):535-546. doi:10.1093/toxsci /kfq049. Рад приказује резултате теоријских истраживања
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и других елемената) на људски организам. Заштита
заваривача се може обезбедити специјалном опре-
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редуковањем квалитативне и квантитативне коли–
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вљача са диодама омогућава пренос метала са елек-
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шава за 46% а пренос метала за 28%, пренос леги-
рајућих елемената на шав за 6% и губитак са линије
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негативног утицаја аеросола од заваривања (мангана
и других елемената) на људски организам. Заштита
заваривача се може обезбедити специјалном опре-
мом (вентилацијом и индивидуалном заштитом),
редуковањем квалитативне и квантитативне коли–
чине мангана у аеросолу (технологијом заваривања,
изворима напајања, савременим материјалима за
заваривање), коришћењем медикамената за редуко-
вање садржаја мангана у организму. [55] Onishchenko G.G., Zaitseva N.V., Zemlyanova
M.A. Hygienic indication of health consequences
for external exposure of chemical factors. Perm:
The book format. 2011;532 p. Експериментална истраживања су показала да
употреба инвертера извора напајања уместо испра-
вљача са диодама омогућава пренос метала са елек-
троде кап-по-кап чиме се редукује време формирања
шава за 46% а пренос метала за 28%, пренос леги-
рајућих елемената на шав за 6% и губитак са линије
спајања за 6%, из зоне утицаја топлоте за 3%,
интензитет обуке и њених компонената се редукује
за 23%, а расподела средстава такође за 23%. [56] Dobson A., Erikson K., Aschner M. Manganese
Neurotoxicity. Annals of the New York Academy
of Science. 2004;1012:115-128. [57] Il'Yaschenko D.P., Chinakhov D.A., Mamadaliev
R.A. Effect of Inverter Power Source Characteris-
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English
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Optimisation of exposure parameters using a phantom for thoracic spine radiographs in antero-posterior and lateral views
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Radiography
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C. Sa dos Reis a, *, M. Caso a, L. Dolenc b, K. Howick c, R. Lemmen d, A. Meira e, F. Shatku a,
E. Aymon f, S.S. Ghotra a, g C. Sa dos Reis a, *, M. Caso a, L. Dolenc b, K. Howick c, R. Lemmen d, A. Meira e, F. Shatku
E. Aymon f, S.S. Ghotra a, g a School of Health Sciences (HESAV), University of Applied Sciences and Arts Western Switzerland (HES-SO), Lausanne, 1011, Switz
b Medical Imaging and Radiotherapy Department, Faculty of Health Sciences, University of Ljubljana, Slovenia
c Radiography and Diagnostic Imaging, School of Medicine, University College Dublin, Dublin, Ireland
d Department of Medical Imaging and Radiation Therapy, Hanze University of Applied Sciences, Groningen, the Netherlands
e Medical Imaging and Radiation Therapy, Lisbon School of Health Technology (ESTeSL)/IPL, Lisbon, Portugal
f Department of Radiology, Hospital of Sion, Avenue Du Grand-Champsec 80, 1950, Sion, Switzerland
g Department of Radiology, Hospital of Yverdon-les-Bains (eHnv), 1400 Yverdon-les-Bains, Switzerland * Corresponding author.
E-mail addresses: claudia.sadosreis@hesav.ch (C. Sa dos Reis), melanie.caso@hesav.ch (M. Caso), dolencl@zf.uni-lj.si (L. Dolenc), kayleigh.howick@ucdconnect.ie
(K. Howick), r.s.lemmen@st.hanze.nl (R. Lemmen), 2020493@alunos.estesl.ipl.pt (A. Meira), floi.shatku@hesav.ch (F. Shatku), emmanuel.aymon@hopitalvs.ch (E. Aymon),
switindersingh.ghotra@hesav.ch (S.S. Ghotra). Contents lists available at ScienceDirect Contents lists available at ScienceDirect a r t i c l e
i n f o Article history:
Received 3 April 2023
Received in revised form
19 June 2023
Accepted 20 June 2023 Introduction: To investigate the exposure parameters for thoracic spine/(TS) radiography that allows the
image acquisition at the lowest dose possible, while maintaining an adequate image quality/(IQ) to
identify all relevant anatomical criteria. Methods: An experimental phantom study was conducted, and 48 different radiographs of TS (24 AP/24
lateral) were acquired. The Automatic Exposure Control/(AEC) with the central sensor was used to select
the beam intensity, while Source-to-Detector-Distance/(SDD) (AP:115/125 cm; Lateral:115/150 cm), tube
potential (AP:70/81/90 kVp; Lateral: 81/90/102 kVp), use of grid/no grid and focal spot (fine/broad) were
manipulated. IQ was assessed by observers with ViewDEX. Effective Dose (ED) was estimated using
PCXMC2.0 software. Descriptive statistics paired with intraclass correlation coefficient (ICC) were applied
to analyse data. Keywords:
Plain radiography
Source-to-detector distance
Beam energy
Beam intensity
Radiographers Keywords:
Plain radiography
Source-to-detector distance
Beam energy
Beam intensity
Radiographers Results: The ED increased with a greater SDD for lateral-view, presenting a significant difference
(p ¼ 0.038), however IQ was not affected. For both AP and lateral, the use of grid had a significant effect
on ED (p < 0.001). Despite the images acquired without grid had lower IQ scores, the observers
considered the IQ adequate for clinical use. A 20% reduction in ED (0.042mSve0.033 mSv) was observed
when increasing the beam energy from 70 to 90 kVp for AP grid in. The observers ICC ranged from
moderate to good (0.5e0.75) in lateral and good to excellent (0.75e0.9) for AP views. Conclusions: The optimised parameters in this context were 115 cm SDD, 90 kVp with grid for the best IQ
and lowest ED. Further studies in clinical setting are necessary to enlarge the context and cover different
body habitus and equipment. Implications for practice: The SDD impacts on dose for TS; Higher kVp and grid are necessary to better
image quality. © 2023 The Author(s). Published by Elsevier Ltd on behalf of The College of Radiographers. This is an
open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). Radiography 29 (2023) 870e877 Radiography 29 (2023) 870e877 https://doi.org/10.1016/j.radi.2023.06.009
1078-8174/© 2023 The Author(s). Published by Elsevier Ltd on behalf of The College of Radiographers. This is an open access article under the CC BY license (http://creativeco
mmons.org/licenses/by/4.0/). * Corresponding author. Introduction the
clinical
situation,
available
resources
and
treatment
options.2 Computed tomography (CT) and magnetic resonance
imaging (MRI) are sensitive in assessing spinal disorders.3 However,
these imaging methods are not always available, MRI is expensive
and requires longer acquisition times,2,4 while CT involves typically
higher radiation dose compared to plain radiography.5,6 Plain The thoracic spine is often affected by trauma, arthropathy,
neoplasms, metabolic diseases and congenital deformities, which
require medical imaging examinations to diagnose and to follow up
these disorders.1 The choice of the imaging modality depends on 9
hed by Elsevier Ltd on behalf of The College of Radiographers. This is an open access article under the CC BY license (http://creativeco C. Sa dos Reis, M. Caso, L. Dolenc et al. Radiography 29 (2023) 870e877 Figure 1. Phantom positioning during image acquisition for antero-posterior projec-
tion at Source-to-Detector Distance of 125 cm. radiography is an advantageous modality for thoracic spine
assessment due its accessibility, low cost and low dose required and
it
is
useful
in
assessing
skeletal
alignment,
trauma
and
arthropathies.2,7 It is important to highlight that patients who re-
quires thoracic spine radiography due to spinal arthropathy sus-
picion are typically older (60e70 years old),7 while cases of spinal
trauma usually affect the younger populations (20e50 years-
old),8,9 which is a concern because the younger population is more
radiosensitive. Therefore, it is critical to optimise the image
acquisition to ensure that patients receive the lowest possible ra-
diation dose, while the relevant anatomical structures are still
visible to detect the abnormalities.10 Additionally, the thoracic
spine is located near by the breast tissue, and this is one of the most
radiosensitive anatomical structures, particularly in the female
population. Furthermore, the images must be of adequate quality to
allow the detection of fracture lines, particularly in the older pop-
ulation with less dense bones and less tissue contrast.11 To achieve
this balance between dose and image quality, patient positioning,
beam energy and intensity, focal spot, use of a grid, collimation,
source-to-detector distance (SDD) and additional filtration must be
carefully selected and combined.12 The objective of this study was to identify the ideal exposure
parameters (SDD, focal spot, use of grid, and beam energy) for
thoracic spine radiography, while optimising the dose and the IQ
balance. Effective dose estimation The effective dose (ED) was estimated using the PCXMC 2.0
software, a Monte Carlo based program, and the tissue weighting
factors were based on the International Commission on Radio-
logical Protection (ICRP) 103.15e18 The estimation was performed
considering the different parameters that were manipulated
during the acquisition as explained in the above section. These
include beam energy (kVp), average beam intensity (average
mAs), focal-skin distance, beam size (collimation field), beam
placement and inherent filtration (1.5 mm Al þ 1 mm in the
collimator), different positioning (AP and Lateral), SDD and
“adult” setting.15 All these variables were established using
mathematical probabilities of the interactions between photons
and the phantom's body, such as the photoelectric effect, coherent
scattering and incoherent scattering.17 For the simulations, a
maximum energy of 150 keV and 20,000 photons were used. Methodology For this research, an experimental phantom study was con-
ducted. The research was performed in 4 phases: 1. Image acqui-
sition, 2. Effective dose estimation, 3. Perceptual image quality
assessment and 4. Data analysis. Figure 1. Phantom positioning during image acquisition for antero-posterior projec-
tion at Source-to-Detector Distance of 125 cm. Ethical approval was not required by the University of Ljubljana
to carry out this study as it was conducted with a phantom in a
controlled environment and not in a clinical setting with real pa-
tients. For the third phase (perceptual image assessment), all par-
ticipants signed an informed consent form prior to taking part in
the assessment. All responses were anonymous, and submissions
were in compliance with the General Data Protection Regulation
(GDPR). to observe the consistency of the system. The central sensor of the
Automatic Exposure Control (AEC) was used for the acquisition of all
radiographs (Table 1; Fig.1). The exposure parameters were selected
according to the published literature,13e15 namely for the selection
of beam energy range (see Fig. 2). Image acquisition An anthropomorphic phantom PBU 60 (Kyotokagaku Co., Ltd,
Japan), which represents a human of 160 cm in height and 50 kg in
weight, was used to produce anteroposterior (AP) and lateral
thoracic spine radiographs (Fig. 1). The images were acquired on a
Siemens Multix/Vertix (Siemens, Germany) unit. Prior to the study,
quality control (QC) tests of tube voltage accuracy, reproducibility,
half-value layer, current time product linearity, tube output, total
filtration and the dose area-product (DAP) were performed. In the
study, a fine focal spot of 0.6 mm and a broad focal spot of 1.0 mm
were used. Total filtration of the beam was 2.5 mm of Aluminium. Images were obtained using two different grids, one focused for SDD
115 cm, with a ratio of 12:1, 40 lines per centimetre and one focused
for SDD 150 cm with a ratio 13:1 and 70 lines per centimetre. The
digital detector was composed by Amorphous Silicon with a thin-
film transistor (TFT) array and a pixel matrix of 2560 3072 with
pitch of 140 mm. A constant collimation for the AP projection
(43.0 13.9 cm) was applied. Due to the diverging X-ray beam, the
collimation had to be adjusted for the lateral projection, using
17.1 43cm with 115 cm SDD and 16.8 41.7 cm with 150 cm SDD. By manipulating different parameters (SDD, kVp, focal spot size, use
of grid), 48 different radiographs of thoracic spine (24 AP and 24
lateral) were acquired, and each exposure was repeated three times Table 1
l COM images, this is another reason to choose this program for
is research.19,20 Before the assessment, the program was set up
ith the images and the criteria used to analyse them (Table 2) The
gure 2. Example of radiography used for image quality assessment (acquired in
eral position at 115 cm SDD, broad focus, 81 kVp beam energy, with grid in). Protocols used for image acquisition (positioning, beam energy and intensity, focus,
grid in/out). Protocols used for image acquisition (positioning, beam energy and intensity, focus,
grid in/out). Projection
(AP/lateral)
SDD
(cm)
kVp
Focus
mAs
(AEC)
Grid
AP
115
70
fine
11.7
In
3.13
Out
broad
11.9
In
3.14
Out
81
fine
6.12
In
1.72
Out
broad
6.25
In
1.7
Out
90
fine
4.26
In
1.27
Out
broad
4.34
In
1.24
Out
125
70
fine
14.7
In
3.76
Out
broad
1.5
In
3.78
Out
81
fine
7.65
In
2.03
Out
broad
7.82
In
2.04
Out
90
fine
5.33
In
1.48
Out
broad
5.45
In
1.47
Out
Lateral
115
81
fine
4.14
In
1.57
Out
broad
4.13
In
1.52
Out
90
fine
3.31
In
1.16
Out
broad
3.34
In
1.1
Out
1.24
Out
102
fine
2.17
In
0.87
Out
broad
2.18
In
0.8
Out
150
81
fine
6.27
In
2.56
Out
broad
7.39
In
2.56
Out
90
fine
5.03
In
1.82
Out
broad
4.31
In
1.81
Out
102
fine
3.22
In
1.25
Out
broad
3.25
In
1.23
Out Protocols used for image acquisition (positioning, beam energy and intensity, focus,
grid in/out). Lateral observers decide if the evaluated image was clinically acceptable
(Table 2). Criteria used for VGA are developed based on the
assumption that the capacity to reproduce normal anatomical
structures is correlated with the possibility of clinically detecting a
lesion in radiography, which is normally evaluated with the
Receiver Operating Characteristics (ROC) study.23 However, VGA
represents a much easier and faster method to evaluate IQ. Figure 2. Example of radiography used for image quality assessment (acquired in
lateral position at 115 cm SDD, broad focus, 81 kVp beam energy, with grid in). Three experienced radiographers and three radiography stu-
dents were recruited as observers for the image assessment. These
observers were chosen instead of other medical professionals as in
a clinical setting, the radiographer is the professional responsible
for deciding if an image is diagnostically acceptable by ensuring the
clinical question posed in the request can be answered. Image quality assessment Perceptual IQ assessment was performed using the software
“Viewer
for
digital
evaluation
of
X-ray
images”
(ViewDEX).19e21 ViewDEX is an image viewer that can be used for
different image evaluation tasks and, in this study, it was used for
Visual Grading Analysis (VGA).19,20 The program was developed to
support studies for protocol's optimisation. ViewDEX supports 871 Radiography 29 (2023) 870e877 C. Sa dos Reis, M. Caso, L. Dolenc et al. Figure 2. Example of radiography used for image quality assessment (acquired in
lateral position at 115 cm SDD, broad focus, 81 kVp beam energy, with grid in). Table 2 energies was calculated using KruskaleWallis in SPSS. IBM SPSS
was used to calculate the IQ assessment score from the ViewDEX
program. The Intraclass Correlation Coefficient (ICC) was calculated
to verify the performance of the observers26 (Table 3). program. Participants were asked to focus on the middle region of
the thoracic spine during the assessment because that was where
IQ was homogeneous to ensured consistency across the observers
with regard to the area of assessment. All participants were
required to sign a consent form to inform them about the study
objective, the rules and asking their permission to use the data for
research purposes. During the assessment, four computers with the
same diagnostic monitor model (DELL P2414H) were used. Both
ambient lighting (10 lux)24 and resolution (1920 1080 pixels)
conditions were kept constant to simulate clinical conditions and to
ensure consistency between observers. Participants were not able
to zoom in or out of the images and they were not able to change
the window level or width to prevent variability in the study. y
p
(
)
Visual Grading Characteristics (VGC) analysis were also per-
formed to compare the answers given by the observers for each
image acquired with a different protocol, to identify which one was
the best regarding quality, being considered a non-parametric
rank-invariant statistical method.28,29 With the VGC Analyser
software,30e33 data was analysed and VGC curves were created. These curves show the comparison of the IQ rating for two different
parameters manipulated. In this study, the curves compared the
images acquired at a SDD of 115 and 125 cm for the AP views and
115 and 150 cm for the lateral views. Additionally, the curves
compare the two distances using Area Under the VGC Curve (AUC). If the AUCVGC 95% Interval Confidence (IC) contains 0.5, it means
that the two distances are showing similar outcomes regarding IQ. When the 95% IC AUCVGC does not include 0.5, then the distances
are statistically different. In this test, an AUC below 0.5 suggests
preference for SDD of 115 cm.34 It was required that all participants sit in front of the monitor
approximately 1 m away.25 The observers assessed 50 images (each
radiograph once and one image repeated three times, to observe
the consistency of the observer). They were informed that they
could take a break when they reached 30 min to help reduce the
influence of fatigue on their evaluation. Table 1
l This is done
by assessing overall image quality and the visibility of all relevant
anatomical structures.15 DICOM images, this is another reason to choose this program for
this research.19,20 Before the assessment, the program was set up
with the images and the criteria used to analyse them (Table 2). The
criteria were based on previous studies15,17,22 and a 4-Likert scale
was selected to score the images, which was adapted to the four
groups of details analysed (anatomy, noise, overall image quality
and contrast). Finally, a close-ended question was used to let DICOM images, this is another reason to choose this program for
this research.19,20 Before the assessment, the program was set up
with the images and the criteria used to analyse them (Table 2). The
criteria were based on previous studies15,17,22 and a 4-Likert scale
was selected to score the images, which was adapted to the four
groups of details analysed (anatomy, noise, overall image quality
and contrast). Finally, a close-ended question was used to let Before starting the task, the observers were asked about their
eye's health and their experience in radiography. They were also 872 C. Sa dos Reis, M. Caso, L. Dolenc et al. Radiography 29 (2023) 870e877 informed and coached about how the assessment would be carried
out, the assessment method and the functionalities of ViewDEX informed and coached about how the assessment would be carried
out, the assessment method and the functionalities of ViewDEX values for the distance, grid in/out and focus were calculated using
the ManneWhitney U test in SPSS. The p-value for the different Table 2
Observer study assessment criteria for thoracic spine with a 4 points Likert-scale. Table 2
Observer study assessment criteria for thoracic spine with a 4 points Likert-scale. Table 2
Observer study assessment criteria for thoracic spine with a 4 points Likert-scale. Anatomical details
1. Superior endplates of thoracic vertebrae are:
1. Not defined
2. Slightly defined
3. Defined
4. Clearly defined
2. Inferior endplates of thoracic vertebrae are:
1. Not defined
2. Slightly defined
3. Defined
4. Clearly defined
3. Spinous process of thoracic vertebrae are:
1. Not defined
2. Slightly defined
3. Defined
4. Clearly defined
4. Pedicles of thoracic vertebrae are:
1. Not defined
2.Slightly defined
3. Defined
4. Clearly defined
5. Intervertebral disc spaces are:
1. Not defined
2. Slightly defined
3. Defined
4. Clearly defined
Noise
6. Results The results chapter will be divided into two different sections -
effective dose and image quality assessment. Table 3
ICC score interpretation applied in this study.27
< 0.5
Poor reliability
0.5e0.75
Moderate reliability
0.75e0.9
Good reliability
>0.9
Excellent reliability Table 2 A pilot was conducted prior
to the real assessment to ensure the software setup worked and the
questions were clear and understandable, while also answering the
objective of the study. The feedback and suggestions to improve the
assessment were integrated. During the real assessment, the ob-
servers were supported by two members of the research team if
necessary. Once the study was completed, the results were saved in
a log file and used for the statistical analysis, respecting the data
security.20 Table 1
l How would you estimate the noise on this radiograph in general? A. Excessive noise
B. Prominent image noise
C. Some image noise
D. Limited perceptual image noise
Overall image quality
7. How would you estimate the image quality in general? a. Very poor
b. Poor
c. Adequate
d. Very good
Contrast
8. How would you estimate the contrast in general? a. Very poor
b. Poor
c. Adequate
d. Very good
Clinical acceptance
9. Would you accept the radiograph? Yes
No Effective dose (ED) Regarding the impact of SDD on ED for AP views, it was observed
that the average dose was 0.023 for a SDD 115 cm and 0.024 mSv for
a SDD 125 cm, which is not statistically significant different
(p ¼ 0.453), while for lateral views the difference was significant
(p ¼ 0.038), with an increased ED from 0.007 (SDD 115 cm) to
0.009 mSv (SDD 150 cm). The AP and Lateral projections anatomical criterion analysis by
the VGC showed that AUCVGC were approximately 0.5 for the AP and
0.51, which means that they are similar and neither SSD overrides
the other. In addition, AUCVGC 95% confidence interval ([0.400856,
0.546690] for the AP and [0.449097, 0.589907] for the lateral) of the
two views includes the 0.5 value, which means that they are not
statistically different (Fig. 3). The p-value was estimated as 0.478 for
the AP and 0.910 for the Lateral views which is statistically no
significant. In a similar way, for the IQ criterion of the AP and Lateral
views, the results of the AUCVGC curves were estimated at 0.523704
(AP) and 0.467222 (LAT), near to 0.5. As for the anatomical criteria,
the IQ criteria values are inside the 95% confidence interval
([0.347802, 0.593452] for the AP [0.443890, 0.652503] for the
Lateral). The two results are similar. The p-value corresponds to
0.593 for the AP and 0.337 for the Lateral one, which means that the
result is statistically no significant due to the value being above 0.05
for a 95% confidence interval (Fig. 4). (
)
For the AP projection with SDD 115 cm and grid in, the ED lied
between 0.033 and 0.042 mSv with an average dose of 0.037 mSv. Similarly, the ED for a SDD 125 cm and grid in lied between 0.043
and 0.034 mSv. Without grid at 115 cm SDD, the average ED was
0.010 mSv varying between 0.010 and 0.009 mSv. For AP with SDD
125 cm without a grid, the ED ranged from 0.011 to 0.009 mSv. For
AP grid in and grid out there was a significant difference
(p < 0.001). The grid removal for image acquisition promoted a
decrease of 74% in ED (Fig. 3). For the lateral projection with SDD
115 cm and a grid in, the ED varied between 0.011 and 0.010 mSv;
while for a SDD of 150 cm, the ED ranged between 0.013 and
0.012 mSv. Discussion The objective of this study was to find a set of exposure pa-
rameters (SDD, focus, use of grid and beam energy) for thoracic
spine radiography that allows the acquisition of images with the
lowest dose possible, while keeping an IQ that shows all relevant
information to facilitate an adequate diagnosis and/or follow up. Regarding the impact of focal spot-on ED, for the AP and Lateral
projections,
no
significant
difference
was
observed
when
comparing broad with fine focal spot (p ¼ 0.977 and p ¼ 0.954,
respectively). The results of this study showed that for AP projection, the ED
remained almost the same (increase of 0.001 mSv) when increasing
the SDD to 125 cm, presenting no significant difference (p ¼ 0.453),
which is expectable considering that SDD did not largely varied
(115 cme125 cm). For the Lateral projection, the difference be-
tween the two SDD was larger, 115 and 150 cm, conducting to a
significant difference (p ¼ 0.038) of ED. The ED increased from
0.010 mSv (at 115 cm SDD) to 0.013 mSv (at 150 cm SDD) when the Effective dose (ED) Without a grid at 115 cm SDD and 150 cm SDD, the ED
was constant (0.004 mSv and 0.005 mSv, respectively). The ED
increased with the distance but decreased by 61% when the grid
was removed (Table 4). For the lateral grid in and grid out there was
also a significant difference (p < 0.001). The ICC values ranged from poor (0.151) to excellent ICC (0.906)
(Table 6), with higher values observed to the AP view, while for
Lateral views the ICC was always inferior, being worst for the
criteria 1 and 2 (see Table 7). The observers ranked IQ from 3.490 to 3.504 for AP views ac-
quired with the grid and 2.587 to 2.729 when the grid was
removed. For the Lateral views, the scores were lower and ranged
from 3.462 to 3.569 when the grid was in and between 3.073 and
3.087 when the grid was removed (see Table 7). Increasing the beam energy while keeping the same distance,
reduced the ED. The ED for the AP view, with grid in and SDD
115 cm decreased from 0.042 mSv to 0.033 mSv (20% reduction)
when the beam energy was increased from 70 to 90 kVp; but the ED
difference for the AP view, was not significant (p ¼ 0.145). For the
lateral view, with the SDD 115 cm and grid in, the ED increased from
0.010 to 0.011 mSv when manipulating the kVp. Although, with grid
out, the ED remained constant (0.004 mSv). For SDD 150 cm and
grid in, the ED decreased from 0.013 mSv to 0.012 mSv when the
energy was manipulated from 81 to 102 kVp. For lateral views,
there was not observed a significant difference when beam energy
was changed (p ¼ 0.983). Most of the observers accepted most of the images acquired (40
out of 48 images) for clinical purposes (Fig. 5). Data analysis Descriptive statistics analysis was performed using MS Excel
program for dose estimation. IBM SPSS statistics version 28.0.1.0
was used to calculate the mean and standard deviation. The p- 873 C. Sa dos Reis, M. Caso, L. Dolenc et al. Radiography 29 (2023) 870e877 to 30 years. The participant observers have previously had their
eyes tested, being capable of evaluating the images. Table 4 BF
0.035
0.036
0.035
0.010
0.012
0.011
SF
0.035
0.036
0.036
0.010
0.014
0.012
90
Average
0.033
0.035
0.034
0.011
0.013
0.012
BF
0.033
0.034
0.034
0.011
0.013
0.012
SF
0.034
0.035
0.034
0.011
0.013
0.012
102
Average
e
e
e
0.010
0.012
0.011
BF
e
e
e
0.010
0.012
0.011
SF
e
e
e
0.010
0.013
0.011
Out
Average
OUT
0.010
0.010
0.010
0.004
0.005
0.004
70
Average
0.010
0.011
0.011
e
e
e
BF
0.010
0.011
0.010
e
e
e
SF
0.011
0.011
0.011
e
e
e
81
Average
0.009
0.009
0.009
0.004
0.005
0.004
BF
0.009
0.009
0.009
0.004
0.005
0.004
SF
0.009
0.009
0.009
0.004
0.005
0.004
90
Average
0.009
0.009
0.009
0.004
0.005
0.004
BF
0.010
0.010
0.010
0.004
0.005
0.004
SF
0.009
0.009
0.009
0.004
0.005
0.004
102
Average
e
e
e
0.004
0.005
0.005
BF
e
e
e
0.004
0.005
0.005
SF
e
e
e
0.004
0.005
0.004
Total Average
0.023
0.024
0.023
0.007
0.009
0.008 Table 4 Table 4
Estimated effective dose (mSv) for all parameters. (
)
p
Grid
Energy (kVp)
Focus
Antero-Posterior
Lateral
SDD 115 cm
SDD 125 cm
Average (mSv)
SDD 115 cm
SDD 150 cm
Average (mSv)
In
Average
IN
0.037
0.038
0.037
0.010
0.013
0.012
70
Average
0.042
0.043
0.042
e
e
e
BF
0.041
0.043
0.042
e
e
e
SF
0.042
0.043
0.042
e
e
e
81
Average
0.035
0.036
0.035
0.010
0.013
0.012
BF
0.035
0.036
0.035
0.010
0.012
0.011
SF
0.035
0.036
0.036
0.010
0.014
0.012
90
Average
0.033
0.035
0.034
0.011
0.013
0.012
BF
0.033
0.034
0.034
0.011
0.013
0.012
SF
0.034
0.035
0.034
0.011
0.013
0.012
102
Average
e
e
e
0.010
0.012
0.011
BF
e
e
e
0.010
0.012
0.011
SF
e
e
e
0.010
0.013
0.011
Out
Average
OUT
0.010
0.010
0.010
0.004
0.005
0.004
70
Average
0.010
0.011
0.011
e
e
e
BF
0.010
0.011
0.010
e
e
e
SF
0.011
0.011
0.011
e
e
e
81
Average
0.009
0.009
0.009
0.004
0.005
0.004
BF
0.009
0.009
0.009
0.004
0.005
0.004
SF
0.009
0.009
0.009
0.004
0.005
0.004
90
Average
0.009
0.009
0.009
0.004
0.005
0.004
BF
0.010
0.010
0.010
0.004
0.005
0.004
SF
0.009
0.009
0.009
0.004
0.005
0.004
102
Average
e
e
e
0.004
0.005
0.005
BF
e
e
e
0.004
0.005
0.005
SF
e
e
e
0.004
0.005
0.004
Total Average
0.023
0.024
0.023
0.007
0.009
0.008 Table 5
Observer questionnaire results. Type of observer
How many years of experience in the field of radiography do you have
(since beginning your studies)? (years)
Have you ever had your eyes checked? Radiographers
18
Yes
30
Yes
25
Yes
Radiography Students
2
Yes
1.5
Yes
2
Yes
Figure 4. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD 150 cm) for Lateral projections. Image quality assessment The image assessment was completed by the six observers
(Table 5). The experience of the observers ranged from 1.5 years up Figure 3. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD125cm) for AP projections. 874 Figure 3. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD125cm) for AP projections Figure 3. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD125cm) for AP projections. 874 874 Radiography 29 (2023) 870e877 C. Sa dos Reis, M. Caso, L. Dolenc et al. distance increased. No previous studies were done, considering the
expected considering the inverse square law,12,35 even if the AEC
Table 5
Observer questionnaire results. Type of observer
How many years of experience in the field of radiography do you have
(since beginning your studies)? (years)
Have you ever had your eyes checked? Radiographers
18
Yes
30
Yes
25
Yes
Radiography Students
2
Yes
1.5
Yes
2
Yes
Figure 4. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD 150 cm) for Lateral projections. Table 4
Estimated effective dose (mSv) for all parameters. Image quality assessment Grid
Energy (kVp)
Focus
Antero-Posterior
Lateral
SDD 115 cm
SDD 125 cm
Average (mSv)
SDD 115 cm
SDD 150 cm
Average (mSv)
In
Average
IN
0.037
0.038
0.037
0.010
0.013
0.012
70
Average
0.042
0.043
0.042
e
e
e
BF
0.041
0.043
0.042
e
e
e
SF
0.042
0.043
0.042
e
e
e
81
Average
0.035
0.036
0.035
0.010
0.013
0.012
BF
0.035
0.036
0.035
0.010
0.012
0.011
SF
0.035
0.036
0.036
0.010
0.014
0.012
90
Average
0.033
0.035
0.034
0.011
0.013
0.012
BF
0.033
0.034
0.034
0.011
0.013
0.012
SF
0.034
0.035
0.034
0.011
0.013
0.012
102
Average
e
e
e
0.010
0.012
0.011
BF
e
e
e
0.010
0.012
0.011
SF
e
e
e
0.010
0.013
0.011
Out
Average
OUT
0.010
0.010
0.010
0.004
0.005
0.004
70
Average
0.010
0.011
0.011
e
e
e
BF
0.010
0.011
0.010
e
e
e
SF
0.011
0.011
0.011
e
e
e
81
Average
0.009
0.009
0.009
0.004
0.005
0.004
BF
0.009
0.009
0.009
0.004
0.005
0.004
SF
0.009
0.009
0.009
0.004
0.005
0.004
90
Average
0.009
0.009
0.009
0.004
0.005
0.004
BF
0.010
0.010
0.010
0.004
0.005
0.004
SF
0.009
0.009
0.009
0.004
0.005
0.004
102
Average
e
e
e
0.004
0.005
0.005
BF
e
e
e
0.004
0.005
0.005
SF
e
e
e
0.004
0.005
0.004
Total Average
0.023
0.024
0.023
0.007
0.009
0.008 Table 4
Estimated effective dose (mSv) for all parameters. Table 7 Table 7
Observer study - mean scores for questions 1e8 (anatomy, overall IQ, noise,
contrast). SDD
(cm)
Grid In
Grid Out
IQ score
mean ± STD
ED
(mSv)
IQ score
mean ± STD
ED
(mSv)
AP
115
3.504 ± 0.666
0.037
2.729 ± 0.571
0.010
125
3.490 ± 0.678
0.038
2.587 ± 0.667
0.010
Lateral
115
3.462 ± 0.672
0.010
3.087 ± 0.748
0.005
150
3.569 ± 0.598
0.013
3.073 ± 0.692
0.005 This study faced several limitations. Firstly, the use of an
anthropomorphic phantom restricted the simulation of diverse
body types, such as individuals with higher Body Mass Index or
paediatric patients. The simulation solely focused on the radiation
absorption properties and scatter production of a standard adult
body habitus. Consequently, the impact of these factors on image
quality
could
not
be
accurately
assessed. Furthermore,
the
anthropomorphic phantom did not incorporate any pathological
features and in the majority of contexts, not only the anatomy is
assessed but also the low contrast lesions that can be present on the
images. To address these limitations, it is recommended to include
real patient images featuring different types of lesions (low contrast
details). Additionally, conducting receiver operating characteristic
analysis can help determine the system's capabilities and limita-
tions in detecting major pathologies. The X-ray equipment utilised in the study was not “ceiling
mounted,” which limited the ability to manipulate different SDDs. Only two focused grids (115 cm and 150 cm) were available for use,
preventing the evaluation of the influence of different grid prop-
erties (such as type, grid frequency, interspace materials, and grid
ratio). Another constraint is the observers’ study, as it was con-
ducted in a controlled “reading lab” environment. This setting is not
fully representative of the working conditions in a real radiography
department. Figure 5. Answers provided by the observers when asked if they accept the image for
clinical purposes. GO corresponds to Grid Out and GI to Grid In; AP e Antero-Posterior
and LAT e Lateral Views. IQ. This scatter radiation adds to the patient's dose but does not
contribute to the signal to form the radiographic image. The higher
the energy used, the higher the amount of radiation that scatters. Therefore, it was expected that the use of a grid had a significant
impact on IQ (P < 0.001 for AP and lateral). Table 7 In the AP projection, the
dose decreased by 74% when there was no grid being used, while
for the lateral projection, the decrease was 61%. IQ. This scatter radiation adds to the patient's dose but does not
contribute to the signal to form the radiographic image. The higher
the energy used, the higher the amount of radiation that scatters. Therefore, it was expected that the use of a grid had a significant
impact on IQ (P < 0.001 for AP and lateral). In the AP projection, the
dose decreased by 74% when there was no grid being used, while
for the lateral projection, the decrease was 61%. Table 5 SDD
(cm)
Grid In
Grid Out
IQ score
mean ± STD
ED
(mSv)
IQ score
mean ± STD
ED
(mSv)
AP
115
3.504 ± 0.666
0.037
2.729 ± 0.571
0.010
125
3.490 ± 0.678
0.038
2.587 ± 0.667
0.010
Lateral
115
3.462 ± 0.672
0.010
3.087 ± 0.748
0.005
150
3.569 ± 0.598
0.013
3.073 ± 0.692
0.005
Table 6
ICC values for each criterion assessed during image quality evaluation.27
Criteria
View
ICC
Confidence interval
ICC Meaning
Question 1
AP view
0.894
0.815
0.946
good to excellent
LAT view
0.466
0.050
0.739
poor to moderate
Question 2
AP view
0.885
0.800
0.942
good to excellent
LAT view
0.407
0.056
0.710
poor to moderate
Question 3
AP view
0.906
0.836
0.952
good to excellent
LAT view
0.742
0.540
0.874
moderate to good
Question 4
AP view
0.872
0.777
0.935
good to excellent
LAT view
0.673
0.417
0.840
poor to good
Question 5
AP view
0.857
0.751
0.928
good to excellent
LAT view
0.504
0.118
0.757
poor to good
Question 6
AP view
0.894
0.815
0.946
good to excellent
LAT view
0.588
0.267
0.799
poor to good
Question 7
AP view
0.846
0.732
0.922
moderate to excellent
LAT view
0.676
0.423
0.841
poor to good
Question 8
AP view
0.682
0.446
0.839
poor to good
LAT view
0.610
0.307
0.809
poor to good
Figure 5. Answers provided by the observers when asked if they accept the image for
clinical purposes. GO corresponds to Grid Out and GI to Grid In; AP e Antero-Posterior
and LAT e Lateral Views. Table 6 The ICC values showed mostly good to excellent reliability (0.75e0.9)
in the AP images, but in the lateral images the reliability of the ob-
servers was mostly poor to moderate (0.5e0.75). This may be caused
by the difficulty in clear visualization of structures on the lateral
images, as they overlap,37 making it difficult for observers to assess
the image quality or the difference reference point used to evaluate
the images.38 Another reason for the moderate reliability among
observers could be the differences in clinical experience and
knowledge amongst the recruited participants in this study.39 The AP and Lateral projection AUCvgc curve values were
0.523704 and 0.467222 respectively, meaning that the images are
similar in quality even if the distance has been changed. These
results are similar to other studies in literature.40 Their study found
that the increased SDD from 100 to 130 cm reduced the radiation
dose significantly with no significant effect on image quality. Tug-
well and co-workers (2014)41 demonstrated in their study of AP
pelvis imaging that there is no significant difference between IQ
when increasing source to image distance (SID) from 110 to 140 cm. Table 7
Observer study - mean scores for questions 1e8 (anatomy, overall IQ, noise,
contrast). SDD
(cm)
Grid In
Grid Out
IQ score
mean ± STD
ED
(mSv)
IQ score
mean ± STD
ED
(mSv)
AP
115
3.504 ± 0.666
0.037
2.729 ± 0.571
0.010
125
3.490 ± 0.678
0.038
2.587 ± 0.667
0.010
Lateral
115
3.462 ± 0.672
0.010
3.087 ± 0.748
0.005
150
3.569 ± 0.598
0.013
3.073 ± 0.692
0.005
Figure 5. Answers provided by the observers when asked if they accept the image for
clinical purposes. GO corresponds to Grid Out and GI to Grid In; AP e Antero-Posterior
and LAT e Lateral Views. Regarding the impact of the grid, the average score of IQ without
the grid was lower (2.73) than the score given for the images with
the grid (3.50). However, the result of question 9 (do you accept the
image for clinical purpose?) showed that in the case of the radio-
graphs without the grid, most of the images (40/48) were accepted
by the observers even if the quality was not as good as the one with
the grid. Table 5 Table 6
ICC values for each criterion assessed during image quality evaluation.27
Criteria
View
ICC
Confidence interval
ICC Meaning
Question 1
AP view
0.894
0.815
0.946
good to excellent
LAT view
0.466
0.050
0.739
poor to moderate
Question 2
AP view
0.885
0.800
0.942
good to excellent
LAT view
0.407
0.056
0.710
poor to moderate
Question 3
AP view
0.906
0.836
0.952
good to excellent
LAT view
0.742
0.540
0.874
moderate to good
Question 4
AP view
0.872
0.777
0.935
good to excellent
LAT view
0.673
0.417
0.840
poor to good
Question 5
AP view
0.857
0.751
0.928
good to excellent
LAT view
0.504
0.118
0.757
poor to good
Question 6
AP view
0.894
0.815
0.946
good to excellent
LAT view
0.588
0.267
0.799
poor to good
Question 7
AP view
0.846
0.732
0.922
moderate to excellent
LAT view
0.676
0.423
0.841
poor to good
Question 8
AP view
0.682
0.446
0.839
poor to good
LAT view
0.610
0.307
0.809
poor to good Table 6
ICC values for each criterion assessed during image quality evaluation.27 Table 6 This result shows that, according to the observers, even if
the image is scored lower, it could still be used for clinical practice.42 Table 7
Observer study - mean scores for questions 1e8 (anatomy, overall IQ, noise,
contrast). SDD
(cm)
Grid In
Grid Out
IQ score
mean ± STD
ED
(mSv)
IQ score
mean ± STD
ED
(mSv)
AP
115
3.504 ± 0.666
0.037
2.729 ± 0.571
0.010
125
3.490 ± 0.678
0.038
2.587 ± 0.667
0.010
Lateral
115
3.462 ± 0.672
0.010
3.087 ± 0.748
0.005
150
3.569 ± 0.598
0.013
3.073 ± 0.692
0.005
Figure 5. Answers provided by the observers when asked if they accept the image for
clinical purposes. GO corresponds to Grid Out and GI to Grid In; AP e Antero-Posterior
and LAT e Lateral Views. Table 7
Observer study - mean scores for questions 1e8 (anatomy, overall IQ, noise,
contrast). Table 5 Table 5
Observer questionnaire results. Type of observer
How many years of experience in the field of radiography do you have
(since beginning your studies)? (years)
Have you ever had your eyes checked? Radiographers
18
Yes
30
Yes
25
Yes
Radiography Students
2
Yes
1.5
Yes
2
Yes Table 5
Observer questionnaire results. Type of observer
How many years of experience in the field of radiography do you have
(since beginning your studies)? (years)
Have you ever had your eyes checked? Radiographers
18
Yes
30
Yes
25
Yes
Radiography Students
2
Yes
1.5
Yes
2
Yes
Figure 4. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD 150 cm) for Lateral projections. Figure 4. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD 150 cm) for Lateral projections. Figure 4. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD 150 cm) for Lateral projections. Figure 4. AUCVGC curves comparing a reference condition (SDD 115 cm) to a test condition (SDD 150 cm) for Lateral projections. DD 115 cm) to a test condition (SDD 150 cm) for Lateral projections distance increased. No previous studies were done, considering the
ideal SDD for thoracic spine radiography, only for lumbar spine
radiography,15 but considering the results for ED, the shorter SDD
(115 cm) seems more adequate for radioprotection than a higher
SDD. The increasing ED is probably due to a small collimation
variation which could explain these results. The increasing ED is not expected considering the inverse square law,12,35 even if the AEC
system must adjust the exposure parameters to keep the same
signal level on the detector.36 Scatter radiation is a known contributor to reducing contrast
which impacts the detection of low contrast lesions such as frac-
tures in elderly populations and a grid should be used to improve 875 C. Sa dos Reis, M. Caso, L. Dolenc et al. Radiography 29 (2023) 870e877 IQ Thi
tt
di ti
dd
t
th
ti
t'
d
b t d
t
Table 7
Observer study - mean scores for questions 1e8 (anatomy, overall IQ, noise,
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15. Conclusion The objective of this study was to investigate the ideal exposure
parameters for thoracic spine radiography for optimal image
quality and dose, with a focus on the SDD. It was shown that dose
increases when the SDD increases, however, image quality does not
degrade when the SDD increased. It can therefore be concluded
that the preferable SDD for thoracic spine radiography is 115 cm as
the image quality is the same and the patient dose is lower than
with an SDD of 125 cm or 150 cm. In this study, when the kVp was increased, while maintaining the
same distance and grid in, there was no significant difference
(p ¼ 0.145 for APand p ¼ 0.983 for Lateral) in the effective dose, which
means that to select the ideal parameters IQ assessment is critical,
being necessary to choose the images with higher contrast to be able
to detect fractures. Concerning the IQ data analysis with VGC analy-
ser, it was shown that there were no significant differences when the
distances were changed in lateral (p ¼ 0.593) and in AP (p ¼ 0.337). Future research should consider analysing real patient data,
including different body habitus mainly obese, different pathol-
ogies, and applying real clinical conditions for a more realistic
assessment. 876 C. Sa dos Reis, M. Caso, L. Dolenc et al. Radiography 29 (2023) 870e877 Acknowledgements 20. B€orjesson S, Håkansson M, Båth M, Kheddache S, Svensson S, Tingberg A, et al. A software tool for increased efficiency in observer performance studies in
radiology. Radiat Prot Dosimetry 2005;114(1e3):45e52. https://doi.org/10. 1093/rpd/nch550. PMID: 15933080. The authors would like to thank the support from all students
and tutors that participated in our study, namely to the observers,
to Prof. Nejs Mekis for his availability during the Optimax summer
school and to the University of Ljubljana for making this summer
school possible. 21. Svensson S, Zachrisson S, Svalkvist A, Ba M, Ha M, Radiology D, et al. ViewDEX:
an efficient and easy-to-use software for observer performance studies. Radiat
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22. Gorham S, Brennan PC. Impact of focal spot size on radiologic image quality: a
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23. Boita J, van Engen RE, Mackenzie A, Tingberg A, Bosmans H, Bolejko A, et al. Validation of a candidate instrument to assess image quality in digital
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109686. https://doi.org/10.1016/j.ejrad.2021.109686. Conflict of interest statement None of the authors has a conflict of interest. g
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109686. https://doi.org/10.1016/j.ejrad.2021.109686. 24. Brennan PC, McEntee M, Evanoff M, Phillips P, O'Connor WT, Manning DJ. Ambient lighting: effect of illumination on soft-copy viewing of radiographs of
the wrist. AJR Am J Roentgenol 2007;188(2):177e80. Claudia Sa dos Reis (on behalf of all the authors). References Lai ZH, Sa dos Reis C, Sun Z. Effective dose and image optimisation of lateral
lumbar spine radiography: a phantom study. Eur Radiol Exp 2020;4(1). 40. Karami V, Zabihzadeh M, Shams N, Gilavand A. Optimization of radiological
protection in pediatric patients undergoing common conventional radiological
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Pediatr 2017;5(4):4771e82. 16. Tapiovaara M, Siiskonen T. PCXMC A Monte Carlo program for calculating patient
doses in medical x-ray examinations [Internet]. 2nded. 2008 Availablefrom:.https://
www.julkari.fi/bitstream/handle/10024/124342/stuk-a231.pdf?sequence¼1. 41. Tugwell J, Everton C, Kingma A, Oomkens DM, Pereira GA, Pimentinha DB,
et al. Increasing source to image distance for AP pelvis imaging e impact
on radiation dose and image quality. Radiography [Internet] 2014;20(4):
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S1078817414000674. 17. Pazanin A, Skrk D, Zalokar N, Mekis N. Proper collimation effect on radiation
dose and image quality in thoracic spine radiography. Nucl Technol Radiat
Protect 2020;35(2):181e8. 18. Valentin J, editor. The 2007 recommendations of the international commission on
radiological protection [internet]; 2007. Available from: http://www.elsevier. com/wps/find/bookdescription.cws_home/713998/
description#descriptionhttp://www.elsevier.com/wps/find/bookdescription. cws_home/714371/description%23description. 42. Moore CS, Wood TJ, Avery G, Balcam S, Needler L, Smith A, et al. Investigating the
use of an antiscatter grid in chest radiography for average adults with a computed
radiography imaging system. Br J Radiol [Internet] 2015;88(1047):20140613. Available from: http://www.birpublications.org/doi/10.1259/bjr.20140613. /
p /
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description#descriptionhttp://www.elsevier.com/wps/find/bookdescription. cws_home/714371/description%23description. 19. Svalkvist A, Svensson S, Håkansson M, Bath M, Mansson LG. ViewDEX : a status
report, vol 169; 2016. p. 38e45 (1). 877
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Sport Courage, Worry and Fear in Relation to Success of Alpine Ski Learning
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Received: 5 August 2018; Accepted: 6 September 2018; Published: 14 September 2018 Abstract: (1) Background: Individuals’ psychological traits can influence not just success in sport but
also the ability to learn new motor skills. We investigated whether sport courage, worry and
fear differ between alpine ski-naive and basic level skiers and how they affect ski learning. (2): A total of 337 students (249 ski-naive and 88 basic level) participated in research consisting
of a four-part questionnaire and structured skiing program. (3) Results: For beginners, lower fear
(r = −0.30, p < 0.01) and higher Self-efficiency (r = 0.28, p < 0.05) and mastery (r = 0.20, p < 0.01)
were associated with better performance; reducing fear and increasing self-efficiency and worry
increased performance. Experienced skiers were better in determination, mastery, and self-efficiency
(all p < 0.05). In case of lower score in worry (r = −0.28, p < 0.01) and higher in self-efficiency
(r = 0.22, p < 0.05) performance was better. Males scored higher in sport courage scale-31 (all
p < 0.05). In particular, self-efficiency was associated with better (r = 0.39, p < 0.01), and higher
fear with poorer performance (r = −0.33, p < 0.01). Moreover, self-efficiency was a predictor of ski
success (p < 0.001). On the other hand, females like ski beginners scored higher in fear (p < 0.001). In females, determination, mastery and self-efficiency had a positive correlation with skiing (r = 0.21,
p < 0.05, r = 0.28, p < 0.01, and r = 0.33, p < 0.01, respectively), while association between Fear
and skiing (r = −0.46, p < 0.01) was negative, and fear (p < 0.001) was inversely related to success. (4): Conclusions: Psychological factors and gender differences need to be considered during learning
phases of alpine skiing. There is a positive association between self-efficiency and performance of
male ski beginners, and negative association between fear and achieved results in basic alpine ski
school in case of female ski beginners. Keywords: sport courage; self-efficiency; worry; fear; skiing performance Article Vjekoslav Cigrovski 1,*, Ivan Radman 1
, Erkut Konter 2, Mateja Oˇci´c 1 and Lana Ruži´c 1
1
Faculty of Kinesiology Horvacanski zavoj, University of Zagreb, Zagreb 10000, Croatia;
ivan.radman@kif.hr (I.R.); mateja.ocic@kif.hr (M.O.); lana.ruzic@kif.hr (L.R.)
2
Sport Science and Technology High School, Dokuz Eylül University, Izmir 35220, Turkey;
erkut.konter@gmail.com Vjekoslav Cigrovski 1,*, Ivan Radman 1
, Erkut Konter 2, Mateja Oˇci´c 1 and Lana Ru
1
Faculty of Kinesiology Horvacanski zavoj, University of Zagreb, Zagreb 10000, Croatia;
ivan.radman@kif.hr (I.R.); mateja.ocic@kif.hr (M.O.); lana.ruzic@kif.hr (L.R.)
2
Sport Science and Technology High School, Dokuz Eylül University, Izmir 35220, Turkey;
erkut.konter@gmail.com *
Correspondence: vjekoslav.cigrovski@kif.hr; Tel.: +38-591-544-2202 sports sports sports Sports 2018, 6, 96; doi:10.3390/sports6030096 sports sports 1. Introduction Courage, self-efficiency, worry, anxiety, and fear have clinical importance and can be associated
with different medical conditions, alter every-day life, but can also greatly influence individual’s
success in sport [1]. The latter has received much attention in sport literature, primarily related
to investigation of factors influencing success on a professional level. The literature suggests fear,
anxiety and worry usually have detrimental influence on performance, while self-efficiency and
courage, historically perceived as virtue, are nowadays treated as skills and instruments that can have
a dramatically positive effect on competitive sport [2–6]. Moreover, the term sport courage relates to
the ability of a person to be competent, determined, assertive, and make sacrifice on voluntary basis
and in challenging circumstances to achieve a sport success/result [7]. www.mdpi.com/journal/sports www.mdpi.com/journal/sports Sports 2018, 6, 96; doi:10.3390/sports6030096 2 of 13 Sports 2018, 6, 96 Sport courage is a dynamic process, influenced by numerous factors such as situations (e.g.,
danger, fear, risk), type of sport, personal traits, previous experience, and type of the task the athlete
is confronted with [8]. Currently, sports psychology focuses on different psychological variables
influencing athletes’ performance, with the intention of maximizing efficiency [6]. Psychological
outcomes have social foundations that are sometimes gender-related, which means that gender can
have an important role in linking psychological factors and performance. According to this view, boys
are taught to be more competitive and encouraged to take risks; also they are more self-confident
and thus conditioned to be efficacious [9]. Teachers and coaches may contribute to mentioned
factors by having different expectations from females and males, and by providing different practice
opportunities, which finally can lead to men being more confident in their sport abilities and prone to
participate in activities promoting development of motor skills associated with sports. Moreover, boys
and men can be more encouraged to take part in adventurous and extreme sports [2,10,11]. From recreational aspect, alpine skiing is one of the most popular winter sports with millions
of people participating world-wide [12,13]. Contemporary research literature is mainly related to
investigation of physical and physiological characteristics of elite alpine skiers, while research in the
field of psychology is scarce, especially in case of recreational alpine skiing [14]. In the present study
we examined the influence of fear, worry, self-efficiency and courage on learning alpine skiing in
students of kinesiology. 1. Introduction It is a popular sport in Croatia, but due to specific environment and weather
conditions it is generally considered demanding and challenging [15,16]. Therefore, fear related to new
activity and worry about the mentioned exogenous factors can become a psychological barrier that
prevents successful learning. Furthermore, in case of inexperienced learners, the mentioned factors
can reduce motivation to learn and lead to withdrawal from the activity at an early stage. Finally, fear
of injury may cause a behavioral block with refusal to perform specific elements of a ski technique. Therefore, it is important to address different variables such as fear, anxiety, motivation and courage
when studying factors influencing sport success and performance [6]. We hypothesized that self-efficiency and courage would help students to better acquire new
knowledge of alpine skiing. In order to test the psychological factors it is important to have reliable
testing possibilities. We therefore used questionnaires with previously quantified metric characteristics
in different populations and different countries and adopted them to the Croatian language. For this
research, we also specifically developed an instrument Alpine skiing fear inventory [17–19]. We tested
the metric characteristics of used instruments on young physically capable people, Croatian students
of kinesiology with no or only basic previous knowledge of alpine skiing. 2.1. Design and Participants The present study included a total of 340 participants (252 alpine ski naive students and
88 students with existing basic knowledge of skiing). They were all third-year students of Faculty of
Kinesiology at the University of Zagreb; 106 (31.2%) were females and 234 (68.8%) males (mean age
21.98 ± 1.41 years). We collected data during three consecutive academic years. An absence of any
experience in alpine skiing was a definition for an alpine ski naive person. Prior to being enrolled
in the study, all participants were informed about the study protocol and aim and gave their written
consent to participation. The study was approved by the Ethics Committee of Faculty of Kinesiology. 2.2. Variables Alpine ski knowledge was assessed through demonstration of eight previously selected elements
of alpine ski technique. Upon completion of structured alpine ski school program, the following
elements were graded: traversing left (TL), traversing right (TR), uphill turn to the left (UTL),
uphill turn to the right (UTR), snow-plough turn (SPT), basic turn (BT), parallel turn (PT), and short
turn (ST). Grades ranged from 1 to 5; where 1 meant an unacceptable level of knowledge and 5 was Sports 2018, 6, 96 3 of 13 given for a superb demonstration of an element of the ski technique. Each participant received eight
grades and this represented overall knowledge of alpine skiing. Cronbach alpha for skiing performance
was 0.91. Moreover, each eligible participant fulfilled a four-part questionnaire consisting of 65 items. The
questionnaire tested the way courage, worry, self-efficiency and fear influence the success in alpine ski
learning. Each item was rated on a Likert scale. The questionnaire was not anonymous due to the need
for comparison of its results with acquired knowledge of alpine skiing. The questionnaire consisted of
the following parts: (1)
Sports Courage Scale (SCS-31) [19]. SCS-31 is a 31-item self-report instrument with 5 subscales
titled “Determination, Assertiveness, Mastery, Venturesome and Sacrificial Behavior”. Each item
could be answered on a 5-point Likert based scale graded by 1 (I strongly disagree) to 5 (I strongly
agree). SCS-31 scale showed adequate validity and reliability in previous research [19]. (2)
Penn State Worry Questionnaire (PSWQ) [17]. PSWQ is a 16-item instrument measuring
participants’ worry in particular situation. Each item is rated on a four-point scale, ranging
from 1 (little) to 4 (strong). The questionnaire has proven validity and reliability in previous
research [17]. Cronbach alpha for this research 0.87. (3)
Self-Efficiency in Physical Activity and Alpine Skiing (S-EFPA) [18]. S-EFPA is constructed to
assess participants’ self-efficiency. It consists of ten items. Each item is rated on a four-point scale;
ranging from 1 (not at all) to 4 (strongly). The questionnaire has a satisfactory validity, but there
is a need for more similar studies for its further validation [18]. Cronbach alpha for this research
0.84. (4)
Alpine skiing fear inventory (F & S). F & S is a 9-item instrument used to test fear of alpine skiing. Each item is rated on a 4-point scale ranging from 1 (not at all) to 4 (strongly). Cronbach alpha for
this research 0.85. 2.2. Variables (4)
Alpine skiing fear inventory (F & S). F & S is a 9-item instrument used to test fear of alpine skiing. Each item is rated on a 4-point scale ranging from 1 (not at all) to 4 (strongly). Cronbach alpha for
this research 0.85. Since all questionnaires were originally in English, they were translated into Croatian and then
translated back to English by a bilingual professional. Moreover, this was the first use of SCS-31 in
Croatian population. For this reason, in addition to double translation, we tested the questionnaire on
a large sample of participants with similar characteristics (all young, healthy kinesiology students of
similar age). We also performed CFA to test the latent structure is in line with original model of SCS-31
(more details available under sections statistical methods and results). Sport Courage Scale (SCS-31) questionnaire was used for the first time in Croatian population. In order to further explore the factor loadings, exploratory principal axis factoring analysis with
oblimin rotation was used. Kaiser–Meyer–Olkin measure of adequacy suggested that the data matrix
was suitable for extraction (KMO = 0.926), with Bartlett’s Test of Sphericity Approx. Chi-square =
3900.46; df = 465, p < 0.001. Five factors were extracted, explaining a total of 50.4% of variance of SCS; data presented in
Figure 1. 4 of 13
4 of 13 Sports 2018, 6, 96
S
2018 6 Figure 1. Five factor model of SCS score (sc-mastery; dt-determination; ve-venturesome; as-
assertiveness; aa-sacrificial behavior). The first factor explains the largest proportion of the variance (31.1%), and items loaded ar
l
f
h
b
l
(
“I f
l h
I h
h
h
b
f l
Figure 1. Five factor model of SCS score (sc-mastery; dt-determination; ve-venturesome; as-assertiveness;
aa-sacrificial behavior). The first factor explains the largest proportion of the variance (31.1%), and items loaded are
ostly from the Determination subscale (e.g., “I feel that I have the strength to be successful in difficult Figure 1. Five factor model of SCS score (sc-mastery; dt-determination; ve-venturesome; as-
assertiveness; aa-sacrificial behavior). Figure 1. Five factor model of SCS score (sc-mastery; dt-determination; ve-venturesome; as-assertiveness;
aa-sacrificial behavior). 2.2. Variables The first factor explains the largest proportion of the variance (31.1%), and items loaded are
mostly from the Determination subscale (e.g., “I feel that I have the strength to be successful in
difficult conditions”), but also some items form Assertiveness are loading on this factor (“I like to
take initiative in the face of difficulties”), indicating less than clear factor structure. The first factor explains the largest proportion of the variance (31.1%), and items loaded are
mostly from the Determination subscale (e.g., “I feel that I have the strength to be successful in difficult
conditions”), but also some items form Assertiveness are loading on this factor (“I like to take initiative
in the face of difficulties”), indicating less than clear factor structure. Sports 2018, 6, 96 5 of 13 The second factor explains 6.6% of variance, and it consists of items regarding mastery (e.g.,
“I have limited success because I get frightened”, with reverse coding). The third factor explains additional 4.7% of variance and it should consist with Assertiveness
items (“I have no problems responding to opponent’s sudden attacks”), but it has only 4 items loaded
on this factor. The fourth factor explained additional 4% and contains majority of Venturesome items (e.g.,
“I would take any type of risks to become successful”), but additional items load on this factor (like
Assertiveness “I assert myself even when facing hazards”, Sacrificial Behavior “I compete even if I
have much more to lose than to gain”). The fifth factor also additionally explains 4% of variance, but it is a combination of items from
Assertiveness “I continue to compete without panicking even when faced with a danger”, Venturesome
“Even when facing the possibility of injury, I perform to the best of my ability” and Determination
“I perform to the best of my ability no matter how negative the current conditions”. This unclear factor
structure indicates the need for further validation of this scale in Croatian athletes. Since the sample is limited to athletes involved in skiing, and not various other sports as in the
original validation procedure, and due to possible cultural differences, the original model of latent
structure is used in this research and the subscales were formed based on the original factors extracted
by the authors of the scale. Factors analysis if SPS-31 questionnaire is presented in Table 1. Table 1. Factor analysis of SPS-31 questionnaire. 2.2. Variables Questionnaire
Factor
1
2
3
4
5
scs8
0.720
scs29
0.619
scs17
0.617
scs18
0.552
scs30
0.541
scs25
0.528
−0.223
scs28
0.506
−0.239
scs7
0.374
scs15
scs16
0.694
scs6
0.664
scs21
0.645
scs1
0.523
scs24
−0.319
0.474
scs11
0.399
scs22
−0.343
0.235
−0.263
scs13
−0.254
0.452
−0.212
scs2
0.381
scs3
−0.208
0.283
−0.234
scs27
0.266
scs4
−0.677
scs9
−0.539
0.231
scs14
−0.502
0.259
scs10
−0.493
scs23
0.298
−0.451
scs31
−0.398
scs5
0.306
−0.382
scs19
−0.267
0.555
scs26
0.370
0.385
scs12
0.219
−0.201
0.365
scs20
−0.230
0.351 Table 1. Factor analysis of SPS-31 questionnaire. Questionnaire
Factor
1
2
3
4
5
scs8
0.720
scs29
0.619
scs17
0.617
scs18
0.552
scs30
0.541
scs25
0.528
−0.223
scs28
0.506
−0.239
scs7
0.374
scs15
scs16
0.694
scs6
0.664
scs21
0.645
scs1
0.523
scs24
−0.319
0.474
scs11
0.399
scs22
−0.343
0.235
−0.263
scs13
−0.254
0.452
−0.212
scs2
0.381
scs3
−0.208
0.283
−0.234
scs27
0.266
scs4
−0.677
scs9
−0.539
0.231
scs14
−0.502
0.259
scs10
−0.493
scs23
0.298
−0.451
scs31
−0.398
scs5
0.306
−0.382
scs19
−0.267
0.555
scs26
0.370
0.385
scs12
0.219
−0.201
0.365
scs20
−0.230
0.351 Table 1. Factor analysis of SPS-31 questionnaire. Table 1. Factor analysis of SPS-31 questionnaire. Table 1. Factor analysis of SPS-31 questionnaire. Sports 2018, 6, 96 6 of 13 Confirmatory factor analysis (CFA) conducted on data from the current research to verify the
latent structure shows rather good model fit (24.6% of variance explained, CFI (confirmatory factor
analysis) = 0.965, RMSA (root mean square error for approximation) = 0.06), justifying the use of
originally defined 5 subscales. Both confirmatory and exploratory factor analysis show that one item
(“I can take criticism of my principles or believes”) is not saturated on any of the five factors and
is excluded from the analysis because it decreases overall subscale internal consistency. Therefore,
all except one subscale show very good internal consistency: Cronbach alpha for determination,
assertiveness, mastery, venturesome are 0.82, 0.80, 0.77, 0.75, respectively. Only subscale sacrificial
behavior shows poor Cronbach alpha 0.47 (only 3 items after item 15 is removed). Sacrificial behavior
should be considered and used with caution. Moreover, Cronbach alpha for PSWQ, SEFPA, F & S and
skiing performance were as follows 0.87, 0.84, 0.85 and 0.91. 2.4. Statistical Analysis Data was analyzed using SPSS software (IBM SPSS Statistics for Windows, Version 24.0. IBM
Corp., Armonk, NY, USA). Descriptive statistics was used to describe the basic features of the sample
in this study: proportions for categorical data, and mean+/−standard deviation for normally
distributed continuous variables, or median and interquartile range for variables deviating from
normal distribution. Spearman’s coefficient of correlation was used within subsamples. Respecting
statistical significance of Levene’s homogeneity of variance test, T-test for independent samples was
used to test the significance of differences between two independent groups. Reliability analysis for scales used in the research was done using Cronbach alpha indicator,
while factor structure for SCS-31 was verified using factor analysis: to further explore the factor
loadings, exploratory principal axis factoring analysis with oblimin rotation was used. Additionally,
confirmatory factor analysis (CFA) was conducted on data to verify the latent structure corresponding
to five expected subscales defined by the authors of the SCS-31. Regression analysis was calculated to
find independent predictors of skiing performance. 2.3. Investigational Protocol Research was conducted during three consecutive academic years. Participants were included in
a 10-day structured program of alpine skiing school and then filled in the four-part questionnaire. Alpine ski school program was identical for all participants; conducted in the same ski center on
identical ski slopes. During the ski school, participants were in groups of ten, and learned alpine skiing
6 h each day. Participants rented ski equipment of similar quality, adjusted to specific morphological
characteristics. After completing the program of alpine ski school, participants’ knowledge was graded
by independent judges. Grades were given for the demonstration of previously selected elements of
the alpine ski technique according to the detailed instructions of demonstration layout. Each grade
presented the knowledge of demonstrated element of the ski technique and was given by judges with
many years of experience in evaluation of ski knowledge. Judges’ objectivity and homogeneity were
determined and allowed the use of obtained grades for the assessment of alpine ski knowledge [20]. Each participant received in total eight grades, which was the basis for overall level of acquired alpine
ski knowledge. 3. Results Prior to data analysis, descriptive statistics were calculated for all measures used, together with
indicators of normality of data distribution. Descriptive statistics for used measures is shown in
Table 2. 7 of 13 Sports 2018, 6, 96 Table 2. Mean values and standard deviations, median and interquartile range, and normality of
distribution indicator are shown for each scale and subscale. Questionnaire
M
SD
Min
Max
Median Percentile 25
Percentile 75
P (Kolmogorov-
Smirnov Z)
SCS-31
Determination
34.62
4.76
19
45
35
32
38
<0.001
Mastery
22.77
4.78
10
34
23
19
26
<0.001
Venturesome
13.21
3.06
4
20
13
11
15
<0.001
Assertiveness
26.12
3.71
14
35
26
24
28
<0.001
Sacrificial Behavior
11.50
1.84
5
15
12
10
13
<0.001
PSWQ
45.24
9.69
22
72
44
39
51
<0.001
SEFPA
38.64
5.89
22
50
39
35
43
<0.001
F & S
14.55
5.40
8
39
13
11
16
<0.001
Skiing Performance
29.77
5.17
18
40
30
26
33.50
<0.05
SCS-31-Sports courage scale; PSWQ: Penn State Worry Questionnaire; SEFPA: Self-efficacy in physical activity and
alpine skiing; F & S: Alpine skiing fear inventory; Skiing performance: knowledge of demonstrated element of
ski technique. SCS-31-Sports courage scale; PSWQ: Penn State Worry Questionnaire; SEFPA: Self-efficacy in physical activity and
alpine skiing; F & S: Alpine skiing fear inventory; Skiing performance: knowledge of demonstrated element of
ski technique Although data are not normally distributed (all p < 0.05), due to large sample size and data
measured on an interval scale, parametric statistic T-test for independent samples was used to test
the significance of differences between males and females, and between beginners and skiers. When
Levene’s test of equality of variances is statistically significant, option “variances not assumed” was
used to determine statistically significant differences between given groups. Data are shown in Table 3. Table 3. Differences in skiing performance and psychological variables regarding gender and previous
skiing experience. Table 3. Differences in skiing performance and psychological variables regarding gender and previous
skiing experience. Gender
n
Mean
Std. 3. Results Deviation
p
Determination
male
234
35.20
4.36
0.002 *
female
105
33.31
5.34
Mastery
male
234
23.20
4.52
0.019 *
female
106
21.81
5.22
Venturesome
male
234
13.60
2.91
<0.001 *
female
105
12.34
3.22
Assertiveness
male
234
26.53
3.42
0.005 *
female
106
25.22
4.15
Sacrificial Behavior
male
234
11.63
1.76
0.054
female
106
11.22
1.99
PSWQ
male
230
44.92
9.57
0.378
female
104
45.93
9.94
SEFPA
male
230
39.57
5.53
<0.001 *
female
106
36.60
6.15
F & S
male
232
13.40
4.27
<0.001 *
female
106
17.08
6.64
Skiing Performance
male
238
29.70
5.03
0.690
female
106
29.94
5.49 8 of 13 Sports 2018, 6, 96 Table 3. Cont. Skiing Previously
n
Mean
Std. Deviation
p
Determination
beginners
249
34.31
5.06
0.017 *
skiers
88
35.52
3.66
Mastery
beginners
250
22.41
5.02
0.012 *
skiers
88
23.75
3.97
Venturesome
beginners
250
13.34
3.15
0.217
skiers
87
12.87
2.75
Assertiveness
beginners
250
25.98
3.84
0.192
skiers
88
26.58
3.28
Sacrificial Behavior
beginners
250
11.55
1.90
0.511
skiers
88
11.40
1.66
PSWQ
beginners
248
45.33
9.96
0.791
skiers
85
45.01
8.95
SEFPA
beginners
246
37.68
5.76
<0.001 *
skiers
88
41.23
5.37
F & S
beginners
248
15.36
5.82
<0.001 *
skiers
88
12.33
3.11
Skiing Performance
beginners
250
28.35
4.62
<0.001 *
skiers
89
34.00
4.22
PSWQ—Penn State Worry Questionnaire; SEFPA—Self-efficacy in physical activity and alpine skiing; F & S—Alpine
skiing fear inventory; Skiing performance—knowledge of demonstrated element of ski technique. * p < 0.05. There are statistically significant differences between males and females on several variables. Males score higher on average in Determination, Mastery, Venturesome, Assertiveness and
Self-efficiency (all p < 0.05). Females have higher average scores in fear inventory (p < 0.001). There are
statistically significant differences between beginners and skiers on several variables. Experienced
skiers score higher on average inn determination, mastery, self-efficiency and skiing performance
(all p < 0.05), while beginners score higher in fear inventory (p < 0.001). Since there are statistically
significant differences between males and females and between beginners and skiers, correlations
between variables are calculated separately for those subgroups. Data are shown in Table 4. Spearman’s
coefficient of correlation was used because it is more appropriate when data is not normally distributed
or the relationship between variables is not linear. 3. Results Data in Table 5 show correlations between psychological traits and skiing performance in skiers
and ski novices. Data in Table 5 show correlations between psychological traits and skiing performance in skiers
and ski novices. Table 5. Spearman’s coefficient of correlation between variables for skiers (n = 88) and ski beginners
(n = 249). Skiers (n = 88)
Mastery
Venturesome
Assertiveness
Sacrificial
Behavior
PSWQ
SEFPA
Fear
Skiing
Performance
Determination
0.42 **
0.47 **
0.76 **
0.34 **
−0.17
0.47 **
−0.21 *
0.15
Mastery
0.24 *
0.42 **
0.15
−0.35 **
0.52 **
−0.17
0.05
Venturesome
0.56 **
0.42 **
−0.08
0.26 *
−0.19
0.05
Assertiveness
0.38 **
−0.32 **
0.44 **
−0.19
0.09
Sacrificial
Behavior
−0.08
0.22 *
−0.12
0
PSWQ
−0.32 **
0.28 *
−0.28 **
SEFPA
−0.32 **
0.22 *
F & S
−0.13
Ski Beginners
(n = 249)
Mastery
Venturesome
Assertiveness
Sacrificial
Behavior
PSWQ
SEFPA
Fear
Skiing
Performance
Determination
0.58 **
0.59 **
0.81 **
0.55 **
−0.24 **
0.49 **
−0.46 **
0.103
Mastery
0.40 **
0.59 **
0.28 **
−0.51 **
0.40 **
−0.51 **
0.20 **
Venturesome
0.57 **
0.52 **
−0.12
0.27 **
−0.38 **
0.03
Assertiveness
0.44 **
−0.26 **
0.41 **
−0.43 **
0.07
Sacrificial
Behavior
−0.07
0.26 **
−0.27 **
−0.07
PSWQ
−0.31 **
0.30 **
−0.02
SEFPA
−0.48 **
0.28 **
F & S
−0.30 **
PSWQ: Penn State Worry Questionnaire; SEFPA: Self-efficacy in physical activity and alpine skiing; F & S: Alpine
skiing fear inventory; Skiing performance: knowledge of demonstrated element of ski technique. ** p < 0.01,
* p < 0.05. ble 5. Spearman’s coefficient of correlation between variables for skiers (n = 88) and ski beginners
249) Table 5. Spearman’s coefficient of correlation between variables for skiers (n = 88) and ski beginners
(n = 249). PSWQ: Penn State Worry Questionnaire; SEFPA: Self-efficacy in physical activity and alpine skiing; F & S: Alpine
skiing fear inventory; Skiing performance: knowledge of demonstrated element of ski technique. ** p < 0.01,
* p < 0 05 For skiers, lower score in PSWQ (r = −0.28, p < 0.01) and higher score in Self-efficiency (r = 0.22,
p < 0.05) are weakly associated with better skiing performance. 3. Results In females, Determination and Mastery from SCS-31 have a weak positive correlation with skiing
performance (r = 0.21, p < 0.05 and 0.28, p < 0.01, respectively), and so does Self-efficiency (r = 0.33,
p < 0.01). Moderate association between Fear Inventory and skiing (r = −0.46, p < 0.01) is negative
in direction, suggesting females with higher fear have poorer skiing performance. In males, only
Self-efficiency is associated with better skiing performance (weak correlation, r = 0.39, p < 0.01), and as
in females, higher fear is related to poorer performance (r = 0.33, p < 0.01). Table 4. Spearman’s coefficient of correlation between variables for female sample (n = 105) and for
male sample (n = 235). Table 4. Spearman’s coefficient of correlation between variables for female sample (n = 105) and for
male sample (n = 235). Females (n = 105)
Mastery
Venturesome
Assertiveness
Sacrificial
Behavior
PSWQ
SEFPA
Fear
Skiing
Performance
Determination
0.69 **
0.57 **
0.80 **
0.43 **
−0.29 **
0.56 **
−0.48 **
0.21 *
Mastery
0.44 **
0.66 **
0.29 **
−0.49 **
0.55 **
−0.46 **
0.28 **
Venturesome
0.66 **
0.41 **
−0.16
0.28 **
−0.35 **
0.07
Assertiveness
0.41 **
−0.38 **
0.47 **
−0.44 **
0.13
Sacrificial
Behavior
−0.05
0.18
−0.18
−0.07
PSWQ
−0.34 **
0.24 *
−0.17
SEFPA
−0.46 **
0.33 **
F & S
−0.46 ** 9 of 13 Sports 2018, 6, 96 Table 4. Cont. Table 4. Cont. Males (n = 235)
Mastery
Venturesome
Assertiveness
Sacrificial
Behavior
PSWQ
SEFPA
Fear
Skiing
Performance
Determination
0.45 **
0.50 **
0.80 **
0.52 **
−0.19 **
0.44 **
−0.36 **
0.12
Mastery
0.27 **
0.49 **
0.22 **
−0.46 **
0.34 **
−0.44 **
0.14 *
Venturesome
0.49 **
0.52 **
−0.07
0.18 **
−0.25 **
−0.03
Assertiveness
0.41 **
−0.21 **
0.36 **
−0.31 **
0.09
Sacrificial
Behavior
−0.074
0.23 **
−0.23 **
−0.06
PSWQ
−0.27 **
0.33 **
−0.02
SEFPA
−0.47 **
0.39 **
F & S
−0.33 **
PSWQ—Penn State Worry Questionnaire; SEFPA—Self-efficacy in physical activity and alpine skiing; F & S-Alpine
skiing fear inventory; Skiing performance- knowledge of demonstrated element of ski technique. ** p < 0.01,
* p < 0.05. PSWQ—Penn State Worry Questionnaire; SEFPA—Self-efficacy in physical activity and alpine skiing; F & S-Alpine
skiing fear inventory; Skiing performance- knowledge of demonstrated element of ski technique. ** p < 0.01,
* p < 0.05. 3. Results For beginners, lower score in Fear
(r = −0.30, p < 0.01) and higher score in self-efficiency (r = 0.28, p < 0.05) and in mastery (r = 0.20,
p < 0.01) are weakly associated with better skiing performance. Regression analysis was calculated to see which variables are independent predictors of skiing
performance. Since correlation matrix differs for subgroups according to gender and previous skiing
experiences, series of regressions are done on each subsample. Method stepwise forward was used,
with 0.05 level for entry and 0.10 level of significance for removing the variable, variables entered
in blocks: first block subscales from SCS-31, and the second block FEAR, SEFPA and PSWQ (as
psychological characteristics). Data are presented in Table 6. In both subsamples linear regression analysis shows that entered variables explain significant
proportion of variance of the dependent variable, skiing performance (9% of variance in beginners and
12% in skiers, p < 0.01 and p < 0.05, respectively). For beginners, regression model was analyzed in
4 steps, finally including 3 significant predictors. Independent predictors of skiing performance are:
SEFPA (p = 0.004), PSWQ (p = 0.014), and Fear score (p = 0.043) reducing fear and increasing scores 10 of 13 Sports 2018, 6, 96 on SEFPA and PSWQ increase skiing performance. For skiers, only one predictor has a role in skiing
performance: PSWQ score (p = 0.004), meaning decreasing worry increases skiing performance. Table 6. Regression coefficients for predictors of skiing performance, regarding previous skiing
experience (beginners and skiers) and gender. Table 6. Regression coefficients for predictors of skiing performance, regarding previous skiing
experience (beginners and skiers) and gender. 3. Results Questionnaire
Beginners (n = 244)
Skiers (n = 84)
Standardized
Coefficients (Beta)
t
p
Standardized
Coefficients (Beta)
t
p
Mastery
0.120
1.518
0.130
SEFPA
0.213
2.933
0.004 *
PSWQ
0.160
2.271
0.024 *
−0.313
−2.982
0.004 *
F & S
−0.156
−2.035
0.043 *
R2 = 0.13, R2Adj = 0.12, F = 8.996, p < 0.001
R2 = 0.10, R2Adj = 0.09, F = 8.893, p = 0.004
Males (n = 227)
Females (n = 102)
Standardized
Coefficients (Beta)
t
p
Standardized
Coefficients (Beta)
t
p
Mastery
0.021
0.319
0.750
0.079
0.762
0.448
SEFPA
0.402
6.141
<0.001 *
F & S
−0.396
−3.804
<0.001 *
R2 = 0.17, R2Adj = 0.16, F = 22.663, p < 0.001
R2 = 0.20, R2Adj = 0.18, F = 11.986, p < 0.001
* p < 0.05. In gender related subsamples linear regression analysis shows that entered variables explain
significant proportion of variance of the dependent variable, skiing performance (16% of variance in
males and 18% in females, both p < 0.01). For males, regression model was analyzed in 2 steps, finally
including 2 predictors, but only one being statistically significant for predicting skiing performance
for males: SEFPA (p < 0.001)—increasing self-efficiency increases skiing performance. For females,
regression model was analyzed in 2 steps, finally including 2 predictors, but only one remaining
significant independent predictor of skiing performance for females, and that is Fear score (p < 0.001)
decreasing scores in fear increase skiing performance. 4. Discussion Alpine skiing, although enjoyed by millions of people worldwide, is perceived as a “high-risk”
sport [21], demanding for both learning and teaching [22]. Among different factors that can affect
skiing, personality traits are highly important. Up to date, they were more often investigated in
competitive level skiers, and recently more widely with respect to skiing-related injuries. Research by
Johansson and co-authors (2015) suggests that alpine skiers with previous injuries differ from those
not injured in the trait stress susceptibility, which is higher among the latter and thus perceived as a
protective trait [16]. It seems that psychological traits defined in male skiers correlate with more risky
behavior and therefore higher burden of injury [23]. This broadened the investigation of risk-taking
behaviors and characteristics of participants in winter sports to set educational campaigns and reduce
accident rates [24]. We believe it is an important area to investigate also from the perspective of future
ski instructors, who will set example for their alpine ski school participants on how to learn alpine
skiing more efficiently and safely. Our previous research suggests traits such as self-efficiency and confidence can help in learning
alpine skiing, especially more complex elements of the ski technique [15]. Others have shown how
anxiety diminishes effectiveness of alpine ski instruction [25]. In this research we investigated sport courage defined through determination, mastery,
assertiveness, venturesome, altruistic behavior and self-efficiency as positive traits known to influence
attitude towards sport and sport success on one hand [7] and on the other hand fear and worry as
traits with potential to limit sports performance [6,26]. The study included only kinesiology students, 11 of 13 Sports 2018, 6, 96 with either no previous knowledge or only basic knowledge in alpine skiing. From the aspect of
alpine skiing, to be daring, venturesome, and willing to participate in difficult situations could be an
important trait to differ skiers with better performance. Similarly, from the aspect of alpine skiing,
determination is an important trait because it is a quality that makes one continue in trying, even
though it might be difficult, especially during initial phases of ski learning. Mastery is power, control
and in skiing it might be control over speed for example, which is important for ski beginners as it
can lead to reduction of fear. 4. Discussion Assertiveness is defined as confidence, self-assurance which from the
skiing perspective relates to better success in learning process and is also important for advancement. Self-efficiency is closely related to capability of succeeding in a specific task and is important for ski
beginners as well as competitors. Fear is an unpleasant emotion caused by being aware of danger,
and as skiing is by its nature a sport that tests the boundaries of thrill and fear, it can be under
negative influence of fear. When addressing fear in terms of alpine skiing, it can encompass many
different things and situations; from fear of speed, fear of injury to fear of crowded ski-slopes or
ski-lifts, fear of snow conditions or weather to fear of looking foolish while trying to learn alpine skiing. Worry means being anxious or troubled about actual or potential problems, which again in skiing can
be multifactorial. Understanding and overcoming undesirable traits or in case of positive traits their accentuation
may help in selection of adequate didactic methods and result in more efficient learning [6]. In our
research we found differences in psychological traits between alpine ski naive athletes and those with
previous basic knowledge of alpine skiing. Already experienced skiers scored higher in determination,
mastery, self-efficiency. At the same time beginners scored higher in fear inventory. The mentioned
facts suggest how positive previous experience influences attitude towards new activity and can help in
better learning. Research by Kintschera and coworkers showed that ski beginners worry about falling
and keeping balance during initial phases of alpine ski learning, which is influenced by fear and lower
self-efficiency [27] and may limit the speed and results of ski learning. On the other hand, experienced
skiers with positive psychological traits achieve better results during evaluation of skiing performance. p
p y
g
g
g p
According to published data, one can also expect differences between males and females in
psychological traits linked to sport success [18,27]. Literature suggests males to be more self-efficient
and confident and ready to take more risks, all of which transfers to success and better sport
performance. Gender-related differences are especially notable in traits self-efficiency and fear, which
according to Giulianou and co-workers has its roots in cultural stereotypes where males receive more
attention and support from family, coaches and trainers than their female counterparts, predisposing
them to better sport results and taking part in high-risk sports [28]. 4. Discussion Although our research included young athletes of similar interest in sport and one would not
expect such pronounced differences in psychological traits, we did notice statistically significant
differences between male and female athletes in sport courage related traits. The mentioned facts
were all more pronounced in males, and represent potentially more positive attitude of males towards
sport and skiing. Similarly, Konter describes male sport participants to be more determined, assertive
and venturesome than female counterparts [29]. Moreover, our female participants expressed more
fear than males, which is also noted in other sports [18]. Fear is an important trait which might
limit learning a new motor activity [30]. According to our results, female participants with more
expressed fear had poorer skiing performance than those who were more determined and had greater
scores in mastery, and self-efficiency. Interestingly, in males, only self-efficiency was associated with
better skiing performance, but as in females, higher fear was related to poorer performance. Similar,
detrimental effects of fear on sport success were also described in other sports [31,32]. Regardless of gender, participants with previous ski experience who scored lower in worry
and higher in self-efficiency had better skiing performance. At the same time, ski beginners with
lower score in fear and higher score in self-efficiency and mastery were better at alpine ski learning. Independent predictors of skiing performance for ski beginners are self-efficiency and worry which
when increased relate to better performance and at the same time fear which when reduced also 12 of 13 Sports 2018, 6, 96 helps skiing performance. For skiers, only worry had an influence on skiing performance, meaning
decreasing worry increases skiing performance. For males, significant predictor of skiing performance
is self-efficiency which when increased leads to better skiing performance. For females, significant
independent predictor of skiing performance was fear, suggesting that decreasing fear might improve
skiing performance. Similar results showing positive effects of self-efficiency and confidence on
reduction of fear of injury are shown by different studies and can help in learning a new motor activity
and motivation [33]. Finally, it is important to mention study limitations, primarily related to a sample
consisting of only young motorically capable people. In the future research and for the research to
have a more practical impact on a broader population it would be necessary to include participants of
different age and motor abilities. References 1. Dugas, M.J.; Gosselin, P.; Ladouceur, R. Intolerance of uncertainty and worry: Investigating specificity in a
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skiing. To improve ski learning, teachers and ski instructors would need to help ski beginners increase
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performance, while in female ski beginners it is important to reduce fear to achieve better results
in basics alpine ski school program, as there is a negative association between the two. Mentioned
approach could not only improve but also speed up the learning process. Author Contributions: Conceptualization V.C. and Y.Y.; Methodology, V.C. and E.K.; Software, V.C., L.R. and
I.R.; Validation, V.C., E.K., and L.R.; Formal Analysis, V.C. and I.R.; Investigation, V.C., I.R. and M.O.; Resources,
V.C. and E.K.; Data Curation, V.C. and L.R.; Writing: Original Draft Preparation, V.C., M.O. and I.R.; Writing:
Review and Editing, V.C., E.K. and L.R.; Visualization, V.C.; Supervision, V.C. and L.R.; Project Administration,
M.O. and I.R. Funding: This research received no external funding. Funding: This research received no external funding. Funding: This research received no external funding. Conflicts of Interest: The authors declare no conflict of interest. Conflicts of Interest: The authors declare no conflict of interest. Author Contributions: Conceptualization V.C. and Y.Y.; Methodology, V.C. and E.K.; Software, V.C., L.R. and
I.R.; Validation, V.C., E.K., and L.R.; Formal Analysis, V.C. and I.R.; Investigation, V.C., I.R. and M.O.; Resources,
V.C. and E.K.; Data Curation, V.C. and L.R.; Writing: Original Draft Preparation, V.C., M.O. and I.R.; Writing:
Review and Editing, V.C., E.K. and L.R.; Visualization, V.C.; Supervision, V.C. and L.R.; Project Administration,
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of Zagreb, Faculty of Kinesiology: Zagreb, Croatia, 2008; pp. 513–517. 21. Castanier, C.; Le Scanff, C.; Woodman, T. Who takes risks in high-risk sports? A typological personality
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Lombardo, M.P. Sex Differences in Sports Interest and Motivation:
An Evolutionary Perspective. Evol. Behav. Sci. 2016, 10, 73–97. [CrossRef] 33. Perkos, S.; Theodorakis, Y.; Chroni, Y. Enhancing performance and skill acquisition in novice basketball
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article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/).
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Advancements in Hydrogel-Based Drug Sustained Release Systems for Bone Tissue Engineering
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Frontiers in pharmacology
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cc-by
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Edited by:
Jianxun Ding,
Chinese Academy of Sciences,
China Edited by:
Jianxun Ding,
Chinese Academy of Sciences,
China Bone defects caused by injury, disease, or congenital deformity remain a major health
concern, and efficiently regenerating bone is a prominent clinical demand worldwide. However, bone regeneration is an intricate process that requires concerted participation
of both cells and bioactive factors. Mimicking physiological bone healing procedures, the
sustained release of bioactive molecules plays a vital role in creating an optimal osteogenic
microenvironment and achieving promising bone repair outcomes. The utilization of
biomaterial scaffolds can positively affect the osteogenesis process by integrating cells
with bioactive factors in a proper way. A high water content, tunable physio-mechanical
properties, and diverse synthetic strategies make hydrogels ideal cell carriers and
controlled drug release reservoirs. Herein, we reviewed the current advancements in
hydrogel-based drug sustained release systems that have delivered osteogenesis-
inducing peptides, nucleic acids, and other bioactive molecules in bone tissue
engineering (BTE). Reviewed by:
Juan Wang,
Sichuan University, China
Dan Wang,
Institute of Process Engineering (CAS),
China
Wenjing Yu,
University of Pennsylvania,
United States Reviewed by:
Juan Wang,
Sichuan University, China
Dan Wang,
Institute of Process Engineering (CAS),
China
Wenjing Yu,
University of Pennsylvania,
United States *Correspondence:
Tingting Yu
tingtingyu@bjmu.edu.cn
Yan Wei
kqweiyan@bjmu.edu.cn
Bing Han
kqbinghan@bjmu.edu.cn
†These authors have contributed
equally to this work and share first
authorship *Correspondence:
Tingting Yu
tingtingyu@bjmu.edu.cn
Yan Wei
kqweiyan@bjmu.edu.cn
Bing Han
kqbinghan@bjmu.edu.cn *Correspondence:
Tingting Yu
tingtingyu@bjmu.edu.cn
Yan Wei
kqweiyan@bjmu.edu.cn
Bing Han
kqbinghan@bjmu.edu.cn Advancements in Hydrogel-Based
Drug Sustained Release Systems for
Bone Tissue Engineering 1 Department of Orthodontics, Peking University School and Hospital of Stomatology & National Engineering Laboratory for
Digital and Material Technology of Stomatology & Beijing Key Laboratory of Digital Stomatology, Beijing, China, 2 Department
of Geriatric Dentistry, Peking University School and Hospital of Stomatology, Beijing, China REVIEW REVIEW
published: 06 May 2020
doi: 10.3389/fphar.2020.00622 INTRODUCTION Specialty section:
This article was submitted to
Experimental Pharmacology
and Drug Discovery,
a section of the journal
Frontiers in Pharmacology
Received: 12 March 2020
Accepted: 20 April 2020
Published: 06 May 2020 Specialty section:
This article was submitted to
Experimental Pharmacology
and Drug Discovery,
a section of the journal
Frontiers in Pharmacology Bone defects may be caused by various events, including trauma, inflammation, neoplasm resection,
congenital deformity, and degeneration (Crane et al., 1995; Spicer et al., 2012). Despite numerous
solutions being applied to tackle this issue, clinical demands remain unmet. Bone defects may be caused by various events, including trauma, inflammation, neoplasm resection,
congenital deformity, and degeneration (Crane et al., 1995; Spicer et al., 2012). Despite numerous
solutions being applied to tackle this issue, clinical demands remain unmet. To date, autologous bone grafts are still the gold standard and most considered therapeutic
strategy for critical-sized bone defects among all restoration methods due to their remarkable
osteoconductive and osteoinductive properties. However, de novo problems might arise, such as a
limited amount of donor tissue, an excessive harvest procedure, and the possibility of postoperative
infection of the donor site (Langer and Vacanti, 1993; Betz, 2002; Ahlfeld et al., 2019). Allografts or
xenografts usually serve as secondary alternatives, as slower incorporation, immune rejection, and
pathogen transmission might occur (Crane et al., 1995; Haugen et al., 2019). Utilizing biocompatible
scaffold materials, such as mesenchymal stem cells (MSCs) and/or bioactive factors (Meijer et al.,
2007), bone tissue engineering can offer more possibilities. Achieving sufficient and qualified bone
formation via artificial composites is the grand aim of bone tissue engineering. Keywords: hydrogel, sustained drug release, bone tissue engineering, growth factors, mesenchymal stem cells Citation: Zhang Y, Yu T, Peng L, Sun Q, Wei Y
and Han B (2020) Advancements in
Hydrogel-Based Drug Sustained
Release Systems for Bone
Tissue Engineering. Front. Pharmacol. 11:622. doi: 10.3389/fphar.2020.00622 Compared with bone harvest operations, MSCs are relatively easy to obtain. These cells exhibit
self-renewal, multipotentiality (Prockop, 1997), and immunomodulatory properties (Keating, 2008), May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org Hydrogel-Based Drug Release for BTE Zhang et al. which are imperative for bone regeneration. In addition, bioactive
factors, for example, cytokines and growth factors (GFs), play a
crucial role in new bone formation. Bone morphogenetic proteins
(BMPs) are a group of GFs that have been substantially
investigated. Recombinant human BMP-2 and BMP-7 is
commercially available for limited clinical usage (Nauth et al.,
2011). However, naked GFs are vulnerable in vivo, and to achieve
optimal osteogenic effects, a supraphysiological dose of GFs is
required. Paradoxically, diffusion or uncontrolled release of GFs
may lead to ectopic bone formation and other complications,
including carcinogenicity (Carragee et al., 2011; de Melo Pereira
and Habibovic, 2018). Hence, attaining sustained release of
bioactive factors is an essential objective for scaffold design to
promote the therapeutic efficacy of bone tissue engineering. The
scaffold materials not only create a congenial microenvironment
to promote MSC biological behaviors but also help to maintain
bioactive molecules in situ. To date, the controlled release of
bioactive factors in bone tissue engineering has been realized by a
wide range of biomaterials of different natures and configurations,
which provide diverse release profiles in different treatment
scenarios (Lee and Shin, 2007). it was reported that MSC-derived exosomes combining scaffolds
achieved preferable osteogenesis outcomes (Li et al., 2018),
indicating the promising prospect of exosomes-based cell-free
bone regeneration. MSCs from different sources, such as bone marrow and dental
tissue, are available for bone tissue engineering. The stem cell
niche, 3D microenvironments containing specific biophysical
and biochemical signals, maintains the stemness of stem cells
in vivo (Scadden, 2006; Jones and Wagers, 2008). However,
maintaining the viability and stemness of MSCs as well as
controlling stem cell fate is a fairly critical issue in regenerative
medicine. Substrate-derived stimuli are able to prolong the
stemness of stem cells and guide stem cell fate into specific
lineages (Fisher et al., 2010; Marklein and Burdick, 2010; Lee
et al., 2015). Citation: Moreover, as the proliferation and differentiation of
MSCs may drive into specific lineages depending on different
microenvironmental cues, biochemical stimuli, including
cytokines and GFs, are used in a spatiotemporal sequence
during the complex and continuous reparative procedure
(Samorezov and Alsberg, 2015; Farokhi et al., 2016). Successful
bone regeneration requires the proper combination of stimuli
that can trigger MSC differentiation and matrix deposition. As
the scaffold material itself is capable of combining substrate-
derived and biochemical stimuli, biomimetic and bioinspired
synthetic materials with sustained drug release systems should be
designed to facilitate bone tissue regeneration. Due to the
constraints of current knowledge in this field, the research is
far from sufficient. Hydrogels are a category of highly hydrated 3-dimensional
(3D) crosslinked homopolymer, copolymer, or macromer
networks that can be cast into different shapes and sizes
(Slaughter et al., 2009; Seliktar, 2012). The application of
hydrogels in tissue engineering, bone tissue engineering in
particular, has been garnering increasing attention. Laden with
osteogenic-inducing drugs and sustained release profiles,
hydrogels have been suggested to be promising bone tissue
engineering biomaterials. In this review, we discuss the
progress and limitations of current bone tissue engineering, the
advantages of hydrogel-based bone regeneration biomaterials
and recent advancements in hydrogel-based drug sustained
release systems for bone tissue engineering. Natural bone fracture healing requires the coordinated
participation of osteogenesis and angiogenesis (Collin-Osdoby,
1994; Marsell and Einhorn, 2011). Bioactive factors and signal
pathway crosstalk, which mediates the interplay between
epithelial cells and osteoprogenitors, has been well summarized
(Ramasamy et al., 2016). Likewise, vascularization in bone
substitutes is vital for successful bone tissue engineering. Insufficient blood supply may result in undernutrition,
hypoxia, and inadequate cell recruitment, leading to the failure
of bone tissue engineering. Varieties of assessments and solutions
have been summarized (Rouwkema et al., 2008; Das and
Botchwey, 2011), yet there is no convincing evidence that the
strategies are ample to sustain large tissue constructs,
encouraging the proposal of more promising methods. THE PRESENT CHALLENGES OF BONE
TISSUE ENGINEERING To date, substantial progress has been made in bone regenerative
medicine. A variety of biomimetic polymers and inorganic
materials with bone-like microarchitecture have been designed
with advanced manufacturing methods (Wei et al., 2011; Kim
et al., 2017b; Yin et al., 2019), including 3D printing, aiming to
achieve superb osteogenic properties as well as accuracy and
spatial fitness of critical-sized defects. Light-cured, thermal-
setting, pH- or enzyme-sensitive, and other smart biomaterials
enable bone tissue engineering to serve in many on-demand
circumstances. Varieties of seed cells from different origins
including umbilical cord MSCs (UCMSCs), induced
pluripotent stem cell-derived MSCs (iPSC-MSCs), and
embryonic stem cell-derived MSCs (ESC-MSCs) are
successfully applied (Xie et al., 2016; Chen et al., 2018). Multifarious drugs or bioactive factors are delivered in situ
with different strategies and tailored release profiles, offering
osteogenic-friendly environments for relevant cells. Noteworthy, Frontiers in Pharmacology | www.frontiersin.org BMP With the promoted understanding of the underlying mechanism
of osteogenesis (Chen et al., 2012), BMP, as a prominent member
of the TGF-b superfamily, has always been a favored candidate
for bone tissue engineering applications. Since some hydrogels are believed to possess inferior
osteoconductive properties, Olthof et al. (2018) modified an
oligo[poly(ethylene glycol) (PEG) fumarate] (OPF) hydrogel
with bisphosphate. BMP-2 was encapsulated in poly(lactic-co-
glycolic acid) (PLGA) microspheres. The additional BMP-2 and
drug-laden PLGA microspheres were entrapped in the hydrogel
matrix. The researchers believed that the anionic groups would
produce a strong interaction between the matrix and inorganic
phase of the bone as well as enhance BMP-2-induced bone
formation. The hydrogel matrix could be functionalized by
peptides, which might be beneficial to reduce the dose of
encapsulated BMP. In addition, nanofibrous mesh-hydrogel
hybrid composites have been applied to reach a proper
spatiotemporal release profile (Kolambkar et al., 2011). Shekaran et al. (2014) modified a matrix metalloproteinase
(MMP)-degradable peptide crosslinked PEG with an a2b1
integrin-specific peptide (GFOGER). The interaction between
integrin and collagen I has been proposed to be vital in
osteogenic differentiation and mineralization. It was suggested
that the modified matrix is able to support cell adhesion and
proliferation and upregulate osteogenic gene expression. Laden
with the low dose of BMP-2, robust bone healing was achieved. Along with BMP-2, BMP-7 is considered to be a promising GF in
bone formation. An injectable chitosan/b-glycerophosphate (CS/
b-GP) hydrogel laden with BMP-7 and antibiotic exhibited
preferable reparative effects towards infection-induced bone
loss (Zang et al., 2019). Growth differentiation factor-5 (GDF-5),
also known as BMP-14, regulates the development of numerous
tissue and cell types, including limbs and teeth. Bae et al. (2014)
mixed different concentrations of GDF-5 with a light-cured
hydrogel matrix. The results showed that GDF-5 improved the
osteogenic ability in a dose-dependent manner, as the strongest On the other hand, bioactive molecules play an important
role in bone regenerative medicine. During bone formation,
numerous cytokines and GFs are orchestrated in a
spatiotemporal manner (Farokhi et al., 2016), which would
provide a suitable microenvironment for MSC proliferation
and differentiation, as well as recruit progenitors from
surrounding tissue and peripheral blood for further
restoration. Apart from competent cell carriers, hydrogels can
also be employed as promising local drug reservoirs. Multiple
schemes have been applied to reach desirable and smart drug
delivery kinetics (Lee and Shin, 2007; Slaughter et al., 2009). THE PREPONDERANCE OF HYDROGELS
IN BONE TISSUE ENGINEERING Ideal bone tissue engineering scaffolds should meet the following
criteria: (1) biocompatible, nontoxic and nonimmunogenic; (2)
porous-structured; (3) proper mechanical properties, load-
bearing ability, and sufficient dimensional stability; and (4)
fully degradable, with a degradation rate that matches
neotissue formation (Lee and Shin, 2007; Slaughter et al., 2009;
Haugen et al., 2019). Numerous inorganic scaffolds, such as
metals and bioceramics, have been applied in bone regeneration,
yet their lack of cell affinity, unbalanced mechanical properties, May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org 2 Zhang et al. Hydrogel-Based Drug Release for BTE HYDROGEL-BASED DRUG SUSTAINED
RELEASE SYSTEMS FOR BONE TISSUE
ENGINEERING and rather poor degradation cannot be ignored (Pearlin
et al., 2018). According to types of raw materials, hydrogels can be briefly
categorized into natural and synthetic. It is usually considered
that natural hydrogels are more biocompatible and bioactive,
while synthetic ones possess more tunable mechanical and
degradation properties. 3D-structured, highly water-containing,
and biocompatible hydrogels act as excellent extracellular matrix
(ECM) analogs. The porous structure of the hydrogel enables
substance exchange and cell entrance at the initial stage as well as
vascular ingrowth in the follow-up stage. It has been substantially
shown that cells are easily suspended within hydrogels, and the
viability of the encapsulated cells is highly preserved (Gao et al.,
2020; Paez et al., 2020). Extensive drug and sustained release strategies have been
designed for bone tissue engineering. Herein, we introduce
studies on hydrogel-based controlled release systems according
to the category of bioactive molecule loaded within. Peptides The majority of cytokines, GFs, and hormones that stimulate
bone formation are peptides. These biomolecules are produced
through the autocrine, paracrine, and endocrine systems, acting
concertedly to regulate the complex cascade of bone-related gene
expression (Lee and Shin, 2007; Farokhi et al., 2016). Hence, a
well-orchestrated sustained release system of these peptides has
been pursued in order to present a more biomimetic approach. MSCs are highly sensitive to physical parameters (Higuchi
et al., 2013), including viscoelasticity Engler et al. (2006) and
topography (Fiedler et al., 2013), in the surrounding milieus. The
stiffness (elastic modulus) of the matrix is believed to contribute
greatly to determining stem cell fate. As Engler et al. (2006)
demonstrated, 2D-cultured MSCs exhibited osteogenic
characteristics when the microenvironmental stiffness was
relatively rigid, at 20–40 kPa. However, osteogenesis occurred
at 11–30 kPa when MSCs were cultivated 3-dimensionally
(Huebsch et al., 2010). Due to flexible synthetic strategies and
the range of constituents, hydrogels possess tunable physio-
mechanical properties, which could match the desirable ranges
of material elasticity, porosity, and degradation rate required for
bone tissue engineering (Slaughter et al., 2009). Meanwhile,
photodegradable (Lunzer et al., 2018), thermal-sensitive, or
pH-sensitive (GhavamiNejad et al., 2016) linkages as well as
other advantageous materials could be subtly introduced into
hydrogels, which may fabricate a versatile and intelligent
composite system to fulfill the growing clinical demands. Frontiers in Pharmacology | www.frontiersin.org BMP Non-covalent immobilization strategies are the most commonly
used in hydrogel-based drug depots, the drug release rate was
mainly determined by parameters such as crosslink density,
carrier affinity for drugs, and the matrix degradation profile
(Dimatteo et al., 2018). Bioactive factors also could be linked
covalently to polymers by which a longer drug retention time
would be achieved, and covalent linkages could be broken as
reactions of specific external cues, leading an on-demand drug
controlled release. Moreover, other sustained release systems like
microspheres could be introduced to hydrogel matrix, enabling
multiple drug molecules sustained release in sequential or
spatiotemporal manners (Chen et al., 2010). May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org 3 Hydrogel-Based Drug Release for BTE Zhang et al. augmentation was achieved by the hydrogel loaded with the
highest concentration. properties observed in vitro, significant enhancement by VEGF
on new bone formation has not been detected. Amirian et al. (2015) coated VEGF and BMP-2 separately onto gelatin-pectin-
biphasic calcium phosphate composites. The results revealed that
composites coated with VEGF mainly aided in woven bone
formation, and the percent of new bone formation was not
greater than those coated with BMP-2. Apart from adsorption or physical entrapment, electrostatic,
hydrophobic, or other interactions have been introduced into the
systems to prolong the release of BMPs. Heparin was reported to
be a strong binder to BMPs, yet the side effects were not
negligible. Heparin mimics, which are usually negatively
charged, are supposed to be capable of controlling BMP
release. Chondroitin sulfate (Anjum et al., 2016), 2-N,6-O-
sulfated chitosan (26SCS)-based nanoparticles (Cao et al.,
2014), alginate sulfate (Park et al., 2018) were synthesized by
researchers, and satisfactory results were achieved both in vitro
and in vivo. When higher concentrations of heparin mimics were
introduced, the release rate of BMP became slower. Seo et al. (2015, 2017) harnessed the ionic and hydrophobic interactions
provided by polyphosphazene nanoparticles. They found that
release rate of BMPs were controlled by the types and amounts of
pendants. Thus, the optimal release profile and osteogenesis
outcomes rely on a reasonable proportion of BMP-
tethering molecules. Since exclusive delivery of VEGF performed barely
satisfactorily in GF-induced osteogenesis, dual or multidrug
delivery is warranted. When accompanied by BMP-2, VEGF
exhibited a significant improvement in bone formation
compared with hydrogels encapsulating BMP-2 alone. BMP VEGF
combined with BMP-2 has been used routinely as a GF formula
in bone tissue engineering. Similar loading strategies were
applied by Barati et al. (2016) and Kader et al. (2019) for
spatiotemporal release of BMP-2 and VEGF. MSCs and BMP-
tethered nanoparticles were embedded in the outer space, while
endothelial colony-forming cells (ECFCs) and VEGF-tethered
nanoparticles were dispersed inside the microchannel-patterned
hydrogel, as illustrated in Figure 1. Degradation and GF release
could be tuned by altering stoichiometric ratio chain-extended
molecules and proteolysis. According to the data, the release of
VEGF and BMP-2 could last over 10 days and 21 days,
respectively. It was observed that the patterned hydrogel dual
delivery system performed significantly better than that of single
delivery systems, which was attributed to paracrine crosstalk. During bone repair, VEGF expression peaks appear in the early
period, while BMP peaks later. Thus, consisting of a PLGA
microsphere-incorporating poly(propylene fumarate) (PPF) rod
surrounded by a rapidly degrading gelatin hydrogel, the
composite was designed as a GF delivery vehicle (Kempen
et al., 2009). VEGF was encapsulated in the hydrogel, whereas
BMP-2 was immobilized by microspheres inside the rod in order
to achieve an ideal GF sequential release pattern. VEGF exhibited
a large initial burst release within the first 3 days, and BMP-2
showed sustained release over 8 weeks. Likewise, although VEGF
did not induce neo-bone formation, it significantly enhanced
BMP-induced osteogenesis. Organic-inorganic modular scaffolds
are able to optimally orchestrate dual GF release and serve as an
“anatomy-structure-function” trinity system in regenerating
weight-bearing bones (Bao et al., 2017). Mesoporous bioactive
glass (MBG) with hollowed channels and hierarchical porous
structures was introduced in a controlled release system as a
scaffold (Tang et al., 2020). VEGF was carried by hydrogel inside
the channel, and BMP-2 was adsorbed by the MBG scaffolds. 26SCS acted as an analog of ECM, which exhibits super-affinity
to GFs. In vitro experiments showed that 26SCS promoted the
bioactivity of BMP-2 and VEGF. It could be assumed that the
VEGF hydrogel column in the hollowed channels might induce
chemotaxis of vascular endothelial cells, thus regulating cell
migration and vascular infiltration. Moreover, increased type H
vessels and neotissue ingrowth were observed. Genetic engineering is another option to obtain long-lasting
BMP release. As Lin et al. Vascular Epithelial Growth Factor (VEGF) p
(
)
Vascularization plays a crucial role in both bone development
and bone regeneration (Collin-Osdoby, 1994; Olsen et al., 2000). Blood vessels do not solely work as substance exchange
pathways; they are also regarded as highly active paracrine
organs targeting various progenitors during bone formation
and reconstruction (Collin-Osdoby, 1994). VEGF, a key
angiogenic growth factor (Carmeliet and Jain, 2011), has been
widely used in bone tissue engineering. The cooperation between VEGF receptors and integrin
adhesion receptors has been elucidated in angiogenic
regulation. Garcia et al. (2016) engineered a protease-
degradable, GFOGER-modified PEG hydrogel as a VEGF
depot. They found that covalently linked VEGF remained
highly bioactive during a prolonged release period. Whereas it
was shown that a GFOGER hydrogel augmented
neovascularization regardless of exogenous VEGF, micro
computed tomography (micro-CT) showed delivering
exogenous VEGF failed to enhance critical-sized bone repair. Heterogenous material composites are manufactured by which
we can juggle both timed drug release and osteoconduction. Composed of a 3D multichannel calcium phosphate cement
(CPC) and alginate/gellan gum (AlgGG) hydrogel, the CPC/
AlgGG biphasic scaffold tethers VEGF via the interaction with
heparin (Ahlfeld et al., 2019). Despite some remarkable Frontiers in Pharmacology | www.frontiersin.org BMP (2019) described in a manuscript, the
BMP gene was transduced into human bone marrow-derived
stem cells (BMMSCs), obtaining a continuous (up to 56 days)
and economical BMP supply. Using visible light-based
projection stereolithography (VL-PSL) technology to
encapsulate the transduced cells, the researchers were able to
fabricate more structurally and geometrically compatible
constructs for individualized bone defects, which would be
conducive to achieving tissue fusion and bone tissue
engineering long-term success. Fibroblast Growth Factor (FGF) FGF signaling is a dominant regulator during bone development
and fracture repair (Bourque et al., 1993; Kronenberg, 2003). However, contradictory results have implied that FGF signaling May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org Frontiers in Pharmacology | www.frontiersin.org 4 Zhang et al. Hydrogel-Based Drug Release for BTE A
B
FIGURE 1 | Schematic illustration of (A) nanogel (NG) assembly and peptide grafting. (B) Achievement of BMP-2 and VEGF spatiotemporal release profiles via a
patterned hydrogel-based sustained release system. Reprinted from a previous article by Barati et al. (2016) with permission. B FIGURE 1 | Schematic illustration of (A) nanogel (NG) assembly and peptide grafting. (B) Achievement of BMP-2 and VEGF spatiotemporal release profiles via a
patterned hydrogel-based sustained release system. Reprinted from a previous article by Barati et al. (2016) with permission. to VEGF and FGF9 to obtain a covalent connection with the
fibrin hydrogel. BMP-2 was transfected into BMSCs, endowing a
greater osteogenic ability and resistance of the osteogenic
differentiation inhibition induced by fusion with FGF9. Less
bone was formed in the FGF9 groups compared to the groups
treated with only VEGF, whereas VEGF/FGF9-loaded
composites performed the best among the groups. may exert dual-directional effects on osteogenic procedures,
probably in a dose-dependent manner (Kato et al., 1998;
Quarto and Longaker, 2006). Thus, sustained release should be
achieved when FGF is delivered in bone tissue engineering. Two Japanese groups encapsulated FGF in gelatin hydrogels
for controlled release (Kodama et al., 2009; Furuya et al., 2014). A
longer FGF release period may improve cell proliferation, the
expression levels of osteogenic markers and BMP-2 as well as
bone mineral density (BMD) at defect sites. However, these
enhancements vanished, and side effects occurred when a high
dose of FGF was delivered (Kodama et al., 2009). In order to
achieve bone-like biomechanical properties and slower release of
FGF, a stiffer hydrogel matrix, poly(2-hydroxyethyl
methacrylate) copolymerized with 2-vinyl pyrrolidone, was
engineered (Mabilleau et al., 2008). The data suggested that in
the first 4 days, the FGF release rate was approximately 1% per
day, which was relevant to hydrogel swelling. Unfortunately, no
significant difference between the FGF and control groups was
noted in bone mass, but the poorly mineralized woven bone area
was significantly larger in the FGF group. Other Peptides Other peptides that regulate the bone regeneration cascade,
including osteoprotegerin (OPG) (Jayash et al., 2017), stromal
cell-derived factor-1a (SDF-1a) (Ratanavaraporn et al., 2011;
Cipitria et al., 2017; Mi et al., 2017), platelet-derived growth
factor (PDGF) (Wang et al., 2020), and parathyroid hormone
(PTH) (Erten Taysi et al., 2019), etc., might also be worthy of an
attempt. The selected studies of hydrogel-based peptide
sustained release systems for bone regeneration and their
findings are concluded in Table 1. Frontiers in Pharmacology | www.frontiersin.org Nucleic Acids (Wang et al.,
2020)
PTH
CS microsphere suspended in
poloxamer hydrogel
43% of PTH released in first week,
sustained release lasted over 27
days
New bone formation was found to be significantly
higher compared to other groups after 10 days, but
on day 21 a significant difference exists only when
compared with the no treatment group
(Erten Taysi
et al., 2019)
(Continu TABLE 1 | Summary of selected studies of hydrogel-based peptide sustained release systems
Peptide
Carrier material
Release pattern
BMP-2
PLGA microspheres + bisphosphate
modified oligo OPF hydrogel
Burst and sustained
BMP-2
Nanofibrous mesh + peptide
modified alginate hydrogel
Spatiotemporal controlled release
BMP-2
GFOGER-modified MMP-degradable
PEG-maleimide hydrogel
More than 20% BMP-2 remained
after 14 days
BMP-2
VL-PSL-manufactured live-cell
hydrogel scaffold
Sustained expression for 56 days
BMP-7
CS/b-GP hydrogel
Release 46% in first 12 h, 84% by
the end of 336 h
GDF-5
Photo-cured hyaluronic acid (HA)
hydrogel
Release profiles varies with the
initial drug concentration, sustained
release period over 25 days
BMP-2 + VEGF
Acrylate-functionalized lactide-chain-
extended star polyethylene glycol
(SPELA) hydrogel + gelatin
methacryloyl (GelMA) hydrogel + PEG
nanogel
Release of VEGF and BMP-2
lasted over 10 days and 21 days,
respectively (tunable release
kinetics)
VEGF + BMP-2
PLGA microsphere + PPF rod +
gelatin hydrogel
A large initial burst was shown in
vivo, which changed significantly
from ex vivo release profiles
VEGF + BMP-2
Hydroxyapatite (HA)/
polycaprolactone (PCL) scaffold +
PLGA-PEG-PLGA hydrogel
Burst release in first 3 days,
sustained release for 3 weeks
BMP-2 + VEGF
MBG-based matrix + GelMA/26SCS
hydrogel
The release rates of BMP-2 and
VEGF were 24.01% and 34.47%
respectively within 24 h, 67.90%
and 82.73% respectively in 14
days
BMP-2 + bFGF
Gelatin microspheres + n-HA/PU40
scaffold + F-127 hydrogel
Pronounced burst release
occurred in first 24 h, linear release
in following 29 days
VEGF + FGF9
Nanocalcium sulfate + fibrin hydrogel
Addition of the peptide sequence
decreased GFs release in an
enzyme concentration-dependent
manner
Osteoprotegerin
(OPG)
CS hydrogel
Lasts 28 days, release profile could
be adjusted by CS molecular
weight
SDF-1a
CS/carboxymethyl CS nanoparticles
+ CS/b-GP hydrogel
20% initial burst release, a
cumulative release of 40% over 28
days
SDF-1a + BMP-
2
Gelatin hydrogel
Large initial burst release of SDF-
1a in first 3 days, which may due
to BMP-2 combination
SDF-1a
RGD-modified alginate hydrogel
Sustained release over 42 days
PDGF-BB
+BMP-9
Sericin hydrogel (genetically
incorporated)
Almost 48% released within 17
days, intermittent rapid and slow
release phases
PTH
CS microsphere suspended in
poloxamer hydrogel
43% of PTH released in first week,
sustained release lasted over 27
days . (Kempen et al.,
2009) Nucleic Acids Ectopic bone formation in
hindlimb ischemia model suggested type H vessels
and neobone formation significantly increased
(Tang et al.,
2020)
BMP-2 + bFGF
Gelatin microspheres + n-HA/PU40
scaffold + F-127 hydrogel
Pronounced burst release
occurred in first 24 h, linear release
in following 29 days
Micro-CT analysis indicated dual-delivery reached
significantly higher bone volume (BV). Quantitative
histological analysis showed remarkable tissue
response
(Chen et al.,
2016)
VEGF + FGF9
Nanocalcium sulfate + fibrin hydrogel
Addition of the peptide sequence
decreased GFs release in an
enzyme concentration-dependent
manner
Radiographical and quantitative analysis of micro-CT
showed the highest BV in dual-delivery hybrid
composite. Quantification of blood vessels in
explanted tissue suggested more neovessels were
obtained
(Yuan et al.,
2016)
Osteoprotegerin
(OPG)
CS hydrogel
Lasts 28 days, release profile could
be adjusted by CS molecular
weight
An almost-complete recovery was observed,
osteocalcin and osteopontin were upregulated
(Jayash et al.,
2017)
SDF-1a
CS/carboxymethyl CS nanoparticles
+ CS/b-GP hydrogel
20% initial burst release, a
cumulative release of 40% over 28
days
Micro-CT showed most new bone formation within
the defect area
(Mi et al., 201
SDF-1a + BMP-
2
Gelatin hydrogel
Large initial burst release of SDF-
1a in first 3 days, which may due
to BMP-2 combination
Better new bone formation was observed in the dual-
delivery group. SDF-1a enhanced BMP-2 osteogenic
effects
(Ratanavarapo
et al., 2011)
SDF-1a
RGD-modified alginate hydrogel
Sustained release over 42 days
Improvements induced by SDF-1a or hydrogel
stiffness levelled within 8 weeks. Higher number of
cells were recruited by SDF-1a, but the difference
was not significant in vivo
(Cipitria et al.,
2017)
PDGF-BB
+BMP-9
Sericin hydrogel (genetically
incorporated)
Almost 48% released within 17
days, intermittent rapid and slow
release phases
Biocompatible compared with other materials and
stimulated cell proliferation. Osteogenic markers were
significantly upregulated, and greater bone formation
when accompanied by BMP-9. Nucleic Acids It is a preferable strategy for other GFs to accompany FGF in
order to obtain a promising outcome. Chen et al. (2016) chose
gelatin microspheres as BMP-2 and basic FGF (bFGF) carriers,
which were further embedded in a commercialized injectable
thermal-sensitive hydrogel. The hydrogel was injected into a
porous cell-loading scaffold before use. Micro-CT revealed that
the dual-loaded composites achieved the best reparative results. As expected, composites loading bFGF alone regenerated less
bone and neobone at the margin of the defect areas, while the
dual-loaded composites showed much more central area bone
formation. FGF9 has been indicated to be a stabilizing factor for
neovessels, thus, Yuan et al. (2016) introduced FGF9 as an
assistant for VEGF, exerting synergetic effects on angiogenesis
in bone tissue engineering. A specific peptide segment was fused Since GFs and cytokines are required for weeks during new bone
formation, gene therapy might be a feasible alternative. Delivering DNA or RNA locally to increase or knockdown
target gene expression, gene therapy is capable of manipulating
the microenvironment and determining cell fate in bone
regenerative medicine. Fang et al. (1996) utilized collagen sponges as BMP-4 and PTH
plasmid DNA carriers to regenerate nonunion rat femur defects
early in 1996. Bonadio et al. (1999) confirmed that non-viral DNA
delivery possesses numerous advantages compared with the protein
strategy. Hydrophilic nucleic acids and hydrogels could provide
stable and sequestered environments for gene delivery. Komatsu
et al. (2016) demonstrated that gelatin hydrogels could transduce
BMP-2 plasmid DNA efficiently, facilitating local bone regeneration. May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org 5 Hydrogel-Based Drug Release for BTE Zhang et al. TABLE 1 | Summary of selected studies of hydrogel-based peptide sustained release systems. Nucleic Acids Peptide
Carrier material
Release pattern
Findings (ex/in vivo)
Reference
BMP-2
PLGA microspheres + bisphosphate
modified oligo OPF hydrogel
Burst and sustained
Osteoconductivity and osteoinductivity were
significantly improved
(Olthof et al.,
2018)
BMP-2
Nanofibrous mesh + peptide
modified alginate hydrogel
Spatiotemporal controlled release
Micro-CT showed more bone regeneration, superior
mechanical properties of neobone were achieved
(Kolambkar
et al., 2011)
BMP-2
GFOGER-modified MMP-degradable
PEG-maleimide hydrogel
More than 20% BMP-2 remained
after 14 days
GFOGER-modified hydrogel exhibited intrinsic
osteogenic activities, micro-CT demonstrated;
improved bone repair
(Shekaran et a
2014)
BMP-2
VL-PSL-manufactured live-cell
hydrogel scaffold
Sustained expression for 56 days
Micro-CT and histological evidence indicated mature
and robust bone formation
(Lin et al., 201
BMP-7
CS/b-GP hydrogel
Release 46% in first 12 h, 84% by
the end of 336 h
Radiographical and histological observation
suggested better periodontal regeneration
(Zang et al.,
2019)
GDF-5
Photo-cured hyaluronic acid (HA)
hydrogel
Release profiles varies with the
initial drug concentration, sustained
release period over 25 days
Hydrogel with the highest drug concentration
displayed promoted osteogenic potential both in vitro
and in vivo
(Bae et al., 20
BMP-2 + VEGF
Acrylate-functionalized lactide-chain-
extended star polyethylene glycol
(SPELA) hydrogel + gelatin
methacryloyl (GelMA) hydrogel + PEG
nanogel
Release of VEGF and BMP-2
lasted over 10 days and 21 days,
respectively (tunable release
kinetics)
Patterned constructs significantly increase osteogenic
and vasculogenic differentiation of precursors, bFGF
expression was upregulated
(Barati et al.,
2016)
VEGF + BMP-2
PLGA microsphere + PPF rod +
gelatin hydrogel
A large initial burst was shown in
vivo, which changed significantly
from ex vivo release profiles
Micro-CT and histological section demonstrated co-
delivery significantly enhanced osteogenesis and
angiogenesis ectopically, but it did not reach
significant results orthotopically
(Kempen et al
2009)
VEGF + BMP-2
Hydroxyapatite (HA)/
polycaprolactone (PCL) scaffold +
PLGA-PEG-PLGA hydrogel
Burst release in first 3 days,
sustained release for 3 weeks
Micro-CT showed newly-formed callus in co-delivery
group almost covered defect areas, histological
analysis showed no significant difference between
co-delivery group and autologous group
(Bao et al.,
2017)
BMP-2 + VEGF
MBG-based matrix + GelMA/26SCS
hydrogel
The release rates of BMP-2 and
VEGF were 24.01% and 34.47%
respectively within 24 h, 67.90%
and 82.73% respectively in 14
days
In vitro osteogenic and angiogenic has been
markedly improved. Nucleic Acids (Wang et al.,
2020)
New bone formation was found to be significantly
higher compared to other groups after 10 days, but
on day 21 a significant difference exists only when
compared with the no treatment group
(Erten Taysi
et al., 2019) May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org Hydrogel-Based Drug Release for BTE Zhang et al. TABLE 1 | Continued
Peptide
Carrier material
Release pattern
Findings (ex/in vivo)
Reference
Abaloparatide
(analog of PTH)
Photo-crosslinked methacrylated
gelatin hydrogel
25% released within 24 h,
remaining was released steadily
over next 10 days
Drug-loaded hydrogel showed significantly higher
rate of bone regeneration
(Ning et al.,
2019)
Oxytocin
PLGA microsphere + poloxamer
hydrogel + b‐tricalcium phosphate (b-
TCP) and hydroxyapatite
42% released in first week,
complete release within 32 days
4 weeks after operation, the lowest residual graft and
highest BMD and BV was obtained among all groups
(Akay et al.,
2020)
Calcium
accumulating
peptide
(artificially
synthesized)
Gelatin-derived hydrogel
Sustained release over 7 days,
collagenase accelerated release
Bone formation markers expression levels were
enhanced. Micro-CT and histology showed the
regenerative effect was superior to that of BMP-2
hydrogels
(Jo et al., 2018) Findings (ex/in vivo) sustained delivery of Mg2+ have been applied to hydrogel-based
scaffolds. Lin et al. (2018b) coated MgO nanoparticles with
PLGA and an alginate hydrogel, constructing a monodisperse
core-shell delivery system. The release profile of Mg2+ revealed a
significant suppression of the initial burst, and its release rate was
stable and programmed. Enhancement of progenitor cell viability
and proliferation, upregulation of osteogenic gene expression
levels, and increased bone regeneration volume in vivo were
attributed to the stable and precise Mg2+ supply. Bisphosphonates (BPs) possess two adjacent phosphonic
groups, which are propitiously bind to divalent metal ions. Zhang and colleagues (Zhang et al., 2017) developed acrylated-
BP-Mg nanoparticles to deliver Mg2+ as well as strengthen the
acellular hydrogel composite, which serves as a matrix for in situ
bone formation, via multivalent crosslinked domains. They also
utilized Mg2+ to fulfill on-demand intelligent drug release in
bone tissue engineering (Zhang et al., 2018). Intriguingly, Mg2+
played multiple roles in this research. First, BP-Mg nanoparticles
enabled hydrogel formation and stabilized the prodrug. Second,
Mg2+ promoted osteogenic differentiation, resulting in increased
alkaline phosphatase (ALP) expression. However, and more
importantly, Mg2+ is also a critical cofactor of ALP. Nucleic Acids Findings (ex/in vivo)
Reference
Osteoconductivity and osteoinductivity were
significantly improved
(Olthof et al.,
2018)
Micro-CT showed more bone regeneration, superior
mechanical properties of neobone were achieved
(Kolambkar
et al., 2011)
GFOGER-modified hydrogel exhibited intrinsic
osteogenic activities, micro-CT demonstrated;
improved bone repair
(Shekaran et al.,
2014)
Micro-CT and histological evidence indicated mature
and robust bone formation
(Lin et al., 2019)
Radiographical and histological observation
suggested better periodontal regeneration
(Zang et al.,
2019)
Hydrogel with the highest drug concentration
displayed promoted osteogenic potential both in vitro
and in vivo
(Bae et al., 2014)
Patterned constructs significantly increase osteogenic
and vasculogenic differentiation of precursors, bFGF
expression was upregulated
(Barati et al.,
2016) Micro-CT and histological section demonstrated co-
delivery significantly enhanced osteogenesis and
angiogenesis ectopically, but it did not reach
significant results orthotopically
(Kempen et al.,
2009)
Micro-CT showed newly-formed callus in co-delivery
group almost covered defect areas, histological
analysis showed no significant difference between
co-delivery group and autologous group
(Bao et al.,
2017)
In vitro osteogenic and angiogenic has been
markedly improved. Ectopic bone formation in
hindlimb ischemia model suggested type H vessels
and neobone formation significantly increased
(Tang et al.,
2020)
Micro-CT analysis indicated dual-delivery reached
significantly higher bone volume (BV). Quantitative
histological analysis showed remarkable tissue
response
(Chen et al.,
2016)
Radiographical and quantitative analysis of micro-CT
showed the highest BV in dual-delivery hybrid
composite. Quantification of blood vessels in
explanted tissue suggested more neovessels were
obtained
(Yuan et al.,
2016)
An almost-complete recovery was observed,
osteocalcin and osteopontin were upregulated
(Jayash et al.,
2017)
Micro-CT showed most new bone formation within
the defect area
(Mi et al., 2017) Better new bone formation was observed in the dual-
delivery group. SDF-1a enhanced BMP-2 osteogenic
effects
(Ratanavaraporn
et al., 2011)
Improvements induced by SDF-1a or hydrogel
stiffness levelled within 8 weeks. Higher number of
cells were recruited by SDF-1a, but the difference
was not significant in vivo
(Cipitria et al.,
2017)
Biocompatible compared with other materials and
stimulated cell proliferation. Osteogenic markers were
significantly upregulated, and greater bone formation
when accompanied by BMP-9. Ions or Small Molecules To date, a number of metal ions and artificially synthesized
compounds have been found to be beneficial in bone regeneration. Achieving a sustained release pattern and longer duration of drug
function may lead to promising therapeutic outcomes. Nucleic Acids ALP
enzymatic hydrolysis was promoted; thus, more bioactive drug
molecules were generated, which introduced positive feedback
(Figure 2). According to the results, this strategy significantly
enhanced bone regeneration. CS or polyethyleneimine (PEI) is usually introduced as the carrier
due to the electrostatic interaction between the negatively charged
nucleic acids and the polycations. It was reported that branched
PEI-HA-DNA complexes were entrapped in bilayered OPF
hydrogels to restore osteochondral defects (Needham et al., 2014). Moreover, BMP-2 plasmid DNA conjugated with CS nanoparticles
exhibited significant augmentation in hydrogel-mediated rat
calvaria bone regeneration (Li et al., 2017). Due to the low
stability of liposomes and electrostatic disturbance of other
charged compounds, calcium phosphate (CaP) can also be used
for DNA incorporation and transfection in bone tissue engineering
(Krebs et al., 2010). MicroRNAs (miRNAs) and small interfering RNAs (siRNAs)
are groups of short single-stranded RNA fragments that
downregulate target gene expression post-translationally. Various miRNAs associated with bone formation have been
reported (Fang et al., 2015), shedding new light on future bone
tissue engineering. Nguyen et al. (2014) synthesized an 8-arm
PEG in situ-forming hydrogel loaded with siRNA/PEI
nanocomplexes. siRNA remained bioactive during the
prolonged release period. The in vitro results showed that
siNoggin and siNoggin/miRNA-20a sustained release
promoted hMSC osteogenic differentiation in 3D hydrogel
cultivation. As mentioned previously, a stiffer substrate may
lead to MSC osteogenic differentiation. Carthew et al. (2020)
incorporated PEG/gelatin norbornene hydrogels with
mechanosensitive miRNAs. MSCs encapsulated in hydrogels
were transfected in situ, which predominantly enhanced
osteogenic gene expression and mineralization. Researchers
presumed that the higher transfection efficacy might be
ascribed to longer cell exposure times to the transfection agent. Other metal ions, such as strontium ions (Sr2+) and cobalt
ions (Co2+), may act synergistically in bone reconstruction. A Sr2+-
crosslinked RGD-alginate hydrogel combined with Sr-doped
hydroxyapatite microspheres was engineered, showing a sustained
release of Sr2+ from two sources (Lourenco et al., 2019). The
researchers elaborated that this Sr-hybrid system facilitated MSC
osteogenic differentiation, inhibited the functions of osteoclasts and
modulated the inflammatory response. As a pro-vasculogenic
factor, Co2+ was incorporated into the alginate hydrogel shell,
while BMP-2 was laden into the collagen core (Perez et al., 2015). Co2+ released relatively rapidly, as expected. VEGF secretion and
qPCR revealed that Co2+ not only stimulated angiogenesis but also
elevated osteogenic gene expression. These results indicated an
appealing prospect for applying metal ions bone tissue
engineering in the future. Small Molecules necrosis factor-a (TNF-a) and interferon-g (IFN-g) pathways
(Liu et al., 2011). Statins are inhibitors of a key enzyme of
cholesterol synthesis and are widely used to lower serum lipids. Researchers have reported that osteogenesis was enhanced
concomitant with promoted BMP-2 expression in bone cells
when treating cells and rodents with statins (Mundy et al., 1999). Localized and sustained delivery of these drugs via hydrogels has
pointed to a new direction in bone tissue engineering. Many
relevant studies are listed and outlined in Table 2. A range of pharmaceutical molecules were designed or
discovered to be effective in bone regeneration. Highlighted as
chelating agents, BPs are utilized as antiresorptive drugs
frequently in clinics. BPs mainly target osteoclasts, impeding
the differentiation and maturation of osteoclast progenitors. Increasing evidence has shown that BPs directly or indirectly
take part in other bone-forming mechanisms and are capable of
targeting various cells (Corrado et al., 2017). Since bone healing
and regeneration is known to consist of three consecutive phases
of inflammation, repair, and remodeling, a proper scale of
immune response is indispensable (Claes et al., 2012). However, excess or aberrant immune activation may
jeopardize bone repair procedures (Claes et al., 2012; Gibon
et al., 2017). Therefore, immunomodulatory drugs, such as
nonsteroidal anti-inflammatory drugs (NSAIDs), have been
applied in bone tissue engineering. Evidence has shown that
aspirin elevates MSC osteogenic potency by inhibiting the tumor Metal Ions Since magnesium ions (Mg2+) play an important role in bone
metabolism and mineralization, a variety of strategies for the May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org Zhang et al. Hydrogel-Based Drug Release for BTE A
B
FIGURE 2 | Schematic illustration of (A) positive feedback mediated by a cofactor-assisted smart hydrogel drug release system and (B) in situ application to
promote bone regeneration. Reprinted from a previous article by Zhang et al. (2018) with permission. A A B GURE 2 | Schematic illustration of (A) positive feedback mediated by a cofactor-assisted smart hydrogel drug release system and
omote bone regeneration. Reprinted from a previous article by Zhang et al. (2018) with permission. tration of (A) positive feedback mediated by a cofactor-assisted smart hydrogel drug release system and (B) in situ application to
Reprinted from a previous article by Zhang et al. (2018) with permission. CONCLUSION AND FUTURE
PERSPECTIVES Hydrogel loaded with 10 mg of drug formed the largest
area of bone
(Tanigo et al., 2010)
Rosuvastatin
chitosan/chondroitin
sulfate nanoparticles+
Pluronic F127/hyaluronic
acid hydrogel composite
Release rate significantly slower than
control groups. 60% released from
composite in 48 h
Low cytotoxicity, more calcium deposits were observed
(Rezazadeh et al.,
2019) sustained release systems. Findings (ex/in vivo)
Reference
Hydrogel containing 10-6 M showed the best
augmentation in cell proliferation, osteogenic
differentiation, and bone regeneration. (Kim et al., 2017a)
Drug bioactivity was preserved by the hydrogel. nSi
may contribute to in vivo osteogenesis whereas Dex
showed limited effects. (Basu et al., 2018)
Percent of mineralized tissue was significantly higher
compared to control group. (Fang et al., 2018)
Low cytotoxicity, significantly improved expression of
osteogenic markers and calvarial defect regeneration. Relatively low local inflammation status might be
attributed to being laden with aspirin. (Zhang et al., 2019)
Osteoblasts grew and mineralized significantly
regardless of drug exhaustion. Osteogenic genes
increased over time, while osteogenic suppressing gene
expression decreased. (Lin et al., 2018a)
More newly-formed bone and blood vessels were
observed
(Nabavi et al., 2020)
Good biocompatibility, stimulated ALP activity and
mineralization
(Yan et al., 2018)
Hydrogel loaded with 10 mg of drug formed the largest
area of bone
(Tanigo et al., 2010) Findings (ex/in vivo) Released 50% and 90% in 2.5 h and 8
h, respectively Hydrogel loaded with 10 mg of drug formed the largest
area of bone
(Tanigo et al., 2010) Low cytotoxicity, more calcium deposits were observed
(Rezazadeh et al.,
2019) and drugs, such as electrostatic interactions and covalent bonds,
and other drug reservoirs could be introduced into hydrogels,
providing more efficacious drug protection and immobilization. However, negative results have been reported from sustained
release systems that did not facilitate bone formation mainly
because the carrier exhibited an extremely strong affinity towards
the growth factor, resulting in a low level of drug concentration in
the surrounding tissue (Hettiaratchi et al., 2017). Thus, optimal
drug concentrations should also be determined to achieve a more
reasonable and effective release profile. rate, which makes them qualified bone tissue engineering
scaffolds. Due to their high water content, chemical inertness
and relatively sequestered and stable internal environment, they
are also excellent in preserving the viabilities of the laden cells
and bioactive factors. CONCLUSION AND FUTURE
PERSPECTIVES In this review, we summarized a series of investigations focused
on hydrogel-based drug sustained release systems in bone tissue
engineering. The hydrogels possess a porous microarchitecture,
tunable biophysical parameters, and an adjustable degradation May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org 8 Hydrogel-Based Drug Release for BTE Zhang et al. TABLE 2 | Summary of selected studies on hydrogel-based small bioactive compound sustained release systems. Drug
Carrier material
Release pattern
Findings (ex/in vivo)
Reference
Alendronate
Fibrin hydrogel
Steady release rate, cumulative release
of approximately 45% over 10 days
Hydrogel containing 10-6 M showed the best
augmentation in cell proliferation, osteogenic
differentiation, and bone regeneration. (Kim et al., 2017a)
Dexamethasone
(Dex)
DNA- 2D silicate
nanodisks (nSi) hybrid
hydrogel
Release rate decreased with higher nSi
concentration. Half-time of release was
measured from 2.5 to 5.5 days
Drug bioactivity was preserved by the hydrogel. nSi
may contribute to in vivo osteogenesis whereas Dex
showed limited effects. (Basu et al., 2018)
Aspirin
Thermo-sensitive alginate/
b-TCP hydrogel composite
20% drug released in the first day,
40% in 3 days, slowdown in day 5. Percent of mineralized tissue was significantly higher
compared to control group. (Fang et al., 2018)
Aspirin
Tetra-PEG hydrogel
Released approximately 40% in first 2
days, cumulative release of 80% in 14
days
Low cytotoxicity, significantly improved expression of
osteogenic markers and calvarial defect regeneration. Relatively low local inflammation status might be
attributed to being laden with aspirin. (Zhang et al., 2019)
Diclofenac
CS-coated alginate
hydrogel
Released 50% and 90% in 2.5 h and 8
h, respectively
Osteoblasts grew and mineralized significantly
regardless of drug exhaustion. Osteogenic genes
increased over time, while osteogenic suppressing gene
expression decreased. (Lin et al., 2018a)
Tacrolimus
Type I collagen hydrogel
21 days release profile remained similar
for different concentrations. Steady
release rate. More newly-formed bone and blood vessels were
observed
(Nabavi et al., 2020)
Simvastatin
Maltodextrin micelle-CHO/
hydrogel composite
Slow release profile, exhibiting a slight
difference according to different
degrees of oxidation
Good biocompatibility, stimulated ALP activity and
mineralization
(Yan et al., 2018)
Simvastatin
L-lactic acid oligomer (LAo)
modified gelatin micelle/
gelatin hydrogel composite
Drug released faster as hydrogel
crosslinking degree decreased. Release
rate showed a good correlation with
hydrogel degradation rate. Low cytotoxicity, more calcium deposits were observed
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PERSPECTIVES With the combination of biophysical and
biochemical cues, researchers are able to facilely establish an
osteo-friendly microenvironment, which would be beneficial for
osteoprogenitors to obtain better bone regeneration. Thus,
hydrogel-based biomaterials are strong candidates for current
or future bone tissue engineering. As mentioned above, cells from different origins are involved
in the bone formation process. A vast number of GFs and
cytokines collaboratively trigger the repair cascade. Extensive
studies have already been conducted on multiple bioactive
factors controlled release. Spatiotemporal sequence release of
bioactive factors might be a better mimic of complex
regeneration procedures as well as exert extraordinary
synergistic effects on bone regeneration. Various of multiple
GFs delivery strategies was coherently summarized (Chen et al.,
2010). Nevertheless, controlling dose ratio of drugs to maximize
the synergistic effects and manipulating multiple bioactive factors
release kinetics to mimic physiological release profile in different
phases of bone regeneration are obstacles in nowadays bone tissue
regeneration which needs further investigation. Evidence has shown that hydrogel-based drug sustained release
systems are highly biocompatible and versatile drug deliverers,
obtaining satisfactory osteogenesis results both in vitro and in vivo. The drug release profile varies according to the loading strategy,
degradation ability of the matrix and drug concentration. Among
these studies, physical entrapment and diffusion are the most
applied drug loading and release strategies, respectively. In
particular, the dispersion of drugs, ions and small molecules
largely depends on hydrogel pore size and crosslinking density. Although it is quite simple and easy to operate, there are difficulties
in initial burst release management. Swelling or degradation of
hydrogel matrices contributes to polymer mesh size enlargement,
resulting in drug release acceleration, especially for
macromolecular drugs. Stronger interactions between matrices May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org 9 Hydrogel-Based Drug Release for BTE Zhang et al. FUNDING YZ and TY contributed equally to this review. TY, YW, and BH
designed and revised this article. YZ, TY, LP, and QS collected This work was supported by the National Natural Science
Foundation of China (51972005, 51672009, 51903003),
National Natural Science Foundation of China Youth Fund
(81922019), and National Youth Top-notch Talent Support
Program (QNBJ2019-3). This work was supported by the National Natural Science
Foundation of China (51972005, 51672009, 51903003), National Natural Science Foundation of China Youth Fund
(81922019), and National Youth Top-notch Talent Support
Program (QNBJ2019-3). the literatures, arranged the outline of collected documents, and
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j.msec.2019.02.024 Conflict of Interest: The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could be construed as a
potential conflict of interest. Zhang, K., Lin, S., Feng, Q., Dong, C., Yang, Y., Li, G., et al. (2017). Nanocomposite
hydrogels stabilized by self-assembled multivalent bisphosphonate-magnesium
nanoparticles mediate sustained release of magnesium ion and promote in-situ
bone regeneration. Acta Biomater. 64, 389–400. doi: 10.1016/j.actbio.2017.09.039 Copyright © 2020 Zhang, Yu, Peng, Sun, Wei and Han. This is an open-access article
distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the
original author(s) and the copyright owner(s) are credited and that the original
publication in this journal is cited, in accordance with accepted academic practice. Zhang, K., Lin, S., Feng, Q., Dong, C., Yang, Y., Li, G., et al. (2017). Nanocomposite
hydrogels stabilized by self-assembled multivalent bisphosphonate-magnesium
nanoparticles mediate sustained release of magnesium ion and promote in-situ
bone regeneration. Acta Biomater. 64, 389–400. doi: 10.1016/j.actbio.2017.09.039 Zhang, K., Jia, Z., Yang, B., Feng, Q., Xu, X., Yuan, W., et al. (2018). Adaptable
Hydrogels Mediate Cofactor-Assisted Activation of Biomarker-Responsive
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Sci. (Weinh) 5 (12), 1800875. doi: 10.1002/advs.201800875 Frontiers in Pharmacology | www.frontiersin.org Zhang, K., Lin, S., Feng, Q., Dong, C., Yang, Y., Li, G., et al. (2017). Nanocomposite
hydrogels stabilized by self-assembled multivalent bisphosphonate-magnesium
nanoparticles mediate sustained release of magnesium ion and promote in-situ
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Zhang, K., Jia, Z., Yang, B., Feng, Q., Xu, X., Yuan, W., et al. (2018). Adaptable
Hydrogels Mediate Cofactor-Assisted Activation of Biomarker-Responsive
Drug Delivery via Positive Feedback for Enhanced Tissue Regeneration. Adv.
Sci. (Weinh) 5 (12), 1800875. doi: 10.1002/advs.201800875 REFERENCES No
use, distribution or reproduction is permitted which does not comply with these terms. Zhang, K., Jia, Z., Yang, B., Feng, Q., Xu, X., Yuan, W., et al. (2018). Adaptable
Hydrogels Mediate Cofactor-Assisted Activation of Biomarker-Responsive
Drug Delivery via Positive Feedback for Enhanced Tissue Regeneration. Adv. Sci. (Weinh) 5 (12), 1800875. doi: 10.1002/advs.201800875 May 2020 | Volume 11 | Article 622 Frontiers in Pharmacology | www.frontiersin.org 13
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ANALISIS PENETAPAN HARGA KAYU JATI PLUS PERHUTANI BERDASARKAN STUMPAGE COST DAN WILLINGNESS TO PAY
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ANALISIS PENETAPAN HARGA KAYU JATI PLUS PERHUTANI
BERDASARKAN STUMPAGE COST DAN WILLINGNESS TO PAY Arga Pramudita1*, Suryanaji2
1 Kelompok Peneliti Industri dan Pemasaran, Puslitbang Perhutani
2 Biro Perencanaan dan Pengembangan Bisnis, Divisi Regional Jawa Tengah
Puslitbang Perhutani, Jl. Wonosari, Batokan, Cepu, 58302
* Email: argapram@gmail.com Arga Pramudita1*, Suryanaji2
1 Kelompok Peneliti Industri dan Pemasaran, Puslitbang Perhutani
2 Biro Perencanaan dan Pengembangan Bisnis, Divisi Regional Jawa Tengah
Puslitbang Perhutani, Jl. Wonosari, Batokan, Cepu, 58302
* Email: argapram@gmail.com Kata Kunci: Harga Pokok Produksi, Jati, Konsumen, Market Value, Perhutani. Kata Kunci: Harga Pokok Produksi, Jati, Konsumen, Market Value, Perhutani. Kata Kunci: Harga Pokok Produksi, Jati, Konsumen, Market Value, Perhutani. 2017 seluas ± 200.000 ha. Potensi
kayu penjarangan dari luasan tanaman
JPP dalam kondisi baik, dengan
asumsi hasil penjarangan sebesar 4
m3/ha, dapat mencapai ±170.000 m3. 2017 seluas ± 200.000 ha. Potensi
kayu penjarangan dari luasan tanaman
JPP dalam kondisi baik, dengan
asumsi hasil penjarangan sebesar 4
m3/ha, dapat mencapai ±170.000 m3. Risalah Kebijakan Pertanian dan Lingkungan
Vol. 8 No. 2 Agustus 2021: 53-59
ISSN : 2355-6226
E-ISSN : 2477-0299 Risalah Kebijakan Pertanian dan Lingkungan
Vol. 8 No. 2 Agustus 2021: 53-59
ISSN : 2355-6226
E-ISSN : 2477-0299 RINGKASAN Jati Plus Perhutani (JPP) sebagai salah satu hasil program pemuliaan pohon memiliki
beberapa keunggulan komparatif dibandingkan dengan jenis jati yang dikembangkan melalui
perbanyakan konvensional dengan biji. Keunggulan dalam hal keliling batang dan produktivitas
yang tinggi dalam jangka waktu yang relatif singkat akan memberikan biaya pembangunan
tegakan yang minimal sehingga harga juga tidak terlalu tinggi. Penelitian ini bertujuan untuk
mengetahui harga pokok produksi kayu JPP hasil penjarangan melalui pendekatan biaya selama
daur dan pendekatan pasar (market value). Metode yang digunakan adalah pendekatan nilai
tegakan (stumpage cost) dan metode Willingness to Pay (WTP) yang terdiri dari survei pelanggan
dengan kuisioner dan in depth interview dengan sistematik random sampling. Hasil penelitian
menunjukkan nilai pembangunan tegakan JPP selama 9 tahun sebesar Rp. 20.024.309,-/ha,
dengan harga pokok penjualan sortimen AI sebesar Rp. 642.538,- dan sortimen AII sebesar Rp. 881.214,-. Sedangkan berdasarkan pendekatan pasar, konsumen menginginkan harga kayu JPP
tidak jauh berbeda dengan kayu jati asal hutan rakyat atau penurunan harga sebesar 13% - 22%. Rekomendasi kebijakan yang disarankan terhadap penentuan harga kayu JPP adalah perlu
pemisahan dan pembedaan harga jual dasar kayu JPP supaya dapat bersaing dengan harga kayu
rakyat dan dapat menjadi salah satu sumber pendapatan perusahaan dalam waktu yang tidak
terlalu lama. PERNYATAAN KUNCI •
Jati Plus Perhutani sebagai produk
hasil penelitian dan pengembangan
Puslitbang Perhutani dan sudah
ditanam secara operasional mulai
tahun 2007 sampai dengan tahun 53 53 Rumusan Kajian Strategis Bidang Pertanian dan Lingkungan ISSN : 2355 – 6226
E-ISSN : 2477 – 0299 dan furniture menyebabkan permintaan kayu
jati semakin meningkat. Seiring dengan
permintaan yang semakin meningkat maka
otomatis nilai ekonomi kayu jati juga
semakin tinggi. Perum Perhutani yang
sampai dengan saat ini masih mengelola
hutan jati di Pulau Jawa mempunyai peluang
untuk meningkatkan produktivitas hutan
jatinya yaitu dengan mengembangkan klon
unggul yaitu Jati Plus Perhutani (JPP). •
Harga Jual Dasar (HJD) merupakan
besaran harga yang ditetapkan oleh
Direksi Perum Perhutani untuk
kepentingan penjualan kontrak dan
penjualan langsung secara retail. Harga ini tertera dalam saluran
penjualan di Penjualan Online Toko
Perhutani yang merupakan e- commerce
Perum
Perhutani
dengan
akses
melalui tokopedia.com. Jati Plus Perhutani sebagai produk
hasil pemuliaan pohon yang dilakukan oleh
Puslitbang Perum Perhutani, memiliki
keunggulan di antaranya pertumbuhan
tegakan lebih cepat, seragam dan kayu yang
berkualitas. Dengan keunggulan tanaman
JPP ini maka diharapkan dengan penerapan
umur tebang yang lebih awal dan lebih cepat
bila dibandingkan umur tebang pada jati
konvensional tetap dapat dilakukan tanpa
mengabaikan faktor ekologis tanaman. Kebijakan harga jual dasar kayu JPP
masih disamakan dengan harga jual
kayu jati konvensional sehingga
konsumen kayu jati cenderung lebih
memilih kayu jati konvensional,
sehingga kayu JPP dianggap sebagai
produk slow moving karena kurang
diminati pasar. REKOMENDASI KEBIJAKAN •
Perlu dilakukan penetapan harga
khusus terhadap kayu JPP, dengan
membedakan terhadap kayu jati
konvensional yang ada di Tempat
Pengumpulan Kayu. Penentuan harga kayu jati saat ini
masih dipengaruhi oleh mekanisme pasar
dan belum mencerminkan harga pokok
produksinya. Dari hal tersebut maka perlu
kiranya mengetahui berapa besar biaya yang
dikeluarkan untuk memproduksi kayu JPP
dari kegiatan pengelolaan awal hingga
pemasaran kayunya sampai dengan umur
tertentu. Harga Pokok Produksi kayu JPP
ini bisa berbeda-beda antara KPH satu
dengan yang lain tergantung karakteristik
wilayah dan biaya yang dikeluarkan. Hal ini
sesuai seperti yang dijelaskan oleh Davis •
Dalam hal perencanaan dan penataan
wilayah hutan, perlu dibuat cluster
kayu JPP yang ditujukan untuk
produksi kayu khusus sortimen AI
dan AII untuk memenuhi kebutuhan
kayu di sentra industri kecil dan
menengah. I.
PENDAHULUAN Sifat kayu jati yang banyak diminati
masyarakat sebagai bahan baku bangunan 54 Risalah Kebijakan Pertanian dan Lingkungan Vol. 8 No. 2 Agustus 2021 (1954 dan 1987) dalam Andayani (1998)
bahwa nilai tegakan (stumpage) di masing-
masing lokasi produsen sebenarnya berbeda
(situasional) sehingga tidak dapat ditetapkan
secara sama (satu harga). Perbedaan
tersebut antara lain terletak pada beberapa
parameter
dan
faktor-faktor
yang
mempengaruhi nilai tegakan yaitu daur
ekonomi, keadaan hutan, geografis hutan
dan komponen biaya pembentukannya. Penelitian ini bertujuan untuk mengetahui
harga pokok produksi kayu JPP hasil
penjarangan melalui pendekatan biaya
selama daur dan pendekatan pasar (market
value). penjarangan pada petak lain, sebagaimana
disajikan pada Tabel 1. Tabel 1. Hasil penjarangan perhutanan klon
JPP tahun 2017
No Lokasi Realisasi
(m3)
Produktivitas
(m3)
Keterangan
1. KPH
Ngawi
143,49
30,53
Tahun
tanam 2004
2. KPH
Tuban
134,17
11,37
3. KPH
Cepu
214,12
25,49 Tabel 1. Hasil penjarangan perhutanan klon
JPP tahun 2017 Kayu hasil penjarangan di atas di
dominasi oleh sortimen AI, kelas diameter
16 – 19 cm, mutu T, panjang 2 meter up. Harga rata-rata berdasarkan harga jual dasar
yang ditetapkan oleh Perum Perhutani
tahun 2018 adalah sebesar Rp. 1.225.000,-
untuk sortimen AI dan Rp. 2.100.000,-
untuk sortimen AII (mutu T, kelas diameter
16 – 19 cm dan 24 – 26 cm, panjang 2,00 –
2,90 meter). II.
SITUASI TERKINI Jati
Plus
Perhutani
mulai
dikembangkan secara luas adalah JPP PHT
1 dan PHT 2 mulai tahun 2007. Sampai
dengan tahun 2017, sudah ditanam seluas ±
200.000 ha, di seluruh wilayah Perum
Perhutani, sedangkan yang dikelola oleh
Puslitbang seluas 446,7 ha. Jati Plus
Perhutani klon PHT 1 dan PHT 2 juga telah
mendapatkan hak perlindungan varietas
tanaman dari Kementerian Pertanian tahun
2009. Uji coba penebangan penjarangan Jati
Plus Perhutani dilakukan pada tahun 2013,
di petak 61a, RPH Kedunggalar, BKPH
Kedunggalar,
KPH
Ngawi
dan
menghasilkan ume kayu sebesar 150,5
m3/ha,
jumlah
pohon
rata-rata
597
pohon/ha dengan diameter rata-rata 23,1
cm (Wibowo, 2013). Hasil uji coba III. METODOLOGI Penelitian dilakukan tahun 2017, di
wilayah Kesatuan Pemangkuan Hutan
(KPH) Ngawi, Divisi Regional Jawa Timur. Harga Pokok Produksi ditentukan dengan
melalui pendekatan nilai tegakan (stumpage
value) dalam Rp/ha. Mekanisme analisis
menggunakan metode perhitungan biaya
tetap yang dikeluarkan selama daur. Data
sekunder yang digunakan adalah laporan
keuangan unit kerja pendukung yaitu
Kantor Pusat, Kantor Divisi Regional,
Kantor Puslitbang, Kantor Pusdikbang,
Kantor Biro Perencanaan, dan Kantor
KBM Penjualan. 55 Rumusan Kajian Strategis Bidang Pertanian dan Lingkungan
ISSN : 2355 – 6226
E-ISSN : 2477 – 0299 Rumusan Kajian Strategis Bidang Pertanian dan Lingkungan ISSN : 2355 – 6226
E-ISSN : 2477 – 0299 Seluruh biaya pembentukan tegakan
JPP dibawa ke satuan hektar (Rp/ha) dan
dihitung dengan biaya yang dikeluarkan
selama daur tanaman JPP yang sudah
dilakukan penebangan yaitu umur 9 tahun. Kaitannya dengan konsep joint cost, besarnya
biaya yang dialokasikan untuk JPP dihitung
dengan pendekatan luas penggunaan lahan
untuk tanaman jati. Selanjutnya Untuk
mendapatkan nilai harga pokok produksi
kayu
bundar
tiap
sortimen,
hasil
penghitungan
biaya
tegakan
sebelum
ditebang ditambah dengan biaya eksploitasi,
biaya pemasaran (Rp/m3) dan Provisi
Sumber
Daya
Hutan
tiap
sortimen
(Anonim, 2009). pengolahan dan pemasaran hasil hutan. Perhitungan
untuk
mengetahui
biaya
pembangunan tegakan dilakukan dengan
analisis finansial menggunakan compounding
factor yang disusun dalam bentuk cashflow
selama daur dikaitkan dengan premi
produksi
kayu
yang
dihasilkan
saat
pemanenan hasil hutan. Analisis finansial dilakukan dengan
menghimpun biaya per masing-masing
kegiatan yang dikeluarkan, berdasarkan
pada laporan keuangan di KPH Ngawi. Biaya yang dikeluarkan dibagi dalam 3
kategori yaitu biaya investasi langsung, biaya
prasarana fisik dan biaya rutin. Biaya
investasi
langsung
terdiri
dari
biaya
perencanaan keluar setiap 10 tahun sekali,
biaya persemaian dan biaya tanaman, keluar
1 kali selama daur, biaya pemeliharaan,
keluar mulai tahun pertama sampai tahun ke
empat, dan biaya perlindungan yang
dikeluarkan setiap tahun sampai dengan
umur tebang. Data primer yang digunakan untuk
mengetahui preferensi konsumen terhadap
harga kayu JPP diperoleh dari hasil
wawancara
terhadap
pembeli
yang
melakukan pembelian kayu JPP di wilayah
KBM penjualan Madiun sebanyak 30
responden. Pengambilan sampel responden
dilakukan dengan sistematik random sampling
dengan
pembagian
tipe
pembeli
berdasarkan jenis pembeli perorangan,
perusahaan dan trader/agen. Biaya prasarana fisik terdiri atas biaya
pemeliharaan
dan
penyusutan
sarana
prasarana yaitu kendaraan, tanah dan
bangunan, peralatan kantor dan bengkel
instalasi. Biaya rutin terdiri atas biaya yang
dikeluarkan secara rutin setiap tahun sampai
dengan waktu tebang, yang terdiri atas biaya
pajak, biaya umum dan administrasi, biaya
gaji, premi produksi dan biaya pendidikan
dan penyuluhan. IV. ANALISIS DAN ALTERNATIF
SOLUSI Perhitungan HPP Kayu JPP KPH Ngawi Tahun 2017
Biaya
selama daur
Umur
tebang
HPP
20.024.309
9 tahun
AI
AII
AIII
Harga Pokok Produksi di
tegakan
123.987
272.928
-
Biaya eksploitasi
250.000
250.000
-
Biaya pemasaran
59.043
59.043
-
PSDH
72.000
114.000
-
Biaya pengujian
9.000
9.000
Sharing PHBM (25%)
128.508
176.243
Jumlah
642.538
881.214
- Jumlah biaya rutin, prasarana fisik dan biaya
unit kerja penunjang yang lebih kecil
daripada biaya investasi langsung juga
disebabkan karena dalam tebangan JPP ini
dilakukan pada umur 9 tahun. Tabel 2. Perhitungan HPP Kayu JPP KPH Ngawi Tahun 2017 Biaya
selama daur
Umur
tebang
HPP
20.024.309
9 tahun
AI
AII
AIII
Harga Pokok Produksi di
tegakan
123.987
272.928
-
Biaya eksploitasi
250.000
250.000
-
Biaya pemasaran
59.043
59.043
-
PSDH
72.000
114.000
-
Biaya pengujian
9.000
9.000
Sharing PHBM (25%)
128.508
176.243
Jumlah
642.538
881.214
- rakyat. Hal ini karena anggapan konsumen
bahwa kayu jati JPP secara mutu hampir
sama dengan kayu rakyat. Biaya pembangunan tegakan yang
terbentuk untuk tegakan tanaman JPP umur
9 tahun adalah Rp. 20.024.309,-/ha. Untuk
mengetahui biaya produksi maka biaya
pembangunan tegakan ini dibagi dengan
realisasi produksi kayu JPP yang dihasilkan. Perincian perhitungan ditunjukkan pada
Tabel 2. Tabel 3. Prosentase penurunan harga kayu JPP
di wilayah KBM Jawa Timur
Unit
kerja
Prosentase Penurunan Harga
Berdasarkan Keinginan Konsumen
AI (%)
AII (%)
AIII (%)
Jawa
Timur
15,5 ± 2,34
20,6 ± 2,74
- Tabel 3. Prosentase penurunan harga kayu JPP
di wilayah KBM Jawa Timur
Unit
kerja
Prosentase Penurunan Harga
Berdasarkan Keinginan Konsumen
AI (%)
AII (%)
AIII (%)
Jawa
Timur
15,5 ± 2,34
20,6 ± 2,74
- Tabel 3. Prosentase penurunan harga kayu JPP
di wilayah KBM Jawa Timur
Unit
kerja
Prosentase Penurunan Harga
Berdasarkan Keinginan Konsumen
AI (%)
AII (%)
AIII (%)
Jawa
Timur
15,5 ± 2,34
20,6 ± 2,74
- Tabel 3. Prosentase penurunan harga kayu JPP
di wilayah KBM Jawa Timur Harga Pokok Produksi Kayu JPP di
KPH Ngawi hasil tebangan petak 61a,
berdasarkan perhitungan diperoleh biaya
produksi kayu AI sebesar Rp. 642.538,- dan
sortimen AII sebesar Rp. 881.214,-. Nilai ini
merupakan harga dimana masih merupakan
titik impas, sehingga belum ada keuntungan,
sehingga perlu ditambah dengan profit margin
yang diinginkan. Dari hasil ini maka dapat memberikan
gambaran bahwa harga kayu JPP masih
perlu dikaji untuk diturunkan sampai
dengan batas tertentu di mana pengusaha/
konsumen
masih
mampu
mendapat
membeli dengan perbandingan asal kayu
rakyat. IV. ANALISIS DAN ALTERNATIF
SOLUSI Komponen
biaya
pembangunan
suatu hutan tanaman industri secara garis
besar
meliputi
kegiatan
perencanaan,
penanaman,
penebangan
kayu
atau
pemungutan,
pemeliharaan
hutan, 56 Risalah Kebijakan Pertanian dan Lingkungan
Vol. 8 No. 2 Agustus 2021 Risalah Kebijakan Pertanian dan Lingkungan Risalah Kebijakan Pertanian dan Lingkungan Vol. 8 No. 2 Agustus 2021 Biaya prasarana fisik dan biaya rutin
merupakan biaya bersama atau joint cost. Joint
cost adalah biaya yang dikeluarkan untuk d
b y
y
g d
Gambar 1. Persentase biaya produksi kayu JPP di KPH Ngawi Gambar 1. Persentase biaya produksi kayu JPP di KPH Ngawi menghasilkan dua macam produk atau lebih
yang berlainan. Dalam perhitungan beban
biaya untuk tanaman JPP di KPH Ngawi,
perhitungan joint cost didasarkan pada
kontribusi luas tanaman JPP di KPH Ngawi
terhadap luas hutan produksi di KPH
Ngawi. unit kerja dihitung berdasarkan kontribusi
luas tanaman JPP dibandingkan dengan luas
hutan produksi jati di masing-masing unit
kerja. Bila dilihat besarnya komponen-
komponen penyusun biaya produksi kayu
JPP, biaya paling tinggi yaitu di komponen
biaya investasi langsung yang terdiri dari
biaya penyusunan RPKH, biaya persemaian,
biaya tanaman, biaya pemeliharaan dan
biaya perlindungan. Untuk lebih jelasnya
dalam Gambar 1. Biaya produksi kayu JPP juga
merupakan bagian dari biaya prasarana fisik
dan rutin yang dikeluarkan oleh unit kerja
yang mendukung kegiatan pengelolaan
sumber daya hutan di KPH yaitu Kantor
Pusat,
Puslitbang,
Pusdikbang,
Divisi
Regional Jawa Timur, biro perencanaan dan
divisi komersial kayu. Biaya unit kerja
pendukung ini juga merupakan joint cost
dengan komoditi lainnya. Pembebanan
biaya yang dikeluarkan oleh masing-masing Distribusi
biaya
pembangunan
tegakan JPP dimana komposisi biaya
investasi langsung yang berkaitan erat
dengan kegiatan pengelolaan tanaman JPP
sudah cukup baik, karena hampir sebagian
besar biaya digunakan untuk tanaman JPP. 57 57 Rumusan Kajian Strategis Bidang Pertanian dan Lingkungan Jumlah biaya rutin, prasarana fisik dan biaya
unit kerja penunjang yang lebih kecil
daripada biaya investasi langsung juga
disebabkan karena dalam tebangan JPP ini
dilakukan pada umur 9 tahun. Tabel 2. IV. ANALISIS DAN ALTERNATIF
SOLUSI Standar deviasi yang masih relatif
kecil disebabkan karena variasi responden
tidak terlalu besar, hanya dikerucutkan
kepada pembeli kayu JPP yang merupakan
segmen pembeli menengah sampai dengan
kecil. Untuk
mengetahui
harga
beli
maksimum yang masih bisa diterima oleh Hasil survei responden di wilayah
penjualan Jawa Timur, menunjukkan hasil
bahwa konsumen kayu menginginkan
penurunan harga terhadap kayu JPP agar
minimal tidak berbeda jauh dengan kayu 58 Vol. 8 No. 2 Agustus 2021 Risalah Kebijakan Pertanian dan Lingkungan pembeli secara riil maka nilai persentase
kenaikan harga tersebut dikalikan dengan
harga jual dasar pada setiap kelas diameter,
kelas panjang dan kelas mutu yang
ditetapkan. Asri, M. 1991. Marketing. Unit Penerbit dan
Percetakan
AMP
YKPN. Yogyakarta. Priyanto, R. 2004. Analisis Distribusi Kayu
Jati
Rakyat
di
Kabupaten
Gunungkidul. Skripsi S1. Fakultas
Kehutanan. Universitas
Gadjah
Mada. Yogyakarta. Sofyan, P., Warsito. 1995. Penaksiran Biaya
dan
Pendapatan
Pengusahaan
Hutan Tanaman. Bahan Kuliah
Analisis Bisnis Pengusahaan Hutan
Program Pasca Sarjana. Universitas
Gadjah Mada. Yogyakarta. REFERENSI Andayani, W. 1998. Sistem Distribusi dan
Penetapan Harga Kayu Bulat Jati di
Jawa. Disertasi S3 Program Pasca
Sarjana. Institut Pertanian Bogor. Bogor. Wibowo, A. 2013. Uji Coba Tebangan Kayu
JPP Asal Stek Pucuk (Studi kasus
petak 61a, BKPH Kedunggalar,
KPH
Ngawi). Laporan
Pengamatan. Cepu. Cepu. Anonim. 2009. Analisis Finansial Penerapan
Harga
Pokok
Produksi
Kayu
Bundar Jati Perhutani. Laporan
Penelitian. Puslitbang
Perum
Perhutani Cepu. Cepu. 59 59
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https://openalex.org/W3187247888
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http://ijec.ejournal.id/index.php/counseling/article/download/166/134
|
Indonesian
| null |
Konseling Religi dalam Upaya Menemukan Kebermaknaan Hidup Remaja Korban Broken Home
|
Indonesian journal of educational counseling
| 2,021
|
cc-by
| 5,151
|
1 Corresponding author’s address: Program Studi Interdisciplinary Islamic Studies, Konsentrasi Bimbingan
Konseling Islam Pascasarjana, UIN Sunan Kalijaga Yogyakarta, Indonesia. E-mail: 2020001107@student.uin-
suka.ac.id ABSTRACT RELIGIOUS COUNSELING IN AN EFFFORT TO FIND A MEANINGFULL
LIFE FOR TEENAGER VICTIMS OF BROKEN HOME. The purpose of this
research is to explain the phenomenon of teenage victims of broken home in
their efforts to find meaning in life through religious counseling services. The
type of this research is phenomenological qualitative. Data collection
techniques were carried out by observation and interviews. The data analysis
technique used intra-case and inter-case analysis. The results showed that the
reasons for these adolescents to attend religious counseling in order to find
meaning in their lives included; forgetting the disappointments they face,
finding wisdom from the problems that occur, finding meaning in life, and
preparing for a new life. Religious counseling is carried out through several
activities such as reading the Koran, listening to lectures, reading Islamic and
motivational books. Received: 28.05.2021
Received in revised form:
01.07.2021
Accepted: 08.07.2021
Available online: 30.07.2021 KEYWORDS:1Broken Home, Counseling, Meaningfull Life, Religion, Teenager. KEYWORDS:1Broken Home, Counseling, Meaningfull Life, Religion, Teenager. DOI: 10.30653/001.202152.166 This is an open access article distributed under the terms of the Creative Commons
Attribution 4.0 International License, which permits unrestricted use, distribution, and
reproduction in any medium, provided the original work is properly cited. © 2021 Rois Nafi'ul Umam, Maemonah. Indonesian Journal of Educational Counseling is published by
FKIP of Universitas Mathla’ul Anwar Banten Indonesian Journal of Educational Counseling, 2021, Vol. 5 No. 2, Page: 64-74
Open Access | Url: http://ijec.ejournal.id/index.php/counseling/article/view/166 Indonesian Journal of Educational Counseling, 2021, Vol. 5 No. 2, Page: 64-74
Open Access | Url: http://ijec.ejournal.id/index.php/counseling/article/view/166 http://ijec.ejournal.id PENDAHULUAN Masa remaja merupakan bagian dari transisi perjalanan hidup manusia dari masa
kanak-kanak menuju masa dewasa (Diananda, 2019). Dalam masa transisi tersebut, terjadi
perubahan secara mendasar pada sisi jasmani maupun rohani remaja. Menurut Rice,
remaja dapat diartikan dengan masa peralihan individu yang tumbuh dari masa kanak-
kana menuju masa dewasa yang penuh dengan kematangan, baik secara fisik maupun
psikis (Saputro, 2018). Secara fisik, masa remaja dapat ditandai dengan beberapa ciri
seperti mulai matangnya organ reproduksi, mengalami menstruasi/mimpi basah,
perubahan suara menjadi lebih berat dan munculnya jerawat pada area wajah. Adapun
secara psikis, remaja dapat ditandai dengan ciri yang diantaranya memiliki sikap agresif,
senang mencoba hal-hal baru, mengalami emosi yang tidak stabil, mulai mencari identitas
diri serta memiliki perasaan yang sensitif (Jannah, 2017). Secara psikologis, masa remaja Indonesian Journal of Educational Counseling is published by
FKIP of Universitas Mathla’ul Anwar Banten 64 Rois Nafi'ul Umam, Maemonah dapat dikatakan sebagai masa yang labil, yaitu mereka belum sepenuhnya dapat
mengendalikan dirinya sendiri secara emosi dan hanya mengikuti naluri dan
keingintahuannya tanpa memikirkan dampak yang akan terjadi setelah melakukan hal
tersebut. Oleh karenanya dalam masa perkembangannya, remaja perlu untuk dibimbing
dan diarahkan oleh orang tuanya agar terhindar dari sikap dan perilaku yang buruk tanpa
membatasi dirinya untuk bisa maju dan berkembang (Verawati, 2017). Salah satu faktor yang berperan penting dalam perkembangan remaja adalah orang
tua. Sosok orang tua yang telah melahirkan dan membesarkannya dengan sepenuh hati
menjadi pihak yang dapat menentukan perilaku atau arah masa depan yang hendak
dicapai oleh remaja tersebut. Keluarga yang bahagia dan harmonis merupakan keluarga
yang diidamkan oleh remaja, karena ia dapat merasakan kasih sayang dan perhatian yang
selayaknya ia dapatkan dari orang tuanya (Muniriyanto & Suharnan, 2014). Selain itu
dalam keluarga yang bahagia dan harmonis, remaja tersebut diperlakukan dengan
selayaknya seorang anak yang mendapatkan bimbingan dan pengasuhan yang baik dari
orang tua. Untuk mencapai kebahagiaan dan keharmonisan dalam keluarga, setiap
anggota keluarga memiliki peran penting untuk mewujudkan hal tersebut, baik itu ayah,
ibu maupun anak-anaknya (Arintina & Fauziah, 2015). Peran tersebut dijalankan
sebagaimana kedudukan mereka dalam rumah tangga, ayah yang berusaha untuk
mencari penghidupan yang layak, seorang ibu yang mengurus rumah tangga dan
membimbing anak-anaknya, serta anak-anak yang menjalankan tugas dan tanggung
jawab yang diberikan dengan baik dan membahagiakan kedua orang tuanya. PENDAHULUAN Melalui
keluarga yang bahagia dan harmonis, setiap anggota keluarga di dalamnya akan
merasakan damai dan penuh cinta kasih antar sesama anggota keluarga serta dapat
mencapai tujuan hidup yang telah ditetapkan oleh masing-masing anggota keluarga
(Isminayah & Supandi, 2016). Akan tetapi tidak semua keluarga berada pada kondisi yang diliputi dengan
kebahagiaan dan keharmonisan. Dalam keluarga tersebut terjadi konflik internal yang
menyebabkan renggangnya hubungan antar anggota keluarga. Konflik tersebut dapat
diakibatkan oleh permasalahan dalam keluarga yang tidak terselesaikan dengan baik
(Isminayah & Supandi, 2016). Dari konflik internal dalam keluarga tersebut, dapat
menghantarkan keluarga tersebut pada perceraian akibat masalah-masalah yang ada
tersebut tidak menemui titik temu solusinya. Akibat dari perceraian tersebut, remaja akan
kehilangan salah satu sosok dari orang tuanya karena mereka akan tinggal dengan salah
satu diantara keduanya (Dewanti & Ediati, 2016). Secara psikis, remaja tersebut merasa
kehilangan orang yang penting dalam hidupnya dan merasakan dampak negatif dari
perceraian tersebut. Akibat dari broken home yang terjadi, remaja tersebut akan
mengalami gangguan psikologis, seperti merasa insecure, perasaan takut menjadi
penyebab perceraian yang dilakukan oleh orang tuanya, merasa dirinya tidak berharga
dan depresi yang dapat berujung pada bunuh diri (Ramadhani & Krisnani, 2019). Remaja
yang menjadi korban broken home tersebut dapat kehilangan motivasi untuk hidup serta
mencari pelarian lain untuk mengekspresikan perasaannya dengan melakukan perilaku
yang merugikan dirinya sendiri atau orang lain. Fenomena remaja yang menjadi korban dari broken home dan berdampak secara
psikologis terhadap pribadi mereka, menjadi sebuah fenomena yang amat perlu untuk
dikaji secara mendalam berkaitan dengan kebermaknaan hidup dari remaja tersebut
(Wulandri & Fauziah, 2019). Hal ini dikarenakan para remaja yang menjadi korban broken
home tersebut merasa hidupnya hampa, kosong dan tidak bermakna karena kehilangan
salah satu sosok orang tua yang selama ini telah mendampinginya dan membimbingnya 65 INDONESIAN JOURNAL OF EDUCATIONAL COUNSELING 2021, 5(1), 64-74 dari kecil. Kehampaan dan ketidak berhargaan dalam diri dapat ditunjukkan dalam
bentuk hilangnya semangat untuk menjalani hidup, munculnya perasaan yang absurd,
merasa tidak berdaya dan putus asa (Aziz, 2015). Dalam diri remaja tersebut, ia berusaha
untuk tetap kuat untuk menghadapi kondisi yang tidak ia harapkan tersebut dan
berusaha mencari nilai positif dari kejadian tersebut. Selain itu remaja tersebut akan
mencari makna dari setiap perjalanan hidupnya. Akan tetapi tidak semua remaja yang
menjadi korban broken home memiliki kemampuan dan menjalani usaha tersebut
sehingga ia larut dalam permasalahan yang dapat berdampak secara psikologis terhadap
pribadinya tersebut (Hafiza & Mawarpury, 2018). PENDAHULUAN Kebermaknaan hidup merupakan sebuah nilai yang mendorong terbentuknya
motivasi yang kuat untuk dapat menjalani hidupnya dengan baik, melakukan kegiatan
yang berguna serta memberi manfaat kepada diri sendiri dan orang lain. Kebermaknaan
hidup dapat dicapai apabila mencapai beberapa indikator meliputi menemukan arti
dalam kehidupan yang cocok dengan dirinya, mampu memberi dan menerima cinta serta
memiliki tekad yang kuat untuk menjalani hidup (Siddik, Oclaudya, Ramiza, & Nashori,
2018). Upaya untuk menemukan keberhargan diri dapat dimaknai dengan mengubah
kondisi hidup yang tidak bermakna dan mengarahkannya pada hidup yang bermakna. Untuk mencapai kebermaknaan hidup, remaja harus melewati beberapa tahap penting
yang diantaranya tahap derita, tahap penerimaan diri, tahap pemaknaan hidup, tahap
realisasi makna dan tahap kehidupan bermakna. Kebermaknaan hidup individu dapat
dipengaruhi oleh dua faktor yaitu secara internal maupun eksternal. Faktor internal
meliputi pola berpikir, sikap, konsep diri, kepercayaan dan kepribadian. Adapun faktor
eksternal diantaranya adalah pekerjaan, pengalaman, hubungan dalam keluarga dan
kebudayaan lokal (Mazaya & Supradewi, 2020). Dalam konteks remaja yang menjadi
korban broken home, hubungan antar anggota keluarga yang tidak harmonis dan
berujung pada perceraian membuat remaja korban broken home tidak memiliki
kebermaknaan dalam hidup, merasa hampa dan tidak semangat dalam menjalani hidup. Para remaja tersebut menghendaki kehidupan yang berjalan dengan semestinya, akan
tetapi hal tersebut tidak ia dapatkan ketika remaja tersebut menghadapi kondisi yang
tidak ideal yaitu broken home. Dalam penelitian lainnya yang dilakukan oleh Deviana Citra Dewi Widyawati
menunjukkan bahwa tidak semua remaja larut dalam kesedihan dan menjadikan dirinya
hidup dengan ketidakbermaknaan ketika mengalami broken home. Mayoritas remaja
yang menjadi subjek penelitian tersebut memiliki makna hidup yang baik dan tetap
semangat dalam menjalani hidupnya (Widyawati, 2009). Hal ini berbanding terbalik
dengan penelitian yang dilakukan oleh Yuniar Asih yang menyatakan bahwa
kebermaknaan hidup bagi remaja broken home hanya dirasakan oleh sedikit remaja dan
lebih banyak remaja yang merasakan ketidak bermaknaan dalam hidupnya (Mustikarani,
2013). Adapun perbedaan dengan beberapa penelitian diatas, penelitian yang dilakukan
oleh peneliti berfokus pada upaya menemukan keberhargaan diri bagi remaja korban
broken home melalui konseling religi yang dilaksanakan di rumah konseling mutiara hati. Dalam hal ini remaja yang menjadi subjek penelitian akan berusaha untuk menemukan
keberhargaa melalui tahapan-tahapan dalam konseling religi yang dilaksanakan di
rumah konseling mutiara hati. Konseling religi merupakan proses pemberian bantuan yang dilakukan oleh
konselor kepada konseli yang sedang mengalami permasalahan agar konseli tersebut
dapat menyelesaikan masalah yang sedang dihadapinya. METODE Jenis penelitian yang digunakan adalah penelitian lapangan (field research) yang
dilaksanakan di rumah konseling keluarga mutiara hati. Adapun metode yang digunakan
dalam penelitian ini adalah kualitatif fenomenologis. Hal ini dikarenakan peneliti
berfokus untuk mengamati dan menggambarkan dari kebermaknaan hidup yang
dikehendaki oleh remaja korban broken home melalui pendekatan kualitatif
fenomenologi ini. Teknik pengumpulan data dilakukan dengan observasi dan wawancara
secara singkat terhadap subjek penelitian ini yaitu remaja korban broken home. Melalui
teknik pengumpulan data tersebut, peneliti dapat mengetahui riwayat pribadi dan
permasalahan yang dihadapinya, serta proses yang dihadapi oleh subjek penelitian untuk
menemukan kebermaknaan hidup. Jumlah subjek dalam penelitian ini sebanyak 2 orang,
yaitu konselor berjumlah 1 orang dan konseli sebanyak 1 orang. Teknik analisis data
dilakukan dengan teknik analisis intra kasus dan antar kasus untuk memperoleh
gambaran umum mengenai kehidupan pribadi subjek penelitian, permasalahan yang
dialami oleh subjek penelitian, dan upaya dalam menemukan kebermaknaan hidup
melalui konseling religi yang diikutinya. PENDAHULUAN Dalam konseling religi, nilai-
nilai dasar dalam agama dan kepercayaan akan diserap dan diadaptasi dalam layanan 66 Rois Nafi'ul Umam, Maemonah konseling, sehingga dapat membentuk nilai yang dapat dipahami oleh konseli dan dapat
mengembangkan dirinya berlandaskan nilai-nilai moral keagamaan (Maulana &
Hernisawati, 2017). Melalui konseling religi, diharapkan konseli dapat mengembangkan
potensi diri, kemampuan berpikir, keimanan dan ketakwaan untuk mencapai pribadi
yang baik berlandaskan pada nilai-nilai keagamaan. Dari latar belakang di atas, penelitian ini akan berfokus pada alasan dari remaja yang
menjadi korban broken home tersebut untuk mengikuti konseling religi dalam rangka
menemukan kebermaknaan hidup dan proses yang dilalui dalam konseling religi untuk
mencapai kebermaknaan hidup. Adapun rumusan masalah dalam penelitian ini adalah
apa alasan dari remaja tersebut untuk mengikuti konseling religi dalam mencapai
kebermaknaan hidup? Bagaimana proses dalam konseling religi untuk mencapai
kebermaknaan hidup? HASIL DAN PEMBAHASAN Broken home merupakan retaknya rumah tangga/keluarga yang telah terbangun
yang terjadi akibat anggota keluarga yang tidak dapat menjalankan peran sosial atau
menjalankan kewajibannya. Broken home terjadi dalam bentuk tanpa hadirnya salah satu
dari orang tua, baik ayah ataupun ibu akibat dari perceraian atau meninggalkan rumah. Broken home dalam keluarga dapat terjadi akibat pertengkaran antar anggota keluarga
dan tidak adanya sikap dan bentuk kasih sayang dalam lingkungan keluarga (Novianto,
Zakso, & Salim, 2017). Keluarga yang mengalami broken home dapat diidentifikasi
dengan beberapa ciri yang diantaranya adalah orang tua bercerai, hubungan antara ayah
dan ibu atau antara orangtua dengan anak yang kurang baik, serta suasana dalam rumah
yang tegang karena penuh dengan masalah serta tidak ada upaya untuk bersama-sama
menyelesaikan masalah tersebut (Muttaqin & Sulistyo, 2019). Kondisi keluarga yang
mengalami broken home yang diliputi dengan ketegangan dan konflik antar anggota
keluarga berdampak secara psikologis terhadap anggota keluarga lainnnya. Salah satu
yang merasakan dampak langsung secara psikologis dari kasus broken home adalah
anak/remaja. Seorang remaja yang seharusnya mendapatkan arahan dan bimbingan dari 67 INDONESIAN JOURNAL OF EDUCATIONAL COUNSELING 2021, 5(1), 64-74 orang tuanya, justru dihadapkan dengan kasus broken home yang terjadi pada
keluarganya. Dampak negatif yang dirasakan oleh remaja dari kasus broken home adalah
menjadi tidak percaya diri, memiliki emosi yang tidak stabil, rentan menjadikannya
berperilaku agresif, kehilangan motivasi dan tujuan hidup serta tidak menemukan
kebermaknaan dalam hidupnya. Jika hal ini tidak ditangani dengan cepat dan tepat, maka
sikap dan perilaku ini akan terbawa hingga dewasa dan membuatnya menjadi individu
yang bersikap temperamental serta mencari pelampiasan dengan cara yang buruk (Fitria
& Ifdil, 2020). Salah seorang remaja yang menjadi subjek dalam penelitian ini merasakan hal-hal
yang dimaksud diatas seperti tidak menemukan kebermaknaan dalam hidupnya. Keluarga dari remaja tersebut baru saja mengalami broken home yang berujung pada
perceraian. Akibat dari perceraian tersebut, remaja itu hanya tinggal dengan salah
seorang dari orang tuanya dan kehilangan salah satu sosok orang tua yang selama ini
telah membimbing dan mengarahkannya dalam perjalanan hidupnya. Remaja tersebut
merasa tidak percaya diri dengan kondisinya saat ini, merasa tidak bersemangat untuk
menjalani hidup, kehilangan kebermaknaan dalam hidup serta berpikir untuk
mengakhiri hidup. Dari wawancara yang dilakukan oleh peneliti dengan subjek
penelitian, diketahui bahwa remaja tersebut mengikuti konseling religi atas rekomendasi
dari pihak keluarga agar remaja tersebut kembali pulih secara psikologis dan tidak lagi
trauma dengan kasus broken home yang terjadi. HASIL DAN PEMBAHASAN Menurut subjek penelitian, kesediaannya
untuk mengikuti konseling religi di rumah konseling mutiara hati didasarkan pada
keinginan untuk mengalihkan kesedihan yang dirasakannya menjadi pribadi yang lebih
baik. Berikut akan dijelaskan alasan kesediaan dari remaja tersebut untuk mengikuti
konseling religi yang dilaksanakan di rumah konseling mutiara hati. Menemukan Hikmah dari Masalah yang Terjadi Kasus broken home yang terjadi pada keluarga subjek, menimbulkan dampak secara
psikologis terhadap subjek. Subjek merasa bahwa hidupnya telah berubah dan tak akan
lagi sama seperti sebelum broken home terjadi. Menurut subjek, ia selalu berpikir negatif
atas kasus broken home yang terjadi, baik itu penyebab dari perceraian yang dilakukan
orang tuanya maupun masa depannya setelah kasus broken home ini terjadi. Setelah
beberapa waktu, ia menyadari bahwa di balik sebuah kejadian yang tidak menyenangkan,
terdapat hikmah atau nilai positif yang dapat diambil. Untuk dapat menemukan hikmah
dari kejadian tersebut, ia memutuskan untuk mengikuti konseling religi tersebut. Berikut
adalah ungkapan dari subjek penelitian: “Ketika orang tua saya memilih untuk bercerai, saya merasa bahwa kehidupan saya
akan berbeda antara sebelum dan sesudah perceraian ini. Di masa lalu saya
merasakan kasih sayang yang sepenuh hati dari kedua orang tua saya, dan
kedepannya mungkin saya tidak merasakan itu lagi, kasih sayang yang saya dapat
tidak akan seperti dulu atau saya mungkin tidak akan merasakan kasih sayang itu
lagi. Terkadang saya juga berpikir negatif tentang penyebab perceraian orang tua
saya, dan saya merasa menjadi penyebab dari perceraian diantara keduanya. Saya
kadang menceritakan ini kepada sahabat yang saya percaya dan dia menyarankan
agar saya berusaha untuk berpikir positif dan mengambil hikmah positif dari
masalah broken home yang terjadi. Setelah ada kesempatan untuk mengikuti
konseling, disitu saya sekaligus mengusahakan untuk mencari hikmah positif dari
kejadian tersebut dengan konselor yang sangat-sangat baik menurut saya. Setelah
mengikuti konseling religi yang dilaksanakan di rumah konseling mutiara hati ini,
dengan bantuan dari konselor, saya udah berdamai dengan masa lalu dan siap untuk
menjalani hidup yang baru di masa yang akan datang. Selain itu saya sudah
menemukan hikmah positif dari kejadian broken home di keluarga saya dan sudah
menerima kejadian dan keputusan itu dengan ikhlas” (IN, wawancara 13 April 2021). Melupakan Kekecewaan yang Dihadapinya Rasa kecewa yang dirasakan oleh subjek penelitian berkenaan dengan kasus broken
home yang terjadi pada keluarganya membuatnya larut dalam suasana yang kesedihan
dan suasana yang tidak menentu. Subjek tersebut belum dapat menerima keputusan yang
diambil oleh orang tuanya yang memutuskan untuk bercerai. Subjek merasakan
kekecewaan yang menurutnya belum pernah ia rasakan sebelumnya. Akan tetapi
kemudian subjek menyadari bahwa ia tidak bisa untuk terus berada dalam kondisi yang
penuh dengan kekecewaan. Subjek tersebut memiliki keinginan untuk melakukan
konseling agar ia dapat melupakan kekecewaan yang dirasakannya selama ini akibat dari
kasus broken home tersebut. Subjek tersebut berkata: “Setelah mengetahui keluarga saya mengalami broken home dan kedua orang tua
memutuskan untuk bercerai, disitu saya langsung kecewa sejadi-jadinya dengan
orang tua saya yang mengambil keputusan seperti itu. Ketika itu saya masih
berharap orang tua saya tidak berpisah, akan tetapi Tuhan berkehendak lain dan
akhirnya keluarga saya hancur saat itu juga. Saya khawatir tidak dapat beradaptasi
ketika tidak lagi tinggal bersama salah satu dari orang tua saya dan saya takut jika
nanti mengalami bullying karena broken home yang terjadi pada keluarga saya. Setelah beberapa lama, saya menyadari bahwa hidup terus berjalan dan saya tidak
bisa secara terus menerus berada dalam kondisi yang seperti ini. Saya ingin menjadi
lebih baik dan saya perlu dukungan untuk menjadi lebih baik dan dukungan itu yang
saya dapatkan di rumah konseling mutiara hati ini” (IN, wawancara 13 April 2021). Menurut subjek tersebut dengan melakukan konseling berbasis religi di rumah
konseling mutiara hati, ia merasa lebih tenang dan berusaha untuk melupakan 68 Rois Nafi'ul Umam, Maemonah kekecewaan yang pernah dirasakannya, sebagaimana yang diungkapkan oleh subjek
berikut: kekecewaan yang pernah dirasakannya, sebagaimana yang diungkapkan oleh subjek
berikut: “saya merasa bahwa konseling religi yang dilaksanakan di rumah konseling mutiara
hati ini saya belajar tentang ikhlas, berpikir lebih dewasa dalam menghadapi
masalah serta dapat membantu mengatasi masalah saya” (IN, wawancara 13 April
2021). “saya merasa bahwa konseling religi yang dilaksanakan di rumah konseling mutiara
hati ini saya belajar tentang ikhlas, berpikir lebih dewasa dalam menghadapi
masalah serta dapat membantu mengatasi masalah saya” (IN, wawancara 13 April
2021). Menyiapkan Diri untuk Kehidupan yang Baru Setelah subjek telah menerima masalah broken home yang terjadi pada keluarganya,
ia berharap dapat menjalani hidup baru dengan baik, tidak lagi terbawa dengan masalah
di masa lalu serta yakin dan optimis bahwa kehidupan berikutnya akan menjadi lebih
baik. Dengan dasar keyakinan tersebut, ia mencoba untuk mengikuti konseling religi
dengan konselor di rumah konseling mutiara hati.Harapannya adalah, konselor tersebut
bisa membantunya untuk menyiapkan diri untuk menjalani kehidupan yang baru dan
suasana yang baru. Berikut adalah pemaparan dari subjek penelitian: “Saya berusaha untuk menerima kejadian tersebut dan berusaha untuk beradaptasi
dengan kehidupan yang baru setelah terjadi perceraian kedua orang tua saya itu. Saya berusaha untuk mencoba menjalani hidup yang baru dan tidak lagi mengingat-
ingat masalah yang pernah terjadi di masa lalu. Akan tetapi rasanya berat untuk
menjalani hal itu, dan saya butuh bantuan dan orang yang dapat mengerti perasaan
saya dan dapat membantu saya untuk merencanakan dan memulai kehidupan yang
baru ini. Oleh karenanya saya kemudian mengikuti konseling disini agar saya
kedepannya dapat menjalani hidup dengan baik dan tanpa ada beban pikiran lagi
perihal broken home yang pernah terjadi” (IN, wawancara 13 April 2021). “Saya berusaha untuk menerima kejadian tersebut dan berusaha untuk beradaptasi
dengan kehidupan yang baru setelah terjadi perceraian kedua orang tua saya itu. Saya berusaha untuk mencoba menjalani hidup yang baru dan tidak lagi mengingat-
ingat masalah yang pernah terjadi di masa lalu. Akan tetapi rasanya berat untuk
menjalani hal itu, dan saya butuh bantuan dan orang yang dapat mengerti perasaan
saya dan dapat membantu saya untuk merencanakan dan memulai kehidupan yang
baru ini. Oleh karenanya saya kemudian mengikuti konseling disini agar saya
kedepannya dapat menjalani hidup dengan baik dan tanpa ada beban pikiran lagi
perihal broken home yang pernah terjadi” (IN, wawancara 13 April 2021). Kebermaknaan hidup sebagai sebuah nilai fundamental bagi setiap manusia dalam
menjalani hidupnya agar manusia dapat menemukan arti dan makna dari setiap
perjalanan hidupnya serta menikmati hidup yang dijalaninya tersebut. Kebermaknaan
dalam hidup dimaknai dengan upaya untuk menjadikan hidup yang lebih berarti dan
bermanfaat, baik untuk diri sendiri maupun lingkungan sekitarnya. Melalui
kebermaknaan hidup tersebut, individu dapat mencapai tujuan dan motivasi hidup,
memberi kebermanfaatan bagi orang lain dan memperoleh kebahagiaan bagi individu
tersebut (Andriyan & Rumtianing, 2020). Menemukan Kebermaknaan Hidup Masalah broken home yang terjadi pada keluarga subjek membuatnya kehilangan
arah, motivasi dan tujuan hidup serta tidak mengetahui arti dan makna dari kehidupan
yang dijalaninya saat ini. Hal ini dikarenakan ketidaksiapan dari subjek dalam
menghadapi masalah yang terjadi dan suasana baru yang dirasakan setelah adanya kasus
broken home tersebut. Subjek kemudian menyadari bahwa ia harus menjadi lebih baik
dan menemukan makna atau arti dalam perjalanan hidupnya. Hal ini dilakukannya agar
subjek tidak terjebak dalam suasana kesedihan yang selama ini telah ia jalani. Menurutnya
melalui kebermaknaan hidup, ia dapat menghayati setiap perjalanan hidup yang dijalani,
mengambil makna dari setiap kejadian, selalu berpikir positif jika ada hal yang tidak
dikehendaki untuk mencapai kebahagiaan dalam hidupnya. Hal inilah yang mendorong
subjek untuk berusaha menemukan kebermaknaan hidupnya melalui konseling religi
yang dilaksanakan di rumah konseling mutiara hati. Ia berharap setelah mengikuti 69 INDONESIAN JOURNAL OF EDUCATIONAL COUNSELING 2021, 5(1), 64-74 konseling religi tersebut, ia dapat menjadi pribadi yang lebih baik, menemukan
kebermaknaan dalam hidupnya, serta memiliki motivasi dan tujuan hidup di masa yang
akan datang. Berikut adalah pemaparan selengkapnya dari subjek penelitian: “Hidup yang bermakna menurut saya yaitu ketika saya dapat menghayati, meresapi
dan mendalami setiap bagian dari perjalanan hidup yang dijalani. Selain itu hidup
kita menjadi lebih bermakna ketika kita dapat mengambil sisi positif dari sebuah
kejadian selalu meyakini bahwa hal tersebut telah diatur oleh yang maha kuasa dan
kita meyakini dibalik hal itu ada nilai-nilai positif yang bisa diambil. Untuk
menemukan kebermaknaan hidup, saya memutuskan untuk mengikuti konseling
religi ini agar hidup dengan bantuan konselor di rumah konseling mutiara hati ini. Setelah mengikuti konseling religi ini, saya jadi mengerti bahwa kebermaknaan
hidup saya dijalankan dengan memberi manfaat untuk diri saya sendiri dan orang
lain, dan ketika saya tidak bisa melakukan itu, rasanya seperti menyia-nyiakan
kesempatan yang mungkin tidak setiap waktu akan datang. Selain itu saya
menyadari bahwa kebermaknaan dalam hidup saya ditunjukkan dengan membuat
rencana dan tujuan yang baik dan sesuai dengan kemampuan saya serta dapat
mengisi waktu yang luang dengan sesuatu yang bermanfaat seperti mempelajari
ilmu agama” (IN, wawancara 13 April 2021). Menyiapkan Diri untuk Kehidupan yang Baru Dalam konteks keagamaan, khususnya dalam
Islam, kebermaknaan hidup dapat dicapai ketika manusia dapat mengetahui arti dari
kehidupannya, menerima atas segala yang telah ditetapkan dan terjadi padanya serta
selalu yakin bahwa dibalik sebuah ujian, akan ada kebahagiaan yang akan diberikan 70 Rois Nafi'ul Umam, Maemonah olehnya sebagai balasan atas kesabaran dan usahanya dalam menjalani ujian yang
diberikan oleh Tuhannya tersebut (Ningsih, Reza, & Uyun, 2017). Untuk dapat mencapai kebermaknaan dalam hidupnya, manusia perlu menyadari
bahwa hidup yang dijalaninya perlu dimaknai secara mendalam dan senantiasa berusaha
untuk selalu berbuat baik dan memberi manfaat. Terkadang manusia menyadari
pentingnya kebermaknaan hidup saat ia berada dalam kondisi yang sulit dan diliputi
dengan permasalahan yang rumit. Saat mengalami hal tersebut, manusia akan berusaha
untuk mencari solusi atas permasalahannya dan juga menemukan kebermaknaan dalam
hidupnya ketika ia merasa tak lagi memiliki motivasi dan tujuan dalam hidupnya
(Hidayat, 2019). Ketika manusia berada dalam kondisi yang sulit tersebut, ia memerlukan
dukungan dan orang lain di sekitarnya yang mampu mengarahkannya untuk
menemukan kebermaknaan dalam hidupnya. Salah satu upaya yang dapat dilakukan
untuk menemukan kebermaknaan dalam hidup adalah dengan layanan konseling. Konseling merupakan pemberian bantuan yang diberikan oleh konselor kepada konseli
dalam rangka untuk membantu penyelesaian masalah yang dihadapi oleh konseli. Dalam
layanan konseling, konseli dapat menyampaikan permasalahan yang dihadapinya dan
konselor membantu mengarahkan konseli tersebut untuk mencari alternatif penyelesaian
masalahnya (Sukma, 2018). Dalam konteks remaja yang menjadi korban broken home diatas, layanan konseling
dilakukan untuk membantu dirinya dalam menyelesaikan masalah dan menemukan
kebermaknaan dalam hidupnya untuk mencapai kebermanfaatan dan kebahagiaan
dalam hidupnya. Adapun jenis konseling yang diikuti oleh subjek penelitian ini yaitu
konseling religi yaitu layanan konseling yang menginternalisasi nilai-nilai religiusitas
dalam keagamaan. Adapun penggunaan konseling religi ini didasarkan pada
kebermaknaan hidup yang dikehendaki tidak terlepas dari nilai-nilai spiritual dan nilai
tersebut menjadi pendukung dalam membantu subjek untuk mencapai kebermaknaan
dalam hidupnya. Layanan konseling religi ini dijalankan dalam empat tahap dan berikut
adalah tahapan-tahapan dari konseling religi: Tahap Awal Dalam tahap awal ini, konselor akan menyambut konseli dan memperkenalkan
dirinya kepada konseli. Setelah itu konselor memperkenalkan layanan konseling secara
umum seperti tujuan dari konseling, asas konseling, aturan-aturan yang harus ditaati
dalam proses konseling serta tujuan yang hendak dicapai dari konseling. Selain itu,
konselor juga memperkenalkan mengenai konseling religi yang dilandasi oleh nilai-nilai
spiritual dalam Alquran. Untuk dapat membangun hubungan yang bersifat kekeluargaan
dan tidak memberikan suasana yang tegang bagi konseli, suasana konseling yang
dibangun bersifat santai dan memberikan rasa tenang bagi konseli untuk dapat
menceritakan permasalahan yang dihadapinya serta dapat mengikuti layanan konseling
dengan perasaan yang nyaman. Setelah terbangun hubungan yang baik antara konselor
dan konseli, kemudian keduanya menegosiasikan kontrak perihal waktu, tugas dan
kerjasama selama proses konseling. Setelah menandatangani kontrak perjanjian, konseli
akan diberi kesempatan untuk menceritakan masalah yang dihadapinya kepada konselor. Setelah konseli menyampaikan masalah yang dihadapinya, konselor kemudian
memperjelas masalah yang dihadapi oleh konseli dan mendefinisikan masalah yang
disampaikan konseli untuk menentukan langkah-langkah yang akan diambil pada tahap
berikutnya. 71 INDONESIAN JOURNAL OF EDUCATIONAL COUNSELING 2021, 5(1), 64-74 Tahap Kegiatan Tahap ini merupakan tahap inti dari layanan konseling religi ini, karena fokus dari
konseling yang dijalankan berada pada tahap ini. Setelah mengetahui dan mendalami
permasalahan yang dihadapi oleh konseli, kemudian konselor menyampaikan rencana
intervensi yang akan dilakukan dalam konseling. Mengingat bahwa konseling yang
dilakukan merupakan konseling religi, maka kegiatan yang dilakukan berlandaskan pada
nilai-nilai dan ajaran agama, khususnya Islam. Adapun pada tahap ini kegiatan yang
dilakukan meliputi membaca Alquran, mendengar ceramah , membaca buku-buku
motivasi dan keislaman serta membuat rencana dan tujuan hidup yang dikehendaki
ketika konseli telah selesai melakukan layanan konseling. Semua kegiatan yang
dilaksanakan tersebut, didampingi konselor untuk memastikan setiap prosesnya dapat
berjalan dengan baik. Pada kegiatan membaca Alquran, konseli dihadapkan dengan ayat-
ayat yang memiliki makna yang mendalam dan motivasi seperti surah yusuf ayat 87,
surah At-Taubah ayat 40, surah Ali Imran ayat 139, surah A-Baqarah ayat 216 dan surah
Ath-Thalaq ayat 2-3 yang kemudian dapat diambil hikmahnya oleh konseli tersebut. Kegiatan berikutnya adalah dengan mendengar ceramah yang disampaikan oleh ustadz,
dalam hal ini ustadz yang menjadi rujukan bagi konseli dalam belajar agama. Konseli
mendengarkan ceramah dengan tema yang sesuai dengan permasalahan yang
dihadapinya, upaya dalam menemukan kebermaknaan hidup serta motivasi bagi dirinya
untuk dapat menjadi pribadi yang lebih baik. Setelah kegiatan membaca Alquran dan
mendengarkan ceramah telah dilakukan, kegiatan berikutnya yang dilakukan pada tahap
ini yaitu konseli dapat membaca buku-buku mengenai motivasi dan keislaman. Adapun
buku-buku yang dibaca sesuai dengan karakteristik konseli dan mengandung isi yang
dapat menguatkan sisi positif konseli dan membantunya dalam mencapai kebermaknaan
hidup serta dapat memberikan motivasi bagi konseli untuk menjadi lebih baik. Kemudian
masuk pada kegiatan terakhir yaitu menuliskan upayanya untuk berdamai dengan masa
lalu, cita-cita tujuan dan arah hidupnya di masa yang akan datang, serta kebermaknaan
hidup yang telah ia temukan dari kegiatan yang telah dilaksanakan sebelumnya, yaitu
membaca alquran, mendengar ceramah dan membaca buku motivasi dan keislaman. SIMPULAN Masalah broken home yang terjadi pada keluarga, dapat berdampak pada anggota
keluarga yang lain, salah satunya adalah remaja dalam keluarga tersebut. Dampak negatif
dari broken home berpengaruh terhadap kondisi psikologis remaja yang sedang berada
pada masa perkembangan, dan membuatnya kehilangan arah, tujuan, motivasi hidup
serta kebermaknaan yang ada dalam hidupnya. Untuk dapat membantu remaja tersebut
dalam menemukan kebermaknaan hidupnya, dapat dilakukan melalui konseling religi
yang dilandasi oleh nilai-nilai keagamaan. Konseling religi diisi dengan kegiatan seperti
membaca Alquran, mendengar ceramah dan membaca buku-buku motivasi dan
keislaman yang dapat membantunya dalam menemukan kebermaknaan hidup dan
upayanya untuk berdamai dengan masalah broken home yang dihadapinya. Setelah
mengikuti konseling religi, remaja tersebut telah menemukan kebermaknaan dalam
hidupnya, menerima dengan lapang dada permasalahan broken home yang terjadi dan
mengambil sisi positifnya serta menyiapkan diri untuk menjalani kehidupan yang baru
dengan rencana yang telah ditetapkan. Tahap Akhir Setelah melalui rangkaian kegiatan yang dilaksanakan pada tahap sebelumnya
dalam proses konseling, konseli merasakan perubahan positif yang ada pada dirinya dan
menemukan kebermaknaan hidup yang dikehendakinya. Perubahan-perubahan tersebut
meliputi kesediaan untuk menerima masalah yang terjadi dan mengambil sisi positif dari
masalah tersebut, menemukan kebermaknaan dalam hidupnya yaitu konseli menyadari
bahwa setiap bagian dari perjalanan hidupnya merupakan sesuatu yang bernilai dan
memiliki makna yang mendalam dan menghayati setiap detik-detik dalam kehidupannya
agar menjadikannya sebagai manusia yang bersyukur, memberi manfaat bagi diri sendiri
dan orang lain. Adapun rencana dalam kehidupan yang baru menurut konseli diisi
dengan kegiatan-kegiatan yang bermanfaat bagi dirinya, memberi manfaat bagi orang
lain serta tidak bertentangan dengan ketentuan Agama. Selain itu, konseli telah
mempersiapkan rencana untuk membahagiakan orang tuanya walaupun salah satu dari
mereka tidak lagi tinggal bersama dengan konseli tersebut. Sebelum konseling diakhiri,
konselor dan konseli menyampaikan evaluasi atas konseling yang telah dilakukan dan
mengatur pertemuan apabila konseli kembali mengalami kendala berhubungan dengan
kebermaknaan dalam hidupnya. 72 Rois Nafi'ul Umam, Maemonah REFERENSI Andriyan, & Rumtianing, I. (2020). Kebermaknaan hidup Lansia (Studi kasus di cabang
UPT Pelayanan Sosial Tresna Werdha Magetan Jawa Timur). Rosyada: Islamic
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berperilaku agresif pada siswa SMK. Jurnal Empati, 4(1), 208-212. Aziz, M. (2015). Perilaku sosial anak remaja korban broken home dalam berbagai
perspektif (Suatu penelitian di SMPN 18 kota Banda Aceh). Jurnal Al-Ijtimaiyyah, 1(1),
30-50. Dewanti, W., & Ediati, A. (2016). Sikap remaja laki-laki dan perempuan terhadap
perceraian: Studi komparasi pada remaja siswa SMA Negeri 6 Semarang. Jurnal
Empati, 5(3), 594-597. Diananda, A. (2019). Psikologi remaja dan permasalahannya. Journal ISTIGHNA, 1(1), 116-
133. Fitria, L., & Ifdil. (2020). Kecemasan remaja pada masa pandemi Covid-19. Jurnal
EDUCATIO: Jurnal Pendidikan Indonesia, 6(1), 1-4. Hafiza, S., & Mawarpury, M. (2018). Pemaknaan kebahagiaan oleh remaja broken home. Psympathic : Jurnal Ilmiah Psikologi, 5(1), 59-66. Hidayat, V. (2019). Kebermaknaan hidup pada mahasiswa semester akhir. Jurnal Psikologi
Integratif, 6(2), 141. Isminayah, A., & Supandi. (2016). Relasi tingkat keharmonisan keluarga dengan konsep
diri remaja. Al-Balagh : Jurnal Dakwah dan Komunikasi, 1(2), 233-245. Jannah, M. (2017). Remaja dan tugas-tugas perkembangannya dalam Islam. Psikoislamedia : Jurnal Psikologi, 1(1), 243-256. Maulana, A., & Hernisawati. (2017). Konseling religi untuk mengembangkan karakter
konseling. Jurnal Islamic Counseling, 1(2), 55-65. 73 INDONESIAN JOURNAL OF EDUCATIONAL COUNSELING 2021, 5(1), 64-74 Mazaya, K. N., & Supradewi, R. (1970). Konsep diri dan kebermaknaan hidup pada remaja
di panti asuhan. Proyeksi, 6(2), 103-112 Mazaya, K. N., & Supradewi, R. (1970). Konsep diri dan kebermaknaan hidup pada remaja
di panti asuhan. Proyeksi, 6(2), 103-112 Muniriyanto, & Suharnan. (2014). Keharmonisan keluarga, konsep diri dan kenakalan
remaja. Persona: Jurnal Psikologi Indonesia, 3(2), 156-164. Mustikarani, A. Y. (2013). Makna hıdup pada mahasıswa yang mengalamı broken home. Skripsi. Bandung: Universitas Pendidikan Indonesia Muttaqin, I., & Sulistyo, B. (2019). Analisis faktor penyebab dan dampak keluarga broken
home. Studi Gender dan Anak, 6, 245-256. Ningsih, D. A., Reza, I. F., & Uyun, M. (2017). Kebermaknaan hidup lansia pemulung yang
beragama Islam di tempat pembuangan akhir (TPA) Sukajaya Kecamatan Sukarame
Palembang. Psikis: Jurnal Psikologi Islami, 3(1), 52-59. Novianto, R., Zakso, A., & Salim, I. (2017). Analisis dampak broken home terhadap minat
belajar siswa SMA Santun Untan Pontıanak. Jurnal Pendidikan dan Pembelajaran
Khatulistiwa, 8(3), 1-8. Ramadhani, P. E., & Krisnani, H. (2019). Analisis dampak perceraian orang tua terhadap
anak remaja. Focus: Jurnal Pekerjaan Sosial, 2(1), 109-119. Saputro, K. Z. (2018). REFERENSI Memahami ciri dan tugas perkembangan masa remaja. Aplikasia:
Jurnal Aplikasi Ilmu-Ilmu Agama, 17(1), 25. Siddik, I. N., Oclaudya, K., Ramiza, K., & Nashori, F. (2018). Hubungan antara ıkhlas dan
dukungan sosial dengan kebermaknaan hidup pada orang dengan HIV/AIDS. Psikoislamedia Jurnal Psikologi, 3(1), 98-114. Sukma, D. (2018). Concept and application group guidance and group counseling base on
Prayitno’s paradigms. Konselor, 7(2), 49-54. Verawati, I. (2017). Dukungan sosial orangtua dalam mengikutsertakan anaknya berlatih
di Krakatau Taekwondo klub Medan. Jurnal Edutech, 3(2), 22-28. Widyawati, D. C. D. (2009). Studi kasus tentang kebermaknaan hidup remaja yang orangtuanya
bercerai. Yogyakarta: Universitas Sanata Dharma. Wulandri, D., & Fauziah, N. (2019). Pengalaman remaja korban broken home (Studi
kualitatif fenomenologis). Empati, 8(1), 1-9. 74 74
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Robustness of STDP to spike timing jitter
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To cite this version: Yihui Cui, Ilya Prokin, Alexandre Mendes, Hugues Berry, Laurent Venance. Robustness of STDP
to spike timing jitter. Scientific Reports, 2018, 8 (1), pp.8139:1-15. 10.1038/s41598-018-26436-y. hal-01788826v2 Robustness of STDP to spike timing jitter
Yihui Cui, Ilya Prokin, Alexandre Mendes, Hugues Berry, Laurent Venance
To cite this version:
Yihui Cui, Ilya Prokin, Alexandre Mendes, Hugues Berry, Laurent Venance. Robustness of STDP
to spike timing jitter. Scientific Reports, 2018, 8 (1), pp.8139:1-15. 10.1038/s41598-018-26436-y.
hal-01788826v2 Yihui Cui, Ilya Prokin, Alexandre Mendes, Hugues Berry, Laurent Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 In Hebbian plasticity, neural circuits adjust their synaptic weights depending on patterned firing. Spike-
timing-dependent plasticity (STDP), a synaptic Hebbian learning rule, relies on the order and timing
of the paired activities in pre- and postsynaptic neurons. Classically, in ex vivo experiments, STDP is
assessed with deterministic (constant) spike timings and time intervals between successive pairings,
thus exhibiting a regularity that differs from biological variability. Hence, STDP emergence from noisy
inputs as occurring in in vivo-like firing remains unresolved. Here, we used noisy STDP pairings where
the spike timing and/or interval between pairings were jittered. We explored with electrophysiology
and mathematical modeling, the impact of jitter on three forms of STDP at corticostriatal synapses:
NMDAR-LTP, endocannabinoid-LTD and endocannabinoid-LTP. We found that NMDAR-LTP was highly
fragile to jitter, whereas endocannabinoid-plasticity appeared more resistant. When the frequency or
number of pairings was increased, NMDAR-LTP became more robust and could be expressed despite
strong jittering. Our results identify endocannabinoid-plasticity as a robust form of STDP, whereas the
sensitivity to jitter of NMDAR-LTP varies with activity frequency. This provides new insights into the
mechanisms at play during the different phases of learning and memory and the emergence of Hebbian
plasticity in in vivo-like activity. Received: 13 February 2018
Accepted: 9 May 2018
Published: xx xx xxxx Long-term changes in synaptic efficacy between neurons are thought to underlie learning and memory1 and can
be assessed with a synaptic Hebbian paradigm such as spike timing-dependent plasticity (STDP)2–8. In STDP,
the occurrence of timing-dependent-long-term potentiation (tLTP) or -depression (tLTD) is the result of three
factors. The two first ones are the two coincident activity patterns on either side of the synapse, which depends on
(1) the relative timing between pre- and postsynaptic spikes (ΔtSTDP)4,8, (2) the dendritic and axonal propagation
delays9, (3) the number of paired spikes (Npairings)10–13, (4) the frequency of the paired spikes (Fpairings)6,7,10,11,14 and
(5) membrane depolarization14,15; the third factor involves in STDP expression mainly refers to neuromodula-
tors16,17 and glial cells18. STDP has been attracting a lot of interest in computational and experimental neurosciences because it relies
on spike correlation and has emerged as a candidate mechanism for experience-dependent changes in neural
circuits, including map plasticity8,19–23. STDP is classically investigated using regular repetitions of the same spike
timing and fixed intervals between successive paired stimulations. HAL Id: hal-01788826
https://inria.hal.science/hal-01788826v2
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abroad, or from public or private research centers. www.nature.com/scientificreports www.nature.com/scientificreports Robustness of STDP to spike timing
jitter
OPEN Robustness of STDP to spike timing
jitter
OPEN Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 A typical experimental protocol consists in
pairing pre- and postsynaptic stimulations with a fixed ΔtSTDP (ranging from −30 to +30 ms for plasticity induc-
tion); ΔtSTDP < 0 when the postsynaptic stimulation occurs before the paired presynaptic one (post-pre pairings),
whereas ΔtSTDP > 0 when the presynaptic stimulation occurs before the postsynaptic one (pre-post pairings). These pairings are then repeated between 15 and 200 times (between 0.1 and 5 Hz) with the spike timing and the
time interval between successive pairings being kept constant. Regular stimulation paradigms produce patterns of activity that are likely to differ from the variability
expected in in vivo-like firing. In vivo paired natural visual stimulations with postsynaptic action potential lead
to Hebbian STDP with specific characteristics when compared to ex vivo STDP (broader membrane potential
changes and STDP window), which can be explained by jittered presynaptic inputs occurring in vivo22. A theoreti-
cal study has started to explore naturalistic stimulations and showed that when neurons fire irregularly, the impact
of spike timing in plasticity becomes weaker than the influence of the firing rate24. However, whether STDP emer-
gence and maintenance is robust against biological variability remains to be investigated. To address this question,
it is important to take into account various forms of STDP, involving distinct intracellular signal transduction 1Dynamics and Pathophysiology of Neuronal Networks Team, Center for Interdisciplinary Research in Biology (CIRB),
College de France, CNRS, INSERM, PSL Research University, Paris, France. 2INRIA, Villeurbanne, France. 3University
of Lyon, LIRIS UMR5205, Villeurbanne, France. Yihui Cui and Ilya Prokin contributed equally to this work. Hugues
Berry and Laurent Venance jointly supervised this work. Correspondence and requests for materials should be
addressed to H.B. (email: hugues.berry@inria.fr) or L.V. (email: laurent.venance@college-de-france.fr) SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 1 www.nature.com/scientificreports/ Figure 1. NMDAR- and eCB-mediated STDP induced with fixed ΔtSTDP. (a) STDP protocol: a single
postsynaptic spike was paired with a single cortical stimulation, 10 or 100 times at 1 Hz. (b) Example of 5
successive pairings (#4–8, from c1) exhibiting relatively fixed ΔtSTDP (#4–8: −8.0, −7.8, −7.4, −8.8 and
−8.0 ms). Plotting the successive ΔtSTDP illustrates their low variance σΔt (2.6 ms). (c) Corticostriatal NMDAR
tLTP induced by 100 post-pre pairings with fixed ΔtSTDP. (c1) Example of tLTP induced by 100 post-pre
pairings (mΔt = −9 ms, σΔt = 2.6 ms). Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 The mean baseline EPSC amplitude, 95 ± 3 pA, increased by 77% to
168 ± 6 pA 45 minutes after pairings. Upper panel, time course of Ri (baseline: 172 ± 2 MΩ and 35–45 min
after pairings: 170 ± 3 MΩ; change of 1%) and holding current (Ihold). Inset: distribution of the 100 ΔtSTDP. (c2)
Averaged time-courses of tLTP induced by 100 post-pre pairings; this tLTP was NMDAR-mediated because
prevented by D-AP5 (50 μM). (c3) Relationship between the STDP magnitude and the STDP jitter for each
of the recorded neurons. (d) Corticostriatal eCB-tLTD induced by 100 pre-post pairings with fixed ΔtSTDP. (d1) Example of tLTD induced by 100 pre-post pairings (mΔt = +21 ms, σΔt = 2.1 ms); the mean baseline Figure 1. NMDAR- and eCB-mediated STDP induced with fixed ΔtSTDP. (a) STDP protocol: a single
postsynaptic spike was paired with a single cortical stimulation, 10 or 100 times at 1 Hz. (b) Example of 5
successive pairings (#4–8, from c1) exhibiting relatively fixed ΔtSTDP (#4–8: −8.0, −7.8, −7.4, −8.8 and
−8.0 ms). Plotting the successive ΔtSTDP illustrates their low variance σΔt (2.6 ms). (c) Corticostriatal NMDAR-
tLTP induced by 100 post-pre pairings with fixed ΔtSTDP. (c1) Example of tLTP induced by 100 post-pre
pairings (mΔt = −9 ms, σΔt = 2.6 ms). The mean baseline EPSC amplitude, 95 ± 3 pA, increased by 77% to
168 ± 6 pA 45 minutes after pairings. Upper panel, time course of Ri (baseline: 172 ± 2 MΩ and 35–45 min
after pairings: 170 ± 3 MΩ; change of 1%) and holding current (Ihold). Inset: distribution of the 100 ΔtSTDP. (c2)
Averaged time-courses of tLTP induced by 100 post-pre pairings; this tLTP was NMDAR-mediated because
prevented by D-AP5 (50 μM). (c3) Relationship between the STDP magnitude and the STDP jitter for each
of the recorded neurons. (d) Corticostriatal eCB-tLTD induced by 100 pre-post pairings with fixed ΔtSTDP. (d1) Example of tLTD induced by 100 pre-post pairings (mΔt = +21 ms, σΔt = 2.1 ms); the mean baseline SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 2 www.nature.com/scientificreports/ EPSC amplitude, 206 ± 4 pA, decreased by 25%, to 154 ± 8 pA, 45 minutes after pairings. Upper panel, time
course of Ri (baseline: 195 ± 1 MΩ and 45–55 min after pairings: 196 ± 2 MΩ; change <1%) and Ihold. Inset:
distribution of the 100 ΔtSTDP. Results Corticostriatal synapses exhibit a bidirectional STDP in which NMDAR-tLTP, eCB-tLTD29,32–34 or eCB-tLTP12,13
are induced depending on the number of pairings (Npairings) and the spike timing (ΔtSTDP). While STDP is classi-
cally investigated using fixed ΔtSTDP, we examine here the effect of noisy ΔtSTDP, closer to in vivo-like firing. Here,
we define the Inter-Pairing-Interval (IPI) as the time interval between two consecutive presynaptic stimulations,
whereas ΔtSTDP is the time interval between the post- and the presynaptic stimulation times within a given paired
stimulation (in a “standard” STDP protocol both IPIs and ΔtSTDP are constant). NMDAR- and eCB-mediated STDP triggered with fixed ΔtSTDP. We observed bidirectional asym-
metric STDP in MSNs (Fig. 1a): 100 post-pre pairings (ΔtSTDP < 0) induced tLTP whereas 100 pre-post pairings
(ΔtSTDP > 0) induced tLTD. An example of the tLTP induced by 100 post-pre pairings with fixed ΔtSTDP (Fig. 1b)
is illustrated in Fig. 1c1. The input resistance Ri remained stable over this period. Although we fixed the values
of ΔtSTDP, they slightly vary from one pairing to another (Fig. 1b). This low jitter can be written formally as
ΔtSTDP = mΔt + ξΔt, were mΔt is the mean spike timing and ξΔtis a random variable with mean 0 and stand-
ard deviation σΔt. Overall, 100 post-pre pairings induced tLTP (mean EPSC amplitude recorded 60 min after
protocol induction: 149 ± 13% of baseline, p = 0.0028, n = 13) (Fig. 1c2). σΔt was 1.4 ± 0.2 ms (Fig. 1c3). This
tLTP was NMDAR-mediated since prevented by D-AP5 (50 μM), a NMDAR antagonist, (88 ± 12%, p = 0.375,
n = 5; p < 0.0001 when compared with tLTP in control) (Fig. 1c2). Conversely, pre-post pairings induced tLTD,
as shown in the example in Fig. 1d1. Overall, 100 pre-post pairings induced tLTD (76 ± 7%, p = 0.0070, n = 10)
(Fig. 1d2). σΔt was 1.4 ± 0.2 ms (Fig. 1d3). This tLTD was CB1R-mediated since prevented by AM251 (3 μM), a
CB1R specific inhibitor (99 ± 2%, p = 0.816, n = 5; p < 0.0001 when compared with tLTD in control) (Fig. 1d2). We thus find an anti-Hebbian polarity for corticostriatal STDP. We previously showed that GABA operates as a
Hebbian/anti-Hebbian switch at corticostriatal synapses34,35 and corticostriatal STDP polarity depends on the
presence32,33 or the absence12,29,36,37 of GABAA receptor antagonists. Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 Our results further indicate that the robustness of NMDAR-tLTP
to jitter is also strengthened by the irregularity of the spike-train stimulations. Yihui Cui1, Ilya Prokin2,3, Alexandre Mendes1, Hugues Berry 2,3 & Laurent Venance 1 (d2) Averaged time-courses of tLTD induced by 100 pre-post pairings; this tLTD
was CB1R-mediated because prevented by AM251 (3 μM). (d3) Relationship between the STDP magnitude
and the jitter. (e) Corticostriatal eCB-tLTP induced by 10 post-pre pairings with fixed ΔtSTDP. (e1) Example
of tLTP induced by 10 post-pre pairings (mΔt = −11 ms, σΔt = 3.1 ms); the mean baseline EPSC amplitude,
147 ± 3 pA, increased by 43% to 209 ± 9 pA 45 minutes after pairings. Upper panel, time course of Ri (baseline:
227 ± 2 MΩ and 45–55 min after pairings: 239 ± 2 MΩ; change of 5%) and Ihold. Inset: distribution of the 10
ΔtSTDP. (e2) Averaged time-courses of tLTP induced by 10 post-pre pairings; this tLTP was CB1R-mediated
because prevented by AM251. (e3) Relationship between the STDP magnitude and the jitter. (c1-d1-e1) Insets:
average of 12 successive EPSC amplitude at baseline (black traces) and at 40–50 min after pairings (grey traces). (f) Predictions of the mathematical model in the absence of added jitter. The model expresses the biochemical
pathways schematized in (f1). (f2) Changes of Wtotal with various ΔtSTDP and Npairings. (f3) Removing NMDAR/
CaMKII signaling suppresses the tLTP observed for ΔtSTDP > 0 and Npairings > 50. (f4) Removing CB1R
suppresses the tLTP observed for ΔtSTDP > 0 and Npairings < 20 and the tLTD observed for ΔtSTDP < 0. pathways, i.e. NMDAR-, mGluR- or endocannabinoid(eCB)-mediated STDP25–29 (for reviews see8,30,31). It is thus
expected that those various STDP forms might exhibit different robustness to spike train variability. expected that those various STDP forms might exhibit different robustness to spike train variability. Here, we show with patch-clamp recordings and mathematical modeling that corticostriatal NMDAR- and
eCB-STDP do not exhibit the same sensitivity to noisy spike timings: eCB-plasticity (eCB-tLTD and eCB-tLTP)
appeared robust to jittering whereas NMDAR-tLTP was fragile. However, increasing the number or frequency of
pairings improved NMDAR-tLTP robustness. Our results further indicate that the robustness of NMDAR-tLTP
to jitter is also strengthened by the irregularity of the spike-train stimulations. p
gf
p
y
Here, we show with patch-clamp recordings and mathematical modeling that corticostriatal NMDAR- and
eCB-STDP do not exhibit the same sensitivity to noisy spike timings: eCB-plasticity (eCB-tLTD and eCB-tLTP)
appeared robust to jittering whereas NMDAR-tLTP was fragile. However, increasing the number or frequency of
pairings improved NMDAR-tLTP robustness. Results eCB-tLTP (red full line) was triggered with 10 pairings of average spike
timing mΔt = −15 ms and eCB-tLTD (blue full line) with 100 pairings of mΔt = +20 ms. NMDAR-tLTP (red
dashed line) corresponded to 100 pairings of mΔt = −15 ms. eCB-tLTD and eCB-tLTP are weakly affected by
the jitter. NMDAR-tLTP is less robust. Full and dashed lines show averages and light swaths show ± 1 sem. (b)
Evolution of NMDAR-tLTP, eCB-tLTD and eCB-tLTP domains for σΔt = 0 (b1), 5 (b2) and 10 ms (b3) for the
STDP protocol of (a). Note the persistence of the eCB-tLTD domain (blue box) while eCB-tLTP (red box) and
NMDAR-tLTP domain (red dashed box) disappeared for 5 and 10 ms, respectively. (c–e) Similar results are
obtained with other jitter distributions, including Gaussian-distributed jitter (c) and triangularly-distributed
jitter (d). We also considered a combination of triangularly-distributed jitter with random IPIs distributed
according to an exponential distribution with refractory period τr = 0.95 s (see Methods) in (e). The plots of
the amplitude of NMDAR-tLTP, eCB-tLTD and eCB-tLTP for increasing jitter amplitude σΔt (c2, d2, e2) show ii Figure 2. The mathematical model predicts that NMDAR-tLTP is less robust to jitter than eCB-tLTP and
eCB-tLTD. (a) Using a jittered STDP protocol combining uniformly-distributed jitter of the spike timing
with deterministic IPIs at 1 Hz (a1), the amplitudes of NMDAR-tLTP, eCB-tLTD and eCB-tLTP decrease with
increasing jitter amplitudes σΔt (a2). eCB-tLTP (red full line) was triggered with 10 pairings of average spike
timing mΔt = −15 ms and eCB-tLTD (blue full line) with 100 pairings of mΔt = +20 ms. NMDAR-tLTP (red
dashed line) corresponded to 100 pairings of mΔt = −15 ms. eCB-tLTD and eCB-tLTP are weakly affected by
the jitter. NMDAR-tLTP is less robust. Full and dashed lines show averages and light swaths show ± 1 sem. (b)
Evolution of NMDAR-tLTP, eCB-tLTD and eCB-tLTP domains for σΔt = 0 (b1), 5 (b2) and 10 ms (b3) for the
STDP protocol of (a). Note the persistence of the eCB-tLTD domain (blue box) while eCB-tLTP (red box) and
NMDAR-tLTP domain (red dashed box) disappeared for 5 and 10 ms, respectively. (c–e) Similar results are
obtained with other jitter distributions, including Gaussian-distributed jitter (c) and triangularly-distributed
jitter (d). Results p
p
g
Besides this bidirectional STDP (NMDAR-tLTP and eCB-tLTD) induced for 100 pairings, we reported
that low numbers of pairings (Npairings = 5–15) induce an eCB-tLTP, dependent on the activation of CB1R12,13. Figure 1e1 shows an example of tLTP induced by 10 post-pre pairings. Overall, 10 post-pre STDP pairings
induced tLTP (145 ± 9%, p = 0.0005, n = 15), which was prevented by AM251 (3 μM) (76 ± 10%, p = 0.080,
n = 4; p < 0.0001 when compared with tLTP in control) (Fig. 1e2) as recently reported12. σΔt for 10 post-pre par-
ings STDP was 1.8 ± 0.3 ms (n = 15) (Fig. 1e3). Note that there was no difference between the σΔt measured for
NMDAR-tLTP, eCB-tLTD and eCB-tLTP (one way ANOVA, p = 0.4094). A computer model emulates NMDAR- and eCB-mediated STDP. To help interpret our experimental
results, we used a mathematical model of the signaling pathways implicated in corticostriatal STDP, including
NMDAR- and CB1R-plasticity12,29,32,33 (Fig. 1f1). The model is detailed in S1 Text, with parameter values listed
in S1 and S2 Tables. We include in the Supporting Information a thorough description of the mechanisms by
which eCB- and NDMAR-plasticity are expressed in the model when STDP protocol (Npairings = 100) is applied
(S3 Text). Figure 1f2 shows the value of the total synaptic weight (Wtotal) predicted by the model with various
Npairings and ΔtSTDP. In agreement with the experimental data, with small σΔt (3 ms), the model features three main
plasticity regions: tLTD was observed for short pre-post pairings (0 < ΔtSTDP < +30 ms), whereas short post-pre
pairings (−30 < ΔtSTDP < 0 ms) induced a first tLTP region for low numbers of pairings (3 < Npairings < 25) and a
second tLTP region emerging for Npairings > 50. Blocking the NMDAR-CaMKII pathway in the model (Fig. 1f3)
suppresses the second tLTP region whereas blocking CB1R activation (Fig. 1f4) prevents the expression of tLTD SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 3 3 www.nature.com/scientificreports/ ntificreports/
Figure 2. The mathematical model predicts that NMDAR-tLTP is less robust to jitter than eCB-tLTP and
eCB-tLTD. (a) Using a jittered STDP protocol combining uniformly-distributed jitter of the spike timing
with deterministic IPIs at 1 Hz (a1), the amplitudes of NMDAR-tLTP, eCB-tLTD and eCB-tLTP decrease with
increasing jitter amplitudes σΔt (a2). Results (b) Example of 5 successive pairings (#5–9, from c1) exhibiting jittered values
of ΔtSTDP (#5–9: −10.3, −15.2, −30.5, −20.7 and −17.3 ms); these pairings are indicated with the grey frame
within the 100 pairings displaying jittered ΔtSTDP. The distribution of the successive ΔtSTDP illustrate their
standard deviation σΔt = 7.9 ms. (c) NMDAR-tLTP is prevented with σΔt > 3 ms. (c1) Example of the absence of
plasticity after 100 post-pre pairings with mΔt = −18 ms and σΔt = 7.9 ms; the mean baseline EPSC amplitude,
194 ± 3 pA, did not show significant change 45 minutes after pairing, 186 ± 4 pA. Upper panel, time course of
Ri (baseline: 198 ± 2 MΩ and 45–55 min after pairings: 194 ± 2 MΩ; change of 2%) and holding current (Ihold). Inset: distribution of the 100 ΔtSTDP. (c2) Averaged time-courses of the absence of plasticity after 100 post-pre
pairings with 3 < σΔt < 10 ms. (c3) Relationship between the STDP magnitude and the jitter. (d) eCB-tLTD is
not affected by 3 < σΔt < 10 ms. (d1) Example of tLTD induced by 100 pre-post pairings with mΔt = +18 ms
and σΔt = 7.1 ms; the mean baseline EPSC amplitude, 160 ± 3 pA, decreased by 50%, to 81 ± 5 pA, 45 minutes
after pairings. Upper panel, time course of Ri (baseline: 112 ± 1 MΩ and 35–45 min after pairings: 118 ± 1 MΩ;
change of 5%) and Ihold. Inset: distribution of the 100 ΔtSTDP. (d2) Averaged time-courses of tLTD induced
by 100 pre-post pairings with 3 < σΔt < 10 ms. (d3) Relationship between the STDP magnitude and the jitter. (e) eCB-tLTP is not affected by 3 < σΔt < 10 ms. (e1) Example of tLTP induced by 10 post-pre pairings with
mΔt = −21 ms and σΔt = 6.4 ms; the mean baseline EPSC amplitude, 113 ± 2 pA, increased by 115% to 243 ± 6
pA 45 minutes after pairings. Upper panel, time course of Ri (baseline: 185 ± 1 MΩ and 45–55 min after
pairings: 197 ± 2 MΩ; change of 6%) and Ihold. Inset: distribution of the 10 ΔtSTDP. (e2) Averaged time-courses of Figure 3. Differential effect of jitter on NMDAR- and eCB-mediated plasticity. (a) STDP protocol; ΔtSTDP
indicates the time between pre- and postsynaptic stimulations. ΔtSTDP < 0 and ΔtSTDP > 0 refer to post-pre and
pre-post pairings, respectively. Results We also considered a combination of triangularly-distributed jitter with random IPIs distributed
according to an exponential distribution with refractory period τr = 0.95 s (see Methods) in (e). The plots of
the amplitude of NMDAR-tLTP, eCB-tLTD and eCB-tLTP for increasing jitter amplitude σΔt (c2, d2, e2) show
similar profiles and are similar to panel (a2), indicating that the model predictions are not qualitatively modified
by the probability distribution function of the jitter. SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y ntificreports/
Figure 3. Differential effect of jitter on NMDAR- and eCB-mediated plasticity. (a) STDP protocol; ΔtSTDP
indicates the time between pre- and postsynaptic stimulations. ΔtSTDP < 0 and ΔtSTDP > 0 refer to post-pre and
pre-post pairings, respectively. (b) Example of 5 successive pairings (#5–9, from c1) exhibiting jittered values
of ΔtSTDP (#5–9: −10.3, −15.2, −30.5, −20.7 and −17.3 ms); these pairings are indicated with the grey frame
within the 100 pairings displaying jittered ΔtSTDP. The distribution of the successive ΔtSTDP illustrate their
standard deviation σΔt = 7.9 ms. (c) NMDAR-tLTP is prevented with σΔt > 3 ms. (c1) Example of the absence of
plasticity after 100 post-pre pairings with mΔt = −18 ms and σΔt = 7.9 ms; the mean baseline EPSC amplitude,
194 ± 3 pA, did not show significant change 45 minutes after pairing, 186 ± 4 pA. Upper panel, time course of
Ri (baseline: 198 ± 2 MΩ and 45–55 min after pairings: 194 ± 2 MΩ; change of 2%) and holding current (Ihold). Inset: distribution of the 100 ΔtSTDP. (c2) Averaged time-courses of the absence of plasticity after 100 post-pre
pairings with 3 < σΔt < 10 ms. (c3) Relationship between the STDP magnitude and the jitter. (d) eCB-tLTD is
not affected by 3 < σΔt < 10 ms. (d1) Example of tLTD induced by 100 pre-post pairings with mΔt = +18 ms
d
7 1
h
b
li
EPSC
li
d
160±3 A d
d b 50%
81±5 A 45
i www.nature.com/scientificreports/ Figure 3. Differential effect of jitter on NMDAR- and eCB-mediated plasticity. (a) STDP protocol; ΔtSTDP Figure 3. Differential effect of jitter on NMDAR- and eCB-mediated plasticity. (a) STDP protocol; ΔtSTDP
indicates the time between pre- and postsynaptic stimulations. ΔtSTDP < 0 and ΔtSTDP > 0 refer to post-pre and
pre-post pairings, respectively. Results Insets: average of 12 successive EPSC amplitude at baseline (black trace) and at 40–50 min after STDP
pairings (grey trace). Error bars represent the SEM. and of the first tLTP region. Therefore, the model emulates both CB1R-tLTD and -tLTP, in agreement with the
eCB-tLTD and eCB-tLTP illustrated in the experimental data of Fig. 1d and e, respectively. The model also faith-
fully emulates the appearance of NMDAR-tLTP for Npairings > 50. Predicting the effects of jittered STDP with a mathematical model. The small jitter observed in the
above experiments was the result of biological variability inherent to the experimental setup. Our next goal was
to extend this study to larger jitters (3 < σΔt < 10 ms) that we controlled experimentally. We first used constant
intervals between two consecutive pairings (Inter-Pairing Interval, IPI) and used the same definition as above
for jittered ΔtSTDP (Methods): ΔtSTDP = mΔt + ξΔt, where ξΔt is a random variable with zero mean and mΔt is the
expected value of the spike timing ΔtSTDP over the Npairings. Next, we adjusted the probability distribution function
and the variance of ξΔt to perform a parametric exploration of the impact of jitter. p
p
p
p
j
In Fig. 2, the jitter ξΔt was sampled from a uniform distribution in [−σΔt, +σΔt] (Fig. 2a1). STDP in the model
was globally not robust to jitter, since the three forms of plasticity vanished when σΔt was large enough (Fig. 2a2). However, the model delivered the prediction that the three forms of plasticity do not display the same sensitivity
to jitter. In particular, NMDAR-tLTP is predicted to be very fragile when subjected to σΔt: its amplitude decreases
as soon as σΔt > 2 ms and completely vanishes for σΔt > 4 ms (Fig. 2a2 and b2). In contrast, eCB-plasticity is
much less sensitive: eCB-tLTP is still present for jitters as large as 7–8 ms and eCB-tLTD is still expressed with
σΔt = 10 ms (Fig. 2b3). We next evaluated the validity of this prediction when the probability distribution function of the jitter
changes. Figure 2 illustrates the results we obtained with Gaussian (Fig. 2c) or triangular (Fig. 2d) distributions
of σΔt (see Methods). In both cases, the profiles of the robustness curves (Fig. 2c2 and d2) are almost identical to
each other and similar to the robustness curves obtained with a uniform distribution (Fig. 2a2). In Fig. Results 2e, we
combined a triangular distribution for σΔt with random IPIs, i.e. in this case, stochasticity is applied not only to
the timing between the two stimulations of a given pairing, but also to the time interval between two consecutive
pairings: IPI distribution was Poisson with refractory period τr and rate λ (see Methods). Despite this increased
stochasticity, we found that eCB-tLTD was again more robust, tolerating jitters up to 6–7 ms (eCB-tLTP) or 10 ms
(eCB-tLTD) whereas NMDAR-tLTP was fragile, vanishing for σΔt > 4 ms (Fig. 2e2). Therefore, according to our
model, the observation that eCB-STDP is more robust to jitter than NMDAR-tLTP could be a general property of
the response of the signaling pathways to noisy paired stimulations. Differential effect of jitter on NMDAR- and eCB-mediated plasticity. Based on the model pre-
dictions, we next investigated experimentally the sensitivity to jitter of NMDAR-tLTP induced by 100 post-pre
pairings (Fig. 3a), using stochastic ΔtSTDP, distributed according to Fig. 2c1. As aforementioned, NMDAR-tLTP
can be induced with a σΔt < 3 ms (Fig. 1c3). Increasing σΔt beyond 3 ms was sufficient to obliterate tLTP for
100 post-pre pairings. As exemplified (Fig. 3b and c1), 100 post-pre pairings with σΔt = 7.9 ms and centered on
mΔt = −18 ms failed to induce plasticity. Overall, we did not observe tLTP expression for σΔt > 3 ms (92 ± 2%,
p = 0.560, n = 7; p < 0.0001 when compared with tLTP observed without jitter) (Fig. 3c2 and c3) in agreement
with the model prediction (Fig. 2c2). p
g
We next investigated the robustness of the eCB-tLTD to the variability of ΔtSTDP. We observed potent
tLTD even with large values of σΔt. As shown in Fig. 3d1, 100 pre-post pairings with σΔt = 7.0 ms centered on
mΔt = +18 ms induced a large tLTD. Overall, we observed that tLTD was still induced for 100 pre-post pairings
with 3 < σΔt < 10 ms (70 ± 6%, p < 0.0001, n = 14; p = 0.1661 when compared with tLTD observed without jitter)
(Fig. 3d2 and d3). The mean value of tLTD, observed for 3 < σΔt < 10 ms, was not different from the one observed
in control, i.e. for 0 < σΔt < 3 ms (p = 0.400). Results (b) Example of 5 successive pairings (#5–9, from c1) exhibiting jittered values
of ΔtSTDP (#5–9: −10.3, −15.2, −30.5, −20.7 and −17.3 ms); these pairings are indicated with the grey frame
within the 100 pairings displaying jittered ΔtSTDP. The distribution of the successive ΔtSTDP illustrate their
standard deviation σΔt = 7.9 ms. (c) NMDAR-tLTP is prevented with σΔt > 3 ms. (c1) Example of the absence of
plasticity after 100 post-pre pairings with mΔt = −18 ms and σΔt = 7.9 ms; the mean baseline EPSC amplitude,
194 ± 3 pA, did not show significant change 45 minutes after pairing, 186 ± 4 pA. Upper panel, time course of
Ri (baseline: 198 ± 2 MΩ and 45–55 min after pairings: 194 ± 2 MΩ; change of 2%) and holding current (Ihold). Inset: distribution of the 100 ΔtSTDP. (c2) Averaged time-courses of the absence of plasticity after 100 post-pre
pairings with 3 < σΔt < 10 ms. (c3) Relationship between the STDP magnitude and the jitter. (d) eCB-tLTD is
not affected by 3 < σΔt < 10 ms. (d1) Example of tLTD induced by 100 pre-post pairings with mΔt = +18 ms
and σΔt = 7.1 ms; the mean baseline EPSC amplitude, 160 ± 3 pA, decreased by 50%, to 81 ± 5 pA, 45 minutes
after pairings. Upper panel, time course of Ri (baseline: 112 ± 1 MΩ and 35–45 min after pairings: 118 ± 1 MΩ;
change of 5%) and Ihold. Inset: distribution of the 100 ΔtSTDP. (d2) Averaged time-courses of tLTD induced
by 100 pre-post pairings with 3 < σΔt < 10 ms. (d3) Relationship between the STDP magnitude and the jitter. (e) eCB-tLTP is not affected by 3 < σΔt < 10 ms. (e1) Example of tLTP induced by 10 post-pre pairings with
mΔt = −21 ms and σΔt = 6.4 ms; the mean baseline EPSC amplitude, 113 ± 2 pA, increased by 115% to 243 ± 6
pA 45 minutes after pairings. Upper panel, time course of Ri (baseline: 185 ± 1 MΩ and 45–55 min after
pairings: 197 ± 2 MΩ; change of 6%) and Ihold. Inset: distribution of the 10 ΔtSTDP. (e2) Averaged time-courses of SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 5 www.nature.com/scientificreports/ tLTP induced by 10 post-pre pairings with 3 < σΔt < 10 ms. (e3) Relationship between the STDP magnitude and
the jitter. Results Finally, we investigated the robustness of the eCB-tLTP to the variability of ΔtSTDP and observed potent tLTP
even with large values of jitter, i.e. up to σΔt ~ 8 ms. An example of tLTP induced by 10 post-pre pairings with
σΔt = 6.4 ms centered on mΔt = −21 ms is illustrated in Fig. 3e1. In summary, tLTP could be induced even for
σΔt = 8 ms (160 ± 12%, p = 0.0021, n = 9; p = 0.0732 when compared with tLTP observed without jitter) (Fig. 3e2
and e3); the mean value of tLTP was not different from the one observed in control (p = 0.1660).f f
p
We ensured that the mΔt absolute values were not different in control and jittered conditions for 100 post-pre
pairings (18 ± 2 ms, n = 13, vs 21 ± 3 ms, n = 7, p = 0.4711), 100 pre-post pairings (21 ± 2 ms, n = 10, vs 21 ± 2 ms,
n = 14, p = 0.8984), 10 post-pre pairings (17 ± 2 ms, n = 15, vs 20 ± 2 ms, n = 9, p = 0.2297) or between these dif-
ferent STDP protocols (one-way ANOVA: p = 0.4851) (Supplementary Fig. 1). p
y
p
pp
y
g
In conclusion, in agreement with our model prediction, whereas NMDAR-tLTP appears fragile against the
variability of ΔtSTDP, eCB-plasticity (eCB-tLTD as well as eCB-tLTP) exhibits a large robustness for the temporal
imprecision of ΔtSTDP. Model-based analysis of the effects of jittering. The above experimental results provide us with a
validation of our mathematical model. We next used this validated model as a tool to investigate the molecular
mechanisms behind these differences of robustness. f
We have shown in a previous study that eCB-tLTP was expressed when the amount of eCBs produced was
large enough that the fraction of activated CB1R, yCB1R, overcomes a threshold θLTP
start13. In the absence of added
noise, σΔt = 0, our experimental data shows that as few as 5 pairings at 1 Hz and ΔtSTDP = −15 ms are enough to SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 6 www.nature.com/scientificreports/ Figure 4. Model-based analysis of the effects of jittering. Model-predicted time-courses of (a) CB1R activation
yCB1R and (b) the concentration of active CaMKII during a protocol with jitter amplitude σΔt = 0 (a1,b1), 5
(a2,b2) or 1 0 (a3,b3) ms. Results The STDP protocol was the same as Fig. 2a1 i.e. uniformly-distributed jitter of the
spike timing and deterministic IPIs at 1 Hz. eCB-tLTP (a, red full line) and NMDAR-tLTP (b, red dashed line)
were triggered with average spike timing mΔt = −15 ms and eCB-tLTD (a, blue full line) with mΔt = +20 ms. In
(a) the shaded boxes locate the areas where yCB1R triggers eCB-tLTP or eCB-tLTD. Those areas are defined by the
thresholds θLTP
start and (θLTD
start, θLTD
stop), respectively (see Methods). For mΔt = −15 ms (red), increasing jitter
amplitudes progressively hinders the build up of the yCB1R transients. yCB1R still reaches the LTP area for 5–25
pairings with σΔt = 0 (a1) and 5 ms (a2) thus triggering eCB-tLTP but fails to do so for σΔt = 10 ms (a3). Conversely, eCB-tLTD is robust to jitter because yCB1R remains in the LTD region for most of the pairings with
mΔt = +20 ms (blue), for all tested jitter amplitudes. In (b), with mΔt = −15 ms (red) the switch from the low
activation to the high activation state of CaMKII is obtained in the absence of jittering σΔt = 0 ms (b1). The
progressive build up of activated CaMKII is efficiently suppressed as soon as σΔt = 5 ms (b2) thus effectively
suppressing NMDAR-tLTP (b3). Figure 4. Model-based analysis of the effects of jittering. Model-predicted time-courses of (a) CB1R activation
yCB1R and (b) the concentration of active CaMKII during a protocol with jitter amplitude σΔt = 0 (a1,b1), 5
(a2,b2) or 1 0 (a3,b3) ms. The STDP protocol was the same as Fig. 2a1 i.e. uniformly-distributed jitter of the
spike timing and deterministic IPIs at 1 Hz. eCB-tLTP (a, red full line) and NMDAR-tLTP (b, red dashed line)
were triggered with average spike timing mΔt = −15 ms and eCB-tLTD (a, blue full line) with mΔt = +20 ms. In
(a) the shaded boxes locate the areas where yCB1R triggers eCB-tLTP or eCB-tLTD. Those areas are defined by the
thresholds θLTP
start and (θLTD
start, θLTD
stop), respectively (see Methods). For mΔt = −15 ms (red), increasing jitter
amplitudes progressively hinders the build up of the yCB1R transients. yCB1R still reaches the LTP area for 5–25
pairings with σΔt = 0 (a1) and 5 ms (a2) thus triggering eCB-tLTP but fails to do so for σΔt = 10 ms (a3). Results Conversely, eCB-tLTD is robust to jitter because yCB1R remains in the LTD region for most of the pairings with
mΔt = +20 ms (blue), for all tested jitter amplitudes. In (b), with mΔt = −15 ms (red) the switch from the low
activation to the high activation state of CaMKII is obtained in the absence of jittering σΔt = 0 ms (b1). The
progressive build up of activated CaMKII is efficiently suppressed as soon as σΔt = 5 ms (b2) thus effectively
suppressing NMDAR-tLTP (b3). trigger eCB-tLTP12. Accordingly, our model produces large amounts of eCBs in the first 5–10 pre-post pairings so
that yCB1R reaches θLTP
start after as few as 5 pairings (Fig. 4a1). When σΔt = 5 ms jitter was added to mΔt = −15 ms,
some of the pairings failed to deliver yCB1R transients of maximal amplitude (Fig. 4a2). But for σΔt < 7–8 ms, a
number of yCB1R transients still had sufficient amplitude to overcome θLTP
start, which was enough to trigger eCB-tLTP
(Fig. 4a2). When σΔt was large though (e.g. 10 ms), the yCB1R transients failed to reach the LTP zone and eCB-tLTP
was not expressed (Fig. 4a3). p
g
In experiments without added jitter, eCB-tLTD progressively accumulates when Npairings increases and starts
to be significant for Npairings > 2512. In the model with mΔt = +20 ms and σΔt = 0, the yCB1R transients remain in
the LTD area for the most part of the STDP pairings (Fig. 4a3). To account for the progressive accumulation of
eCB-tLTP with Npairings, each transient in the model contributes a small decrease of the synaptic weight. Even with
large amounts of jitter (see e.g. σΔt = 5 or 10 ms in Fig. 4a2,a3) yCB1R transients remain mostly inside LTD area so
eCB-tLTD remains expressed even with large jitter. The robustness of NMDAR-tLTP is predicted to be frequency-dependent. Experimentally,
NMDAR-tLTP (mΔt = −15 ms and σΔt = 0) was observed at Fpairings = 1 Hz when Npairings > 50, beyond which its
amplitude did not depend much on Npairings
13. To account for this feature, the steady-state concentration of acti-
vated CaMKII, that sets Wpost in the model, is bistable: 45 min after the stimulation, CaMKII was either almost
fully inactivated (“no plasticity” state) or almost fully activated (“LTP” state)13,38. When the frequency of post-pre
pairings (−30 < ΔtSTDP < 0 ms) was large enough (i.e. Results Fpairings ≥ 1 Hz), the IPI was smaller than the decay time
of the CaMKII activation transient triggered by each post-pre pairings (Fig. 4b1). As a result, in the absence of
added noise (σΔt = 0), the CaMKII activation transients progressively built up on top of each other. The accu-
mulated CaMKII activation overcomes the threshold between the “no plasticity” and the “LTP” states only for
Npairings > 50 (Fig. 4b1)13. With jitter (e.g. σΔt = 5 ms in Fig. 4b2 or 10 ms in Fig. 4b3), many of the IPIs were either
too long or too short to trigger maximal amplitude transients of activated CaMKII. As a result, CaMKII activation
never reached the threshold, thus explaining the fragility of NMDAR-tLTP with respect to jittering (at 1 Hz). p
g
g
y
p
j
g
A major indication from the above analysis is the importance of the IPI frequency for the robustness of
NMDAR-tLTP to jitter. Figure 5 compares the robustness to jitter for constant IPI with Fpairings = 1 Hz (Fig. 5a)
and 1.05 Hz (Fig. 5b). The size of the green area that locates eCB-tLTD on Fig. 5a2,b2 was not much altered by the
increase of the pairing frequency. Likewise, the size of the red area that locates eCB-tLTP in Fig. 5a1,b1 did not p
g
g
y
p
j
g
A major indication from the above analysis is the importance of the IPI frequency for the robustness of
NMDAR-tLTP to jitter. Figure 5 compares the robustness to jitter for constant IPI with Fpairings = 1 Hz (Fig. 5a)
and 1.05 Hz (Fig. 5b). The size of the green area that locates eCB-tLTD on Fig. 5a2,b2 was not much altered by the
increase of the pairing frequency. Likewise, the size of the red area that locates eCB-tLTP in Fig. 5a1,b1 did not SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 7 www.nature.com/scientificreports/ Figure 5. Model-based analysis of the change of robustness with frequency. (a) Summary of the robustness
of the model plasticity at 1 Hz. The jittered STDP protocol combined uniformly-distributed jitter of the spike
timing and constant IPIs at 1 Hz. Two-dimensional maps show alterations of the plasticity obtained with 10 (a1)
or 100 (a2) pairings, when the average spike timing mΔt and the jitter amplitude σΔt are changed. Results vary much when the frequency was increased. However, the size of the red area locating NMDAR-tLTP increased
drastically with frequency (Fig. 5a2,b2). Therefore, our model predicts that NMDAR-tLTP should become more
robust to jitter at larger frequencies, whereas eCB-tLTP and eCB-tLTD are not significantly changed. Several aspects of glutamate signaling at the corticostriatal synapse are known to display complex
frequency-dependence39. Our model is calibrated with experimental data at 1 Hz and features none of the above
frequency dependencies. Therefore, we cannot expect a precise quantitative match between experiments and
model when frequency is varied below or above 1 Hz. However, the model still yields correct predictions of the
main qualitative trends observed in the experiments, since the effects of a small change of Fpairings in the model
(1.00 to 1.05 Hz) were similar to the effects of larger changes (1 to 3 Hz) in the experiments13. Therefore, in the
experiments, we expect to observe a change of robustness pattern similar to Fig. 5b, around Fpairings = 3 Hz rather
than 1.05 Hz. j
g
q
,
gi
y
g
Several aspects of glutamate signaling at the corticostriatal synapse are known to display complex
frequency-dependence39. Our model is calibrated with experimental data at 1 Hz and features none of the above
frequency dependencies. Therefore, we cannot expect a precise quantitative match between experiments and
model when frequency is varied below or above 1 Hz. However, the model still yields correct predictions of the
main qualitative trends observed in the experiments, since the effects of a small change of Fpairings in the model
(1.00 to 1.05 Hz) were similar to the effects of larger changes (1 to 3 Hz) in the experiments13. Therefore, in the
experiments, we expect to observe a change of robustness pattern similar to Fig. 5b, around Fpairings = 3 Hz rather
than 1.05 Hz. Increasing Fpairings and Npairings stabilized NMDAR-tLTP against jitter. We next tested experimentally
whether an increase from 1 to 3 Hz of STDP pairings would protect the NMDAR-tLTP against jitter, as predicted
by the model. As shown in Fig. 6a1, 100 post-pre pairings at 3 Hz with σΔt = 9 ms and centered on mΔt = −22 ms
induced tLTP. Overall, we observed tLTP for 100 post-pre pairings applied at 3 Hz with 4 < σΔt < 9 ms (181 ± 30%,
p = 0.0274, n = 9) (Fig. Results The locations
of the plasticity blotches are indicated with horizontal lines: full red for eCB-tLTP, full blue for eCB-tLTD and
dashed red for NMDAR-tLTP. The length of each of these blotches along the y-axis reflects their robustness
to jitter. (a3) A cross-section along the dashed and full horizontal lines of (a1,a2) illustrates the robustness of
NMDAR-tLTP, eCB-tLTD and eCB-tLTP. (b) Robustness of the model with larger frequency (1.05 Hz) and
same jittered STDP protocol as in (a). Note the distortion between the pairing frequency in the model and
in the experiments. The behavior of the model for Fpairings = 1.05 Hz corresponds to experimental results with
Fpairings ≈ 3 Hz13. The two-dimensional maps for 10 (b1) or 100 (b2) pairings as well as the cross-sections (b3)
predict that for frequencies larger than 1 Hz, the robustness changes, with NMDAR-tLTP becoming more
robust. Figure 5. Model-based analysis of the change of robustness with frequency. (a) Summary of the robustness
of the model plasticity at 1 Hz. The jittered STDP protocol combined uniformly-distributed jitter of the spike
timing and constant IPIs at 1 Hz. Two-dimensional maps show alterations of the plasticity obtained with 10 (a1)
or 100 (a2) pairings, when the average spike timing mΔt and the jitter amplitude σΔt are changed. The locations
of the plasticity blotches are indicated with horizontal lines: full red for eCB-tLTP, full blue for eCB-tLTD and
dashed red for NMDAR-tLTP. The length of each of these blotches along the y-axis reflects their robustness
to jitter. (a3) A cross-section along the dashed and full horizontal lines of (a1,a2) illustrates the robustness of
NMDAR-tLTP, eCB-tLTD and eCB-tLTP. (b) Robustness of the model with larger frequency (1.05 Hz) and
same jittered STDP protocol as in (a). Note the distortion between the pairing frequency in the model and
in the experiments. The behavior of the model for Fpairings = 1.05 Hz corresponds to experimental results with
Fpairings ≈ 3 Hz13. The two-dimensional maps for 10 (b1) or 100 (b2) pairings as well as the cross-sections (b3)
predict that for frequencies larger than 1 Hz, the robustness changes, with NMDAR-tLTP becoming more
robust. vary much when the frequency was increased. However, the size of the red area locating NMDAR-tLTP increased
drastically with frequency (Fig. 5a2,b2). Therefore, our model predicts that NMDAR-tLTP should become more
robust to jitter at larger frequencies, whereas eCB-tLTP and eCB-tLTD are not significantly changed. Results Increasing frequency and Npairings stabilized NMDAR-tLTP against jitter. (a) 100 post-pre pairings at
d
d
T
h h
ff
d b
(
)
l
f
T
d
d b We next investigated whether a higher Npairings would secure the expression of NMDAR-tLTP even with
σΔt > 3 ms (Fig. 6b). When we doubled Npairings we did not observe expression of significant tLTP. Indeed, 200
post-pre pairings (−30 < mΔt < 0 ms with 3 < σΔt < 10 ms) did not induce plasticity (112 ± 9%, p = 0.236, n = 6;
Fig. 6b3) but when we increased Npairings up to 250–300, tLTP was reliably observed with σΔt > 3 ms. Figure 6b1
shows an example of tLTP induced by 250 pairings with σΔt = 5 ms and centered on −20 ms. Overall, 250–300
post-pre STDP pairings (−30 < mΔt < 0 ms with 3 < σΔt < 10 ms) induced tLTP (170 ± 10%, p = 0.0012, n = 6;
Fig. 6b2 and b3); the mean values of these tLTP observed for Npairings = 250–300 at 1 Hz with 4 < σΔt < 9 ms were not
different compared to tLTP induced with 100 post-pre pairings at 1 Hz with σΔt < 3 ms (p = 0.1952). We observed
different synaptic efficacy changes following 100, 200 and 250–300 post-pre pairings (ANOVA, p < 0.0001). Transitions from spike-timing- to frequency-dependent plasticity. We next tested with our mathe-
matical model whether the robustness of STDP to jitter of the spike timing depends on the regularity of the spike
train. In Fig. 2e, we show an example of the combination of stochastic spike timings (triangular distribution)
with stochastic IPIs (Poisson distributed) with a constant and relatively large refractory period τr (see Methods-
Mathematical model, Protocol 4). However, when τr decreases, short IPIs are more frequently sampled and the
train of presynaptic stimulation becomes more irregular. Therefore, through variations of the refractory period τr,
the presynaptic stimulation can be progressively switched from highly irregular to regular spike trains. Figure 7 shows model predictions for the alteration of the robustness curves when τr varies. With high refrac-
tory period, i.e. when τr ≈ Fpairings, the IPIs are weakly noisy, so one recovers the robustness curves of Fig. 2e,
with eCB-STDP being more robust than NMDAR-tLTP. Results 6a2 and a3); the mean value of these tLTP observed at 3 Hz with 4 < σΔt < 9 ms were not
significantly different from the one observed with stimulation-related σΔt < 3 ms at 1 Hz (p = 0.520). When we
plotted the magnitude of plasticity against σΔt for 100 post-pre pairings at 3 Hz, we observed that tLTP was still
induced even for σΔt = 9 ms (Fig. 6a3). We observed different synaptic efficacy changes following 100 post-pre
pairings at 1 Hz with and without jitter and at 3 Hz with jitter (ANOVA, p < 0.0001). The NMDAR-tLTP thus
acquires certain robustness to the variability of ΔtSTDP with increasing frequency of pairings. 8 SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y www.nature.com/scientificreports/ Figure 6. Increasing frequency and Npairings stabilized NMDAR-tLTP against jitter. (a) 100 post-pre pairings at
3 Hz induced a tLTP, which is not affected by 3 < σΔt < 10 ms. (a1) Example of tLTP induced by 100 post-pre
pairings at 3 Hz with mΔt = −22 ms and σΔt = 9 ms; the mean baseline EPSC amplitude, 194 ± 4 pA, increased
by 212%, to 606 ± 7 pA, 45 min after pairings. Upper panel, time course of Ri (baseline: 123 ± 1 MΩ and
45–55 min after pairings: 138 ± 1 MΩ; change of 12%) and Ihold for this cell. Inset: distribution of ΔtSTDP for the
100 pairings. (a2) Averaged time-courses of tLTP induced by 100 post-pre pairings at 3 Hz with 3 < σΔt < 10 ms. (a3) Relationship between the STDP magnitude and the jitter for each of the recorded neurons after 100
pairings at 1 Hz (n = 7; blue circles) or at 3 Hz (n = 10; red circles). (b) 250–300 post-pre pairings induced
tLTP, which is not affected by 3 < σΔt < 10 ms. (b1) Example of tLTP induced by 250 post-pre pairings with
mΔt = −20 ms and σΔt = 5 ms; the mean baseline EPSC amplitude, 126 ± 3 pA, increased by 57% to 198 ± 4 pA
45 minutes after pairings. Upper panel, time course of Ri (baseline: 65 ± 1 MΩ and 45–55 min after pairings:
65 ± 2 MΩ; change < 1%) and Ihold. Inset: distribution of ΔtSTDP for the 250 pairings. (b2) Averaged time-courses
of tLTP induced by 250–300 post-pre pairings with 3 < σΔt < 10 ms. Results (b3) Relationship between the STDP
magnitude and the jitter for each of the recorded neurons after 100 (n = 7; blue circles), 200 (n = 6; black circles)
or 250–300 (n = 7; red circles) post-pre pairings. Insets: average of 12 successive EPSC amplitude at baseline
(black trace) and at 40–50 min after STDP pairings (grey trace). Error bars represent the SEM. Figure 6. Increasing frequency and Npairings stabilized NMDAR-tLTP against jitter. (a) 100 post-pre pairings at
3 Hz induced a tLTP, which is not affected by 3 < σΔt < 10 ms. (a1) Example of tLTP induced by 100 post-pre
pairings at 3 Hz with mΔt = −22 ms and σΔt = 9 ms; the mean baseline EPSC amplitude, 194 ± 4 pA, increased
by 212%, to 606 ± 7 pA, 45 min after pairings. Upper panel, time course of Ri (baseline: 123 ± 1 MΩ and
45–55 min after pairings: 138 ± 1 MΩ; change of 12%) and Ihold for this cell. Inset: distribution of ΔtSTDP for the
100 pairings. (a2) Averaged time-courses of tLTP induced by 100 post-pre pairings at 3 Hz with 3 < σΔt < 10 ms. (a3) Relationship between the STDP magnitude and the jitter for each of the recorded neurons after 100
pairings at 1 Hz (n = 7; blue circles) or at 3 Hz (n = 10; red circles). (b) 250–300 post-pre pairings induced
tLTP, which is not affected by 3 < σΔt < 10 ms. (b1) Example of tLTP induced by 250 post-pre pairings with
mΔt = −20 ms and σΔt = 5 ms; the mean baseline EPSC amplitude, 126 ± 3 pA, increased by 57% to 198 ± 4 pA
45 minutes after pairings. Upper panel, time course of Ri (baseline: 65 ± 1 MΩ and 45–55 min after pairings:
65 ± 2 MΩ; change < 1%) and Ihold. Inset: distribution of ΔtSTDP for the 250 pairings. (b2) Averaged time-courses
of tLTP induced by 250–300 post-pre pairings with 3 < σΔt < 10 ms. (b3) Relationship between the STDP
magnitude and the jitter for each of the recorded neurons after 100 (n = 7; blue circles), 200 (n = 6; black circles)
or 250–300 (n = 7; red circles) post-pre pairings. Insets: average of 12 successive EPSC amplitude at baseline
(black trace) and at 40–50 min after STDP pairings (grey trace). Error bars represent the SEM. igure 6. Discussion Here, we have introduced jittered STDP protocols as noisy variants of STDP protocols where the spike tim-
ing, i.e. the delay between a presynaptic and paired postsynaptic stimulation (or vice-versa), is a random varia-
ble. Our results show that the magnitude of variability that STDP can tolerate varies depending on the pairing
frequency and on the STDP form, i.e. NMDAR-tLTP, eCB-tLTP or eCB-tLTD. At 1 Hz pairing frequency,
eCB-STDP (eCB-tLTD and eCB-tLTP) is much more robust than NMDAR-dependent tLTP. However, this
robustness depends on the number and frequency of pairings. Indeed, increasing the average pairing frequency
to 3 Hz improved the robustness of NMDAR-tLTP. We observed a similar improvement of the robustness of
NMDAR-tLTP to jitter when the number of pairings increased more than two folds.hih j
p
g
These results have wide-ranging implications for STDP expression in in vivo-like firing. They indicate that
eCB-STDP could be more likely responsible for fast learning involving few trials (eCB-tLTP) or for learning at
lower activity frequency than NMDAR-STDP. NMDAR-STDP could however take part in learning for larger fre-
quencies or when the same spike timing recurs a large number of times during sustained activity. It also implies
that in synapses where NMDAR is the main coincidence detector for tLTD or tLTP3,5,6,40–42, a high degree of
precision of the spike timing, and/or an increased number or frequency of pairings, are required for the emer-
gence of the NMDAR-mediated plasticity. This can be viewed as a mechanism preventing the occurrence of spu-
rious plasticity for noisy neural network activity. At synapses in which NMDAR and endocannabinoids are both
involved in STDP expression12,13,25–27,33,43 and in which eCB-LTP can occur, as it is the case in the striatum12,13,
the hippocampus44–48 or the neocortex49 the emergence of STDP would be possible even in noisy conditions and
could thus serve subsequently as a priming for the subsequent expression of NMDAR-tLTP. h
d
l
d h
b
f T
h
ff
l
h
k
b
l q
y
p
g
q
p
In the present study, we explored the robustness of STDP when noise affects not only the spike timing but also
the inter-pairing interval (IPI), i.e. the delay between two successive pairings. In the absence of jitter of the spike
timing, our model predicts that IPI irregularity tends to consolidate tLTP at the expense of tLTD. Results Hence,
when the presynaptic spike train is progressively switched from regular to very irregular spike trains, two main
changes occur: (i) the three plasticity forms (NMDAR-tLTP, eCB-tLTD and eCB-tLTP) become increasingly
robust to jittering (at the limit of highly irregular spike trains, jittering hardly affects plasticity amplitudes), and
(ii) eCB-tLTD is progressively changed to LTP, so that highly irregular paired stimulations with Fpairings = 1 Hz
only produce LTP regardless of the sign of the spike timing. τr, NMDAR-tLTP becomes more robust than eCB-tLTP. Moreover, eCB-tLTD (Fpairings = 1 Hz, mΔt = +20 ms,
Npairings = 100) progressively stops triggering LTD when τr decreases, even inducing LTP for very low τr. Hence,
when the presynaptic spike train is progressively switched from regular to very irregular spike trains, two main
changes occur: (i) the three plasticity forms (NMDAR-tLTP, eCB-tLTD and eCB-tLTP) become increasingly
robust to jittering (at the limit of highly irregular spike trains, jittering hardly affects plasticity amplitudes), and
(ii) eCB-tLTD is progressively changed to LTP, so that highly irregular paired stimulations with Fpairings = 1 Hz
only produce LTP regardless of the sign of the spike timing. SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y Results However, when τr is 70% of Fpairings, the robustness of
both eCB-tLTP and NMDAR-tLTP increases and both exhibit similar robustness. With even smaller values of SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 9 www.nature.com/scientificreports/ Figure 7. The robustness of plasticity depends on the regularity of the presynaptic spike train. The figure shows
the robustness of the model plasticity (eCB-tLTP with red, eCB-tLTD with blue and NMDAR-tLTP with dashed
red) with the jittered STDP protocol of Fig. 2e1, i.e. exponentially-distributed IPIs with refractory period τr
and triangularly-distributed jitter ξΔt. From right to left, the refractory period τr is gradually decreased. When
τr → 1/Fpairings where Fpairings = 1 Hz is the average pairings frequency, one gets regular presynaptic spike trains,
similar to classical STDP protocols. In this case (see e.g. τr = 0.95/Fpairings), the robustness pattern is similar
to the previous observations at 1 Hz: eCB-STDP is robust, whereas NMDAR-tLTP is fragile. However, when
τr → 0, the presynaptic spike train becomes highly irregular. In this case, the robustness patterns changes totally,
with NMDAR-LTP becoming the most robust, and eCB-tLTD turning into a LTP. Hence, irregular presynaptic
spike trains are predicted to produce robust LTP at the expense of LTD. Figure 7. The robustness of plasticity depends on the regularity of the presynaptic spike train. The figure shows
the robustness of the model plasticity (eCB-tLTP with red, eCB-tLTD with blue and NMDAR-tLTP with dashed
red) with the jittered STDP protocol of Fig. 2e1, i.e. exponentially-distributed IPIs with refractory period τr
and triangularly-distributed jitter ξΔt. From right to left, the refractory period τr is gradually decreased. When
τr → 1/Fpairings where Fpairings = 1 Hz is the average pairings frequency, one gets regular presynaptic spike trains,
similar to classical STDP protocols. In this case (see e.g. τr = 0.95/Fpairings), the robustness pattern is similar
to the previous observations at 1 Hz: eCB-STDP is robust, whereas NMDAR-tLTP is fragile. However, when
τr → 0, the presynaptic spike train becomes highly irregular. In this case, the robustness patterns changes totally,
with NMDAR-LTP becoming the most robust, and eCB-tLTD turning into a LTP. Hence, irregular presynaptic
spike trains are predicted to produce robust LTP at the expense of LTD. τr, NMDAR-tLTP becomes more robust than eCB-tLTP. Moreover, eCB-tLTD (Fpairings = 1 Hz, mΔt = +20 ms,
Npairings = 100) progressively stops triggering LTD when τr decreases, even inducing LTP for very low τr. www.nature.com/scientificreports/ plasticity, SDDP) has been reported in the neocortex51, striatum52 and hippocampus53. In the striatum, SDDP dif-
fers from STDP on two aspects: (i) SDDP was induced in a larger temporal window (−100 < ΔtSTDP < +100 ms)
and (ii) tLTP and tLTD were induced regardless of the spike timing. Therefore, the action potential would not be
necessary for plasticity induction, but determinant for the STDP polarity and the width of the ΔtSTDP. The effects
of jitter remain to be determined on SDDP, i.e. subthreshold events. j
,
The polarity of plasticity depends greatly on the frequency of stimulus presentation. Indeed, prolonged
firing activity at low and high frequency promote LTD and LTP, respectively, at the same synapses in hip-
pocampus54–56. However, alteration of a regular low-frequency stimulation (900 stimulations at 1 Hz) with
Poisson-distributed interstimulus intervals (as an approximation of naturalistic patterns) prevents LTD expres-
sion in the neocortex57. Interestingly, this LTD induced with regular low-frequency stimulation has been reported
to be NMDAR-mediated58, and we show here that NMDAR-plasticity is fragile to jitter especially for low firing
rate (1 Hz). It remains thus to investigate in a rate-based paradigm whether Poisson stimulation would erase LTD
induced by higher firing rate (3–5 Hz54). Various forms of naturalistic patterns as an attempt to mimic in vivo
activity have been experimentally tested for plasticity expression: from single burst of dendritic spikes, which
induced NMDAR-LTP in hippocampus59 and NMDAR-LTD in neocortex60, to noisy STDP-like protocols7,19,20,22
and burst-timing-dependent plasticity paradigms61. Based on data harvested in the neocortex, a phenomeno-
logical computer modeling showed that for constant frequencies but random spiking, LTP wins over LTD with
increasing frequency7. Activity patterns recorded in vivo during specific behavioral tasks have been replayed ex
vivo to test their ability to induce plasticity19,56,62. When sequences of spike trains recorded in vivo from visual
cortex of the anaesthetized cat in response to natural scenes were replayed ex vivo in layer 2/3 pyramidal cells
of rat visual cortex, both LTP and LTD was observed experimentally and modeling suggested the importance of
suppressive interactions between spikes19. In the dentate gyrus, in vivo patterns recorded in granule cell during a
task of delayed nonmatch-to-sample were able in most of the cases to induce LTP at mossy fibers56. Similarly, when
hippocampal place cells activity recorded in vivo were replayed ex vivo, NMDAR-LTP (also dependent on cholin-
ergic tone) was observed62. www.nature.com/scientificreports/ Natural stimuli recruit various neuronal circuits leading to presynaptic jitter, which can
have important physiological consequences. Indeed, in vivo visually driven presynaptic inputs induce presynaptic
jitter leading to broader membrane potential events, and can consequently promote a spatial reorganization of the
responses of neurons to neighboring stimuli within a receptive field, as demonstrated in the rat visual cortex22.h p
g
g
pi
,
The debate about the nature of the neural coding, i.e. rate versus time codes63–65, is prominent when studying STDP. It remains to be tested with mathematical models and experiments whether activity patterns of various vigilance states
differently result in the emergence of spike- or rate-coding plasticity. At most of the synapses both rate and time codes
co-exist; for example at CA1 synapses, stimuli induced LTD at 2–3 Hz, no plasticity at 10 Hz and LTP at 50 Hz demon-
strating a rate code54, and time codes are also supported as illustrated by the expression of various forms of STDP at the
same synapse11. An answer to this debate could be that both rate and time codes work together for plasticity expression,
as recently shown in the somatosensory cortex66, but their relative contribution in vivo would depend on the ongoing
activity (low or high), the cerebral structures, the noise level, the neuromodulatory systems at play and the signaling
pathways involved in plasticity expression (as shown in the present study). Briefly, it appears that STDP can emerge
form low frequency activity whereas rate codes would take over for higher frequency ranges7,19,20,22. q
y
y
g
q
y
g
A major conclusion from our present work is that the robustness of STDP to jitter depends on the underlying
signaling pathways. For instance, the robustness of NMDAR-tLTP in our model is dependent on the amplitude
of the activated CaMKII transients triggered by each post-pre pairings or on alterations of the ratio between
the decay time of these transients and the IPIs. The robustness of STDP to jitter could be similarly controlled by
quantitative variations in the underlying pathways. Such variations are expected to occur between two neuronal
subtypes or brain regions but also as a result of the activation of a neuromodulatory pathway. www.nature.com/scientificreports/ Therefore, our work
suggests that the expression of STDP in in vivo-like firing might appear or disappear as a result of modulations of
its robustness to jitter, depending on the properties of the incoming patterns (reflecting for example fast learning
or heavy training), but also on the brain region and, more transiently, on the activation of neuromodulatory
pathways. Discussion This prediction
that irregular IPIs strengthen tLTP when the spike timing is constant confirms the result obtained independently
and with different mathematical models7,19,22,24. When IPIs are regular, adding jitter of the spike timing in our
model progressively suppresses STDP. However, with more irregular IPIs our model predicts that the three STDP
forms should exhibit larger robustness to noise. Therefore our result suggests that jittering of the spike timing has
larger impact on STDP when the IPIs are regular, but that jittering has much less consequence with irregular IPIs. Experimental testing of this model prediction would greatly improve our understanding of the induction and
maintenance of STDP in in vivo-like firing.h i
g
Subthreshold postsynaptic depolarization is a key factor in the induction of plasticity50. The existence of
plasticity induced by STDP-like protocols without postsynaptic spikes (subthreshold-depolarization-dependent SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 10 www.nature.com/scientificreports/ Animals and brain slice preparation Current- and voltage-clamp recordings were filtered at 5 kHz
and sampled at 10 kHz, with the Patchmaster v2 × 32 program (HEKA Elektronik). Spike timing-dependent plasticity protocols: regular and jittered patterns. Electrical stimulations
were performed with a concentric bipolar electrode (Phymep, Paris, France) placed in layer 5 of the somatosen-
sory cortex. Electrical stimulation was monophasic, at constant current (ISO-Flex stimulator, AMPI, Jerusalem,
Israel). Cortical stimulations evoked glutamatergic excitatory postsynaptic currents (EPSCs) (inhibited by CNQX
10 μM and D-AP5 50 μM, n = 6) and not significantly affected by GABAergic events; Indeed, blocking GABAARs
with picrotoxin (50 μM) did not significantly affect EPSC amplitude at corticostriatal synapses (123 ± 31 pA
before and 118 ± 26 pA after picrotoxin, p = 0.500, n = 5). MSNs were kept at their resting membrane potential
(−74.6 ± 0.6 mV, n = 90) and repetitive control stimuli were applied at 0.1 Hz. Currents were adjusted to evoke
50–200 pA EPSCs. STDP protocols consisted of pairings of pre- and postsynaptic stimulations (at 1 Hz) separated
by a specific time interval (ΔtSTDP). The paired stimulations were applied at 1 Hz throughout the study except in
Fig. 6a1 and a2 in which 3 Hz stimulations were tested. Presynaptic stimulations corresponded to cortical stimu-
lations and the postsynaptic stimulation of an action potential evoked by a depolarizing current step (30 ms dura-
tion) in MSNs. ΔtSTDP < 0 ms for post-before-pre pairings, and ΔtSTDP > 0 ms for pre-before-post pairings. Neuronal
recordings on neurons were made over a period of 10 minutes at baseline, and for at least 60 minutes after the STDP
protocols. EPSC baseline stability was assessed by comparing the average of the EPSC magnitude of the two first
and the two last minutes of baseline; if the variation of amplitude exceeded 10%, the experiment was discarded4. Recordings were stopped if the injected current was larger than 50 or 100–150 pA during the baseline and after pair-
ings, respectively. We individually measured and averaged the amplitude of 60 successive EPSCs from both baseline
and 45–55 minutes after STDP protocol, in which the latter was normalized by the former to calculate long-term
synaptic efficacy changes. Neuron recordings were made in voltage-clamp mode during baseline and for the 60 min-
utes of recording after the STDP protocol, and in current-clamp mode during STDP protocol. Animals and brain slice preparation Animals and brain slice preparation. OFA (Oncins France Strain A) rats (Charles River, L’Arbresle,
France) were used at postnatal day 25–32 for brain slice electrophysiology. All experiments were performed in
accordance with the guidelines of the local animal welfare committee (Center for Interdisciplinary Research
in Biology Ethics Committee) and the EU (directive 2010/63/EU). The experimental protocol (ref. 2017–02 N°
2016120110255176) was approved by the Ethics Committee in Charge of Animal Experimentation (Paris Centre
et Sud). Every precaution was taken to minimize stress and the number of animals used in each series of experi-
ments. Animals were housed in standard 12 hours light/dark cycles and food and water were available ad libitum. Horizontal brain slices containing the somatosensory cortical area and the corresponding corticostriatal projec-
tion field were prepared as previously described36. Corticostriatal connections (between somatosensory cortex
layer 5 and the dorso-lateral striatum) are preserved in the horizontal plane. Brain slices (330 µm-thick) were
prepared with a vibrating blade microtome (VT1200S, Leica Microsystems, Nussloch, Germany). Brains were
sliced in an ice-cold cutting solution (125 mM NaCl, 2.5 mM KCl, 25 mM glucose 25 mM NaHCO3, 1.25 mM
NaH2PO4, 2 mM CaCl2, 1 mM MgCl2, 1 mM pyruvic acid) through which 95% O2/5% CO2 was bubbled. The slices
were transferred to the same solution at 34 °C for one hour and then to room temperature. Patch-clamp recordings. Patch-clamp recordings were performed as previously described12,18. Briefly,
for whole-cell recordings in borosilicate glass pipettes of 5–7 MΩ resistance were filled with (in mM): 122
K-gluconate, 13 KCl, 10 HEPES, 10 phosphocreatine, 4 Mg-ATP, 0.3 Na-GTP, 0.3 EGTA (adjusted to pH 7.35
with KOH). The composition of the extracellular solution was (mM): 125 NaCl, 2.5 KCl, 25 glucose, 25 NaHCO3, SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 11 www.nature.com/scientificreports/ 1.25 NaH2PO4, 2 CaCl2, 1 MgCl2, 10 μM pyruvic acid bubbled with 95% O2 and 5% CO2. Signals were amplified
using EPC10–2 amplifiers (HEKA Elektronik, Lambrecht, Germany). All recordings were performed at 34 °C
(Bath-controller V, Luigs&Neumann, Ratingen, Germany) and slices were continuously superfused with extracel-
lular solution, at a rate of 2 ml/min. Slices were visualized under microscope (BX51WI Olympus, Rungis, France),
with a 4x/0.13 objective for the placement of the stimulating electrode and a 40x/0.80 water-immersion objective
for the localization of cells for whole-cell recordings. Animals and brain slice preparation Experiments were
excluded if the input resistance (Ri), measured every 10 sec all along the experiment, varied by more than 20%
during the very same time period where the EPSC amplitude were measured for plasticity assessment: 10 minutes of
baseline and the 10 last minutes of the recording (generally 45–55 minutes after pairings). After recording of 10 min
control baseline, drugs were applied in the bath. A new baseline with drugs was recorded after a time lapse of 10 min
(to allow the drug to be fully perfused) for 10 min before the STDP protocol. Drugs were present until the end
of the recording. All chemicals were purchased from Tocris (Ellisville, MO, USA), except for picrotoxin (Sigma). DL-2-amino-5-phosphono-pentanoic acid (D-AP5; 50 μM) and 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX;
10 μM) were dissolved directly in the extracellular solution and bath applied. N-(piperidin-1-yl)-5-(4-iodophenyl)-
1-(2,4-dichlorophenyl)-4-methyl-1H-pyrazole-3-carboxamide (AM251; 3 μM) and picrotoxin (50 μM) were dis-
solved in ethanol and added to the external solution, such that the final concentration of ethanol was 0.01–0.1%. Δ i
For the jittered ΔtSTDP patterns, we used the following algorithm (see Protocol 2, below). We note tprei and tposti
the times of the presynaptic and postsynaptic stimulations in the paired stimulation number i, respectively. To set
them, we first fixed the times of each presynaptic stimulation using the pairing frequency, i.e. =
t
i
F
prei
pairings. The
postsynaptic times were then chosen randomly according to tposti = tprei + mΔt + ξΔt,i where mΔt is the average spike
timing and ξΔt is a random variable with mean 0 and variance σΔt
2 . We defined the Inter-Pairing-Interval (IPI) as the
time interval between the presynaptic stimulation times of two consecutive paired stimulations
(
=
−
+
t
t
IPIi
pre
pre
i
i
1
), whereas the spike timing ξt is the time interval between the postsynaptic and the presynaptic
stimulation times within a given paired stimulation (Δ
=
−
t
t
t
STDP
post
pre
i
i
i). In a standard STDP protocol both the
spike timing and the IPI are constant. The above algorithm yields constant IPIs and stochastic spike timings. Electrophysiological data analysis. Off-line analysis was performed with Fitmaster (Heka Elektronik)
and Igor-Pro 6.0.3 (Wavemetrics). Statistical analysis was performed with Prism 5.02 software. In all cases “n”
refers to an experiment on a single cell from a single slice. Animals and brain slice preparation We set
ζ
=
+
t
i
F
i
pre
pre,
i
pairings
and
ζ
δ
=
+
+
−
∆
t
m
i
F
t
i
post
post,
i
pairings
, where ζpre and ζpost are i.i.d. random variables with the uniform
distribution of eq. (1). The resulting spike timing is ΔtSTDP = mΔt + ξΔt where ξΔt = ζpost − ζpre has a
triangular distribution. Note that in this case, one still has
≈
∀i
IPIi
F
1
pairings
as long as 2|mΔt|Fpairings ≪ 1,
which can safely be assumed here since we used |mΔt| < 50 ms and Fpairings < 2 Hz.i y
|
Δt|
pairings
(5) Protocol 4 includes both a stochastic spike timing and stochastic IPIs (Fig. 2e1). We first sampled Npairings
IPIs according to an exponential distribution with rate λ and refractory period τr: P(I-
PI) = Θ (IPI − τr)λ exp (−λ(IPI − τr)), where Θ(x) is the Heaviside function Θ (x) = 1 if x ≥ 0; 0 otherwise. The spike trains defined with protocol 4 are thus Poisson process. We used these IPIs to fix the stimulation
times, adding a triangularly distributed jitter as in Protocol 3 above:
ζ
= ∑
+
<
t
IPI
j
i
j
i
pre
pre,
i
and
ζ
δ
= ∑
+
+
−
<
∆
t
m
IPI
j
i
j
t
i
post
post,
i
. To keep the average stimulation frequency at 1/Fpairings, we con-
strained the value of the rate λ as λ = (1/Fpairings − τr)−1. y
|
Δt|
pairings
(5) Protocol 4 includes both a stochastic spike timing and stochastic IPIs (Fig. 2e1). We first sampled Npairings
IPIs according to an exponential distribution with rate λ and refractory period τr: P(I-
PI) = Θ (IPI − τr)λ exp (−λ(IPI − τr)), where Θ(x) is the Heaviside function Θ (x) = 1 if x ≥ 0; 0 otherwise. The spike trains defined with protocol 4 are thus Poisson process. We used these IPIs to fix the stimulation
times, adding a triangularly distributed jitter as in Protocol 3 above:
ζ
= ∑
+
<
t
IPI
j
i
j
i
pre
pre,
i
and
ζ
δ
= ∑
+
+
−
<
∆
t
m
IPI
j
i
j
t
i
post
post,
i
. Animals and brain slice preparation To simulate
Protocol 1, we first set each presynaptic stimulation time using
=
t
i
F
prei
pairings. Postsynaptic times were then fixed by tposti = tprei + mΔt + ξΔt,i − δ. pairings
fixed by tposti = tprei + mΔt + ξΔt,i − δ. p
i
p
i
(3) Protocol 2 (Fig. 2c1) shared the same definition as Protocol 1, except that the jitter followed a norma
distribution with zero mean and variance σΔt
2 :
ξ
σ
=
∆
∆
P(
)
(0,
)
t
t
2 . p
i
p
i
(3) Protocol 2 (Fig. 2c1) shared the same definition as Protocol 1, except that the jitter followed a normal
distribution with zero mean and variance σΔt
2 :
ξ
σ
=
∆
∆
P(
)
(0,
)
t
t
2 . i
p
i
(3) Protocol 2 (Fig. 2c1) shared the same definition as Protocol 1, except that the jitter followed a normal
distribution with zero mean and variance σΔt
2 :
ξ
σ
=
∆
∆
P(
)
(0,
)
t
t
2 . i
i
(3) Protocol 2 (Fig. 2c1) shared the same definition as Protocol 1, except that the jitter followed a normal
distribution with zero mean and variance σΔt
2 :
ξ
σ
=
∆
∆
P(
)
(0,
)
t
t
2 . Δt
ξ∆
∆
(
)
(
)
t
t
(4) For Protocol 3 (Fig. 2d1), we used a triangular distribution of the jitter by adding uniformly distributed
jitter to both presynaptic and postsynaptic times. We set
ζ
=
+
t
i
F
i
pre
pre,
i
pairings
and
ζ
δ
=
+
+
−
∆
t
m
i
F
t
i
post
post,
i
pairings
, where ζpre and ζpost are i.i.d. random variables with the uniform
distribution of eq. (1). The resulting spike timing is ΔtSTDP = mΔt + ξΔt where ξΔt = ζpost − ζpre has a
triangular distribution. Note that in this case, one still has
≈
∀i
IPIi
F
1
pairings
as long as 2|mΔt|Fpairings ≪ 1,
which can safely be assumed here since we used |mΔt| < 50 ms and Fpairings < 2 Hz.i ∆t
(4) For Protocol 3 (Fig. 2d1), we used a triangular distribution of the jitter by adding uniformly distributed
jitter to both presynaptic and postsynaptic times. SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y Animals and brain slice preparation To keep the average stimulation frequency at 1/Fpairings, we con-
strained the value of the rate λ as λ = (1/Fpairings − τr)−1. Stimulations. A detailed account of our mathematical model can be found in S1 Text. We modeled glutamate
concentration in the synaptic cleft, G(t), as a train of exponentially-decaying impulses triggered by presynaptic
stimuli at time tprei. We modeled the electrical response to these stimulations in a postsynaptic element considered
as a single isopotential compartment with AMPAR, NMDAR, VSCC and TRPV1 conductances: = −
−
−
−
−
−
−
C dV dt
g V
V
I
V G t
I
V G t
I
V
I
V AEA
I
t
/
(
)
( ,
( ))
( ,
( ))
( )
( ,
)
( )
(2)
L
m
L
AMPAR
NMDAR
VSCC
TRPV1
action (2) where V is membrane potential; gL and VL are leak conductance and reversal potential respectively; IAMPAR,
INMDAR, IVSCC and ITRPV1 are currents through AMPAR, NMDAR, L-type VSCC (v1.3) and TRPV1 respectively. Iaction is the action current resulting from the postsynaptic stimulation (backpropagating action potential on top of
a 30 ms depolarization). Details about the analytical expressions of these currents are given in S1 Text. Biochemical signaling. We modeled the kinetics of the biochemical pathways activated by the above electrical
stimulations using the model of ref.13. We give below a quick overview of this model and refer to S1 Text for the
details. Free cytosolic calcium is one of the main signaling actors in the model. To model its dynamics, we assumed
calcium can be transferred from/to two main sources: (i) extracellular calcium, via the plasma membrane channels
of eq. (2) above and (ii) the endoplasmic reticulum (ER), via the IP3-dependent calcium-induced calcium release
system. These calcium transients activated a network of biochemical pathways that collectively set the synaptic
weight. Hence, in this model, the synaptic weight is entirely fixed by the underlying biochemical signaling network13. More precisely, we assumed that the total synaptic weight is the product of a pre- and a postsynaptic contribution
Wtotal = WpreWpost. Postsynaptic plasticity was based on the activation by calcium of calmodulin and CaMKII67,68. We
assumed that postsynaptic plasticity is directly proportional to the calcium-dependent activation of CaMKII: =
+
. Animals and brain slice preparation All results are expressed as mean ± SEM in the text
and as mean ± SD in the figures. Statistical significance was assessed in unpaired t tests, one way ANOVA, or in
one-sample t tests, as appropriate, using the indicated significance threshold (p). Mathematical model. Jittered STDP protocols. In the model, we define the spike timing as ∆
=
tSTDPi
δ
−
+
t
t
post
pre
i
i
where δ accounts for the time elapsed between the onset of the postsynaptic step current and the
action potential it triggers (~3 ms in MSNs). In agreement with the experiment protocol above, we introduced
stochasticity of the spike timing by adding a random jitter ξΔt to the spike timing: ΔtSTDP = mΔt + ξΔt, were mΔt is
the average spike timing and ξΔt is a random variable whose distribution is given by the STDP protocol. Here, we
explored five STDP protocols: (1) Protocol 0 consisted of pairings with no jittering, i.e. =
IPIi
F
1
pairings and ξΔt = 0. (1) Protocol 0 consisted of pairings with no jittering, i.e. =
IPIi
F
1
pairings and ξΔt = 0. Fpairings
(2) Protocol 1 consisted of deterministic IPIs and uniformly distributed spike timings (Fig. 2a1), i.e. =
∀i
IPI
,
i
F
1
pairings
, and the probability distribution function of the jitter is given by pairings
(2) Protocol 1 consisted of deterministic IPIs and uniformly distributed spike timings (Fi
=
∀i
IPI
,
i
F
1
pairings
, and the probability distribution function of the jitter is given by SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 12 www.nature.com/scientificreports/ ξ
ξ
σ
=
|
| <
σ
Δ
Δ
Δ
Δ
P(
)
if
,
0
otherwise
t
t
t
1
2
t (1) In eq. (1), σΔt defines the maximal jitter amplitude i.e. the level of noise in the protocol. To simulate
Protocol 1, we first set each presynaptic stimulation time using
=
t
i
F
prei
pairings. Postsynaptic times were then
fixed by tposti = tprei + mΔt + ξΔt,i − δ. In eq. (1), σΔt defines the maximal jitter amplitude i.e. the level of noise in the protocol. To simulate i
i In eq. (1), σΔt defines the maximal jitter amplitude i.e. the level of noise in the protocol. References Natural Firing Patterns Imply Low Sensitivity of Synaptic Plasticity to Spike Timing
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g
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p
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d d
l
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d h
d
d
l . E. The spike-timing dependence of plasticity. Neuron 75, 556–571 h
p
g
p
p
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(
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0. Froemke, R. C., Tsay, I. A., Raad, M., Long, J. D. & Dan, Y. SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y Animals and brain slice preparation ⁎
⁎
W
CaMKII
CaMKII
1
3 5
(3)
post
max (3) where CaMKII* and
⁎
CaMKIImax are the current concentration of activated (phosphorylated) CaMKII and its
maximum value. Our model also accounts for the biochemical pathways leading to the production of the endocannabinoids
2-arachidonoylglycerol (2-AG) and AEA, and their subsequent activation of CB1R (see S1 Text for further details). In our model, CB1R activation (yCB1R) controls the presynaptic weight Wpre according to the following rule: Ω
=
−
Θ
<
< Θ
+
Θ
<
y
A
y
A
y
(
)
1
if
1
if
1
otherwise
(4)
CB1R
LTD
LTD
start
CB1R
LTD
stop
LTP
LTP
start
CB1R Ω
=
−
Θ
<
< Θ
+
Θ
<
y
A
y
A
y
(
)
1
if
1
if
1
otherwise
(4)
CB1R
LTD
LTD
start
CB1R
LTD
stop
LTP
LTP
start
CB1R (4) and Ω
τ
=
−
W
dt
y
W
d
/
( (
)
)/ W
pre
CB1R
pre
pre (5) Details about model implementation and numerics are given in the supplementary information. Details about model implementation and numerics are given in the supplementary information. SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 13 www.nature.com/scientificreports/ Acknowledgements g
We thank the L.V. lab members for helpful suggestions and critical comments, in particular Sebastien Valverde
for proofreading of the manuscript. This work was supported by grants from INSERM, Collège de France, the
Agence Nationale pour la Recherche (Grant Dopaciumcity ANR-14-NEUC-0003), INRIA, CNRS, and the Ecole
des Neurosciences de Paris. www.nature.com/scientificreports/ & Le Novère, N. DARPP-32 is a robust integrator of dopamine and glutamate signals. PLoS
Comput. Biol. 2, e176 (2006). Author Contributions L.V. and H.B. designed the study; Y.C. and A.M. performed electrophysiological experiments and analysis; H.B. carried out the conception and the design of the mathematical model; H.B. and I.P. performed the acquisition and
analysis of data from the mathematical model; H.B. and L.V. wrote the manuscript and all authors have edited and
corrected the manuscript; L.V. supervised the whole study. www.nature.com/scientificreports/ www.nature.com/scientificreports/ 5. Wang, W. et al. A Primary Cortical Input to Hippocampus Expresses a Pathway-Specific and Endocannabinoid-Dependent Form o
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y
p
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ti T
i i
t R t B
l C
t
C
b C
t
1 14 htt
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h
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k
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via endocannabinoids. J. Physiol. (Lond.) 590, 2305–2315 (2012). SCIENTIFIC RePorTS | (2018) 8:8139 | DOI:10.1038/s41598-018-26436-y 14 Additional Information Supplementary information accompanies this paper at https://doi.org/10.1038/s41598-018-26436-y Competing Interests: The authors declare no competing interests. Competing Interests: The authors declare no competing interests. Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and
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Identification and evaluation of circulating small extracellular vesicle microRNAs as diagnostic biomarkers for patients with indeterminate pulmonary nodules
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Zheng et al. Journal of Nanobiotechnology (2022) 20:172
https://doi.org/10.1186/s12951-022-01366-0 Zheng et al. Journal of Nanobiotechnology (2022) 20:172
https://doi.org/10.1186/s12951-022-01366-0 Journal of Nanobiotechnology Open Access Identification and evaluation of circulating
small extracellular vesicle microRNAs
as diagnostic biomarkers for patients
with indeterminate pulmonary nodules Di Zheng1†, Yuming Zhu2†, Jiyang Zhang3†, Wei Zhang3†, Huizhen Wang3, Hao Chen3, Chunyan Wu4,
Jian Ni1, Xiaoya Xu3, Baoning Nian3, Sheng Chen3, Beibei Wang3, Xiaofang Li3, Yanan Zhang3, Jiatao Zhang5,
Wenzhao Zhong5, Lei Xiong3, Fugen Li3, Dadong Zhang3*, Jianfang Xu1* and Gening Jiang2* Abstract Background: The identification of indeterminate pulmonary nodules (IPNs) following a low-dose computed tomog-
raphy (LDCT) is a major challenge for early diagnosis of lung cancer. The inadequate assessment of IPNs’ malignancy
risk results in a large number of unnecessary surgeries or an increased risk of cancer metastases. However, limited
studies on non-invasive diagnosis of IPNs have been reported. Methods: In this study, we identified and evaluated the diagnostic value of circulating small extracellular vesicle
(sEV) microRNAs (miRNAs) in patients with IPNs that had been newly detected using LDCT scanning and were sched-
uled for surgery. Out of 459 recruited patients, 109 eligible patients with IPNs were enrolled in the training cohort
(n = 47) and the test cohort (n = 62). An external cohort (n = 99) was used for validation. MiRNAs were extracted from
plasma sEVs, and assessed using Small RNA sequencing. 490 lung adenocarcinoma samples and follow-up data were
used to investigate the role of miRNAs in overall survival. Results: A circulating sEV miRNA (CirsEV-miR) model was constructed from five differentially expressed miRNAs
(DEMs), showing 0.920 AUC in the training cohort (n = 47), and further identified in the test cohort (n = 62) and in
an external validation cohort (n = 99). Among five DEMs of the CirsEV-miR model, miR-101-3p and miR-150-5p were
significantly associated with better overall survival (p = 0.0001 and p = 0.0069). The CirsEV-miR scores were calcu-
lated, which significantly correlated with IPNs diameters (p < 0.05), and were able to discriminate between benign *Correspondence: dadong.zhang@3dmedcare.com; xujianfang63@aliyun. com; geningjiang@tongji.edu.cn †Di Zheng, Yuming Zhu, Jiyang Zhang and Wei Zhang contributed
equally to this work 1 Department of Medical Oncology, Shanghai Pulmonary Hospital, Tongji
University School of Medicine, No. 507 Zhengmin Road, Yangpu District,
Shanghai 200433, China 2 Department of Thoracic Surgery, Shanghai Pulmonary Hospital, Tongji
University School of Medicine, No. 507 Zhengmin Road, Yangpu District,
Shanghai 200433, China g
3 3D Medicines Inc, Building 2A, No. 158 Xin Junhuan Road, Pujiang
Hi‑tech Park, Shanghai 201114, China
Full list of author information is available at the end of the article © The Author(s) 2022. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the
original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or
other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line
to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory
regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this
licence, visit http://creativecommons.org/licenses/by/4.0/. The Creative Commons Public Domain Dedication waiver (http://creativeco
mmons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated in a credit line to the data. Page 2 of 17 Zheng et al. Journal of Nanobiotechnology (2022) 20:172 and malignant PNs (diameter ≤ 1 cm). The expression patterns of sEV miRNAs in the benign, adenocarcinoma in situ/
minimally invasive adenocarcinoma, and invasive adenocarcinoma subgroups were found to gradually change with
the increase in aggressiveness for the first time. Among all DEMs of the three subgroups, five miRNAs (miR-30c-5p,
miR-30e-5p, miR-500a-3p, miR-125a-5p, and miR-99a-5p) were also significantly associated with overall survival of
lung adenocarcinoma patients. Conclusions: Our results indicate that the CirsEV-miR model could help distinguish between benign and malignant
PNs, providing insights into the feasibility of circulating sEV miRNAs in diagnostic biomarker development. Trial registration: Chinese Clinical Trials: ChiCTR1800019877. Registered 05 December 2018, https://www.chictr.org.cn/
showproj.aspx?proj=31346. Keywords: Indeterminate pulmonary nodule, Low-dose computed tomography, Small extracellular vesicle,
microRNA, Small RNA sequencing Graphical Abstract LDCT to improve its ability of distinguishing between
benign and malignant PNs among IPNs is urgently
needed. Background The training cohort consisted of 47 patients with IPNs,
including 17 benign PNs (benign group) and 30 malig-
nant PNs (malignant group) identified using pathological
diagnosis; the test cohort consisted of 62 patients with
IPNs, including 24 benign PNs and 38 malignant PNs. Benign PNs were used as controls. In addition, an exter-
nal validation cohort (n = 99), including 20 patients with
benign PNs and 79 patients with malignant PNs, was
used to validate the model (Fig. 1 and Table 1), and 11
healthy people were as healthy control. More than 90% of
LUAD patients (92.6%, 63/68) were in the early stage (no
lymph node metastasis), and 33.0% (36/109) of patients
had IPNs with a diameter less than 1 cm (Table 1). The
demographic and clinicopathologic characteristics of the
patients are shown in Table 1. Benign PNs included cases
of atypical adenomatous hyperplasia (AAH), fibrosis,
granulomas, hamartoma, organizing pneumonias (OPs),
cyst, and other benign subtypes, while malignant PNs
included AIS, MIA, and invasive adenocarcinoma. The
pathology subtype composition of the three cohorts is
shown in (Additional file 1: Figure S2). The representative
imaging features of benign and malignant PNs are shown
in (Additional file 1: Figure S3). of lung cancer patients have also been found to be signifi-
cantly different from those of healthy people, indicating
that serum/plasma sEV miRNAs as potential biomarkers
of lung cancer [13, 14]. Some studies have attempted to
distinguish malignant PNs using plasma sEV miRNAs
[15–17], however, these studies included healthy people
as controls to develop a diagnostic model, and lacks fur-
ther investigation in biological difference between benign
and malignant PNs. Zhang and colleagues managed to
distinguish malignant ground-glass nodules and benign
nodules using plasma sEV miRNAs, but the sample sizes
were relatively small [18]. The discrimination between
benign and malignant PNs are quite difficult to identify,
yet very important for accurate diagnosis of lung cancer
after LDCT scanning in clinical practice. g
In the present study, we assessed the expression levels
of circulating sEV miRNAs using small RNA sequenc-
ing to detect the differences between patients with
benign PNs and patients with malignant PNs (early- to
mid-stage lung adenocarcinoma). We then developed a
CirsEV-miR model to differentiate between benign and
malignant PNs in a training cohort (n = 47) and further
confirmed the model in a test cohort (n = 62) and in an
external validation cohort (n = 99). Participants and clinical characteristics
Th
kfl
f
d
ll The workflow of our study is illustrated in Fig. 1 To
explore circulating sEV miRNAs as diagnostic biomarkers
in patients with lung IPNs, a total of 199 and 260 patients
were recruited in the training phase and the test phase,
respectively. After the elimination of hemolysis, non-
lung adenocarcinoma (non-LUAD), failure of sequencing Background Lung cancer is the most frequent malignancy with the
highest morbidity and mortality worldwide [1]. In the
U.S. National Lung Cancer Screening Test (NLST) trial
[2], low-dose computed tomography (LDCT) has been
proven to be a highly sensitive method to detect early
stage lung cancer in patients with a smoking history. LDCT scanning has dramatically improved the abil-
ity to detect pulmonary nodules (PNs); however, a total
of 96.4% of false positive screening results mandates
improvements in the management of indeterminate pul-
monary nodules (IPNs), which are difficult to diagnose. The inadequate assessment of IPNs’ malignancy risk is a
major cause of misdiagnosis and mistreatment. Benign
PNs require no surgical resection, however, the rates of
benign PNs undergoing surgical resection have been
reported to range from 1 to 30% in suspected lung cancer
cases [3]. The invasive thoracic procedure of unnecessary
operations performed in patients with benign PNs results
in unnecessary costs, societal burden, and morbidity and
mortality risk to the patient, with no therapeutic benefit
[4, 5]. Hence, a non-invasive auxiliary diagnostic test for Extensive efforts using blood biomarkers, such as
DNA, RNA, and proteins to distinguish malignant from
benign PNs have yielded novel insights into lung cancer
diagnosis [6]. Small extracellular vesicles (sEVs), secreted
by a variety of cells into the blood, contain bioactive mol-
ecules such as proteins, lipids, and nucleic acids that can
mirror the cellular origin and the physiological state,
and these molecules are attractive potential biomark-
ers representing the “fingerprint” or “signature” of the
donor cell [7]. Moreover, the membranous structure of
the sEVs protects the luminal contents, avoiding degra-
dation by extracellular enzymes. The remarkable stability
and activity strengthen the potential of circulating sEVs
to be reservoirs for biomarker development [8, 9]. Micro-
RNAs (miRNAs) have been found to be the most abun-
dant species among plasma-derived sEVs RNAs [10], and
are also notably stable under different storage conditions
[11]. Lung cancer cells secrete more sEVs into blood than
normal tissue cells [12], and miRNAs derived from sEVs Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 3 of 17 Zheng et al. Journal of Nanobiotechnology (2022) 20:172 library construction, and limitation of the ratio of benign
and malignant PNs, a total of 109 patients were included
in two independent cohorts (Additional file 1: Figure S1). Circulating sEV characterization Circulating sEVs were successfully isolated from patient
plasma samples and characterized using western blot
(WB) analysis, nanoparticle tracking analysis (NTA), and
transmission electron microscopy (TEM). In accordance
with the Minimal Information for Studies of Extracellular
Vesicles (MISEV) 2018 [19], several protein markers were
evaluated using WB in eight representative sEV samples
from the patients with benign and malignant PNs. The
expression levels of TSG101, CD63, CD9, and Syntenin
were detected in the eight sEV samples, while a negative
marker, Calnexin, was absent in all eight sEV samples
(Fig. 2a). Furthermore, the majority of the isolated sEVs
were around 100 nm in diameter, which is the typical size
of sEVs (Fig. 2b). The TEM result from a representative
sample showed that the isolated sEVs were cup-shaped
(Fig. 2c), which is the typical morphology of sEVs. Fur-
thermore, we also detected sEV transmembrane proteins
(CD63, CD81, CD9) using ExoView platform. As shown
in Fig. 2d, CD63, CD81, and CD9 were all detected in
plasma sEVs of patients with benign or malignant PNs. Background Furthermore, we cal-
culated CirsEV-miR scores and for the first time explored
the relationship between CirsEV-miR scores and clini-
cal characteristics, including the diameter of IPNs and
the PN aggressiveness in the benign, adenocarcinoma
in situ (AIS)/minimally invasive adenocarcinoma (MIA),
and invasive adenocarcinoma. We also found that the
circulating sEV miRNA signature was able to discrimi-
nate between benign and malignant PNs with the diam-
eter ≤ 1 cm, which are otherwise difficult to distinguish
in clinical practice. In total, we analyzed circulating sEV
miRNAs of 208 patients with IPNs, providing the larg-
est sample size among the available studies. Addition-
ally, 490 lung adenocarcinoma samples and follow-up
data from The Cancer Genome Atlas (TCGA) were used
to investigate the role of miRNAs in overall survival. The
development of such non-invasive diagnostic test, i.e., the
CirsEV-miR model, may complement the highly sensitive
but insufficiently specific LDCT and be integrated into
the diagnostic algorithm to achieve higher diagnostic
accuracy for patients with IPNs. Construction and validation of a circulating sEV miRNA
(CirsEV‑miR) model for IPN diagnosis Construction and validation of a circulating sEV miRNA
(CirsEV‑miR) model for IPN diagnosis
We then extracted the RNA from sEVs and per-
formed small RNA sequencing. Raw data of small RNA We then extracted the RNA from sEVs and per-
formed small RNA sequencing. Raw data of small RNA Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 4 of 17 Training Cohort
(n = 47)
Shanghai Pulmonary Hospital
4/2019 - 5/2019
Benign (n = 17)
AIS/MIA (n = 13)
Invasive (n = 17)
Exosome miRNAs
small RNA sequencing
Candidate miRNAs
(6 miRNAs)
CirsEV-miR model
(5 miRNAs)
Target genes function analysis
Survival analysis (TCGA)
Test Cohort
(n = 62)
Shanghai Pulmonary Hospital
9/2019 - 10/2019
Benign (n = 24)
AIS/MIA (n = 17)
Invasive (n = 21)
CirsEV-miR model perfomance
(ROC analysis)
Perfomance in IPNs ≤ 1 cm
CirsEV-miR model perfomance
(ROC analysis)
Discrimination of AIS/MIA
(CirsEV-miR score analysis)
miRNA expression differences
Survival analysis (TCGA)
External Validation Cohort
(n = 99)
Guangdong Provincial
People’s Hospital
7/2017 - 3/2018
Benign (n = 20)
AIS/MIA (n = 15)
Invasive (n = 64)
Model perfomance validation
(ROC analysis)
Model application in IPNs ≤ 1 cm
Different
combinations
Fig. 1 Overall study design and patients in the training, test, and external validation cohorts. AIS adenocarcinoma in situ; MIA minimally invasive
carcinoma, ROC receiver operating characteristic Training Cohort
(n = 47)
Shanghai Pulmonary Hospital
4/2019 - 5/2019
Benign (n = 17)
AIS/MIA (n = 13)
Invasive (n = 17)
Exosome miRNAs
small RNA sequencing
Candidate miRNAs
(6 miRNAs)
CirsEV-miR model
(5 miRNAs)
Target genes function analysis
Survival analysis (TCGA)
CirsEV-miR model perfomance
(ROC analysis)
Different
combinations Fig. 1 Overall study design and patients in the training, test, and external validation cohorts. AIS adenocarcinoma in situ; MIA minimally invasive
carcinoma, ROC receiver operating characteristic Fig. Integration of five miRNAs achieved an even higher AUC
of 0.904 (let-7b-3p, miR-125b-5p, miR-197-3p, miR-
150-5p, and miR-3168), while the AUC of the integra-
tion of all six miRNAs dropped to 0.794. We then used
LASSO-penalized regression to develop a classifier of
five miRNAs (let-7b-3p, miR-125b-5p, miR-150-5p, miR-
101-3p, and miR-3168), called the CirsEV-miR model,
which exhibited an AUC of 0.920 in the training cohort
(Fig. 3d, Additional file 1: Table S2, Additional file 1:
Table S3), which was the highest AUC among all clas-
sifiers. The sensitivity reached 0.900, and the specificity
was 0.882. Construction and validation of a circulating sEV miRNA
(CirsEV‑miR) model for IPN diagnosis Based on DEMs, CirsEV-miR scores were also
generated through LASSO analysis. CirsEV-miR scores
of malignant PNs were significantly higher than those
of benign PNs in the training cohort (Fig. 3e, p < 0.0001),
suggesting that CirsEV-miR scores increase with malig-
nancy. The CirsEV-miR model was further confirmed in
the test cohort, showing an AUC of 0.763 (Additional
file 1: Figure S4b) and significant differences in CirsEV-
miR scores between benign and malignant IPNs (Addi-
tional file 1: Figure S4c, p = 0.0004). The CirsEV-miR
model was also validated in the external cohort con-
sisting of 20 patients with benign PNs and 79 patients
with malignant PNs; the AUC in that cohort was 0.781 sequencing were filtered and normalized. Unsupervised
hierarchical clustering based on the expression levels of
circulating sEV miRNAs categorized the samples in a
similar pattern as the clinicopathologic classifications
did, both in the training cohort (Fig. 3a) and in the test
cohort (Additional file 1: Figure S4a), thereby suggest-
ing the feasibility of predicting clinicopathologic clas-
sifications using circulating sEV miRNA expression. Six
DEMs between benign and malignant PNs were identi-
fied in the training cohort (Fig. 3b), and these DEMs were
selected to construct a model for distinguishing benign
PNs from malignant PNs. To verify the potential of miR-
NAs as a suitable classifier, receiver operating character-
istic (ROC) analysis was performed, and the area under
curve (AUC) was calculated (Fig. 3c, Additional file 1:
Table S1, Additional file 1: Table S2). Let-7b-3p exhib-
ited an AUC of 0.875, which was the best performance of
an individual miRNA. Next, we integrated two or more
miRNAs to further improve the performance. Integration
of let-7b-3p and miR-125b-5p led to a slightly improved
AUC of 0.886; integrating three miRNAs achieved an
AUC of 0.892 (let-7b-3p, miR-125b-5p, and miR-197-3p);
and integrating four miRNAs achieved an AUC of 0.882
(let-7b-3p, miR-125b-5p, miR-197-3p, and miR-150-5p). Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 5 of 17 Table 1 The demographic and clinicopathologic characteristics of the patients in three cohorts
AIS adenocarcinoma in situ, MIA minimally invasive adenocarcinoma, AAH atypical adenomatous hyperplasia, OP organizing pneumonias, NA not available
Characteristic
Training cohort
(n = 47)
Testing cohort
(n = 62)
External cohort
(n = 99)
Categories—no. (%)
Benign
17 (36.2)
24 (38.7)
20 (20.2)
Malignant
30 (63.8)
38 (61.3)
79 (79.8)
Age, mean (SD)
58.7 (12.5)
58.1 (10.7)
57.9 (11.9)
Age, no. Construction and validation of a circulating sEV miRNA
(CirsEV‑miR) model for IPN diagnosis (%)
< 60
24 (51.1)
28 (45.2)
49 (49.5)
≥ 60
23 (48.9)
34 (54.8)
50 (50.5)
Gender, no. (%)
Female
20 (42.6)
37 (59.7)
55 (55.6)
Male
27 (57.4)
25 (40.3)
44 (44.4)
Smoking status, no. (%)
NA
Yes
12 (25.5)
6 (9.7)
No
35 (74.5)
56 (90.3)
Pathology, no. (%)
AIS
3 (6.4)
6 (9.7)
10
MIA
10 (21.3)
11 (17.7)
5
Invasive
17 (36.2)
21 (33.9)
66
AAH
2 (4.3)
2 (3.2)
0
Fibrosis
2 (4.3)
4 (6.5)
0
Granulomas
4 (8.5)
3 (4.8)
0
Hamartoma
2 (4.3)
3 (4.8)
3
OP
4 (8.5)
4 (6.5)
0
Other benign subtypes
3 (6.4)
8 (12.9)
15
Nodule diameter (cm), no. (%)
≤ 1
12 (25.5)
24 (38.7)
21 (21.2)
> 1
35 (74.5)
38 (61.3)
78 (78.8)
Malignant stages, no. (%)
NA
0
3 (10.0)
6 (15.8)
IA1
12 (40.0)
13 (34.2)
IA2
5 (16.7)
11 (28.9)
IA3
6 (20.0)
4 (10.5)
IB
1 (3.3)
0
IIA
2 (6.7)
0
IIB
0
1 (2.6)
IIIA
1 (3.3)
3 (7.9) Table 1 The demographic and clinicopathologic characteristics of the patients in three cohorts n situ, MIA minimally invasive adenocarcinoma, AAH atypical adenomatous hyperplasia, OP organizing pneumonias, NA not available (Additional file 1: Figure S4d). We were also interested
in finding the difference in the circulating sEV miR-
NAs between healthy participants and lung cancer and
benign pulmonary nodule patients. Therefore, to exam-
ine the discrimination value of CirsEV-miR score, the
blood specimens from 11 healthy participants were col-
lected and used to analyze the expression levels of the
circulating sEV miRNAs using small RNA sequencing. The results showed that healthy controls could be signifi-
cantly different from the lung cancer (Additional file 1:
Figure S4e, p = 3.07E-06) and benign pulmonary nodule (Additional file 1: Figure S4d). We were also interested
in finding the difference in the circulating sEV miR-
NAs between healthy participants and lung cancer and
benign pulmonary nodule patients. Therefore, to exam-
ine the discrimination value of CirsEV-miR score, the
blood specimens from 11 healthy participants were col-
lected and used to analyze the expression levels of the
circulating sEV miRNAs using small RNA sequencing. Construction and validation of a circulating sEV miRNA
(CirsEV‑miR) model for IPN diagnosis a Western blotting identified sEV proteins of eight representative samples,
including the sEV positive markers TSG101, CD63, CD9, Systenin, and the negative marker Calnexin. b Nanoparticle tracking analysis (NTA)
results from representative sEV samples. c Transmission electron microscopic (TEM) images of representative sEVs. d Representative spots and
immunofluorescence staining of CD63 (red), CD81 (green), and CD9 (blue) d
CD63
CD81
Benign
10 μm
10 μm
10 Merge
10 μm d CD9
10 μm 10 μm 10 μm 10 μm Fig. 2 Characterization of circulating small extracellular vesicles (sEVs). a Western blotting identified sEV proteins of eight representative samples,
including the sEV positive markers TSG101, CD63, CD9, Systenin, and the negative marker Calnexin. b Nanoparticle tracking analysis (NTA)
results from representative sEV samples. c Transmission electron microscopic (TEM) images of representative sEVs. d Representative spots and
immunofluorescence staining of CD63 (red), CD81 (green), and CD9 (blue) Construction and validation of a circulating sEV miRNA
(CirsEV‑miR) model for IPN diagnosis The results showed that healthy controls could be signifi-
cantly different from the lung cancer (Additional file 1:
Figure S4e, p = 3.07E-06) and benign pulmonary nodule patients (Additional file 1: Figure S4e, p = 0.015), indi-
cating that the CirsEV-miR score has great capability of
discriminating IPNs compared to healthy people. Expres-
sion of the five miRNAs of the CirsEV-miR model was
also verified by quantitative reverse-transcription PCR in
26 patients with benign or malignant PNs, showing simi-
lar patterns with sequencing results (Additional file 1:
Figure S4f). These results revealed that the CirsEV-miR
model may be a new promising approach to assist in the
differential diagnosis of lung IPNs. Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 6 of 17 a
0
100
200
300
400
500
Diameter (nm)
0
0.5
1.0
1.5
2.0
Concentration (particles / ml)
80
115
180
272
428
b
c
TSG101
44KD
Syntenin
32KD
CD63
26KD
CD9
25KD
Calnexin
90KD
Benign
Malignant
1
2
3
4
1
2
3
4
d
10 μm
CD63
Merge
CD81
CD9
Benign
Malignant
10 μm
10 μm
10 μm
10 μm
10 μm
10 μm
10 μm
200 nm
Fig. 2 Characterization of circulating small extracellular vesicles (sEVs). a Western blotting identified sEV proteins of eight representative samples,
including the sEV positive markers TSG101, CD63, CD9, Systenin, and the negative marker Calnexin. b Nanoparticle tracking analysis (NTA)
results from representative sEV samples. c Transmission electron microscopic (TEM) images of representative sEVs. d Representative spots and
immunofluorescence staining of CD63 (red), CD81 (green), and CD9 (blue) a
TSG101
44KD
Syntenin
32KD
CD63
26KD
CD9
25KD
Calnexin
90KD
Benign
Malignant
1
2
3
4
1
2
3
4 0
100
200
300
400
500
Diameter (nm)
0
0.5
1.0
1.5
2.0
Concentration (particles / ml)
80
115
180
272
428
b c
200 nm b a c 0
100
200
300
400
500
Diameter (nm)
0
0.5
1.0
1.5
2.0
Concentration (particles / ml)
80
115
180
272
428
TSG101
44KD
Syntenin
32KD
CD63
26KD
CD9
25KD
Calnexin
90KD
Benign
Malignant
1
2
3
4
1
2
3
4
d
10 μm
CD63
Merge
CD81
CD9
Benign
Malignant
10 μm
10 μm
10 μm
10 μm
10 μm
10 μm
10 μm
200 nm
Fig. 2 Characterization of circulating small extracellular vesicles (sEVs). Biological function enrichment and overall survival
analysis of five DEMs in the CirsEV‑miR model Biological function enrichment and overall survival
analysis of five DEMs in the CirsEV‑miR model Biological function enrichment and overall survival
analysis of five DEMs in the CirsEV‑miR model to tumorigenesis and progression processes, such as
MAPK, TGFβ, Hippo, p53 signaling pathways, cell
cycle, and adherence junction processes (Fig. 4c, d). Moreover, 490 lung adenocarcinoma samples and their
follow-up data from the TCGA showed that patients
with high expression of miR-101-3p or miR-150-5p
in tumor tissue samples had better overall survival
(Fig. 4e, f, p = 0.0001; p = 0.0069). However, the other
three miRNAs, let-7b-3p, miR-125b-5p, and miR-3168,
were not significantly associated with overall survival
(data not shown). The functional enrichment analysis
indicated that five miRNAs were involved in tumori-
genesis and progression of lung cancer. To evaluate the potential functions of the five miR-
NAs used in the CirsEV-miR model, we performed
Gene Ontology (GO) and pathway analysis of their
target genes (Additional file 1: Table S4). We found
that 2238 mRNAs were targeted by the five miRNAs
through the analysis of the miRNA target database. The target genes were enriched in biological processes
(BPs), such as cellular nitrogen compound metabolic
process, biosynthetic process, and cellular protein
modification processes (Fig. 4a), and in molecular
functions (MFs), such as ion binding, enzyme bind-
ing, and RNA binding (Fig. 4b). In addition, the target
genes were significantly enriched in pathways related Page 7 of 17 Zheng et al. Biological function enrichment and overall survival
analysis of five DEMs in the CirsEV‑miR model Journal of Nanobiotechnology (2022) 20:172 Cluster
Histology
Cluster
Cluster 1
Cluster 2
Histology
Benign
AIS/MIA
Invasive
-2
0
2
Normalized
Expression
Benign Malignant
0
3
6
9
let-7b-3p
Log2CPM
****
Benign Malignant
0
6
12
18
miR-101-3p
**
Log2CPM
Log2CPM
Benign Malignant
3
6
9
12
miR-125b-5p
***
Benign Malignant
1
4
7
10
miR-150-5p
Log2CPM
***
Benign Malignant
1
3
5
7
miR-197-3p
Log2CPM
**
Benign Malignant
1
4
7
10
miR-3168
Log2CPM
*
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.886
Two miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.892
Three miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.882
Four miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.904
Five miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.794
Six miRNA classifier
a
b
c
d
e
One miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.875
0.0
0.5
1.0
1.5
Benign
Malignant
CirsEV miR score
****
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.920
CirsEV-miR model
Fig. 3 Classifier development process for IPNs. a Heatmap of sEV-miRNA expression in the training cohort by unsupervised hierarchical clustering. b Expression levels of six identified DEMs in the training cohort. c ROCs of classifiers constructed with different combinations of the six DEMs. Only
ROCs with the best diagnostic powers of different miRNA combinations are shown. d ROC curve of the CirsEV-miR model in the training cohort. Biological function enrichment and overall survival
analysis of five DEMs in the CirsEV‑miR model e
CirsEV-miR scores of benign and malignant PNs in the training cohort Cluster
Histology
Cluster
Cluster 1
Cluster 2
Histology
Benign
AIS/MIA
Invasive
-2
0
2
Normalized
Expression
a Benign Malignant
0
3
6
9
let-7b-3p
Log2CPM
****
Benign Malignant
0
6
12
18
miR-101-3p
**
Log2CPM
Log2CPM
Benign Malignant
3
6
9
12
miR-125b-5p
***
Benign Malignant
1
4
7
10
miR-150-5p
Log2CPM
***
Benign Malignant
1
3
5
7
miR-197-3p
Log2CPM
**
Benign Malignant
1
4
7
10
miR-3168
Log2CPM
*
b
d b a 3
6
Log2CPM d c
One miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.875 c d d
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.920
CirsEV-miR model Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.892
Three miRNA classifier Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.886
Two miRNA classifier
c
One miRNA classifier
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.875 Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.886
Two miRNA classifier Specificity
0.6
0.4 e
0.0
0.5
1.0
1.5
Benign
Malignant
CirsEV miR score
****
Specificity e p
y
Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.794
Six miRNA classifier Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.882
Four miRNA classifier Specificity
Sensitivity
1.0
0.8
0.6
0.4
0.2
0.0
0.0
0.2
0.4
0.6
0.8
1.0
AUC: 0.904
Five miRNA classifier Six miRNA classifier Fig. 3 Classifier development process for IPNs. a Heatmap of sEV-miRNA expression in the training cohort by unsupervised hierarchical clustering. b Expression levels of six identified DEMs in the training cohort. c ROCs of classifiers constructed with different combinations of the six DEMs. Only
ROCs with the best diagnostic powers of different miRNA combinations are shown. d ROC curve of the CirsEV-miR model in the training cohort. e
CirsEV-miR scores of benign and malignant PNs in the training cohort The signature of circulating sEV miRNAs discriminated correlation between the nodule size and risk score
of the CirsEV-miR model. All 109 eligible samples
were recategorized into two subgroups: the IPNs
with diameter ≤ 1 cm and IPNs with diameter > 1 cm,
and re-ranked according to the CirsEV-miR scores. The CirsEV-miR scores were significantly higher in
the subgroup of IPNs with diameter > 1 cm (Fig. 5a,
p = 0.0170), suggesting that the CirsEV-miR scores The signature of circulating sEV miRNAs discriminated
between benign and malignant PNs with diameter ≤ 1 cm. The size of IPNs is a key factor that is associated with
their malignant potential and patients’ long term sur-
vival [20, 21]. In clinical practice, IPNs with a diameter
> 1 cm are thought to need resection, whereas IPNs
with a diameter ≤ 1 cm require repeated scans and a
long-term follow-up [22]. Thus, we also analyzed the g
g
between benign and malignant PNs with diameter ≤ 1 cm.h The size of IPNs is a key factor that is associated with
their malignant potential and patients’ long term sur-
vival [20, 21]. In clinical practice, IPNs with a diameter
> 1 cm are thought to need resection, whereas IPNs
with a diameter ≤ 1 cm require repeated scans and a
long-term follow-up [22]. Thus, we also analyzed the Zheng et al. The signature of circulating sEV miRNAs discriminated Journal of Nanobiotechnology (2022) 20:172 Page 8 of 17 endosome
focal adhesion
cytoplasmic stress granule
microtubule organizing center
cytosol
nucleoplasm
protein complex
SMAD binding
small conjugating protein binding
virus receptor activity
translation factor activity, nucleic acid binding
enzyme regulator activity
cytoskeletal protein binding
poly(A) RNA binding
nucleic acid binding transcription factor activity
protein binding transcription factor activity
RNA binding
enzyme binding
ion binding
0
25
50
−log10(P−value)
Number
200
400
600
Category
Biological Process
Cellular Component
Molecular Function
b innate immune response
immune system process
cell cycle
transcription, DNA−templated
nucleobase−containing compound catabolic process
response to stress
cell death
catabolic process
cellular component assembly
small molecule metabolic process
neurotrophin TRK receptor signaling pathway
cellular protein modification process
gene expression
biosynthetic process
cellular nitrogen compound metabolic process
0
30
60
90
−log10(P−value)
tra
nuc
a
b innate immune response
immune system process
cell cycle
transcription, DNA−templated
nucleobase−containing compound catabolic process
response to stress
cell death
catabolic process
cellular component assembly
small molecule metabolic process
neurotrophin TRK receptor signaling pathway
cellular protein modification process
gene expression
biosynthetic process
cellular nitrogen compound metabolic process
0
30
60
90
−log10(P−value)
endosome
focal adhesion
cytoplasmic stress granule
microtubule organizing center
cytosol
nucleoplasm
protein complex
SMAD binding
small conjugating protein binding
virus receptor activity
translation factor activity, nucleic acid binding
enzyme regulator activity
cytoskeletal protein binding
poly(A) RNA binding
nucleic acid binding transcription factor activity
protein binding transcription factor activity
RNA binding
enzyme binding
ion binding
0
25
50
−log10(P−value)
Number
200
400
600
Category
Biological Process
Cellular Component
Molecular Function
a
b
Fatty acid metabolism
Biosynthesis of unsaturated fatty acids
Hippo signaling pathway
TGF−beta signaling pathway
MAPK signaling pathway
Adherens junction
Cell cycle
Regulation of actin cytoskeleton
p53 signaling pathway
Endocytosis
Focal adhesion
mRNA surveillance pathway
12
Fatty acid biosynthesis
Lysine degradation
0
6
−log10(P−value)
Number
10
25
50
Categrey
Genetic Information Processing
Cellular Processes
Environmental Information Processing
Metabolism
Viral carcinogenesis
Proteoglycans in cancer
Transcriptional misregulation in cancer
Alcoholism
MicroRNAs in cancer
Epstein−Barr virus infection
Pathways in cancer
Herpes simplex infection
PI3K−Akt signaling pathway
Hippo signaling pathway
AMPK signaling pathway
FoxO signaling pathway
Wnt signaling pathway
Rap1 signaling pathway
cGMP−PKG signaling pathway
MAPK signaling pathway
Ras signaling pathway
cAMP signaling pathway
Adherens junction
Cell cycle
Regulation of actin cytoskeleton
Focal adhesion
Signaling pathways regulating pluripotency of stem cells
Endocytosis
Tight junction
Neurotrophin signaling pathway
Dopaminergic synapse
Adrenergic signaling in cardiomyocytes
RNA transport
52
50
37
33
52
31
75
28
65
39
25
27
30
38
29
51
39
26
30
37
49
45
30
41
25
25
27
25
39
0
20
40
60
Annotated Genes
Genetic Information Processing
Organismal Systems
Cellular Processes
Environmental
Information
Processing
Human Diseases
c
d
e
f
+++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++
+++
++++++
+
+
++++++++++++++++++++++++++++++++++++++++++++++++++++++
+
++++++
+
miR-101-3p
0.00
0.25
0.50
0.75
1.00
0
50
100
150
200
Time (months)
Overall survival
+
Top 25% (n=122)
+
Bottom 25% (n=122)
122
21
4
3
1
122
15
1
0
0
Bottom 25%
Top 25%
p = 0.0001
+++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++
+++++++++
+++++++
+
++
+++++++++++++++++++++++++++++++++++++++++++++++++++++++++
++++++++
++++
0.00
0.25
0.50
0.75
1.00
0
50
100
150
200
Time (months)
122
17
6
3
1
122
13
0
0
0
Overall survival
+
Top 25% (n=122)
+
Bottom 25% (n=122)
p = 0.0069
Bottom 25%
Top 25%
miR-150-5p
Fig. The signature of circulating sEV miRNAs discriminated e, f Overall
survival analysis of miR-101-3p (e) and miR-150-5p (f) from the TCGA database b a log10(P value)
Fatty acid metabolism
Biosynthesis of unsaturated fatty acids
Hippo signaling pathway
TGF−beta signaling pathway
MAPK signaling pathway
Adherens junction
Cell cycle
Regulation of actin cytoskeleton
p53 signaling pathway
Endocytosis
Focal adhesion
mRNA surveillance pathway
12
Fatty acid biosynthesis
Lysine degradation
0
6
−log10(P−value)
Number
10
25
Categrey
Genetic Information Processing
Cellular Processes
S
c
d log10(P value)
Fatty acid metabolism
Biosynthesis of unsaturated fatty acids
Hippo signaling pathway
TGF−beta signaling pathway
MAPK signaling pathway
Adherens junction
Cell cycle
Regulation of actin cytoskeleton
p53 signaling pathway
Endocytosis
Focal adhesion
mRNA surveillance pathway
12
Fatty acid biosynthesis
Lysine degradation
0
6
−log10(P−value)
Number
10
25
50
Categrey
Genetic Information Processing
Cellular Processes
Environmental Information Processi
Metabolism
Signa
c
d
e
+++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++
++++++++++++++++++++++++++++++++++++
miR-101-3p
0.75
1.00
+
Top 25% (n=122)
+
Bottom 25% (n=122) d sing
Viral carcinogenesis
Proteoglycans in cancer
Transcriptional misregulation in cancer
Alcoholism
MicroRNAs in cancer
Epstein−Barr virus infection
Pathways in cancer
Herpes simplex infection
PI3K−Akt signaling pathway
Hippo signaling pathway
AMPK signaling pathway
FoxO signaling pathway
Wnt signaling pathway
Rap1 signaling pathway
cGMP−PKG signaling pathway
MAPK signaling pathway
Ras signaling pathway
cAMP signaling pathway
Adherens junction
Cell cycle
Regulation of actin cytoskeleton
Focal adhesion
Signaling pathways regulating pluripotency of stem cells
Endocytosis
Tight junction
Neurotrophin signaling pathway
Dopaminergic synapse
Adrenergic signaling in cardiomyocytes
RNA transport
52
50
37
33
52
31
75
28
65
39
25
27
30
38
29
51
39
26
30
37
49
45
30
41
25
25
27
25
39
0
20
40
60
Annotated Genes
Genetic Information Processing
Organismal Systems
Cellular Processes
Environmental
Information
Processing
Human Diseases
d 25
50
Cellular Processes
Environmental Information Processing
Metabolism
e
+++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++
+++
++++++
+
+
++++++++++++++++++++++++++++++++++++++++++++++++++++++
+
++++++
+
miR-101-3p
0.00
0.25
0.50
0.75
1.00
0
50
100
150
200
Time (months)
Overall survival
+
Top 25% (n=122)
+
Bottom 25% (n=122)
122
21
4
3
1
122
15
1
0
0
Bottom 25%
Top 25%
p = 0.0001 +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++
+++++++++
+++++++
+
++
+++++++++++++++++++++++++++++++++++++++++++++++++++++++++
++++++++
++++
0.00
0.25
0.50
0.75
1.00
0
50
100
150
200
Time (months)
122
17
6
3
1
122
13
0
0
0
Overall survival
+
Top 25% (n=122)
+
Bottom 25% (n=122)
p = 0.0069
Bottom 25%
Top 25%
miR-150-5p f e Fig. The signature of circulating sEV miRNAs discriminated 4 Target analysis and survival analysis of the five DEMs used for CirsEV-miR model development. A bubble plot of enriched GOs ((Biological
process, (a), and (Cellular component and Molecular Function, (b)) of target genes of the five DEMs. c A bubble plot of enriched KEGG pathways of
target genes of the five DEMs. d A bar plot showing the number of DEM target genes in each of the KEGG pathways (target genes > 20). Circulating sEV miRNA analysis reveals that AIS/
MIA can alleviate the differentiation between benign
and malignant PNs remarkably increase with the diameter of IPNs. More-
over, among the patients with small IPNs (diameter
≤ 1 cm), malignant PNs also exhibited higher CirsEV-
miR scores than benign PNs (Fig. 5b, p = 0.0088). The
AUC of the CirsEV-miR model in patients with small
IPNs (diameter ≤ 1 cm) was 0.767 in our cohort (Addi-
tional file 1: Figure S5a), and 0.721 in the external vali-
dation cohort (Additional file 1: Figure S5b). Moreover,
we achieved the diagnostic specificity of 91.7% for the
IPNs with diameter ≤ 1 cm in our cohort, and speci-
ficity of 75% in the external validation cohort (Fig. 5c). Among the IPNs with diameter ≤ 1 cm, benign and
malignant subgroups showed different miRNA expres-
sion patterns (Fig. 5d). Twelve DEMs between benign
and malignant PNs with diameter ≤ 1 cm were identi-
fied (Fig. 5e), and eight of them were involved in lung
cancer tumorigenesis and progression, suggesting that
even small IPNs were different from benign PNs. These
results suggest that even in IPNs with diameter ≤ 1 cm,
the circulating sEV miRNA signature can separate
benign PNs from malignant PNs. remarkably increase with the diameter of IPNs. More-
over, among the patients with small IPNs (diameter
≤ 1 cm), malignant PNs also exhibited higher CirsEV-
miR scores than benign PNs (Fig. 5b, p = 0.0088). The
AUC of the CirsEV-miR model in patients with small
IPNs (diameter ≤ 1 cm) was 0.767 in our cohort (Addi-
tional file 1: Figure S5a), and 0.721 in the external vali-
dation cohort (Additional file 1: Figure S5b). Moreover,
we achieved the diagnostic specificity of 91.7% for the
IPNs with diameter ≤ 1 cm in our cohort, and speci-
ficity of 75% in the external validation cohort (Fig. 5c). Among the IPNs with diameter ≤ 1 cm, benign and
malignant subgroups showed different miRNA expres-
sion patterns (Fig. 5d). Twelve DEMs between benign
and malignant PNs with diameter ≤ 1 cm were identi-
fied (Fig. 5e), and eight of them were involved in lung
cancer tumorigenesis and progression, suggesting that
even small IPNs were different from benign PNs. These
results suggest that even in IPNs with diameter ≤ 1 cm,
the circulating sEV miRNA signature can separate
benign PNs from malignant PNs. AIS and MIA represent early-stage lung cancer, and both
AIS and MIA patients possess superior prognoses to
invasive adenocarcinoma [23, 24]. The signature of circulating sEV miRNAs discriminated (b) The CirsEV-miR
scores of benign and malignant subgroups of IPNs ≤ 1 cm. c Model sensitivity and specificity of IPNs ≤ 1 cm. d Gene expression heatmap of benign
and malignant subgroups of IPNs ≤ 1 cm. e Differentially expressed miRNAs between the benign and malignant subgroups of IPNs ≤ 1 cm The signature of circulating sEV miRNAs discriminated 4 Target analysis and survival analysis of the five DEMs used for CirsEV-miR model development. A bubble plot of enriched GOs ((Biological
process, (a), and (Cellular component and Molecular Function, (b)) of target genes of the five DEMs. c A bubble plot of enriched KEGG pathways of
target genes of the five DEMs. d A bar plot showing the number of DEM target genes in each of the KEGG pathways (target genes > 20). e, f Overall
survival analysis of miR-101-3p (e) and miR-150-5p (f) from the TCGA database Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 9 of 17 Malignant
Benign
Pathology
a
b
c
d
e
Normalized
Expression
-2
0
2
0
1
2
3
miR−125a−5p
miR−125b−5p
miR−181a−3p
miR−195−3p
miR−197−3p
miR−199a−5p
miR−328−3p
miR−378d
miR−4508
miR−450b−5p
miR−500a−3p
miR−6087
−1.0
−0.5
0.0
0.5
1.0
-Log10P-value
Log2FC
IPNs ≤1 cm
Pathology
Predicted
Benign
Predicted
Malignant
Sensitivity
Specificity
Internal
Benign (12)
11
1
54.2%
91.7%
Malignant (24)
11
13
External
Benign (4)
3
1
82.4%
75.0%
Malignant (17)
3
14
−0.3
0.0
0.3
0.6
0.9
1.2
<=1
>1
*
CirsEV-miR score
**
−0.3
0.0
0.3
0.6
0.9
1.2
Benign Malignant
CirsEV-miR score
Fig. 5 The CirsEV-miR scores increased with the increase in the diameter of IPNs. a CirsEV-miR scores of IPNs ≤ 1 cm and > 1 cm. (b) The CirsEV-miR
scores of benign and malignant subgroups of IPNs ≤ 1 cm. c Model sensitivity and specificity of IPNs ≤ 1 cm. d Gene expression heatmap of benign
and malignant subgroups of IPNs ≤ 1 cm. e Differentially expressed miRNAs between the benign and malignant subgroups of IPNs ≤ 1 cm c
IPNs ≤1 cm
Pathology
Predicted
Benign
Predicted
Malignant
Sensitivity
Specificity
Internal
Benign (12)
11
1
54.2%
91.7%
Malignant (24)
11
13
External
Benign (4)
3
1
82.4%
75.0%
Malignant (17)
3
14 a
b
−0.3
0.0
0.3
0.6
0.9
1.2
<=1
>1
*
CirsEV-miR score
**
−0.3
0.0
0.3
0.6
0.9
1.2
Benign Malignant
CirsEV-miR score b
**
−0.3
0.0
0.3
0.6
0.9
1.2
Benign Malignant
CirsEV-miR score b a c e
0
1
2
3
miR−125a−5p
miR−125b−5p
miR−181a−3p
miR−195−3p
miR−197−3p
miR−199a−5p
miR−328−3p
miR−378d
miR−4508
miR−450b−5p
miR−500a−3p
miR−6087
−1.0
−0.5
0.0
0.5
1.0
-Log10P-value
Log2FC d e e Fig. 5 The CirsEV-miR scores increased with the increase in the diameter of IPNs. a CirsEV-miR scores of IPNs ≤ 1 cm and > 1 cm. Circulating sEV miRNA analysis reveals that AIS/
MIA can alleviate the differentiation between benign
and malignant PNs We further
compared all DEMs between the benign, AIS/MIA, and
invasive adenocarcinoma subgroups (Fig. 6d, e, Addi-
tional file 1: Table S5, Additional file 1: Table S6). We only
found two overlapping upregulated DEMs (let-7b-3p and
miR-125b-5p), which were also elements of our CirsEV-
miR model. Thirteen miRNAs (six upregulated and seven
downregulated) with repeated emergence were found
in Benign_vs_AIS/MIA and Benign_vs_Invasive adeno-
carcinoma subgroups, showing some similarity in sEV
miRNA profiles of AIS/MIA and invasive adenocarci-
noma. Meanwhile, three upregulated miRNAs and four
downregulated miRNAs were found in Benign_vs_AIS/
MIA alone, while nine miRNAs showed altered expres-
sion in invasive adenocarcinoma compared with AIS/
MIA (three upregulated and six downregulated), sug-
gesting difference in sEV-miRNA profiles between AIS/
MIA and invasive adenocarcinoma. Among all DEMs,
five miRNAs (miR-30c-5p, miR-30e-5p, miR-500a-3p,
miR-125a-5p, and miR-99a-5p) were significantly asso-
ciated with overall survival from the TCGA database
(Fig. 6f, p = 0.0008; p = 0.0090; p = 0.0110; p = 0.0310,
p = 0.0007). Taken together, the results of the circulating
sEV miRNA analysis demonstrated that the presence of
AIS/MIA could affect the differentiation between benign
and malignant PNs, and that AIS/MIA presented the
intermediate molecular features of circulating sEV miR-
NAs between benign and malignant PNs. we found that this score remarkably increased with the
diameter of IPNs, and then gradually changed with the
increase in aggressiveness. Moreover, we found that the
molecular features of circulating sEV miRNAs of AIS/
MIA were between benign and invasive adenocarcino-
mas, meanwhile, AIS/MIA also possessed unique circu-
lating sEV miRNA profiles different from the other two
subgroups. Furthermore, 490 lung cancer samples and
follow-up data from TCGA were used to investigate
the role of miRNAs in overall survival. In summary, the
CirsEV-miR model developed using circulating sEV miR-
NAs could serve as a non-invasive auxiliary test to help
physicians to manage the IPNs for early-stage lung can-
cer diagnosis. The differentially expressed circulating sEV
miRNAs identified in this study might further serve as
potential therapeutic targets in future investigations.h This study focused on IPNs and enrolled the patients
with highly suspicious malignant PNs undergoing sur-
gery (404/459, confirmed by pathology diagnosis after
surgery) from the routine work-ups in a tertiary hospital. Unlike previous studies that focused on early lung can-
cer screening in low-prevalence populations, our study
recruited patients with a high prevalence, and we used
benign PNs as controls. (See figure on next page.)
Fig. 6 AIS/MIA are intermediate between benign and malignant PNs. a The CirsEV-miR scores of the benign, AIS/MIA, and invasive adenocarcinoma
subgroups categorized according to the pathologies. b Gene expression heatmap of all genes in the benign, AIS/MIA and invasive adenocarcinoma
subgroups. c The expression levels of the five DEMs in the benign, AIS/MIA, and invasive adenocarcinoma subgroups. d, e Venn diagrams of
upregulated (d) or downregulated (e) genes of the benign, AIS/MIA, and IA subgroups. f Overall survival analysis of miR-30c-5p, miR-30e-5p,
miR-500a-3p, miR-125a-5p, and miR-99a-5p from the TCGA database. ***, p < 0.001; **, p < 0.01; *, p < 0.05; ns, not significant Circulating sEV miRNA analysis reveals that AIS/
MIA can alleviate the differentiation between benign
and malignant PNs The discrimination between
malignant and benign PNs was more difficult than that
between the nodule population and non-nodule con-
trols [25]. Benign PNs in this study contained a variety of
pathological subtypes without selection, reflecting real-
world clinical situations. The benign PNs in the exter-
nal validation cohort contained different pathological
subtypes from those in the training and test cohorts, yet
yielding no significant impairment in the model perfor-
mance, which indicates that our CirsEV-miR model may
have broad clinical application potential. Circulating sEV miRNA analysis reveals that AIS/
MIA can alleviate the differentiation between benign
and malignant PNs Therefore, we specu-
lated that the possible reason for the relatively low sen-
sitivity of the current model was the misprediction of
the clinicopathologically equivocal AIS/MIA samples. To test this hypothesis, 109 eligible samples were catego-
rized into three subgroups: benign (41, 37.6%), AIS/MIA
(30, 27.5%), and invasive adenocarcinoma (38, 34.9%). The proportions of AIS/MIA samples in the training
and the test cohorts were 27.7% and 27.4%, respectively
(Table 1). For all 109 samples, we calculated the CirsEV-
miR scores and ranked each subgroup. The CirsEV-miR
scores in the benign, AIS/MIA and invasive adenocar-
cinoma subgroups gradually changed with the increase
in aggressiveness (Fig. 6a). We also tested the miRNA
expression levels; we found that the expression pattern
of the AIS/MIA subgroup was intermediate between
benign and invasive adenocarcinoma subgroups (Fig. 6b). The expression levels of the five DEMs in the AIS/MIA Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 10 of 17 subgroup were located between the benign and the inva-
sive adenocarcinoma subgroups (Fig. 6c). We further
compared all DEMs between the benign, AIS/MIA, and
invasive adenocarcinoma subgroups (Fig. 6d, e, Addi-
tional file 1: Table S5, Additional file 1: Table S6). We only
found two overlapping upregulated DEMs (let-7b-3p and
miR-125b-5p), which were also elements of our CirsEV-
miR model. Thirteen miRNAs (six upregulated and seven
downregulated) with repeated emergence were found
in Benign_vs_AIS/MIA and Benign_vs_Invasive adeno-
carcinoma subgroups, showing some similarity in sEV
miRNA profiles of AIS/MIA and invasive adenocarci-
noma. Meanwhile, three upregulated miRNAs and four
downregulated miRNAs were found in Benign_vs_AIS/
MIA alone, while nine miRNAs showed altered expres-
sion in invasive adenocarcinoma compared with AIS/
MIA (three upregulated and six downregulated), sug-
gesting difference in sEV-miRNA profiles between AIS/
MIA and invasive adenocarcinoma. Among all DEMs,
five miRNAs (miR-30c-5p, miR-30e-5p, miR-500a-3p,
miR-125a-5p, and miR-99a-5p) were significantly asso-
ciated with overall survival from the TCGA database
(Fig. 6f, p = 0.0008; p = 0.0090; p = 0.0110; p = 0.0310,
p = 0.0007). Taken together, the results of the circulating
sEV miRNA analysis demonstrated that the presence of
AIS/MIA could affect the differentiation between benign
and malignant PNs, and that AIS/MIA presented the
intermediate molecular features of circulating sEV miR-
NAs between benign and malignant PNs. subgroup were located between the benign and the inva-
sive adenocarcinoma subgroups (Fig. 6c). Discussion With the development of LDCT scanning for early
screening and early diagnosis of lung cancer, the number
of detected IPNs has dramatically increased. The inad-
equate assessment of IPNs’ malignancy risk is a major
cause of a large number of unnecessary surgeries or an
increased risk of cancer metastases; hence, the accurate
diagnosis of IPNs becomes one of the major challenges
in clinic. However, a non-invasive diagnostic test to
assist LDCT to distinguish benign PNs from malignant
PNs in IPN diagnosis is not yet available. In this study,
we included 208 patients with IPNs, which is the largest
sample size among the available studies; and we devel-
oped, tested, and validated the CirsEV-miR model to
discriminate between benign and malignant PNs using
small RNA sequencing of circulating sEV miRNAs. We
also calculated a CirsEV-miR score from the model, and The diameter is an important parameter in assess-
ing the malignant potential of an IPN. The possibility
of malignancy positively correlates with the diameter. Lung cancer probability was low in IPNs smaller than
5 mm, intermediate in IPNs with a diameter of
5–10 mm, and high in IPNs larger than 10 mm [22]. Clinically, IPNs > 10 mm would be considered to need
resection, while IPNs ≤ 10 mm would require repeated
scans. We found that CirsEV-miR score was signifi-
cantly higher in IPNs > 10 mm than in IPNs ≤ 10 mm. Even in IPNs ≤ 10 mm, CirsEV-miR score of malignant
PNs was significantly higher than that of benign PNs
(Fig. 5a, b), suggesting that malignancy may increase Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 11 of 17 Fig. 6 (See legend on previous page.) Fig. 6 (See legend on previous page.) Fig. 6 (See legend on previous page ) Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 12 of 17 miRNAs bind to the 3ʹ-untraslated region (UTR) of tar-
get mRNA, resulting in post-translational gene silencing
either by mRNA degeneration or by inhibition of transla-
tion. Thousands of miRNAs have been linked to various
human diseases, including cancers [34]. In this study,
we identified differentially expressed miRNAs between
benign and malignant PNs and constructed the diagnos-
tic model composed of five DEMs using LASSO-penal-
ized regression. Discussion Functional enrichment analysis revealed
that the target genes of the five miRNAs of our diagnostic
model were involved in many cancer-related pathways,
such as MAPK, TGFβ, Hippo, p53 signaling pathway,
focal adhesion, and cell cycle (Fig. 4c, d). We also found
sEV-miRNAs enriched in immune system processes and
metabolic processes, in accordance with the previous
report that intratumor heterogeneity and immunoedit-
ing are early phenomena in PNs [29]. Moreover, we found
seven miRNAs (miR-101-3p, miR-150-5p, miR-30c-5p,
miR-30e-5p, miR-500a-3p, miR-125a-5p, and miR-
99a-5p) that were associated with better overall survival
based on the TCGA databank (Figs. 4e, f and 6f). Interest-
ingly, these miRNAs have all been reported to function as
tumor suppressors [35–40], although some of them were
upregulated in tumor tissues [37, 39], and in circulating
sEVs of patients with malignant PNs in our study, indicat-
ing that further research is required to clarify the molecu-
lar mechanism underlying benign and malignant PNs. with IPNs diameter. Our CirsEV-miR model also had
good performance in IPNs with a diameter of less
than 1 cm, exhibiting AUC of 0.767 and specificity of
91.7% in our cohorts, and AUC of 0.721 and specific-
ity of 75.0% in the external validation cohort (Fig. 5c,
Additional file 1: Figure S5). Taken together, these
findings indicate that our model has the ability to dis-
tinguish between benign and malignant PNs, even in
IPNs ≤ 10 mm, thereby expanding the applications
widely.i According to IASLC/ATS/ERS and 2015 WHO classifi-
cation of lung tumors, AIS is a tumor smaller than 3 cm
with no invasion, and MIA denotes tumor smaller than
3 cm with invasion less than 0.5 cm [26]. Both AIS and
MIA patients achieve 100% DFS with complete resec-
tion [27]. It is believed that LUAD develops stepwise from
benign AAH to AIS, to MIA, and eventually to overt
invasive adenocarcinoma [28]. However, most studies
have focused on histopathological subtypes, while little is
known about the genomic evolution from preinvasive to
invasive adenocarcinoma of LUAD. In 2019, Zhang et al. reported that mutations in EGFR, ERBB2, NRAS, and
BRAF, as well as genomic intratumor heterogeneity and
immunoediting, are common and early phenomena har-
bored by AIS, MIA, and IA, and that mutations in TP53, as
well as cell mobility and metastasis-related genes, may be
later events during neoplastic progression [29]. Discussion Our results
also demonstrated that the expression profiles of circulat-
ing sEV miRNAs in patients with AIS/MIA were inter-
mediate between the patients with benign PNs and the
patients with invasive adenocarcinoma (Fig. 6b), support-
ing the stepwise development hypothesis. Yet, we found
seven expression-altered miRNAs unique to AIS/MIA
compared with benign PNs, and nine miRNAs unique
to AIS/MIA compared with invasive adenocarcinoma
(Fig. 6d, e, Additional file 1: Table S5, Additional file 1:
Table S6). Among them, miR-30e-5p, a well-known tumor
suppressor that suppresses tumorigenesis via the Sirt1/
JAK/STAT3 signaling pathway [30], was downregulated
in both AIS/MIA and invasive adenocarcinoma compared
with benign PNs. However, miR-25-3p and miR-128-3p,
which have been reported to promote metastasis, exhib-
ited higher expression in AIS/MIA than in invasive adeno-
carcinoma [31, 32]. These findings suggest that AIS/MIA
exhibit unique features different from benign PNs or inva-
sive adenocarcinoma. It is also proposed that due to a lack
of signals in AIS/MIA specimens, the difficulty in distin-
guishing benign PNs from AIS/MIA IPNs is an intrinsic
feature [33]. This study is the first report that revealed the
circulating sEV miRNA signatures of benign, AIS/MIA,
and invasive adenocarcinoma, suggesting that the pres-
ence of AIS/MIA could impact the differentiation between
benign and malignant PNs. Several limitations of this study are worth mention-
ing. First, this was a single-center case control study that
was performed in a high prevalence population. Thus,
the performance of the CirsEV-miR model needs to be
explored further in a community where a low prevalence
is expected. Second, a prospective validating study was
not conducted. Third, the study lacks in vitro experi-
ments to verify the miRNA expression level in lung ade-
nocarcinoma tissues. To overcome the above limitations,
a prospective multicenter clinical trial in a larger popula-
tion-based setting is ongoing, and the expression profiles
and biological functions of the identified miRNAs need
to be validated in vitro in lung adenocarcinoma tissues or
in lung adenocarcinoma cell lines. Conclusion In summary, our study profiled circulating sEV miRNAs
in patients with IPNs and provided a diagnostic model,
the CirsEV-miR model, based on the measurement of
circulating sEV miRNAs to distinguish benign PNs from
malignant PNs to assist LDCT scanning for early-stage
lung cancer diagnosis. Five circulating sEV miRNAs
(let-7b-3p, miR-101-3p, miR-125b-5p, miR-150-5p, and
miR-3168) were revealed, and the CirsEV-miR model
consisting of these miRNAs was established and vali-
dated in 208 patients with IPNs, which is the largest Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 13 of 17 not a malignant disease. Considering the perspective of
the pathological progression of lung adenocarcinoma,
we still classified AIS as a type of malignant nodule. The
pathological subtypes of benign PNs were without selec-
tion and comprised more than 10 subtypes. The patho-
logical information of all of the samples was obtained
from surgically resected tissue sections in accordance
with the 2015 WHO Histological Classification of Lung
Cancer [26]. The pathological diagnosis of each patient
was confirmed by two pathologists. The tumor–node–
metastasis (TNM) stage was determined in accordance
with the 8th edition International Association for the
Study of Lung Cancer (IASLC) lung cancer staging sys-
tem [41]. The pathological subtypes of our training and
test cohorts, and those of the external validation cohort,
are shown in (Additional file 2: Figure S2). sample size so far. Moreover, CirsEV-miR could discrimi-
nate between benign and malignant PNs with diam-
eter ≤ 1 cm, which are mostly difficult to distinguish in
clinical setting. We primarily revealed that the molecular
feature of AIS/MIA was intermediate of benign and inva-
sive PNs, while exhibiting its own characteristics. These
results suggest that circulating sEV miRNAs as diagnos-
tic biomarkers could be integrated with LDCT scan to
obtain further evaluation work-ups for IPNs. Patient enrollment and study designh This study was approved by the Institutional Review
Board in Shanghai Pulmonary Hospital affiliated with
Tongji University (K18-199Y) and registered at the Chi-
nese Clinical Trial Registry (http://www.chictr.org.
cn/) with registration number ChiCTR1800019877. All
patients were from Shanghai Pulmonary Hospital affili-
ated with Tongji University and had signed written con-
sents for their blood samples and clinical information to
be used in this study. i
The accuracy of a diagnostic test is usually measured by
its sensitivity and specificity [42]. In this study, sensitivity
represented the model’s ability to correctly identify indi-
viduals with malignant PNs, and specificity represented
the model’s ability to correctly identify individuals with
benign PNs. y
Patients with IPNs detected using LDCT scanning,
who subsequently underwent surgical resections and
diagnosed with LUAD (malignant PNs) and various
benign PNs, were enrolled in this study. 199 patients with
IPNs were recruited in the training phase from April to
May 2019, including 20 patients with benign PNs and
179 with malignant PNs diagnosed using pathological
examination. 260 patients with IPNs were recruited in
the test phase from September to October 2019, includ-
ing 35 patients with benign PNs and 225 patients with
malignant PNs. Plasma samples had been prospectively
collected in a vacutainer with anticoagulant (REF367863;
Becton Dickinson, Franklin Lakes, NJ, USA) prior to
surgical operation. After the elimination of pathology
samples from patients with non-LUAD (n = 65), seri-
ous hemolysis (above or equal to grade 5, n = 107), fail-
ure to meet the ratio of benign and malignant PNs set at
1:2 (n = 155), and failure in construction of sequencing
libraries (n = 23), finally, the training cohort consisted of
47 patients with IPNs (17 benign PNs and 30 malignant
PNs), and the test cohort consisted of 62 patients with
IPNs (24 benign PNs and 38 malignant PNs). In addition,
an external cohort consisting of 20 patients with benign
PNs and 79 patients with malignant PNs was used for
validation. In this case–control study, a variety of benign
PNs without selection served as controls. Additionally,
11 healthy people were enrolled as healthy control.The
whole study design and inclusion/exclusion criteria are
depicted in Fig. 1 and (Additional file 1: Figure S1). Methods Patient enrollment and study designh Western blot analysish y
The isolated sEVs were lysed in 200 μL lysis buffer
(P0013B, Beyotime, Shanghai, China); next, the pro-
teins were extracted using an isolation reagent (N3525,
3DMed, Shanghai, China). The protein concentration
of the sEVs was measured using a Pierce™ BCA Protein
Assay Kit (Thermo Fisher Scientific, USA). 20 µg of total
protein was resolved on a 12% SDS-PAGE gel, electro-
transferred onto a PVDF membrane (Millipore, USA). The membranes were blocked in 5% non-fat milk for
60 min, and incubated with anti- CD9 antibody (diluted
1:500; cat. no. ab92726; Abcam, Cambridge, UK), anti-
CD63 antibody (1:2000, ab216130; Abcam, Cambridge,
UK), anti-Syntenin antibody (diluted 1:500; cat. no. ab19903; Abcam, Cambridge, UK), anti-TSG101 poly-
clonal antibody (diluted 1:500; cat. no. abs115706; Absin
Bioscience Inc., Shanghai, China), and anti-Calnexin
antibody (diluted 1:1000; cat. no. 2679; Cell Signaling
Technology, Danvers, MA, USA) primary antibodies
overnight at 4 °C. Horseradish peroxidase-conjugated
goat anti-rabbit IgG and goat anti-mouse IgG antibodies
(Beyotime Biotechnology, China) were used as second-
ary antibodies. Antibody binding was detected using an
enhanced chemilluminescence system according to the
manufacturer’s protocol (Tanon-5200 Multi; Tanon Sci-
ence & Technology Co. Ltd., Shanghai, China). RNA isolation from sEVs RNA was extracted from sEVs using the miRNeasy
Serum/Plasma Kit (217184; QIAGEN, Shanghai, China)
in accordance with the manufacturer’s protocol. The
miRNA quality, yield, and distribution were analyzed
using the Agilent 2100 Bioanalyzer with Small RNA
Chips (5067-1548; Agilent, Savage, MD, USA). Transmission electron microscopy (TEM) Transmission electron microscopy (TEM) Transmission electron microscopy (TEM) added to the supernatant; gently inverted and incubated
for 30 min at 4 °C and then centrifuged at 4700×g for
30 min at 4 °C. Finally, the supernatant was removed and
the pellets containing the total sEVs were re-suspended
in 0.2 mL phosphate-buffered saline (PBS). For TEM analysis, plasma sEVs were suspended in PBS
prior to fixing in 4% paraformaldehyde and transferred to
the carbon-coated electron microscopy grids. They were
washed with PBS twice, and the third time with PBS con-
taining glycine (50 mM), each for 3 min; then, they were
incubated with PBS containing BSA (0.5%) for 10 min. Finally, the grids were stained with 2% uranyl acetate. After the staining, TEM (H-7650, Hitachi High-Tech-
nologies, Japan) was used to analyze the morphology of
sEVs. ExoView analysis Plasma sEVs were detected using ExoView chips
(NanoView Biosciences, Brighton, MA) printed with
antibodies against CD63, CD81, CD9, and mouse IgG1
as a negative control. 35 μL samples were dropped onto
the chip and incubated for 16 h. After washing, chips
were incubated with a fluorescence antibody cocktail of
anti-CD9 (CF® 488), anti- CD81 (CF® 555), and anti-
CD63 (CF® 647) for 1 h at room temperature. Chips were
then imaged in the ExoView R100 Scanner (NanoView
Biosciences, Brighton, MA). Data were analysed using
NanoViewer Software (NanoView Biosciences, Brighton,
MA). Plasma isolation and sEVs isolation Blood samples were collected from patients in 10-mL
vacutainer tubes containing an anticoagulant of K2EDTA
(REF367863; Becton Dickinson, Franklin Lakes, NJ,
USA), mixed by gently inverting several times, stored
with the tubes placed upright, and then transported on
ice within 1 h after collection. To harvest the plasma, the
samples were centrifuged at 1600×g for 10 min at 4 °C,
after which the hemolysis level was determined and
recorded. Samples with hemolysis grade of no more than
4 were used [43]. The collected supernatant was centri-
fuged again at 16,000×g for 15 min at 4 °C, and then the
1 mL supernatant was transferred into a fresh 1.5 mL
tube and stored at − 80 °C prior to use. For the sEV isolation from plasma, a polyethylene
glycol-based 3D Medicine isolation reagent [18] (L3525;
3DMed, Shanghai, China) was used. This isolation rea-
gent has been modified and improved based on the work
of Rider [44], and has been registered to the National
Medical Products Administration as a Class I medical
device (#HMXB20190091), specifically for the isolation
of sEVs in the clinical setting. The plasma samples were
centrifuged at 12,000×g for 10 min at 4 °C after a static
water bath incubation at 37 ℃ for 5 min. The superna-
tant was transferred to a 0.45 µm tube filter (CLS8163-
100EA; Costar, Corning, NY, USA), followed by transfer
to a 0.22 µm tube filter (CLS8161-100EA; Costar) and
then centrifuged at 12,000×g for 5 min at 4 °C. The fil-
tered supernatant was transferred to a fresh 1.5 mL tube. One-quarter volume of an isolation reagent (L3525) was i
The LUAD consisted of three pathological subtypes,
namely, AIS, MIA, and invasive adenocarcinoma. Nine
patients were diagnosed with AIS, which is technically Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Page 14 of 17 Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Zheng et al. Journal of Nanobiotechnology (2022) 20:172 Quantitative reverse‑transcription PCR Quantitative reverse‑transcription PCR
Total RNA extraction from sEVs were as previously
described. miRNA were reverse transcribed using
TaqMan™ Advanced miRNA cfDNA Synthesis Kit
(A28007, Applied Biosystems™, USA) according to
the manufacturer’s protocol. qPCR was performed on
Applied Biosystems 7500 Fast Real-Time PCR systems
with specific (miR-451a, miR-125b-5p, miR-101-3p,
miR-3168, miR-150-5p and let-7b-3p) probes (A25576,
Applied Biosystems™, USA). The expression level of miR-
451a were used as control as previously reported [52]. Relative expression were calculated with mean Ct values
using 2−ΔΔCt method. Supplementary Information The online version contains supplementary material available at https://doi.
org/10.1186/s12951-022-01366-0. The online version contains supplementary material available at https://doi.
org/10.1186/s12951-022-01366-0. Additional file 1: Figure S1. Cohorts’ details and inclusion/exclusion
criteria. Figure S2. Pathology subtypes of the training (a), test (b), and
external validation (c) cohorts. Figure S3. Representative pathological
images of the benign and malignant pulmonary nodule subtypes. Malig-
nant nodule subtypes: adenocarcinoma in situ (AIS), minimally invasive
adenocarcinoma (MIA), and invasive adenocarcinoma (IA); Benign nodule
subtypes: granulomas, atypical adenomatous hyperplasia (AAH), hamar-
toma, cyst, fibrosis, organizing pneumonias (OP). Magnification, ×400;
Formalin Fixed Paraffin-Embedded (FFPE) tissues. Figure S4. CirsEV-miR
model performance in the test cohort and the external cohort. (a) Circu-
lating sEV miRNA heatmap of the test cohort by unsupervised hierarchical
clustering. (b) ROC curve of the CirsEV-miR model in the test cohort. (c)
The CirsEV-miR scores of benign and malignant PNs in the test cohort. (d) ROC curve of the CirsEV-miR model in the external validation cohort. (e) The CirsEV-miR scores of healthy people and patients with benign or
malignant PNs. (f) Expression level of the five miRNAs used in CirsEV-miR Bioinformatics analysis of small RNA sequencing datah informatics analysis of small RNA sequencing datah The 3′ adaptors of reads were cleaved using a custom
program. Subsequently, the reads were aligned to the
human genome hg19 assembly (http://hgdownload.soe.
ucsc.edu/goldenPath/hg19/bigZips/) using BWA 0.7.12
[45]. An individual Small RNA-Seq dataset is required
to have a minimum of 5,000,000 reads with minimum
mapping rate 80% that mapped with any annotated RNA
transcript in the human genome. The annotations were
generated from Gencode v25 [46] and miRBase v21 [47]
for statistical analysis and to determine expression levels. The annotation includes all small RNAs, such as miR-
NAs, rRNAs, tRNAs, and piRNAs, as well as long tran-
scripts from GENCODE, which includes both protein
coding genes and long non-coding RNAs (lincRNAs). The percentage of reads that mapped to the annotated
miRNAs should be greater than 25% (Additional file 1:
Table S7). The miRNA expressions were determined by
counting the number of reads mapped to the regions
annotated by mature miRNAs. The miRNA mapped by
at least two reads in each of the samples and with length
less than 30 nt was saved for miRNA expression analy-
sis. The miRNA expression analysis was performed using
the voom function in the limma package [48], with nor-
malization by Trimmed Mean of M-values (TMM) via
the edgeR package, and the miRNA expression level was
converted to log2-counts-per-million (logCPM) [49]. The Empirical Bayes algorithm implemented in ComBat
was applied to the training and the test cohort data sets
adjusted for batch effects [50, 51]. Statistical analysis was performed using the statistical
programming language R (version 3.6). The dendextend
package [53] in R was used to perform average linkage
hierarchical clustering of genes and cases. The heatmap
was constructed using the ComplexHeatmap package
[54] in R/Bioconductor. The biological processes of Gene
Ontology (GO) and the Kyoto Encyclopedia of Genes and
Genomes (KEGG) pathways enrichment of the experi-
mentally validated targets of miRNAs were examined
using mirPath v.3, which provided the Expression Analy-
sis Systematic Explorer (EASE) score and false-discovery
rates using the Fisher’s exact tests and unbiased empirical
distributions [55]. The Kaplan–Meier plot analysis of the
TCGA data was performed using OncoLnc [56]. Small RNA libraries preparation and sequencing Nanosight NS 300 system (NanoSight Technology, Mal-
vern, UK) was used to characterize the number and size
of EVs. Isolated sEVs were resuspended in PBS at a con-
centration of 5 μg/mL and were further diluted 100- to
1000-fold, to achieve between 20 and 100 objects/frame. Samples were manually injected into the sample cham-
ber at ambient temperature. Each sample was config-
ured using a 488 nm laser and a high-sensitivity scientific
complementary metal-oxide semiconductor camera, and
the measurements were performed in triplicate at a cam-
era setting of 13 with an acquisition time of 30 s and a
detection threshold setting of 7. At least 200 completed
tracks were analyzed and obtained per video. Finally, the
NTA analytical software (version 2.3) was used to analyze
the nanoparticle tracking data of the sEV samples in this
study. Small RNA libraries preparation and sequencing
To prepare and construct the small RNA sequencing
libraries, a NEB Next Multiplex Small RNA Library Prep
Set for Illumina (E7300L; New England Biolabs, Ipswich,
MA, USA) was used in accordance with the manufac-
turer’s protocol. Briefly, the reverse transcription primer
was hybridized after 3ʹ adaptor ligation of 100 ng RNA
per sample, following 5ʹ adaptor ligation. A total of 18
PCR cycles were performed with Illumina feasible bar-
code primers after the first strand cDNA synthesis. The
prepared libraries were resolved on NucleoSpin Gel and
PCR Clean-up (740609.50; MACHEREY–NAGEL, Ger-
many) and recovered in 30 μL DNase- and RNase-free
water. The DNA quality, yield, and distribution were
analyzed using the LabChip® GX Touch™ HT Nucleic
Acid Analyzer with DNA High Sensitivity Reagent Kit
(CLS760672; PerkinElmer, Waltham, MA, USA) and
the DNA Extended Range LabChip (CLS138948; Perki-
nElmer). A total of 20–25 libraries were pooled into a Page 15 of 17 Zheng et al. Journal of Nanobiotechnology (2022) 20:172 single sequencing lane and sequenced using an Illumina
HiSeq PE150 analyzer. and selection operator (LASSO) in the training cohort. The test cohort and the external cohort were used to test
and validate the diagnostic model. We selected the dif-
ferentially expressed sEV-miRNAs (DEMs) determined
according to the stringent statistical threshold (Student’s
t-test p-value ≤ 0.05, 1.5-fold change, and the mean
expression CPM ≥ 50) between the benign and malignant
PNs. Based on DEMs, the risk scores were generated
using LASSO analysis, and the best parameters of the
model constructed using LASSO were ultimately selected
using tenfold cross-validation. Abbreviations AAH: Atypical adenomatous hyperplasia; AIS: Adenocarcinoma in situ; AUC
: Area under curve; BPs: Biological functions; CPM: Counts-per-million; DEMs:
Differentially expressed miRNAs; GO: Gene ontology; IPNs: Indeterminate pul-
monary nodules; ISEV: International Society for Extracellular Vesicles; LASSO:
Least absolute shrinkage and selection operator; LDCT: Low-dose computed
tomography; LUAD: Lung adenocarcinoma; MFs: Molecular functions; MIA:
Minimally invasive adenocarcinoma; miRNAs: MicroRNAs; MISEV: Minimal
Information for Studies of Extracellular Vesicles; NTA: Nanoparticle tracking
analysis; OPs: Organizing pneumonias; OS: Overall survival; PNs: Pulmonary
nodules; ROC: Receiver operating characteristic; sEV: Small extracellular vesi-
cles; TCGA: The Cancer Genome Atlas; TEM: Transmission electron microscopy;
UTR: Untranslated region; WB: Western blot. Funding 11. Ge Q, Zhou Y, Lu J, Bai Y, Xie X, Lu Z. miRNA in plasma exosome is stable
under different storage conditions. Molecules. 2014;19(2):1568–75. 11. Ge Q, Zhou Y, Lu J, Bai Y, Xie X, Lu Z. miRNA in plasma exosome is stable
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**p < 0.01; ***, p < 0.001. Figure S5. CirsEV-miR model performance of
IPNs ≤ 1 cm. (a) CirsEV-miR model performance in our cohorts. (b) CirsEV-
miR model performance in the external validation cohort. Table S1. Clas-
sifiers constructed from the six identified DEMs. Table S2. Performance of
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Cancer Res. 2017;23(17):5311–9. Authors’ contributions Conceived and designed the experiments: GN-J, D-Z, DD-Z, JF-X. Performed
the experiments: YM-Z, CY-W, J-N, XX-X, BN-N, S-C, YN-Z, XF-L. Analyzed the
data: JY-Z, HZ-W, W-Z, H-C, CY-W, BB-W, DD-Z, FG-L. Contributed reagents/
materials/analysis: D-Z, L-X, JT-Z, WZ-Z, JF-X, GN-J. All authors read and
approved the final manuscript. 9. Boukouris S, Mathivanan S. Exosomes in bodily fluids are a highly stable
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Not applicable. Consent for publication
Not applicable. Declarations 14. Smolarz M, Widlak P. Serum exosomes and their mirna load-a potential
biomarker of lung cancer. Cancers. 2021;13(6):1373. Acknowledgements
h
h
ld l k The authors would like to thank Huiyan Li, and Chao Sun for their assistance
with sample collection for this study. The authors appreciate the support and
participation of the physicians and patients in this study. The authors would
like to thank LetPub (www.letpub.com.cn) for English language editing. 7. Zhang Y, Liu Y, Liu H, Tang WH. Exosomes: biogenesis, biologic function
and clinical potential. Cell Biosci. 2019;9:19. 8. Kalra H, Adda CG, Liem M, Ang CS, Mechler A, Simpson RJ, et al. Compara-
tive proteomics evaluation of plasma exosome isolation techniques and
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validation of extracellular vesicle-associated miRNAs as noninvasive
detection biomarkers for early-stage non-small-cell lung cancer. Mol
Oncol. 2021;15(9):2439–52. Authors affiliated with 3D Medicines Inc. are current or former employees. No
other disclosures were reported. 1.
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et al. Global cancer statistics 2020: GLOBOCAN estimates of incidence Ethics approval and consent to participate 15. Gao S, Guo W, Liu T, Liang N, Ma Q, Gao Y, et al. Plasma extracellular vesi-
cle microRNA profiling and the identification of a diagnostic signature for
stage I lung adenocarcinoma. Cancer Sci. 2022;113(2):648–59. All patients enrolled signed informed consent. This study was approved by
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https://openalex.org/W2946714351
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https://www.frontiersin.org/articles/10.3389/fgene.2019.00450/pdf
|
English
| null |
Personalized Interpretation and Clinical Translation of Genetic Variants Associated With Cardiomyopathies
|
Frontiers in genetics
| 2,019
|
cc-by
| 9,737
|
ORIGINAL RESEARCH
published: 15 May 2019
doi: 10.3389/fgene.2019.00450 Citation: Campuzano O,
Fernandez-Falgueras A,
Sarquella-Brugada G, Cesar S,
Arbelo E, García-Álvarez A, Jordà P,
Coll M, Fiol V, Iglesias A,
Perez-Serra A, Mates J, del Olmo B,
Ferrer C, Alcalde M, Puigmulé M,
Mademont-Soler I, Pico F, Lopez L,
Tiron C, Brugada J and Brugada R
(2019) Personalized Interpretation
and Clinical Translation of Genetic
Variants Associated With
Cardiomyopathies. Front. Genet. 10:450. doi: 10.3389/fgene.2019.00450 Keywords: sudden cardiac death, arrhythmias, cardiomyopathies, genetics, genetic counseling Campuzano O,
Fernandez-Falgueras A,
Sarquella-Brugada G, Cesar S,
Arbelo E, García-Álvarez A, Jordà P,
Coll M, Fiol V, Iglesias A,
Perez-Serra A, Mates J, del Olmo B,
Ferrer C, Alcalde M, Puigmulé M,
Mademont-Soler I, Pico F, Lopez L,
Tiron C, Brugada J and Brugada R
(2019) Personalized Interpretation
and Clinical Translation of Genetic
Variants Associated With
Cardiomyopathies.
Front. Genet. 10:450.
doi: 10.3389/fgene.2019.00450 Reviewed by: Reviewed by:
Serena Sanna,
University Medical Center Groningen,
Netherlands
Natasa Djordjevic,
University of Kragujevac, Serbia
Richard Hauer,
ICIN Netherlands Heart Institute
(KNAW), Netherlands Reviewed by:
Serena Sanna,
University Medical Center Groningen,
Netherlands
Natasa Djordjevic,
University of Kragujevac, Serbia
Richard Hauer,
ICIN Netherlands Heart Institute
(KNAW), Netherlands 1 Cardiovascular Genetics Center, Biomedical Research Institute of Girona, University of Girona, Girona, Spain, 2 Department
of Medical Science, School of Medicine, University of Girona, Girona, Spain, 3 Centro Investigación Biomédica Red
Enfermedades Cardiovasculares, Madrid, Spain, 4 Department of Biochemistry and Molecular Genetics, Hospital Clinic,
Barcelona, Spain, 5 Arrhythmias Unit, Hospital Sant Joan de Déu, University of Barcelona, Barcelona, Spain, 6 Arrhythmias
Unit, Hospital Clinic, University of Barcelona, Barcelona, Spain, 7 Cardiology Service, Hospital Josep Trueta, University
of Girona, Girona, Spain *Correspondence:
Oscar Campuzano
oscar@brugada.org
Ramon Brugada
ramon@brugada.org
†These authors have contributed
equally to this work Cardiomyopathies
are
a
heterogeneous
group
of
inherited
cardiac
diseases
characterized by progressive myocardium abnormalities associated with mechanical
and/or electrical dysfunction. Massive genetic sequencing technologies allow a
comprehensive genetic analysis to unravel the cause of disease. However, most
identified genetic variants remain of unknown clinical significance due to incomplete
penetrance and variable expressivity. Therefore, genetic interpretation of variants
and translation into clinical practice remain a current challenge. We performed
retrospective comprehensive clinical assessment and genetic analysis in six families, four
diagnosed with arrhythmogenic cardiomyopathy, and two diagnosed with hypertrophic
cardiomyopathy (HCM). Genetic testing identified three rare variants (two non-sense and
one small indel inducing a frameshift), each present in two families. Although each variant
is currently classified as pathogenic and the cause of the diagnosed cardiomyopathy,
the onset and/or clinical course differed in each patient. New genetic technology allows
comprehensive yet cost-effective genetic analysis, although genetic interpretation, and
clinical translation of identified variants should be carefully done in each family in a
personalized manner. Specialty section:
This article was submitted to
Genomic Medicine,
a section of the journal
Frontiers in Genetics
Received: 13 January 2019
Accepted: 30 April 2019
Published: 15 May 2019 Specialty section:
This article was submitted to
Genomic Medicine,
a section of the journal
Frontiers in Genetics Received: 13 January 2019
Accepted: 30 April 2019
Published: 15 May 2019 Edited by:
Lili Milani,
University of Tartu, Estonia Edited by:
Lili Milani,
University of Tartu, Estonia Oscar Campuzano1,2,3,4*†, Anna Fernandez-Falgueras1†, Georgia Sarquella-Brugada2,5,
Sergi Cesar5, Elena Arbelo3,6, Ana García-Álvarez6, Paloma Jordà6, Monica Coll1,
Victoria Fiol5, Anna Iglesias1,3, Alexandra Perez-Serra1,3, Jesus Mates1,
Bernat del Olmo1, Carles Ferrer1, Mireia Alcalde1,3, Marta Puigmulé1,
Irene Mademont-Soler1, Ferran Pico1, Laura Lopez1, Coloma Tiron7, Josep Brugada3,5,6
and Ramon Brugada1,2,3,7* INTRODUCTION Sudden cardiac death (SCD) is a major cause of death with an incidence ranging from
approximately 1:1000 to 1:3000 individuals per year in the general population (Deo and
Albert, 2012). In population <35 years old, inherited cardiomyopathies are the leading cause
of SCD, mainly hypertrophic cardiomyopathy (HCM), dilated cardiomyopathy (DCM), and May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 1 Cardiomyopathies: Personalized Diagnosis Campuzano et al. arrhythmogenic
cardiomyopathy
(ACM)
(Straus
et
al.,
2004). Cardiomyopathies
are
characterized
by
structural
heart abnormalities that induce mechanical and/or electrical
dysfunction, leading to ventricular arrhythmias and even SCD. Sometimes the first manifestation of disease is SCD, which usually
occurs in a healthy young individual while doing exercise. These
disorders induce progressive morphological and histological
myocardial features, but abnormalities may be minimal or even
absent at young ages (Gopinathannair et al., 2015). families. A multidisciplinary group including cardiologists,
pediatricians, genetic counselors, molecular geneticists, and
pathologists performed personalized genetic interpretation and
clinical translation in each family. Families diagnosed with
the same cardiomyopathy due to the same mutation were
included in our study. Clinical evaluation of index cases and all
available relatives included a complete physical examination, 12-
lead electrocardiogram (ECG), 2-dimentional echocardiography
(ECHO), cardiac magnetic resonance imaging (MRI), exercise
stress test, 24-h Holter monitoring, and comprehensive genetic
testing. Regarding post-mortem samples, a complete autopsy
was performed according to current international regulations,
including macroscopic, microscopic, and toxicological analyses
(Basso et al., 2008; Sinard, 2013). When macroscopic autopsy
was labeled negative, forensic pathologists performed a complete
histological and toxicological investigation, and collected a blood
sample for genetic investigation. We excluded cases in which
the autopsy was labeled as violent death, including death from
drug overdose. We also obtained family history of syncope and
unexplained deaths. In
post-mortem
cases,
current
guidelines
recommend
molecular autopsy in all cases without a conclusive cause of
death and without any cardiac alteration, but also if a cardiac
alteration suspicious of cardiomyopathy is identified (Priori
and Blomstrom-Lundqvist, 2015). In addition, ongoing clinical
surveillance for family members of affected individuals is
recommended because some relatives might have inherited
the genetic alteration and thus be at risk of SCD. Therefore,
establishing a proper pedigree including both clinical and
genetic data is crucial to provide a clear view of diagnosed
and asymptomatic relatives, focused on adopting preventive
therapeutic measures in each individual. Variant Analysis Genomic DNA was obtained from peripheral blood or saliva. DNA analyzed using NGS technology (MiSeq, Illumina). We
screened the most prevalent genes involved in cardiomyopathies
and other pathologies associated with SCD (ABCC9, ACTC1,
ACTN2,
AKAP9,
ANK2,
ANKRD1,
BAG3,
CACNA1C,
CACNA2D1,
CACNA1G,
CACNA1H,
CACNA1I,
CACNB2,
CALM1, CALM2, CALM3, CALR3, CASQ2, CAV3, CRYAB,
CSRP3, CTNNA3, GJA1, CTF1, DES, DMD, DMPK, DPP6, DSC2,
DSG2, DSP, DTNA, ECE1, EMD, EN1, EYA4, FHL2, FKTN,
FLNA, FLNC, GAA, GJA5, GLA, GPD1L, HCN1, HCN2, HCN4,
JPH2, JUP, KCNA5, KCND3, KCNE1, KCNE2, KCNE3, KCNE4,
KCNE5, KCNH2, KCNJ2, KCNJ5, KCNJ8, KCNQ1, LAMA4,
LAMP2, LDB3, LMNA, MYBPC3, MYH6, MYH7, MYL2, MYL3,
MYLK2, MYOZ2, MYPN, NEBL, NEXN, NOS1AP, NOTCH1,
NPPA, NUP155, PDLIM3, PHOX2A, PHOX2B, PITX2, PKP2,
PLN, PRKAG2, RANGRF, RBM20, RYR2, SCN1B, SCN2B,
SCN3B, SCN4B, SCN5A, SCN10A, SDHA, SGCD, SLC22A5,
SLC6A4, SLC8A1, SLMAP, SLN, SNTA1, TAZ, TCAP, TGFB3,
TLX3, TMEM43, TMPO, TNNC1, TNNI3, TNNT2, TP63, TPM1,
TRDN, TRIM63, TRPM4, TTN, TTR, and VCL) (Campuzano
et al., 2014a). All gene isoforms described in Ensembl 751
that have been linked at least with either a RefSeq code2 or
CCDS3 were included. Massive sequencing technologies such as next generation
sequencing (NGS) now allow comprehensive genetic analysis
of families in a cost-effective way – genetic data can be
obtained in reduced time and at low cost in comparison to
traditional Sanger sequencing (Morini et al., 2015). The main
current challenge is genetic interpretation as well as clinical
translation of these results, as most identified genetic variants
remain of unknown/ambiguous significance (Richards et al.,
2015; Amendola et al., 2016). This means genetic data are
not conclusive and do not help clinicians in the diagnosis,
risk stratification, or adoption of potential therapeutic measures
to prevent malignant arrhythmias. Consequently, genetic data
should be carefully translated into clinical practice. Information of family history, segregation of genetic variants,
and genotype-phenotype correlations are crucial to elucidate
the role of each genetic variant (Campuzano et al., 2015a). Further, the same genetic variant may be associated with
incomplete penetrance and variable expressivity, intrafamilial
as well as interfamilial, and complicating clinical interpretation
(Giudicessi and Ackerman, 2013). INTRODUCTION Hence, incorporation of
genetic testing and genetic counseling is now considered standard
of care for cardiomyopathies (Marian and Braunwald, 2017). Frontiers in Genetics | www.frontiersin.org 1http://www.ensembl.org
2http://www.ncbi.nlm.nih.gov/refseq
3https://www.ncbi.nlm.nih.gov/CCDS MATERIALS AND METHODS Alignment of DNA sequences for different species was
also performed for novel variations using Uniprot database16. Finally, protein structure and domains were consulted at
STRING17 and SMART databases18. Finally, each variant was classified in one of the following
current recommendations of the American College of Medical
Genetics and Genomics and the Association for Molecular
Pathology
(ACMG/AMP)
(Richards
et
al.,
2015). This
recommendations
describe
several
items
of
pathogenicity
(PVS, evidence of pathogenicity very strong; PS, evidence of
pathogenicity strong; PM, evidence of pathogenicity moderate;
and PP, evidence of pathogenicity supporting), and benignity
(BA, evidence of benign impact stand-alone; BS, evidence
of benign impact strong; and BP, evidence of benign impact
supporting), enabling a final score and consequent classification
of variants as: pathogenic (P), likely pathogenic (LP), variant of
uncertain significance (VUS), likely benign (LB), or benign (B). The PM2 item in the ACMG score was considered fulfilled if
MAF ≤0.1% in relevant population databases (Lek et al., 2016). Concerning frequency of disease-causing variants, the majority
of pathogenic variants are extremely rare (<0.01%) (Kobayashi
et al., 2017). Concerning potential PVS1 variants, it should
be only used for variants in genes where loss of function is a
previously established disease mechanism19 (Abou Tayoun et al.,
2018). In addition, some items of ACMG/AMP may underlie a
lack of specificity or ambiguous or contradictory interpretations,
so we check the parameters using Sherloc (semiquantitative,
hierarchical evidence-based rules for locus interpretation)
(Nykamp et al., 2017). Finally, to avoid bias, three authors with
PhDs in human genetics independently and comprehensively
investigated published genetic data concerning each analyzed
variant. In addition, four independent expert cardiologists
(including
two
pediatric
cardiologists)
comprehensively
reviewed available clinical data to reconfirm diagnoses following
current guidelines (Priori and Blomstrom-Lundqvist, 2015). All
investigators discussed data included in each item of the ACMG
and consensus and the final classification of all variants. Regarding copy number variation (CNV), our approach
focused on capturing significant differences between expected
normalized coverage and obtained normalized coverage for a
given sample in a region of interest. Several samples were
analyzed to corroborate similar levels of coverage between
samples, as already published by our group (Campuzano et al.,
2014b). All CNVs were compared with ExAC, including recently
added data concerning CNVs. Genetic analysis in relatives was
performed using the Sanger method. Only rare genetic variants
confirmed in the index case were analyzed in relatives to perform
a comprehensive genetic study in each family. MATERIALS AND METHODS Coordinates of sequence data were based on UCSC human
genome version hg19 (NCBI GRCh37 built). Biotinylated
cRNA probe solution was used as a capture probe (Agilent
Technologies, Santa Clara, CA, United States). Bioinformatic
processing of fastq files consisted of low-quality base-trimming
followed by a mapping step with BWA-MEM. After removal
of duplicates and low-quality reads, variant call was performed
with SAMtools v.1.2, and an ad hoc pipeline to generate raw This study was approved by the Ethics Committee of the
Hospital Josep Trueta (Girona, Spain), following the Helsinki
II declaration. Written informed consent was obtained from all
relatives included in the study. Alive individuals were clinically
evaluated at Hospital Josep Trueta (Girona, Spain), Hospital
Clinic of Barcelona (Barcelona, Spain), or Hospital Sant Joan
de Deu (Barcelona, Spain). All patients were Caucasian and
native from Spain. We analyzed six families diagnosed with cardiomyopathies
caused by 3 variants in 6 families. Each variant in two May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 2 Cardiomyopathies: Personalized Diagnosis Campuzano et al. VCF files. The final annotation steps provided information
included in public databases. Identified variations were compared
with DNA sequences from 500 healthy Spanish individuals
(individuals not related to any index case and of the same
ethnicity) as control cases in order to identify potential founder
mutations. In addition, in order to determinate the frequency
of variants identified, they were contrasted with the Human
Gene Mutation Database (HGMD4), HapMap5, 1000 Genomes
Project6, Exome Variant Server (EVS7), Genome Aggregation
Database (gnomAD8), and Exome Aggregation Consortium
(ExAC9). Non-common genetic variants [minor allele frequency
(MAF) <1%] identified in NGS analysis were confirmed by
Sanger sequencing. Exons and exon-intron boundaries of each
gene were amplified (Verities PCR, Applied Biosystems, Austin,
TX, United States). PCR products were purified (Exosap-IT,
Affymetrix Inc., USB Products, Cleveland, OH, United States),
and then were directly sequenced in both directions (Big
Dye Terminator v3.1 and 3130XL Genetic Analyzer, Applied
Biosystems) with posterior SeqScape Software v2.5 analysis
(Life Technologies, Carlsbad, CA, United States), comparing
obtained results with the hg19 reference sequence. Sequence
variants were described following rules outlined by the Human
Genome Variation Society (HGVS10), and checked in Mutalyzer11. In silico prediction of pathogenicity of genetic variations was
assessed in Functional Annotations for Non Synonymous
SNVs (FannsDB12), Mutation Taster13, Protein Variation Effect
Analyzer (PROVEAN14), and Polymorphism Phenotyping v2
(PPH215). 19https://www.ncbi.nlm.nih.gov/projects/dbvar/clingen Frontiers in Genetics | www.frontiersin.org RESULTS Individual III.4 was diagnosed with ACM -abundant fibro-fatty
infiltration in epicardium and myocardium of right ventricle
(residual myocytes <60%), moderate decrease of right ventricular
ejection fraction (45%), and mild right ventricle dilatation-, but
with a slow clinical evolution during follow-up and negligible
symptoms to date. No other relatives were diagnosed with
ACM except one family member (II.1 -74 years old-) who
showed scattered cardiac fibrosis but not a definite diagnosis
of ACM. She remained asymptomatic thus far, without any
episode of palpitations, arrhythmia, or syncope. Neither SCD
nor syncope episodes occurred in this family. The grandparents
(I.1 and I.2) died at old age (>75 years old both), both due to
oncologic diseases, and without documented episodes of syncope. Genetic analysis identified the already reported pathogenic
variant -PKP2_p.(Leu92∗)- in three cases, II.1, III.1, and III.3. In addition, a rare variant in DSG2 (c.527C > T, p.T176I –
p.Thr176Ile) was also identified in the index case (III.1), with
the amino acid threonine (polar, uncharged side chain) changed
to isoleucine (non-polar, hydrophobic side chain). This variant
is reported in global databases with a low frequency (2/249296,
MAF: 0.0008023%). However, in silico analysis predicts an
ambiguous role, and alignment between species showed no
highly conserved domain. The variant in DSG2 was present
in the index case (III.1, clinically diagnosed with ACM), her
sister (III.3 -44 years old-, not diagnosed with ACM), and her
mother (II.1, not diagnosed with ACM but showing scattered
cardiac abnormalities). One relative clinically diagnosed with
ACM (III.4) did not carry this variant. Therefore, we disregarded
DSG2_p.(Thr176Ile) as pathogenic, at least as the cause of
ACM in this family. The PKP2_p.(Leu92∗) variant was reported first as a cause
of ACM in a cohort of cases (Fressart et al., 2010). Currently,
global population databases have reported this variant with
very low frequency (2/251424, MAF: 0.0007955%), and in silico
analysis predicts this genetic alteration is deleterious. The
PKP2_p.(Arg413∗) variant was reported for the first time
in 2006 in a cohort of cases with ACM (Syrris et al.,
2006). To date, more than 10 reports describe families with
ACM due to this pathogenic variant, and in silico, analysis
predicts the variant is deleterious. Despite identification of
this variant in global databases (rs372827156), the frequency
is very low (2/121408 alleles, MAF: 0.001647%). The third
variant, MYBPC3_p.(Arg891Ala_fr∗160), is novel. Current global
population databases do not include this variant, and in silico
analyses predict a deleterious role. RESULTS Our study retrospectively analyzed six families diagnosed with
cardiomyopathy after comprehensive clinical assessment –
families A, B, C, and D included relatives clinically diagnosed
with ACM, and families E and F included relatives diagnosed
with HCM. Comprehensive NGS analysis was performed
on at least one patient clinically diagnosed with ACM or
HCM in each family. Concerning NGS data, all samples
showed <4% duplicates of amplified regions; a minimum
of 96% reads mapped uniquely and 55% reads of insert. Finally, the median coverage was 300× and call rate at
30× was 99.9%. After genetic interpretation, three pathogenic
rare variants – each present in two different families –
were identified as the cause of disease. Families A and
B carried the variant PKP2_p.(Leu92∗), families C and D
carried the variant PKP2_p.(Arg413∗), and families E and F
carried the variant MYBPC3_p.(Arg891Ala_fr∗160). Variants
PKP2_p.(Leu92∗) and PKP2_p.(Arg413∗) are previously reported
non-sense variants that are widely accepted as pathogenic. Both 4http://www.hgmd.cf.ac.uk/ac/index.php
5http://hapmap.ncbi.nlm.nih.gov
6http://www.1000genomes.org
7http://evs.gs.washington.edu/EVS
8http://gnomad.broadinstitute.org
9http://exac.broadinstitute.org
10http://www.hgvs.org
11https://mutalyzer.nl
12http://bg.upf.edu/fannsdb
13www.mutationtaster.org
14http://provean.jcvi.org/index.php
15http://genetics.bwh.harvard.edu/pph2
16http://www.uniprot.org
17https://string-db.org
18https://smart.embl.de May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 3 Cardiomyopathies: Personalized Diagnosis Campuzano et al. FIGURE 1 | Electropherogram of pathogenic variants identified in our families and closest proteins interactions. (A) Families A and B: c.275T > A (TTG > TAG),
p.L92X (p.Leu92∗) in the PKP2 gene. Families C and D: c.1237C > T (CGA > TGA) and p.R413X (p.Arg413∗) in the PKP2 gene. Families E and F: c.2670dupG,
p.R891A_frX160 (p.Arg891Ala_fs∗160) in the MYBPC3 gene. (B) Network of ten closest proteins to PKP2. (C) Network of ten closest proteins to MYBPC3. FIGURE 1 | Electropherogram of pathogenic variants identified in our families and closest proteins interactions. (A) Families A and B: c.275T > A (TTG > TAG),
p.L92X (p.Leu92∗) in the PKP2 gene. Families C and D: c.1237C > T (CGA > TGA) and p.R413X (p.Arg413∗) in the PKP2 gene. Families E and F: c.2670dupG,
p.R891A_frX160 (p.Arg891Ala_fs∗160) in the MYBPC3 gene. (B) Network of ten closest proteins to PKP2. (C) Network of ten closest proteins to MYBPC3. genetic abnormalities radically change the protein, resulting in
a final protein that is shorter than wildtype PKP2 (Figure 1
and Tables 1, 2). suffered a syncope due to ventricular arrhythmia, and an
implantable cardioverter defibrillator (ICD) was implanted. RESULTS It is a small indel that
induces a frameshift, resulting in a longer than wildtype protein. Therefore, all parameters analyzed indicate a pathogenic role
(Figure 1 and Tables 1, 2). Frontiers in Genetics | www.frontiersin.org Families A and B Families A and B were clinically diagnosed with ACM (Table 1). The cause of the disease was a pathogenic variant (CM102825) in
PKP2 (c.275T > A, p.L92X – p.Leu92∗) (Table 2). Our NGS panel
included genes encoding the functionally closest PKP2-related
proteins, but no abnormalities were identified except in Family
A. In addition, no CNVs were identified in either family. Family A included two members clinically diagnosed with
ACM, III.1 -48 years old- and III.4 -42 years old- (Figure 2). Individual III.1 had repeated episodes of palpitations. During
clinical assessment (right ventricle ejection fraction: 38%; dilated
right ventricle: 4.3 cm; no left ventricle affectation), III.1 Family
B
also
carried
the
same
non-sense
variant,
PKP2_p.(Leu92∗). Relative III.1-39 years old- was definitely Frontiers in Genetics | www.frontiersin.org May 2019 | Volume 10 | Article 450 4 Cardiomyopathies: Personalized Diagnosis Campuzano et al. TABLE 1 | Clinical data. Family
Individual
Age
Gender
Gene_Variant
Diagnosis
Criteria
A
II.1
74
F
PKP2_p.(Leu92∗)
No
–
III.1
48
F
PKP2_p.(Leu92∗)
ACM
Definite (2 major and 2 minor)
III.4
42
F
PKP2_p.(Leu92∗)
ACM
Definite (1 major and 2 minor)
B
II.1
68
F
PKP2_p.(Leu92∗)
ACM
(1 major and 1 minor)
III.1
39
F
PKP2_p.(Leu92∗)
ACM
Definite (1 major and 2 minor)
C
I.1†
44
M
? Families C and D diagnosed
with
ACM
(Figure
3). She
suffered
syncope
during
exercise. Clinical
assessment
showed
dyskinesia
with a moderate dilation of the right ventricle (39 mm), a
minor reduction of ejection fraction (40%), and no criteria
of fibro-adipose infiltration in the myocardium (residual
myocytes nearly 75% in right ventricle). The left ventricle
was not affected. Given the risk of lethal arrhythmias, this
individual had an ICD implanted 6 years ago, with three
appropriate shocks registered to date. Individual II.1 -68 years
old-, despite old age, showed no symptoms during most of
her life (punctual palpitations in the past 8 years during
moderate
physical
activity). One
relative
(grandfather,
I.2) died suddenly at a young age -36 years old- while
working on a farm. However, neither complete autopsy nor
molecular autopsy was performed. No other rare variants were
identified in this family. diagnosed
with
ACM
(Figure
3). She
suffered
syncope
during
exercise. Clinical
assessment
showed
dyskinesia
with a moderate dilation of the right ventricle (39 mm), a
minor reduction of ejection fraction (40%), and no criteria
of fibro-adipose infiltration in the myocardium (residual
myocytes nearly 75% in right ventricle). The left ventricle
was not affected. Given the risk of lethal arrhythmias, this
individual had an ICD implanted 6 years ago, with three
appropriate shocks registered to date. Individual II.1 -68 years
old-, despite old age, showed no symptoms during most of
her life (punctual palpitations in the past 8 years during
moderate
physical
activity). One
relative
(grandfather,
I.2) died suddenly at a young age -36 years old- while
working on a farm. However, neither complete autopsy nor
molecular autopsy was performed. No other rare variants were
identified in this family. Families C and D
Families C and D were also diagnosed with ACM (Table 1), and
both carried a previous reported pathogenic variant (CM060431)
in PKP2 (c.1237C > T, p.R413X – p.Arg413∗), associated with
ACM (Table 2). Our NGS panel included genes encoding
the closest PKP2-related proteins, but no abnormalities were
identified in these genes. In addition, no CNVs were identified
in either family. Family C included six family members diagnosed with
ACM, four of which were alive (II.2 -42 years old-, II.4 -
40 years old-, II.6 -37 years old-, and III.2 -9 years old-) and
carried the same genetic variant, PKP2_p.(Arg413∗) (Figure 4). Families A and B ACM
Possible (2 minor)
II.2
42
F
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 2 minor)
II.3†
21
M
PKP2_p.(Arg413∗)
ACM
Definite (1 major and 2 minor)
II.4
40
M
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 1 minor)
II.6
37
M
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 2 minor)
III.2
9
M
PKP2_p.(Arg413∗)
ACM
Definite (1 major and 2 minor)
D
II.1
74
F
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 1 minor)
II.5
71
M
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 1 minor)
II.7
68
M
PKP2_p.(Arg413∗)
ACM
Definite (1 major and 2 minor)
III.2
39
F
PKP2_p.(Arg413∗)
ACM
Definite (1 major and 2 minor)
III.4
35
F
PKP2_p.(Arg413∗)
No
–
III.6
30
M
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 1 minor)
III.8
38
M
PKP2_p.(Arg413∗)
ACM
Definite (2 major and 1 minor)
IV.1†
10
M
PKP2_p.(Arg413∗)
ACM
(2 minor)
IV.2
11
M
PKP2_p.(Arg413∗)
No
–
IV.5
6
F
PKP2_p.(Arg413∗)
No
–
E
II.2
68
F
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 19 mm
II.3
66
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 20 mm
III.2
39
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 17 mm
III.4
34
F
MYBPC3_p.(Arg891Alafs∗160)
No
–
III.6†
38
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
Myocardial disarray
III.8
36
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 24 mm
III.11
30
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 16 mm
IV.2
11
M
MYBPC3_p.(Arg891Alafs∗160)
No
–
IV.4
8
F
MYBPC3_p.(Arg891Alafs∗160)
No
–
IV.6
3
F
MYBPC3_p.(Arg891Alafs∗160)
No
–
IV.8
2
F
MYBPC3_p.(Arg891Alafs∗160)
No
–
F
II.2
73
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 25mm
III.1
46
F
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 23mm
III.2
41
M
MYBPC3_p.(Arg891Alafs∗160)
HCM
LV wall thickness 22mm
ACM, arrhythmogenic cardiomyopathy; F, female; HCM, hypertrophic cardiomyopathy; LV, left ventricle; M, male. Sign †means sudden death. ACM, arrhythmogenic cardiomyopathy; F, female; HCM, hypertrophic cardiomyopathy; LV, left ventricle; M, male. Sign †means sudden death. genic cardiomyopathy; F, female; HCM, hypertrophic cardiomyopathy; LV, left ventricle; M, male. Sign †means sudden death. Families C and D Families C and D Concerning diagnosis, inverted T-wave in right precordial leads
were observed in three cases (II.2, II.4, and II.6); in addition,
sustained ventricular tachycardia were observed in II.4, and II.6. Image analysis in II.2 showed regional RV akinesia and 38% RV May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 5 Cardiomyopathies: Personalized Diagnosis Campuzano et al. ejection fraction. In III.2, regional RV akinesia and PLAX RV
outflow tract of 31 mm were identified. The other two relatives
died suddenly at a young age and both were diagnosed with ACM
post-mortem (both showing residual myocytes <60% in fibro-
fatty infiltration); I.1 died working in a building site -44 years
old-, and II.3 died at young age during exercise -21 years old-
. Molecular autopsy indicated individual II.3 carried the same
genetic variant. Unfortunately, no sample was obtained from
I.1 to perform genetic analysis. No other rare variants were
identified in this family. Family
Gene
Nucleotide
Protein
dbSNP
EVS
ExAC
gnomAD
PP2
PROVEAN
MuTa
FansDB
HGMD (disease)
ClinVar (pathology)
ACMG
score
A
PKP2
c.275T > A
p.(Leu92∗)
rs763639737
NA
NA
2/251424
NA
NA
NA
D
CM102825 (ACM)
P (ACM)
P
DSG2
c.527C > T
p.(Thr176Ile)
rs536617217
NA
2/120200
2/28069
B
N
DC
N
NA
VUS
VUS
B
PKP2
c.275T > A
p.(Leu92∗)
rs763639737
NA
NA
2/251424
NA
NA
NA
D
CM102825 (ACM)
P (ACM)
P
C
PKP2
c.1237C > T
p.(Arg413∗)
rs372827156
1/13005
2/121408
4/282766
NA
NA
NA
D
CM060431 (ACM)
P (ACM)
P
D
PKP2
c.1237C > T
p.(Arg413∗)
rs372827156
1/13005
2/121408
4/282766
NA
NA
NA
D
CM060431 (ACM)
P (ACM)
P
MYH7
c.4772T > A
p.(Leu1591Gln)
rs730880808
NA
7/121931
10/251476
B
N
DC
N
CM1413450 (SCD)
VUS
VUS
TTN
c.60754G > C
p.(Ala20252Pro)
rs72646880
27/12015
236/120342
513/279434
PD
D
DC
N
NA
VUS
LB
E
MYBPC3
c.2670dupG
p.(Arg891Alafs∗160)
Novel
NA
NA
NA
NA
NA
DC
D
NA
NA
P
F
MYBPC3
c.2670dupG
p.(Arg891Alafs∗160)
Novel
NA
NA
NA
NA
NA
DC
D
NA
NA
P
ACM, arrhythmogenic cardiomyopathy; ACMG, American College of Medical Genetics and Genomics; B, benign; D, deleterious; DC, disease causing; EVS, exome variant server; ExAC, exome aggregation
consortium; HGMD, human gene mutation database; MuTa, mutation taster; NA, not available; N, neutral; P, pathogenic; PD, possibly damaging; PP2, PolyPhen2; SCD, sudden cardiac death; VUS, variant
uncertain/unknown significance. Families C and D netics |
f ontie sin o g y
Family D included six alive family members diagnosed with
ACM (II.1 -74 years old-, II.5 -71 years old-, II.7 -68 years old-,
III.2 -39 years old-, III.6 -30 years old-, and III.8 -38 years old-)
(Figure 5). Inverted T-wave in right precordial leads were
observed in five relatives (II.1, II.5, III.2, III.6, and III.8). III.7
also showed inverted T-wave in right precordial leads but in the
presence of complete right bundle-branch block. Non-sustained
ventricular tachycardia of left bundle-branch morphology were
observed in III.6, and III.8. Image analysis showed RV ejection
fraction of 38% and 39% in II.1 and II.5, respectively. All
other relatives (II.7, II.2, III.6, and III.8) showed RV ejection
fraction between 40 and 45%. Two relatives (III.3 and IV.1)
died at a young age (15 and 10 years old, respectively). The
index case, IV.1, died while sleeping and showed minor post-
mortem cardiac abnormalities (including myocarditis) with no
definite diagnosis of ACM but possible. Neither autopsy nor
molecular autopsy was performed in III.3 despite SCD at a
young age during exercise. No syncope or cardiac pathology
was diagnosed in either young relative who died suddenly. Molecular autopsy of the index case (IV.1) identified three rare
genetic variants –PKP2_p.(Arg413∗), MYH7_p.(Leu1591Gln)
and TTN_p.(Ala20252Pro)-. All clinically affected relatives (II.1,
II.5, II.7, III.2, III.6, and III.8) carried the PKP2_p.(Arg413∗)
variant. Three additional relatives (III.4 -35 years old-, IV.2 -
11 years old-, and IV.5 -6 years old-) carried the same genetic
variant, but all remain asymptomatic so far. Individual III.4
showed some cardiac abnormalities, although they were not
diagnostic of ACM following current guidelines (Marcus et al.,
2010). Neither grandparent (I.1 nor I.2) died suddenly or at
a young age (both more than 75 years old). Concerning the
other two rare variants identified -MYH7_p.(Leu1591Gln) and
TTN_p.(Ala20252Pro)-, previous studies published by our group
disregard a pathogenic role (Campuzano et al., 2015b). Frontiers in Genetics | www.frontiersin.org Families E and F Question mark indicates no
genetic analysis available. The ECG corresponds to index case. The ECG showing negative T waves in V1-3, and prolonged Terminal Activation Duration as major
and respectively, minor diagnostic Task Force Criteria for ACM. FIGURE 2 | Pedigree of family (A). Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are show in gray,
clinically unaffected patients are show in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.1). The genetic carriers of
PKP2_p.(Leu92∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. The ECG corresponds to index case. The ECG showing negative T waves in V1-3, and prolonged Terminal Activation Duration as major
and respectively, minor diagnostic Task Force Criteria for ACM. FIGURE 3 | Pedigree of family B. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.1). The genetic carriers of
PKP2_p.(Leu92∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. ECG showing negative T waves in V1-2 (minor ACM criterion). FIGURE 3 | Pedigree of family B. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.1). The genetic carriers of
PKP2_p.(Leu92∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. ECG showing negative T waves in V1-2 (minor ACM criterion). III.8 -36 years old-, and III.11 -30 years old-) diagnosed
with HCM (Figure 6). Only III.8 (left ventricular wall
thickness 24 mm; ejection fraction 41%), had syncope and
an ICD was implanted. Families E and F Families E and F were diagnosed with HCM (Table 1). The
disease was caused by a novel variant in MYBPC3. This variant
is a duplication of G (c.2670dupG), inducing a frameshift
(p.Arg891Ala_fr∗160) (Table 2). This genetic alteration induces
a radical change in fibronectin type 3 (FN3) domains, changing
the amino acid sequence of wildtype MYBPC3 starting at amino
acid 891. Our NGS panel included genes encoding the closest
MYBPC3-related proteins, but no abnormalities were identified. In addition, neither other rare variants nor CNVs were identified
in either family. Family E included five living family members (II.2 -
68 years old-, II.3 -66 years old-, III.2 -39 years old-, May 2019 | Volume 10 | Article 450 6 Cardiomyopathies: Personalized Diagnosis Campuzano et al. FIGURE 2 | Pedigree of family (A). Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are show in gray,
clinically unaffected patients are show in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.1). The genetic carriers of
PKP2_p.(Leu92∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. The ECG corresponds to index case. The ECG showing negative T waves in V1-3, and prolonged Terminal Activation Duration as major
and respectively, minor diagnostic Task Force Criteria for ACM. FIGURE 3 | Pedigree of family B. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.1). The genetic carriers of
PKP2_p.(Leu92∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. ECG showing negative T waves in V1-2 (minor ACM criterion). FIGURE 2 | Pedigree of family (A). Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are show in gray,
clinically unaffected patients are show in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.1). The genetic carriers of
PKP2_p.(Leu92∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Frontiers in Genetics | www.frontiersin.org Families E and F FIGURE 4 | Pedigree of family C Generations are indicated in the left side Each individual is identified with a number Clinically affected patients are shown in gray FIGURE 4 | Pedigree of family C. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (II.2). The genetic carriers of
PKP2_p.(Arg413∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. Families A and B were diagnosed with ACM due to a
pathogenic variant, PKP2_p.(Leu92∗) (CM102825). Our group
previously published the index case of Family A (Alcalde
et al., 2014), and here we have included family segregation. Hence, Family A showed incomplete penetrance and variable
expressivity
because
one
relative
carried
the
pathogenic
variant but had no diagnosis of ACM (despite showing
some
scattered
fibrotic
heart
abnormalities). Intriguingly,
the relative without an ACM diagnosis is the progenitor of
both clinically affected patients, may be due to age-dependent
penetrance age. The phenotype of both diagnosed patients
differed despite their similar age—neither had early disease
onset, but the evolution was faster and more severe in
one patient in contrast to the other. Further, the variant
in DSG2 (p.T176I) was not present in a relative clinically
diagnosed
with
ACM
(III.4). Therefore,
we
disregarded
DSG2_p.(Thr176Ile) as pathogenic, at least in Family A,
remaining the variant PKP2_p.(Leu92∗) is the most plausible
cause of disease in Family A. syncope. The three living relatives carried the same genetic
variant, MYBPC3_p.(Arg891Ala_fr∗160). Neither grandparent
(I.1 nor I.2) died suddenly or at a young age (both >75 years old). Families E and F All other relatives were diagnosed
with HCM: II.2 -left ventricular wall thickness 19 mm;
ejection fraction 49%-, II.3 -left ventricular wall thickness
20 mm; ejection fraction 45%-, III.2 -left ventricular wall
thickness 17 mm; ejection fraction 49%-, and III.11 -left
ventricular wall thickness 16 mm; ejection fraction 51%-. They did not show severe heart abnormalities (follow-up
showed slow evolution of disease), and no syncope has
been documented thus far. All carried the same rare genetic
variant, MYBPC3_p.(Arg891Ala_fr∗160). Individual III.6 was
diagnosed post-mortem –died at 38 years old-, during an
extreme effort at work. Molecular autopsy identified the same
variant (p.Arg891Ala_fr∗160) and forensic report described
an asymmetric septal hypertrophy and microscopic analysis
identified myocardial disarray with interstitial fibrosis. In addition, five relatives (III.4 -34 years old-, IV.2 -11 ears old-,
IV.4 -8 years old-, IV.6 -3 years old-, and IV.8 -2 years old-)
carried the same genetic variant, but all remained asymptomatic
and without any cardiac alteration, so far. Both grandparents (I.1
and I.2) died at old age (both >75 years old) due to oncologic
and ischemic episodes, respectively. Neither had documented
episodes of arrhythmia or syncope. Family F included three living family members (III.1, III.2,
and II.2) diagnosed with HCM (Figure 7). Individual II.2 -
73 years old- had a syncopal episode several years before,
and an ICD was implanted. He showed a left ventricular wall
thickness of 25 mm, mitral valve regurgitation, and slightly
decreased ejection fraction (48%). The other two relatives
(III.1 and III.2), despite young ages (46 and 41 years old,
respectively), showed enlarged left ventricular wall thickness
(23 mm and 22 mm, respectively), moderately decreased
ejection fraction (45 and 47%, respectively), and mitral valve
regurgitation. Individual III.1 had an ICD implanted due to May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 7 Cardiomyopathies: Personalized Diagnosis Campuzano et al. FIGURE 4 | Pedigree of family C. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (II.2). The genetic carriers of
PKP2_p.(Arg413∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. Frontiers in Genetics | www.frontiersin.org DISCUSSION The “five P” characteristics in current medicine – personalized,
predictive, preventive, participative, and precision – are an
innovative approach to the patient using available technologies to
interpret individually all patient data. Recently updated guidance,
first published in 2009 for the Heart Failure Society of America
and ACMG, supports the use of NGS in genetic testing for
cardiomyopathy to assist patient care and management of at-
risk family members (Hershberger et al., 2018). In our study, we
carried out an individualized interpretation of three pathogenic
variants leading to cardiomyopathy in six families showing high
expression variability even for the same variant and family due to
different onset, evolution, and/or outcome of the disease. May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 8 Cardiomyopathies: Personalized Diagnosis Campuzano et al. FIGURE 5 | Pedigree of family D. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (IV.1). The genetic carriers of
PKP2_p.(Arg413∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to IV.1. FIGURE 5 | Pedigree of family D. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (IV.1). The genetic carriers of
PKP2_p.(Arg413∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to IV.1. FIGURE 5 | Pedigree of family D. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (IV.1). The genetic carriers of
PKP2_p.(Arg413∗) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark indicates no
genetic analysis available. SD means Sudden Death. The ECG corresponds to IV.1. Frontiers in Genetics | www.frontiersin.org DISCUSSION Family
B,
despite
only
two
genetic
carriers
showed
incomplete
penetrance
and
variable
expressivity. The
younger relative (III.1) showed an aggressive phenotype,
with
often-malignant
arrhythmias,
and
had
an
ICD
implanted. The
older
relative
(II.1)
showed
a
slow
evolution
and
benevolent
phenotype
due
to
minimal
symptoms throughout her life. Therefore, considering the
clinical data in both Families A and B, PKP2_p.(Leu92∗)
may
be
considered
the
cause
of
ACM,
although
other
genetic
abnormalities
may
modify
onset,
evolution,
and
outcome of the disease. affected relatives carried the non-sense variant in PKP2, and
young relatives showed cardiac abnormalities associated with
ACM, suggesting an early onset of disease due to this genetic
variant. In 2015, our group also published part of Family D
(Campuzano et al., 2015b), and here we have included more
relatives to confirm the pathogenic role of PKP2_p.(Arg413∗)
as cause of disease. Genotype-phenotype correlation suggests
that PKP2_p.(Arg413∗) is the cause of disease in these families,
whereas additional variants could play a role as phenotype
modulators. We observed incomplete penetrance and variable
expressivity, as occurs in other reported families affected by
ACM (Kannankeril et al., 2006). Asymptomatic patients carrying
this genetic variant were young relatives. However, it is well
accepted that ACM is a progressive structural disease, so
cardiac abnormalities may not yet be evident in early ages. Families C and D were also diagnosed with ACM and
both carried a pathogenic variant (CM060431) in PKP2
(c.1237C > A, p.R413X - p.Arg413∗). Our group published
on Family C in 2014 (Alcalde et al., 2014). All clinically May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 9 Cardiomyopathies: Personalized Diagnosis Campuzano et al. FIGURE 6 | Pedigree of family E. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (III.2). The genetic carriers of
MYBPC3_p.(Arg891Alafs∗160) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark
indicates no genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. FIGURE 6 | Pedigree of family E. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. DISCUSSION Index case is indicated with an arrow (III.2). The genetic carriers of
MYBPC3_p.(Arg891Alafs∗160) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark
indicates no genetic analysis available. SD means Sudden Death. The ECG corresponds to index case. Families E and F were diagnosed with HCM due to a
novel frameshift alteration in MYBPC3 that results in a
different protein than wildtype MYBPC3. All analyses suggest
a pathogenic role. Family E showed incomplete penetrance
and variable expressivity, with clinically diagnosed relatives
carrying the MYBPC3_p.(Arg891Alafs∗160) variant and showing
different phenotypic evolution of heart abnormalities. This
agrees with several reports showing different phenotypes from
one family diagnosed with HCM due to a single pathogenic
variant (Wang et al., 2018). In Family E, asymptomatic
patients were young relatives, suggesting a delay in the
onset of HCM that makes cardiac manifestations not yet
evident at younger ages. This agrees with previous reports
suggesting
that
pathogenic
variants
in
genes
associated Indeed, a middle-aged relative in Family D showed some
cardiac abnormalities that were not diagnostic of ACM following
current guidelines (Marcus et al., 2010). Therefore, the same
non-sense variant induces pathology in both Families C and
D, but the onset in each family is different, especially in
young individuals (<40 years old). In Family C, relatives
showed evident heart abnormalities at an early age, suggesting
a highly aggressive role of the variant; however, in Family D,
the abnormalities seemed to occur progressively, suggesting a
pathogenic role of the variant but not with the same temporal
onset. Therefore, despite PKP2_p.(Arg413∗) being the cause
of ACM, other genetic abnormalities (although not in any of
the closest related proteins) may modify the onset, evolution,
and outcome of ACM. Indeed, a middle-aged relative in Family D showed some
cardiac abnormalities that were not diagnostic of ACM following
current guidelines (Marcus et al., 2010). Therefore, the same
non-sense variant induces pathology in both Families C and
D, but the onset in each family is different, especially in
young individuals (<40 years old). In Family C, relatives
showed evident heart abnormalities at an early age, suggesting
a highly aggressive role of the variant; however, in Family D,
the abnormalities seemed to occur progressively, suggesting a
pathogenic role of the variant but not with the same temporal
onset. DISCUSSION Therefore, despite PKP2_p.(Arg413∗) being the cause
of ACM, other genetic abnormalities (although not in any of
the closest related proteins) may modify the onset, evolution,
and outcome of ACM. May 2019 | Volume 10 | Article 450 Frontiers in Genetics | www.frontiersin.org 10 Cardiomyopathies: Personalized Diagnosis Campuzano et al. FIGURE 7 | Pedigree of family F. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (II.2). The genetic carriers of
MYBPC3_p.(Arg891Alafs∗160) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark
indicates no genetic analysis available. The ECG corresponds to index case. FIGURE 7 | Pedigree of family F. Generations are indicated in the left side. Each individual is identified with a number. Clinically affected patients are shown in gray,
clinically unaffected patients are shown in white, and slashes indicate a deceased relative. Index case is indicated with an arrow (II.2). The genetic carriers of
MYBPC3_p.(Arg891Alafs∗160) are represented with a plus sign inside rounds/squares. Minus sign indicates non-carriers of the genetic variant. Question mark
indicates no genetic analysis available. The ECG corresponds to index case. with
HCM
do
not
induce
structural
abnormalities
at
early years (Brion et al., 2009, 2012). Family F showed
complete penetrance and variable expressivity. All relatives
who
carried
the
MYBPC3_p.(Arg891Alafs∗160)
variant
had a clinical diagnosis of HCM, including young family
members. Therefore,
MYBPC3_p.(Arg891Alafs∗160)
is
the
likely cause of HCM in both Families E and F, and other
genetic abnormalities (although not in the closest related
proteins) may modify onset, evolution, and outcome of the
disease in each family. with
HCM
do
not
induce
structural
abnormalities
at
early years (Brion et al., 2009, 2012). Family F showed
complete penetrance and variable expressivity. All relatives
who
carried
the
MYBPC3_p.(Arg891Alafs∗160)
variant
had a clinical diagnosis of HCM, including young family
members. Therefore,
MYBPC3_p.(Arg891Alafs∗160)
is
the
likely cause of HCM in both Families E and F, and other
genetic abnormalities (although not in the closest related
proteins) may modify onset, evolution, and outcome of the
disease in each family. to high risk of lethal arrhythmias, accordingly to recent
published manuscripts (Kapplinger and Ackerman, 2016). DISCUSSION In addition, proper genetic counseling is essential for all
family
members
despite
the
high
complexity
of
genetic
heterogeneity, incomplete penetrance, and variable expressivity
related to inherited cardiomyopathies (De Bortoli et al.,
2017). In this prospective, further genetic studies aiming to
identify modifying factors are needed to improve prevention
of malignant arrhythmias and guide appropriate treatment
in each patient. Autopsy in young individuals who die suddenly may be
not conclusive and may not identify cardiac abnormalities. However, potential cardiomyopathy should not be disregarded,
as ultramicroscopic changes that are not evident at autopsy may
nonetheless induce arrhythmia responsible for SCD (Campuzano
et al., 2014a). Molecular autopsy may identify the genetic
alteration in a gene associated with cardiomyopathy but
without anatomic heart abnormalities. This fact is important
in sudden deaths occurring in infants, children, and even
young people (Sarquella-Brugada et al., 2016). A recent
study states that, carriers of a pathogenic variant may be
at 2.5-fold greater risk of SCD compared to non-carriers
(Milano et al., 2016). Nonetheless, it should be communicated
to the patient that despite being a carrier, the individual
may never express the disease phenotype (Kapplinger and
Ackerman, 2016). Similar to our study, a 2010 publication
of a cohort of 60-year-old patients diagnosed with ACM and
carrying a pathogenic radical variant indicates that only 60%
of genetic carriers had experienced any symptoms, without
differences between genders (van der Zwaag et al., 2010). It is well-known that cardiomyopathies may display a broad
range of phenotype variability, from asymptomatic individuals
to severe heart alterations; it is also important to remark
that sometimes the first manifestation of disease is SCD,
which usually occurs in a healthy young individual while
doing exercise, main trigger of life-threatening ventricular
arrhythmias. Considering all data, we believe that clinical
follow-up including periodic tests of risk should be performed
in all genetic carriers, despite remaining asymptomatic due In
conclusion,
genetic
testing
has
been
progressively
incorporated into clinical diagnosis, mainly due to improved
NGS technology. Although genetic interpretation may classify
rare variants as pathogenic, translation into clinical practice
sometimes shows both interfamilial and intrafamilial phenotypic
differences. We report here families carrying the same pathogenic
variant associated with cardiomyopathies but with differential
onset and/or clinical progression. Therefore, additional factors
may
modify
the
disease
phenotype
in
each
individual. Nonetheless, all individuals carrying a pathogenic variant,
despite being asymptomatic, are at risk of disease therefore,
clinical follow-up should be performed. DISCUSSION Clinical translation
remains a main challenge for cardiovascular genetic units and
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Deo, R., and Albert, C. M. (2012). Epidemiology and genetics of sudden
cardiac death. Circulation 125, 620–637. doi: 10.1161/CIRCULATIONAHA. FUNDING This work was supported by Obra Social “La Caixa Foundation”
(ID 100010434), Fondo Investigacion Sanitaria (FIS, PI14/01773
and PI17/01690) from Instituto Salud Carlos III (ISCIII),
and “Fundacio Privada Daniel Bravo Andreu.” CIBERCV is
an initiative of the ISCIII, Spanish Ministry of Economy
and Competitiveness. Limitations A main limitation of this study is the lack of understanding
of the reasons implicated in high expression variability even
for the same variant and family. Additional analysis of the
assessed genes as well as in other genes not included in our NGS
custom-panel is necessary to identify additional genetic variants
that could modify the phenotype. In addition, other genetic
analyses focused on transcriptional/proteomic modifications are
necessary to clarify the role of each genetic variant in each
individual. It is important to remark the absence of data on
exercise, an important contributor to phenotypic manifestation. Finally, in vivo and/or in vitro studies of each genetic variant
may help clarify pathophysiological mechanisms associated
with the pathology. May 2019 | Volume 10 | Article 450 11 Cardiomyopathies: Personalized Diagnosis Campuzano et al. ETHICS STATEMENT MP, LL, FP, IM-S, and CT acquired, pre-processed, and
analyzed
the
data. OC,
AF-F,
and
GS-B
prepared
the
manuscript. OC, JB, and RB supervised the study. All authors
contributed to manuscript revision, read, and approved the
submitted version. This study was approved by the Ethics Committee of the
Hospital Josep Trueta (Girona, Spain), following the Helsinki
II declaration. Written informed consent was obtained from all
relatives included in the study. Alive individuals were clinically
evaluated at Hospital Josep Trueta (Girona, Spain), Hospital
Clinic of Barcelona (Barcelona, Spain), or Hospital Sant Joan de
Deu (Barcelona, Spain). This study was approved by the Ethics Committee of the
Hospital Josep Trueta (Girona, Spain), following the Helsinki
II declaration. Written informed consent was obtained from all
relatives included in the study. Alive individuals were clinically
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Clinic of Barcelona (Barcelona, Spain), or Hospital Sant Joan de
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Crafting, Crafting, Crafting - Extreme Programming in Classroom?
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149 149 Informatics in Education, 2008, Vol. 7, No. 2, 149–158
© 2008 Institute of Mathematics and Informatics, Vilnius © 2008 Institute of Mathematics and Informatics, Vilnius Received: April 2008 Abstract. Extreme Programming (XP) shows several interesting approaches which are very attrac-
tive for education. It is centered around early and incremental creation of working software. In the
following, the chances XP offers for class are shown – especially for use in a class project, but also
for practical phases in all lessons where programming is useful. Finally several common problems
which can occur with XP will be shown as well as how to deal with them to make the use in class
as smooth as possible. Keywords: extreme programming, XP, education, pair programming, project class, independent
work. Crafting, Crafting, Crafting – Extreme
Programming in Classroom? Marcus BITZL
Technical University of Munich, Department of Informatics, Didactics in Informatics
Boltzmannstr. 3, D-85748 Garching
e-mail: marcus@bitzl.com Marcus BITZL
Technical University of Munich, Department of Informatics, Didactics in Informatics
Boltzmannstr. 3, D-85748 Garching
e-mail: marcus@bitzl.com Marcus BITZL
Technical University of Munich, Department of Informatics, Didactics in Informatics
Boltzmannstr. 3, D-85748 Garching
e-mail: marcus@bitzl.com 1Here we can already see an important point of Extreme Programming: instead of dictating a schedule
and forcing its adherence at any price, one rather aims for the best effort estimation with inclusion of the
programmers. 1. Introduction Extreme Programming (XP) is an agile software development methodology founded by
Kent Beck. On a closer examination it shows several very interesting approaches for use
in classroom which will be shown in the following. Because of XP’s educational values,
Brüggemann et al. chose it for their projects in professional education in IT companies
(Brüggemann et al., 2006). But scientific papers concerning the use in schools are very
rare. Thus, XP will be explained below and interesting aspects for the use in school will
be especially emphasized. In addition, different applications and solutions for important
problems concerning its use in class will be shown. But first: what does “use in school” mean? The term school is cleared at once: where
this paper relates to a special type of school, the Bavarian Gymnasium is meant because
of its large amount of informatics lessons, but everything in this paper can be easily
generalized to other school types. When it comes up to “use”, there are mainly three
possibilities: first, one could apply XP in practical phases in a lesson; second, one could
use XP in a class project, and finally, one could make an own elective creating software
with XP (e.g., an “XP-Team”). The main source for this paper is “Extreme Programming Explained. Embrace
Change” from Kent Beck (Beck, 2000), in which he presents XP. All statements on how
XP works are from this book. 150 M. Bitzl 2. Roles For XP, Beck defines several roles (Beck, 2000) which are important for its understand-
ing: the central role is the one of the programmer and is similar to the one of other
programmers, but with slightly shifted priorities. The programmers’ main objective is
communication between each other and with all other roles as shown later. The customer
is in XP also part of the team – no matter if it is an external customer or the marketing
section of the same company. It is crucial that the customers are real users which would
really work with the software that is built. Beside these, there are more roles: the tester helps the customer to create function
tests and is responsible for the regular execution of these test, if it is not already au-
tomated (unit- and integration-tests are part of the development and therefore tasks of
the programmers). The time manager is responsible for the project’s schedule. The time
manager is kind of the team’s conscience as he (or she) is responsible for adhering the
schedule or for its correction if adhering is impossible. He also watches the team’s per-
formance to help them making better effort estimations1. The time manager should stay
in the background for not to disturb work more than necessary. The Coach is responsible for the overall process. His job is to direct people in a
certain direction if necessary. His interferences should be as few as possible. Moreover,
it is recommended to prefer an indirect guidance over a direct one, so people (mainly
programmers) could find a better way by themselves. As XP programmers are rather rarely specialists in their customers’ area of application
such specialists are taken for a certain time as advisors. The Big-Boss is responsible for the team’s motivation, but also for all organizational
concerns like staff or scheduling. The team takes its problems to him and expects his help
to solve them. If everything works fine, the Big-Boss should stay in the background – his
intervention would perturb the team’s productivity. What do these roles mean for use in class? The students get in any case the program-
mers’s role, some of the could also be customers instead. The teacher should take the
other roles. In a class project or a practice class it would be possible for some roles (e.g.,
the advisor) to invite other persons for some time. 3. Values and Principles 3.1. Values 3.1. Values 3.1. Values According to (Beck, 2000), Extreme Programming is based on four values which can be
found in all aspects: open and honest communication has to take place throughout the
whole project, between the pair-programming partners, between all developers, the man-
agement and also with the customers. This is necessary to produce high quality software. 151 Crafting, Crafting, Crafting – Extreme Programming in Classroom? All solutions have to follow the principle of simplicity: an implemented solution may
be only as complex as necessary to meet the requirements. If an existing solution is too
limited a refactoring, that is a change of the code structure, is applied. To ensure high-quality solutions, some feedback is needed: from the customer to meet
his requirements, from the other developers to find the best solutions and to learn. And
also from the software itself feedback is demanded by tests to ensure that all requirements
are fulfilled and to avoid defects. The fourth value is courage: it takes courage to address
problems and conflicts frankly and openly. It also takes courage to replace already work-
ing parts of the software with new ones when it is necessary. XP always takes courage
for change and for experiments, if an appropriate solution is not clear enough, and at last
courage to take the responsibility for the different tasks. If one considers these values together with the constitution of the free state of Bavaria,
or the profile of the Bavarian Gymnasium, one can find several interesting parallels: the
constitution already emphasizes the importance of a sense of responsibility and the dispo-
sition to accept responsibility as an educational objective. This is taken up in the profile
of the Bavarian Gymnasium and supplemented with skills like the ability to communi-
cate, the ability to work in a team, conflict handling skills and the ability to make sound
judgements. 3.2. Basic Principles To use the values above practically, Beck derives five basic principles: mainly based on
behavioristic learning theories, but also to recognize problems as early as possible, rapid
feedback is demanded. Technically, one gets this with automated tests. For people, stand
up meetings and personal discussions are used. Second, the developers should assume simplicity, that is solving any task only as
it is given at the moment – not as it will be later or even as it could be later. This is
exactly what the customer wants and pays for in the next release. Also, this is the easiest
solution to understand and to maintain. For class, this is good because of three reasons:
the students get solutions in less time, realize an immediate relation between their work
and the resulting use (there is nothing that is not used at the moment) and with a latter
extension, they can see the limitations of their present solution and learn new approaches. As the project is under constant development, the third principle is incremental
change: large and extensive changes are error prone and therefore difficult to control. If large changes are split into sequences of many small changes, every single one could
be very well controlled and realized. This is also very useful for school, as it supports
students in realizing changes and understanding their consequences. Moreover, they get
working versions even before all changes are implemented. This can only work, if everybody encourages change. This should also be visible in
the way how solutions are chosen: the best ones are those solving the problem (without
violating another principle) and offer the most options for future development. In every single aspect, high quality work is demanded: if the created software is of
poor quality, it is no fun for the developers to work on it. This has negative consequences
for the quality of their further work and therefore for the success of the whole project. 152 M. Bitzl In school it would surely be the same: software of poor quality (as one would see from
many errors and instabilities) would harm the motivation of all participants seriously. 3.3. More Principles Besides the five base principles, there are ten more principles from which some special
ones get further explained: software development is always also considered as a learning
process. Therefore, teaching to learn is a crucial task for the coach. Systematic experi-
ments are also part of the XP’s everyday life helping to understand concepts or to select
solutions (thus, learning again). Furthermore, each developer has to take responsibility, a
competence already demanded by the Bavarian constitution. Technically, small initial investments are demanded: the project is naturally quite
small when it is started (as it didn’t exist before). Therefore, starting with too much re-
sources can lead to problems. How to solve this will be shown later. The principle “playing to win” has to be understood in contrast to “playing not to
loose”: our goal has to be to develop a project in intensive communication with our
customer and delivering it in high quality. Everything else would harm the project and
therefore also the developers’ motivation and the effectiveness of software development. This is also very important for school because students expect a well working result at
the end of a project. A low quality product would be certainly demotivating, and a failed
project would be disastrous, regardless if the rules of a certain process model have always
been followed correctly. In every aspect of the development process, we need open and honest communication,
also one has to adopt XP to local conditions, use people’s instincts (not work against
them) and travel light – that means, not to use any unnecessary processes or software. When using tests, metrics or simply when speaking with developers and customers
honest measurement is important: if measurement results or information retrieved from
discussions are not “nice”, this is an important hint that something doesn’t work properly
or could be done better. 4.1. Activities According to Beck, the four main activities for XP programmers are coding, testing,
listening and designing. Coding is the central point in XP, because it quickly leads to
measurable results, one can easily quantify implemented features and clean code is suit-
able for communication about solutions. The implementation of an idea, an algorithm or
a solution can be tested if they meet the requirements. Different solutions can be com-
pared. This enables the programmers not only to find the best solutions or to improve the
existing ones, but also to learn with every comparison, selection or solution they make. In school, the students can sample theoretical concepts with concrete problems and solu-
tions. The short times till students achieve results and the fact that they can easily quantify
their work and understand their code, support a positive feeling of success and therefore
supports independent work. 153 Crafting, Crafting, Crafting – Extreme Programming in Classroom? Permanent testing ensures that all implementations fulfil their requirements. There-
fore, one creates tests before creating the corresponding implementation. The execution
of these tests should be automated, so the programmers can execute them easily and of-
ten. In addition, Extreme Programming uses these tests as parts of the documentation,
because one can clearly see there the expected behaviors. A meta-analysis done by Jef-
fries und Melnik (Jeffries and Melnik, 2007) shows the use of this approach: out of 16
studies 11 showed a considerable improvement of software quality. Only one states a
negative influence on software quality and only four do not show any effect at all. For school, this approach is interesting for different reasons: first, the students get
a positive (or negative) feedback on their implementations immediately. Therefore, their
erroneous work is still present and an immediate feedback after they corrected the mistake
leads to an increased learning effect according to behavioristic learning theories (see
also the principle of immediate feedback). With unit tests, errors get narrowed down to
the smallest possible code regions. Feedback from tests also supports the teacher and
also encourages the students’ independent activities with concrete information on errors. Moreover, the exact way of working as demanded in the profile of informatics education
(see (Fachprofil Informatik G8, 2004)) is supported as tests only accept exact solutions
to their requirements. 4.1. Activities The third activity – listening – is based on a basic problem in software engineer-
ing: misunderstandings between the customer and the developers. Therefore, developers
have to learn how to listen and have to learn listening and how to do this actively in
a discussion, to individuals as well as to large groups. Part of this is to ask questions
and to develop a basis for common communication. Moreover, developers help the cus-
tomers to understand which things are hard and which ones are easy to achieve. In school,
this strong focus on communication supports students by improving their communication
skills as for example demanded in Bavarian curriculums. These activities alone could help us only for a short time, because if good design is
neglected, code soon gets unmaintainable and almost impossible to change. Therefore,
the last activity is design. This is not a phase, but a continuous process interrupted by
other activities or melted with programming to one unit. This is very useful for class
as implementations immediately follow on design, changing in very short cycles. This
allows students to experience the consequences of their good (or bad) design immediately
and to realize how appropriate design simplifies their solutions. Designing in XP is different from other software engineering methods: first, program-
mers implement a test case and a piece of code solving the current task in the simplest
possible way. For the next task, they extend the test case and change the existing code to
pass the tests. Doing so, they take care that the code doesn’t contain anything which is
not necessary at the moment. For the third, fourth and any further tasks they do the same,
until all tasks are done. This is also expedient for use in lessons: breaking up the whole
problem into many pieces reduces it to a set of small problems which are easy to solve,
and every one is independent from the following changes. Therefore, students can focus
on their problems without being distracted by much stuff not necessary at the moment
and whose purpose is yet hard to understand. Astrachan et. al suggest to give students (in 154 M. Bitzl his case from university, but for this technique this doesn’t really matter) a task for which
they should find a solution which is as simple as possible. 4.1. Activities The following tasks demand
a gradual extension of the existing solutions leading slowly to a certain, very elegant de-
sign. He reports a strongly improved learning success compared to assignments solved
with a design the teacher had only presented his students (Astrachan et al., 2003). 4.2. Workflow If the customer already needs some feature for the next release
which is not in the plan yet, he can write a story card for this and replace another story
card needing the same effort. If the programmers get the impression that the plan might
lose touch with reality, the plan has to be estimated again. Therefore, in XP the customer
is always a member of the team and has always to be available for developers’ questions. This is called ‘customer in place’, although physical presence is not needed all the time. For the creation of software several techniques are used: a metaphore is used for com-
munication, for an intuitive understanding of the project’s aim and also for a common
design. For an accounting software it could look for example like the following: “a vir-
tual accountant collects and manages all accounting orders, evaluates them and makes
nice charts”. The metaphor should support a simple design. In the case of our virtual ac-
countant it could be like this: one package for managing transactions (the “accountant”
himself), one for analysis and one for making charts. After some time it could prove use-
ful to move the “accounts” from our accountant to a new package of its own – refactoring
simplifies design even more. If this software would need interfaces to other accounting
solutions, the introduction of a “translator” package could solve this. To do refactoring easily without accidentally destroying other features, continuous
testing (with unit tests and continuous integration) is necessary. A very central aspect of XP is programming in pairs: at any time, two programmers
do the programming together. One of them has the active part and writes code, the other
one follows his line of thought, asks questions and tries if he (or she) can find a solution
that is simpler. If they are not sure if a solution is good, they give it a try and maybe
they also try another one until it is clear which solution is the simplest. The partner helps
to identify problems earlier, which saves time and energy. Together, pair programmers
might find solutions each of them alone wouldn’t. The partners can switch their roles
anytime. For school all these aspects are advantages, from debugging to creative problem solv-
ing and, of course, learning from each other. Werner et. al. 4.2. Workflow An XP-project’s workflow consist of two parts: during the planning game, the project
plan is created or adapted together with the customers and so the software development
is controlled. On the planning game follows the creation of the software. If requirements
change we repeat the planning game for the changed requirements (also during a cre-
ational phase) and the project plan is adapted. Afterwards, the creation can be continued
without any problems. If a new version of the software should be developed, the planning
game is done again. How is the extent of a version determined? According to XP, this can only be done
with coarse communication with the customers. This happens in the so called planning
game, a process consisting of three phases (see Fig. 1) which will be further explained
in the following: in the exploration phase the customers create story cards describing the
desired features. The developers estimate for every story card, how long it would take
to implement the features. If this is not possible, they talk to the customers which will
explain their ideas in a more detailed way or split the story card into several smaller ones. Fig. 1. Phases of the planning game. Fig. 2. A story card. Fig. 1. Phases of the planning game. Fig. 1. Phases of the planning game. Fig. 2. A story card. Fig. 2. A story card. Fig. 2. A story card. 155 Crafting, Crafting, Crafting – Extreme Programming in Classroom? In commitment phase all story cards are sorted into three stacks: first, by the customer
by value (absolutely necessary, important and nice, but less important), and second, by
the developers by risk. This is measured by how closely the necessary time can be calcu-
lated (almost certain, closely and hardly to not at all). Afterwards, the developers tell the
customer, how fast they can be in ideal development time per month. At last, the steering phase follows: the customers choose from the selected story cards
those, which should be implemented in the next iteration (usually a period from one to
three weeks). After the first iteration the system has to work, but doesn’t have to be fully
developed. If the developers realize that they have overestimated their development speed,
they can correct the schedule together with the customers and remove less important
features from the plan. M. Bitzl Common responsibility is another instrument for quality assurance and part of a XP
working culture. Every developer is responsible for the whole project. Regularly changing
the partners of the programming pairs ensures that all programmers get knowledge of
the whole project by and by. If one sees something which he (or she) could do simpler
or nicer, he (or she) has to make this change (except somebody else is working on the
same piece of software). Especially there are no “private” parts of the software which
are off-limits for other programmers. Because of this and also because of design through
refactoring, it is very common that parts of the software are completely replaced by others
within a few weeks. The tests ensure that changes can be performed without any danger
of breaking the software. For students, this is, of course, a chance to learn, but also to
take responsibility. The rule “40-hour-week” demands a project plan which allows the programmers to
fulfill it without making overtime regularly. This would reduce their manpower as well
as their motivation and therefore harm the project. For school, this is very important as
overtime is hardly possible while the schedule is very tight. As many programmers work on the same project, XP demands coding standards. These can help students to develop a clean structure for their code. 4.2. Workflow describe a pair-programming-
session with middle school girls and introduce some rules for successful pair program-
ming with students (Werner et al., 2004). A survey from McDowell and Werner (Mc-
Dowell et al., 2003) examines the effects of pair programming in programming courses
for beginners. The participants using pair programming showed better results than those
who did not. 156 M. Bitzl 5. Problems and Solutions Several circumstances at school make the use of Extreme Programming difficult: the
group size, the available amount of time, the experience the developers have, the customer
in place and the immediate assignment of all developers. The group size has an important role in XP, because the integration process is a bottle
neck. With estimated four minutes for each integration process, ten pairs (twenty pupils)
would need 40 minutes to integrate their results. Smaller groups and projects with low
interference between the developer pairs can speed this up. The elective solution has one
crucial advantage: perhaps the group size one can expect is in acceptable limits, and if
not, one could control these factors from the outset. Another critical factor is time: from the 45 minutes of a lesson, a certain amount of
time is already lost for other activities (pupils have to come to the computer room, take
their places and make pairs, there might be organizational work to do from the school,
etc.). In the remaining time the pupils have to take a story card, extract their tasks, imple-
ment their tests and solutions and finally integrate their solutions. The teacher as coach
supports them when there are problems, but otherwise stay in the background. Here it becomes apparent that the working techniques cannot be introduced only for
a class project, but have to be familiar to the students at least partly. For this, there are
different approaches: in practical phases, the complexity can be easily controlled, for the
used working techniques as well as for the tasks. In class projects, this is more difficult,
but with an appropriate control of the planning game it is possible in certain limits. An-
other solution is to split the class and give together with another teacher of this class two
double periods at the same time. This way, one could get double time and half group size. To use this method, it has to be checked how to deal with the teachers’ overtime. 157 Crafting, Crafting, Crafting – Extreme Programming in Classroom? The pupils’ lack of experience could be faced with appropriate tasks and a stepwise
introduction of the XP-techniques. Before pupils can work independent in a class project,
they should have made first experiences with pair programming and writing software tests
in practical phases in class. 5. Problems and Solutions In an elective, they could make first experiences with software
tests together with the teacher and first experiences with pair-programming directly. For
the introduction of pair programming, a version of pair programming proposed by Astra-
chan et al. (Astrachan et al., 2003) would be appropriate. In this version, the teacher works
as active and the whole class together as passive programmer in one programming pair. The class would take part in the solution with ideas, proposals, but also with questions. The customer in place could be replaced with students taking the customer’s role. Doing so, it is also possible to make pairs with an odd number of students and perhaps to
slightly lower the number of developer pairs (probably by one or two pairs). It also gives
a chance to assign certain, more appropriate tasks to some students (e.g., for motivational
purposes). An immediate assignment of all “developers” is hard to avoid in school as we always
have to deal with whole classes (or at least large parts of them if they are split), where all
students have to be present. Obviously, each of them (or rather, each pair of them) needs
an expedient task. One solution might be to give a small code basis to the students at
the beginning of the project to avoid that they would disturb each other. Part of this code
basis could be developed in practical phases of past lessons or through the teacher-class-
pair-programming described above. It would also be possible to start with a part of the
students programming, while others do some investigation work (or other not interfering
tasks). At last, projects with low coupling between its parts are better. References Astrachan, O., Duvall, R.C. and Wallingford, E. (2003). Bringing extreme programming to the classroom. In M. Marchesi, G. Succi, D. Wells and L. Williams (Eds.), Extreme Programming Perspectives. Addison
Wesley, Boston, MA, 237–250. Beck, K. (2000). Extreme Programming Explained. Embrace Change. Addison-Wesley, Reading, Mass. Brüggemann, A., Rohs, M. and Schwill, A. (2006). Extreme programming – extreme learning? Erfahrungen
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and Advanced Technology in Education, 301–305. Werner, L., Denner, J. and Bean, S. (2004). Pair programming strategies for middle school girls. In Computers
and Advanced Technology in Education, 301–305. M. Bitzl is a teacher student at the Technical University of Munich. He has studied com-
puter science, physics and educational science and is now doing his first state examination
for qualifying as a teacher at Bavarian Gymnasium. His main research interests include
the implementation of independent learning and ontologies both for e-learning and se-
mantic web. 6. Conclusion Examining the values, the principles, the basic working techniques up to the whole project
flow, it is obvious that Extreme Programming offers many interesting aspects for class. XP emphasizes learning as an important part of the principles and practices in many
points and therefore is very appropriate for use in schools. Above three basic scenarios
have been presented and will now be summarized shortly. The use in practical phases in a lesson takes the least time and gives the teacher the
strongest control. Here, the students can learn the practices together with their teacher. With design through refactoring, students can learn the advantages of certain designs,
and programming in pairs supports the students in finding good solutions. The creation
of tests supports them in recognizing their mistakes. For the use in a class project, one needs more time and the students have to know
the working techniques. The teacher’s control is much smaller and in many cases limited
to support at problems. Software tests ensure always a working version of the program. Small iterations allow a fine risk-control and a smooth change of the project plan, so one
can ensure that the students will have working software. An XP-elective offers several
other opportunities: because it has longer period than a class project and smaller group
sizes, the techniques can be learned more intensively and also can be applied much better. This allows larger projects. 158 M. Bitzl No matter in which form Extreme Programming is used – the student’s activities will
always be the most important aspect. Acknowledgments. I would like to thank Daniela Drexler and Elisabeth Rothe for
their valuable comments on a manuscript of this paper, and also Prof. Dr. Peter Hubwieser
for guiding me to research. Marcus BITZL Ekstreminis (kraštutinis) programavimas (angl u k. santrumpa XP) turi idomi u savybi u, ypaˇc
tinkam u mokymui. Šis programavimas iš esm˙es susij es su programin˙es irangos, k¯urimo darbais. Straipsnyje aptariami b¯udai, kaip taikyti ekstremini programavim a mokant klas˙eje, ypaˇc atliekant
klas˙es projektus, taˇciau tuo neapsiribojama – parodoma, kaip panaudoti ir kitose pamokose, kur tik
programavimas gal˙et u b¯uti naudingas. Aptariamos kelios svarbiausios problemos, kurios gali iškilti
naudojant ekstremini programavim a, taip pat pateikiami ši u problem u sprendimo b¯udai.
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Hepatitis B virus reactivation associated with CAR T-cell therapy
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Holistic integrative oncology
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© The Author(s) 2024. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the
original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or
other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line
to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory
regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this
licence, visit http://creativecommons.org/licenses/by/4.0/. REVIEW Open Access Abstract Patients with hematological malignancies who also have a hepatitis B virus (HBV) infection need to be aware
of the potential risk of HBV reactivation when undergoing anti-cancer treatments. Among these treatments, CAR
T-cell therapy has gained significant attention as a promising option, but it also raises concerns regarding HBV reac-
tivation. This review aims to provide an overview of published reports on HBV reactivation during CAR T-cell therapy,
along with an assessment of the effectiveness of prophylactic antiviral therapy. Additionally, we propose a systematic
approach for monitoring and managing HBV reactivation during CAR T-cell therapy to enhance the safety of this treat-
ment for patients with HBV infection. Keywords Hepatitis B virus, Reactivation, CAR T-cell therapy, Hematological malignancies Haolong Lin1,2, Zigang Dai1,2, Liang Huang1,2 and Xiaoxi Zhou1,2* Haolong Lin1,2, Zigang Dai1,2, Liang Huang1,2 and Xiaoxi Zhou1,2* Holistic Integrative Oncology Holistic Integrative Oncology Lin et al. Holistic Integrative Oncology (2024) 3:16
https://doi.org/10.1007/s44178-024-00082-z 1 Background to proactively address the potential challenge of HBV
reactivation and its associated, potentially life-threaten-
ing complications. This endeavor is grounded in the lat-
est advancements in research and clinical experience in
this domain. In recent times, a body of research has emerged indicat-
ing the potential occurrence of hepatitis B virus (HBV)
reactivation during chimeric antigen receptor (CAR)
T-cell therapy, which, fortunately, can be effectively miti-
gated through the administration of nucleotide analogs
(NAs). It is noteworthy that China is situated within
regions of high HBV endemicity, and individuals afflicted
with HBV infection are at an increased risk of developing
non-Hodgkin lymphoma (NHL) and diffuse large B-cell
lymphoma (DLBCL). 1.1 HBV infection and B‑cell malignancies Hepatitis B virus (HBV) is a global health concern, with
a pervasive presence across the world. Approximately
257 million individuals are currently affected by HBV on
a global scale, with a staggering 68% of these cases con-
centrated in the regions of Africa and the West Pacific
[2]. In China, HBV presents a significant public health
challenge, contributing to approximately one-third of
the total global burden of HBV infections. Despite wide-
spread vaccination efforts, the national prevalence of
hepatitis B surface antigen (HBsAg) has declined to 5.2%. However, HBV continues to impact a substantial popu-
lation, affecting roughly 70 million individuals annually,
with 20–30 million cases evolving into chronic infections
and tragically resulting in more than 880,000 deaths each
year [3, 4]. Studies have shown that chronic HBV infec-
tion significantly increases the risk of NHL and DLBCL
[5]. DLBCL patients with concomitant HBV infec-
tion have distinct clinical, genomic, and transcriptomic Drawing upon insights from prior clinical investiga-
tions and expert consensus, our objective is to provide an
actionable framework for healthcare professionals, ena-
bling them to judiciously administer CAR T-cell therapy
to patients grappling with hematological malignancies
concurrent with HBV infection [1]. Our ultimate aim is *Correspondence:
Xiaoxi Zhou
cello316@163.com
1 Department of Hematology, Tongji Hospital, Tongji Medical College,
Huazhong University of Science and Technology, Wuhan 430030, Hubei,
China
2 Immunotherapy Research Center for Hematologic Diseases of Hubei
Province, Wuhan 430030, Hubei, China *Correspondence:
Xiaoxi Zhou
cello316@163.com
1 Department of Hematology, Tongji Hospital, Tongji Medical College,
Huazhong University of Science and Technology, Wuhan 430030, Hubei,
China
2 Immunotherapy Research Center for Hematologic Diseases of Hubei
Province, Wuhan 430030, Hubei, China *Correspondence:
Xiaoxi Zhou
cello316@163.com
1 Department of Hematology, Tongji Hospital, Tongji Medical College,
Huazhong University of Science and Technology, Wuhan 430030, Hubei,
China
2 Immunotherapy Research Center for Hematologic Diseases of Hubei
Province, Wuhan 430030, Hubei, China *Correspondence:
Xiaoxi Zhou
cello316@163.com
1 Department of Hematology, Tongji Hospital, Tongji Medical College,
Huazhong University of Science and Technology, Wuhan 430030, Hubei,
China
2 Immunotherapy Research Center for Hematologic Diseases of Hubei
Province, Wuhan 430030, Hubei, China Lin et al. Holistic Integrative Oncology (2024) 3:16 Page 2 of 7 Lin et al. Holistic Integrative Oncology patients with chronic HBV infection are at high risk of
HBV reactivation when they receive CAR T-cell therapy
targeting B cells or plasma cells. characteristics, indicating that HBV infection plays a
contributory role in the development of B-cell malignan-
cies [6]. In China, 9%-30.2% of NHL patients and approx-
imately 25% of DLBCL patients harbor chronic HBV
infections, and about 20%-44% of DLBCL patients have
resolved HBV infections [7, 8]. Given the prevalence of
HBV infection among patients with NHL or DLBCL, it
is essential to recognize that the risk of HBV reactivation
significantly escalates following exposure to immunosup-
pressive or cytotoxic agents [9].h While in the patients who had resolved HBV infec-
tion, no cases of HBV reactivation were reported during
NAs administration, even in case of HBV DNA-positive. However, 2 independent studies each reported 1 patient
(3.4–12.5%), who were HBV DNA-negative and did not
receive prophylactic NAs, experienced asymptomatic
HBV reactivation at 2 months and 6 months after CAR
T-cell infusion, respectively [16, 21]. HBV reactivation in
these 2 patients presented as an increase in HBV DNA
copy number or a reverse seroconversion from HBsAg
negativity to positivity and was promptly controlled by
the preemptive use of NAs. These data demonstrate that
patients with resolved HBV infection might be also at
high risk of HBV reactivation after CAR T-cell infusion,
and the administration of NAs can prevent HBV reac-
tivation in these patients, regardless of the HBV DNA
status. The risk of HBV reactivation varies for each patient,
depending on their HBV serological status as well as the
specific anti-tumor treatment they receive. Because of
the persistent B-cell aplasia induced by chemoimmuno-
therapy, the incidence of HBV reactivation ranges from
21 to 60% in patients with chronic HBV infection and
varies from 2 to 25% in patients with resolved HBV infec-
tions [10, 11]. The risk of reactivation can be considered
as high risk if the incidence of HBV reactivation reaches
or exceeds 10%. Therefore, patients with either chronic or
resolved HBV infection are at high risk of HBV reactiva-
tion when receiving hematopoietic stem cell transplan-
tation or immunochemotherapy (including rituximab
and glucocorticoid chemotherapy) [12, 13]. However,
randomized controlled clinical studies have shown that
the prophylactic use of NAs can reduce the risk of HBV
reactivation in this patient cohort. Similarly, since CAR
T cells targeting B cells or plasma cells can lead to long-
lasting B-cell depletion and humoral immunodeficiency,
patients with contaminant HBV infection might also face
a high risk of HBV reactivation when undergoing CAR
T-cell therapy. Furthermore, HBV can be activated in these high-risk
patients after improper withdrawal of prophylactic NAs. Four patients, involving 2 with chronic HBV infection
and 2 with resolved HBV infection, experienced HBV
reactivation after self-discontinuation of ETV [15, 17, 19,
26]. At the time of HBV reactivation, B-cell depletion or
complete remission of disease was still sustained. One
patient with chronic HBV infection self-discontinued
the use of ETV only 1 month after CAR T-cell infusion,
leading to a tragic outcome as the patient succumbed to
fulminant hepatic failure just two months later [26]. This
mournful incident underscores the critical importance
prophylactic NAs are essential for high-risk patients. Similarly, another patient with resolved HBV infection
ceased antiviral treatment five months after the CAR
T-cell infusion, resulting in HBV reactivation just one
month later. Despite prompt intervention with entecavir
and tenofovir, the patient eventually succumbed to com-
plications of hepatitis and cerebral hemorrhage [19]. The
remaining two patients had been on ETV prophylaxis for
more than 1 year before the discontinuation of ETV. They
presented asymptomatic elevation of HBV DNA titer or
hepatitis flare. After prompt recognition of these signs
and the subsequent resumption of ETV treatments, HBV
reactivation was successfully resolved in both patients
[15, 17]. These outcomes provide compelling evidence
that prophylactic NAs therapy may serve as a protective
measure against HBV reactivation in patients undergo-
ing CAR T-cell therapy, whether they have chronic HBV
infection or resolved HBV infection. 1.2 HBV reactivation in the setting of CAR T‑cell therapy
Recently, HBV reactivation has been described in
patients with chronic HBV infection or resolved HBV
infection after anti-CD19, CD20, CD22, or B-cell matu-
ration antigen (BCMA) CAR T-cell therapy (Table 1). In
a study including 19 patients with B-cell malignancies
and concomitant chronic HBV infection, HBV reactiva-
tion and HBV-related severe hepatitis were detected in
1 (5.3%) patient at 4 months after anti-CD19 CAR T-cell
infusion, although they had received entecavir (ETV)
[14]. Holistic Integrative Oncology Page 3 of 7 Table 1 Summary of HBV reactivation cases in CAR T-cell therapy
Study
Disease
Targets of CAR T-cell
therapy
HBV characteristic
at baseline
Number
reactivat
total
Strati P 2019 [17]
R/R-DLBCL
CD19
Resolved HBV infec-
tion
1/3
Yang C 2020 [15]
R/R-DLBCL
CD19
Chronic HBV infection
3/15
Lai P 2020 [18]
R/R B-ALL
CD19
Chronic HBV infection
0/3
Cui R 2020 [19]
7 R/R B-ALL and 13
R/R DLBCL
CD19
Chronic HBV infection
0/5
Liu W 2020 [20]
R/R B-cell lymphoma
CD19
Chronic HBV infection
0/6
Resolved HBV infec-
tion
0/11
Han L 2020 [21]
R/R MM
BCMA
Chronic HBV infection
0/1
Resolved HBV infec-
tion
1/8
Wang Y 2020 [16]
advanced B-cell
cancers
CD19, CD20 or BCMA
Chronic HBV infection
2/12
Resolved HBV infec-
tion
1/29
Cao W 2020 [14]
R/R B-ALL or DLBCL
CD19 and CD22 CAR T
cocktail
Chronic HBV infection
1/19
Resolved HBV infec-
tion
0/37
Ma Y 2021 [22]
One stomach lymph-
oblastic lymphoma
and one R/R DLBCL
CD19 and CD22 CAR T
cocktail
Chronic HBV infection
2/2
Li P 2021 [23]
R/R B-cell malignan-
cies
CD19
Resolved HBV infec-
tion
2/30
Kong D 2023 [24]
R/R DLBCL
CD19, CD19 com-
bined with CD20
or CD22, tandem
CD19/CD22
Chronic HBV infection
1/6
Resolved HBV infec-
tion
1/25
Fu S 2023 [25]
R/R MM
BCMA
Chronic HBV infection
1/7
Resolved HBV infec-
tion
3/43 Table 1 Summary of HBV reactivation cases in CAR T-cell therapy
Study
Disease
Targets of CAR T-cell
therapy
HBV characteristic
at baseline
Numbers of
reactivation/
total
Strati P 2019 [17]
R/R-DLBCL
CD19
Resolved HBV infec-
tion
1/3
Yang C 2020 [15]
R/R-DLBCL
CD19
Chronic HBV infection
3/15
Lai P 2020 [18]
R/R B-ALL
CD19
Chronic HBV infection
0/3
Cui R 2020 [19]
7 R/R B-ALL and 13
R/R DLBCL
CD19
Chronic HBV infection
0/5
Liu W 2020 [20]
R/R B-cell lymphoma
CD19
Chronic HBV infection
0/6
Resolved HBV infec-
tion
0/11
Han L 2020 [21]
R/R MM
BCMA
Chronic HBV infection
0/1
Resolved HBV infec-
tion
1/8
Wang Y 2020 [16]
advanced B-cell
cancers
CD19, CD20 or BCMA
Chronic HBV infection
2/12
Resolved HBV infec-
tion
1/29
Cao W 2020 [14]
R/R B-ALL or DLBCL
CD19 and CD22 CAR T
cocktail
Chronic HBV infection
1/19
Resolved HBV infec-
tion
0/37
Ma Y 2021 [22]
One stomach lymph-
oblastic lymphoma
and one R/R DLBCL
CD19 and CD22 CAR T
cocktail
Chronic HBV infection
2/2
Li P 2021 [23]
R/R B-cell malignan-
cies
CD19
Resolved HBV infec-
tion
2/30
Kong D 2023 [24]
R/R DLBCL
CD19, CD19 com-
bined with CD20
or CD22, tandem
CD19/CD22
Chronic HBV infection
1/6
Resolved HBV infec-
tion
1/25
Fu S 2023 [25]
R/R MM
BCMA
Chronic HBV infection
1/7
Resolved HBV infec-
tion
3/43 Table 1 Summary of HBV reactivation cases in CAR T-cell therapy Table 1 Summary of HBV reactivation cases in CAR T-cell therapy
R/R Relapsed/refractory HBV Hepatitis B virus CAR T Chimeric antigen receptor T cell DLBCL Diffuse large B cell lymphoma B ALL B cell acute lymphoblastic leukemia
Study
Disease
Targets of CAR T-cell
therapy
HBV characteristic
at baseline
Numbers of
reactivation/
total
Prophylactic
antiviral therapy
Clinical
manifestations and
outcomes
Strati P 2019 [17]
R/R-DLBCL
CD19
Resolved HBV infec-
tion
1/3
All received entecavir
Successfully treated
with entecavir
Yang C 2020 [15]
R/R-DLBCL
CD19
Chronic HBV infection
3/15
All received entecavir
or Lamivudine
Asymptomatic HBV
reactivation
Lai P 2020 [18]
R/R B-ALL
CD19
Chronic HBV infection
0/3
All received entecavir
or tenofovir
No HBV reactivation
Cui R 2020 [19]
7 R/R B-ALL and 13
R/R DLBCL
CD19
Chronic HBV infection
0/5
All received pro-
phylactic entecavir
or tenofovir
No HBV reactivation
Liu W 2020 [20]
R/R B-cell lymphoma
CD19
Chronic HBV infection
0/6
All 6 patients were
on prophylactic
entecavir
No HBV reactivation
Resolved HBV infec-
tion
0/11
Only 2 of the 11 were
on prophylactic ente-
cavir or lamivudine
No HBV reactivation
Han L 2020 [21]
R/R MM
BCMA
Chronic HBV infection
0/1
All received entecavir
or
lamivudine
No HBV reactivation
Resolved HBV infec-
tion
1/8
Only 2 of the 8 were
on prophylactic ente-
cavir or lamivudine
One asymptomatic HBV
reactivation in no pro-
phylactic group
Wang Y 2020 [16]
advanced B-cell
cancers
CD19, CD20 or BCMA
Chronic HBV infection
2/12
All received entecavir,
tenofovir disoproxil,
or lamivudine
Asymptomatic HBV
reactivation
Resolved HBV infec-
tion
1/29
No
Asymptomatic HBV
reactivation
Cao W 2020 [14]
R/R B-ALL or DLBCL
CD19 and CD22 CAR T
cocktail
Chronic HBV infection
1/19
All received prophy-
lactic entecavir
Severe hepatitis
and died of septic
shock
Resolved HBV infec-
tion
0/37
Only 2 of the 37
received prophylactic
entecavir
No HBV reactivation
Ma Y 2021 [22]
One stomach lymph-
oblastic lymphoma
and one R/R DLBCL
CD19 and CD22 CAR T
cocktail
Chronic HBV infection
2/2
All received prophy-
lactic entecavir
Both successfully
treated with entecavir
and tenofovir
Li P 2021 [23]
R/R B-cell malignan-
cies
CD19
Resolved HBV infec-
tion
2/30
No antiviral prophy-
laxis
One asymptomatic
HBV reactivation
and one severe hepa-
titis, both successfully
treated with entecavir
Kong D 2023 [24]
R/R DLBCL
CD19, CD19 com-
bined with CD20
or CD22, tandem
CD19/CD22
Chronic HBV infection
1/6
All received pro-
phylactic entecavir
or adefovir combined
with entecavir
Hepatic malignant
tumor
Resolved HBV infec-
tion
1/25
19 received prophy-
lactic entecavir
One asymptomatic HBV
reactivation in no pro-
phylactic group
Fu S 2023 [25]
R/R MM
BCMA
Chronic HBV infection
1/7
6 patients received
entecavir and 1
received tenofovir
One developed to liver
failure. R/R Relapsed/refractory, HBV Hepatitis B virus, CAR T Chimeric antigen receptor T-cell, DLBCL Diffuse large B-cell lymphoma, B-ALL B-cell acute lymphoblastic leukemia,
MM Multiple myeloma, BCMA B cell maturation antigen Similarly, in another 2 studies which included 15
and 12 patients with B-cell or plasma-cell malignancies
and concomitant chronic HBV infection, respectively, 3
(20%) and 2 (16.7%) patients experienced HBV reactiva-
tion [15, 16]. Even with the continuous administration of
NAs, HBV reactivation in these patients occurred mainly
within 6 months after CAR T-cell infusion and mani-
fested asymptomatically as an increase in HBV DNA
copy number while without hepatitis flare. Because of
the high incidence of HBV reactivation in these studies, It remains uncharted territory to fully understand the
impact of CAR T-cell therapy on HBV reactivation in
T-cell malignancies. A noteworthy case involves a patient Lin et al. Holistic Integrative Oncology (2024) 3:16 Lin et al. then died,
Resolved HBV infec-
tion
3/43
No
One hepatitis flare
and successfully treated
with entecavir, and two
asymptomatic HBV
reactivation Kong D 2023 [24]
R/R DLBCL
CD19, CD19 com-
bined with CD20
or CD22, tandem
CD19/CD22
Chronic HBV infection
1/6
Resolved HBV infec-
tion
1/25
Fu S 2023 [25]
R/R MM
BCMA
Chronic HBV infection
1/7
Resolved HBV infec-
tion
3/43 Lin et al. Holistic Integrative Oncology (2024) 3:16 Page 4 of 7 resolved HBV infection who test positive for HBV DNA
should receive prophylactic NAs therapy before initiat-
ing CAR T-cell therapy. It’s essential to recognize that
even HBV DNA-negative patients with resolved HBV
infection must adopt a preemptive treatment approach,
as lethal HBV reactivation can still occur following
CAR T-cell infusion [19]. In patients with resolved HBV
infection, baseline HBsAb seronegativity may serve as a
potential risk factor for HBV reactivation, particularly
in the absence of antiviral prophylaxis [23]. Addition-
ally, an HBsAb titer of 56.48 IU/ml has been identified
as a critical threshold for predicting HBV reactivation, a
finding validated in both rituximab-containing chemo-
therapy and CAR T-cell therapy [20, 28]. HBcAb repre-
sents another important biomarker, with a cutoff value
of 6.41 IU/ml serving as a cautionary indicator of poten-
tial HBV recurrence [29]. Furthermore, the presence of
HBeAg positivity is considered a risk factor for reactiva-
tion and is associated with elevated viral loads and infec-
tivity levels [15]. with T-cell acute lymphoblastic leukemia (T-ALL) who
exhibited high HBV DNA levels. This patient underwent
treatment with donor-derived anti-CD7 CAR T-cells, fol-
lowed by allogeneic hematopoietic stem cell transplanta-
tion while receiving continuous antiviral therapy. There
was a transient surge in HBV DNA levels observed two
weeks following CAR T-cell infusion. However, these
levels quickly diminished to a safe range. Impressively,
by the 10-month mark following CAR T-cell reinfusion
(which was 8 months after transplantation), the patient
achieved complete remission with undetectable HBV
DNA levels [27]. This case underscores the significance of
not overlooking the possibility of HBV reactivation in the
context of CAR T-cell therapy for different malignancies
beyond B-cells or plasma cells. In addition, although higher serum interleukin-6 had
been observed in patients with chronic HBV infection
when receiving CAR T-cell therapy, there is no clear evi-
dence that HBV infection affects the safety or efficacy of
CAR T-cell therapy [17]. 2.2 Prophylactic NA therapyh y
y
The aforementioned populations with a high risk of HBV
reactivation should receive highly potent and low-resist-
ance prophylactic NAs before undergoing CAR T-cell
therapy. These NAs include entecavir (ETV), tenofovir
disoproxil fumarate (TDF), and tenofovir alafenamide
fumarate tablets (TAF). Commencement of NA treat-
ment typically initiates one week before, or at the latest,
concurrently with prechemotherapy procedures. Cur-
rently, most guidelines recommend that patients with
hematologic malignancies should receive prophylactic
NAs for 12–18 months after immunochemotherapy. Because the sustained duration of B lymphocyte deple-
tion is not consistent between different individuals after
receiving a CAR T-cell infusion, the course of prophy-
lactic NAs should be determined by the duration of B
lymphocyte depletion in patients [30]. We recommend
monitoring the levels of peripheral blood lymphocyte
subpopulations and serum immunoglobulins (IgG, IgM,
and IgA) in patients with prophylactic NAs. Prophylactic
NAs should be given at least 12–18 months after the lev-
els of peripheral blood B lymphocytes and serum immu-
noglobulins are restored. fi
Taken together, these findings collectively underscore
the indispensable and efficacious role of NAs prophylaxis
in averting HBV reactivation among patients with con-
comitant chronic HBV infection or resolved HBV infec-
tion when undergoing CAR T-cell therapy. Importantly,
it appears that the presence of chronic or resolved HBV
infection does not impair the safety or efficacy of CAR
T-cell therapy. Given the retrospective nature and limited
scale of these studies, validation in multicenter prospec-
tive trials is warranted. Both animal studies and clinical
studies have shown that HBV infection did not increase
the incidence or severity of hepatotoxicity, cytokine
release syndrome (CRS), or neurotoxicity during CAR
T-cell therapy and did not compromised the activity and
clinical efficacy of CAR T-cells [14, 18, 24, 25].i 2 Prevention and Treatments of HBV reactivation
during CAR T‑cell therapy 2.1 Identification of high‑risk populations
HBV serological markers including HBsAg, anti-hep-
atitis B surface antigen (HBsAb) and anti-hepatitis B
core antigen (HBcAb), HBV DNA, and liver function
indexes (alanine aminotransferase, aspartate aminotrans-
ferase, bilirubin, albumin) should be routinely assessed
in patients with hematologic malignancies who receive
CAR T-cell therapy. To effectively mitigate the risk of
HBV reactivation in these patients, a comprehensive
evaluation of risk factors is crucial, guiding the adop-
tion of appropriate prevention and treatment strategies
(Fig. 1). Notably, both HBsAg positivity and HBV DNA
positivity represent virological risk factors for HBV reac-
tivation. Patients with either chronic HBV infection or 2.3 Monitoring and treatment of HBV reactivation While undergoing NAs treatment, it is crucial to main-
tain regular monitoring of HBV DNA levels, HBV sero-
logical markers, and liver function. Monitoring intervals
should occur at least every one to three months, ensur-
ing early intervention when necessary. HBV reactivation
often occurs within 6 months after CAR T-cell infusion, Page 5 of 7 Lin et al. Holistic Integrative Oncology (2024) 3:16 Lin et al. Holistic Integrative Oncology Fig. 1 Prevention and management strategies of HBV reactivation in CAR T-cell therapy. The necessity for prophylactic Nucleotide Analogue (NAs)
treatment hinges upon the baseline HBV status of patients prior to CAR T-cell reinfusion. Patients with chronic HBV infection (HBsAg +) or resolved
HBV infection (HBsAg- HBcAb +) who exhibit high-risk characteristics (such as a low HBsAb titer or the presence of positive HBeAg) should continue
prophylactic NAs until 12–18 months after the restoration of peripheral blood B lymphocytes and serum immunoglobulin levels. HBV status should
be monitored regularly until more than one year after NAs withdrawal, with monthly HBV monitoring in the first 6 months and once every 3 months
thereafter. In the event of HBV recurrence, immediate initiation of antiviral therapy with NAs that do not share cross-resistance is paramount. CAR
T-cell: chimeric antigen receptor T-cell; NAs: nucleotide analogs. HBsAg: hepatitis B surface antigen; HBsAb: anti-hepatitis B surface antigen; HBcAb:
anti-hepatitis B core antigen; HBeAg: hepatitis B e antigen; HBV: hepatitis B virus Fig. 1 Prevention and management strategies of HBV reactivation in CAR T-cell therapy. The necessity for prophylactic Nucleotide Analogue (NAs)
treatment hinges upon the baseline HBV status of patients prior to CAR T-cell reinfusion. Patients with chronic HBV infection (HBsAg +) or resolved
HBV infection (HBsAg- HBcAb +) who exhibit high-risk characteristics (such as a low HBsAb titer or the presence of positive HBeAg) should continue
prophylactic NAs until 12–18 months after the restoration of peripheral blood B lymphocytes and serum immunoglobulin levels. HBV status should
be monitored regularly until more than one year after NAs withdrawal, with monthly HBV monitoring in the first 6 months and once every 3 months
thereafter. In the event of HBV recurrence, immediate initiation of antiviral therapy with NAs that do not share cross-resistance is paramount. CAR
T-cell: chimeric antigen receptor T-cell; NAs: nucleotide analogs. 2.3 Monitoring and treatment of HBV reactivation HBsAg: hepatitis B surface antigen; HBsAb: anti-hepatitis B surface antigen; HBcAb:
anti-hepatitis B core antigen; HBeAg: hepatitis B e antigen; HBV: hepatitis B virus viral resistance mutations should be considered, particu-
larly in patients with lamivudine resistance who are more
susceptible to entecavir resistance mutations [32, 33]. Alongside drug-resistance gene testing, adjustments to
antiviral treatment regimens should be made, including
switching to or adding NAs drugs with no cross-resist-
ance, such as ETV, TDF and TAF. However, potential
adverse reactions associated with NAs should be moni-
tored. Multidisciplinary therapy may be initiated if nec-
essary. Additionally, based on the patient’s liver function,
hepatoprotective drugs and supportive treatments should
be administered accordingly. during which the above indicators should be monitored
monthly and once every three months thereafter. Because
HBV reactivation can still recur after discontinuation of
NAs, the above indicators should continue to be moni-
tored for 12 months after NA withdrawal, at a frequency
of once every one to three months, especially within
six months. For patients with resolved HBV infection
who receive preemptive treatment, a similar monitor-
ing regimen is advised [18, 31]. This vigilant monitoring
approach is essential for maintaining patient safety and
addressing any potential HBV reactivation promptly. If HBV reactivation is suspected, prompt assessment
and treatment protocols should be initiated to prevent
the occurrence of fatal acute liver failure. For patients
who have not received prophylactic NAs, immediate
initiation of NAs therapy, even before confirmation of
HBV reactivation, is essential to prevent further increase
in HBV DNA copies. In cases where HBV reactivation
occurs despite prophylactic NAs use, the possibility of Funding 17. Strati P, Nastoupil LJ, Fayad LE, Samaniego F, Adkins S, Neelapu SS. Safety
of CAR T-cell therapy in patients with B-cell lymphoma and chronic
hepatitis B or C virus infection. Blood. 2019;133(26):2800–2. 18. Lai P, Chen X, Qin L, et al. The efficacy and safety of CAR-T cell therapy in
patients with refractory ALL and concomitant HBV infection. Leukemia. 2020;34(10):2790–3. 3 Conclusion CAR T-cell therapy represents a groundbreaking
advancement in the treatment of hematological malig-
nancies and is gaining increasing traction. However, it is
essential to conduct meticulous evaluations of the safety
and efficacy of CAR T-cell therapy, especially in specific Lin et al. Holistic Integrative Oncology (2024) 3:16 Page 6 of 7 patient subgroups. A substantial portion of patients
with hematologic malignancies presents concurrent
chronic or resolved HBV infection. Consequently, when
administering CAR T-cell therapy to such individu-
als, a heightened awareness of the risk of HBV reactiva-
tion is paramount. The continuous prophylactic use of
NAs has demonstrated its effectiveness in curbing HBV
recurrence. Equally important is the early identifica-
tion of high-risk populations and the implementation
of long-term, regular monitoring of HBV DNA levels. In addition, based on several independent clinical stud-
ies, we concluded that the infection status of HBV does
not compromise the safety and efficacy of CAR T-cell
therapy. Therefore, CAR T-cell therapy for patients with
hematological malignancies and concomitant HBV infec-
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(CAR)-T-cell therapy in persons with advanced B-cell cancers and hepati-
tis B virus-infection. Leukemia. 2020;34(10):2704–7. References Abbreviations
B-ALL
B-cell acute lymphoblastic leukemia
BCMA
B cell maturation antigen
CAR
Chimeric antigen receptor
CRS
Cytokine release syndrome
DLBCL
Diffuse large B-cell lymphoma
ETV
Entecavir
HBcAb
Anti-hepatitis B core antigen
HBeAg
Hepatitis B e antigen
HBsAb
Anti-hepatitis B surface antigen
HBsAg
Hepatitis B surface antigen
HBV
Hepatitis B virus
MM
Multiple myeloma
NAs
Nucleotide analogs
NHL
Non-Hodgkin lymphoma
R/R
Relapsed/refractory
T-ALL
T-cell acute lymphoblastic leukemia
Acknowledgements
Not applicable
Authors’ contributions
HL and ZD searched the literatures and wrote this manuscript. LH and XZ
revised the manuscript. All authors read and approved the final manuscript. Funding
This work was supported by the National Natural Science Foundation of China
(Grant No. 82170167 to Xiaoxi Zhou). Availability of data and materials
Not applicable. Declarations
Ethics approval and consent to participate
Not applicable. Consent for publication
Not applicable. Competing interests
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cell therapy in DLBCL patients with concomitant chronic HBV infection. Leukemia. 2020;34(11):3055–9. Competing interests Prior exposure to lamivudine increases
entecavir resistance risk in chronic hepatitis B Patients without detectable
lamivudine resistance. Antimicrob Agents Chemother. 2014;58(3):1730–7. Competing interests 22. Ma Y, Yang L, Bao Y, Yang Y, Chen L, Zheng M. Case Report: Post-CAR-T
Infusion HBV Reactivation in Two Lymphoma Patients Despite Entecavir
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Did an ancient chlamydial endosymbiosis facilitate the establishment of primary plastids?
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Did an ancient chlamydial endosymbiosis facilitate the
establishment of primary plastids?
Jinling Huang*†‡ and Johann Peter Gogarten‡ Addresses: *Department of Biology, Howell Science Complex, East Carolina University, Greenville, NC 27858, USA. †NASA Astrobiology
Institute at Marine Biological Laboratory, Woods Hole, MA 02543, USA. ‡Department of Molecular and Cell Biology, University of Connecticut,
91 North Eagleville Road, Storrs, CT 06269-3125, USA. Correspondence: Jinling Huang. Email: huangj@ecu.edu Received: 30 November 2006
Revised: 6 March 2007
Accepted: 4 June 2007 The electronic version of this article is the complete one and can be
found online at http://genomebiology.com/2007/8/6/R99 © 2007 Huang and Gogarten; licensee BioMed Central Ltd. This is an open access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Primary plastid evolution
<p>Phylogenomic analyses of the red alga <it>Cyanidioschyzon merolae </it>shows that at least 21 genes were transferred between
chlamydiae and primary photosynthetic eukaryotes suggesting an ancient chlamydial endosymbiosis with the ancestral primary photosyn-
thetic eukaryote </p> Abstract Background: Ancient endosymbioses are responsible for the origins of mitochondria and plastids,
and they contribute to the divergence of several major eukaryotic groups. Although chlamydiae, a
group of obligate intracellular bacteria, are not found in plants, an unexpected number of chlamydial
genes are most similar to plant homologs, which, interestingly, often contain a plastid-targeting
signal. This observation has prompted several hypotheses, including gene transfer between
chlamydiae and plant-related groups and an ancestral relationship between chlamydiae and
cyanobacteria. Results: We conducted phylogenomic analyses of the red alga Cyanidioschyzon merolae to identify
genes specifically related to chlamydial homologs. We show that at least 21 genes were transferred
between chlamydiae and primary photosynthetic eukaryotes, with the donor most similar to the
environmental Protochlamydia. Such an unusually high number of transferred genes suggests an
ancient chlamydial endosymbiosis with the ancestral primary photosynthetic eukaryote. We
hypothesize that three organisms were involved in establishing the primary photosynthetic lineage:
the eukaryotic host cell, the cyanobacterial endosymbiont that provided photosynthetic capability,
and a chlamydial endosymbiont or parasite that facilitated the establishment of the cyanobacterial
endosymbiont. Conclusion: Our findings provide a glimpse into the complex interactions that were necessary to
establish the primary endosymbiotic relationship between plastid and host cytoplasms, and thereby
explain the rarity with which long-term successful endosymbiotic relationships between
heterotrophs and photoautotrophs were established. Our data also provide strong and
independent support for a common origin of all primary photosynthetic eukaryotes and of the
plastids they harbor. dria and plastids are derived respectively from an α-proteo-
bacterial and a cyanobacterial endosymbiont in early
eukaryotes [1]. It also has been suggested that the nucleus, a Open Access Open Access Open Access Results and discussion Chlamydiae are a group of obligate intracellular bacteria of
uncertain evolutionary position [13-15]. Many chlamydiae,
including Chlamydophila pneumoniae and Chlamydia tra-
chomatis, are important pathogens in humans and other ani-
mals [16] whereas others such as Protochlamydia,
Neochlamydia, and Fritschea are endosymbionts in environ-
mental amoebae and insects [17,18]. Although the available
evidence suggests increasing chlamydial diversity in free-liv-
ing amoebae and in the environment [19], thus far no chlamy-
dial species has been reported in photosynthetic eukaryotes
or plastid-containing lineages. However, chlamydial genome
analyses revealed an unexpected number of genes that are
most similar to plant homologs [20,21], which, interestingly,
often contain a plastid-targeting signal [13]. This observation
has prompted several hypotheses, notably an ancestral evolu-
tionary relationship between cyanobacteria (plastids) and
chlamydiae [13] and gene transfer between the two groups
with the donor being either chlamydiae [22,23] or plant-
related groups [21,24,25]. Additionally, it has also been sug-
gested that plants might have acquired these genes from
mitochondria [26] or through intermediate vectors such as
insects [17]. y
g
The nuclear genome of Cyanidioschyzon contains 4,771 pre-
dicted protein-coding genes [32]. Phylogenomic screen and
subsequent phylogenetic analyses identified 16 probable
chlamydiae-related genes in Cyanidioschyzon, 14 of which
were also found in green plants. Five other previously
reported genes [13,23] from green plants were also classified
as chlamydiae-related after careful re-analyses. The genome
sequences of glaucophytes are currently not publicly availa-
ble, but the gene encoding ATP/ADP translocase is reportedly
present in the glaucophyte Cyanophora paradoxa and the
diatom Odontella sinensis [25]. In our search of the Taxo-
nomically Broad EST Database (TBestDB) [33], ATP/ADP
translocase homologs were also found in another glaucophyte
(Glaucocystis nostochinearum), euglenids (Astasia longa
and Euglena gracilis), and a haptophyte (Pavlova lutheri). This suggests that chlamydiae-related genes are present in all
primary photosynthetic eukaryotic lineages and that the
ADP/ATP translocase has been retained in at least some sec-
ondary photosynthetic groups (eukaryotic lineages that
emerged by engulfing another algal cell as endosymbiont). Therefore, a total of 21 genes from primary photosynthetic
eukaryotes are listed here as chlamydiae-related (Table 1). Sequences that are not exclusively related to chlamydial
homologs and those that form a monophyletic group with
chlamydial homologs but with insufficient bootstrap support
(<80%) are not included. These very stringent criteria
excluded a large portion (18/37) of previously reported
chlamydiae-related plant sequences [13,23]. Background Background
Ancient symbioses are responsible for some of the major
eukaryotic innovations. It is widely accepted that mitochon- Genome Biology 2007, 8:R99 Genome Biology 2007, 8:R99 R99.2 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.2 Genome Biology 2007, Volume 8, Issue 6, Article R9 http://genomebiology.com/2007/8/6/R99 hallmark of eukaryotic cells, either arose directly from or was
mediated by an ancient symbiosis between archaeal and bac-
terial partners [2-7]. Additionally, secondary and tertiary
symbioses through engulfment of a plastid-containing cell
played an important role in the evolution of several major
eukaryotic lineages, including heterokonts, apicomplexans,
dinoflagellates, euglenids, and others [8-12]. Undoubtedly,
the evolution of extant eukaryotes was significantly shaped by
past symbioses. chlamydial homologs. Our data suggest a likely ancient sym-
biosis (sensu deBary; including mutualism, commensalisms,
and parasitism) [31] between a chlamydial endosymbiont and
the ancestor of primary photosynthetic eukaryotes. The
ancient chlamydial endosymbiont contributed genes to the
nuclear genome of primary photosynthetic eukaryotes and
might have facilitated the early establishment of plastids. Table 1 Table 1
Chlamydiae-like genes detected in red algae and green plants
Gene name or gene product
Presence
Putative function
Phosphoglycerate mutase
G
Glycolysis
CMP-KDO synthetase
G
Cell envelope formation
4-diphosphocytidyl-2-C-methyl-D-erythritol kinase (ispE)
R and G
Isoprenoid biosynthesis
Polynucleotide phosphorylase
R and G
RNA degradation
Aspartate transaminase
R and G
Amino acid metabolism
Tyrosyl-tRNA synthetase
R and G
Translation
Oligoendopeptidase F
R
Amino acid biosynthesis
2-C-methyl-D-erythritol 4-phosphate cytidylyltransferase (ispD)
R and G
Isoprenoid biosynthesis
Enoyl-ACP reductase (fabI)
R and G
Fatty acid biosynthesis
23S rRNA (Uracil-5-)-methyltransferase
R and G
RNA modification
Glycerol-3-phosphate acyltransferase
R and G
Phospholipid biosynthesis
ADT/ATP translocase
R and G
ATP/ADP transport
Isoamylase
R and G
Starch biosynthesis
Phosphate transporter
G
Phosphate transport
Hypothetical protein
R
Unknown
β-Ketoacyl-ACP synthase (fabF)
R and G
Fatty acid biosynthesis
Malate dehydrogenase
G
Energy conversion
Sodium:hydrogen antiporter
R and G
Ion transport
4-hydroxy-3-methylbut-2-en-1-yl diphosphate synthase (gcpE) (GcpE)
R and G
Isoprenoid biosynthesis
Sugar phosphate isomerase
G
Sugar interconversion
Cu-ATPase
R and G
Ion transport
The genome sequences of glaucophytes are not available for comparison, except for the ATP/ADP translocase, which is reportedly present in the
glaucophyte Cyanophora paradoxa [25] and was also identified in another glaucophyte Glaucocystis from the Taxonomically Broad EST Database
(TBestDB). G, green plants; R, red algae. The genome sequences of glaucophytes are not available for comparison, except for the ATP/ADP translocase, which is reportedly present in the
glaucophyte Cyanophora paradoxa [25] and was also identified in another glaucophyte Glaucocystis from the Taxonomically Broad EST Database
(TBestDB). G, green plants; R, red algae. The genome sequences of glaucophytes are not available for comparison, except for the ATP/ADP translocase, which is reportedly present in the
glaucophyte Cyanophora paradoxa [25] and was also identified in another glaucophyte Glaucocystis from the Taxonomically Broad EST Database
(TBestDB). G, green plants; R, red algae. to function in plastids in plants [13]. However, the Arabidop-
sis CMP-KDO synthetase homolog (GenBank: NP_175708),
although containing a weak plastid-targeting signal (score
0.504 and 0.610 from ChloroP and TargetP, respectively), is
believed to be associated with the endomembrane of plant
cells [23]. because of insufficient phylogenetic signal as evidenced by
low internal bootstrap support, and sometimes because of
possible differential gene losses or other evolutionary scenar-
ios (for instance, see Figure 2). Results and discussion Reconstructing possible evolutionary scenarios that explain
the chlamydial and plant sequence similarity requires an
understanding of the taxonomic distribution and the origin of
all involved genes. However, available phylogenetic data from
chlamydial genome analyses often suffer from small taxo-
nomic sample size [20,21]. Most other relevant studies are
heavily biased toward the gene encoding ATP/ADP translo-
case, which has an uncertain evolutionary origin and a nar-
row distribution, mainly in obligate intracellular bacteria
(chlamydiae and rickettsiae) and photosynthetic eukaryotes
[22,25-28]. The evolutionary history of a single gene, even if
correctly interpreted, might not reflect those of others. If a
single evolutionary event underlies the current observation of
chlamydial and plant sequence similarity, then a compatible
evolutionary history of multiple genes should provide more
convincing evidence. The chlamydiae-like genes identified in this study does not
constitute an accurate list of all chlamydiae-related genes in
primary photosynthetic eukaryotes, but rather is an estimate
from our phylogenomic analyses. This number is probably an
underestimate, because the evolutionary origin of many
genes is difficult to ascertain using available phylogenetic
algorithms, and because some other chlamydiae-like genes
may exist in glaucophytes and other red algae but are not
retained in the smaller genome of Cyanidioschyzon. With the
exception of the genes encoding sugar-phosphate isomerase
and a hypothetical protein, the protein sequences of all other
genes listed in Table 1 contain a plastid-targeting signal as
predicted by ChloroP [34] or TargetP [35], or experimentally
determined to be plastid localized; this is consistent with the
previous report that chlamydiae-related gene products tend Given the common belief that all primary photosynthetic
eukaryotes, including glaucophytes, red algae, and green
plants, share a common ancestry [11,29,30], we undertook a
phylogenomic analysis of Cyanidioschyzon merolae (the only
red alga whose complete genome sequence is currently avail-
able) to search for genes that are evolutionarily related to Genome Biology 2007, 8:R99 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.3 http://genomebiology.com/2007/8/6/R99 Table 1 These chlamydial and primary
photosynthetic eukaryotic sequences also do not appear to be
particularly related to homologs from other eukaryotes (Fig-
ures 1 and 2, and Additional data file 1). The chlamydial and plant sequence similarities were previ-
ously suggested to be an indication of gene transfer from
plants or plant-related groups to chlamydiae [21,24,25,28]. However, the genes listed in Table 1 are predominantly dis-
tributed in bacteria, indicating a likely bacterial origin (Fig-
ures 1 and 2, and Additional data file 1). In all cases,
sequences from primary photosynthetic eukaryotes and
sometimes from other plastid-containing lineages form a well
supported monophyletic group with chlamydial homologs. In
most cases they are more closely related to homologs of Can-
didatus Protochlamydia amoebophila UWE25 (a chlamydial
species that is found in free-living acanthamoebae and envi-
ronmental samples and was previously classified as a member
of Parachlamydia or Parachlamydia-related [36]) than to
chlamydial sequences as a whole (Figures 1 and 2, and Addi-
tional data file 1). However, the sequence relationships
among primary photosynthetic eukaryotes vary slightly and
differ from the expected organismal relationship, mostly The bacterial nature of chlamydiae-like genes in primary pho-
tosynthetic eukaryotes suggests that they were transferred
either from chlamydiae to these eukaryotes or from plastids
to chlamydiae. The latter scenario (plastid-to-chlamydiae
transfer) implies a plastidic (cyanobacterial) origin for the
transferred genes listed (Table 1). For many of the genes, this
scenario can be rejected because it does not account for the
specific relationship of the chlamydiae-like genes in primary
photosynthetic eukaryotes to the Protochlamydia homologs
(Figure 1a-d, Figure 2a, and Additional data file 1), and
because it is incompatible with the cyanobacterial homologs
forming a well supported group that is distinct from the
chlamydial homologs (Figures 1 and 2, and Additional data
file 1). After all, plastids evolved from a cyanobacterial ances-
tor, and therefore any gene acquired by chlamydiae from
plastids should also be more closely related to cyanobacterial
than to other bacterial sequences. Is there an ancestral relationship between chlamydiae
and cyanobacteria? The relationship between chlamydiae and other bacterial
groups remains largely unresolved. Phylogenetic analyses of
16S rRNA suggested that chlamydiae form a sister group with
either planctomycetes and verrucomicrobia [14,15,20,37] or
cyanobacteria [13,38], without significant support. An ances-
tral relationship between chlamydiae and cyanobacteria was
hypothesized by Brinkman and coworkers [13], largely based
on the possession of a predicted plastid-targeting signal in
chlamydiae-like plant sequences. Those authors explicitly
excluded the possibility of horizontal gene transfer between
chlamydiae and their hosts, and assumed that these plant
plastid-targeted sequences were of cyanobacterial origin. According to Brinkman and coworkers, these plant sequences
are similar to chlamydial homologs because chlamydiae and
cyanobacteria (plastids) are evolutionarily related. A few
additional characters uniquely shared by cyanobacteria and
chlamydiae were identified in the usually structurally con-
served ribosomal superoperon [13], including the absence of
S10 and S14 genes, which are present in different chromo-
somal locations in chlamydiae and cyanobacteria. However, a
more detailed phylogenetic and comparative study of S14 sug-
gested that this gene was probably independently transferred
from α-proteobacteria to cyanobacteria, chlamydiae, and
actinomycetes [39]. Therefore, the absence of S14 from the
cyanobacterial and chlamydial ribosomal superoperons
might be due to relaxed selection to maintain redundant
homologs in the genome, rather than an indication of evolu-
tionary relatedness between the two groups. In all studies of gene transfer, there are always alternative
explanations for each individual gene tree (for example, see
the discussion of Figure 2 in section "Further evidence for an
ancient chlamydial endosymbiosis with primary photosyn-
thetic eukaryotes") [40]. Overall, however, the pattern from
our phylogenetic analyses does not support the hypothesis
that cyanobacteria (plastids) and chlamydiae share a close
ancestral relationship. This consistent phylogenetic signal
from multiple genes should not be dismissed lightly as arti-
facts of phylogenetic reconstruction, but rather suggests a
clear evolutionary link between chlamydiae and primary
photosynthetic eukaryotes. Furthermore, given their often
specific
affinity
with
environmental
Protochlamydia
homologs (Figures 1 and 2, and Additional data file 1), if all of
these primary photosynthetic eukaryotic sequences were Although the chlamydiae-cyanobacteria hypothesis offers a
popular explanation for the sequence similarity between
chlamydiae and plants [20,22,27], it has not been rigorously
tested. The major shortcoming of this hypothesis is that all
plastids certainly are derived from a past cyanobacterial
rather than a chlamydial endosymbiont. Table 1 Additionally, five of the Genome Biology 2007, 8:R99 R99.4 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.4 Genome Biology 2007, Volume 8, Issue 6, Article R99 http://genomebiology.com/2007/8/6/R99 gy
g
g
p g
gy
gure 1 (see legend on next page)
0.2
Rhodopseudomonas
Agrobacterium
Haemophilus
Escherichia
Nitrosomonas
Streptomyces
Frankia
Mycobacterium
Thermus
Deinococcus
Thermotoga
Anabaena
Crocosphaera
Gloeobacter
Synechococcus
Prochlorococcus
Pelobacter
Syntrophomonas
Solibacter
Moorella
Blastopirellula
Aquifex
Leptospira
Chlamydia
Chlamydophila
Cyanidioschyzon
Protochlamydia
Arabidopsis
Oryza
Staphylococcus
Staphylococcus
Bacteroides
Cytophaga
Clostridium
Thermoanaerobacter
Bacillus
Chlorobium
Fusobacterium
100/95
99/98
88/93
95/90
76/86
100/100
66/65
91/93
100/100
63/51
100/100
87/86
100/100
56/*
52/53
63/*
88/86
Cyanobacteria
Chlamydiae
Red algae
Chalmydiae
Green plants
Proteobacteria
High G+C gram+
Thermus/Deinococci
Delta-proteobacteria
Thermotogae
Low G+C gram+
Acidobacteria
Low G+C gram+
Planctomycetes
Aquificae
Spirochaetes
Low G+C gram+
Low G+C gram+
Chlorobi
Bacteroidetes
Fusobacteria
Rhodopseudomonas
Mesorhizobium
Deinococcus
Nitrosomonas
Rubrivivax
Pseudomonas
Haemophilus
Desulfotalea
Chlorobium
Aquifex
Bacteroides
Fusobacterium
Chlamydomonas
Oryza
Cyanidioschyzon
Protochlamydia
Chlamydophila
Chlamydia
Solibacter
Listeria
Bacillus
Nocardia
Streptomyces
Synechococcus
Prochlorococcus
Nostoc
Trichodesmium
Crocosphaera
Gloeobacter
Dehalococcoides
Blastopirellula
Treponema
Thermotoga
0.2
94/96
93/78
97/99
86/96
88/100
*/53
96/100
62/70
100/100
57/54
99/100
100/100
100/100
89/92
92/98
57/*
Chlamydiae
Cyanobacteria
Red algae
Green plants
Alpha-proteobacteria
Beta, gamma-proteobacteria
Deinococci
Delta-proteobacteria
Chlorobi
Aquificae
Bacteroidetes
Fusobacteria
Acidobacteria
Low G+C gram+
High G+C gram+
Chloroflexi
Planctomycetes
Spirochaetes
Thermotogae
Aedes
Homo
Neurospora
Dictyostelium
Arabidopsis
Tetrahymena
Rhodospirillum
Rickettsia
Campylobacter
Solibacter
Solibacter
Coxiella
Pseudomonas
Rubrivivax
Treponema
Thermus
Deinococcus
Chlamydia
Chlamydophila
Arabidopsis
Ostreococcus
Ostreococcus
Arabidopsis
Cyanidioschyzon
Protochlamydia
Brachyspira
Chlorobium
Cytophaga
Flavobacteria
Toxoplasma
Clostridium
Nostoc
Nostoc
Synechocystis
Crocosphaera
Rhodopirellula
Chloroflexus
Chloroflexus
Mycobacterium
Nocardioides
Thermotoga
Eukaryotic mt
Green plants
Red algae
Chlamydiae
Chlamydiae
Cyanobacteria
Alpha-proteobacteria
Planctomycetes
Epsilon-proteobacteria
Thermus/Deinococcus
Bacteroidetes
Gamma, beta-proteobacteria
Acidobacteria
Apicomplexan
High G+C gram+
Low G+C gram+
Chlorobi
Spirochaetes
Chloroflexi
Spirochaetes
Thermotogales
0.2
88/88
89/90
95/70
80/86
92/93
100/100
59/*
62/84
67/*
100/100
100/100
100/100
100/100
94/98
94/89
92/94
100/100
100/100
99/100
67/70
79/90
99/100
Pseudomonas
Escherichia
Ralstonia
Chlorobium
Campylobacter
Rhodospirillum
Mesorhizobium
Crocosphaera
Anabaena
Desulfovibrio
Methanosarcina
Archaeoglobus
Aquifex
Anabaena
Trichodesmium
Moorella
Dehalococcoides
Bacillus
Oryza
Chloroflexus
Pyrococcus
Aeropyrum
Clostridium
Desulfitobacterium
Bacteroides
Methanothermobacter
Geobacter
Synechococcus
Nostoc
Crocosphaera
Trichodesmium
Prochlorococcus
Leptospira
Cyanidioschyzon
Oryza
Arabidopsis
Oryza
Arabidopsis
Protochlamydia
Chlamydophila
Chlamydia
98/98
100/100
100/100
100/100
100/99
59/59
59/72
60/64
100/100
100/96
100/100
69/86
66/56
100/100
87/89
71/59
65/61
100/100
100/99
100/100
59/7599/100
97/100
55/51
95/91
100/100
Gamma, beta-proteobacteria
Chlorobi
Alpha, epsilon-proteobacteria
Cyanobacteria
Cyanobacteria
Cyanobacteria
Red algae
Green plants
Green plants
Chlamydiae
Spirochaetes
Delta-proteobacteria
Delta-proteobacteria
Aquificae
Chloroflexi
Euryarchaeota
Low G+C gram+
Chloroflexi
Low G+C gram+
Crenarchaeota
Euryarchaeota
Low G+C gram+
Bacteroidetes
Euryarchaeota
0.2
(a)
(b)
(c)
(d) 0.2
Rhodopseudomonas
Agrobacterium
Haemophilus
Escherichia
Nitrosomonas
Streptomyces
Frankia
Mycobacterium
Thermus
Deinococcus
Thermotoga
Anabaena
Crocosphaera
Gloeobacter
Synechococcus
Prochlorococcus
Pelobacter
Syntrophomonas
Solibacter
Moorella
Blastopirellula
Aquifex
Leptospira
Chlamydia
Chlamydophila
Cyanidioschyzon
Protochlamydia
Arabidopsis
Oryza
Staphylococcus
Staphylococcus
Bacteroides
Cytophaga
Clostridium
Thermoanaerobacter
Bacillus
Chlorobium
Fusobacterium
100/95
99/98
88/93
95/90
76/86
100/100
66/65
91/93
100/100
63/51
100/100
87/86
100/100
56/*
52/53
63/*
88/86
Cyanobacteria
Chlamydiae
Red algae
Chalmydiae
Green plants
Proteobacteria
High G+C gram+
Thermus/Deinococci
Delta-proteobacteria
Thermotogae
Low G+C gram+
Acidobacteria
Low G+C gram+
Planctomycetes
Aquificae
Spirochaetes
Low G+C gram+
Low G+C gram+
Chlorobi
Bacteroidetes
Fusobacteria
(a) Rhodopseudomonas
Mesorhizobium
Deinococcus
Nitrosomonas
Rubrivivax
Pseudomonas
Haemophilus
Desulfotalea
Chlorobium
Aquifex
Bacteroides
Fusobacterium
Chlamydomonas
Oryza
Cyanidioschyzon
Protochlamydia
Chlamydophila
Chlamydia
Solibacter
Listeria
Bacillus
Nocardia
Streptomyces
Synechococcus
Prochlorococcus
Nostoc
Trichodesmium
Crocosphaera
Gloeobacter
Dehalococcoides
Blastopirellula
Treponema
Thermotoga
0.2
94/96
93/78
97/99
86/96
88/100
*/53
96/100
62/70
100/100
57/54
99/100
100/100
100/100
89/92
92/98
57/*
Chlamydiae
Cyanobacteria
Red algae
Green plants
Alpha-proteobacteria
Beta, gamma-proteobacteria
Deinococci
Delta-proteobacteria
Chlorobi
Aquificae
Bacteroidetes
Fusobacteria
Acidobacteria
Low G+C gram+
High G+C gram+
Chloroflexi
Planctomycetes
Spirochaetes
Thermotogae
(b) (b) Desulfotalea
Chlorobium Bacteroides Pseudomonas
Escherichia
Ralstonia
Chlorobium
Campylobacter
Rhodospirillum
Mesorhizobium
Crocosphaera
Anabaena
Desulfovibrio
Methanosarcina
Archaeoglobus
Aquifex
Anabaena
Trichodesmium
Moorella
Dehalococcoides
Bacillus
Oryza
Chloroflexus
Pyrococcus
Aeropyrum
Clostridium
Desulfitobacterium
Bacteroides
Methanothermobacter
Geobacter
Synechococcus
Nostoc
Crocosphaera
Trichodesmium
Prochlorococcus
Leptospira
Cyanidioschyzon
Oryza
Arabidopsis
Oryza
Arabidopsis
Protochlamydia
Chlamydophila
Chlamydia
98/98
100/100
100/100
100/100
100/99
59/59
59/72
60/64
100/100
100/96
100/100
69/86
66/56
100/100
87/89
71/59
65/61
100/100
100/99
100/100
59/7599/100
97/100
55/51
95/91
100/100
Gamma, beta-proteobacteria
Chlorobi
Alpha, epsilon-proteobacteria
Cyanobacteria
Cyanobacteria
Cyanobacteria
Red algae
Green plants
Green plants
Chlamydiae
Spirochaetes
Delta-proteobacteria
Delta-proteobacteria
Aquificae
Chloroflexi
Euryarchaeota
Low G+C gram+
Chloroflexi
Low G+C gram+
Crenarchaeota
Euryarchaeota
Low G+C gram+
Bacteroidetes
Euryarchaeota
0.2
(d) Figure 1 (see legend on next page)
Aedes
Homo
Neurospora
Dictyostelium
Arabidopsis
Tetrahymena
Rhodospirillum
Rickettsia
Campylobacter
Solibacter
Solibacter
Coxiella
Pseudomonas
Rubrivivax
Treponema
Thermus
Deinococcus
Chlamydia
Chlamydophila
Arabidopsis
Ostreococcus
Ostreococcus
Arabidopsis
Cyanidioschyzon
Protochlamydia
Brachyspira
Chlorobium
Cytophaga
Flavobacteria
Toxoplasma
Clostridium
Nostoc
Nostoc
Synechocystis
Crocosphaera
Rhodopirellula
Chloroflexus
Chloroflexus
Mycobacterium
Nocardioides
Thermotoga
Eukaryotic mt
Green plants
Red algae
Chlamydiae
Chlamydiae
Cyanobacteria
Alpha-proteobacteria
Planctomycetes
Epsilon-proteobacteria
Thermus/Deinococcus
Bacteroidetes
Gamma, beta-proteobacteria
Acidobacteria
Apicomplexan
High G+C gram+
Low G+C gram+
Chlorobi
Spirochaetes
Chloroflexi
Spirochaetes
Thermotogales
0.2
88/88
89/90
95/70
80/86
92/93
100/100
59/*
62/84
67/*
100/100
100/100
100/100
100/100
94/98
94/89
92/94
100/100
100/100
99/100
67/70
79/90
99/100
Pseudomonas
Escherichia
Ralstonia
Chlorobium
Campylobacter
Rhodospirillum
Mesorhizobium
Crocosphaera
Anabaena
Desulfovibrio
Methanosarcina
Archaeoglobus
Aquifex
Anabaena
Trichodesmium
Moorella
Dehalococcoides
Bacillus
Oryza
Chloroflexus
Pyrococcus
Aeropyrum
Clostridium
Desulfitobacterium
Bacteroides
Methanothermobacter
Geobacter
Synechococcus
Nostoc
Crocosphaera
Trichodesmium
Prochlorococcus
Leptospira
Cyanidioschyzon
Oryza
Arabidopsis
Oryza
Arabidopsis
Protochlamydia
Chlamydophila
Chlamydia
98/98
100/100
100/100
100/100
100/99
59/59
59/72
60/64
100/100
100/96
100/100
69/86
66/56
100/100
87/89
71/59
65/61
100/100
100/99
100/100
59/7599/100
97/100
55/51
95/91
100/100
Gamma, beta-proteobacteria
Chlorobi
Alpha, epsilon-proteobacteria
Cyanobacteria
Cyanobacteria
Cyanobacteria
Red algae
Green plants
Green plants
Chlamydiae
Spirochaetes
Delta-proteobacteria
Delta-proteobacteria
Aquificae
Chloroflexi
Euryarchaeota
Low G+C gram+
Chloroflexi
Low G+C gram+
Crenarchaeota
Euryarchaeota
Low G+C gram+
Bacteroidetes
Euryarchaeota
0.2
(c)
(d) Figure 1 (see legend on next page)
Aedes
Homo
Neurospora
Dictyostelium
Arabidopsis
Tetrahymena
Rhodospirillum
Rickettsia
Campylobacter
Solibacter
Solibacter
Coxiella
Pseudomonas
Rubrivivax
Treponema
Thermus
Deinococcus
Chlamydia
Chlamydophila
Arabidopsis
Ostreococcus
Ostreococcus
Arabidopsis
Cyanidioschyzon
Protochlamydia
Brachyspira
Chlorobium
Cytophaga
Flavobacteria
Toxoplasma
Clostridium
Nostoc
Nostoc
Synechocystis
Crocosphaera
Rhodopirellula
Chloroflexus
Chloroflexus
Mycobacterium
Nocardioides
Thermotoga
Eukaryotic mt
Green plants
Red algae
Chlamydiae
Chlamydiae
Cyanobacteria
Alpha-proteobacteria
Planctomycetes
Epsilon-proteobacteria
Thermus/Deinococcus
Bacteroidetes
Gamma, beta-proteobacteria
Acidobacteria
Apicomplexan
High G+C gram+
Low G+C gram+
Chlorobi
Spirochaetes
Chloroflexi
Spirochaetes
Thermotogales
0.2
88/88
89/90
95/70
80/86
92/93
100/100
59/*
62/84
67/*
100/100
100/100
100/100
100/100
94/98
94/89
92/94
100/100
100/100
99/100
67/70
79/90
99/100
Pseudomonas
Escherichia
Ralstonia
Chlorobium
Campylobacter
Rhodospirillum
Mesorhizobium
Crocosphaera
Anabaena
Desulfovibrio
Methanosarcina
Archaeoglobus
Aquifex
Anabaena
Trichodesmium
Moorella
Dehalococcoides
Bacillus
Oryza
Chloroflexus
Pyrococcus
Aeropyrum
Clostridium
Desulfitobacterium
Bacteroides
Methanothermobacter
Geobacter
Synechococcus
Nostoc
Crocosphaera
Trichodesmium
Prochlorococcus
Leptospira
Cyanidioschyzon
Oryza
Arabidopsis
Oryza
Arabidopsis
Protochlamydia
Chlamydophila
Chlamydia
98/98
100/100
100/100
100/100
100/99
59/59
59/72
60/64
100/100
100/96
100/100
69/86
66/56
100/100
87/89
71/59
65/61
100/100
100/99
100/100
59/7599/100
97/100
55/51
95/91
100/100
Gamma, beta-proteobacteria
Chlorobi
Alpha, epsilon-proteobacteria
Cyanobacteria
Cyanobacteria
Cyanobacteria
Red algae
Green plants
Green plants
Chlamydiae
Spirochaetes
Delta-proteobacteria
Delta-proteobacteria
Aquificae
Chloroflexi
Euryarchaeota
Low G+C gram+
Chloroflexi
Low G+C gram+
Crenarchaeota
Euryarchaeota
Low G+C gram+
Bacteroidetes
Euryarchaeota
0.2
(c)
(d) Aedes
Homo
Neurospora
Dictyostelium
Arabidopsis
Tetrahymena
Rhodospirillum
Rickettsia
Campylobacter
Solibacter
Solibacter
Coxiella
Pseudomonas
Rubrivivax
Treponema
Thermus
Deinococcus
Chlamydia
Chlamydophila
Arabidopsis
Ostreococcus
Ostreococcus
Arabidopsis
Cyanidioschyzon
Protochlamydia
Brachyspira
Chlorobium
Cytophaga
Flavobacteria
Toxoplasma
Clostridium
Nostoc
Nostoc
Synechocystis
Crocosphaera
Rhodopirellula
Chloroflexus
Chloroflexus
Mycobacterium
Nocardioides
Thermotoga
Eukaryotic mt
Green plants
Red algae
Chlamydiae
Chlamydiae
Cyanobacteria
Alpha-proteobacteria
Planctomycetes
Epsilon-proteobacteria
Thermus/Deinococcus
Bacteroidetes
Gamma, beta-proteobacteria
Acidobacteria
Apicomplexan
High G+C gram+
Low G+C gram+
Chlorobi
Spirochaetes
Chloroflexi
Spirochaetes
Thermotogales
0.2
88/88
89/90
95/70
80/86
92/93
100/100
59/*
62/84
67/*
100/100
100/100
100/100
100/100
94/98
94/89
92/94
100/100
100/100
99/100
67/70
79/90
99/100
(c) (d) (c) 0.2 0.2 Figure 1 (see legend on next page) Figure 1 (see legend on next page) Genome Biology 2007, 8:R99 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.5 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.5 Genome Biology 2007, Volume 8, Issue 6, Article R99 http://genomebiology.com/2007/8/6/R99 Phylogenetic analyses of chlamydiae-like genes in primary photosynthetic eukaryotes
Figure 1 (see previous page)
Phylogenetic analyses of chlamydiae-like genes in primary photosynthetic eukaryotes. Table 1 Numbers above the branch show bootstrap values for maximum
likelihood and distance analyses, respectively. Asterisks indicate values lower than 50%. (a) 2-C-methyl-D-erythritol 4-phosphate cytidylyltransferase
(ispD). (b) 4-diphosphocytidyl-2-C-methyl-D-erythritol kinase (ispE). (c) β-Ketoacyl-ACP synthase (fabF). (d) Aspartate transaminase. Note that red algal
and green plant sequences form a well supported monophyletic group with environmental Protochlamydia homologs. mt, mitochondrial precursor. Colors
represent different phylogenetic affiliations. chlamydiae-like genes in photosynthetic eukaryotes lack
identifiable cyanobacterial homologs. (The genes encoding
ATP/ADP translocase, glycerol-3-phosphate acyltransferase,
oligoendopeptidase F, sodium-hydrogen antiporter, and the
malate dehydrogenase chloroplast precursors in green plants
lack significant hits to cyanobacterial sequences in GenBank
searches.) This suggests that the majority of the genes listed
in Table 1 were probably transferred from chlamydiae to pri-
mary photosynthetic eukaryotes. cyanobacterial than to chlamydial homologs, unless these
sequences diverged so rapidly as to generate long-branch
attraction artifacts or lateral gene transfer was involved. Based on this reasoning, we have paid particular attention to
the relationships among homologous sequences from
chlamydiae, cyanobacteria, and primary photosynthetic
eukaryotes in our analyses. All red algal and green plant genes listed in Table 1 clearly are
more closely related to chlamydial than to cyanobacterial
homologs, with the exception of the genes encoding 4-
hydroxy-3-methylbut-2-en-1-yl diphosphate synthase (gcpE)
and enoyl-ACP reductase (fabI), which include distinct gene
copies in red algae, glaucophytes, and other plastid-contain-
ing eukaryotes (Figure 2 and Additional data file 1). Some of
the identified chlamydiae-like genes (for instance, that
encoding polyribonucleotide phosphorylase) also contain a
number of conserved amino acid residues uniquely shared by
chlamydiae, red algae, and green plants. In our molecular
phylogenetic analyses of 12 of the genes that contain both
cyanobacterial and chlamydial homologs, the cyanobacterial
sequences form a clade that is clearly distinct from the
chlamydial homologs (Figures 1 and 2, and Additional data
file 1). Added to this observation is the fact that several
chlamydiae-like genes are not found in cyanobacteria and
that the gene encoding glycerol-3-phosphate acyltransferase
has identifiable homologs (using Arabidopsis [GenBank:
NP_849738] and Protochlamydia [GenBank: CAF24042]
sequences as queries) only in chlamydiae, red algae, green
plants, and the apicomplexan Plasmodium, which also con-
tains a nonphotosynthetic plastid. The closer relationship
between certain chlamydial and plant sequences was also
observed in independent studies [20]. Random horizontal gene transfer versus ancient
chlamydial endosymbiosis Conceivably, if horizontal gene transfer occurred from
chlamydiae to the earlier cyanobacterial progenitor of
plastids, then chlamydial genes could end up in the nuclear
genomes of photosynthetic eukaryotes following subsequent
intracellular transfer from plastids to the nucleus. Given the
number of chlamydiae-like genes detected in our analyses
and the fact that most of the original plastidic (cyanobacte-
rial) genes were lost in modern photosynthetic eukaryotes
[41], this scenario entails massive gene transfers from ancient
chlamydiae to the cyanobacterial progenitor of plastids. Although gene transfer indeed occurs frequently in prokaryo-
tes [42-44], thus far no chlamydiae-like genes have been
reported in any extant cyanobacterium to suggest such mas-
sive transfer events. Rickettsia
Mesorhizobium
Campylobacter
Rhodopirellula
Chloroflexus
Geobacter
Solibacter
Listeria
Clostridium
Nostoc
Prochlorococcus
Cyanidioschyzon
Synechococcus
Gloeobacter
Salmonella
Haemophilus
Ralstonia
Aquifex
Ralstonia
Symbiobacterium
Cytophaga
Bacteroides
Chlorobium
Frankia
Mycobacterium
Deinococcus
Thermus
Oryza
Arabidopsis
Chlamydophila
Chlamydia
Phaeodactylum
Protochlamydia
Toxoplasma
Bigelowiella
0.2
100/100
97/85
100/89
57/51
95/90
100/100
76/95
100/100
72/63
100/99
100/99
50/64
50/*
60/59
74/94
63/85
Red algae
Cyanobacteria
Cyanobacteria
Green plants
Chlamydiae
Diatom
Apicomplexan
Chlamydiae
Chlorarachniophyte
Thermus/Deinococcus
Gamma, beta-proteobacteria
Aquificae
Beta-proteobacteria
Alpha-proteobacteria
Epsilon-proteobacteria
Chloroflexi
Delta-proteobacteria
High G+C gram+
Acidobacteria
Low G+C gram+
High G+C gram+
Planctomycetes
Bacteroidetes
Chlorobi
(b) Because chlamydiae are found in insects, it has also been sug-
gested that plants might have acquired chlamydial genes
through insect feeding activities [17]. However, the presence
of chlamydiae-like genes in red algae and glaucophytes,
which are not a favorable food source for insects, makes this
scenario less likely. Furthermore, red algae, glaucophytes,
and green plants represent one of the major deep lineages of
eukaryotes [45,46]. Such an insect-to-plant transfer scenario
would also push the emergence of insects to before the split of
primary photosynthetic eukaryotes, which contradicts all
available molecular and fossil evidence [47,48]. An unexpectedly high number of chlamydial genes that are
most similar to plant homologs has been reported in several
independent studies [13,20,21]. For example, most eukary-
ote-related sequences of Chlamydia trachomatis tend to
group with plant homologs in phylogenetic analyses [21]. In a
similarity-based genome survey, sequences of rickettsia,
cyanobacteria, and chlamydiae represented only 14% of the
analyzed genes, but they accounted for 65% of bacterial genes
that were most similar to eukaryotic homologs; these cyano-
bacterial and chlamydial sequences disproportionately corre-
spond to plant proteins [13]. Is there an ancestral relationship between chlamydiae
and cyanobacteria? Rhodospirillum
Zymomonas
Silicibacter
Pseudomonas
Erwinia
Vibrio
Haemophilus
Symbiobacterium
Helicobacter
Fusobacterium
Clostridium
Bacillus
Dehalococcoides
Aquifex
Frankia
Treponema
Thermotoga
Rhodopirellula
Brucella
Deinococcus
Solibacter
Anabaena
Trichodesmium
Crocosphaera
Cyanophora
Glaucocystis
Cyanidioschyzon
Prochlorococcus
Synechococcus
Bacteroides
Porphyromonas
Cytophaga
Chlorobium
Pelodictyon
Leptospira
Ostreococcus
Isochrysis
Bigelowiella
Chlamydomonas
Arabidopsis
Plasmodium
Protochlamydia
Chlamydophila
Chlamydia
0.2
100/100
100/97
79/87
69/*
99/64
76/62
97/100
77/*
93/87
100/100
100/100
56/*
83/71
80/84
67/72
90/94
*/74
53/86
100/100
100/100
79/87
57/57
76/74
52/*
100/99
100/99
100/100
79/65
100/100
90/72
100/99
Proteobacteria
Epsilon-proteobacteria
High G+C gram+
High G+C gram+
Low G+C gram+
Planctomycetes
Fusobacteria
Chloroflexi
Acidobacteria
Alpha-proteobacteria
Deinococci
Thermotogales
Spirochaetes
Aquificae
Cyanobacteria
Cyanobacteria
Red algae
Glaucophytes
Bacteroidetes
Chlorobi
Spirochaetes
Green plant
Green plants
Chlamydiae
Apicomplexan
Chlorarachniophyte
Haptophyte
Rickettsia
Mesorhizobium
Campylobacter
Rhodopirellula
Chloroflexus
Geobacter
Solibacter
Listeria
Clostridium
Nostoc
Prochlorococcus
Cyanidioschyzon
Synechococcus
Gloeobacter
Salmonella
Haemophilus
Ralstonia
Aquifex
Ralstonia
Symbiobacterium
Cytophaga
Bacteroides
Chlorobium
Frankia
Mycobacterium
Deinococcus
Thermus
Oryza
Arabidopsis
Chlamydophila
Chlamydia
Phaeodactylum
Protochlamydia
Toxoplasma
Bigelowiella
0.2
100/100
97/85
100/89
57/51
95/90
100/100
76/95
100/100
72/63
100/99
100/99
50/64
50/*
60/59
74/94
63/85
Red algae
Cyanobacteria
Cyanobacteria
Green plants
Chlamydiae
Diatom
Apicomplexan
Chlamydiae
Chlorarachniophyte
Thermus/Deinococcus
Gamma, beta-proteobacteria
Aquificae
Beta-proteobacteria
Alpha-proteobacteria
Epsilon-proteobacteria
Chloroflexi
Delta-proteobacteria
High G+C gram+
Acidobacteria
Low G+C gram+
High G+C gram+
Planctomycetes
Bacteroidetes
Chlorobi
(a)
(b) Rhodospirillum
Zymomonas
Silicibacter
Pseudomonas
Erwinia
Vibrio
Haemophilus
Symbiobacterium
Helicobacter
Fusobacterium
Clostridium
Bacillus
Dehalococcoides
Aquifex
Frankia
Treponema
Thermotoga
Rhodopirellula
Brucella
Deinococcus
Solibacter
Anabaena
Trichodesmium
Crocosphaera
Cyanophora
Glaucocystis
Cyanidioschyzon
Prochlorococcus
Synechococcus
Bacteroides
Porphyromonas
Cytophaga
Chlorobium
Pelodictyon
Leptospira
Ostreococcus
Isochrysis
Bigelowiella
Chlamydomonas
Arabidopsis
Plasmodium
Protochlamydia
Chlamydophila
Chlamydia
0.2
100/100
100/97
79/87
69/*
99/64
76/62
97/100
77/*
93/87
100/100
100/100
56/*
83/71
80/84
67/72
90/94
*/74
53/86
100/100
100/100
79/87
57/57
76/74
52/*
100/99
100/99
100/100
79/65
100/100
90/72
100/99
Proteobacteria
Epsilon-proteobacteria
High G+C gram+
High G+C gram+
Low G+C gram+
Planctomycetes
Fusobacteria
Chloroflexi
Acidobacteria
Alpha-proteobacteria
Deinococci
Thermotogales
Spirochaetes
Aquificae
Cyanobacteria
Cyanobacteria
Red algae
Glaucophytes
Bacteroidetes
Chlorobi
Spirochaetes
Green plant
Green plants
Chlamydiae
Apicomplexan
Chlorarachniophyte
Haptophyte
(a) ema
pirellula
Epsilon-proteobacteria
High G+C gram+
High G+C gram+
Low G+C gram+
Planctomycetes
Fusobacteria
Chloroflexi
Acidobacteria
Alpha-proteobacteria
Deinococci
Thermotogales
Spirochaetes
Aquificae Is there an ancestral relationship between chlamydiae
and cyanobacteria? Even if chlamydiae
and cyanobacteria indeed shared a common ancestry, any
sequences of plastidic origin should be more closely related to Genome Biology 2007, 8:R99 R99.6 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.6 Genome Biology 2007, Volume 8, Issue 6, Article R99 http://genomebiology.com/2007/8/6/R99 indeed of plastidic origin (even though they do not group with
cyanobacterial homologs), then this would make chlamydiae
a paraphyletic group. Such an observation contradicts the
common belief that chlamydiae are monophyletic [14,15] and
weighs further against the hypothesis that chlamydiae and
cyanobacteria are sister taxa. Random horizontal gene transfer versus ancient
chlamydial endosymbiosis Although our focus is to
elucidate the cause of chlamydial and plant sequence similar-
ity rather than to reiterate the previous conclusion, our anal-
yses yielded similar findings. For all likely transferred genes
in Cyanidioschyzon whose origins can be reliably inferred,
chlamydiae-like genes (n = 16) account for the greatest
number from any single group other than cyanobacteria
(plastids) and α-proteobacteria (mitochondria), and are fol-
lowed by five genes from γ-proteobacteria and β-proteobacte-
ria. The latter are also the most represented bacterial groups Primary ph
and chlamy
Figure 2 About 30 genes of plastidic origin (<1% of the
nuclear genome) were reported in the human malaria para-
site Plasmodium falciparum [55] and only two such genes
were identified in Cryptosporidium that probably lost the
plastid entirely [49]. None of these intracellularly transferred
genes in apicomplexans are related to photosynthesis. A sim-
ilar scenario was suggested for Entamoeba, which contains a
reduced mitochondrion-derived organelle and appears to
have lost most mitochondrial pathways [56]. Therefore, the
number of chlamydiae-like genes identified in photosynthetic
eukaryotes, albeit being a small fraction of the cyanobacterial
genes reported in Arabidopsis and still lower than reported in
some heterotrophic apicomplexans, is many times higher
than in Cryptosporidium. This lower number of chlamydiae-
like genes in primary photosynthetic eukaryotes is in accord-
ance with the seeming absence of chlamydial endosymbionts
in modern plastid-containing lineages. in GenBank (the taxonomy browser in Entrez of the National
Center for Biotechnology Information [NCBI] reported
1,692,357 protein sequences from γ and β-proteobacteria, and
only 28,831 from the chlamydiae/verrucomicrobia group as
of 4 November 2006). Because of a greater level of stringency,
the number of chlamydiae-like genes identified in our phylo-
genetic analyses is much lower than previously reported (19
versus the 37 reporrted by Brinkman and coworkers [13] in
green plants), but this number is still striking, given that
using similar methods only a total of 31 genes acquired from
all other sources (including those likely from plants and plas-
tids) were identified in the apicomplexan Cryptosporidium
parvum [49] and about 50 genes in the kinetoplastid
Trypanosoma brucei [50]. The high number of genes transferred between chlamydiae
and photosynthetic eukaryotes is probably due to a more sta-
ble association of these two groups in the past. In general,
such an association could theoretically occur in the form of
symbiosis or physical contact between donor and recipient
organisms. However, given the distribution of chlamydiae-
like genes mainly in primary photosynthetic eukaryotes and
the fact that all extant chlamydial species are obligate endo-
symbionts, we propose that these genes resulted from an
ancient chlamydial endosymbiosis with the ancestor of pri-
mary photosynthetic eukaryotes, rather than multiple inde-
pendent
horizontal transfer events or
an ancestral
relationship between chlamydiae and cyanobacteria. Presum-
ably, the chlamydial symbiotic partner was similar to extant
environmental Protochlamydia. Primary ph
and chlamy
Figure 2 We use the term 'symbiont'
(or 'endosymbiont' in the case of chlamydiae) in the sense of
deBary [31] to include mutualistic, parasitic, and commensal
associations. Such an ancient endosymbiotic association,
similar to those giving rise to mitochondria and plastids,
would allow ample time for intracellular (or endosymbiotic)
gene transfer from the chlamydial endosymbiont to the
nucleus of its eukaryotic hosts (either the ancestor of all pri-
mary photosynthetic eukaryotes or individual plastid-con-
taining lineages such as green plants or red algae) and
occasionally between the chlamydial endosymbiont and the
plastids. Additionally, because the protein products of these
intracellularly transferred genes often target to the original
organelles, the co-existence of chlamydial and cyanobacterial
endosymbionts within the same eukaryotic cell also led to the
targeting of chlamydial gene products into plastids and vice
versa. Primary ph
and chlamy
Figure 2 Primary photosynthetic eukaryotes contain gene copies of both plastidic
and chlamydial origin. Numbers above the branch show bootstrap values
for maximum likelihood and distance analyses, respectively. Asterisks
indicate values lower than 50%. (a) 4-hydroxy-3-methylbut-2-en-1-yl
diphosphate synthase (gcpE). (b) Enoyl-ACP reductase (fabI). Note that in
panel (a) sequences from red algae and glaucophytes are of plastidic origin,
whereas those from green plants, apicomplexans, haptophytes, and
chlorarachniophytes are of chlamydial origin. Also note that that in panel
(b) sequences from green plants, diatoms, chlorarachniophytes, and
apicomplexans form a strongly supported group, whereas cyanobacterial
and red alga Cyanidioschyzon homologs form another group. Colors
represent different phylogenetic affiliations. Genome Biology 2007, 8:R99 Genome Biology 2007, Volume 8, Issue 6, Article R99 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.7 Huang and Gogarten R99.7 http://genomebiology.com/2007/8/6/R99 To answer this question, it should be re-emphasized that all
extant chlamydial species are obligate intracellular bacteria. Therefore, if these chlamydiae-like genes in primary photo-
synthetic eukaryotes resulted from gene transfer from
chlamydiae, then they are probably derived from a chlamy-
dial endosymbiont. It should also be noted that although gene
transfer from organelles to the nucleus occurs frequently in
eukaryotes, the actual scope of transfer might vary among lin-
eages. For example, up to 18% of the nuclear genome was
interpreted as derived from plastids in Arabidopsis, but a
much lower percentage of intracellular gene transfer has been
found in the glaucophyte Cyanophora paradoxa (9.1%) [54]
and in the red alga Cyanidioschyzon merolae (Huang and
Gogarten, unpublished data). Given the relatively smaller
genome of Cyanidioschyzon (4,771 predicted protein-coding
genes) and a lower level of intracellular gene transfer, it
would probably not be possible to identify thousands of genes
of any organellar or endosymbiont origin (mitochondrial,
plastidic, or chlamydial) in our genome analyses. Most
importantly, the retention of transferred genes is often
related to the retention and functionality of the organelles in
eukaryotic cells [41]. Because the protein products of intrac-
ellularly transferred genes often function in the original
organelles, loss of certain biochemical functions or even of the
organelles themselves will certainly lead to the loss of related
transferred genes. For instance, even though thousands of
genes were reportedly transferred from chloroplasts in Arabi-
dopsis [53], the number of such genes is significantly lower in
apicomplexan parasites that harbor a relict, nonphotosyn-
thetic plastid. Counting the numbers: can endosymbiosis be inferred
from 21 genes? The sec-
ond possibility is that independent transfer events occurred
from plastids to the nucleus of green plants, apicomplexans,
other plastid-containing groups, and the ancestor of extant
chlamydiae. In either of the alternative scenarios, a chlamy-
dial endosymbiont and its co-existence with plastids in the
same host cell would provide a favorable intracellular envi-
ronment for transfer between chlamydiae and other
organelles. The hypothesis of an ancient chlamydial endosymbiosis is
consistent with the available data. For example, the gene
encoding ATP/ADP translocase is a key innovation by obli-
gate intracellular bacteria (chlamydiae and rickettsiae) that
live as energy parasites. Instead of making ATP on their own,
these bacterial parasites gain ATP from their host cells and
transport ADP back for recycling. Aside from these obligate
intracellular bacteria, recognizable homologs of the ATP/
ADP translocase gene are only found in the microsporidial
Encephalitozoon (another obligate intracellular parasite) and
photosynthetic eukaryotes, where they provide plastids (the
original cyanobacterial endosymbiont) with the ATP neces-
sary for starch and fatty acid biosynthesis or as an energy sup-
plement for carbon dioxide fixation [58-60]. The common
origin of plastidic and chlamydial ATP/ADP translocases was
confirmed by all available phylogenetic analyses [22,25-28],
and various evolutionary scenarios have been proposed
[13,22,25-28]. However, gene transfer from a chlamydial
endosymbiont to its photosynthetic eukaryotic hosts offers a
more logical and parsimonious explanation (also see
Schmitz-Esser and coworkers [27] for related discussions). The phylogenies of gcpE and fabI (Figure 2) also are in agree-
ment with our hypothesis of an ancient chlamydial endosym-
biont in the ancestor of primary photosynthetic eukaryotes. Like the genes encoding 2-C-methyl-D-erythritol 4-phos-
phate cytidylyltransferase (ispD) and 4-diphosphocytidyl-2-
C-methyl-D-erythritol kinase (ispE; Figure 1a,b), gcpE is
related to isoprenoid biosynthesis (see section "Implications
for plastid and eukaryotic evolution" for discussion). The
gcpE sequences from green plants, apicomplexans, hapto-
phytes, and chlorarachniophytes form a strongly supported
monophyletic group with chlamydial (in particular Protoch-
lamydia) homologs. On the other hand, the gcpE sequences
from red algae and glaucophytes form another strongly sup-
ported group with cyanobacterial homologs. These two
groups are not particularly related (Figure 2a) and it is highly
unlikely that the chlamydial gcpE gene was acquired from
cyanobacteria or plastids. Counting the numbers: can endosymbiosis be inferred
from 21 genes? The residence of plastids and mitochondria within eukaryotic
cells led to frequent gene transfers from these organelles to
the nucleus [51-53]. Indeed, it has been reported that thou-
sands of genes were transferred from chloroplasts to the
nucleus in Arabidopsis [53]. Therefore, an apparent question
related to our hypothesis is, can we infer the chlamydial endo-
symbiosis event based on 21 genes? Because chlamydiae are found in diverse eukaryotes such as
acanthamoebae and animals [17,57], it is tempting to specu-
late that the chlamydial endosymbiosis might have existed
before the split of the primary photosynthetic eukaryotes
from other eukaryotic groups. In this study, we searched the
GenBank database, which includes genome sequences of Genome Biology 2007, 8:R99 R99.8 Genome Biology 2007, Volume 8, Issue 6, Article R99 R99.8 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten http://genomebiology.com/2007/8/6/R99 probably resulted from independent losses of the plastidic
(cyanobacterial) gcpE gene copy in these taxa. many early-branching eukaryotes, and the TBestDB, which
covers diverse groups of protists. Chlamydiae-like genes were
found to be restricted mainly to primary photosynthetic
eukaryotes and other plastid-containing lineages, supporting
specifically an association between chlamydiae and the
ancestor of primary photosynthetic eukaryotes. An associa-
tion with an even earlier eukaryote is not supported. FabI is another chlamydiae-related gene, aside from the gene
encoding β-ketoacyl-ACP synthase (fabF; Table 1 and Figure
1c), which is involved in type II fatty acid biosynthesis. Simi-
lar to the gcpE gene phylogeny, cyanobacterial and red algal
Cyanidioschyzon fabI sequences form one group, whereas
homologs from chlamydiae, green plants, apicomplexans,
chlorarachniophytes, and diatoms form another group. These
two very distinct sequence groups differ in several highly con-
served insertions and deletions, but the relationship between
them is less certain because of insufficient internal bootstrap
support on the gene tree. Therefore, although it is more likely
that the two sequence groups in photosynthetic eukaryotes
are derived from cyanobacterial and chlamydial endosymbi-
onts, respectively, it is also theoretically possible that chlamy-
diae acquired their fabI from the plastids of a plant-related
group [63]. Because of the distinct sequence difference of the
two fabI copies, this second scenario entails one of the two
following possibilities. The first is the existence of two fabI
paralogs in the cyanobacterial progenitor of plastids and sub-
sequent independent losses in extant cyanobacteria, and in
red algae and other plastid-containing eukaryotes. Counting the numbers: can endosymbiosis be inferred
from 21 genes? The most plausible explanation for
this observation is that two distinct gcpE gene copies were
originally contributed by chlamydial and cyanobacterial
(plastidic) endosymbionts to the nuclear genome of the
ancestral primary photosynthetic eukaryote and differentially
retained in green plants, red algae, and glaucophytes. The
chlorarachniophyte Bigelowiella and apicomplexan Plasmo-
dium, and the haptophyte Isochrysis are believed to contain a
green and a red algal endosymbiont, respectively [10,61,62];
their proximity in the same chlamydial group (Figure 2a) Implications for plastid and eukaryotic evolution Implications for plastid and eukaryotic evolution
Ancient endosymbionts gave rise to organelles, including
mitochondria, hydrogenosomes, and plastids [1]. Thus far, no
chlamydial endosymbiont has been reported in photosyn-
thetic eukaryotes. Whether a relict organelle derived from the
proposed ancient chlamydial endosymbiont exists in extant
plastid-containing lineages remains to be further investi-
gated. On the other hand, it also would not be surprising if the
chlamydial endosymbiont had degenerated entirely during
the evolution of photosynthetic eukaryotes. As obligate intra-
cellular bacterial parasites, chlamydiae depend on their hosts
for certain nutrients, and consequently they have a relatively
reduced genome [20,21]. Many dispensable genes were prob-
ably lost as a result of their parasitic lifestyle. Given the lack
of apparent benefit to the host cell, such gene losses in an iso-
lated intracellular system could gradually lead to deteriora-
tion of the endosymbiont genome and ultimately the
endosymbiont itself [64]. Almost all chlamydiae-like genes identified in red algae and
green plants (Table 1) contain a predicted plastid-targeting
signal, although the example of CMP-KDO synthetase sug-
gests that this prediction may not always be reliable. The Genome Biology 2007, 8:R99 Genome Biology 2007, Volume 8, Issue 6, Article R99 H Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.9 http://genomebiology.com/2007/8/6/R99 Huang and Gogarten R99.9 In the first stage, a chlamydial bacterium, similar to the
extant Protochlamydia, entered a mitochondrion-containing
eukaryote as a bacterial parasite. This chlamydial endosymbi-
ont possessed a necessary transport system to gain nutrients
and other metabolites from the host cell. At about the same
time, a once free-living photoautotrophic cyanobacterium
was captured by the eukaryotic host by chance, initially possi-
bly as a food source (Figure 3a,b). chlamydiae-like genes are involved in a variety of biochemical
activities in plastids, including fatty acid biosynthesis, ion
transport, nitrogen metabolism, and RNA processing, among
others. Notably, three of these genes (ispD, ispE, and gcpE)
are key enzymes of the deoxyxylulose 5-phosphate (DXP)
pathway, which leads to the formation of isopentenyl diphos-
phate, a major metabolite for isoprenoid biosynthesis in bac-
teria and plastids [65-67]. The chlamydial origin of ispD and
ispE was also confirmed by independent studies [68], which
hypothesized that the DXP pathway in primary photosyn-
thetic eukaryotes was probably derived from plastids. Implications for plastid and eukaryotic evolution In the fourth and final stage, once the plastid organelle was
fully established in the host cell, the benefits of the chlamydial
endosymbiont to the host became less apparent. It is possible
that the chlamydial endosymbiont remained in the host cell
mostly as a bacterial parasite. Such a parasitic relationship
might not be sustained over a long period of time and the
chlamydial endosymbiont might have gradually degenerated. It is also possible that the chlamydial endosymbiont was
transformed into an organelle yet to be recognized in photo-
synthetic eukaryotes (Figure 3e). In the fourth and final stage, once the plastid organelle was
fully established in the host cell, the benefits of the chlamydial
endosymbiont to the host became less apparent. It is possible
that the chlamydial endosymbiont remained in the host cell
mostly as a bacterial parasite. Such a parasitic relationship
might not be sustained over a long period of time and the
chlamydial endosymbiont might have gradually degenerated. It is also possible that the chlamydial endosymbiont was
transformed into an organelle yet to be recognized in photo-
synthetic eukaryotes (Figure 3e). We suggest that three organisms were involved in establish-
ing the primary photosynthetic lineage: the eukaryotic host
cell, the cyanobacterial endosymbiont that provided photo-
synthetic capability, and a chlamydial endosymbiont or para-
site that facilitated the establishment of the cyanobacterial
endosymbiont. The coexistence of three partners with differ-
ent biological requirements and capabilities might have
offered an opportunity for some transient mutualistic inter-
actions, and the acquisition of genes such as those encoding
ATP/ADP translocase and sodium:hydrogen antiporter from
the chlamydial endosymbiont might have facilitated the suc-
cessful endosymbiosis of cyanobacteria by allowing energy
flux into the protoplastid organelle and effective regulation of
ion composition. Specifically, we hypothesize that the origin
and the establishment of primary plastids might have
involved the following stages. Once a rich repertoire of transporters were in place in pri-
mary photosynthetic eukaryotes, transport of photosynthetic
products and other metabolites across the photosynthetic
organelle might be more easily adapted to the different mem-
branes in secondary or tertiary endosymbiotic hosts [70]. This also explains the observation that secondary endosymbi-
osis is more frequent than primary endosymbiosis in eukary-
otic evolution. The likely ancient chlamydial endosymbiosis with primary
photosynthetic eukaryotes has some other important impli-
cations for eukaryotic evolution. Implications for plastid and eukaryotic evolution In our
phylogenetic analyses of ispD and ispE, the chlamydial and
primary photosynthetic eukaryotic sequences form a mono-
phyletic group that is distinct from the cyanobacterial
homologs (Figure 1a,b), suggesting probable gene displace-
ment after the endosymbiotic origin of plastids. In the second stage, gene transfer between the chlamydial
endosymbiont and the host cell ensued because of their phys-
ical association. As a result, the eukaryotic host acquired
transporters from the chlamydial endosymbiont, facilitating
its communication with the cyanobacterial captive. At this
stage, the relationship between chlamydial endosymbiont
and the host cell might be considered transiently mutualistic. Gene transfer between the cyanobacterial captive and the
host cell might also have occurred during this stage (Figure
3c). Secondary endosymbioses between photoautotrophic algae
and heterotrophic host cells occurred several times during
eukaryotic evolution [9-12]. In contrast, the formation of the
primary cyanobacterial endosymbiosis appears to have been
unique (but also see the report by Marin and coworkers [69]). Our finding of the probable participation of a third symbiotic
partner offers an explanation for this rarity. As free-living
photoautotrophic cells, in which ATP is generated in the main
cytoplasmic compartment, cyanobacteria do not need mech-
anisms to transport energy-rich metabolites between mem-
brane-enclosed compartments. However, for an enslaved
cyanobacterium in a heterotrophic host to transform into a
photosynthetic organelle, new transport systems are neces-
sary. Therefore, the initial adaptation of a photoautotrophic
cyanobacterium toward a photosynthetic organelle was prob-
ably a difficult process contingent on the simultaneous pres-
ence of suitable transport systems. At least in part these
transporters might have evolved in a chlamydial parasite that
was present within the same eukaryotic host cell. These trans-
porters enabled chlamydiae to parasitize energy and other
molecules from the host cell, but also allowed for ATP/ADP
equilibration with the cyanobacterium. In the third stage, the cyanobacterial captive was gradually
transformed into a photosynthetic organelle (plastid) in the
host cell and a stable, mutualistic relationship between the
plastid and the host cell was in place. The plastid provided
photosynthetic products to the host, whereas the host offered
shelter and also transported protein products of the intracel-
lularly transferred genes (both from the chlamydial endosym-
biont and the cyanobacterial captive) and other necessary
metabolites to the plastid organelle (Figure 3d). Hypotheti
Figure 3 (d)
? X? X
X
(e) (d) Implications for plastid and eukaryotic evolution Ancient endosymbiosis
events, such as those that gave rise to mitochondria and plas-
tids, to a large degree defined the evolution of eukaryotes. Our
data suggest that these ancient endosymbiosis events might Genome Biology 2007, 8:R99 Huang and Gogarten R99.10 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.10 Genome Biology 2007, Volume 8, Issue 6, Article R99 http://genomebiology.com/2007/8/6/R99 Figure 3
(a)
(b)
? ? (c)
? ? (d)
? X? X
X
(e) have occurred more frequent
been contingent on others. S
subsequent intracellular gen
evolution of host organisms
regardless of whether an org
symbionts is retained in exta
The finding of this study also
primary photosynthetic euk
origin of these groups and
ported by many studies [29,
ticularly those of nuclear gen
conflicting results [71,72]. Th
biosis at the root of the prim
vides strong and independen
of all primary photosyntheti
they harbor. Conclusion
The availability of a comple
alga Cyanidioschyzon and ex
for diverse deep eukaryotes
the distribution and evoluti
primary photosynthetic euka
ogenomic analyses indicate t
are unlikely to have derive
gene transfer events or evo
chlamydiae and cyanobacter
photosynthetic eukaryotes pr
endosymbiosis event between
primary photosynthetic euka
ner being similar to extant e
This ancient chlamydial end
synthetic eukaryotes might
establishment of plastids by p
functions and by allowing eff
h
b
i l
d
b
Figure 3
(a)
(b)
? ? (c)
? ? (d)
? X? X
X
(e)
Hypothetic stages of plastid origin a
Figure 3
Hypothetic stages of plastid origin a
discussed in the text) are displayed
stage; (d) third stage; and (e) fourt
colors show α-proteobacterial (mit
cyanobacterial endosymbionts, as w
respective origins. Arrows directly
symbiotic partner that receives the
indicates the degree of benefit. Das
intracellular gene transfer, whereas
the transferred genes. Crosses indic
transfer processes that might not e
eukaryotes. Note that chlamydial e
parasite in the first stage, but it had
with the host cell in the second and
degenerated in modern photosynth
cyanobacterial endosymbiont was i
host cell (panel b), and then receive
process facilitated by the chlamydia
transformed into a plastid organelle (a) Hypotheti
Figure 3 Hypothetic stages of plastid origin and establishment
Figure 3
Hypothetic stages of plastid origin and establishment. The stages (as
discussed in the text) are displayed as follows: (a,b) first stage; (c) second
stage; (d) third stage; and (e) fourth stage. White, yellow, and green
colors show α-proteobacterial (mitochondrial), chlamydial, and
cyanobacterial endosymbionts, as well as genes and proteins of their
respective origins. Arrows directly from the endosymbiont point to the
symbiotic partner that receives the benefit, and the thickness of the arrow
indicates the degree of benefit. Dashed lines indicate directions of
intracellular gene transfer, whereas solid lines show protein targeting of
the transferred genes. Crosses indicate chlamydial endosymbiont and gene
transfer processes that might not exist in extant photosynthetic
eukaryotes. Note that chlamydial endosymbiont was initially a bacterial
parasite in the first stage, but it had a transient mutualistic relationship
with the host cell in the second and third stages, and then might have
degenerated in modern photosynthetic eukaryotes. Note also that the
cyanobacterial endosymbiont was initially captured to solely benefit the
host cell (panel b), and then received metabolites from the host cell (a
process facilitated by the chlamydial endosymbiont) and was gradually
transformed into a plastid organelle in the host cell (panels d and e). (a) (b) (b) have occurred more frequently, and some of them might have
been contingent on others. Such ancient endosymbioses and
subsequent intracellular gene transfers contributed to the
evolution of host organisms and their descendent lineages,
regardless of whether an organelle derived from past endo-
symbionts is retained in extant species. have occurred more frequently, and some of them might have
been contingent on others. Such ancient endosymbioses and
subsequent intracellular gene transfers contributed to the
evolution of host organisms and their descendent lineages,
regardless of whether an organelle derived from past endo-
symbionts is retained in extant species. ? ? (c) (c) The finding of this study also weighs into the relationship of
primary photosynthetic eukaryotes. Although a common
origin of these groups and of their plastids has been sup-
ported by many studies [29,30], several other analyses, par-
ticularly those of nuclear genes, have provided ambiguous or
conflicting results [71,72]. The ancient chlamydial endosym-
biosis at the root of the primary photosynthetic lineages pro-
vides strong and independent evidence for a common origin
of all primary photosynthetic eukaryotes and of the plastids
they harbor. ? ? (d) ? ? Phylogenomic analyses y
g
y
Phylogenomic analyses of Cyanidioschyzon, which is the only
red algal species whose complete genome sequence is yet
available, were performed using PhyloGenie [75] and the
NCBI nonredundant protein sequence database to identify
chlamydiae-related genes. The results were compared with
those of previously published analyses and further detailed
phylogenetic analyses were performed. For detailed phyloge-
netic analyses, sequences were selected from major groups
within each domain of life. Multiple protein sequence align-
ments were performed using MUSCLE [76] and clustalx [77],
followed by cross-comparisons and manual refinement. Only
unambiguously aligned sequence portions were used. The
alignments are detailed in Additional data file 1. Phylogenetic
analyses were performed with a maximum likelihood method
using PHYML [78] and a distance method using the program
neighbor of PHYLIP version 3.65 [79], with maximum likeli-
hood distances calculated using TREE-PUZZLE [80]. Branch
lengths and topologies of the trees depicted in all figures (Fig-
ures 1 and 2, and Additional data file 1) were calculated with
PHYML [78]. All maximum likelihood calculations were
based on the JTT substitution matrix and a mixed model of
four gamma-distributed rate classes plus invariable sites. Maximum likelihood distances for bootstrap analyses were
calculated using TREE-PUZZLE [80] and PUZZLEBOOT
v1.03 (ME Holder and AJ Roger; available on the web [81]). The plastid-targeting signal of identified chlamydiae-like pro-
tein sequences was predicted using the web-based ChloroP
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h
l b l Co c us o
The availability of a complete genome sequence of the red
alga Cyanidioschyzon and expressed sequence tag (EST) data
for diverse deep eukaryotes allows a more detailed study of
the distribution and evolution of chlamydiae-like genes in
primary photosynthetic eukaryotes. Our very stringent phyl-
ogenomic analyses indicate that these chlamydiae-like genes
are unlikely to have derived from independent horizontal
gene transfer events or evolutionary relatedness between
chlamydiae and cyanobacteria. The chlamydiae-like genes in
photosynthetic eukaryotes probably resulted from an ancient
endosymbiosis event between chlamydiae and the ancestor of
primary photosynthetic eukaryotes, with the chlamydial part-
ner being similar to extant environmental Protochlamydia. This ancient chlamydial endosymbiosis with primary photo-
synthetic eukaryotes might also have played a role in the
establishment of plastids by providing genes that possess new
functions and by allowing effective communications between
the cyanobacterial endosymbiont and the eukaryotic host cell. ? X? X
X
(e) (e) Figure 3 Genome Biology 2007, 8:R99 http://genomebiology.com/2007/8/6/R99 Genome Biology 2007, Volume 8, Issue 6, Article R99 Huang and Gogarten R99.11 Genome Biology 2007, Volume 8, Issue 6, Article R99 H Additional data files The following additional data are available with the online
version of this paper. Additional data file 1 contains protein
sequence alignments used for phylogenetic analyses and
resulting phylogenetic trees. Additi
l Fil 1
P
t i
li
t
d
h l
ti t
Th d
t
t i
t i
li
t
d f
h
l
ti
l
d
lti
h l
ti t
E
h
i
l d
G
B
k GI
b
f ll
d b th
i
Cli k h
f
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https://riunet.upv.es/bitstream/10251/39854/2/Zarzo-sensors-13-00463.pdf
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English
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What is a Fresh Scent in Perfumery? Perceptual Freshness is Correlated with Substantivity
|
Sensors
| 2,012
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cc-by
| 12,105
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What is a Fresh Scent in Perfumery? Perceptual Freshness is
Correlated with Substantivity Manuel Zarzo Sensors 2013, 13, 463-483; doi:10.3390/s130100463 Sensors 2013, 13, 463-483; doi:10.3390/s130100463 sensors
ISSN 1424-8220
www.mdpi.com/journal/sensors
OPEN ACCESS sensors
ISSN 1424-8220
www.mdpi.com/journal/sensors
OPEN ACCESS 1. Introduction Fresh is an odor character descriptor commonly used in perfumery. Actually, it is the one most
frequently encountered in a semantic olfactory database of 119 perfume materials reported by
Thiboud [1]. In a recent study of this database, it was found that warm, oriental and powdery were the
descriptors most dissimilar to fresh (see Figure 6 of [2]). Consistent with this result, fresh and oriental
are regarded as opposite families of perfumes in the Fragrance Wheel proposed by Edwards [3]. The
fresh category comprises citrus, green, water, and fruity subfamilies. The Odor Effects Diagram is an olfactory representation of perfumery notes based on two basic
polarities: (i) erogenous vs. antierogenous (refreshing) and (ii) narcotic vs. stimulating [4]. Citrus,
green, watery, and aldehydic are regarded as refreshing scents, while erogenous, animal, musk, vanilla
and powdery appear at the opposite pole (see Figure 5 of [2]). Fruity is located in this diagram between
floral and fresh, and the same criterion was considered by Edwards [3]. Perfume is a complex mixture of odorants with different volatilities. The parameter that measures
the lasting property of a material when applied on the skin is called substantivity or tenacity. It is well
known by perfumers that olfactory notes perceived as fresh tend to evaporate quickly, while the
opposite applies to those most dissimilar to fresh. Actually, fresh and green are attributes commonly
encountered in the description of top notes (i.e., the ones that are perceived firstly when smelling a
fragrance) [2]. Light refers to scents with high volatility, while heavy, rich or tenacious is applied to
materials with high substantivity. Light fragrances are those perceived as non-sweet with a
predominant fresh note that is often associated with citrus, greens or aldehydes [5]. Conversely, the
least volatile ingredients such as mosses and animal scents dominate in heavy perfumes [1]. Vapor
pressure is the basic factor that determines the volatility of a specific compound [6], but the vapor
composition in equilibrium with the liquid is difficult to predict in mixtures due to the complex
molecular interactions that occur [7–10]. Sensory ratings on a scale of freshness are difficult to obtain because the fresh dimension of
olfactory perception is not well understood yet. Probably for this reason, psychophysical studies aimed
at quantifying the relationship between this odor quality and tenacity have not received much attention
yet. Manuel Zarzo Department of Applied Statistics, Operations Research and Quality, Universidad Politécnica de Valencia,
Cmno. Vera s/n, 46022 Valencia, Spain; E-Mail: mazarcas@eio.upv.es;
Tel.: +34-963-877-490; Fax: +34-963-877-499 Received: 26 November 2012; in revised form: 19 December 2012 / Accepted: 20 December 2012 /
Published: 28 December 2012 Received: 26 November 2012; in revised form: 19 December 2012 / Accepted: 20 December 2012 /
Published: 28 December 2012 Abstract: Perfumes are manufactured by mixing odorous materials with different
volatilities. The parameter that measures the lasting property of a material when applied on
the skin is called substantivity or tenacity. It is well known by perfumers that citrus and
green notes are perceived as fresh and they tend to evaporate quickly, while odors most
dissimilar to ‘fresh’ (e.g., oriental, powdery, erogenic and animalic scents) are tenacious. However, studies aimed at quantifying the relationship between fresh odor quality and
substantivity have not received much attention. In this work, perceptual olfactory ratings
on a fresh scale, estimated in a previous study, were compared with substantivity
parameters and antierogenic ratings from the literature. It was found that the correlation
between fresh odor character and odorant substantivity is quite strong (r = −0.85). ‘Fresh’
is sometimes interpreted in perfumery as ‘cool’ and the opposite of ‘warm’. This
association suggests that odor freshness might be somehow related to temperature. Assuming that odor perception space was shaped throughout evolution in temperate
climates, results reported here are consistent with the hypothesis that ‘fresh’ evokes scents
typically encountered in the cool season, while ‘warm’ would be evoked by odors found in
nature during summer. This hypothesis is rather simplistic but it may provide a new insight
to better understand the perceptual space of scents. Keywords: fragrance; tenacity; odor descriptor; perceptual dimension; olfactory map Sensors 2013, 13 Sensors 2013, 13 464 1. Introduction Several sensory maps of scents reported in the literature are investigated in the present work
attempting to further understand the psychological aspects involved in the perception of refreshing
odor character. The main target of the present work is to study the correlation between this odor
quality and odorant substantivity. This relationship is well established in perfumery, but only at
descriptive level. 2.1. Classification of Odor Descriptors on a Scale of Perceptual Freshness Hedonic tones of odor descriptors are available from the literature, and they can be used to interpret
underlying dimensions of odor profile databases [11]. Similarly, it is possible to assign a refreshing
tone to odor descriptors as described ahead. Boelens and Haring [12] asked a panel of six perfumers to
smell 309 aroma chemicals and to rate on a 0–9 scale the odor similarity to 30 reference materials. Each one was selected as a standard for a certain odor character descriptor. The resulting sensory
dataset, which will be referred to hereafter as B-H database, was analyzed in previous studies using
principal components analysis (PCA) [2,11]. The first principal component (PC1) is the linear
combination of variables (reference materials) that explains the maximum amount of data variability
(17.5% in this case). The contributions of variables in the formation of a given PC are called loadings,
being p1 and p2 the loadings corresponding to PC1 and PC2, respectively. PC1 was interpreted as a
dimension of freshness because fresh was the descriptor with highest p1 loading. The p1 values can be
regarded as (i) indirect assessments of the 30 reference materials on a scale of odor freshness, and (ii)
refreshing tones of the odor descriptors associated to the reference materials. Thiboud [1] reproduces a two-dimensional projection of a similarity matrix developed from
interviews with consumers in England, USA, Spain, Brazil, and Japan relating to defined perfume
bases and to verbal descriptions. The horizontal axis of this fragrance mapping discriminates
masculine vs. feminine scents, and fresh is one of the descriptors that determine the vertical axis. The
same interpretation was proposed for PC2 and PC1 of the B-H database [2]. Thus, a scatterplot of
p2 vs. p1, which is usually referred to as loading plot, was superimposed with Thiboud’s sensory map,
after being properly scaled and rotated in order to achieve the best matching between both odor
representations. Chastrette et al. [13] analyzed semantic odor profiles of 628 pure odorous substances commonly
encountered in perfumery. The database was obtained by Firmenich SA from a team of seven
perfumers who assessed each compound and assigned two to four notes chosen among 32 possible
descriptors. The three most frequent ones were considered as the odor profile. A transformed matrix
derived from this database was analyzed with PCA, and the PC1/PC2 loading plot was proposed as a
sensory map of scents. 2. Methods In order to characterize the connection between perceived fresh odor character and substantivity,
three stages have been carried out: (i) to estimate the freshness of odorant materials and descriptors on
a numeric scale, (ii) to obtain substantivity values from the literature, and (iii) to study the correlation
between them by means of simple linear regression. Each one of these stages is detailed below in a
different section. In the present work, ‘fresh’ and ‘refreshing’ are used as synonyms. The different
connotations of both terms are discussed in Section 4.4. Sensors 2013, 13 Sensors 2013, 13 465 2.1. Classification of Odor Descriptors on a Scale of Perceptual Freshness 2.2. Fresh Odor Quality and Antierogenic Character of Odorant Materials Thiboud [1] compiled semantic odor profiles for 44 natural odorants and 75 synthetic chemicals. Each material was labeled with three or four main odor descriptors and with a set of ancillary attributes
ranging from 0 to 14 (average = 6.6). This database was analyzed in a previous study [2] and it was
found that freshness was an underlying dimension. Based on a few representative descriptors of this
dimension, I obtained estimated scores on a fresh scale for the 44 natural odorants as described below. Jellinek [4] conducted a sensory study attempting to classify perfume materials as erotic or not
erotic. Three experienced perfumers assessed the erogenic character of 61 essential oils, 34 extracts
and 105 aroma chemicals. Each material was added to four perfume compositions at a level that was
just sufficient to create a clearly noticeable difference, and the panel had to decide whether the added
material increased or decreased the erotic character of the perfume. From these sensory ratings, an
antierogenic index (AI) was derived ranging from −100 to 100. The maximum value 100 indicates that
the material was regarded by the panel as clearly antierogenic, while −100 corresponds to a noticeably
erogenic scent. Jellinek [4] considered that antierogenic could also be interpreted as refreshing in odor
description. To study this issue, I checked the correlation between p1 and AI of reference materials in
the B-H database. 2.1. Classification of Odor Descriptors on a Scale of Perceptual Freshness I obtained the projections over the first factorial axis from this plot (Figure 1
of [13]) and compared these projections with p1 loadings of equivalent descriptors in the B-H database. Different sensory maps of scents have been developed by chemical companies that supply perfume
raw materials. One of them is the Rosace of Firmenich [14], which is comprised by 13 categories of
scents properly arranged according to the opinion of professional perfumers. Citrus and balsamic are
located at opposite positions, which suggests that this direction can be interpreted as a dimension of
freshness. I obtained the orthogonal projections of all odor classes over this direction. Next, the
correlation between these projections and p1 loadings from the B-H database was studied. Another olfactory representation is the Field of Odors, which displays in a semicircle different
descriptors and odor categories [15]. It was derived from an odor profile database of perfume
materials. Attempting to study if freshness is a salient dimension of this odor map, I obtained the polar
coordinates for the different odor classes (i.e., angular position in the semicircle and distance to the
center). The angular coordinates were compared with p1 loadings of equivalent attributes in the
B-H database. Sensors 2013, 13 Sensors 2013, 13 466 2.2. Fresh Odor Quality and Antierogenic Character of Odorant Materials 2.4. Relationship between Fresh Odor Character and Substantivity The p1 loadings derived from the B-H database can be regarded as assessments of the 30 reference
materials on a perceptual scale of odor freshness. These values can also be interpreted as refreshing
tones of the odor descriptors associated to reference materials (i.e., the degree of perceptual similarity
or dissimilarity of a given odor quality with respect to the fresh odor character). The correlation
between p1 loadings and the substantivity index of equivalent descriptors was studied by means of
linear regression. The evaporation index and SGS are available for most reference materials in the B-H
database. Thus, the correlation between p1, EI and SGS was also checked. CS values are available for the 44 odorants used by Thiboud [1]. I studied the correlation of CS
with the scores of freshness derived from odor profiles, as well as with AI and other substantivity
parameters. Additional materials commonly used in perfumery were also taken into consideration. 3. Results The comparison of refreshing tones derived from odor profiles and sensory maps yields consistent
results, as indicated in the next section. This issue is of interest to further understand the psychological
issues involved in the perception of fresh odor character. Section 3.2 describes the correlation between
the fresh odor character and substantivity. Sensors 2013, 13 Sensors 2013, 13 467 Another substantivity parameter was reported by Appell [7], who calculated the volatility of 81
essential oils by obtaining the amount of oil evaporated in various time intervals. The quantity (mg)
evaporated in one hour from one gram of oil was called evaporation index. Another substantivity parameter was reported by Appell [7], who calculated the volatility of 81
essential oils by obtaining the amount of oil evaporated in various time intervals. The quantity (mg)
evaporated in one hour from one gram of oil was called evaporation index. The website of The Good Scents Company (www.thegoodscentscompany.com) provides useful
information about suppliers, safety, organoleptic properties, and physical parameters of aroma
chemicals and natural perfume materials. Substantivity values, which will be referred as SGS, are
available for most odorants. This parameter is measured in hours and ranges from 1 to 400 h. The
maximum SGS in fact indicates a censored value higher than 400 (i.e., ≥16.7 days). Actually, certain
materials can smell for months [8]. 2.3. Substantivity Parameters of Perfume Materials After conducting an extensive literature review, I found four substantivity parameters of perfume
materials as described next. These parameters were not obtained in the same way and may not be
accurate, but they are correlated and can be compared with perceptual freshness. Perfumes are described according to their top, middle and base notes, which are also called head,
heart and bottom notes, respectively. The top note is the first odor perceived when smelling a fragrance
product, and it usually consists of the most volatile portion of the composition [1]. Middle notes,
which represent the main body of a blend, are perceived after the top notes fade away [5]. Base notes
are basically determined by fixative materials with a very low volatility and great substantivity that
yield the characteristic lasting note of any fragrance. The three stages of evaporation are often
illustrated with a pyramid [16]. The H&R Fragrance Guide [17] contains the semantic odor description
of 820 commercial perfumes (367 men’s and 453 women’s). I counted the total number of times that a
given attribute was applied to describe top (Ntop), middle (Nmid) and base notes (Nbase). A substantivity
index was calculated according to Equation (1), which takes the value 0, 50 and 100 for attributes that
are only applied to describe top, middle and base notes, respectively. Thus, it provides an estimation of
the substantivity associated to a given descriptor on a 0-100 scale: Substantivity index =
base
mid
top
base
mid
N
N
N
N
N
+
+
⋅
+
⋅
100
50
(1) (1) Poucher [18] obtained the duration of evaporation by olfaction for 332 odorous materials. A certain
quantity of each odorant was placed on a paper strip, and he determined the time that the smelling strip
retained the typical odor note. Based on the results, a coefficient of substantivity (CS) from 1 to 100
was assigned to each material. Sensors 2013, 13 Sensors 2013, 13 468 Regarding the vertical axis of Figure 1, men-husbands appears at the bottom and feminine is found
at the opposite side. This result further supports the hypothesis of a dimension influenced by
psychological and cultural aspects that discriminates feminine vs. masculine cosmetic odors, as
discussed elsewhere [2,19]. Figure 1. Loading plot (p2 vs. p1) of the database obtained by Boelens & Haring [12]
(white triangles). Data were mean-centered and scaled to unit variance prior to the PCA. Thiboud’s fragrance map [1] is superimposed (filled diamonds; labels in italics) after being
properly scaled and rotated (DIM1 and DIM2 stand for dimension 1 and 2, respectively, in
the original publication). Dotted lines group the floral descriptors. Equivalent or related
odor attributes located close to each other are joined with dashed lines. Another sensory map of scents was obtained by Chastrette et al. [13]. Most descriptors in this odor
map have a direct correspondence with attributes in the B-H database. Herbaceous was paired with
vegetable because the reference for the latter smells like herbs [2]. Medicinal was matched with
phenolic because a significant similarity between both descriptors was found in a previous study [22]. Amber was paired with erogenic because a mixture of ambergris and costus oil was the reference
material for erogenic in the B-H database. Figure 2 compares the projection of olfactory notes over the
first factorial axis obtained by Chastrette et al. [13] with p1 loadings of equivalent descriptors in the
B-H database. The correlation is statistically significant (r = 0.86, p < 10−4), which reveals that
freshness is the most salient dimension. Sensors 2013, 13 green
vegetable
minty
medicinal
powdery
jasmine
tuberose
aromatic
clinging
erogenic
aldehyde
citrusy
sourish
night
evening
oriental
fresh
spicy
narcissus
-0.4
-0.3
-0.2
-0.1
0.0
0.1
0.2
0.3
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0.4
sweet
sweet
fruity coconut
feminine
dusty
lemons
fresh
earthy
woody
smoky
spicy
balsamic
animal
honey
buttery
floral
fruity
fatty
watery
lavender
coniferous
metallic
tart (dry)
men-husbands
fern
aftershave
sharp-tangy
camphoraceous-pine
cologne
green
mountains
minty
agrestic-herbal
oily
honey
animal
autumn
carnation
winter
medicinal
woody
aldehydic
fruity raspberry
fruity
young wives-mothers
orris-violet
chypre
fruity apple
lily-of-the-valley
spring
morning
rose
lilac
hyacinth
gardenia
summer
afternoon
musky
anisic
p1
p2
DIM1
DIM2 green
vegetable
minty
medicinal
powdery
jasmine
tuberose
aromatic
clinging
erogenic
aldehyde
citrusy
sourish
night
evening
oriental
fresh
spicy
narcissus
-0.4
-0.3
-0.2
-0.1
0.0
0.1
0.2
0.3
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0.4
sweet
sweet
fruity coconut
feminine
dusty
lemons
fresh
earthy
woody
smoky
spicy
balsamic
animal
honey
buttery
floral
fruity
fatty
watery
lavender
coniferous
metallic
tart (dry)
men-husbands
fern
aftershave
sharp-tangy
camphoraceous-pine
cologne
green
mountains
minty
agrestic-herbal
oily
honey
animal
autumn
carnation
winter
medicinal
woody
aldehydic
fruity raspberry
fruity
young wives-mothers
orris-violet
chypre
fruity apple
lily-of-the-valley
spring
morning
rose
lilac
hyacinth
gardenia
summer
afternoon
musky
anisic
p1
p2
DIM1
DIM2 fruity apple Another sensory map of scents was obtained by Chastrette et al. [13]. Most descriptors in this odor
map have a direct correspondence with attributes in the B-H database. Herbaceous was paired with
vegetable because the reference for the latter smells like herbs [2]. Medicinal was matched with
phenolic because a significant similarity between both descriptors was found in a previous study [22]. Amber was paired with erogenic because a mixture of ambergris and costus oil was the reference
material for erogenic in the B-H database. Figure 2 compares the projection of olfactory notes over the
first factorial axis obtained by Chastrette et al. [13] with p1 loadings of equivalent descriptors in the
B-H database. The correlation is statistically significant (r = 0.86, p < 10−4), which reveals that
freshness is the most salient dimension. Another sensory map of scents was obtained by Chastrette et al. [13]. Most descriptors in this odor
map have a direct correspondence with attributes in the B-H database. Herbaceous was paired with
vegetable because the reference for the latter smells like herbs [2]. 3.1. Freshness as an Underlying Dimension in Odor Maps Figure 1 shows the PC1/PC2 loading plot of the B-H database, superimposed with a rotated
fragrance map [1]. Although the position of comparable descriptors is not exactly coincident, both
sensory maps are strikingly similar. An odor map based on equivalent dimensions has also been
reported [19]. Certain disparity exists in Figure 1 for some attributes (e.g., spicy, aldehydic, honey, or
animal) probably due to the lack of consensus in their interpretation. For example, eugenol and
cinnamon are frequently chosen by perfumers as a reference for spicy [20], but the latter smells
sweeter. Decanal was the reference for aldehydic in the B-H database. It smells antierogenic, but most
aldehydes are perceived as erogenic [4], which would explain the disparity of aldehydic. Honey scents
are often described in perfumery as sweet-medicinal, and Abe et al. [21] found a similarity between
honey and animal, which would clarify the discrepancy of honey in Figure 1. Sensors 2013, 13 Sensors 2013, 13 469 Figure 2. Plot of fitted regression analysis of p[1] (projections on the first factorial axis)
from the Firmenich database analyzed by Chastrette et al. [13] vs. p1 of the B-H database. Empty diamonds correspond to pairs of similar descriptors (e.g., ‘acidic—sourish’ are
assumed to be equivalent attributes in the Firmenich and the B-H databases, respectively). Figure 3. (left) Olfactory representation adapted from the Rosace of Firmenich [14]. (right) Projections of dots over the solid line (Sfreshness) are compared with p1 loadings of
equivalent attributes in the B-H database. -8
-6
-4
-2
0
2
4
6
8
10
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
powdery
balsamic
green
fruity
anisic
honey
minty
fatty
citrus
spicy
pyrogenous - smoky
phenolic - medicinal
animal
woody
amber - erogenic
floral
humus - earthy
piney - coniferous
acidic - sourish
herbaceous - vegetable
p1 (Boelens & Haring, 1981)
p[1] (Chastrette et al., 1991) Figure 3. (left) Olfactory representation adapted from the Rosace of Firmenich [14]. (right) Projections of dots over the solid line (Sfreshness) are compared with p1 loadings of
equivalent attributes in the B-H database. powdery
-2
0
2
4
6
8
10
12
14
16
18
20
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
balsamic
spicy
floral
fruity
earthy
animal
woody
pyrogen. green
herbaceous
aldehydic
citrus
p1 (Boelens & Haring, 1981)
Sfreshness
aldehydic
citrus
herbaceous
green
floral
spicy
fruity
pyrogeneous
animal
balsamic
powdery
woody
moss-earthy Figure 3 shows the 13 odor classes contained in the Rosace of Firmenich [14]. Citrus and aldehydic
are located in opposite positions with respect to balsamic and powdery, which suggests that this
direction (solid line in Figure 3 left) can be regarded as the fresh dimension. Scores of freshness
(Sfreshness) were obtained as the distance between the orthogonal projection of odor classes (dots in the
figure) over the solid line and the right end of this line, measured in a metric arbitrary scale. Most of
these odor classes can be directly matched with attributes in the B-H database. Herbaceous and
powdery
-2
0
2
4
6
8
10
12
14
16
18
20
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
balsamic
spicy
floral
fruity
earthy
animal
woody
pyrogen. Sensors 2013, 13 Medicinal was matched with
phenolic because a significant similarity between both descriptors was found in a previous study [22]. Amber was paired with erogenic because a mixture of ambergris and costus oil was the reference
material for erogenic in the B-H database. Figure 2 compares the projection of olfactory notes over the
first factorial axis obtained by Chastrette et al. [13] with p1 loadings of equivalent descriptors in the
B-H database. The correlation is statistically significant (r = 0.86, p < 10−4), which reveals that
freshness is the most salient dimension. Another sensory map of scents was obtained by Chastrette et al. [13]. Most descriptors in this odor
map have a direct correspondence with attributes in the B-H database. Herbaceous was paired with
vegetable because the reference for the latter smells like herbs [2]. Medicinal was matched with
phenolic because a significant similarity between both descriptors was found in a previous study [22]. Amber was paired with erogenic because a mixture of ambergris and costus oil was the reference
material for erogenic in the B-H database. Figure 2 compares the projection of olfactory notes over the
first factorial axis obtained by Chastrette et al. [13] with p1 loadings of equivalent descriptors in the
B-H database. The correlation is statistically significant (r = 0.86, p < 10−4), which reveals that
freshness is the most salient dimension. Sensors 2013, 13 green
herbaceous
aldehydic
citrus
p1 (Boelens & Haring, 1981)
Sfreshness
aldehydic
citrus
herbaceous
green
floral
spicy
fruity
pyrogeneous
animal
balsamic
powdery
woody
moss-earthy powdery
-2
0
2
4
6
8
10
12
14
16
18
20
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
balsamic
spicy
floral
fruity
earthy
animal
woody
pyrogen. green
herbaceous
aldehydic
citrus
p1 (Boelens & Haring, 1981) herbaceous Figure 3 shows the 13 odor classes contained in the Rosace of Firmenich [14]. Citrus and aldehydic
are located in opposite positions with respect to balsamic and powdery, which suggests that this
direction (solid line in Figure 3 left) can be regarded as the fresh dimension. Scores of freshness
(Sfreshness) were obtained as the distance between the orthogonal projection of odor classes (dots in the
figure) over the solid line and the right end of this line, measured in a metric arbitrary scale. Most of
these odor classes can be directly matched with attributes in the B-H database. Herbaceous and 470 Sensors 2013, 13 pyrogeneous were paired with vegetable and smoky, respectively, based on their odor similarity. If
Sfreshness is compared with p1 loadings from the B-H database, it turns out that the correlation is
statistically significant (r = 0.84, p = 0.0003), which indicates that freshness is an underlying
dimension of this odor map. Interestingly, the orthogonal direction to the solid line in Figure 3 is
defined by floral vs. earthy, which are also located at opposite positions in Figure 1. However, the
position of fruity is arguable taking into account that this descriptor is usually regarded as fresh [3]. The Field of Odors [15] displays different odor classes from citrus in one end to animal at the other,
which suggests that categories are arranged according to perceptual freshness. In order to study this
issue, I compared the angular position (α) of each class with the p1 loading of equivalent attributes in
the B-H database. The descriptors vegetable, earthy, powdery, erogenic, aromatic, coniferous, watery,
and sweet from the B-H database were paired with herbaceous, mossy, musk, amber, vanilla, terpenic,
marine and caramel, respectively, from the Field of Odors, which refer to similar smells. The
correlation between α and p1 is statistically significant (r = 0.82, p < 10−4), which confirms that
freshness is also the most salient dimension in the Field of Odors. Curiously, a similar correlation
coefficient is obtained in Figure 2 (r = 0.86) and Figure 3 (r = 0.84). It is important to keep in mind
that odor descriptors are not always interpreted in the same way, which would partly explain the
residual variability observed in Figures 2–4. For example, eugenol is usually regarded as the reference
material for spicy, but some perfumers may choose cinnamon [20], which smells sweeter. Figure 4. Plot of fitted regression analysis of αJaubert (angular coordinate of odor classes in
the semicircular Field of Odors [15], measured in degrees) vs. p1 loadings of equivalent
descriptors in the B-H database. 3.2. Sensors 2013, 13 smoky—leathery [14,21,24]; powdery—warm [2]. The p1 loadings and substantivity index (SI) of these
descriptors are shown in Table 1. Table 2 displays all reference materials in the B-H database (except
buttery) and indicates p1, AI, SI, SGS, and evaporation index (EI). Strikingly, p1 is tightly correlated
with SI (Table 1: r = −0.81, p < 10−4) and with EI0.5 (Table 2: r = 0.70, p = 0.005). The square root
transformation (i.e., EI0.5) is required to normalize the data. Moreover, AI is also significantly
correlated with p1 (r = 0.75, p < 10−4), SI (r = −0.63, p = 0.001), SGS (r = −0.68, p = 0.0003) and with
EI0.5 (r = 0.72, p = 0.006). Taking into account that (i) antierogenic and refreshing are equivalent
concepts [4], and (ii) p1 loadings can be interpreted as sensory ratings on a scale of freshness, the
observed correlation among p1, AI, SI, SGS, and EI reveals that materials with a low substantivity tend
to be perceived as fresh or refreshing. Table 1. Correspondence between attributes/reference materials in the B-H database and
descriptors in the H&R Fragrance Guide [17]. A substantivity index (SI) is calculated
according to Equation (1) based on the frequency of occurrence of odor descriptors used to
describe the top (NT), middle (NM) and base note (NB) of 820 commercial perfumes. a
H&R Fragrance Guide
H&R Fragrance Guide
Attribute a
p1 Descriptor
NT
NM
NB
SI
Attribute a
p1 Descriptor
NT
NM
NB
SI
bergamot oil
0.341 bergamot
722
0
0
0
floral
−0.010 floral
86 681
47
48
fresh
0.341 fresh
576
63
8
6
honey
−0.064 honey
3
50
38
69
green
0.279 green
240
35
1
7
anisic
−0.081 anise
50
11
0
9
watery
0.260 watery
0
1
0
cedarwood oil
−0.124 cedarwood
0 159
364
85
lemon oil
0.189 lemon
401
1
0
0
woody
−0.124 woody
3 166
305
82
citrusy
0.189 citrusy
57
0
0
0
civet absolute
−0.142 civet
0
0
199
100
galbanum res. Sensors 2013, 13 Relationship between Fresh Odor Character and Substantivity
buttery
20
40
60
80
100
120
140
160
180
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0.4
floral
honey
anisic
spicy
balsamic
musk
mossy
animal
woody
amber
vanilla
sweet
coniferous
lavender
minty
fruity
herbaceous
citrus
aldehyde
marine
green
fresh
p1 (Boelens & Haring)
αJaubert buttery
20
40
60
80
100
120
140
160
180
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0.4
floral
honey
anisic
spicy
balsamic
musk
mossy
animal
woody
amber
vanilla
sweet
coniferous
lavender
minty
fruity
herbaceous
citrus
aldehyde
marine
green
fresh
p1 (Boelens & Haring)
αJaubert 3.2. Relationship between Fresh Odor Character and Substantivity Most reference materials and attributes in the B-H database have a direct correspondence with
descriptors used by the H&R guide (Table 1). Some attributes were paired taking into account their
similarity in perfumery: erogenic—sensual; animal—castoreum [23]; earthy—mossy [21]; 471 a Reference materials (in italics) and descriptors are listed by decreasing order of p1 (loadings in the formation of the first
principal component). The term ‘res.’ stands for resinoid. The correspondence between descriptors and references used by
Boelens & Haring [12] is indicated in Table 2. b Peach is the fruity descriptor most frequently encountered in the H&R
guide. c Pine and fir trees are conifers and their essential oil smells alike. d Peppermint or spearmint. e Mixture of
ambergris and costus oil. f Clove oil contains >85% of eugenol [25]. g Tonka is the second attribute (after vanilla) most
frequently associated to sweet in the H&R guide. h Mixture of musk ketone and coumarin. Sensors 2013, 13 Table 2. Substantivity (SI: substantivity index from Table 1; SGS: substantivity value from
www.thegoodscentscompany.com; EI: evaporation index [7]) and fresh odor character
(p1: loadings in the formation of PC1; AI: antierogenic index [4]) of reference materials
used by Boelens & Haring [12]. Values in bold appear as outliers in Figure 5. Freshness
Substantivity
Reference material
Attribute
p1
AI
SI
SGS
EI
Bergamot oil
Fresh
0.341
23
0
450
Methyl 2-octynoate
Green
0.279
100
7
240
Styrallyl acetate
Sourish
0.277
100
8
Cyclamen aldehyde
Watery
0.260
100
72
Lemon oil
Citrusy
0.189
83
0
4
800 d
Galbanum resinoid
Tart (dry)
0.180
100
3
45
Aldehyde C-10
Aldehyde
0.177
50
0
36
10-undecen-1-ol
Fatty
0.166
60
152
Bay oil
Metallic
0.165
−20
34
364
140
Lavender oil
Lavender
0.128
50
6
12
410
Clary sage oil
Vegetable
0.126
7
21
36
180
Hexadecanal
Fruity
0.085
−100
8
Fir needle oil
Coniferous
0.061
100
82
24
580
Peppermint oil
Minty
0.045
100 a
3
16
300
Oakmoss resinoid
Earthy
−0.008
−35
100
400
Jasmine absolute
Floral
−0.010
−100
50
280
50
Methyl salicylate
Medicinal
−0.052
−15
8
Ethyl phenylacetate
Honey
−0.064
69
312
Fennel oil
Anisic
−0.081
9
24
150
Cedarwood oil
Woody
−0.124
10
85
388
30
Civet absolute
Animal
−0.142
−100
100
400
Cade oil
Smoky
−0.144
−80 b
100
400
10
Ambergris + costus oil
Erogenic
−0.161
−100
100
316c
Eugenol
Spicy
−0.182
−100
50
52
20 e
Patchouli oil
Dusty
−0.197
−100
83
400
4
Vanillin
Aromatic
−0.221
−100
100
400
Olibanum resinoid
Balsamic
−0.249
−100
99
284
Heliotropin
Sweet
−0.280
−23
88
212
Musk ketone+coumarin
Powdery
−0.323
−60
100
400
a Value of spearmint (both plants are botanically related). b Value of birch tar oil (both smell smoky and are
obtained by destructive distillation of wood). c Value of costus oil (SGS of ambergris is not available). d Value
of lime oil. e Value of clove oil. a Value of spearmint (both plants are botanically related). b Value of birch tar oil (both smell smoky and are
obtained by destructive distillation of wood). c Value of costus oil (SGS of ambergris is not available). d Value
of lime oil. e Value of clove oil. In Thiboud’s database, certain material was described with a given attribute as main or ancillary
descriptor if that odor character was clearly recognizable or just noticeable, respectively. Sensors 2013, 13 0.180 galbanum
113
7
0
3
animal
−0.142 castoreum
0
0
83
100
tart (dry)
0.180 dry
19
63
0
38
smoky
−0.144 leathery
0
1
104
100
aldehyde
0.177 aldehydic
140
0
0
0
amber.+costus e
−0.161 ambery
0
0
137
100
bay oil
0.165 bay
5
11
0
34
erogenic
−0.161 sensual
0
0
70
100
lavender oil
0.128 lavender
204
26
0
6
eugenol
−0.182 clove f
0
26
0
50
clary sage oil
0.126 clary sage
83
62
0
21
spicy
−0.182 spicy
163 244
1
30
vegetable
0.126 herbaceous
165
7
2
3
patchouli oil
−0.197 patchouli
0 136
253
83
fruity
0.085 fruity
151
24
3
8
vanillin
−0.221 vanilla
0
0
301
100
fruity
0.085 peach b
149
7
0
2
olibanum res. −0.249 olibanum
0
3
117
99
fir needle oil
0.061 fir c
0
24
41
82
balsamic
−0.249 balsamic
0
0
76
100
coniferous
0.061 pine c
1
87
0
49
sweet
−0.280 sweet
0
50
157
88
peppermint oil
0.045 peppermint d
29
2
0
3
sweet
−0.280 tonka g
0
1
287
100
oakmoss res. −0.008 oakmoss
0
0 139
100
musk+coumarin h
−0.323 musk
0
0
698
100
earthy
−0.008 mossy
0
0 252
100
powdery
−0.323 powdery
0
0
376
100
jasmine
absolute
−0.010 jasmine
0 671
0
50
powdery
−0.323 warm
0
0
137
100 472 Sensors 2013, 13 Sensors 2013, 13 Because of
this, odor descriptors can be coded numerically as 0 if that term is not applied, 1 if it is applied as
ancillary descriptor, and 2 for the main attributes. These coded descriptions are shown in Table 3 for 6
relevant attributes: (i) fresh and citrus, which account for the fresh odor character, (ii) balsamic and
oriental, which are dissimilar to fresh, and (iii) floral and agrestic, with an intermediate refreshing
tone (Figure 1). By adding the values in columns fresh, citrus, floral and agrestic, and subtracting the
columns balsamic and oriental, it results a numeric variable from −3 to 6 that can be interpreted as
sensory scores on a fresh scale (Sfresh in Table 3). Interestingly, Sfresh is significantly correlated with SI
(r = −0.85, p < 10−4), CS (r = −0.77, p < 10−4) and SGS (r = −0.61, p < 10−4). The tight correlation once
again evidences that fresh odor character reflects odorant tenacity. 473 Sensors 2013, 13 Sensors 2013, 13 Figure 5. Relationship between fresh odor character and substantivity of reference
materials used by Boelens and Haring [12]: scatterplot of p1 (loadings in the formation of
PC1) vs. antierogenic index (AI) and substantivity parameters: SI, SGS and EI (values in
Table 2). AI, SGS and EI were conveniently transformed as indicated in the legend to range
approximately on a 0–100 scale. The fitted regression line (r = −0.81) was obtained after
discarding 8 outliers (filled points, highlighted in bold in Table 2). Dashed lines: prediction
limits with a confidence level of 95%. Substantivity and AI values of additional materials not included in Table 3 are shown in Table 4. If
both tables are merged, it turns out that SI is strongly correlated with CS (r = 0.88, p < 10−4) and
SGS (r = 0.83, p < 10−4), but SGS yields a weaker correlation with EI0.5 (r = −0.61, p = 0.0001) probably
because many EI values are missing for the least volatile materials. h
l
ifi d d
(
)
iddl
(
)
d b
0
10
20
30
40
50
60
70
80
90
100
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0.4
100-3.5·EI0.5
SI
SGS/4
(100-AI)/2
p1 (Boelens & Haring, 1981) 0
10
20
30
40
50
60
70
80
90
100
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0.4
100-3.5·EI0.5
SI
SGS/4
(100-AI)/2
p1 (Boelens & Haring, 1981) Substantivity and AI values of additional materials not included in Table 3 are shown in Table 4. If
both tables are merged, it turns out that SI is strongly correlated with CS (r = 0.88, p < 10−4) and
SGS (r = 0.83, p < 10−4), but SGS yields a weaker correlation with EI0.5 (r = −0.61, p = 0.0001) probably
because many EI values are missing for the least volatile materials. Substantivity and AI values of additional materials not included in Table 3 are shown in Table 4. If
both tables are merged, it turns out that SI is strongly correlated with CS (r = 0.88, p < 10−4) and
SGS (r = 0.83, p < 10−4), but SGS yields a weaker correlation with EI0.5 (r = −0.61, p = 0.0001) probably
because many EI values are missing for the least volatile materials. Substantivity and AI values of additional materials not included in Table 3 are shown in Table 4. Sensors 2013, 13 If
both tables are merged, it turns out that SI is strongly correlated with CS (r = 0.88, p < 10−4) and
SGS (r = 0.83, p < 10−4), but SGS yields a weaker correlation with EI0.5 (r = −0.61, p = 0.0001) probably
because many EI values are missing for the least volatile materials. Poucher [18] classified odorants as top notes (CS < 15), middle notes (15 ≤ CS ≤ 60) and base notes
(CS >60). Interestingly, most materials listed in Table 3 up to elemi oil are basically encountered in
top notes (SI << 50) and their CS is below 15. Moreover, materials from vetiver oil to tolu can be
regarded as base notes (SI ≈ 100) and they exhibit a high substantivity (CS >> 60). Thus, Poucher’s
classification based on CS values seems adequate. Some materials with a low CS (top notes) like
mimosa absolute, narcissus absolute or copaiba present a high substantivity according to The Good
Scents Company (SGS = 400), which suggests that such particular values might not be reliable. Actually, the website does not indicate how this parameter was obtained. Moreover, 5 out of the 8
outliers highlighted in Figure 5 correspond to SGS values. Sensors 2013, 13 Sensors 2013, 13 Sensors 2013, 13 474 Table 3. Substantivity parameters and fresh odor character of the 44 natural materials
contained in the semantic odor profile database compiled by Thiboud [1]. Table 3. Substantivity parameters and fresh odor character of the 44 natural materials
contained in the semantic odor profile database compiled by Thiboud [1]. contained in the semantic odor profile database compiled by Thiboud [1]. Sensors 2013, 13 Substantivity
Odor Character Descriptors f
Freshness
Material a
CS b
SI c SGS
d
EI e
Ifresh
Icitrus
Iagrest
Ifloral Ibalsam Ioriental
Sfresh
g
AI h
Petitgrain bergamot oil
3
4
28
170
2
2
1
1
0
0
6
100
Bergamot oil
6
0
450
2
2
1
0
0
0
5
23
Jonquil absolute
24
2
0
1
2
0
0
5
−100
Lime oil
2
0
20
800
2
2
0
0
0
0
4
80
Mandarin oil
2
4
8
2
2
0
0
0
0
4
−27
Coriander oil
3
13
8
200
2
0
2
0
0
0
4
−5
Grapefruit oil
6
264
2
2
0
0
0
0
4
Thyme oil
7
39
172
220
2
0
2
0
0
0
4
100
Lemon oil
8
0
4
2
2
0
0
0
0
4
83
Orange Florida oil
11
1
140
970
2
2
0
0
0
0
4
10
Mimosa absolute
14
400
0
0
2
2
0
0
4
−30
Mugwort oil
9
4
16
1
0
2
0
0
0
3
Chamomile oil, Roman
10
112
530
1
0
2
0
0
0
3
−100
Wormwood oil (absinthe)
10
212
500
0
0
2
1
0
0
3
Violet leaf absolute
18
23
400
20
1
0
0
2
0
0
3
0
Cumin oil
4
7
0
0
2
0
0
0
2
−40
Myrtle oil
4
400
0
0
2
2
2
0
2
Rose oil (Bulgarian)
8
50
168
10
1
0
0
1
0
0
2
−80
Galbanum oil
11
3
72
540
1
0
1
0
0
0
2
100
Carrot seed oil
11
96
80
0
0
2
0
0
0
2
−60
Elemi oil
13
20
720
2
1
1
0
2
0
2
100
Clary sage oil
20
21
36
180
0
0
2
0
0
0
2
7
Geranium Bourbon oil
29
50
28
120
1
0
0
1
0
0
2
−70
Tuberose absolute
43
50
304
0
0
0
2
0
0
2
−100
Cardamom oil
30
16
320
0
0
2
0
0
1
1
75
Rose absolute (French)
43
50
168
0
0
0
1
0
0
1
−90
Cedarwood oil Virginia
8
85
388
30
0
0
1
0
0
0
1
10
Narcissus absolute
11
49
400
0
0
0
2
1
0
1
Nutmeg oil
11
52
550
0
0
1
0
0
1
0
−40
Ginger
7
292
150
0
0
0
0
0
0
0
0
Copaiba
6
400
0
0
0
2
2
0
0
Clove bud oil
22
50
188
20
0
0
0
1
0
1
0
−100
Cinnamon leaf oil
22
56
304
30
0
0
1
0
0
1
0
Vetiver oil Bourbon
100
86
400
0
0
0
0
0
0
0
100
Patchouli oil
100
83
400
4
0
0
1
0
0
1
0
−100
Cistus oil (labdanum)
100
100
400
0
0
2
0
2
0
0
−80
Styrax
100
98
400
0
0
0
2
2
0
0
−100
Pepper (black) oil
100
48
400
0
0
0
0
0
1
−1
Opoponax
90
100
400
20
0
1
0
0
2
1
−2
10
Allspice (pimento berry)
100
400
50
0
0
0
0
2
0
−2
−40
Sandalwood oil
100
88
400
0
0
0
0
2
1
−3
−63
Peru
100
400
0
0
0
0
2
1
−3
−100
Benjoin
100
100
400
0
0
0
0
2
1
−3
−100
Tolu
100
100
400
0
0
0
0
2
1
−3
−100
a Sorted by decreasing value of Sfresh. Sensors 2013, 13 b Coefficient of substantivity [18]. c Substantivity index deduced from
the H&R Fragrance Guide according to Equation (1). d Substantivity (hours) according to
www.thegoodscentscompany.com. e Evaporation index [7]. f Indicator variables corresponding to 6 relevant
descriptors. The value 2 indicates a main descriptor in the semantic odor profile, and the value 1 indicates an
ancillary descriptor. g Scores of freshness calculated as: Ifresh + Icitrus + Iagrestic + Ifloral − Ibalsamic − Ioriental. h Antierogenic index [4]. a Sorted by decreasing value of Sfresh. b Coefficient of substantivity [18]. c Substantivity index deduced from
the H&R Fragrance Guide according to Equation (1). d Substantivity (hours) according to
www.thegoodscentscompany.com. e Evaporation index [7]. f Indicator variables corresponding to 6 relevant
descriptors. The value 2 indicates a main descriptor in the semantic odor profile, and the value 1 indicates an
ancillary descriptor. g Scores of freshness calculated as: Ifresh + Icitrus + Iagrestic + Ifloral − Ibalsamic − Ioriental. h Antierogenic index [4]. Sensors 2013, 13 Sensors 2013, 13 475 Table 4. Substantivity parameters and antierogenic index (AI) of materials not included in
Table 3 (sorted approximately by increasing order of substantivity). Substantivity a
Substantivity a
Material
CS
SI SGS
EI
AI
Material
CS
SI SGS
EI
AI
Lavender oil
4
6
12 410
50
Cascarilla oil
29
248 200
0
Neroli oil
5
4 116 210
17
Orange flower absolute
31
15 400
−50
Laurel leaf oil
9
750
Cinnamon bark oil
24
56 372 120 −100
Peppermint oil
8
3
16 300
100
Lily of the valley; orchid
50
Rosewood oil
2
16
12 250
−8
Carnation absolute
50
−30
Hyacinth absolute
11
16
40
Ylang-ylang oil
32
50 140
90
−80
Basil oil
14
6
225 −50
Ambrette seed oil
30
121
20 −100
Palmarosa oil
14
60
50
0
Orris resinoid
40 b
50
−60
Juniper berry oil
18
700
Jasmine absolute
43
50 280
50 −100
Gardenia
14
Heliotrope
70
Rosemary oil
21
5
4 820
100
Civet absolute
79 100 400
−100
Marjoram oil
18
12 600
Tonka resinoid
100 100 400
−55
Verbena resinoid
19
0
Castoreum absolute
100 100 400
−100
a CS: coefficient of subst.; SI: subst. index; SGS: subst. from www.thegoodscentscompany.com;
EI: evaporation index. b Average CS of orris concrete, orris absolute and orris oleo-resin. Sensors 2013, 13 Most materials in Table 2 with p1 > 0 smell antierogenic (AI > 0) and, as a result, the correlation
between p1 and AI is statistically significant (r = 0.75, p < 10−4). In Table 3, AI is positively correlated
with Sfresh (r = 0.42, p = 0.01). By merging Tables 3 and 4, it turns out that AI is also correlated with SI
(r = −0.60, p < 10−4), SGS (r = −0.50, p = 0.0004) and CS (r = −0.45, p = 0.0006). Given the
relationship between substantivity and freshness, the observed correlations support Jellinek’s
interpretation of antierogenous and refreshing as equivalent concepts in odor description. Nonetheless,
most floral odors were regarded by Jellinek [4] as erogenic (AI ≤ −50), which is somewhat unexpected
because floral scents present an intermediate refreshing tone (Figure 1) and substantivity (SI ≈ 50 in
Table 4). The reason might be that the three perfumers who obtained AI values were probably men. Floral scents, which are perceived as feminine, may be regarded by men as erogenic (i.e., arousing
sexual desire for the opposite sex). Thus, a larger panel should have been used with the same number
of male and female individuals. Actually, Jellinek [4] recognized that his results were preliminary
because of the difficulties in reaching an agreement among observers. 4.2. Proposed Hypothesis to Explain the High Substantivity of Erogenic Odors According to Jellinek [4], erogenous scents are those arousing sexual desire. They are basically
reminiscent of the human body smell (i.e., odors released by our skin) and somewhat resemble the
smell of materials obtained from animals (e.g., civet, castoreum, ambergris or Tonkin musk). This
similarity between erogenic and animal is apparent in the B-H database because both descriptors are
correlated (r = 0.46) and determine an independent dimension of odor character (see Figure 3 of [2]). Body odorants are obviously easily retained by the skin and, hence, they present a high substantivity. The same applies to animalic materials. This reasoning is consistent with the observed negative
correlation between antierogenic odor character and substantivity parameters (Tables 2–4). Taking into account that substantivity depends on physicochemical properties, the tight correlation
of substantivity with freshness implies that, in the case of aroma chemicals, their refreshing odor
character could be estimated by means of electronic noses [32] and also based on physical properties. In the B-H database, the projections of odorants over PC1 can be interpreted as estimated scores of
freshness on a numeric scale. Work in progress suggests that these scores can be predicted from a set
of 20 physicochemical parameters and molecular descriptors using multiple linear regression, with a
coefficient of determination R2 ≈ 0.52. This preliminary result further suggests that the correlation of fresh odor character with
substantivity is primarily innate and evolutionarily hardwired. Nonetheless, olfactory perception of
fragrance raw materials can drastically change when applied in a mixture (e.g., a perfume), in a paper
blotter or on the skin, thus possibly changing its substantivity. Further research will be necessary to
study this issue. The development of standard psychophysical procedures to obtain accurate values of
substantivity and scores of freshness for perfumery materials is strongly encouraged. Sensors 2013, 13 476 Sensors 2013, 13 some studies that have discussed several odor maps of perfumery scents [13,28]. Edwards classifies
12% of fragrances in the citrus, green, watery or fruity categories, which are grouped in the fresh
family [2]. But fresh is not considered by most fragrance companies as a main olfactory family of
perfumes (see Table 1 of [29]). This category is also missing in the classification of fragrances
proposed by the French Society of Perfumers [30], Fragrantica, or Osmoz by Firmenich, among many
others. The reason could be the lack of consensus in the interpretation of fresh in odor description, as
discussed below. Many psychophysical studies have reported that pleasantness is the most salient dimension when a
wide range of odors are assessed (for review, see [31]). This dimension did not clearly show up in a
previous study of the B-H database by checking the correlation between loadings and hedonic tones
from the literature [11]. Although the same methodology could have been applied here, there is enough
evidence to support that freshness is the basic construct underlying Figures 2–4. Actually, animal,
mossy or earthy are often described as disagreeable odors, but these descriptors appear close to sweet
or spicy that are regarded as pleasant. 4.1. Freshness as a Salient Dimension of Odor Perception Smell is a complex multidimensional perception difficult to measure and describe [26]. The
underlying constructs of this multivariate space are still poorly understood. A relatively easy procedure
is proposed here to study if freshness is a salient dimension of odor maps, by assigning a refreshing
tone to odor character descriptors. This issue is of interest for developing standard sensory maps of
cosmetic odors. Different works have reported that freshness is a latent dimension in the perceptual space of
fragrances (for review, see [2]). However, this idea is not well established yet in olfactory research. Actually, neither fresh nor refreshing are included in the comprehensive Dravnieks’ list of 146 terms
most commonly used in odor description [27]. Moreover, the fresh dimension was not considered by 4.4. Interpretation of ‘Fresh’ vs. ‘Refreshing’ in Odor Description The Collins Dictionary & Thesaurus [44] considers the following terms as synonyms of refreshing:
new, original, novel, unusual, stimulating, innovative, cooling, invigorating, etc.. Probably based on the
semantic associations, different studies have found that refreshing and invigorating are similar odor
descriptors [19]. Actually, a fresh fragrance is usually considered invigorating, nature inspired,
reminiscent of early morning air or sea breeze and it is typified by green, citrus notes [1]. Chrea et al. [45]
found that refreshed and revitalized were interpreted in the same way by French speakers, and both
terms were classified in an emotional factor of odor perception called energizing/refreshing. The same
association was found in a further study carried out in Liverpool (UK) and Singapore, and the
underlying factor was named energy [42]. Refreshing and stimulating are regarded as orthogonal
dimensions of the Odor Effects Diagram [4], but this issue is arguable. In the odor map proposed by
Tisserand [46], vivacity and stimulant appear next to erogenic, which is also debatable based on the
reported empirical evidence. Given the multiple meanings of refreshing, this term might be interpreted differently according to
culture, language, context, age, and experience. This issue may explain why Jellinek [4] regarded
refreshing and fresh as different odor descriptors. The former was associated with antierogenous and
the latter with green–herbaceous odors, which were supposed to produce antierogenous and
stimulating effects. However, this criterion is arguable given the semantic association between
refreshing and stimulating. Moreover, odors considered by Jellinek [4] as stimulating are better
described as masculine [2]. Although fresh and refreshing may have different connotations in odor
description according to language, their different meaning is not clearly established in perfumery and,
hence, in my opinion both terms can be used as synonyms for clarity purposes. Sensors 2013, 13 477 Sensors 2013, 13 The human fetus starts the learning process of smelling in the prenatal environment [38]. Flavors from
the mother’s diet during pregnancy are transmitted to amniotic fluid and swallowed by the fetus [39],
which may provide the foundation for cultural and ethnic differences in odor preference. The human fetus starts the learning process of smelling in the prenatal environment [38]. Flavors from
the mother’s diet during pregnancy are transmitted to amniotic fluid and swallowed by the fetus [39],
which may provide the foundation for cultural and ethnic differences in odor preference. Despite the general belief that odor perception is basically shaped by learned associations, some
studies reveal that certain aspects of olfactory perception might be innate and hardwired. For example,
rats show an unconditioned defensive response to the odor of predators [40]. Moreover, recent works
have revealed that the hedonic odor character is partly determined by molecular structure [31]. Interestingly, freshness is the most salient perceptual dimension of cosmetic scents and it is also
somewhat encoded by molecular structure given the correlation with substantivity. Although many
efforts have been carried out to relate molecular structure to the perceived smell [41], odor-structure
relationships focused on the fresh olfactory quality have not been reported yet as far as I know. Ferdenzi et al. [42] found that the refreshing/energizing dimension of smell was common to three
tested cultures, but other constructs were different. The neurological basis of innate and learned
responses to odors has been investigated in mice [43]. 4.4. Interpretation of ‘Fresh’ vs. ‘Refreshing’ in Odor Description 4.3. Conditioned and Unconditioned Factors Influencing Odor Quality Perception It is well established that odor quality perception is strongly influenced by memory [33], learning [34]
and culture [35,36]. Social effects affecting the perception of fragrances have also been discussed [37]. Sensors 2013, 13 478 Sensors 2013, 13 Sensors 2013, 13 dimension underlies consumers’ perceptions as to whether a fragrance seems to be more suited for
formal evening wear or for informal daytime wear. People feel more comfortable wearing during the
day fragrances that do not transmit an erotic message (i.e., smelling antierogenic), which seems quite
obvious. Thus, refreshing scents are preferred for daytime wear because they are not perceived as
erotic, and daytime is associated with stimulating activities, energy, vigorous actions, etc. Although fresh was long ago proposed as an independent category of odors [47], this particular
odor quality is rather subjective and it can be interpreted differently. According to Müller [24], fresh is
generally associated in European regions with lemon, lavender, green notes and light floral
components. Different studies support this interpretation [2]. For example, a so-called light dimension
determined by the descriptors fresh and lemon was found by analyzing numeric odor profiles of 40
compounds rated by French individuals [48]. However, Müller states that sweet and powdery perfumes
are also considered as fresh in North America [24]. Taking into account that sweet and powdery are
perceived as dissimilar to fresh [2], it is of interest to discuss this controversy. Fresh can be defined in English as (i) newly made or obtained, novel, original, additional, (ii) not
stale or deteriorated, (iii) not canned, frozen or preserved, etc. [49]. Consistent with this definition,
Schiffman et al. [50] regarded fresh and rotten as two polarities of the same semantic differential scale. By contrast, the first meaning of fresh in French (frais) and Spanish (fresco) is ‘moderately
cool’ [51,52], which is antonym of warm. The semantic association between fresh and cool also exists
in English, but only when applied to weather, wind or breeze. Thus, fresh air means clean and cool,
found outside buildings rather than in a room [49]. In French, the similarity between refreshed, clean
and cooling in odor perception has also been reported [45]. I believe that fresh was used originally in the sense of cool, probably long ago by French perfumers. Unfortunately, fresh and cool cannot be regarded nowadays as synonyms because cool has acquired
many connotations in perfumery [1]. Cool is often applied to the perception of freshness associated
with a trigeminal effect, as it is the case of minty scents. Actually, minty yields the highest correlation
with ‘cool, cooling’ (r = 0.82) in the database of Dravnieks [27]. Sensors 2013, 13 Moreover, Harper [53] selected two
odorants sharing a minty–fresh odor character (menthol and camphor) as references for ‘cool, cooling’. In order to avoid confusion, I would recommend using fresh, refreshing and antierogenic as
synonyms, referred to scents perceived as cooling and not associated with a trigeminal effect. 4.5. Controversies in the Interpretation of ‘Fresh’ in Odor Description The association between refreshing and energizing might correspond to the activation dimension of
emotion (arousal) [45], but another interpretation is possible. According to Jellinek [29], the refreshing Sensors 2013, 13 Sensors 2013, 13 Sfresh (Table 3). In the B-H database, the negative correlation between fresh and erogenic (r = −0.24,
p < 10−4) partly supports Jellineks’ interpretation, but fresh yields the most negative correlation with
powdery (r = −0.58). The reference material for this descriptor smells warm [2], which suggests that
the perceptual dimension of freshness is better interpreted as a fresh vs. warm polarity rather than fresh
vs. erogenous. This interpretation is supported by further evidence from the literature. Warm perfumes have a high
proportion of animalic ingredients [24] and are often described as rich and tenacious [1]. Harper [53]
selected costus and amber as standards for warm. Both materials were regarded by Boelens & Haring [12]
as references for erogenic, which implies that warm, erogenic and animal are similar descriptors in
perfumery. Actually, in a two-dimensional odor map recently reported, warm and sexy appeared close
to each other [45]. The origin of this similarity from an evolutionary perspective is not well understood
yet, but some hypotheses have been proposed [54]. In a reported analysis of the language of French perfume advertising, it was found that the
dimension warm vs. fresh was the most important axis of the semantic field of fragrances [55]. Chrea et al. [45] obtained a two-dimensional odor map from numeric ratings of 24 odorant samples
according to 73 affective terms in French, and it was found that refreshed and warm appeared in
opposite positions within the pleasant cluster of terms. Jellinek [29] developed a two-dimensional map
of fragrances and one of the dimensions was interpreted as warm vs. cool. In a previous study, 140
commercial perfumes were also classified according to a cool vs. warm dimension [56]. Given that
cool and warm are terms referring to temperature, these studies support the interpretation of fresh and
warm as opposite polarities of a perceptual odor dimension somehow associated with temperature. Interestingly, colors can also be described on a warm vs. cool scale [57]. 4.6. Fresh and Warm as Opposite Polarities of a Perceptual Odor Dimension Based on the semantic meaning, French and Spanish people would easily agree that fresh and warm
are two opposite poles of the same perceptual odor dimension, because they will readily associate fresh
with cool. The same link probably exists in Japanese, because a fresh dimension that discriminated
cold and sour with respect to warm, sweet and sexy was found by Japanese researchers who analyzed
with PCA numeric odor profiles of 37 aroma chemicals according to 55 descriptors [32]. Conversely,
in English, not everybody would understand the dissimilarity fresh vs. warm unless it is indicated that
fresh is used as a synonym of cool. The lack of consensus in the semantic interpretation of freshness
might explain why this perception is not easy to define in perfumery [1,24]. Refreshing and erogenous were regarded by Jellinek [4] as opposite polarities of the same
dimension. Consistent with this criterion, AI values are positively correlated with p1 (Table 2) and 479 Sensors 2013, 13 480 The olfactory map shown in Figure 1 might reflect an underlying structure of odor perception that
discriminates those scents most typically encountered in the different seasons in temperate climates:
fresh (winter), floral (spring), warm (summer), and earthy (autumn). The position of summer next to
fruity in Figure 1 might suggest that summer scents are perceived as fresh, which is misleading. The
correct interpretation is that fresh scents are preferred in summer, probably because they evoke cool
temperatures. Conversely, warm and animalic fragrances are preferred in winter [29]. The work of Tisserand [46] leads to the hypothesis that human olfactory perception was shaped
throughout evolution in temperate climates to recognize the characteristic odors of each season. This
theory is debatable because our sense of smell did not evolve under a constant climate. Moreover, it is
uncertain if the fresh odor character is perceived differently by human races that developed in climates
with slight weather variations along the year, or by people living in such climates. The proposed theory
seems very simplistic given the complexity of factors affecting olfactory perception, but it deserves to
be further investigated because it might provide the fundamental basis to better understand certain
psychological aspects of olfactory perception. For example, it is still poorly understood to what extent
the perception of freshness depends on culture, language, context, age, etc. It would also be of interest
to study why sweet fragrances are preferred by women, or why cosmetic odors are perceived as
masculine vs. feminine. 5. Conclusions Odor freshness is an underlying dimension in the perceptual space of perfumery scents, as further
studied here. Fresh odors are typical in top notes, while those most dissimilar to fresh are encountered
in base notes. This relationship between freshness and substantivity is well known by perfumers, but it
has not received much attention yet by the scientific community. The present work reports that the
correlation between perceptual freshness and odorant substantivity is quite strong (p1 vs. SI in Table 1:
r = −0.82; Sfresh vs. SI in Table 3: r = −0.85). This result is important because substantivity depends on
physicochemical properties, which provides the fundamental basis to predict odor freshness based on
molecular structure. The interpretation of fresh in perfumery as synonym of cool and antonym of warm suggests that
this psychological dimension might be associated with temperature and may reflect a structure of odor
perception shaped throughout evolution in temperate climates. Thus, fresh would correspond to scents
typically encountered in the cool season, while warm would be evoked by odors found in nature
during summer. 3.
Edwards, M. Fragrances of the World: Parfums du Monde 2010, 26th ed.; Fragrances of the
World: Sidney, BC, Canada, 2010; pp. 6–8. 4.7. Proposed Hypotheses to Explain the Fresh/Warm Dimension of Odor Character Taking into account the alternation of seasons in temperate climates, with cool temperatures in
winter and warmer conditions in summer, the ability of human olfaction to perceive scents as fresh vs. warm might be explained from an evolutionary perspective. In summer, as a result of the hot weather,
only those odorants with higher substantivity would be found in the natural environment, while light
scents could be encountered at cooler temperatures. This hypothesis is consistent with the fact that
fresh scents are characterized by a lower substantivity, while the opposite applies to warm odors. The idea that olfactory perceptual space in humans might reflect chronobiological annual rhythms
was put forward by Tisserand [46]. Starting from the four elements (air, fire, earth and water) long ago
proposed by Greek philosophers like Hippocrates, he discussed their associations with the four seasons
and with moods evoked by essential oils. It was found that the resulting structure had certain
resemblance to the Odor Effects Diagram [4]. This diagram has been recently discussed in detail [2]. Tisserand suggested that erogenic scents evoked odors typically encountered in summer, while those
associated with winter such as watery were regarded as antierogenic. Autumn was associated with
earthy, which is intuitively appealing because this descriptor has a neutral refreshing tone (p1 ≈ 0). Moreover, earthy is the attribute most dissimilar to floral (Figure 1), and the latter would correspond to
the typical smells of spring. 2.
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(http://creativecommons.org/licenses/by/3.0/).
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Modelling of tropospheric NO2 using WRF-Chem with optimized
temporal NOx emission profiles derived from in-situ observations -
Comparisons to in-situ, satellite, and MAX-DOAS observations over
central Europe For stations within Germany, biases of −5.9 % to +50.3 % were obtained when comparing average noontime NO2
concentrations at different altitudes. Outside of Germany, where lower resolution emission data was used, biases of up to +78.6 % were observed. Overall, the study demonstrates that temporal modulation of emission data is crucial for modelling
15
tropospheric NO2 realistically. Modelling of tropospheric NO2 using WRF-Chem with optimized
temporal NOx emission profiles derived from in-situ observations -
Comparisons to in-situ, satellite, and MAX-DOAS observations over
central Europe Leon Kuhn1, 2, Steffen Beirle2, Vinod Kumar2, Sergey Osipov2, 4, Andrea Pozzer2, Tim Bösch5,
Rajesh Kumar3, and Thomas Wagner1, 2 Rajesh Kumar3, and Thomas Wagner1, 2
1Institute for Environmental Physics, University of Heidelberg, Germany
2Max Planck Institute for Chemistry, Mainz, Germany
3National Center for Atmospheric Research, Boulder, United States of America
4Red Sea Research Center, King Abdullah University of Science and Technology, Thuwal, Saudi Arabia
5Institute for Environmental Physics, University of Bremen, Germany 1Institute for Environmental Physics, University of Heidelberg, Germany
2Max Planck Institute for Chemistry, Mainz, Germany
3National Center for Atmospheric Research, Boulder, United States of America
4Red Sea Research Center, King Abdullah University of Science and Technology, Thuwal, Saudi Arabia
5Institute for Environmental Physics, University of Bremen, Germany 1Institute for Environmental Physics, University of Heidelberg, Germany
2Max Planck Institute for Chemistry, Mainz, Germany
3National Center for Atmospheric Research, Boulder, United States of America
4Red Sea Research Center, King Abdullah University of Science and Technology, Thuwal, Saudi Arabia
5Institute for Environmental Physics, University of Bremen, Germany Correspondence: Leon Kuhn (l.kuhn@mpic.de) Abstract. We present a WRF-Chem simulation over central Europe with a high spatial resolution of 3 km×3 km and a focus on
nitrogen dioxide (NO2). A regional emission inventory, issued by the German Environmental Agency, with a spatial resolution
of 1 km×1 km is used. We demonstrate, that by precise temporal modulation of the emission data (use of "temporal profiles"),
significant improvement in model accuracy over existing simulations is achieved. Simulated NO2 surface concentrations are compared to measurements from a total of 275 in-situ measurement stations in Germany, where the model was able to reproduce
5
average noontime NO2 concentrations with a bias of +0.9 % and R = 0.76. A comparison between modelled NO2 vertical
column densities (VCDs) and satellite observations from TROPOMI (TROPOspheric Monitoring Instrument) is conducted,
where crucial aspects of the observation process, such as altitude-dependent NO2 sensitivity as well as the influence of clouds
and a priori assumptions of the retrieval, are taken into account. Simulations and satellite observations are shown to agree with a 5 10 model bias of −6.6 % and R = 0.84 for monthly means. Lastly, simulated NO2 concentration profiles are compared to profiles
10
obtained from Multiaxis Differential Optical Absorption Spectroscopy (MAX-DOAS) measurements of five European ground
stations using the profile retrieval algorithms from the Mexican MAX-DOAS fit (MMF) and the Mainz Profile Algorithm
(MAPA). https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Modelling of regional atmospheric chemistry and transport (RCT) is an important discipline in the field of air quality research.
While observational data is often only available at coarse spatial or temporal resolutions, model data can be generated on regular 1
Introduction Modelling of regional atmospheric chemistry and transport (RCT) is an important discipline in the field of air quality research. While observational data is often only available at coarse spatial or temporal resolutions, model data can be generated on regular 1 1 (2015); Chowdhury et al. 30
(2021)). Monitoring and predicting realistic distributions of NO2 in the troposphere is therefore of ongoing political and
scientific interest. Past modelling efforts with focus on tropospheric NO2 have typically resulted in very similar general outcomes: while
researchers found good agreement between modelled and observed meteorological data (such as wind speeds and air temper- been widely recognized among the scientific community (see e.g. Faustini et al. (2014); Mills et al. (2015); Chowdhury et al. 30
(2021)). Monitoring and predicting realistic distributions of NO2 in the troposphere is therefore of ongoing political and
scientific interest. Past modelling efforts with focus on tropospheric NO2 have typically resulted in very similar general outcomes: while
researchers found good agreement between modelled and observed meteorological data (such as wind speeds and air temper- been widely recognized among the scientific community (see e.g. Faustini et al. (2014); Mills et al. (2015); Chowdhury et al. 30
(2021)). Monitoring and predicting realistic distributions of NO2 in the troposphere is therefore of ongoing political and
scientific interest. Past modelling efforts with focus on tropospheric NO2 have typically resulted in very similar general outcomes: while
researchers found good agreement between modelled and observed meteorological data (such as wind speeds and air temper- ature), systematic disagreements between modelled surface NO2 concentrations and in-situ observations from ground based
35
measuring stations were found. Visser et al. (2019) report on the results of a WRF-Chem simulation over central Europe in
which noontime NO2 surface concentrations and VCDs are underestimated by 38.5 % and approximately 15 %, respectively. In a comparison of monthly mean NO2 VCDs, R-values between 0.82 and 0.92 were obtained. The authors identify an un-
derestimation of soil emissions in their emission inventory (TNO-MACC-III, short for "Monitoring Atmospheric Composition ature), systematic disagreements between modelled surface NO2 concentrations and in-situ observations from ground based
35
measuring stations were found. Visser et al. (2019) report on the results of a WRF-Chem simulation over central Europe in
which noontime NO2 surface concentrations and VCDs are underestimated by 38.5 % and approximately 15 %, respectively. In a comparison of monthly mean NO2 VCDs, R-values between 0.82 and 0.92 were obtained. The authors identify an un-
derestimation of soil emissions in their emission inventory (TNO-MACC-III, short for "Monitoring Atmospheric Composition and Climate" by the Netherlands Organisation for Applied Scientific Research) as a possible explanation. Kuik et al. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. grids of much higher spatio-temporal resolution. Modern RCT models can therefore be used to systematically investigate the
20
processes of transport and (photo)-chemical conversion that trace gases are subject to upon emission into the atmosphere. Most importantly, however, they allow for predictions of trace gas concentrations when observational data is not available and
they are also used for operational air quality forecasting. Thus, they give valuable insight into the dynamics of air quality in
polluted regions of the earth. RCT models can be "fully coupled", meaning that they include interactions between chemistry grids of much higher spatio-temporal resolution. Modern RCT models can therefore be used to systematically investigate the
20
processes of transport and (photo)-chemical conversion that trace gases are subject to upon emission into the atmosphere. Most importantly, however, they allow for predictions of trace gas concentrations when observational data is not available and
they are also used for operational air quality forecasting. Thus, they give valuable insight into the dynamics of air quality in
polluted regions of the earth. RCT models can be "fully coupled", meaning that they include interactions between chemistry and meteorology, or "offline", meaning that they simulate chemistry and meteorology separately. Examples for state of the art
25
RCT models are WRF-Chem (Grell et al., 2005), COSMO/MESSy (Kerkweg and Jöckel, 2012), Lotos-Euros (Manders et al.,
2017), CAM-chem (Emmons et al., 2020), and CHIMERE (Menut et al., 2021). Nitrogen dioxide (NO2) is one of the most relevant chemical species for air quality in polluted regions. It is toxic to humans
and acts as a precursor for ozone (O3), a key pollutant of urban smog. The hazardous impact of NO2 on human health has and meteorology, or "offline", meaning that they simulate chemistry and meteorology separately. Examples for state of the art
25
RCT models are WRF-Chem (Grell et al., 2005), COSMO/MESSy (Kerkweg and Jöckel, 2012), Lotos-Euros (Manders et al.,
2017), CAM-chem (Emmons et al., 2020), and CHIMERE (Menut et al., 2021). Nitrogen dioxide (NO2) is one of the most relevant chemical species for air quality in polluted regions. It is toxic to humans
and acts as a precursor for ozone (O3), a key pollutant of urban smog. The hazardous impact of NO2 on human health has been widely recognized among the scientific community (see e.g. Faustini et al. (2014); Mills et al. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. the clear consensus that RCT simulations tend to underestimate tropospheric NO2 and that the emission inventories and their
55
preprocessing play a key role in resolving that problem. A number of publications have theorized that an apparent model bias can be explained because groundbased in-situ measure-
ments systematically overestimate NO2 concentrations. Conventional in-situ methods often utilize molybdenum converters,
which were found to be cross sensitive to other reactive nitrogen species, such as NH3 or PAN. This issue was discussed e.g. the clear consensus that RCT simulations tend to underestimate tropospheric NO2 and that the emission inventories and their
55
preprocessing play a key role in resolving that problem. the clear consensus that RCT simulations tend to underestimate tropospheric NO2 and that the emission inventories and their
55
preprocessing play a key role in resolving that problem. A number of publications have theorized that an apparent model bias can be explained because groundbased in-situ measure-
ments systematically overestimate NO2 concentrations. Conventional in-situ methods often utilize molybdenum converters,
which were found to be cross sensitive to other reactive nitrogen species, such as NH3 or PAN. This issue was discussed e.g. the clear consensus that RCT simulations tend to underestimate tropospheric NO2 and that the emission inventories and their
55
preprocessing play a key role in resolving that problem. A number of publications have theorized that an apparent model bias can be explained because groundbased in-situ measure-
ments systematically overestimate NO2 concentrations. Conventional in-situ methods often utilize molybdenum converters,
which were found to be cross sensitive to other reactive nitrogen species, such as NH3 or PAN. This issue was discussed e.g. by Dunlea et al. (2007), Steinbacher et al. (2007), Boersma et al. (2009), and Visser et al. (2019). However, a systematic in-
60
vestigation to prove a general bias of the NO2 in-situ measurements in Europe is still missing. Furthermore, the corresponding
European authority claims that their instruments follow strict regulations (defined by the European Norm EN 14221) and that
the overestimation of NO2 was found to be smaller than 7 % in test measurements (German Environmental Agency (a)). In this paper, we show the results of a WRF-Chem simulation over central Europe for the month of May 2019 with a spatial resolution of 3 km × 3 km. (2016)
40
run a WRF-Chem simulation with the same emission inventory as Visser et al. (2019) over the region of Berlin, Germany,
and observe even stronger underestimations of surface NO2 by more than 50 % during daytime and a strong overestimation
at night-time in a similar comparison. The study reveals that increasing the spatial resolution (including downscaling of the
used emission data) of the model from 15 km × 15 km to 1 km × 1 km slightly improves agreement, but not to a satisfying and Climate" by the Netherlands Organisation for Applied Scientific Research) as a possible explanation. Kuik et al. (2016)
40
run a WRF-Chem simulation with the same emission inventory as Visser et al. (2019) over the region of Berlin, Germany,
and observe even stronger underestimations of surface NO2 by more than 50 % during daytime and a strong overestimation
at night-time in a similar comparison. The study reveals that increasing the spatial resolution (including downscaling of the
used emission data) of the model from 15 km × 15 km to 1 km × 1 km slightly improves agreement, but not to a satisfying degree. In a subsequent publication, the authors attribute the disagreements to underestimations in their emission data (see Kuik
45
et al., 2018). Mar et al. (2016) report that the choice of the chemical mechanism used by WRF-Chem barely impacts the NOx
underestimation by comparing simulations using the RADM2 and MOZART-4 chemical mechanisms. Kumar et al. (2021)
demonstrate in a simulation with the MECO(n) model system over Germany using TNO-MACC-III and regional emission data
from the German Environmental Agency (UBA), that agreement between modelled NO2 concentrations and in-situ observa- tions improves greatly when diurnal and seasonal variability is added to the yearly resolved emission data using reasonably
50
chosen hourly and monthly weighting factors ("temporal profiles"). This temporal upsampling has become common practice
among the air quality modelling community and standard values for such temporal profiles have established (see Crippa et al. (2020b); Kumar et al. (2021) and the references within). Kumar et al. (2021) used improved temporal profiles derived from
empirical proxies, e.g. car counts on highways at different hours of the day. Altogether the contemporary literature comes to 2 CC BY 4.0 License. 30° W
12° E
54° E
62° N
50° N
38° N
26° N
D1
D2
‚
Figure 1. Geographical coverage of the WRF-Chem simulation. The spatial resolutions of the outer domain D1 and the inner domain D2 are
15 km × 15 km, and 3 km × 3 km, respectively. ‚
Figure 1. Geographical coverage of the WRF-Chem simulation. The spatial resolutions of the outer domain D1 and the inner domain D2 are
15 km × 15 km, and 3 km × 3 km, respectively. 2
WRF-Chem simulation setup
85 WRF-Chem (Weather Research and Forecasting model with Chemistry, see Grell et al., 2005) is a mesoscale RCT model. We
run WRF-Chem on a twofold nested domain over central Europe for the month of May, 2019, see Fig. 1. The spatial resolutions
of the outer and inner domain (called D1, and D2 from hereon) are 15 km×15 km, and 3 km×3 km (with 320×245 pixels and
500 × 430 pixels, respectively). The temporal resolution of the simulation is 60 seconds on D1 and 12 seconds on D2. Output files are written daily for D1 and hourly for D2. The simulation uses the Thompson microphysics scheme (see Thompson et al.,
90
2008), the RRTMG (Rapid Radiative Transfer Model for General Circulation Models long- and shortwave radiation scheme,
see Iacono et al., 2008), the Monin-Obukhov similarity scheme for surface layer modelling (see Monin and Obukhov, 1954),
the NOAH Land-Surface Model (see Niu et al., 2011), the YSU boundary layer scheme (see Hong, 2010), and the Grell-
Devenyi ensemble scheme for cumulus modelling (see Grell and Dévényi, 2002). For modelling of chemistry, the MOZART chemical mechanism (see Emmons et al., 2010) is coupled to the GOCART aerosol mechanism (see Chin et al., 2000) along
95
with the TUV full photolysis scheme (Madronich (1987); Tie et al. (2003)), which deploys climatological O3 and O2 columns. Dry deposition is calculated according to Wesely (1989). Spectral nudging (see e.g. Omrani et al., 2012) to ERA5 reanalysis
data (see Hersbach and Dee, 2017) is used. The full list of WRF-Chem simulation options (the "namelist") can be found in
the supplement of this article. The simulation uses a total of 43 vertical levels in terrain-following coordinates (see Table A1 chemical mechanism (see Emmons et al., 2010) is coupled to the GOCART aerosol mechanism (see Chin et al., 2000) along
95
with the TUV full photolysis scheme (Madronich (1987); Tie et al. (2003)), which deploys climatological O3 and O2 columns. Dry deposition is calculated according to Wesely (1989). Spectral nudging (see e.g. Omrani et al., 2012) to ERA5 reanalysis
data (see Hersbach and Dee, 2017) is used. The full list of WRF-Chem simulation options (the "namelist") can be found in
the supplement of this article. The simulation uses a total of 43 vertical levels in terrain-following coordinates (see Table A1 for the layer heights up to 6 km). The goal is to quantify the level of agreement between simulated NO2 concentrations and VCDs
65
and corresponding results from different observational datasets. In particular, we investigate whether the NO2 biases of the
simulation can be resolved by optimization of the temporal profiles. For this purpose our simulation results are compared to
three reference datasets: 1. Surface NO2 concentrations measured by a network of in-situ instruments over Germany, operated by the German
Environmental Agency (UBA, see German Environmental Agency (b)) 2. Tropospheric NO2 vertical column densities (VCDs) measured by TROPOMI (TROPOspheric Monitoring Instrument)
on the Sentinel 5 precursor satellite (specifically the NO2 PAL data product as described in Eskes et al., 2019). This
includes recomputing the air mass factors (AMFs) of the retrieval based on our simulation results. 3. NO2 concentration profiles obtained from five MAX-DOAS instruments that partake in the FRM4DOAS project (see
Fayt et al., 2021) 3. NO2 concentration profiles obtained from five MAX-DOAS instruments that partake in the FRM4DOAS project (see
Fayt et al., 2021) 75 75 We show that when temporal profiles are chosen such that the simulation shows good agreement with in-situ measurements,
the simulation also agrees well with satellite and MAX-DOAS observations. This demonstrates that the total NOx (= NO2 +
NO) budget assumed in the used emission inventories is realistic. As a direct consequence of this finding, improved simulation
strategies like ours could be used for more precise air quality modelling with the focus on NO2, NOx or other species in the future. 80
The paper is structured as follows: Sect. 2 describes the setup of our WRF-Chem simulation and the preprocessing of emis-
sion data in detail. Sect. 3 shows the results of our simulation and compares them to a "classical" simulation with empirically
derived temporal profiles. Furthermore, the simulated data is compared to the above mentioned observational datasets. In sect. 4 the results are discussed and conclusions are drawn. future. 80
The paper is structured as follows: Sect. 2 describes the setup of our WRF-Chem simulation and the preprocessing of emis-
sion data in detail. Sect. 3 shows the results of our simulation and compares them to a "classical" simulation with empirically
derived temporal profiles. Furthermore, the simulated data is compared to the above mentioned observational datasets. In sect. 4 the results are discussed and conclusions are drawn. 3 3 https://doi.org/10.5194/egusphere-2022-1473
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c⃝Author(s) 2023. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. et al., 2020). Additionally, CAM-chem yields the boundary conditions for D1. D2 receives boundary conditions online from
the WRF-Chem simulation running on D1. et al., 2020). Additionally, CAM-chem yields the boundary conditions for D1. D2 receives boundary conditions online from
the WRF-Chem simulation running on D1. et al., 2020). Additionally, CAM-chem yields the boundary conditions for D1. D2 receives boundary conditions online from
the WRF-Chem simulation running on D1. 2.1
Emission preprocessing Outside of Germany the EDGARv5 emission inventory with a moderate spatial resolu-
tion (0.1◦× 0.1◦or ∼11 km × 7 km) is used (see Crippa et al., 2020a). Since UBA-E does not include organic anthropogenic emissions, EDGARv5 is used on the entirety of D1 and D2 for organic species. Non-methane volatile organic compounds
115
(NMVOCs) are provided as lumped species and are speciated according to Huang et al. (2017). An alternative to EDGARv5
would have been the TNO-MAC-III emission inventory (Kuenen et al., 2014) which comes at a higher spatial resolution of
0.0625◦× 0.125◦but was only available for the year 2011 at the time the simulation was run. Since EDGARv5 was available
for 2015 and NOx emissions have steadily decreased over the past years (see e.g. Anenberg et al., 2022), EDGARv5 was considered a more reasonable choice. 120
EDGARv5 (in 2015) and UBA-E have time resolutions of 1 month and 1 year, respectively. However, many emissions follow
strong diurnal and seasonal patterns. For example, emissions from car traffic are expected to be lower at nighttime and higher
at daytime and agricultural emissions typically occur in specific months of the year. EDGARv5 can still resolve emission
variations on a monthly time-resolution but is incapable of resolving diurnal patterns. UBA-E provides only annual emissions, considered a more reasonable choice. 120
EDGARv5 (in 2015) and UBA-E have time resolutions of 1 month and 1 year, respectively. However, many emissions follow
strong diurnal and seasonal patterns. For example, emissions from car traffic are expected to be lower at nighttime and higher
at daytime and agricultural emissions typically occur in specific months of the year. EDGARv5 can still resolve emission
variations on a monthly time-resolution but is incapable of resolving diurnal patterns. UBA-E provides only annual emissions, therefore without any temporal pattern. The solution to this problem is to scale the coarsely resolved emission data to an hourly
125
resolution using presumed hourly, daily, and monthly emission scaling factors, called temporal profiles. The emission rate
EX,k(m,d,h,lat, lon) of a species X from sector k at month m, day d, and hour h at fixed latitude lat and longitude lon is
given as
EX,k(m,d,h,lat, lon) = ˆEX,k(lat, lon) · pmonthly,k(m) · pdaily,k(d) · phourly,k(h)
(1) therefore without any temporal pattern. 2
WRF-Chem simulation setup
85 Both domains D1 and D2 receive initial conditions from the CAM-chem model (Emmons
100 4 2.1
Emission preprocessing The WRF-Chem simulation is driven by different emission inventories for different emission sectors (see also Table 1). Emis-
sions from biomass burning are taken from the Fire Inventory from NCAR (FINN, see Wiedinmyer et al., 2011) with a spatio-
105
temporal resolution of 1 km × 1 km and 24 hours. Biomass burning emissions are assumed to have a diurnal variation with a
peak in the early afternoon (1 pm local time) and are distributed vertically in the model following the plumerise parametrization
of Freitas et al. (2007). Biogenic emissions are computed online using an implementation of the MEGAN model (see Guenther
et al., 2006). For non-organic anthropogenic emissions a combination of two emission inventories is used: Over Germany, an inventory of high spatial resolution (1 km × 1 km, resampled to 0.01◦× 0.01◦or ∼1.1 km × 0.7 km) is provided by the
110
German Environmental Agency (Umweltbundesamt, UBA, see Strogies et al., 2020). From hereon, this emission inventory
will be referenced as UBA-E. Using the UBA-E emission data over Germany enables modelling of NO2 distributions at an
unprecedented high spatial resolution. Outside of Germany the EDGARv5 emission inventory with a moderate spatial resolu-
tion (0.1◦× 0.1◦or ∼11 km × 7 km) is used (see Crippa et al., 2020a). Since UBA-E does not include organic anthropogenic an inventory of high spatial resolution (1 km × 1 km, resampled to 0.01◦× 0.01◦or ∼1.1 km × 0.7 km) is provided by the
110
German Environmental Agency (Umweltbundesamt, UBA, see Strogies et al., 2020). From hereon, this emission inventory
will be referenced as UBA-E. Using the UBA-E emission data over Germany enables modelling of NO2 distributions at an
unprecedented high spatial resolution. Outside of Germany the EDGARv5 emission inventory with a moderate spatial resolu-
tion (0.1◦× 0.1◦or ∼11 km × 7 km) is used (see Crippa et al., 2020a). Since UBA-E does not include organic anthropogenic an inventory of high spatial resolution (1 km × 1 km, resampled to 0.01◦× 0.01◦or ∼1.1 km × 0.7 km) is provided by the
110
German Environmental Agency (Umweltbundesamt, UBA, see Strogies et al., 2020). From hereon, this emission inventory
will be referenced as UBA-E. Using the UBA-E emission data over Germany enables modelling of NO2 distributions at an
unprecedented high spatial resolution. 2.1
Emission preprocessing The solution to this problem is to scale the coarsely resolved emission data to an hourly
125
resolution using presumed hourly, daily, and monthly emission scaling factors, called temporal profiles. The emission rate
EX,k(m,d,h,lat, lon) of a species X from sector k at month m, day d, and hour h at fixed latitude lat and longitude lon is
given as
E
(
d h l
l
)
ˆE
(l
l
)
(
)
(d)
(h)
(1) EX,k(m,d,h,lat, lon) = ˆEX,k(lat, lon) · pmonthly,k(m) · pdaily,k(d) · phourly,k(h)
(1) X,k(lat, lon) · pmonthly,k(m) · pdaily,k(d) · phourly,k(h)
(1) X,k(m,d,h,lat, lon) = ˆEX,k(lat, lon) · pmonthly,k(m) · pdaily,k(d) · phourly,k(h) (1) where ˆEX,k denotes the total emission budget of species X from sector k in the emission inventory and pmonthly, pdaily, and
130
phourly the monthly, daily, and hourly temporal profiles. The individual profiles are normalized to 12 (annual cycle), 7 (weekly
cycle), and 24 (diurnal cycle), respectively. Because different emission sectors follow vastly different temporal patterns, the
temporal profiles are defined for each sector individually. An overview of the emission sectors is given in Table 1. The total where ˆEX,k denotes the total emission budget of species X from sector k in the emission inventory and pmonthly, pdaily, and
130
phourly the monthly, daily, and hourly temporal profiles. The individual profiles are normalized to 12 (annual cycle), 7 (weekly
cycle), and 24 (diurnal cycle), respectively. Because different emission sectors follow vastly different temporal patterns, the
temporal profiles are defined for each sector individually. An overview of the emission sectors is given in Table 1. The total where ˆEX,k denotes the total emission budget of species X from sector k in the emission inventory and pmonthly, pdaily, and
130
phourly the monthly, daily, and hourly temporal profiles. The individual profiles are normalized to 12 (annual cycle), 7 (weekly
cycle), and 24 (diurnal cycle), respectively. Because different emission sectors follow vastly different temporal patterns, the
temporal profiles are defined for each sector individually. An overview of the emission sectors is given in Table 1. The total 5 5 N/A = "not available" N/A = not available
† Relative contribution of this sector to the overall emission budget (yearly for UBA-E, and for the month of May for EDGARv5) of NOx https://doi.org/10.5194/egusphere-2022-1473
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c⃝Author(s) 2023. CC BY 4.0 License. where Xlump is the lump of species that contains species X (i.e. Xlump = NOx when X = NO2). 140
In principle, WRF-Chem also supports vertical distribution of trace gas emissions. This is reasonable to consider, given
that many strong emissions, like those from combustion stacks, take place at elevated altitudes. In analogy to the mentioned
temporal and speciation profiles, this can be modelled using an additional vertical emission profile. Suggestions for vertical
emission profiles are given in Bieser et al. (2011) and Pozzer et al. (2009). In our study we omit the use of vertical profiles and inject all emissions into the lowest model layer (0 - 8 m). This simplification was made seeing that vertical profiles would add
145
another dimension of complexity to our main goal, which is to optimize the temporal profiles. Furthermore, precise information
on the emission height is often not available. Lastly, assuming a well-mixed troposphere, vertical profiles can be expected to
be relevant only in the close vicinity of strong elevated emission sources. inject all emissions into the lowest model layer (0 - 8 m). This simplification was made seeing that vertical profiles would add
145
another dimension of complexity to our main goal, which is to optimize the temporal profiles. Furthermore, precise information
on the emission height is often not available. Lastly, assuming a well-mixed troposphere, vertical profiles can be expected to
be relevant only in the close vicinity of strong elevated emission sources. inject all emissions into the lowest model layer (0 - 8 m). This simplification was made seeing that vertical profiles would add
145
another dimension of complexity to our main goal, which is to optimize the temporal profiles. Furthermore, precise information
on the emission height is often not available. Lastly, assuming a well-mixed troposphere, vertical profiles can be expected to
be relevant only in the close vicinity of strong elevated emission sources. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Table 1. Emission sectors in the UBA-E and EDGARv5 emission inventories
Name
Contribution† (UBA-E) [%]
Contribution† (EDGARv5) [%]
traffic (no resuspension)
43.8
38.7
power industry
18.5
15.0
agricultural soils
10.1
4.5
energy for buildings
7.3
6.6
manufacturing industry
7.2
15.8
non-metallic minerals production
2.5
0.0
production of chemicals
2.4
0.3
shipping
2.1
2.1
iron and steel production
1.8
< 0.1
oil refineries and transformation industry
1.7
1.8
aviation landing and take-off
1.2
1.8
railways, pipelines, and off-road transport
0.9
2.3
production of food, pulp, and paper
0.3
0.3
manure management
0.1
1.1
fuel exploitation
0.1
0.0
solid waste incineration
0.1
0.1
non-ferrous metal production
0
< 0.1
non-energy use of fuels
< 0.1
0.0
agricultural waste burning
N/A
0.3
fossil fuel fires
N/A
< 0.1
aviation climbing and descent
N/A
5.9
aviation cruise
N/A
3.4 Table 1. Emission sectors in the UBA-E and EDGARv5 emission inventories Name
Contribution† (UBA-E) [%]
Contribution† (EDGARv5) [%] N/A = not available
† Relative contribution of this sector to the overall emission budget (yearly for UBA-E, and for the month of May for EDGARv5) of NOx N/A = not available
† Relative contribution of this sector to the overall emission budget (yearly for UBA-E, and for the month of May for EDGARv5) of NOx emission rate of species X is obtained by summation over all emission sectors k emission rate of species X is obtained by summation over all emission sectors k EX(m,d,h,lat, lon) =
X
k
ˆEX,k(lat, lon) · pmonthly,k(m) · pdaily,k(d) · phourly,k(h)
(2)
35 EX(m,d,h,lat, lon) =
X
k
ˆEX,k(lat, lon) · pmonthly,k(m) · pdaily,k(d) · phourly,k(h)
(2)
135 (2) Some species are only implicitly contained in the emission inventories. For example, EDGARv5 and UBA-E specify NOx
emissions, but the partitioning into NO and NO2 must be chosen by the user via a speciation profile pspec(k). Equation (2)
then generalizes to Some species are only implicitly contained in the emission inventories. For example, EDGARv5 and UBA-E specify NOx
emissions, but the partitioning into NO and NO2 must be chosen by the user via a speciation profile pspec(k). Equation (2)
then generalizes to (3) 6 https://doi.org/10.5194/egusphere-2022-1473
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c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2022-1473
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c⃝Author(s) 2023. CC BY 4.0 License. 3. Gradient-based optimization methods depend on hyperparameters, such as the step size. If these are not picked correctly
from the start, the optimization may converge slowly or diverge entirely. This becomes unfeasible, when a single forward
run takes days to compute. 3. Gradient-based optimization methods depend on hyperparameters, such as the step size. If these are not picked correctly
from the start, the optimization may converge slowly or diverge entirely. This becomes unfeasible, when a single forward
run takes days to compute. 3. Gradient-based optimization methods depend on hyperparameters, such as the step size. If these are not picked correctly
from the start, the optimization may converge slowly or diverge entirely. This becomes unfeasible, when a single forward
run takes days to compute. Using a conventional optimization routine, where the gradient of a loss function determines the outcome of a single optimiza-
175
tion step, would require to encode aspects 1-3 in the form of mathematical constraints. This makes a rigorous treatment of the
problem extremely complex. The manual optimization approach, however, does not require a mathematical formulation and
can thus take the discussed aspects into consideration more easily. We run the optimization under the following conditions: 1. Only the most relevant emission sectors are modified during the process. According to Table 1 these are: traffic (no
resuspension), power industry, agricultural soils, energy for buildings, and manufacturing industry. 1. Only the most relevant emission sectors are modified during the process. According to Table 1 these are: traffic (no
resuspension), power industry, agricultural soils, energy for buildings, and manufacturing industry. 2. Speciation of lumped species follows the recommendations of Huang et al. (2017) and is not further optimized. For NOx,
the partitioning is assumed to be 87.5 % NO and 12.5 % NO2. This choice reflects the fact that NOx from combustion
processes is mostly emitted as NO, which mostly oxidizes to NO2 over time. Literature values for typical NO2/NOx
ratios in anthropogenic emissions range from lower values (e.g. 5.3 %, as reported by Wild et al. (2017) and 7.8 %, as
reported by Jimenez et al. (2000)) to much higher values (e.g. 39 %, as reported by Richmond-Bryant et al. (2017) and
36 %, as reported by Costantini et al. (2016)). 3. In order to improve generalization of the temporal profiles, the optimization is performed using simulated and observed
data from May 2018. Additionally, we accelerate the optimization by only using two weeks of simulation time for each
optimization step. The final temporal profiles are evaluated in a full-month simulation for the year 2019, as described in
sect. 2. 190 Figure 2 gives an overview of the optimization process in a total of 3 steps. Figure 2h reveals that the overall NOx budget
assumed for the simulated month is realistic, meaning that the corresponding monthly weighting factor is reasonable. Similarly,
the daily profiles from Kumar et al. (2021) were taken over without changes. What remains are the hourly profiles of each sector. Prior to the optimization, simulated NOx was too low at daytime and too high at night. Therefore, the hourly profiles were Figure 2 gives an overview of the optimization process in a total of 3 steps. Figure 2h reveals that the overall NOx budget
assumed for the simulated month is realistic, meaning that the corresponding monthly weighting factor is reasonable. Similarly,
the daily profiles from Kumar et al. (2021) were taken over without changes. What remains are the hourly profiles of each sector. Prior to the optimization, simulated NOx was too low at daytime and too high at night. Therefore, the hourly profiles were adjusted accordingly. For example, in step 1 (red line), only a single change was made to the traffic sector profile (subfigure a)
195
in order to boost simulated NOx values in the morning (∼6 AM). The principle was extended to the remaining hourly profiles
in the next two steps. The full list of emission profiles used in our simulation is attached to the supplement of this article. 2.2
Optimization of temporal profiles The key difference between previously published RCT simulations and ours is the optimization of the temporal emission
150
profiles. The goal is to find temporal profiles that minimize the model’s mean NOx bias over the course of the day. The mean
relative bias is computed as bias = 1
N
N
X
i=1
xsim,i −xobs,i
xobs,i
(4) bias = 1
N
N
X
i=1
xsim,i −xobs,i
xobs,i (4) measurements of background NOx surface concentrations as reference. A detailed explanation of the dataset is given in sect. 155
3.1. Equation (3) hints towards the complexity of the optimization problem: Because the emission inventories include dozens
of emission sectors, each with their own emission profiles, this poses an optimization problem with many degrees of freedom. In addition, a single WRF-Chem simulation of just one month takes days to finish even on modern supercomputers. This
circumstance makes it nearly impossible to optimize the emission profiles using standard methods like gradient descent. For our simulation we have therefore optimized the emission profiles manually. By "manual optimization" we mean the iterative
160
process of running WRF-Chem, evaluating the simulation results against in-situ observations, and slightly nudging the temporal
profiles in a direction in which better agreement between simulation and observations can be expected. Due to the short lifetime
of NOx the observed concentrations closely follow the temporal profiles. As an initial starting point, the temporal profiles from
Kumar et al. (2021) were used. Although the proposed optimization method is rather unconventional, it has a few important 1. The hourly profiles of many sectors have characteristic shapes, e.g. a peak in the hourly profile of the traffic sector during
the morning rush hour. These should be at least coarsely preserved during the optimization process in order to maintain
realistic emission behaviour. 1. The hourly profiles of many sectors have characteristic shapes, e.g. a peak in the hourly profile of the traffic sector during
the morning rush hour. These should be at least coarsely preserved during the optimization process in order to maintain
realistic emission behaviour. 2. Because the optimization problem is ill-posed, it is often unclear, of which sector the profiles should be tuned further
in order to improve the simulation. Sometimes, the spatial distribution of a specific emission sector matches that of the
170
model error, indicating that this sector should be prioritized. 7 7 2.3
Chemical boundary conditions Overview of the optimization process of the hourly emission profiles based on the simulation results and in-situ measurements of
May 2018. The profiles used by Kumar et al. (2021) are drawn in blue and our final optimized profiles in orange. Therefore, in addition to the tuning of temporal emission profiles, we further conduct a tuning Therefore, in addition to the tuning of temporal emission profiles, we further conduct a tuning of O3 BCs in the same fashion. As a result, a O3 BC scaling factor of ∼0.85 was determined. The results discussed in sect. 3 verify that scaling of the O3 BCs
205
yields an improvement to the resulting O3 concentrations. As a result, a O3 BC scaling factor of ∼0.85 was determined. The results discussed in sect. 3 verify that scaling of the O3 BCs
205
yields an improvement to the resulting O3 concentrations. 2.3
Chemical boundary conditions As described, our simulation uses CAM-chem model results as chemical boundary conditions (BCs) on the outer domain D1
and initial conditions (ICs) on both domains. In practice, it shows that simulated NO2 and NO concentrations are mostly
200
insensitive to BCs and ICs. O3 concentrations, however, are highly sensitive to the O3 BCs. At the same time, O3 is known to
be a crucial reaction partner of NOx chemistry, mostly for oxidizing NO in the reaction As described, our simulation uses CAM-chem model results as chemical boundary conditions (BCs) on the outer domain D1
and initial conditions (ICs) on both domains. In practice, it shows that simulated NO2 and NO concentrations are mostly
200
insensitive to BCs and ICs. O3 concentrations, however, are highly sensitive to the O3 BCs. At the same time, O3 is known to
be a crucial reaction partner of NOx chemistry, mostly for oxidizing NO in the reaction
NO + O3 →NO2 + O2
(R1) (R1) NO + O3 →NO2 + O2 8 AMT paper
6.96986 in * 9.13422 in (full size)
8.3 cm * 5.1297 cm (single col. Fig., golden ratio)
17.5 cm * 10.8156 cm (double col. Fig., golden ratio)
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c⃝Author(s) 2023. CC BY 4.0 License. AMT paper
6.96986 in * 9. 8.3 cm * 5.129
17 5 cm * 10 8
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Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. (a)
(a)
(d)
(g)
(b)
(e)
(h)
(c)
(f)
(i)
[µg m-3]
Initial configuration
Step 1
Step 2
Step 3
In-situ measurements (background stations)
Figure 2. Overview of the optimization process of the hourly emission profiles based on the simulation results and in-situ measurements of
May 2018. The profiles used by Kumar et al. (2021) are drawn in blue and our final optimized profiles in orange. (a)
(a)
(b)
(c) (b)
(c) (e)
(f) (d) (f) (e) (f) (i) (g)
(h)
(i)
[µg m-3]
Initial configuration
Step 1
Step 2
Step 3
In-situ measurements (background station
Figure 2. Overview of the optimization process of the hourly emission profiles based on the simulation results and in-situ measuremen (g)
(h)
(i)
[µg m-3]
I iti l
fi
ti
St
1
St
2
St
3
I
it (i) (h) (h)
[µg m-3] In-situ measurements (background stations) Figure 2. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 3.1
Comparison of surface concentrations Given that the used emission inventories treat NO and NO2 as lumped NOx, this validates our choice of temporal Comparing S-old and S-new shows the vast difference in simulation accuracy achieved by using the optimized temporal
230
profiles. S-old consistently overestimates the NOx concentration at night, but underestimates it at daytime. Particularly around
noontime the simulated NOx concentrations are low-biased by approximately −20 %. S-new achieves much better agreement
with the in-situ measurements: The mean NOx concentration is reproduced with minimal deviations (< 2 µg m−3), regardless
of daytime. Given that the used emission inventories treat NO and NO2 as lumped NOx, this validates our choice of temporal profiles. 235
The individual time series of NO2 and NO also show good agreement with the in-situ observations, with minimal to no
deviations at noontime. However, discrepancies during nighttime between simulation and observations remain (< 4.7 µg m−3
for NO2 and < 4.1 µg m−3 for NO). S-old shows a consistent overestimation of O3 of approximately 20 µg m−3 regardless of daytime, which is reduced to profiles. 235
The individual time series of NO2 and NO also show good agreement with the in-situ observations, with minimal to no
deviations at noontime. However, discrepancies during nighttime between simulation and observations remain (< 4.7 µg m−3
for NO2 and < 4.1 µg m−3 for NO). S-old shows a consistent overestimation of O3 of approximately 20 µg m−3 regardless of daytime, which is reduced to profiles. 235
The individual time series of NO2 and NO also show good agreement with the in-situ observations, with minimal to no
deviations at noontime. However, discrepancies during nighttime between simulation and observations remain (< 4.7 µg m−3
for NO2 and < 4.1 µg m−3 for NO). S-old shows a consistent overestimation of O3 of approximately 20 µg m−3 regardless of daytime, which is reduced to approximately 10 µg m−3 in S-new. The improvements are mostly attributed to the adjustments to the O3 boundary conditions
240
described in sect. 2.3. A version of Fig. 3 showing traffic stations instead of background stations can be found in appendix
A (Fig. A1). The NO2 concentration timeline of the traffic observations has a fundamentally different shape compared to the
background observations, with practically no noontime low and no evening peak. As expected, the modelled and observed
concentration timelines do not agree well for the traffic stations. 3
Results We compare the results of two WRF-Chem simulations for May 2019 (one with the temporal profiles used by Kumar et al. (2021) called "S-old", and one with optimized profiles called "S-new") to the dataset of in-situ surface concentration mea-
surements and show that S-new outperforms S-old in every regard. We then continue to compare S-new to the remaining
210
observational datasets of satellite and MAX-DOAS measurements. 9 9 3.1
Comparison of surface concentrations The left panel (subfigures a-d)
is restricted to the first ten days of the simulated time period (01 May 2019 - 10 May 2019) for easier readability. The right
panel (subfigures e-h) shows the average diurnal concentrations obtained from averaging over all days of the simulation period. ∆xt = max(|xt −xt−1|,|xt −xt+1|)
(5)
225
i.e. the maximal hour-to-hour variation in concentration. Figure 3 shows the average surface concentrations of NO2, NO, NOx,
and O3 of both simulations S-old and S-new as well as the corresponding in-situ measurements. The left panel (subfigures a-d)
is restricted to the first ten days of the simulated time period (01 May 2019 - 10 May 2019) for easier readability. The right
panel (subfigures e-h) shows the average diurnal concentrations obtained from averaging over all days of the simulation period. ∆xt = max(|xt −xt−1|,|xt −xt+1|)
(5)
225
i.e. the maximal hour-to-hour variation in concentration. Figure 3 shows the average surface concentrations of NO2, NO, NOx,
and O3 of both simulations S-old and S-new as well as the corresponding in-situ measurements. The left panel (subfigures a-d)
is restricted to the first ten days of the simulated time period (01 May 2019 - 10 May 2019) for easier readability. The right
panel (subfigures e-h) shows the average diurnal concentrations obtained from averaging over all days of the simulation period. (5) Comparing S-old and S-new shows the vast difference in simulation accuracy achieved by using the optimized temporal
230
profiles. S-old consistently overestimates the NOx concentration at night, but underestimates it at daytime. Particularly around
noontime the simulated NOx concentrations are low-biased by approximately −20 %. S-new achieves much better agreement
with the in-situ measurements: The mean NOx concentration is reproduced with minimal deviations (< 2 µg m−3), regardless
of daytime. Given that the used emission inventories treat NO and NO2 as lumped NOx, this validates our choice of temporal Comparing S-old and S-new shows the vast difference in simulation accuracy achieved by using the optimized temporal
230
profiles. S-old consistently overestimates the NOx concentration at night, but underestimates it at daytime. Particularly around
noontime the simulated NOx concentrations are low-biased by approximately −20 %. S-new achieves much better agreement
with the in-situ measurements: The mean NOx concentration is reproduced with minimal deviations (< 2 µg m−3), regardless
of daytime. 3.1
Comparison of surface concentrations In-situ measurements of trace gas surface concentrations in Germany are conducted by the UBA. They are available for NO2,
NO and O3 as hourly mean values. A total of 434 UBA measuring stations are distributed over Germany. 92 % of the sta-
tions use a chemiluminescence based measuring method. The remaining 8 % use other methods (cavity enhanced phase shift
215
spectroscopy, diffuse sampling, or photolysis conversion). Of all UBA stations, 63 % are classified as "background", 30 % as
"traffic", and 7 % as "industry". In this study we only take "background" stations into consideration. This choice was made
seeing that NO2 concentration can vary strongly near traffic over distances of only 10 - 100 m (see e.g. Beckwith et al., 2019)
and timescales of a few minutes. Given that our simulation runs at a spatial resolution of 3 km × 3 km, it is expected to show poor agreement with the traffic stations. Many of the publications referenced in sect. 1 have also restricted their reference data
220
to "background" in-situ measurements. In order to compare the simulated surface concentrations to the in-situ measurements, they are interpolated from the WRF-
Chem simulation grid to the geolocations of the UBA stations. The uncertainty ∆xt of the simulated surface concentrations
and the in-situ reference values at time step t is computed via poor agreement with the traffic stations. Many of the publications referenced in sect. 1 have also restricted their reference data
220
to "background" in-situ measurements. In order to compare the simulated surface concentrations to the in-situ measurements, they are interpolated from the WRF-
Chem simulation grid to the geolocations of the UBA stations. The uncertainty ∆xt of the simulated surface concentrations
and the in-situ reference values at time step t is computed via In order to compare the simulated surface concentrations to the in-situ measurements, they are interpolated from the WRF-
Chem simulation grid to the geolocations of the UBA stations. The uncertainty ∆xt of the simulated surface concentrations
and the in-situ reference values at time step t is computed via ∆xt = max(|xt −xt−1|,|xt −xt+1|)
(5)
225
i.e. the maximal hour-to-hour variation in concentration. Figure 3 shows the average surface concentrations of NO2, NO, NOx,
and O3 of both simulations S-old and S-new as well as the corresponding in-situ measurements. 3.1
Comparison of surface concentrations Interestingly, although this was not considered during the optimization process, the temporal profiles we have obtained for the traffic sector (see Fig. 2a) have a shape similar to the NOx
245 10 0
10
20
30
NO2 conc. [µg m
3]
(a)
(a)
0
10
20
30
(e)
(e)
Monthly average
0.0
2.5
5.0
7.5
10.0
12.5
NO conc. [µg m
3]
(b)
(b)
0.0
2.5
5.0
7.5
10.0
12.5
(f)
(f)
0
10
20
30
40
NOx conc. [µg m
3]
(c)
(c)
0
10
20
30
40
(g)
(g)
Wed, 05-01 00:00
Thu, 05-02 00:00
Fri, 05-03 00:00
Sat, 05-04 00:00
Sun, 05-05 00:00
Mon, 05-06 00:00
Tue, 05-07 00:00
Wed, 05-08 00:00
Thu, 05-09 00:00
Fri, 05-10 00:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
Date and time
25
50
75
100
125
O3 conc. [µg m
3]
(d)
00:00
04:00
08:00
12:00
16:00
20:00
00:00
Time of day
25
50
75
100
125
(h)
UBA measurements
S-new
S-old
Figure 3. Time series of NO2, NO, NOx, and O3 surface concentrations. The red lines show reference values obtained from the UBA
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c⃝Author(s) 2023. CC BY 4.0 License. 0
10
20
30
NO2 conc. [µg m
3]
(a)
(a)
0
10
20
30
(e)
(e)
Monthly average
0.0
2.5
5.0
7.5
10.0
12.5
NO conc. [µg m
3]
(b)
(b)
0.0
2.5
5.0
7.5
10.0
12.5
(f)
(f)
0
10
20
30
40
NOx conc. [µg m
3]
(c)
(c)
0
10
20
30
40
(g)
(g)
Wed, 05-01 00:00
Thu, 05-02 00:00
Fri, 05-03 00:00
Sat, 05-04 00:00
Sun, 05-05 00:00
Mon, 05-06 00:00
Tue, 05-07 00:00
Wed, 05-08 00:00
Thu, 05-09 00:00
Fri, 05-10 00:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
Date and time
25
50
75
100
125
O3 conc. [µg m
3]
(d)
00:00
04:00
08:00
12:00
16:00
20:00
00:00
Time of day
25
50
75
100
125
(h)
UBA measurements
S-new
S-old
Figure 3. Time series of NO2, NO, NOx, and O3 surface concentrations. The red lines show reference values obtained from the UBA
background in-situ stations. The green and blue lines show the simulation results of S-old and S-new, respectively. 3.1
Comparison of surface concentrations (a) - (d) display a time
series spanning multiple days (01 May 2019 - 10 May 2019) and (e) - (f) display the corresponding average concentration values over the
entire month of May 2019. Monthly average Figure 3. Time series of NO2, NO, NOx, and O3 surface concentrations. The red lines show reference values obtained from the UBA
background in-situ stations. The green and blue lines show the simulation results of S-old and S-new, respectively. (a) - (d) display a time
series spanning multiple days (01 May 2019 - 10 May 2019) and (e) - (f) display the corresponding average concentration values over the
entire month of May 2019. Figure 3. Time series of NO2, NO, NOx, and O3 surface concentrations. The red lines show reference values obtained from the UBA
background in-situ stations. The green and blue lines show the simulation results of S-old and S-new, respectively. (a) - (d) display a time
series spanning multiple days (01 May 2019 - 10 May 2019) and (e) - (f) display the corresponding average concentration values over the
entire month of May 2019. 11 Table 2. Statistical summary for the surface concentration comparison (all hours of the day, "background" stations only) Table 2. Statistical summary for the surface concentration comparison (all hours of the day, "background" stations only) Table 2. Statistical summary for the surface concentration comparison (all hours of the day, "background" sta al summary for the surface concentration comparison (all hours of the day, "background" stations only) y
p
y
g
y
Mean [µg m−3]
RMSE [µg m−3]
Bias
R
Trace gas
UBA
S-new
S-old
S-new
S-old
S-new
S-old
S-new
S-old
NO2
11.9
13.6
15.9
3.7
6.4
+14.3 %
+34.0 %
0.77
0.72
NO
2.5
1.7
1.8
1.9
1.8
-30.0 %
-27.8 %
0.63
0.65
NOx
14.3
15.3
17.7
4.2
6.6
+6.7 %
+23.4 %
0.76
0.67
O3
66.2
73.4
78.5
12.2
16.8
+10.9 %
+18.7 %
0.85
0.82 Table 3. Statistical summary for the surface concentration comparison (at noon, "background" stations only)
Mean [µg m−3]
RMSE [µg m−3]
Bias
R
Trace gas
UBA
S-new
S-old
S-new
S-old
S-new
S-old
S-new
S-old
NO2
8.1
8.1
6.8
1.7
2.0
+0.9 %
-15.7 %
0.76
0.75
NO
2.1
2.1
1.5
0.5
0.8
-1.5 %
-31.8 %
0.72
0.66
NOx
10.2
10.2
8.3
2.0
2.7
+0.4 %
-19.1 %
0.77
0.75
O3
84.9
91.4
100.2
11.1
19.0
+7.6 %
+18.0 %
0.72
0.58 Table 3. Statistical summary for the surface concentration comparison (at noon, "background" stations only) time series of the traffic stations (see Fig. A1g), whereas the corresponding initial profile deviates quite strongly in shape. This
can be seen as further validation of our results. time series of the traffic stations (see Fig. A1g), whereas the corresponding initial profile deviates quite strongly in shape. This
can be seen as further validation of our results. Figure 4 emphasizes the improvement in modelling accuracy via scatter plots for NO2, NO, NOx, and O3 surface concen-
trations at noontime. The results of S-new are consistently closer to the 1:1 line and linear regressions through the point clouds of each plot yield slopes much closer to 1 with similar (small) intercepts compared to S-old. For example, comparing Fig. 4a
250
and Fig. 4b shows slopes of 0.84 (S-new) and 0.73 (S-old) for NO2. A version of Fig. 4 showing values for all hours of the day
can be found in appendix A (see Fig. A2). A summary of corresponding statistical diagnostics is given in Tables 2 and 3 for all
hours of the day and noontime only, respectively. The Root Mean Square Error (RMSE) is computed according to 250 of each plot yield slopes much closer to 1 with similar (small) intercepts compared to S-old. For example, comparing Fig. 4a
250
and Fig. 4b shows slopes of 0.84 (S-new) and 0.73 (S-old) for NO2. A version of Fig. 4 showing values for all hours of the day
can be found in appendix A (see Fig. A2). A summary of corresponding statistical diagnostics is given in Tables 2 and 3 for all
hours of the day and noontime only, respectively. The Root Mean Square Error (RMSE) is computed according to RMSE =
sPN
i=1 (xsim,i −xobs,i)2
N (6) N where xobs,i denotes the i-th observation and xsim,i the corresponding simulated value. Here the RMSE is computed from the
255
averages over all stations used to generate Fig. 3 and 4. where xobs,i denotes the i-th observation and xsim,i the corresponding simulated value. Here the RMSE is computed from the
255
averages over all stations used to generate Fig. 3 and 4. where xobs,i denotes the i-th observation and xsim,i the corresponding simulated value. Here the RMSE is computed from the
255
averages over all stations used to generate Fig. 3 and 4. Comparing the entries of Tables 2 and 3 shows that S-new consistently outperforms S-old by showing lower RMSE, lower
bias, and higher R-values. Most importantly, the biases of total and noontime NOx are reduced from +23.4 % to +6.7 % and
from −19.1 % to +0.4 %, respectively. Similarly, the biases of total and noontime NO2 are reduced from +34.0 % to +14.3 %
and from −15.7 % to +0.9 %. There appears to be only one minor exception to the overall improvement, namely total NO (see
260 Comparing the entries of Tables 2 and 3 shows that S-new consistently outperforms S-old by showing lower RMSE, lower
bias, and higher R-values. Most importantly, the biases of total and noontime NOx are reduced from +23.4 % to +6.7 % and
from −19.1 % to +0.4 %, respectively. Similarly, the biases of total and noontime NO2 are reduced from +34.0 % to +14.3 % Comparing the entries of Tables 2 and 3 shows that S-new consistently outperforms S-old by showing lower RMSE, lower
bias, and higher R-values. Most importantly, the biases of total and noontime NOx are reduced from +23.4 % to +6.7 % and
from −19.1 % to +0.4 %, respectively. Similarly, the biases of total and noontime NO2 are reduced from +34.0 % to +14.3 % and from −15.7 % to +0.9 %. There appears to be only one minor exception to the overall improvement, namely total NO (see
260 12 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 0.0
2.5
5.0
7.5
10.0
12.5
15.0
0
5
10
15
20
Simulated mass
concentration [µg m
3]
(a)
(a)
(a)
(a)
(a)
(a)
(a)
(a)
NO2 (S-new)
f(x) = 0.84x + 1.39
0
2
4
6
8
10
12
14
0
5
10
15
(b)
(b)
(b)
(b)
(b)
(b)
(b)
(b)
NO2 (S-old)
f(x) = 0.73x + 0.87
0
1
2
3
4
0
2
4
Simulated mass
concentration [µg m
3]
(c)
(c)
(c)
(c)
(c)
(c)
(c)
(c)
NO (S-new)
f(x) = 0.89x + 0.20
0.0
0.5
1.0
1.5
2.0
2.5
3.0
0
1
2
3
4
(d)
(d)
(d)
(d)
(d)
(d)
(d)
(d)
NO (S-old)
f(x) = 0.51x + 0.37
0
5
10
15
20
0
10
20
Simulated mass
concentration [µg m
3]
(e)
(e)
(e)
(e)
(e)
(e)
(e)
(e)
NOx (S-new)
f(x) = 0.88x + 1.25
0.0
2.5
5.0
7.5
10.0
12.5
15.0
17.5
0
5
10
15
20
(f)
(f)
(f)
(f)
(f)
(f)
(f)
(f)
NOx (S-old)
f(x) = 0.72x + 0.95
100
150
ed mass
on [µg m
3]
(g)
(g)
(g)
(g)
(g)
(g)
(g)
(g)
O3 (S-new)
f(x) = 0.60x + 40.52
100
150
(h)
(h)
(h)
(h)
(h)
(h)
(h)
(h)
O3 (S-old)
f(x) = 0.52x + 55.92
c⃝Author(s) 2023. CC BY 4.0 License. 0.0
2.5
5.0
7.5
10.0
12.5
15.0
0
5
10
15
20
Simulated mass
concentration [µg m
3]
(a)
(a)
(a)
(a)
(a)
(a)
(a)
(a)
NO2 (S-new)
f(x) = 0.84x + 1.39
0
2
4
6
8
10
12
14
0
5
10
15
(b)
(b)
(b)
(b)
(b)
(b)
(b)
(b)
NO2 (S-old)
f(x) = 0.73x + 0.87
0
1
2
3
4
0
2
4
Simulated mass
concentration [µg m
3]
(c)
(c)
(c)
(c)
(c)
(c)
(c)
(c)
NO (S-new)
f(x) = 0.89x + 0.20
0.0
0.5
1.0
1.5
2.0
2.5
3.0
0
1
2
3
4
(d)
(d)
(d)
(d)
(d)
(d)
(d)
(d)
NO (S-old)
f(x) = 0.51x + 0.37
0
5
10
15
20
0
10
20
Simulated mass
concentration [µg m
3]
(e)
(e)
(e)
(e)
(e)
(e)
(e)
(e)
NOx (S-new)
f(x) = 0.88x + 1.25
0.0
2.5
5.0
7.5
10.0
12.5
15.0
17.5
0
5
10
15
20
(f)
(f)
(f)
(f)
(f)
(f)
(f)
(f)
NOx (S-old)
f(x) = 0.72x + 0.95
0
20
40
60
80
100
120
Observed mass concentration [µg m
3]
0
50
100
150
Simulated mass
concentration [µg m
3]
(g)
(g)
(g)
(g)
(g)
(g)
(g)
(g)
O3 (S-new)
f(x) = 0.60x + 40.52
0
20
40
60
80
100
120
Observed mass concentration [µg m
3]
0
50
100
150
(h)
(h)
(h)
(h)
(h)
(h)
(h)
(h)
O3 (S-old)
f(x) = 0.52x + 55.92
Figure 4. Scatter plots of simulated vs. observed surface concentrations of NO2, NO, NOx and O3. One scatter point resembles the mean
over all "background" stations at noontime of one day. The left panel (a) - (g) shows the results of the new simulation (S-new) and the right
panel (b) - (h) shows the results of the simulation with the original temporal profiles (S-old). The dashed line in each subplot marks the 1:1
line. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. second line of Table 2). However, the difference in RMSE between S-new and S-old is on the scale of ∼0.1 µg m−3 for NO,
which is minuscule second line of Table 2). However, the difference in RMSE between S-new and S-old is on the scale of ∼0.1 µg m−3 for NO,
which is minuscule. 0
2
4
6
8
10
12
14
0
5
10
15
(b)
(b)
(b)
(b)
(b)
(b)
(b)
(b)
NO2 (S-old)
f(x) = 0.73x + 0.87
(d)
(d)
(d)
(d)
(d)
(d)
(d)
(d)
NO (S-old) 0.0
2.5
5.0
7.5
10.0
12.5
15.0
0
5
10
15
20
Simulated mass
concentration [µg m
3]
(a)
(a)
(a)
(a)
(a)
(a)
(a)
(a)
NO2 (S-new)
f(x) = 0.84x + 1.39
0
0
5
10
15
( )
( )
( )
( )
( )
( )
( )
( ) (b)
(b)
(b)
(b)
(b)
(b)
(b)
(b) (d)
(d)
(d)
(d)
(d)
(d)
(d)
(d) (f)
(f)
(f)
(f)
(f)
(f)
(f)
(f) Simulated mass (h)
(h)
(h)
(h)
(h)
(h)
(h)
(h) 0
20
40
60
80
100
120
Observed mass concentration [µg m
3]
0
50
100
150
f(x) = 0.52x + 55.92 Figure 4. Scatter plots of simulated vs. observed surface concentrations of NO2, NO, NOx and O3. One scatter point resembles the mean
over all "background" stations at noontime of one day. The left panel (a) - (g) shows the results of the new simulation (S-new) and the right
panel (b) - (h) shows the results of the simulation with the original temporal profiles (S-old). The dashed line in each subplot marks the 1:1
line. 13 3.2
Comparison of NO2 vertical column densities As a second diagnostic, we compare simulated NO2 vertical column densities (VCDs) to observations of the TROPOMI
(TROPOspheric Monitoring Instrument, see Veefkind et al., 2012) satellite instrument. The satellite overpass occurs daily,
265
typically at 13:30 local time in central Europe, with a pixel size of 3.5 km × 5.5 km in NADIR viewing geometry. TROPOMI
measures spectra of backscattered sunlight from earth’s surface, from which tropospheric slant column densities (SCDs) are
computed using Differential Optical Absorption Spectroscopy (DOAS, see Platt and Stutz, 2008). Tropospheric VCDs are
obtained from tropospheric SCDs via VCDtrop = SCDtrop
AMFtrop
(7)
270
where AMFtrop stands for the tropospheric air mass factor. The tropospheric AMF depends on a number of atmospheric and
surface conditions and assumptions, e.g. cloud properties, the stratospheric column, surface albedo, and the relative a priori
NO2 profile. In the TROPOMI retrieval, the NO2 a priori profiles are taken from the TM5 global CT model (see Krol et al. (2005); Williams et al. (2017)), with a spatial resolution of ∼1◦× 1◦≈110 km × 70 km where our simulation domain D2 is (7) located. A detailed description of the retrieval algorithm can be found in van Geffen et al. (2022) and Eskes et al. (2021). From
275
hereon we exclusively refer to tropospheric VCDs and SCDs, respectively. Simulated VCDs are obtained by vertical integration of box VCDs (= partial columns) from WRF-Chem. The WRF-Chem
output is interpolated to the vertical grid of the TM5 model. The box VCDs are computed by multiplying the NO2 concentration
within each model grid cell with its vertical extent. The partial columns are then summed up: located. A detailed description of the retrieval algorithm can be found in van Geffen et al. (2022) and Eskes et al. (2021). From
275
hereon we exclusively refer to tropospheric VCDs and SCDs, respectively. Si
l t d VCD
bt i
d b
ti
l i t
ti
f b
VCD (
ti l
l
) f
WRF Ch
Th WRF Ch Simulated VCDs are obtained by vertical integration of box VCDs (= partial columns) from WRF-Chem. The WRF-Chem
output is interpolated to the vertical grid of the TM5 model. The box VCDs are computed by multiplying the NO2 concentration
within each model grid cell with its vertical extent. TM5 model. It has been demonstrated that replacing TM5 a priori profiles with profiles from high-resolution RCT
models improves the VCD retrieval significantly (see e.g. Ialongo et al. (2020); Tack et al. (2021); Liu et al. (2021)). TM5 model. It has been demonstrated that replacing TM5 a priori profiles with profiles from high-resolution RCT
models improves the VCD retrieval significantly (see e.g. Ialongo et al. (2020); Tack et al. (2021); Liu et al. (2021)). TM5 model. It has been demonstrated that replacing TM5 a priori profiles with profiles from high-resolution RCT
models improves the VCD retrieval significantly (see e.g. Ialongo et al. (2020); Tack et al. (2021); Liu et al. (2021)). We incorporate the AKs and our simulated high-resolution NO2 a priori profiles by computing corrected observed VCDs
following Eskes et al. (2019): We incorporate the AKs and our simulated high-resolution NO2 a priori profiles by computing corrected observed VCDs
following Eskes et al. (2019): We incorporate the AKs and our simulated high-resolution NO2 a priori profiles by computing corrected observed VCDs
following Eskes et al. (2019): obs · AMFtrop
AMF
·
P
l<13 cl · ∆hl
P
l<13 cl · ∆hl · Al
(9) (9) where VCDobs denotes the observed NO2 VCD, AMFtrop the tropospheric AMF, and A the averaging kernel vector. where VCDobs denotes the observed NO2 VCD, AMFtrop the tropospheric AMF, and A the averaging kernel vector. 2. Each observed VCD has an associated q-value that describes the retrieval quality on a range from 0 (bad) to 1 (good)
(see van Geffen et al., 2022). q-values of ≥0.75 also require the cloud radiance fraction (crf) to be smaller than 0.5,
which effectively acts as a cloud filter. The effect of clouds on the measurement process depends on the height of the
cloud layer: High clouds can shield off lower layers of the atmosphere, decreasing the NO2 sensitivity to almost zero
below the cloud layer. Vice versa, cloud layers directly below NO2 layers can enhance the observed NO2 absorption due
to the increased cloud albedo. If cloud layers and NO2 layers are at the same altitude, NO2 absorption can be increased
by an enhanced light path due to multiple scattering. The influence of clouds on satellite measurements of NO2 has been
discussed e.g. in Martin et al. (2002), Kokhanovsky and Rozanov (2008), and Liu et al. (2021). 3.2
Comparison of NO2 vertical column densities The partial columns are then summed up: (8) sim
X
l<13
l
l
( )
where VCDsim denotes the simulated VCD, l the TM5 model layer index, cl the NO2 concentration in layer l, and ∆hl the
vertical extent of layer l. Usually the summation would be conducted up until layer ltp, the tropopause layer of the TM5 model. Typically, ltp ≈16. WRF-Chem, however, is designed to be a tropospheric model, meaning that model results within the
tropopause region should be used with caution. We observe an increase in modelled NO2 concentrations above 10 km altitude (which equates approximately to layer 13 in the TM5 model, see Fig. A3), which is likely due to stratosphere-troposphere
285
exchange. We therefore limit the summation of box VCDs to the lowest 13 layers. Then, pairs of simulated and observed
VCDs are obtained by interpolating the WRF-Chem data to the horizontal TROPOMI grid in space (linear) and time (nearest
neighbour). In order to make a representative comparison between simulated and observed VCDs, the following further aspects are taken
into consideration:
290 In order to make a representative comparison between simulated and observed VCDs, the following further aspects are taken
into consideration:
290 1. TROPOMI is not equally sensitive to all layers of the troposphere. This circumstance is described by the averaging
kernels (AKs, see van Geffen et al., 2022). Thus the AMF depends on relative NO2 a priori profiles provided by the 1. TROPOMI is not equally sensitive to all layers of the troposphere. This circumstance is described by the averaging
kernels (AKs, see van Geffen et al., 2022). Thus the AMF depends on relative NO2 a priori profiles provided by the 14 We apply a q-filter that
removes all observations with q < 0.75, as recommended by Eskes et al. (2019). Figure 5 shows the result of our comparison in the form of monthly mean values (i.e. the total average over May 2019). Subfigures a-e show the comparison between simulated NO2 VCDs and TROPOMI observations using the TM5 NO2 a priori Subfigures a-e show the comparison between simulated NO2 VCDs and TROPOMI observations using the TM5 NO2 a priori
profiles. Strong enhancements occur in west Germany (Ruhr region) and along the Rhine river. Moderate enhancements occur
310
over logistical hotpots (e.g. the port of Hamburg with a nearby coal power plant), and larger cities (e.g. Berlin and Munich;
see Fig. 8 for an overview of the geographical regions mentioned here). Subfigure c shows the difference between simulated
and observed VCDs and reveals the model’s tendency to overestimate the observed VCDs. The overestimation is close to zero
in rural regions but reaches up to 1016 molec. cm−2 over hotspot regions. The simulated and observed VCDs correlate with profiles. Strong enhancements occur in west Germany (Ruhr region) and along the Rhine river. Moderate enhancements occur
310
over logistical hotpots (e.g. the port of Hamburg with a nearby coal power plant), and larger cities (e.g. Berlin and Munich;
see Fig. 8 for an overview of the geographical regions mentioned here). Subfigure c shows the difference between simulated
and observed VCDs and reveals the model’s tendency to overestimate the observed VCDs. The overestimation is close to zero
in rural regions but reaches up to 1016 molec. cm−2 over hotspot regions. The simulated and observed VCDs correlate with an R-value of 0.83, an RMSE of 1.23 · 1015 molec. cm−2, and a bias of +17.6 %. Subfigures f-j show the same comparison,
315
but with recomputed airmass factors according to eq. (9). The comparison improves significantly with an R-value of 0.84, an
RMSE of 1.11 · 1015 molec. cm−2, and a bias of only −6.6 %. Subfigures k-o show the spatial distribution of simulated and observed surface NO2 concentrations. Model overestimations
of NO2 VCD and surface concentration occur in similar geographic regions, particularly in the Ruhr region, along the Rhine an R-value of 0.83, an RMSE of 1.23 · 1015 molec. cm−2, and a bias of +17.6 %. Subfigures f-j show the same comparison,
315
but with recomputed airmass factors according to eq. (9). The comparison improves significantly with an R-value of 0.84, an
RMSE of 1.11 · 1015 molec. cm−2, and a bias of only −6.6 %. Subfigures k-o show the spatial distribution of simulated and observed surface NO2 concentrations. Model overestimations
of NO2 VCD and surface concentration occur in similar geographic regions, particularly in the Ruhr region, along the Rhine Subfigures k-o show the spatial distribution of simulated and observed surface NO2 concentrations. Model overestimations
of NO2 VCD and surface concentration occur in similar geographic regions, particularly in the Ruhr region, along the Rhine
river and the port of Hamburg This hints towards a possible overestimation of NO emissions in these regions
320 Subfigures k-o show the spatial distribution of simulated and observed surface NO2 concentrations. Model overestimations
of NO2 VCD and surface concentration occur in similar geographic regions, particularly in the Ruhr region, along the Rhine
river, and the port of Hamburg. This hints towards a possible overestimation of NOx emissions in these regions. 320 river, and the port of Hamburg. This hints towards a possible overestimation of NOx emissions in these regions. 320
Figure 6 presents further comparisons of simulated and observed NO2 VCDs in the style of Fig. 5. Here, we only show
comparisons using reprocessed observations according to eq. (9). Subfigures a-e show the comparison restricted to Germany,
where the high-resolution UBA-E emission inventory was used. Compared to the full domain, an increase in RMSE (1.46 ·
1015 molec. cm−2) and decrease in bias (+3.3%) is observed. The increase in RMSE can be attributed to the Ruhr region in ,
p
g
p
x
g
Figure 6 presents further comparisons of simulated and observed NO2 VCDs in the style of Fig. 5. Here, we only show
comparisons using reprocessed observations according to eq. (9). Subfigures a-e show the comparison restricted to Germany,
where the high-resolution UBA-E emission inventory was used. Compared to the full domain, an increase in RMSE (1.46 ·
1015 molec. cm−2) and decrease in bias (+3.3%) is observed. The increase in RMSE can be attributed to the Ruhr region in
west Germany which is one of the most polluted regions within our simulation domain Subfigures f-j show the simulation
325 west Germany, which is one of the most polluted regions within our simulation domain. Subfigures f-j show the simulation
325 15 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 53°N
50°N
47°N
17°E
11°E
5°E
WRF-Chem
(d)
(a)
(b)
(c)
(d)
(e)
(f)
(g)
(h)
Figure 7. Comparison of simulated and observed NO2 VCDs for two exemplary individual days. (a)-(d) shows 23 May 2019, (e)-(h) shows
15 May 2019. Gray regions are filtered due to the strict masking of clouded pixels. (d) 53°N
50°N
47°N
17°E
11°E
5°E
WRF-Chem
(d)
(a)
(b)
(c)
(d) (d) (h) (e)
(f)
(g)
(h) (e)
(f)
(g) (g) Figure 7. Comparison of simulated and observed NO2 VCDs for two exemplary individual days. (a)-(d) shows 23 May 2019, (e)-(h) shows
15 May 2019. Gray regions are filtered due to the strict masking of clouded pixels. domain with Germany excluded. Although there are still some strongly polluted regions where WRF-Chem underestimates the
NO2 VCD (e.g. the Netherlands) there are also large regions with model underestimation (e.g. Paris, France, Poland, Austria,
and Hungary). This results in an overall mean bias of −9.6% on the displayed domain. Fi
6k
h
h
i
h f ll i
l i
d
i
i
h
i i
l
l
fil
b K
l (2021) domain with Germany excluded. Although there are still some strongly polluted regions where WRF-Chem underestimates the
NO2 VCD (e.g. the Netherlands) there are also large regions with model underestimation (e.g. Paris, France, Poland, Austria,
and Hungary). This results in an overall mean bias of −9.6% on the displayed domain. Figure 6k-o shows the comparison on the full simulation domain using the original temporal profiles by Kumar et al. (2021), Figure 6k-o shows the comparison on the full simulation domain using the original temporal profiles by Kumar et al. (2021),
referred to as S-old previously. Although the linear fit in subfigure o is remarkably good with a slope of 1.00, the overall mean
330
bias is −10.2%. This is in agreement with sect. 3.1, where it was shown that S-old suffers from a low bias of −15.7 % for
NO2 surface concentrations around noon time. However, in stark contrast to the surface concentrations, which were 16.6 %
lower in S-old, the simulated NO2 VCDs are reduced by only 3.6 % in comparison to S-new. This shows that surface NO2
concentrations are highly sensitive to the hourly profiles (i.e. the diurnal distribution of emissions), while the integrated NO2
column mostly depends on the total emission budget. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 53°N
50°N
47°N
17°E
11°E
5°E
WRF-Chem
ure 5. Comparison of simulated and observed NO2 VCDs (a)-(j) and surface NO2 concentrations (k)-(o). The first row (a)-(e) shows
comparison to the original observations using low-resolution TM5 a priori NO2 profiles. The second row (f)-(j) shows the comparison
reprocessed observations using high-resolution WRF-Chem a priori profiles and averaging kernels. All satellite observations were
icted to cases with q ≥0.75. The third row (k)-(o) shows a comparison between simulated and observed surface NO2 concentrations
m "background" stations. 53°N
50°N
47°N
17°E
11°E
5°E
WRF-Chem Figure 5. Comparison of simulated and observed NO2 VCDs (a)-(j) and surface NO2 concentrations (k)-(o). The first row (a)-(e) shows
the comparison to the original observations using low-resolution TM5 a priori NO2 profiles. The second row (f)-(j) shows the comparison
with reprocessed observations using high-resolution WRF-Chem a priori profiles and averaging kernels. All satellite observations were
restricted to cases with q ≥0.75. The third row (k)-(o) shows a comparison between simulated and observed surface NO2 concentrations
from "background" stations. 16 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 53°N
50°N
47°N
17°E
11°E
5°E
WRF-Chem
(a)
(b)
(c)
(d)
(e)
(f)
(g)
(h)
(i)
(j)
(k)
(l)
(m)
(n)
(o)
ure 6. Further comparisons of simulated and observed NO2 VCDs. The first row (a)-(e) shows the comparison only for Germany. The
nd row (f)-(j) shows the comparison for the entire simulation domain except for Germany. The third row (k)-(o) shows a comparison
re instead of the newly optimized temporal profiles, the original ones by Kumar et al. (2021) were used. (e) WRF-Chem 53°N
50°N
47°N
17°E
11°E
5°E
WRF-Chem
(a)
(b)
(c)
(d)
(e) (e) 47 N
(f)
(g)
(h)
(i)
(j) (j) (i) (f)
(g)
(h)
(i)
(j) (f)
(g)
(h)
(i) (j) (h) (n)
(o) (k)
(l)
(m)
(n)
(o) (m) Figure 6. Further comparisons of simulated and observed NO2 VCDs. The first row (a)-(e) shows the comparison only for Germany. The
second row (f)-(j) shows the comparison for the entire simulation domain except for Germany. The third row (k)-(o) shows a comparison
where instead of the newly optimized temporal profiles, the original ones by Kumar et al. (2021) were used. 17 17 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2022-1473
Preprint. The two displayed days were chosen because they cover the most pol-
luted regions of the simulation domain well. In comparison to monthly means, the modelled NO2 VCDs are smeared out. This is most likely an artefact of the simulated wind directions and speeds. In a monthly mean, such effects are expected to cancel out
340
statistically. Furthermore, a significant decrease in R-value (0.66 and 0.80) and an increase in RMSE (2.32·1015 molec cm−2
and 1.92·1015 molec cm−2) is observed in comparison to monthly means (R = 0.84, RMSE = 1.11·1015 molec cm−2). This
is expected, seeing that averaging reduces the statistical noise of both datasets. Noise sources that contribute statistically in-
clude the DOAS fit error (from the SCD retrieval), uncertainties of the AMF (e.g. due to clouds) and meteorological model
variables. Table 4 summarizes the results of this section. 345 340 335 referred to as S-old previously. Although the linear fit in subfigure o is remarkably good with a slope of 1.00, the overall mean
330
bias is −10.2%. This is in agreement with sect. 3.1, where it was shown that S-old suffers from a low bias of −15.7 % for
NO2 surface concentrations around noon time. However, in stark contrast to the surface concentrations, which were 16.6 %
lower in S-old, the simulated NO2 VCDs are reduced by only 3.6 % in comparison to S-new. This shows that surface NO2
concentrations are highly sensitive to the hourly profiles (i.e. the diurnal distribution of emissions), while the integrated NO2 column mostly depends on the total emission budget. 335
So far only monthly means were considered in the evaluation. Figure 7 shows the comparison for two exemplary individual
days: 23 May and 15 May 2019. When looking at single days, large regions of the maps are filtered due to the strict masking of So far only monthly means were considered in the evaluation. Figure 7 shows the comparison for two exemplary individual
days: 23 May and 15 May 2019. When looking at single days, large regions of the maps are filtered due to the strict masking of 18 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Table 4. Statistical summary of the results shown in Fig. 5, 6, and 7 Referring to
NO2 a priori profiles
Mean (sim.)
Mean (obs.)
Slope
Intercept
RMSE
Bias
R
Fig. 5a-e
TM5
2.52
2.14
1.39
-0.04
1.23
+17.6 %
0.83
Fig. 5f-j
WRF-Chem
2.52
2.70
1.15
-0.06
1.11
-6.6 %
0.84
Germany only, Fig. 6a-e
WRF-Chem
3.48
3.37
1.38
-0.12
1.46
+3.3 %
0.83
without Germany, Fig. 6f-j
WRF-Chem
2.30
2.54
1.08
-0.04
1.01
-9.6 %
0.83
old temporal profiles, Fig. 6k-o
WRF-Chem
2.41
2.68
1.00
-0.03
0.97
-10.2 %
0.83
single day (23 May 2019, Fig. 7a-d)
WRF-Chem
2.78
3.15
0.80
0.03
2.32
-11.8 %
0.66
single day (15 May 2019, Fig. 7e-h)
WRF-Chem
2.89
2.89
1.13
-0.04
1.92
-0.3 %
0.80
Means and RMSE are given in units of [1015 molec. cm−2] clouded pixels. This is the case for all days of May 2019. The two displayed days were chosen because they cover the most pol-
luted regions of the simulation domain well. In comparison to monthly means, the modelled NO2 VCDs are smeared out. This
is most likely an artefact of the simulated wind directions and speeds. In a monthly mean, such effects are expected to cancel out
40
statistically. Furthermore, a significant decrease in R-value (0.66 and 0.80) and an increase in RMSE (2.32·1015 molec cm−2
and 1.92·1015 molec cm−2) is observed in comparison to monthly means (R = 0.84, RMSE = 1.11·1015 molec cm−2). This
is expected, seeing that averaging reduces the statistical noise of both datasets. Noise sources that contribute statistically in-
clude the DOAS fit error (from the SCD retrieval), uncertainties of the AMF (e.g. due to clouds) and meteorological model
variables. Table 4 summarizes the results of this section. 45 clouded pixels. This is the case for all days of May 2019. The two displayed days were chosen because they cover the most pol-
luted regions of the simulation domain well. In comparison to monthly means, the modelled NO2 VCDs are smeared out. This
is most likely an artefact of the simulated wind directions and speeds. In a monthly mean, such effects are expected to cancel out
40
15
2 clouded pixels. This is the case for all days of May 2019. Means and RMSE are given in units of [1015 molec. cm−2] 3.3
Comparison of NO2 concentration profiles As a third diagnostic we compare modelled NO2 concentration profiles to profiles obtained from MAX-DOAS (Multi-Axis
Differential Optical Absorption Spectroscopy) measurements. In contrast to the data shown in sect. 3.1 and 3.2, profile com-
parison allows for assessment of the model’s capability to capture vertical distributions of NO2. MAX-DOAS measurements As a third diagnostic we compare modelled NO2 concentration profiles to profiles obtained from MAX DOAS (Multi Axis
Differential Optical Absorption Spectroscopy) measurements. In contrast to the data shown in sect. 3.1 and 3.2, profile com-
parison allows for assessment of the model’s capability to capture vertical distributions of NO2. MAX-DOAS measurements
use the DOAS principle (see Platt and Stutz, 2008) to obtain trace gas differential slant column densities (dSCDs) at different
350
elevation angles. By application of an inversion algorithm a discretized concentration vector c is obtained, whose entries de-
note the target gas concentration in different atmospheric layers. An overview of different inversion algorithms can be found in
Frieß et al. (2019). We use data from the FRM4DOAS network (see Fayt et al., 2021) which applies the Mexican MAX-DOAS
fit (MMF, see Friedrich et al., 2019) and the Mainz Profile Algorithm (MAPA, see Beirle et al., 2019). While MMF is based on use the DOAS principle (see Platt and Stutz, 2008) to obtain trace gas differential slant column densities (dSCDs) at different
350
elevation angles. By application of an inversion algorithm a discretized concentration vector c is obtained, whose entries de-
note the target gas concentration in different atmospheric layers. An overview of different inversion algorithms can be found in
Frieß et al. (2019). We use data from the FRM4DOAS network (see Fayt et al., 2021) which applies the Mexican MAX-DOAS
fit (MMF, see Friedrich et al., 2019) and the Mainz Profile Algorithm (MAPA, see Beirle et al., 2019). While MMF is based on optimal estimation (see Rodgers, 2000), MAPA uses Monte Carlo simulation in order to determine profile shape parameters
355
which combine (possibly lifted) box profiles and exponential profiles. Purely exponential concentration profiles, however, can
not be obtained from MAPA. 19 Germany
Belgium
Netherlands
Hamburg
Berlin
Munich
Rhine
river
Ruhr
region
Heidelberg
Mainz
De Bilt
Uccle
Bremen
Paris
France
Czechia
Austria
Figure 8. Part of central Europe covering Germany, Belgium and Netherlands with important geographical regions marked in red. The blue
stars show the locations of the MAX-DOAS measurements reported in sect. 3.3. Figure 8. 3.3
Comparison of NO2 concentration profiles Part of central Europe covering Germany, Belgium and Netherlands with important geographical regions marked in red. The blue
stars show the locations of the MAX-DOAS measurements reported in sect. 3.3. Five MAX-DOAS instruments are operated within our simulation domain D2: Mainz (Germany), Bremen (Germany), Hei-
delberg (Germany), De Bilt (Netherlands), and Uccle (Belgium). Figure 8 shows the locations of these stations. A single station Five MAX-DOAS instruments are operated within our simulation domain D2: Mainz (Germany), Bremen (Germany), Hei-
delberg (Germany), De Bilt (Netherlands), and Uccle (Belgium). Figure 8 shows the locations of these stations. A single station
typically yields 2-4 NO2 profile measurements per hour. In order to compare simulated and observed profiles, the WRF-Chem
360
dataset is interpolated to the geolocations and measurement times of the MAX-DOAS instruments. We use nearest-neighbour
interpolation in space and time. The uncertainty of the simulated NO2 profiles is obtained as the standard deviation of the
surrounding eight WRF-Chem grid cells. The MMF inversion algorithm provides averaging kernels (AKs), represented by a (h × The MMF inversion algorithm provides averaging kernels (AKs), represented by a (h × h) matrix, where h is the number of atmospheric layers considered. Here, h = 20, comprising retrieval altitudes up to 4 km. Using the AKs, the MAX-DOAS
365
measuring sensitivity to different altitudes can be applied to the simulated profiles. For this purpose the simulated NO2 profiles
are vertically interpolated to the layers of the MAX-DOAS retrieval grid (linear interpolation). Then, the AKs are applied via
csim, corr = Acsim + (1l −A)cap
(10) of atmospheric layers considered. Here, h = 20, comprising retrieval altitudes up to 4 km. Using the AKs, the MAX-DOAS
365
measuring sensitivity to different altitudes can be applied to the simulated profiles. For this purpose the simulated NO2 profiles
are vertically interpolated to the layers of the MAX-DOAS retrieval grid (linear interpolation). Then, the AKs are applied via
csim, corr = Acsim + (1l −A)cap
(10) (10) csim, corr = Acsim + (1l −A)cap where csim, corr is the corrected simulated profile, A the averaging kernel matrix, csim the orig where csim, corr is the corrected simulated profile, A the averaging kernel matrix, csim the original simulated profile, 1l the unity matrix, and cap the a priori profile see (see Rodgers and Connor, 2003). MAPA neither provides AKs, nor depends on an
370
a priori profile. 3.3
Comparison of NO2 concentration profiles All profiles flagged as "erroneous" by MAPA were dismissed from the evaluation. Figure 9 shows averaged NO2 profiles from the MAX-DOAS station Mainz, Germany, in the time window from 11 AM to
02 PM for a selection of individual days. The aim is to give an overview of the variety of observed and modelled profile shapes. 20 0.0
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https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 0.0
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NO2 profiles in Mainz (11 AM - 02 PM)
MAX-DOAS (MAPA)
MAX-DOAS (MMF)
A priori profile
WRF-Chem (with AK)
WRF-Chem (without AK)
DERP007 (UBA, Background)
DERP009 (UBA, Background)
DERP010 (UBA, Traffic)
DERP011 (UBA, Traffic)
Figure 9. Comparison of exemplary profiles for Mainz around noontime (11 AM - 02 PM) for four different days: (a) 03 May 2019, (b)
17 May 2019, (c) 01 May 2019, (d) 27 May 2019. Additionally, average NO2 surface concentrations measured by UBA measuring stations
within a radius of 5 km of the MAX-DOAS instrument are drawn as colored scatter markers at 0 m altitude. NO2 profiles in Mainz (11 AM - 02 PM) NO2 profiles in Mainz (11 AM - 02 PM) 0.0
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NO2 concentration [1011 molec. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure 9a shows a typical scenario: On this day (03 May 2019) observation and simulation show good general agreement. All three datasets (MMF, MAPA, and WRF-Chem) yield profiles of similar shape. Above an altitude of ∼2 km the profiles
quickly approach zero, which is characteristic for the transitioning regime between the planetary boundary layer and the free
troposphere. At low altitudes of 0-150 m the simulated profile has a strong exponential tail, typical for the surface layer into
which most NOx emissions are injected (the lowest ∼30 m). Furthermore, the simulated profiles show good agreement with
380
the measurements of the nearby UBA in-situ stations. High vertical resolution turns out to be crucial for the comparison to the
observed surface concentrations: In the FRM4DOAS dataset, MAPA and MMF were operated with a vertical resolution of 200
m. In a direct comparison, the concentrations obtained from MAPA or MMF in the lowest layer (0-200 m) underestimate the
in-situ observations at the surface because of the limited vertical resolution of the MAX-DOAS measurements and the rather coarse retrieval grid used by FRM4DOAS. Meanwhile, the lowest layer of our WRF-Chem simulation only spans 0 - 8 m,
385
which allows for a much more representative comparison. Subfigure 9b shows a day on which an elevated NO2 layer was detected by MMF. This occurs regularly at all MAX-DOAS
stations in this study and is characterized by strongly enhanced NO2 concentrations at higher altitudes (here at ∼500 m). Ele-
vated layers are typically caused by elevated emissions, e.g. from a power plant stack at a few hundred meters height. Addition- coarse retrieval grid used by FRM4DOAS. Meanwhile, the lowest layer of our WRF-Chem simulation only spans 0 - 8 m,
385
which allows for a much more representative comparison. Subfigure 9b shows a day on which an elevated NO2 layer was detected by MMF. This occurs regularly at all MAX-DOAS
stations in this study and is characterized by strongly enhanced NO2 concentrations at higher altitudes (here at ∼500 m). Ele-
vated layers are typically caused by elevated emissions, e.g. from a power plant stack at a few hundred meters height. Addition- ally, transport events which advect NO2 from the surface layer could be the cause. However, no corresponding enhancements
390
can be seen in the simulated NO2 profile. As described in sect. 3.3
Comparison of NO2 concentration profiles cm
3]
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(c)
2019-05-01
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2019-05-27 (c) Altitude [km] NO2 concentration [1011 molec. cm
3] Figure 9. Comparison of exemplary profiles for Mainz around noontime (11 AM - 02 PM) for four different days: (a) 03 May 2019, (b)
17 May 2019, (c) 01 May 2019, (d) 27 May 2019. Additionally, average NO2 surface concentrations measured by UBA measuring stations
within a radius of 5 km of the MAX-DOAS instrument are drawn as colored scatter markers at 0 m altitude. Additionally, the average NO2 surface concentrations measured by UBA stations within a radius of 5 km of the MAX-DOAS
instrument are drawn as colored scatter markers at 0 m altitude. 375 Additionally, the average NO2 surface concentrations measured by UBA stations within a radius of 5 km of the MAX-DOAS
instrument are drawn as colored scatter markers at 0 m altitude. 375 375 21 2.1, we do not use vertical emission profiles in the WRF-Chem
simulation. Furthermore, it is possible that the overall spatial resolution of our simulation limits its ability to model compa-
rable elevated trace gas abundances. MAX-DOAS instruments typically have a field of view of ∼1◦, which is small enough
to resolve e.g. plumes from power plant stacks. Our WRF-Chem simulation has a spatial resolution of 3 km × 3 km × 100 m ally, transport events which advect NO2 from the surface layer could be the cause. However, no corresponding enhancements
390
can be seen in the simulated NO2 profile. As described in sect. 2.1, we do not use vertical emission profiles in the WRF-Chem
simulation. Furthermore, it is possible that the overall spatial resolution of our simulation limits its ability to model compa-
rable elevated trace gas abundances. MAX-DOAS instruments typically have a field of view of ∼1◦, which is small enough
to resolve e.g. plumes from power plant stacks. Our WRF-Chem simulation has a spatial resolution of 3 km × 3 km × 100 m at ∼500 m altitude and therefore dilutes trace gas concentrations into comparably large grid cell volumes. Lastly, horizontal
395
concentration gradients can be expected near strong emission sources. These can lead to apparently elevated profiles, because
the MAX-DOAS profile inversion makes the simplifying assumption of horizontally homogeneous distributions. Comparison
of simulated and observed elevated urban emissions is therefore an advanced problem and not further adressed here. Subfigure 9c shows a day on which the agreement between model and observations was poor, with deviations of > 100 %. at ∼500 m altitude and therefore dilutes trace gas concentrations into comparably large grid cell volumes. Lastly, horizontal
395
concentration gradients can be expected near strong emission sources. These can lead to apparently elevated profiles, because
the MAX-DOAS profile inversion makes the simplifying assumption of horizontally homogeneous distributions. Comparison
of simulated and observed elevated urban emissions is therefore an advanced problem and not further adressed here. Subfigure 9c shows a day on which the agreement between model and observations was poor, with deviations of > 100 %. g
y
g
p
,
Such days are outliers and do not represent the overall quality of the model or the measurements, however, they should not
400
remain unmentioned. Possible reasons for such deviations include e.g. falsely modelled wind directions (affecting trace gas
transport) or cloud cover (affecting photolysis). The scatter plots (subfigures b, d, f) show a strong correlation between simulation and measurements Figure 10 shows the results of the three German MAX-DOAS locations Mainz, Bremen, and Heidelberg. Overall, the three
stations show qualitatively similar profiles. At Bremen (see subfigures c-d), a consistent elevated layer at ∼1000 m altitude
was detected, which could be explained by multiple power plants in the near vicinity of the instrument. This has been discussed
in detail by Bösch (2018). The scatter plots (subfigures b, d, f) show a strong correlation between simulation and measurements and yield linear fits with slopes of 1.32 for Mainz, 1.15 for Bremen, and 0.97 for Heidelberg. The results shown in Fig. 11
420
show similar agreement in De Bilt but larger deviations in Uccle. As a result, the slope of 1.58 for the linear fit in Fig. 11d is
significantly larger than for the other stations. Table 5 gives a statistical summary of the results shown in Fig. 10 and Fig. 11. As an additional diagnostic, a correlation analysis between observed (MMF) and simulated (with AKs applied) NO2 con-
centrations in the lowest 0-200 m above the surface is shown in Appendix A (see Fig. A4). Subfigure 9d shows a day on which the modelled profile has an extremely steep exponential gradient in combination with
a relatively thin boundary layer. At the transition from the boundary layer to the surface layer (∼30 - 50 m above ground), Such days are outliers and do not represent the overall quality of the model or the measurements, however, they should not
400
remain unmentioned. Possible reasons for such deviations include e.g. falsely modelled wind directions (affecting trace gas
transport) or cloud cover (affecting photolysis). Subfigure 9d shows a day on which the modelled profile has an extremely steep exponential gradient in combination with
a relatively thin boundary layer. At the transition from the boundary layer to the surface layer (∼30 - 50 m above ground), a relatively thin boundary layer. At the transition from the boundary layer to the surface layer (
30
50 m above ground),
the simulated profiles shows an increase of almost 400 %. Interestingly, on this day (27 May 2019), one of the background
405
stations (DERP009) measured more than twice the surface concentration compared to a nearby traffic station (DERP011). This
phenomenon was observed on only one other day (10 May 2019) of the simulated month, on which very similar profile shapes
were observed. Furthermore, no MAPA profile is available on this day due to the filtering by error flags. Also, there is good
agreement between modelled and observed profiles above the surface layer. It is therefore plausible to assume that the steep the simulated profiles shows an increase of almost 400 %. Interestingly, on this day (27 May 2019), one of the background
405
stations (DERP009) measured more than twice the surface concentration compared to a nearby traffic station (DERP011). This
phenomenon was observed on only one other day (10 May 2019) of the simulated month, on which very similar profile shapes
were observed. Furthermore, no MAPA profile is available on this day due to the filtering by error flags. Also, there is good
agreement between modelled and observed profiles above the surface layer. It is therefore plausible to assume that the steep 22 22 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Table 5. Statistical summary of the results shown in Fig. 10 and 11 Location
Mean (sim.)
Mean (obs.)
Slope
Intercept
RMSE
Bias
Mainz
3.05
2.03
1.32
0.38
1.35
+50.3 %
Bremen
1.83
1.94
1.15
-0.41
1.42
-5.9 %
Heidelberg
2.57
1.98
0.97
0.66
0.93
+30.1 %
De Bilt
2.40
2.05
1.16
0.02
0.54
+16.9 %
Uccle
4.11
2.30
1.58
0.48
2.44
+78.6 %
Means, intercept, and RMSE are given in units of [1010 molec. cm−3]. NO2 gradient towards the surface is not due to a faulty model, but, on the contrary, indicates the model’s ability to capture
410
scenarios in which e.g. vertical mixing is mostly suppressed and large portions of the emitted NO2 remain in the surface layer. In order to condense the remaining evaluation, we will focus on monthly noontime averages from hereon, i.e. plots of the
same structure as shown in Fig. 9, but averaged over the entire simulation period of May 2019 (Fig. 10 and 11). In both figures
the right-side panel shows scatter plots of averaged NO2 concentrations (WRF-Chem vs. MMF, due to availability of AKs) at
diff
t ltit d
415 NO2 gradient towards the surface is not due to a faulty model, but, on the contrary, indicates the model’s ability to capture
410
scenarios in which e.g. vertical mixing is mostly suppressed and large portions of the emitted NO2 remain in the surface layer. In order to condense the remaining evaluation, we will focus on monthly noontime averages from hereon, i.e. plots of the
same structure as shown in Fig. 9, but averaged over the entire simulation period of May 2019 (Fig. 10 and 11). In both figures
the right-side panel shows scatter plots of averaged NO2 concentrations (WRF-Chem vs. MMF, due to availability of AKs) at different altitudes. 415
Figure 10 shows the results of the three German MAX-DOAS locations Mainz, Bremen, and Heidelberg. Overall, the three
stations show qualitatively similar profiles. At Bremen (see subfigures c-d), a consistent elevated layer at ∼1000 m altitude
was detected, which could be explained by multiple power plants in the near vicinity of the instrument. This has been discussed
in detail by Bösch (2018). 4
Discussion and conclusion
425 We have presented new WRF-Chem simulation results with specific focus on NO2. The simulation was run on a domain over
central Europe with a spatial resolution of 3 km×3 km for the month of May 2019. Over Germany, a new emission inventory
from the German Environmental Agency with a resolution of 1 km × 1 km was used. Outside of Germany, the EDGARv5
emission inventory with a resolution of 11 km×7 km was used. A new set of temporal emission profiles for the most important emission sectors (traffic, power industry, agricultural soils, energy for buildings, and manufacturing industry) was determined
430
by comparing simulated NOx surface concentrations to corresponding in-situ measurements in Germany. Once good agreement
was found, the model results were also compared to satellite measurements of the NO2 VCD and NO2 concentration profiles emission sectors (traffic, power industry, agricultural soils, energy for buildings, and manufacturing industry) was determined
430
by comparing simulated NOx surface concentrations to corresponding in-situ measurements in Germany. Once good agreement
was found, the model results were also compared to satellite measurements of the NO2 VCD and NO2 concentration profiles 23 0
1
2
3
4
Altitude [km]
(MMF)
17.5 cm * 10.8156 cm (double col. Fig., golden ratio)
0
1
2
3
4
Altitude [km]
MAX-DOAS (MAPA)
MAX-DOAS (MMF)
WRF-Chem (with AK)
WRF-Chem (without AK)
Simulated NO2 concentration (with AK)
UBA stations
Figure 10. Comparison of averaged NO2 profiles for Mainz, Bremen, and Heidelberg around noontime (11 AM - 02 PM). The left column
(a), (c), (e) shows the NO2 profiles as obtained by MMF, MAPA, and WRF-Chem. The right column (b), (d), (f) shows the corresponding
scatter plots of averaged NO2 concentrations at different altitudes (here: WRF-Chem vs. MMF). imulated NO2 concentration (with AK) Figure 10. Comparison of averaged NO2 profiles for Mainz, Bremen, and Heidelberg around noontime (11 AM - 02 PM). The left column
(a), (c), (e) shows the NO2 profiles as obtained by MMF, MAPA, and WRF-Chem. The right column (b), (d), (f) shows the corresponding
scatter plots of averaged NO2 concentrations at different altitudes (here: WRF-Chem vs. MMF). 24 0
1
2
3
4
Altitude [km]
0
1
2
3
4
Altitude [km]
MAX-DOAS (MAPA)
MAX-DOAS (MMF)
WRF-Chem (with AK)
WRF-Chem (without AK)
Simulated NO2 concentration (with AK)
(MMF)
Figure 11. Like Fig. 10, but for the stations De Bilt and Uccle. 4
Discussion and conclusion
425 Simulated NO2 concentration (with AK)
(MMF) mulated NO2 concentration (with AK) (MMF) Figure 11. Like Fig. 10, but for the stations De Bilt and Uccle. obtained from MAX-DOAS measurements. The main focus of the study was to investigate, whether RCT simulations can be
improved by fine-tuning of the temporal emission profiles and whether good agreement with all three observational datasets
can be achieved. 5 obtained from MAX-DOAS measurements. The main focus of the study was to investigate, whether RCT simulations can be
improved by fine-tuning of the temporal emission profiles and whether good agreement with all three observational datasets obtained from MAX-DOAS measurements. The main focus of the study was to investigate, whether RCT simulations can be
improved by fine-tuning of the temporal emission profiles and whether good agreement with all three observational datasets can be achieved. 435
In sect. 3.1 modelled surface concentrations of NO2, NO, NOx and O3 were compared to in-situ measurements. Here, we
used simulation results of two WRF-Chem runs: Firstly, a run with conventional temporal profiles used in previously pub-
lished literature, and secondly, a run with our newly optimized temporal profiles. The optimized simulation showed significant
improvements over the conventional simulation. In particular, the diurnal variation of trace gas concentration was much more accurately reproduced using the new temporal profiles. NOx biases were reduced from +23.4 % to +6.7 % (all hours of the day
440
considered) and from -19.1 % to +0.4 % (at noontime). NO2 biases were reduced from +34.0 % to +14.3 % (all hours of the day
considered) and from -15.7 % to +0.9 % (at noontime). The corresponding correlation coefficients also slightly improved. This
way, the noon-time underestimations of NOx and NO2 could be resolved without any changes to the total emission budget. 25 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Underestimations like these were identified in recent literature, but have remained an open issue so far. Despite the significant
improvements in NOx and NO2 modelling accuracy, some discrepancies remain: While NOx concentrations appear to be ac-
445
curate throughout the day, the NO2/NO ratio is too large before and after noontime. It was found that this problem can not be
resolved by changing the speciation ratio of NOx, which can be explained by the rapid conversion of NO to NO2 directly after
emission. It is therefore more likely that this issue relates to chemistry: Both O3 and VOCs oxidize NO to NO2, and a slight
model overestimation of O3 (+10.9 %) was found. Additionally, O3 concentrations are influenced by VOCs. However, the reference data includes no measurements of VOCs, which means no direct validation of the modelled VOC concentrations is
450
possible. If such measurements become available in the future, the methods described in this article can be extended to further
improve the simulation results. In sect. 3.2 modelled tropospheric NO2 VCDs were compared to TROPOMI measurements. In order to make a representative
comparison the air mass factors of the TROPOMI retrieval were recomputed using the high-resolution NO2 profiles from our possible. If such measurements become available in the future, the methods described in this article can be extended to further
improve the simulation results. In sect. 3.2 modelled tropospheric NO2 VCDs were compared to TROPOMI measurements. In order to make a representative
comparison, the air mass factors of the TROPOMI retrieval were recomputed using the high-resolution NO2 profiles from our simulation. The distribution of modelled and observed NO2 VCDs was found to be similar and agrees with previously reported
455
simulation results. Monthly means of simulated and observed VCDs were shown to correlate strongly (R = 0.84) with a model
bias of −6.6%. Recomputing the air mass factors using the NO2 profiles from the regional model was found to be crucial
for good agreement: With the original air mass factors the model bias was much larger (+17.6 %). Comparison between the
spatial distribution of NO2 VCDs and surface concentration revealed that in regions where the simulation overestimated the simulation. The distribution of modelled and observed NO2 VCDs was found to be similar and agrees with previously reported
455
simulation results. Monthly means of simulated and observed VCDs were shown to correlate strongly (R = 0.84) with a model
bias of −6.6%. Recomputing the air mass factors using the NO2 profiles from the regional model was found to be crucial
for good agreement: With the original air mass factors the model bias was much larger (+17.6 %). Comparison between the
spatial distribution of NO2 VCDs and surface concentration revealed that in regions where the simulation overestimated the VCDs, it also tended to overestimate the surface concentrations. This was observed in the strongly polluted regions of west
460
Germany and the Netherlands. On the other hand, a general tendency for model underestimation in less polluted regions was
found. This observation hints towards faulty NOx emissions in the corresponding regions. While surface concentrations were
shown to be highly sensitive to the choice of temporal profiles, only a weak dependence was found for the NO2 VCDs. In a
comparison to the simulation run with conventional temporal profiles, NO2 VCDs were only +3.6 % larger on average. The VCDs, it also tended to overestimate the surface concentrations. This was observed in the strongly polluted regions of west
460
Germany and the Netherlands. On the other hand, a general tendency for model underestimation in less polluted regions was
found. This observation hints towards faulty NOx emissions in the corresponding regions. While surface concentrations were
shown to be highly sensitive to the choice of temporal profiles, only a weak dependence was found for the NO2 VCDs. In a
comparison to the simulation run with conventional temporal profiles, NO2 VCDs were only +3.6 % larger on average. The surface concentrations, on the other hand, were 16.6 % larger. 465
In sect. 3.3 modelled noontime NO2 concentration profiles were compared to profiles from MAX-DOAS measurements at
5 locations in central Europe from the FRM4DOAS network. The inversion algorithms MMF and MAPA were used. General
agreement in the overall shape and magnitude of the profiles was found, with per-station concentration biases of +50.3 %
(Mainz), −5.9 % (Bremen), +30.1 % (Heidelberg), +16.9 % (De Bilt) and +78.6 % (Uccle). Two qualitative differences surface concentrations, on the other hand, were 16.6 % larger. 465
In sect. 3.3 modelled noontime NO2 concentration profiles were compared to profiles from MAX-DOAS measurements at
5 locations in central Europe from the FRM4DOAS network. The inversion algorithms MMF and MAPA were used. General
agreement in the overall shape and magnitude of the profiles was found, with per-station concentration biases of +50.3 %
(Mainz), −5.9 % (Bremen), +30.1 % (Heidelberg), +16.9 % (De Bilt) and +78.6 % (Uccle). Two qualitative differences between observed and simulated profiles (as produced by WRF-Chem, i.e. without application of averaging kernels) were
470
identified: Firstly, the simulated profiles show steep NO2 gradients close to the surface and, furthermore, agree well with
the collocated surface concentration measurements. The observed profiles do not resolve the surface layer, most likely due to
the coarse retrieval grid with 200 m layer height. As a consequence, the observed profiles tend to underestimate the surface
concentration measurements. Secondly, elevated NO2 layers were identified in some of the observed profiles, but not in the between observed and simulated profiles (as produced by WRF-Chem, i.e. without application of averaging kernels) were
470
identified: Firstly, the simulated profiles show steep NO2 gradients close to the surface and, furthermore, agree well with
the collocated surface concentration measurements. The observed profiles do not resolve the surface layer, most likely due to
the coarse retrieval grid with 200 m layer height. As a consequence, the observed profiles tend to underestimate the surface
concentration measurements. Secondly, elevated NO2 layers were identified in some of the observed profiles, but not in the corresponding simulated profiles. Here, the lack of vertical emission profiles, the overall spatial resolution of our WRF-Chem
475
simulation, and the influence of horizontal concentration gradients on the MAX-DOAS inversion procedure were identified as
possible explanations. corresponding simulated profiles. Here, the lack of vertical emission profiles, the overall spatial resolution of our WRF-Chem
475
simulation, and the influence of horizontal concentration gradients on the MAX-DOAS inversion procedure were identified as
possible explanations. 26 profiles can be applied wherever suitable observational data is available, making the method viable for large parts of the world
490
and other seasons. In the future, our method of deriving temporal profiles could be made more rigorous by experimenting with
modern optimization algorithms. This will take time and require the synergy of both, modelling and inversion expertise, but
has the potential to make our method applicable to larger domains and simulation periods with less effort. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. We draw two main conclusions: Firstly, our study shows that the model accuracy for NOx and NO2 in RCT simulations can
be strongly improved by optimizing the corresponding hourly emission profiles. In particular, the typical underestimations of
noontime NOx and NO2 can be resolved. We have described a flexible manual approach for temporal profile tuning, which
480
only requires an iterative chain of model forward runs and comparison to corresponding in-situ observations. Secondly, the
intercomparison of the observational datasets used for model validation (here: in-situ measurements, satellite observations, and
MAX-DOAS measurements) are overall consistent and agree with our simulation results. It is important to emphazise the scope of our study. We have shown exemplary temporal profiles and simulation results
which only represent central Europe in mid-summer. Influences such as regulations for industrial processes, changes in the
485
prevailing energy sources, or reduction of cars as a primary means of personal transport may impact the hourly profiles in the
future. Furthermore, it is likely that the hourly profiles inhibit a seasonal dependence. Therefore, the temporal profiles we have
presented here should not be seen as generally valid, but rather as a specific result obtained in the framework of our case study
for the purpose of proving how sensitive simulation results are to them. However, our general procedure of finding temporal profiles can be applied wherever suitable observational data is available, making the method viable for large parts of the world
490
and other seasons. In the future, our method of deriving temporal profiles could be made more rigorous by experimenting with
modern optimization algorithms. This will take time and require the synergy of both, modelling and inversion expertise, but
has the potential to make our method applicable to larger domains and simulation periods with less effort. profiles can be applied wherever suitable observational data is available, making the method viable for large parts of the world
490
and other seasons. In the future, our method of deriving temporal profiles could be made more rigorous by experimenting with
modern optimization algorithms. This will take time and require the synergy of both, modelling and inversion expertise, but
has the potential to make our method applicable to larger domains and simulation periods with less effort. Data availability. All data are available from the authors upon request. Data availability. All data are available from the authors upon request. Author contributions. LK, TW, SB, and VK developed the question of research. LK and SO carried out the optimization process for the
495
temporal emission profiles. VK, LK, SO, and RK conducted the WRF-Chem simulations. LK, VK, TW, SO, SB, AP, and TB made the
comparison between simulation and observational datasets. LK wrote the paper, with all authors contributing by revising it interactively. Author contributions. LK, TW, SB, and VK developed the question of research. LK and SO carried out the optimization process for the
495
temporal emission profiles. VK, LK, SO, and RK conducted the WRF-Chem simulations. LK, VK, TW, SO, SB, AP, and TB made the
comparison between simulation and observational datasets. LK wrote the paper, with all authors contributing by revising it interactively. Competing interests. The authors declare that they have no conflict of interest. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2022-1473 https://doi.org/10.5194/egusphere-2022-1473
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retrievals: description and validation, Geoscientific Model Development, 10, 721–750, https://doi.org/10.5194/gmd-10-721-2017, 2017. 675 675 retrievals: description and validation, Geoscientific Model Development, 10, 721–750, https://doi.org/10.5194/gmd-10-721-2017, 2017. 675 32 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Appendix A: Additional figures and tables 33 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Table A1. Layer extent of the lowest 24 layers in our WRF-Chem simulation Layer number
Layer bottom [m]
Layer top [m]
Layer number
Layer bottom [m]
Layer top [m]
1
0
8
13
1370
1546
2
8
33
14
1546
1697
3
33
66
15
1697
1841
4
66
125
16
1841
1937
5
125
209
17
1937
2035
6
209
310
18
2035
2183
7
310
429
19
2183
2374
8
429
575
20
2374
2661
9
575
741
21
2661
3142
10
741
935
22
3142
3907
11
935
1178
23
3907
4762
12
1178
1370
24
4762
5643
Exact layer bottoms and tops depend on location and time The values given here are averages r number
Layer bottom [m]
Layer top [m]
Layer number
Layer bottom [m]
Layer top [m] Table A2. Statistical summary of the results shown in Fig. A4 Table A2. Statistical summary of the results shown in Fig. A4 Location
Mean (sim.)
Mean (obs.)
Slope
Intercept
RMSE
Bias
R
Mainz
11.69
7.18
0.07
11.15
10.42
+62.8 %
0.13
Bremen
9.09
4.68
0.64
6.12
8.63
+94.3 %
0.29
Heidelberg
8.21
9.60
0.15
6.78
17.06
-14.5 %
0.38
De Bilt
8.62
6.71
0.69
3.97
6.31
+28.5 %
0.56
Uccle
17.69
11.24
0.57
11.32
13.35
+57.3 %
0.45
Means, intercept, and RMSE are given in units of [1010 molec. cm−3]. 34 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 0
10
20
30
40
50
60
NO2 conc. [µg m
3]
(a)
(a)
0
10
20
30
40
50
60
(e)
(e)
Monthly average
0
20
40
60
80
100
NO conc. [µg m
3]
(b)
(b)
0
20
40
60
80
100
(f)
(f)
120
140
3]
(c)
(c)
120
140
(g)
(g)
UBA measurements
S-new
S-old 0
10
20
30
40
50
60
NO2 conc. [µg m
3]
(a)
(a)
0
10
20
30
40
50
60
(e)
(e)
Monthly average
0
20
40
60
80
100
NO conc. [µg m
3]
(b)
(b)
0
20
40
60
80
100
(f)
(f)
0
20
40
60
80
100
120
140
NOx conc. [µg m
3]
(c)
(c)
0
20
40
60
80
100
120
140
(g)
(g)
Wed, 05-01 00:00
Thu, 05-02 00:00
Fri, 05-03 00:00
Sat, 05-04 00:00
Sun, 05-05 00:00
Mon, 05-06 00:00
Tue, 05-07 00:00
Wed, 05-08 00:00
Thu, 05-09 00:00
Fri, 05-10 00:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
Date and time
25
50
75
100
125
O3 conc. [µg m
3]
(d)
00:00
04:00
08:00
12:00
16:00
20:00
00:00
Time of day
25
50
75
100
125
(h)
UBA measurements
S-new
S-old 0
10
20
30
40
50
60
NO2 conc. [µg m
3]
(a)
(a)
0
10
20
30
40
50
60
(e)
(e)
Monthly average
0
20
40
60
80
100
NO conc. [µg m
3]
(b)
(b)
0
20
40
60
80
100
(f)
(f)
0
20
40
60
80
100
120
140
NOx conc. Table A2. Statistical summary of the results shown in Fig. A4 [µg m
3]
(c)
(c)
0
20
40
60
80
100
120
140
(g)
(g)
Wed, 05-01 00:00
Thu, 05-02 00:00
Fri, 05-03 00:00
Sat, 05-04 00:00
Sun, 05-05 00:00
Mon, 05-06 00:00
Tue, 05-07 00:00
Wed, 05-08 00:00
Thu, 05-09 00:00
Fri, 05-10 00:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
12:00
Date and time
25
50
75
100
125
O3 conc. [µg m
3]
(d)
00:00
04:00
08:00
12:00
16:00
20:00
00:00
Time of day
25
50
75
100
125
(h)
UBA measurements
S-new
S-old
Figure A1. Like Fig. 3, but for traffic stations instead of background stations. Figure A1. Like Fig. 3, but for traffic stations instead of background stations. 35 0
5
10
15
20
25
30
0
10
20
30
Simulated mass
concentration [µg m
3]
(a)
(a)
(a)
(a)
(a)
(a)
(a)
(a)
NO2 (S-new)
f(x) = 0.88x + 3.18
0
5
10
15
20
25
30
0
10
20
30
(b)
(b)
(b)
(b)
(b)
(b)
(b)
(b)
NO2 (S-old)
f(x) = 1.15x + 2.26
0.0
2.5
5.0
7.5
10.0
12.5
15.0
17.5
0
5
10
15
20
Simulated mass
concentration [µg m
3]
(c)
(c)
(c)
(c)
(c)
(c)
(c)
(c)
NO (S-new)
f(x) = 0.46x + 0.58
0.0
2.5
5.0
7.5
10.0
12.5
15.0
17.5
0
5
10
15
20
(d)
(d)
(d)
(d)
(d)
(d)
(d)
(d)
NO (S-old)
f(x) = 0.41x + 0.76
0
10
20
30
40
0
20
40
Simulated mass
concentration [µg m
3]
(e)
(e)
(e)
(e)
(e)
(e)
(e)
(e)
NOx (S-new)
f(x) = 0.71x + 5.17
0
10
20
30
40
0
20
40
(f)
(f)
(f)
(f)
(f)
(f)
(f)
(f)
NOx (S-old)
f(x) = 0.82x + 5.92
0
20
40
60
80
100
120
Observed mass concentration [µg m
3]
0
50
100
150
Simulated mass
concentration [µg m
3]
(g)
(g)
(g)
(g)
(g)
(g)
(g)
(g)
O3 (S-new)
f(x) = 0.74x + 24.35
0
20
40
60
80
100
120
Observed mass concentration [µg m
3]
0
50
100
150
(h)
(h)
(h)
(h)
(h)
(h)
(h)
(h)
O3 (S-old)
f(x) = 0.82x + 24.28
Figure A2. Like Fig. 4, but for all hours of the day instead for noontime only. Error bars were omitted and a color coding (without physical
meaning) applied for easier readability. Table A2. Statistical summary of the results shown in Fig. A4 Simulated mass
3 (h)
(h)
(h)
(h)
(h)
(h)
(h)
(h) Simulated mass
3 Figure A2. Like Fig. 4, but for all hours of the day instead for noontime only. Error bars were omitted and a color coding (without physical
meaning) applied for easier readability. 36 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. [1010 molec cm-3]
Figure A3. Average simulated NO2 profile, vertically interpolated to the TM5 grid. Figure A3. Average simulated NO2 profile, vertically interpolated to the TM5 grid. 37 37 https://doi.org/10.5194/egusphere-2022-1473
Preprint. Discussion started: 6 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. 0
1
2
3
4
5
(a)
Mainz
(b)
Bremen
0
1
2
3
4
5
NO2 concentration (WRF-Chem with AK, 0-200 m) [1011 molec. cm
3]
(c)
Heidelberg
0
1
2
3
4
5
NO2 concentration (MMF, 0-200 m) [1011 molec. cm
3]
(d)
De Bilt
0
1
2
3
4
5
NO2 concentration (MMF, 0-200 m) [1011 molec. cm
3]
0
1
2
3
4
5
(e)
Uccle
Figure A4. Correlation analysis between observed (MMF) and simulated (averaging kernels applied) NO2 concentrations in the lowes
0-200 m above the surface. In each subplot, the dashed line marks the 1:1 line and the solid line is a linear fit through the point cloud. The
R-values and further statistical diagnostics of each plot are found in Table A2. Bremen De Bilt NO2 concentration (MMF, 0-200 m) [1011 molec. cm
3] NO2 concentration (MMF, 0-200 m) [1011 molec. cm
3] NO2 concentration (MMF, 0-200 m) [1011 molec. cm
3] Figure A4. Correlation analysis between observed (MMF) and simulated (averaging kernels applied) NO2 concentrations in the lowest
0-200 m above the surface. In each subplot, the dashed line marks the 1:1 line and the solid line is a linear fit through the point cloud. The
R-values and further statistical diagnostics of each plot are found in Table A2. Figure A4. Correlation analysis between observed (MMF) and simulated (averaging kernels applied) NO2 concentrations in the lowest
0-200 m above the surface. In each subplot, the dashed line marks the 1:1 line and the solid line is a linear fit through the point cloud. The
R-values and further statistical diagnostics of each plot are found in Table A2. 38
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Using Natural Language Processing to Explore Mental Health Insights From UK Tweets During the COVID-19 Pandemic: Infodemiology Study (Preprint)
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Corresponding Author: Email: chris.marshall@io.nihr.ac.uk (JMIR Infodemiology 2022;2(1):e32449) doi: 10.2196/32449 (JMIR Infodemiology 2022;2(1):e32449) doi: 10.2196/32449 Using Natural Language Processing to Explore Mental Health
Insights From UK Tweets During the COVID-19 Pandemic:
Infodemiology Study Christopher Marshall, BSc, PhD; Kate Lanyi, BSc, MRes; Rhiannon Green, BSc, MRes; Georgina C Wilkins, BSc,
PhD; Fiona Pearson, BSc, MRes, PhD; Dawn Craig, BSc, MSc National Institute for Health Research Innovation Observatory, Newcastle University, Newcastle, United Kingdom National Institute for Health Research Innovation Observatory, Newcastle University, Newcas Corresponding Author:
Christopher Marshall, BSc, PhD
National Institute for Health Research Innovation Observatory
Newcastle University
The Catalyst
Newcastle, NE4 5TG
United Kingdom
Phone: 44 0191 2082259
Email: chris.marshall@io.nihr.ac.uk Abstract Background: There is need to consider the value of soft intelligence, leveraged using accessible natural language processing
(NLP) tools, as a source of analyzed evidence to support public health research outputs and decision-making. Objective: The aim of this study was to explore the value of soft intelligence analyzed using NLP. As a case study, we selected
and used a commercially available NLP platform to identify, collect, and interrogate a large collection of UK tweets relating to
mental health during the COVID-19 pandemic. Methods: A search strategy comprised of a list of terms related to mental health, COVID-19, and lockdown restrictions was
developed to prospectively collate relevant tweets via Twitter’s advanced search application programming interface over a 24-week
period. We deployed a readily and commercially available NLP platform to explore tweet frequency and sentiment across the
United Kingdom and identify key topics of discussion. A series of keyword filters were used to clean the initial data retrieved
and also set up to track specific mental health problems. All collated tweets were anonymized. Results: We identified and analyzed 286,902 tweets posted from UK user accounts from July 23, 2020 to January 6, 2021. The
average sentiment score was 50%, suggesting overall neutral sentiment across all tweets over the study period. Major fluctuations
in volume (between 12,622 and 51,340) and sentiment (between 25% and 49%) appeared to coincide with key changes to any
local and/or national social distancing measures. Tweets around mental health were polarizing, discussed with both positive and
negative sentiment. Key topics of consistent discussion over the study period included the impact of the pandemic on people’s
mental health (both positively and negatively), fear and anxiety over lockdowns, and anger and mistrust toward the government. Conclusions: Using an NLP platform, we were able to rapidly mine and analyze emerging health-related insights from UK
tweets into how the pandemic may be impacting people’s mental health and well-being. This type of real-time analyzed evidence
could act as a useful intelligence source that agencies, local leaders, and health care decision makers can potentially draw from,
particularly during a health crisis. (JMIR Infodemiology 2022;2(1):e32449) doi: 10.2196/32449 Marshall et al Marshall et al JMIR INFODEMIOLOGY JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 1
(page number not for citation purposes) KEYWORDS Twitter; mental health; COVID-19; sentiment; lockdown; soft intelligence; artificial intelligence; machine learning; natural
language processing https://infodemiology.jmir.org/2022/1/e32449 https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
RenderX JMIR INFODEMIOLOGY Marshall et al applying NLP can aid health researchers in gaining insights
from large, unstructured data sets, such as Twitter. However,
the true value of this type of work, including the data set itself,
analysis methods, and how it might be integrated into more
formal public health research outputs, is still uncertain. For
example, a lot of previous work so far has focused on the use
of internally developed, bespoke tools or packages, which tend
to require a certain level of technical expertise around machine
learning (ML) in order operate effectively. However, as methods
continue to mature, we are seeing a growing number of
“off-the-shelf” solutions become available, which appear to be
more accessible and require less technical understanding of the
underlying ML concepts. AND mental health OR anxiety OR depression OR anxious OR depressed OR depressing OR trauma OR traumatic OR “obsessive compulsive disorder”
OR OCD OR vulnerable OR loneliness OR lonely OR isolated OR isolation OR sleep OR stress OR stressful OR self-harm OR self-harming OR
suicide OR suicidal OR well-being mental health OR anxiety OR depression OR anxious OR depressed OR depressing OR trauma OR traumatic OR “obsessive compulsive disorder”
OR OCD OR vulnerable OR loneliness OR lonely OR isolated OR isolation OR sleep OR stress OR stressful OR self-harm OR self-harming OR
suicide OR suicidal OR well-being Search terms were identified through discussion within the
research team and scanning recent literature around mental
health. Once the strategy had been agreed upon, it was reviewed
by a topic expert and information specialist. We then began
prospectively searching for and scraping relevant tweets using
Twitter’s advanced search application interface [15]. A
geolocation filter was applied to the search strategy to limit the
collection of tweets to those posted in the United Kingdom only. Search terms were identified through discussion within the
research team and scanning recent literature around mental
health. Once the strategy had been agreed upon, it was reviewed
by a topic expert and information specialist. We then began
prospectively searching for and scraping relevant tweets using
Twitter’s advanced search application interface [15]. A
geolocation filter was applied to the search strategy to limit the
collection of tweets to those posted in the United Kingdom only. In this article, we report the findings from our analysis of
relevant tweets in the United Kingdom collected over a 24-week
period, from July 23, 2020 to January 6, 2021. Introduction COVID-19 was identified as a new type of coronavirus in early
January 2020 [1]. Since then, the disease has rapidly spread to
and affected almost all parts of the world. In the United
Kingdom, the first outbreak was reported on January 31, 2020,
with a national lockdown following on March 26, 2020. Shortly
before this, COVID-19 was declared a global pandemic by the
World Health Organization (WHO) on March 11, 2020 [2,3]. The COVID-19 pandemic continues to have a profound effect
on mental health [4]. In a key position paper published in June
2020, the authors explored the current and future potential
psychological, social, and neuroscientific effects of COVID-19
and set out a series of priorities and longer-term strategies for
mental health research [4]. One of the immediate research
priorities presented in the paper was “surveillance.” In particular,
the authors suggested that finding useful ways to monitor and
analyze data on the mental health effects of the COVID-19
pandemic across the whole population, as well as vulnerable
subgroups, was essential [4]. The aim of this study was to further explore the value of soft
intelligence as a meaningful source of evidence, which, when
analyzed using an accessible NLP platform, can support public
health research activity. In this article, we report the findings
from a case study that examined a large collection of tweets
relating to mental health posted from the United Kingdom during
the COVID-19 pandemic. With over 300 million active monthly users, Twitter is one of
the most popular social media platforms available. Twitter is a
free microblogging service that enables its users to post, read,
and respond to each other’s “tweets” (ie, short messages limited
to 280 characters). Social media data are being increasingly
used as a data source to inform health-related research, with the
potential for offering a more efficient means of data collection
over traditional, time-consuming, and costly survey-based
methods [5]. In particular, Twitter has been used to monitor,
track trends, and disseminate health information during past
viral pandemics [6-9]. Further, previous studies have
successfully leveraged Twitter data for the assessment of public
sentiments, attitudes, and opinions concerning health-related
issues [10,11]. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 2
(page number not for citation purposes) Data Collection An advanced AI-based, text analytics platform using NLP was
used to initially analyze the tweets. The analytics platform,
“Wordnerds,” is described by its developers as a “text analysis
and insights platform using machine learning techniques” [13]. In particular, this off-the-shelf platform supports analysis of
metadata, topic, and sentiment to understand the context of a
tweet and to group tweets together into topic clusters that contain
tweets relating to each other or discussing similar issues. This
facilitates a more accurate and sophisticated insight into the
vaccine conversation on Twitter compared with methodologies
that rely solely on a qualitative count of single words, phrases,
or hashtags [14]. Channels of soft intelligence like Twitter, leveraged using novel
artificial intelligence (AI) techniques (including natural language
processing [NLP]), offer an opportunity for real-time analysis
of public attitudes, sentiments, and key topics of discussion
[12]. As aforementioned, previous case studies have shown that We developed a search strategy comprising a list of terms related
to COVID-19, the lockdown, and mental health to search (or
“scrape”) for relevant tweets (see Textbox 1). Textbox 1. Search strategy for relevant tweets. Corona OR covid OR lockdown
AND
mental health OR anxiety OR depression OR anxious OR depressed OR depressing OR trauma OR traumatic OR “obsessive compulsive disorder”
OR OCD OR vulnerable OR loneliness OR lonely OR isolated OR isolation OR sleep OR stress OR stressful OR self-harm OR self-harming OR
suicide OR suicidal OR well-being Corona OR covid OR lockdown https://infodemiology.jmir.org/2022/1/e32449 Data Analysis We used Wordnerds to interrogate the tweets. The developers
state that their platform uses a range of different technologies
in order to deliver its various analyses, including contextual
word embeddings and collocation methods [13]. To date, we
have found no other published studies coordinated by an
academic research group that have used this specific tool. The NLP platform was then used to explore the sentiment (ie,
positive, neutral, or negative) of the whole corpus of tweets. Sentiment was determined using contextual word embedding
techniques, including classification of grammar to understand
how words interact [16]. Using the platform, we were able to track and determine the
weekly frequency of tweets relevant to our initial search strategy Textbox 3. Keyword filters used to identify specific mental health problems. Textbox 3. Keyword filters used to identify specific mental health problems. Textbox 3. Keyword filters used to identify specific mental health problems. Anxiety: “anxious,” “anxiety”
Depression: “depression,” “depressed,” “depressing”
Stress: “stress,” “stressful,” “stressed”
Loneliness: “loneliness,” “lonely,” “alone” Following sentiment analysis, the platform’s topic analysis
feature was used to identify and cluster key emerging topics of
discussion, both with positive and negative underlying
sentiment. For this analysis, the platform automatically clustered
key topics of positive and negative discussion using topic
collocation methods. This is a probabilistic method of
identifying interesting sentence fragments and words that occur
frequently together within a data set. The results of the
platform’s topic analysis were examined, and its findings were
summarized by 2 of the authors (KL and RG). These summaries
were checked by 2 further authors (CM and GCW). Following sentiment analysis, the platform’s topic analysis
feature was used to identify and cluster key emerging topics of
discussion, both with positive and negative underlying
sentiment. For this analysis, the platform automatically clustered
key topics of positive and negative discussion using topic
collocation methods. This is a probabilistic method of
identifying interesting sentence fragments and words that occur
frequently together within a data set. The results of the
platform’s topic analysis were examined, and its findings were
summarized by 2 of the authors (KL and RG). These summaries
were checked by 2 further authors (CM and GCW). notable events or issues that occurred throughout the study
period are summarized in Table 1 (weeks 1 to 12) and Table 2
(weeks 13 to 24). Textbox 2. Keyword filter used to clean the data set. Textbox 2. Keyword filter used to clean the data set. mood OR “mental health” OR depression OR anxiety OR anxious OR depressed OR depressing OR trauma OR traumatic OR OCD OR compulsive
OR vulnerable OR loneliness OR lonely OR isolated OR sleep OR stress OR stressful OR self-harm OR self-harming OR suicide OR suicidal OR
anorexia OR anorexic OR bulimia OR bulimic OR eating disorder OR binge eating OR OFSED (other specified feeding or eating disorder) from the United Kingdom between July 23, 2020 and January
6, 2021. We also tracked the frequency of subsets of these tweets
that incorporated terms for specific mental health problems, as
listed in Textbox 3. Data Analysis As shown in Figure 1, the highest volume of tweets occurred
week commencing (w/c) October 29, 2020, with 51,340 tweets. The lowest volume was observed w/c December 3, 2020, with
12,622 tweets. The data show a fairly consistent baseline trend
over the study period. Spikes in the volume of tweets occurred
in September, October, and December, typically during periods
leading up to (or during) a major change in social distancing
and lockdown measures across the United Kingdom. Due to the high volume of tweets collected, the topic analysis
was split between 2 equal time periods. The first covered
summer 2020 to autumn 2020, when lockdown restrictions were
relaxed. The second covered the autumn to winter period in
2020, when regional and then further national lockdown
restrictions were introduced. The first peak was observed w/c September 17, 2020, the week
after the introduction of the “rule of 6,” whereby a mix of 6
people from any household could meet indoors or outdoors. A
similar peak was observed w/c October 8, 2020. This was the
week leading up to the government’s introduction of a new
tiered system, whereby regions across the United Kingdom were
allocated to 1 of 3 tiers (and later a fourth tier) based on
prevalence of COVID-19. Higher tiers corresponded with tighter
restrictions, including closing nonessential businesses and limits
placed on social gatherings. Ethical Considerations Institutional review board approval was not sought as this study
used only publicly available data. All posts were de-identified,
and there was no direct interaction with Twitter users. The largest peak was observed w/c October 29, 2020, the week
before the second national lockdown began. The final peak
occurred w/c December 31, 2020, the week leading up to the
start of a third national lockdown. For all 3 national lockdowns,
all nonessential businesses were closed, and UK residents were
restricted from meeting anyone outside of their “social bubble”
(ie, their household or, for people living alone, themselves plus
one other household). Preparing and Cleaning the Data All collated tweets were anonymized. Before analyzing the data,
the retrieved results were run through a final keyword filter. This filter was comprised of a series of terms and keywords
associated with mental health problems to help ensure a more
relevant, cleaner, and less noisy data set for analysis. For XSL•FO
RenderX XSL•FO
RenderX Marshall et al JMIR INFODEMIOLOGY were added alongside the original terms. example, general terms such as “isolation” and “well-being”
were filtered out. Further, terms associated with eating disorders JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 3
(page number not for citation purposes) https://infodemiology.jmir.org/2022/1/e32449 JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 4
(page number not for citation purposes) Tweet Volume We captured and collated 286,902 tweets posted by users in the
United Kingdom from July 23, 2020 to January 6, 2021. The
volume of tweets by week, together with key events taking place
during this study period, is visualized in Figure 1. Further JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 3
(page number not for citation purposes) XSL•FO
RenderX JMIR INFODEMIOLOGY JMIR INFODEMIOLOGY Marshall et al Marshall et al Figure 1. Volume of tweets from July 23, 2020 to January 6, 2021. Key events during the study period included the (1) rule of 6 (up to 6 people from
any number of households could meet indoors or outdoors), (2) tier system (regions across England were assigned a tier from 1 to 3 based on
epidemiological indicators, and these tiers dictated the restrictions in that area, such as which businesses could open and how many individuals could
meet in a group during national lockdown—nonessential businesses were closed and people were prohibited from meeting outside of their support
bubble). Table 1. Notable events that occurred from July 23, 2020 to October 14, 2020. Events
Weeks
Weeks 1-2
•
“Health Protection Regulations 2020” comes into force (ie, mandatory wearing of face masks in most indoor establishments
•
Gyms, swimming pools, and other indoor sports facilities reopen. •
England reports highest number of excess deaths in Europe for February to June. •
Major incident is declared in Greater Manchester after rise in cases. Weeks 3-4
•
Stricter measures are reintroduced in Preston, Lancashire. •
England’s revamped contact-tracing app begins public trials. •
General Certificate of Secondary Education (GCSE) results are published with grades based on teachers’ assessments. •
The Education Secretary confirms free appeals for A-Level and GCSE. Weeks 5-6
•
Ban on property evictions is extended until September 20. •
Greater Manchester Police report breaking up 126 illegal gatherings. •
Thousands of lockdown protesters gather in Trafalgar Square. Weeks 7-8
•
Health experts express doubt about mass testing plan: “Operation Moonshot.”
•
World Suicide Prevention Day is on September 10. •
“Rule of 6” for indoor and outdoor gatherings is announced. Weeks 9-10
•
Local lockdown measures are announced for Newcastle. •
Restrictions are relaxed for childcare purposes between households. •
Couples in established relationships can meet without social distancing. •
Second version of National Health Service (NHS) contact-tracing app becomes publicly available. •
£10,000 (US $13,035) fine for failing to self-isolate is announced. Weeks 11-12
•
Lockdown restrictions are tightened in the Northeast. •
Tier system comes into force, replacing local lockdowns. •
World Mental Health Day occurs on October 10. Table 1. Notable events that occurred from July 23, 2020 to October 14, 2020. https://infodemiology.jmir.org/2022/1/e32449 Marshall et al JMIR INFODEMIOLOGY Table 2. Marshall et al Notable events that occurred from October 15, 2020 to January 6, 2021
Events
Weeks
Weeks 13-14
•
Shielding ends for clinically vulnerable
•
Additional financial support to businesses in Tier 3 is announced. Weeks 15-16
•
Month-long national lockdown in England from November 5, 2020 is announced. •
Clinically vulnerable are asked to shield again. •
Furlough scheme is extended until March 2021. •
Pfizer/BioNTech press release announces vaccine is 90% effective. •
Travel window for university students to return for Christmas is announced. Weeks 17-18
•
Government publishes their “staying mentally well this winter” guidance. •
Plans to ease restrictions for 5 days over Christmas are announced. Weeks 19-20
•
Indoor care home visits can resume subject to lateral flow test. •
Medicines and Healthcare products Regulatory Agency (MHRA) approves Pfizer/BioNTech vaccine for rollout in the
United Kingdom. •
Joint Committee on Vaccination and Immunisation (JCVI) publishes vaccine priority groups. •
National lockdown ends, and Tier system resumes. Most regions in England are placed in Tier 2 or 3. •
Shielding ends for clinically vulnerable. •
First COVID-19 vaccine is administered. Weeks 21-22
•
Self-isolation period is reduced from 14 days to 10 days. •
New variant of the virus is identified. •
Further regions of the United Kingdom enter highest Tier. •
Furlough scheme is extended to April 2021. •
Large parts of Southeast England move into the new, stricter Tier 4. •
Christmas relaxation period is reduced from 5 days to 1 day. •
Travel restrictions for South Africa are enforced. •
Most of England will enter Tier 4 from December 26, 2020 is announced. Weeks 23-24
•
MHRA approves AstraZeneca/Oxford vaccine. •
JCVI publishes updated guidance. •
UK Prime Minister announces third national lockdown from January 4, 2021, with schools remaining closed. December 3, 2020. The “Anxiety” filter returned the highest
total number of tweets, and the “Loneliness” filter returned the
lowest (see Table 3). The trend in tweet frequency for each filter
mirrored the trend reported for the overall data set over the study
period. December 3, 2020. The “Anxiety” filter returned the highest
total number of tweets, and the “Loneliness” filter returned the
lowest (see Table 3). The trend in tweet frequency for each filter
mirrored the trend reported for the overall data set over the study
period. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 5
(page number not for citation purposes) Textbox 4. Examples of positive and negative tweets classified by the NLP platform. Example tweets classified with positive sentiment
...Since lockdown my anxiety has dramatically gotten so much better. I used to get stress spots, panic attacks before
presentations, etc. Na bruh, working from home has been a life saver... If anyone is struggling with lockdown (or wants to reduce stress/anxiety). I would really recommend trying gratitude
journaling. I’ve shared some tips below – I hope they’ll be helpful
…Never been so happy to set foot in a gym today. I feel so much happier getting back into a routine and pleased to
see I haven’t lost much strength through pregnancy and lockdown. It’s made me realise how important it is for my
own mental health... Example tweets classified with negative sentiment
...I’m finding this lockdown harder than the last one. For me it's not just the weather, it’s having no end in sight which
is a struggle when you suffer with anxiety and depression... …genuinely think a second lockdown would completely destroy my mental health. I’m nervous about the next few
weeks/months... Feeling absolute rubbish today, feel like my mental health is debilitating can’t bring myself to get out of bed, my only
escape is drinking with to forget these awful times Sentiment Analysis Of the 286,902 tweets, 34,347 (11.97%) were identified as
having positive sentiment, 217,728 (75.89%) as having neutral
sentiment, and 34,827 (12.14%) as having negative sentiment,
with an overall sentiment score of 50% assigned. A score below
50% suggests negative sentiment, and a score greater than 50% Here, the overall score of 50% indicates neutral sentiment across
all tweets over the study period. Figure 3 visualizes the weekly
change in sentiment over the study period from July 23, 2020
to January 6, 2021. Textbox 4. Examples of positive and negative tweets classified by the NLP platform. Mental Health Problems Figure 2. Volume of tweets from July 23, 2020 to January 6, 2021. suggests positive sentiment. Textbox 4 presents a selection of
example tweets that the NLP platform classified as both positive
and negative. Sentiment Analysis Mental Health Problems Table 3 summarizes the volume of tweets that utilized at least
one of the terms related to anxiety, depression, stress, or
loneliness. In total, 113,312 (39.50%) of the 286,902 tweets
scraped through the initial search strategy related to anxiety,
depression, stress, or loneliness. During the first half of the analysis period (w/c July 23, 2020
to w/c October 22, 2020), tweet volume between the 4 mental
health problem filters varied. In particular, tweets related to the
“Anxiety” filter were consistently posted most often, and tweets
relating to the “Loneliness” filter were consistently posted least. There was a spike in volume across all 4 filters w/c October 29,
2020, the start of the second national lockdown. Following the
spike, the volume of tweets across all of the filters was broadly
similar for the remainder of the analysis period. Figure 2 presents the volume of tweets utilizing terms related
to anxiety, depression, stress, or loneliness around each keyword
filter over the study period. Across all of the mental health problems that were focused on
here, the highest volume of tweets was observed w/c October
29, 2020, and the lowest volume of tweets occurred w/c Table 3. Keyword filters and resultant volume of tweets for specific mental health problems. Tweets (N=113,312)
Keyword terms
Filter
37,213 (32.84%)
“anxious,” “anxiety”
Anxiety
29,523 (26.05%)
“depression,” “depressed,” “depressing”
Depression
26,725 (23.59%)
“stress,” “stressful,” “stressed”
Stress
19,851 (17.52%)
“loneliness,” “lonely,” “alone”
Loneliness https://infodemiology.jmir.org/2022/1/e32449 https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
RenderX JMIR INFODEMIOLOGY Marshall et al JMIR INFODEMIOLOGY Figure 2. Volume of tweets from July 23, 2020 to January 6, 2021. Sentiment Analysis
Of the 286,902 tweets, 34,347 (11.97%) were identified as
having positive sentiment, 217,728 (75.89%) as having neutral
sentiment, and 34,827 (12.14%) as having negative sentiment,
with an overall sentiment score of 50% assigned. A score below
50% suggests negative sentiment, and a score greater than 50%
suggests positive sentiment. Textbox 4 presents a selection of
example tweets that the NLP platform classified as both positive
and negative. Here, the overall score of 50% indicates neutral sentiment across
all tweets over the study period. Figure 3 visualizes the weekly
change in sentiment over the study period from July 23, 2020
to January 6, 2021. Textbox 4. Examples of positive and negative tweets classified by the NLP platform. Example tweets classified with positive sentiment
Marshall et al
JMIR INFODEMIOLOGY Figure 2. Volume of tweets from July 23, 2020 to January 6, 2021. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 6
(page number not for citation purposes) Example tweets classified with positive sentiment ...Since lockdown my anxiety has dramatically gotten so much better. I used to get stress spots, panic attacks before
presentations, etc. Na bruh, working from home has been a life saver... If anyone is struggling with lockdown (or wants to reduce stress/anxiety). I would really recommend trying
journaling. I’ve shared some tips below – I hope they’ll be helpful If anyone is struggling with lockdown (or wants to reduce stress/anxiety). I would really recommend trying gratitude
journaling. I’ve shared some tips below – I hope they’ll be helpful …Never been so happy to set foot in a gym today. I feel so much happier getting back into a routine and pleased to
see I haven’t lost much strength through pregnancy and lockdown. It’s made me realise how important it is for my
own mental health... Example tweets classified with negative sentiment Example tweets classified with negative sentiment
...I’m finding this lockdown harder than the last one. For me it's not just the weather, it’s having no end in sight which
is a struggle when you suffer with anxiety and depression... …genuinely think a second lockdown would completely destroy my mental health. I’m nervous about the next few
weeks/months... Feeling absolute rubbish today, feel like my mental health is debilitating can’t bring myself to get out of bed, my only
escape is drinking with to forget these awful times Feeling absolute rubbish today, feel like my mental health is debilitating can’t bring myself to get out of bed, my only
escape is drinking with to forget these awful times JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 6
(page number not for citation purposes) JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 6
(page number not for citation purposes) https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
RenderX JMIR INFODEMIOLOGY Marshall et al Figure 3. Sentiment of tweets from July 23, 2020 to January 6, 2021. Figure 3. Sentiment of tweets from July 23, 2020 to January 6, 2021. Figure 3. Sentiment of tweets from July 23, 2020 to January 6, 2021. Figure 3. Sentiment of tweets from July 23, 2020 to January 6, 2021. Figure 3. Sentiment of tweets from July 23, 2020 to January 6, 2021. Sentiment remained neutral or positive throughout most of the
study period. The highest assigned sentiment score, 52%,
occurred in weeks 1, 4 to 6, 16, and 18 to 20. The lowest
assigned sentiment score, 49%, occurred in week 9 (w/c
September 17, 2020). Sentiment remained neutral or positive throughout most of the
study period. The highest assigned sentiment score, 52%,
occurred in weeks 1, 4 to 6, 16, and 18 to 20. The lowest
assigned sentiment score, 49%, occurred in week 9 (w/c
September 17, 2020). health throughout the “coronavirus pandemic,” as a critical
health issue, was shared widely by people on Twitter. During
this period, people openly discussed their mental health and
how they had been coping. People also shared praise for specific
local and national mental health services, as well as key public
figures (eg, Marcus Rashford). Overall, the data show a relatively consistent trend in sentiment
over the study period. Example tweets classified with negative sentiment When sentiment fluctuation did occur, it
was similar to the trend observed with tweet frequency and
coincided with major changes to lockdown or social distancing
rules. There was considerable discussion based around “World Mental
Health Day” and “World Suicide Prevention Day.” People were
sharing helpful strategies (eg, videos, charities, help lines,
exercise regimes, healthy eating advice) others could use to
protect and maintain their mental health. There were also calls
from people to be particularly vigilant and make sure they are
checking in with any “vulnerable people” in their life. Topic Analysis In this section, results are reported for 2 time periods, as follows:
(1) July 23, 2020 to October 14, 2020 and (2) October 15, 2020
to January 6, 2021. Positive discussion was observed around “working from home.”
Various users were sharing helpful resources to support working
from home effectively, including strategies that people had
found useful during the previous national lockdown. Some
people reported that mandatory working from home had helped
them to achieve a better work-life balance and reduced their
anxiety. Results of the Topic Analysis for Weeks 1 to 12 This section presents the results of a topic analysis based on
115,700 scraped tweets posted from July 23, 2020 to October
14, 2020. ...Never been so happy to set foot in a gym today. I feel so much happier getting back into a routine and pleased to
see I haven’t lost much strength through pregnancy and lockdown. It’s made me realise how important it is for my
own mental health... ...I eventually took time out from March as it just got too much. My anxiety was through the roof and frankly I was
glad of lockdown - I didn’t want to leave the house anyway. I’m in a much better place now thankfully... ...This coming Thursday is world suicide prevention day and we need to be talking about this in the context of covid,
lockdown, the economy and more. Most of all we need to talk and respond with compassion and support... xtbox 5. Sample of clustered tweets from weeks 1 to 12 with positive sentiment. ...Never been so happy to set foot in a gym today. I feel so much happier getting back into a routine and pleased to
see I haven’t lost much strength through pregnancy and lockdown. It’s made me realise how important it is for my
own mental health...
...I eventually took time out from March as it just got too much. My anxiety was through the roof and frankly I was
glad of lockdown - I didn’t want to leave the house anyway. I’m in a much better place now thankfully...
...This coming Thursday is world suicide prevention day and we need to be talking about this in the context of covid,
lockdown, the economy and more. Most of all we need to talk and respond with compassion and support... https://infodemiology.jmir.org/2022/1/e32449 Summary of Clustered Topics With Negative Sentiment Tables 4 and 5 present the top 10 most discussed topics th
occurred throughout the study period. ...went into the year hopeful and ready to turn my life around...then covid happened and the world became a giant
prison. Depression slowly came back, anxiety eating away and all emotion and feeling slowly dry up. Mind became
a wasteland pretty much. Lifeless and desert... ...I have to get a COVID test done before going back to work at the end of the month and the anxiety I am having from
the thought of them sticking that thing up my nose... ...I’m finding this lockdown harder than the last one. For me it's not just the weather, it’s having no end in sight which
is a struggle when you suffer with anxiety and depression... ...genuinely think a second lockdown would completely destroy my mental health. I’m nervous about the next few
weeks/months... ...I'm all for people following the rules - wear your masks and keep distance when required, but this is very true and
will make people more depressed by lack of communication with real people. Depression will be a huge and sad factor
if this new lockdown does go ahead. Stay safe... ...This bizarre competition between suicide and Covid deaths needs to end. I understand the mental health issues that
can arise from lack of social interaction and other factors from Covid but this weird death competition between the
2 needs to stop... bl 2
i
l
i
f
bl
h Table 2 summarizes a selection of notable events that occurred
during this time period. Summary of Clustered Topics With Negative Sentiment y f
p
g
“Second lockdown” emerged as a key topic of discussion
underpinned with negative sentiment. People were sharing their
fears, concerns, and anxieties over the prospect of a second
national lockdown and the impact this would have on theirs
(and other’s) mental health. People recalled and spoke openly
about how their mental health had suffered during the previous
lockdown, referencing specific problems such as “anxiety,”
“depression,” and posttraumatic stress disorder. Some people
shared that they had been diagnosed with depression for the
first time due to the previous national lockdown. Many people were angry that not enough had been done by the
government to protect “vulnerable people” during the previous
national lockdown. Suicide was also discussed. People were
claiming that suicide rates had increased during lockdown,
particularly among younger people. There was widespread
sharing of warnings from key educational figures that the
pandemic would have long-lasting negative effects on Tweets contained within some of the other topic clusters
generated by the platform, such as “care homes” and “covid
deaths,” did not appear to be related to mental health. Specific examples of mental health tweets underpinned with
negative sentiment from weeks 1 to 12 are presented in Textbox
6. Textbox 6. Sample of clustered tweets from weeks 1 to 12 with negative sentiment" to be consistent with the other textbox p
g
...went into the year hopeful and ready to turn my life around...then covid happened and the world became a giant
prison. Depression slowly came back, anxiety eating away and all emotion and feeling slowly dry up. Mind became
a wasteland pretty much. Lifeless and desert... ...I have to get a COVID test done before going back to work at the end of the month and the anxiety I am having from
the thought of them sticking that thing up my nose... ...I’m finding this lockdown harder than the last one. For me it's not just the weather, it’s having no end in sight which
is a struggle when you suffer with anxiety and depression... ...genuinely think a second lockdown would completely destroy my mental health. I’m nervous about the next few
weeks/months... ...I'm all for people following the rules - wear your masks and keep distance when required, but this is very true and
will make people more depressed by lack of communication with real people. Summary of Clustered Topics With Negative Sentiment Depression will be a huge and sad factor
if this new lockdown does go ahead. Stay safe... ...This bizarre competition between suicide and Covid deaths needs to end. I understand the mental health issues that
can arise from lack of social interaction and other factors from Covid but this weird death competition between the
2 needs to stop... esults of the Topic Analysis for Weeks 13 to 24
his section presents the results of a topic analysis of 171,202
raped tweets posted from October 15, 2020 to January 6, 2021. Table 2 summarizes a selection of notable events that occurred
during this time period. Tables 4 and 5 present the top 10 most discussed topics that
occurred throughout the study period. ...went into the year hopeful and ready to turn my life around...then covid happened and the world became a giant
prison. Depression slowly came back, anxiety eating away and all emotion and feeling slowly dry up. Mind became
a wasteland pretty much. Lifeless and desert... ...I have to get a COVID test done before going back to work at the end of the month and the anxiety I am having from
the thought of them sticking that thing up my nose... ...I’m finding this lockdown harder than the last one. For me it's not just the weather, it’s having no end in sight which
is a struggle when you suffer with anxiety and depression... ...genuinely think a second lockdown would completely destroy my mental health. I’m nervous about the next few
weeks/months... ...I'm all for people following the rules - wear your masks and keep distance when required, but this is very true and
will make people more depressed by lack of communication with real people. Depression will be a huge and sad factor
if this new lockdown does go ahead. Stay safe... ...This bizarre competition between suicide and Covid deaths needs to end. I understand the mental health issues that
can arise from lack of social interaction and other factors from Covid but this weird death competition between the
2 needs to stop... Results of the Topic Analysis for Weeks 13 to 24
This section presents the results of a topic analysis of 171,202
scraped tweets posted from October 15, 2020 to January 6, 2021. Table 2 summarizes a selection of notable events that occurr
during this time period. Summary of Clustered Topics With Positive Sentiment Specific examples of mental health tweets underpinned with
positive sentiment from weeks 1 to 12 are presented in Textbox Specific examples of mental health tweets underpinned with
positive sentiment from weeks 1 to 12 are presented in Textbox
5. “Mental health” emerged as a key topic of discussion
underpinned with positive sentiment. The importance of mental Textbox 5. Sample of clustered tweets from weeks 1 to 12 with positive sentiment. ...Never been so happy to set foot in a gym today. I feel so much happier getting back into a routine and pleased to
see I haven’t lost much strength through pregnancy and lockdown. It’s made me realise how important it is for my
own mental health... ...This coming Thursday is world suicide prevention day and we need to be talking about this in the context of covid,
lockdown, the economy and more. Most of all we need to talk and respond with compassion and support... JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 7
(page number not for citation purposes) https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
RenderX XSL•FO
RenderX JMIR INFODEMIOLOGY Marshall et al “children.” Further, some argued that people were using “mental
health” as an excuse to avoid further lockdown restrictions. Many people were concerned that a second lockdown would
be much worse for people’s mental health than the first (the
lockdown coinciding with winter and students returning to
university were both seen as contributing negative factors). Those who refused to wear masks were a further source of
anxiety for some people, with a high proportion of tweets calling
on others to “wear a mask.” People also discussed COVID-19
tests during this period. In particular, some people shared how
stressful and anxiety-inducing taking the test, and also waiting
for the results, can be. ...went into the year hopeful and ready to turn my life around...then covid happened and the world became a giant
prison. Depression slowly came back, anxiety eating away and all emotion and feeling slowly dry up. Mind became
a wasteland pretty much. Lifeless and desert... ...went into the year hopeful and ready to turn my life around...then covid happened and the world became a giant
prison. Depression slowly came back, anxiety eating away and all emotion and feeling slowly dry up. Mind became
a wasteland pretty much. Lifeless and desert...
...I have to get a COVID test done before going back to work at the end of the month and the anxiety I am having from
the thought of them sticking that thing up my nose...
...I’m finding this lockdown harder than the last one. For me it's not just the weather, it’s having no end in sight which
is a struggle when you suffer with anxiety and depression...
...genuinely think a second lockdown would completely destroy my mental health. I’m nervous about the next few
weeks/months...
...I'm all for people following the rules - wear your masks and keep distance when required, but this is very true and
will make people more depressed by lack of communication with real people. Depression will be a huge and sad factor
if this new lockdown does go ahead. Stay safe...
...This bizarre competition between suicide and Covid deaths needs to end. I understand the mental health issues that
can arise from lack of social interaction and other factors from Covid but this weird death competition between the
2 needs to stop...
Results of the Topic Analysis for Weeks 13 to 24
Table 2 summarizes a selection of notable events that occur
during this time period. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 9
(page number not for citation purposes) Suicide figures are up 200% since lockdown. Could two followers please copy and re-post this tweet? We’re trying
to demonstrate that someone is always listening. Call 116 123 (Samaritans UK). Just two. Any two. Copy, not RT.
#MentalHealth #SuicidePrevention #SuicideAwareness
important to remember with all the 200% increase in suicide tweets. Raising awareness is fantastic, but make sure
it’s factual. #SuicidePrevention. https://infodemiology.jmir.org/2022/1/e32449 Results of the Topic Analysis for Weeks 13 to 24 This section presents the results of a topic analysis of 171,202
scraped tweets posted from October 15, 2020 to January 6, 2021. Tables 4 and 5 present the top 10 most discussed topics that
occurred throughout the study period. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 8
(page number not for citation purposes) https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
RenderX XSL•FO
RenderX Marshall et al JMIR INFODEMIOLOGY Table 4. Top 10 most discussed positive and negative topics from July 23, 2020 to October 14, 2020. Number of tweets
Clustered topic
Ranking
Topics with positive sentiment
5499
“mental health”
1
1513
“vulnerable people”
2
491
“lockdown anxiety”
3
348
“communities... support”
4
327
“health... important”
5
266
“coronavirus pandemic”
6
257
“work from home”
7
246
“support... provide”
8
234
“improved... health”
9
233
“world day”
10
Topics with negative sentiment
5192
“mental health”
1
1421
“people... vulnerable”
2
1037
“children... coronavirus”
3
743
“covid... deaths”
4
566
“wear a mask”
5
419
“test for covid”
6
411
“lockdown... gone”
7
405
“care homes”
8
377
“second lockdown”
9
365
“anxiety... depression”
10 Table 4. Top 10 most discussed positive and negative topics from July 23, 2020 to October 14, 2020. Clustered topic
Ranking https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
RenderX XSL•FO
RenderX Marshall et al JMIR INFODEMIOLOGY Table 5. Top 10 most discussed positive and negative topics from October 15, 2020 to January 6, 2020.. Number of tweets
Clustered topic
Ranking
Topics with positive sentiment
10,096
“suicide lockdown”
1
10,031
“mental health”
2
1860
“people…vulnerable”
3
526
“friends or family”
4
451
“need…support”
5
409
“ones you love”
6
306
“community…helping”
7
295
“stress & anxiety
8
275
“night’s sleep”
9
259
“difficult times”
10
Topics with negative sentiment
10,299
“mental health“
1
1767
“vulnerable people”
2
1025
“going…lockdown”
3
979
“deaths…covid”
4
490
“suicide rates”
5
479
“sleep at night”
6
474
“lockdown…depressing”
7
460
“committed suicide”
8
441
“coronavirus…children”
9
441
“wearing masks”
10 children, young people, disabled people, those with learning
difficulties, and those with any pre-existing mental health
problems. Some people were calling on the government to
provide further support for these groups as lockdown restrictions
tightened. Many people were encouraging those that that were
struggling to stay connected with others and reach out to “ones
you love” and “friends and family.” Summary of Clustered Topics With Positive Sentiment Summary of Clustered Topics With Positive Sentiment During this period, a tweet suggesting suicide rates had risen
by 200% since lockdown was shared widely. The tweet
contained contact details for a registered UK charity, Samaritans,
urging people to reach out for support if needed. This viral tweet
resulted in our analysis platform recognizing “suicide lockdown”
as a key topic of discussion. The information being reported by
this tweet was not accurate [17,18]. “Sleep” emerged as a key topic of discussion with positive
sentiment. Many people discussed how important getting a good
“night’s sleep” was for their mental health, particularly during
these “difficult times.” People shared relaxation techniques they
had used, which had helped them to fall asleep, and which may
help others too. As with the previous 12 weeks, “mental health” remained a key
topic of discussion underpinned with positive sentiment during
this period. People discussed how the lockdown had, in some
ways, had a positive impact on their mental health. Various
people and organizations continued to share practical tips on
how to support one’s mental health, particularly around
strategies to reduce “stress and anxiety.” Specific examples of mental health tweets underpinned with
positive sentiment from weeks 13 to 24 are presented in Textbox Many people continued to express concern for the mental health
of perceived “vulnerable people” during lockdown, including Textbox 7. Sample of clustered tweets from weeks 13 to 24 with positive sentiment. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 10
(page number not for citation purposes) XSL•FO
RenderX JMIR INFODEMIOLOGY Marshall et al My mental health is sooo much better now it’s a real lockdown again. However I know this isn’t a universal experience. For those who find lockdown harder for whatever reason, I’m a) sending virtual hugs if wanted but b) reminding you
it’s STRONG to reach out to a helpline! As we start lockdown 3, a reminder that your situation does not have to be the worst for it to suck and for you to get
help! Reach out to your loved ones & professional mental health support if you need it. Stay safe. Such a tough time at the moment for everyone, another lockdown especially in winter can be devastating for mental
health. My dm’s are always open for anyone who needs a chat – be kind and check on your loved ones. Summary of Clustered Topics With Negative Sentiment Summary of Clustered Topics With Negative Sentiment “Mental health” continued to be widely discussed during this
final 12-week period. Many people reflected on the negative
impact that lockdowns had had on their mental health. There
were concerns from some that any progress they had made with
their mental health would be lost with another lockdown. There
was continued anger toward the UK government about a
perceived lack of support for those struggling with their mental
health. During this period, people shared their thoughts about the
vaccine rollout. In particular, people were concerned about the
length of time between jabs and the number of canceled
vaccination appointments being reported by the media. “Sleep” continued to be a key topic of discussion with many
people sharing how they had not been sleeping well. Some
people shared how they had been increasing their alcohol intake
in an effort to help them sleep. There was a lot of discussion around the looming national
lockdown announced for January 4, 2021. Many people
expressed concern about how long this lockdown would last
and their hope that this would be the final lockdown. Some
shared that they would be defying restrictions in order to
prioritize their mental health. Others continued to argue that Specific examples of mental health tweets underpinned with
negative sentiment from weeks 13 to 24 are presented in
Textbox 8. Textbox 8. Sample of clustered tweets from weeks 13 to 24 with negative sentiment. Textbox 8. Sample of clustered tweets from weeks 13 to 24 with negative sentiment. Well lockdown 3.0 has barely started and I can already feel all the hard work I put balancing my mental health slip
away
Tbh I just hope my mental health doesn’t become as bad as first lockdown
Feeling absolute rubbish today, feel like my mental health is debilitating can’t bring myself to get out of bed, my only
escape is drinking with to forget these awful times
There needs to be more mental health support @BorisJohnson as that will be one of the highest collateral costs of
this. People like me are struggling badly in isolation and with mental health issues and there isn’t enough support. https://infodemiology.jmir.org/2022/1/e32449 Summary of Clustered Topics With Positive Sentiment Thanks to local organisations, community groups and faith institutions that have provided vital services, human
support and companionship, in person and online, to many vulnerable people during #Covid-19. We will keep working
with you all to build stronger and united communities. If anyone is struggling with lockdown (or wants to reduce stress/anxiety). I would really recommend trying gratitude
journaling. I’ve shared some tips below – I hope they’ll be helpful #COVID19 wellbeing tip: make sure you get a good night’s sleep! A good rest is so important for your mental and
physical health, managing stress and much more. If you’re struggling with sleep, try these tips and check out our
Sleep self-help guide. people were using their mental health as an excuse for not
following the rules. There was continued worry about how the
lockdown would affect perceived “vulnerable people.” Summary of Clustered Topics With Negative Sentiment #Uklockdown #Covid
My mental health is already rock bottom due to not seeing my family/friends, being overworked and not sleeping –
god help what another lockdown is gonna do to me
… everyone is talking about lockdown 3 but carehomes haven’t left lockdown since the first lockdown started. Everyone
is ignoring how vulnerable people feel, disables people or careworkers and carers. ...I’ve never had ocd but do have A LOT of anxiety, + have noticed the longer lockdown goes on the more my anxious
behaviours start to look like obsessive ones eg. I check my cooker before bed/before leaving my house when I never
did that pre lockdown, also started counting.! It’s only our second day of lockdown 3.0 going solo & I’m already finding harder than the others. It could be due to
the distinct lack of sleep last night. What are your favourite self care activities to do? I need motivation to get off the
sofa today #lockdown #lockdownblues
...everyone is rotting in their own houses and getting depression I’m not saying they’re less important but clearly this
going in and out of lockdown isn’t helping anyone is it There needs to be more mental health support @BorisJohnson as that will be one of the highest collateral costs of
this. People like me are struggling badly in isolation and with mental health issues and there isn’t enough support. #Uklockdown #Covid My mental health is already rock bottom due to not seeing my family/friends, being overworked and not sleeping –
god help what another lockdown is gonna do to me My mental health is already rock bottom due to not seeing my family/friends, being overworked and not sleeping –
god help what another lockdown is gonna do to me … everyone is talking about lockdown 3 but carehomes haven’t left lockdown since the first lockdown started. Everyone
is ignoring how vulnerable people feel, disables people or careworkers and carers. ...I’ve never had ocd but do have A LOT of anxiety, + have noticed the longer lockdown goes on the more my anxious
behaviours start to look like obsessive ones eg. I check my cooker before bed/before leaving my house when I never
did that pre lockdown, also started counting.! It’s only our second day of lockdown 3.0 going solo & I’m already finding harder than the others. It could be due to
the distinct lack of sleep last night. Limitations In this study, we identified and analyzed 286,902 geolocated
tweets posted from users in the United Kingdom from July 23,
2020 to January 6, 2021 using a commercially available NLP
platform. The findings showed that there was a fairly consistent
trend in the volume of tweets over the study period, with spikes
typically occurring during (or leading up to) a major change in
social distancing measures in the United Kingdom. The NLP
platform calculated an overall sentiment score of 50% indicating
neutral sentiment across all tweets over the study period. Similar
to volume, major fluctuations in sentiment appeared to coincide
with major changes to lockdown rules. Several limitations can be attributed to this study. First, there
are still considerable limitations concerning the reliability,
accuracy, and transparency of the technologies in play. As an
example, on examining the results of the NLP platform’s topic
analysis, some of the tweets collated were not relevant to mental
health (despite being identified as such). For example, tweets
contained within clustered topics like “care homes” and “covid
deaths” were expressing anger at the government, rather than
negatively discussing mental health problems. Some of the tweets included by the platform in its analysis were
posted by businesses or charitable organizations, rather than
members of the public. Such tweets, which often advertised
local or national mental health services or shared
self-improvement strategies, were typically classified by the
platform as having positive sentiment. This created a large
amount of background noise and skewed the overall sentiment
toward positive. Further, tweets were not deduplicated by the
platform, nor was there any formal analysis accounting for
potential bot traffic. These factors will also have impacted the
results. Key topics of discussion that emerged consistently throughout
the study period included (1) the impact that the pandemic and
resulting lockdowns had been having on people’s mental health,
both positive and negative; (2) fear and anxiety around the
prospect of prolonged and subsequent lockdowns and how this
might (or continue to) affect people’s mental health; and (3)
anger and mistrust toward the government concerning a
perceived lack of support for people struggling with their mental
health. Later in (and less consistently discussed throughout) the study
period, other topics linked with mental health emerged,
including sleep difficulties, increased alcohol intake, and
anxieties concerning testing and the vaccine rollout. Summary of Clustered Topics With Negative Sentiment What are your favourite self care activities to do? I need motivation to get off the
sofa today #lockdown #lockdownblues It’s only our second day of lockdown 3.0 going solo & I’m already finding harder than the others. It could be due to
the distinct lack of sleep last night. What are your favourite self care activities to do? I need motivation to get off the
sofa today #lockdown #lockdownblues ...everyone is rotting in their own houses and getting depression I’m not saying they’re less important but clearly this
going in and out of lockdown isn’t helping anyone is it JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 11
(page number not for citation purposes) https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
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RenderX Marshall et al JMIR INFODEMIOLOGY this lockdown is starting to get to me – went off my food, crying more and just generally depressed loool examining the value of this type of analyzed evidence and how
it might support, link to, and (where appropriate) replace more
traditional survey-based methods and data [15,21-24]. Limitations In addition, despite the popularity of Twitter as a social
networking tool, its users are not an accurate representation of
the overall demographic of a population. Therefore, if we are
to consider using Twitter (and similar resources) as a potential
intelligence source, we must be mindful of bias concerning the
key demographic information among its users (such as age,
gender, and socioeconomic status). Before the study, we anticipated that topics of discussion relating
to mental health would be mostly underpinned with negative
sentiment. It was therefore surprising that the findings of the
topic analysis revealed higher levels of positive sentiment across
posts associated with mental health. Consistently over the study
period, people took to Twitter to share practical tips, strategies,
and resources that could be used to support one’s mental health,
and the platform was effective in clustering these types of posts
with positive sentiment. There was also a number of limitations specific to the NLP
platform that we selected. We had originally planned to run the
topic analysis using the platform across all of the tweets as a
single corpus. However, the platform was not able to process
and analyze such a large volume of tweets in one go. Therefore,
we had to split and run the topic analysis over 2 time periods
(weeks 1 to 12 and weeks 13 to 24). Further, it was not possible
to retrospectively search for and collect historic tweets, thus
restricting possible options for analysis. Finally, although the
broader methodologies that power the platform are touched on
by its developers, the finer technical detail is not shared publicly
due to commercial reasons. The viral spread of misinformation and “fake news” has
represented a critical issue generating mass confusion, fear, and
insecurity surrounding COVID-19 [19]. The WHO has
repeatedly used the term “infodemic” to describe the sheer
overabundance of misinformation being shared throughout the
pandemic [20]. Our findings provide further example and insight
into the rapid spread of health-related misinformation,
particularly via channels of soft intelligence like social media. Specifically, in the case of this study, a copy-and-paste tweet
campaign falsely claiming that suicide rates had increased by
200% since the first lockdown was shared widely by users. This
particular cluster of tweets ranked as the topmost discussion
topic during weeks 13 to 24 over the study period. https://infodemiology.jmir.org/2022/1/e32449 JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 12
(page number not for citation purposes) Acknowledgments This project is funded by the National Institute for Health Research (NIHR; HSRIC-2016-1009/Innovation Observatory). The
views expressed are those of the author(s) and not necessarily those of the NIHR or the Department of Health and Social Care. Authors' Contributions CM was responsible for project supervision; the conception, design, and conduct of the study; leading the development of the
initial manuscript; and managing its revision. KL, RG, and GCW were responsible for data analysis, data interpretation, and
critical review and revision of the manuscript. FP and DC provided critical review of the manuscript. Conclusions In this work, we analyzed a large collection of UK tweets
relating to mental health during the COVID-19 pandemic to
further explore the value of soft intelligence leveraged using
NLP. Using a specialist, off-the-shelf, NLP platform, we collated
a large corpus of tweets over a 24-week period and carried out
various analyses to explore the volume, sentiment, and key
trends and topics of discussion. Overall, the results of this study demonstrate that using NLP to
mine and analyze sources of soft intelligence (like Twitter) can
yield useful health-related insights, which agencies, local
leaders, and health care decision makers can potentially draw
from. These findings contribute to a growing body of literature Our findings provide further evidence that this type of research
is potentially a highly useful and efficient means to gain a rapid JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 12
(page number not for citation purposes) JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 12
(page number not for citation purposes) XSL•FO
RenderX JMIR INFODEMIOLOGY Marshall et al real-time analysis and intelligence may be particularly useful
in helping shape rapid and reactive public health engagement
and communication strategies during a health crises like
COVID-19. real-time analysis and intelligence may be particularly useful
in helping shape rapid and reactive public health engagement
and communication strategies during a health crises like
COVID-19. understanding of the key messages, concerns, and issues people
are facing at scale. In the case of this reported study, we were
able to draw insights into how the pandemic may be impacting
people’s mental health and well-being by examining both the
topic and sentiment specific to the UK population. This type of understanding of the key messages, concerns, and issues people
are facing at scale. In the case of this reported study, we were
able to draw insights into how the pandemic may be impacting
people’s mental health and well-being by examining both the
topic and sentiment specific to the UK population. This type of Conflicts of Interest The National Institute for Health Research (NIHR) Innovation Observatory purchased a license to use and evaluate the natural
language processing (NLP) platform, Wordnerds. However, as independent users, the authors have no vested interest in the tool. References The National Institute for Health Research (NIHR) Innovation Observatory purchased a license to use and evaluate the natural
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[Medline: 32750001]
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10.2196/21418] [Medline: 33284783]
Abb
i
i 17. Appleby L, Richards N, Ibrahim S, Turnbull P, Rodway C, Kapur N. Suicide in England in the COVID-19 pandemic: Early
observational data from real time surveillance. Lancet Reg Health Eur 2021 May;4:100110 [FREE Full text] [doi:
10.1016/j.lanepe.2021.100110] [Medline: 34557817] j
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online/suicide-200-percent/ [accessed 2022-03-13] 19. Freiling I, Krause NM, Scheufele DA, Brossard D. Believing and sharing misinformation, fact-checks, and accurate
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(page number not for citation purposes) Abbreviations AI: artificial intelligence
ML: machine learning
NIHR: National Institute for Health Research
NLP: natural language processing
w/c: week commencing
WHO: World Health Organization Edited by T Mackey; submitted 28.07.21; peer-reviewed by H Lee, R Castilla-Puentes, W Zhang; comments to author 04.09.21; revised
version received 15.10.21; accepted 12.01.22; published 31.03.22
Please cite as:
Marshall C, Lanyi K, Green R, Wilkins GC, Pearson F, Craig D
Using Natural Language Processing to Explore Mental Health Insights From UK Tweets During the COVID-19 Pandemic: Infodemiology
Study
JMIR Infodemiology 2022;2(1):e32449
URL: https://infodemiology.jmir.org/2022/1/e32449
doi: 10.2196/32449
PMID: 36406146 ©Christopher Marshall, Kate Lanyi, Rhiannon Green, Georgina C Wilkins, Fiona Pearson, Dawn Craig. Originally published in
JMIR Infodemiology (https://infodemiology.jmir.org), 31.03.2022. This is an open-access article distributed under the terms of
the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original work, first published in JMIR Infodemiology, is properly
cited. The complete bibliographic information, a link to the original publication on https://infodemiology.jmir.org/, as well as
this copyright and license information must be included. JMIR Infodemiology 2022 | vol. 2 | iss. 1 | e32449 | p. 14
(page number not for citation purposes) https://infodemiology.jmir.org/2022/1/e32449 XSL•FO
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Interspecific variation in responses to microclimate by terrestrial isopods: implications in relation to climate change
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ZooKeys
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BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Interspecific
ZooKeys 801: 5–24 (2018)
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http://zookeys.pensoft.net Interspecific
ZooKeys 801: 5–24 (2018)
doi: 10.3897/zookeys.801.24934
http://zookeys.pensoft.net Interspecific
ZooKeys 801: 5–24 (2018)
doi: 10.3897/zookeys.801.24934
http://zookeys.pensoft.net responses to microclima
RESEARCH ARTICLE Interspecific variation in responses to microclimate by
terrestrial isopods: implications in relation
to climate change Mark Hassall1, Anna Moss1,2, Bernice Dixie1, James J. Gilroy1 1 School of Environmental Sciences, University of East Anglia, Norwich, NR4 7TJ, UK 2 School of Social
Sciences, University of Dundee, Dundee, DD1 4HN, UK Corresponding author: Mark Hassall (mhassall49@googlemail.com) Academic editor: Katalin Szlavecz | Received 10 March 2018 | Accepted 28 October 2018 | Published 3 December 2018 Introduction In this review we draw attention to an important contribution that soil biologists, in
particular those who study the biology of terrestrial isopods, can potentially make to
the current debate as to how global climate change may influence components of the
global carbon cycle. Currently the greatest uncertainty in modelling the global carbon
cycle is not the fluxes across the ocean atmosphere interface or fluxes relating to net
primary production but in modelling carbon fluxes within the soil. Globally ten times
more carbon dioxide is emitted from soils than from all anthropogenic sources com
bined (Adger and Brown 1995, IPCC 2013). Anthropogenic induced climate change
has caused shifts in both temperature and rainfall patterns across a range of geographic
scales (Rosenzweig et al. 2008). Climate models predict global surface temperatures
will rise by 0.3–4.8 °C by 2100 (IPCC 2014). A major challenge of current ecological
research is to determine how ecosystem processes will respond to future environmental
conditions (Santotja et al. 2017). It is widely acknowledged that food webs play pivotal
roles in carbon cycles, including emissions of carbon dioxide from soils under climate
change but this is rarely considered in modelling carbon fluxes (Pelini et al. 2015),
while understanding of organismic physiological responses under climate change re
mains very rudimentary (Schmitz 2013). One question that is currently being debated
is the extent to which soil animals can be modelled grouped together in broad trophic
categories or whether they need to be considered at a finer level of functional trait or
taxonomic resolution, depending upon their individual responses to changes in micro-
climate (Jiguet et al. 2011, Dixie et al. 2015). Terrestrial isopods potentially form
a useful model system with which to address this question because their behaviour,
physiology, and life histories have been so extensively studied. Carbon dioxide emissions from soils are mediated predominately by microbial me
tabolism (Fig. 1), which, when not constrained by moisture availability, is a function of
temperature (Schlesinger 1977). There is thus the potential for increased soil temperatures
to accelerate emissions of carbon dioxide from soils. The further accumulation of CO2
in the atmosphere could then lead to further increases in temperature, leading to a posi
tive feedback (Fig. 1) (Crowther et al. 2015, Thakur et al. 2018). This process is of most
concern in regions with the highest pools of soil organic matter, such as tundra regions
and temperate grasslands (Melillo et al. Keywords Aggregation, CO2 emissions from soils, feeding behaviour, future rainfall patterns, life history traits,
norms of reaction, response curves, soil animals Abstracth The importance of considering species-specific biotic interactions when predicting feedbacks between the
effects of climate change and ecosystem functions is becoming widely recognised. The responses of soil an
imals to predicted changes in global climate could potentially have far-reaching consequences for fluxes of
soil carbon, including climatic feedbacks resulting from increased emissions of carbon dioxide from soils. The responses of soil animals to different microclimates can be summarised as norms of reaction, in order
to compare phenotypic differences in traits along environmental gradients. Thermal and moisture reaction
norms for physiological, behavioural and life history traits of species of terrestrial isopods differing in their
morphological adaptations for reducing water loss are presented. Gradients of moisture reaction norms for
respiratory rates and thermal reaction norms for water loss, for a species from the littoral zone were steeper
than those for species from mesic environments. Those for mesic species were steeper than for those from
xeric habitats. Within mesic species, gradients of thermal reaction norms for aggregation were steeper for
Oniscus asellus than for Porcellio scaber or Armadillium vulgare, and moisture reaction norms for sheltering
and feeding behaviours were steeper for Philoscia muscorum than for either P. scaber or A. vulgare. These
differences reflect differences in body shape, permeability of the cuticle, and development of pleopodal
lungs. The implications of differences between different species of soil animals in response to microclimate
on the possible influence of the soil fauna on soil carbon dynamics under future climates are discussed. In
conclusion a modelling approach to bridging the inter-disciplinary gap between carbon cycling and the
biology of soil animals is recommended. Mark Hassall et al. / ZooKeys 801: 5–24 (2018) Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 6 Introduction 2017). However, recent evidence indicates cur
rent carbon stores in other soils may also be changing from sinks to sources (Carrera et al. 2011) due to greater increases in soil respiration than in rates of carbon input to the soils. The possible consequences of this positive feedback cycle could potentially be
reduced if soil moisture were to decrease as a result of changes in rainfall patterns
(Fig. 1). Climate models predict that globally rainfall will increase but it is predicted
to vary strongly both spatially and temporally. Spatially, greater increases are projected Interspecific variation in responses to microclimate by terrestrial isopods... 7 7 Figure 1. A conceptual diagram illustrating some of the pathways by which changes in global climate
could potentially impact on rates of carbon dioxide emissions from soils. Both changes in temperature and
in the levels and patterns of rainfall have strong direct effects on the metabolism of bacteria and fungi but
their ecology and metabolism are also regulated by the extent to which they are stimulated by soil animals. Both functional (e.g., behavioural and physiological) responses and numerical (both life history and popu
lation) responses of soil animals are affected by their microclimate. This is in turn affected by larger scale
changes in temperature and rainfall. Therefore, as well as their direct effect on microbial metabolism, these
climatic variables have a strong indirect effect by influencing the behavioural, physiological, life history,
and population processes of soil animals such as isopods. Figure 1. A conceptual diagram illustrating some of the pathways by which changes in global climate
could potentially impact on rates of carbon dioxide emissions from soils. Both changes in temperature and
in the levels and patterns of rainfall have strong direct effects on the metabolism of bacteria and fungi but
their ecology and metabolism are also regulated by the extent to which they are stimulated by soil animals. Both functional (e.g., behavioural and physiological) responses and numerical (both life history and popu
lation) responses of soil animals are affected by their microclimate. This is in turn affected by larger scale
changes in temperature and rainfall. Therefore, as well as their direct effect on microbial metabolism, these
climatic variables have a strong indirect effect by influencing the behavioural, physiological, life history,
and population processes of soil animals such as isopods. Introduction for high latitudes and mid-latitude wet regions but decreases in subtropical dry regions
and many mid-latitude regions (IPCC 2013). Temporally, summer rainfall in temper
ate regions, such as west and south Europe, is predicted to occur in fewer, but more
intense episodes. These will result in longer periods of summer drought which could
potentially constrain temperature-induced increases in soil carbon fluxes, including by
reducing the stimulation of microbial metabolism by moisture dependent soil animals. g
The microbially mediated emissions of carbon dioxide from soils are strongly influ
enced by soil animals acting as key system regulators (Fig. 1) (Anderson 1991, Lavelle Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 8 and Spain 2001, Krab et al. 2010, Coleman et al. 2017). The significance of direct
and indirect consequences of global climate change on ecosystem processes mediated
by invertebrates, including their indirect effects on heterotroph respiration through
interactions with microorganisms, are widely recognised (Pelini et al. 2015). However,
although they are likely to have a major impact on ecosystems in the future, they are as
yet understudied (Blankinship et al. 2011, Carrera et al. 2011, Del Toro et al. 2015).if It is becoming increasingly apparent that the importance of interspecific differences
in key traits not only leads to different species having different functional roles (Dias et al. 2013, Zimmer et al. 2002) but also causes different species to be affected differently by
climate change (Veldhuis et al. 2017). Because ecosystem science has yet to embrace how
principles of evolutionary ecology explain the ways in which organisms mediate ecosys
tem carbon dynamics (Schmitz 2013), there is an urgent need to increase our understand
ing of how differential impacts of climate change on basic biological traits of individual
species can lead to changes in community structure and function (Jiguet et al. 2011). Traits of many soil animals are sensitive to relative humidity, making the soil fauna
vulnerable to their activities being curtailed by changes in soil moisture (Krab et al. 2010, Crowther et al. 2015). Isopods are the most successful group of crustaceans to
have made the transition to the terrestrial environment in terms of their abundance,
distribution, and diversity, and have colonised a wide range of terrestrial habitats from
the littoral zone to deserts (Edney 1954, Warburg 1987). They have become promi
nent members of the arthropod macro-decomposer guild in many ecosystems (Sutton
1980). Introduction 2014) and their eco-physiology (Spencer and
Edney 1954, Edney 1968, Warburg 1987, 1989, Hornung 2011). We used the selected
species to explore possible implications of inter-specific differences in responses to micro-
climate for the role of soil animals in soil processes, including regulation of microbi
ally mediated soil carbon dynamics (Heemsbergen et al. 2004), under predicted climate
change scenarios. Figure 2. A schematic representation of a typical thermal response curve for enzymes (simplified from
Huey and Kingsolver 1989). The temperature optimum is the temperature at which performance reaches
its maximal level or peak performance. The performance breadth defines how steeply peaked (stenother
mal) or broadly plateaued (eurythermal) the response curve is. Any part or the whole of such a curve can
be considered to be a reaction norm of a genotype representing a range of phenotypes expressed across an
environmental gradient, in this example, of temperature. Schmidt and Wägele 2001, Broly et al. 2014) and their eco-physiology (Spencer and
Edney 1954, Edney 1968, Warburg 1987, 1989, Hornung 2011). We used the selected
species to explore possible implications of inter-specific differences in responses to micro-
climate for the role of soil animals in soil processes, including regulation of microbi
ally mediated soil carbon dynamics (Heemsbergen et al. 2004), under predicted climate
change scenarios. Schmidt and Wägele 2001, Broly et al. 2014) and their eco-physiology (Spencer and
Edney 1954, Edney 1968, Warburg 1987, 1989, Hornung 2011). We used the selected
species to explore possible implications of inter-specific differences in responses to micro-
climate for the role of soil animals in soil processes, including regulation of microbi
ally mediated soil carbon dynamics (Heemsbergen et al. 2004), under predicted climate
change scenarios. Introduction Their morphology, physiology, behaviour, life histories, and ecology are known
to be strongly influenced by micro-climates. Isopods therefore form good model soil
animals for testing hypotheses concerning how inter-specific differences in their physi
ological, behavioural and life history traits will lead to interspecific differences in re
sponses to predicted future changes in climate.f In this review we examine the extent to which traits of different species of terres
trial isopods known to have different strategies for reducing water loss in the terrestrial
environment, respond to temperature and relative humidity. Phenotypic responses to a
gradient in the environment can be summarised as norms of reaction. A reaction norm
is the set of phenotypes produced in a range of environments (Schmalhausen 1949,
Stearns 1992) and represents either the whole or part of a response curve of an organ
ism to any environmental gradient as for example illustrated by the thermal response
curve for enzymes shown in Fig. 2. Differences in the gradients of performance response
curves, such as that in Fig. 2, denote how quickly an organism changes its performance
for a given change in an environmental gradient. A steep gradient indicates a strong
response and a high level of phenotypic plasticity over that region of the environmental
gradient. Reaction norms can be very effective in comparing responses not just between
different species as has been demonstrated very clearly by Dias et al. (2013) but also to
different environmental gradients such as soil moisture and temperature.f We investigated differences in reaction norms of physiological, behavioural, and life
history traits in response to differences in temperature and relative humidity for a range
of isopod species chosen to reflect differences in their eco-morphology (Schmalfuss 1984, Interspecific variation in responses to microclimate by terrestrial isopods... 9 9 Figure 2. A schematic representation of a typical thermal response curve for enzymes (simplified from
Huey and Kingsolver 1989). The temperature optimum is the temperature at which performance reaches
its maximal level or peak performance. The performance breadth defines how steeply peaked (stenother
mal) or broadly plateaued (eurythermal) the response curve is. Any part or the whole of such a curve can
be considered to be a reaction norm of a genotype representing a range of phenotypes expressed across an
environmental gradient, in this example, of temperature. Schmidt and Wägele 2001, Broly et al. Behavioural responses to microclimate: thermal norms of reaction for
aggregating behaviour Case study 1. Here we compare differences in aggregating behaviour of three species
of isopod as described by Dixie et al. (2015) who give full details of the methodology
including those of the statistical analyses used.f Differences in the degree of aggregation, at 90% relative humidity, between three
species of isopods differing in their adaptations to reduce water loss in the terrestrial
environment, are shown in Fig. 3. Oniscus asellus (Linnaeus, 1758) was most suscep
tible to water loss, the Armadillidium vulgare (Latreille, 1804) is morphologically the
best adapted to withstand desiccation while Porcellio scaber (Latreille, 1804) has an
intermediate level of morphological adaptation but also uses aggregation as a key be
havioural tactic in its overall desiccation avoidance strategy. Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 10 Figure 3. Aggregation of isopod species differing in desiccation resistance at different temperatures. Mean ±
1 SE aggregation indices (variance:mean ratio) at 90% relative humidity. a P. scaber (F 4,249 = 3.76, p < 0.01
b A. vulgare (F 4,249 = 1.97, P < 0.01) c O. asellus (F 4, 249 = 12.22, P < 0.001) d thermal reaction norms fo
aggregation expressed as quadratic response curves for: P. scaber (dashed line): y = -11.519 + 1.526× - 0.04×2
A. vulgare (solid line): y = -3.534 + 0.574× 0.016×2; O. asellus (dotted line): y = -5.890 + 0.814× – 0.018×2 Figure 3. Aggregation of isopod species differing in desiccation resistance at different temperatures. Mean ±
1 SE aggregation indices (variance:mean ratio) at 90% relative humidity. a P. scaber (F 4,249 = 3.76, p < 0.01)
b A. vulgare (F 4,249 = 1.97, P < 0.01) c O. asellus (F 4, 249 = 12.22, P < 0.001) d thermal reaction norms for
aggregation expressed as quadratic response curves for: P. scaber (dashed line): y = -11.519 + 1.526× - 0.04×2;
A. vulgare (solid line): y = -3.534 + 0.574× 0.016×2; O. asellus (dotted line): y = -5.890 + 0.814× – 0.018×2. The temperature response curve of P. scaber (Fig. 3a, d) resembles the theoretical
one in Fig. Behavioural responses to microclimate: thermal norms of reaction for
aggregating behaviour 2 in being an upwardly convex curve, with a peak in the aggregation index
of 2.93 at 19 °C, a performance breadth at 90% of the peak of 4.7 °C and a gradient
of the reaction norm from 14 °C to 17 °C of + 0.8 increase in aggregation index for
this 3 °C rise in temperature. At temperatures higher than 19 °C the curve of the ag
gregation index decreases due to more of the animals leaving the clumps and moving
around the arena sometimes climbing up the walls in an attempt to escape to find more
favourable lower temperatures (Dixie et al. 2015). Armadillidium vulgare shows a broadly similar pattern of response (Fig. 3b, d), but
with a lower peak of 1.9 in the aggregation index, again at 19 °C. The response curve is
shallower than for P. scaber with a performance breadth of 6 °C at 90% of the peak value
and a gradient for the reaction norm for the aggregation index between 14 -17 °C of +0.3. As for P. scaber, aggregation decreases at temperatures higher that the peak temperature. Interspecific variation in responses to microclimate by terrestrial isopods... 11 The temperature response curve for aggregation of O. asellus (Fig. 3c, d) has a
higher peak aggregation index of 3.35 at a higher temperature of 21.3 °C, a wide per
formance breadth of 6.2 °C at 90% of the peak value and a gradient for its thermal
norm of reaction between 14 -17 °C of 1.5 in the aggregation index over this 3 °C
range. This was approximately twice as steep as for P. scaber and three times as steep as
for A. vulgare. The decline following the peak for O. asellus was less pronounced than
for either of the other two species. Overall these results show that these three species have substantially different pat
terns of thermal response curves for aggregating behaviours, reflecting differences in
their morphological adaptations to terrestrial life. Behavioural responses to microclimate: moisture norms of reaction for
sheltering and feeding behaviours Many terrestrial isopods aggregate in shelter sites, particularly during the day, often un
der stones or pieces of wood where moisture from the soil maintains a more favourable
relative humidity than in more open sites. They then emerge to forage at night when
temperatures are lower and relative humidity is higher. Sheltering behaviour is thus of
central importance in reducing mortality due to desiccation while also reducing the
risk of being eaten by diurnal predators, such as insectivorous birds.f Case study 2. Effects of substrate moisture content and relative humidity on
sheltering and feeding behaviour were investigated by Moss (2007) under controlled
temperature conditions in the laboratory. Three species were studied: A. vulgare,
Philoscia muscorum (Scopoli, 1763) and P. scaber differing in morphological and
physiological adaptations to the terrestrial environment (Dias et al. 2012). The ex
perimental mesocosms (220mm × 150mm × 80mm deep) were lined with plaster
of Paris covered by 50mm sand with a feeding tray at one end and a shelter at the
other, as described in further detail by Moss (2007). Two simulated rainfall re
gimes were used: high simulated rainfall, representing current mean summer daily
rainfall intensity of 1.65 mm day-1 calculated from British Atmospheric Data Cen
tre records for Lacock, Wiltshire, England (51°43'N, 2°11'W) during 1996–2006
when rainfall events occurred on average once every 72h. The low simulated rainfall
regime was half the intensity of the high rainfall regime, 0.83 mm day-1, represent
ing a scenario with 810 ppm concentration of atmospheric CO2 by 2080 which is
predicted to result in a 50% reduction in summer rainfall in south west England
(Hulme et al. 2002). For both regimes the simulated rainfall was administered at the
start of each 72h period using a seedling watering can with 0.6 mm diameter holes
in the rose with the shelter and food trays temporarily covered. Replicate arenas but
without animals were used to monitor percentage moisture content of the substrate
by taking samples of sand each morning and evening. Differences in the moisture
conditions in the mesocosms were caused by both differences in intensity of simu Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 12 12 lated rainfall and the progressive drying out of mesocosms which occurred with time
between rainfall events. Starting moisture level following high rainfall level ≈ 20%
and for low rainfall level ≈ 10%. Behavioural responses to microclimate: moisture norms of reaction for
sheltering and feeding behaviours Eight replicate mesocosms were used for each rainfall treatment, with six identical
control boxes for monitoring substrate moisture content. Each mesocosm contained
twenty individuals of each species. The behaviours of all animals over a 72-hour period
were classified according to their location in the mesocosm and activity categories,
including sheltering and presence in the feeding area. It was not possible to observe
movements of mouthparts so time spent in the feeding area was assumed to be propor
tional to time spent feeding (Dias et al. 2012). Regression analyses of behavioural traits
on substrate moisture were conducted. The results in Fig. 4 and Table 1 show that Ph. muscorum spent significantly less
time sheltering than either of the two other species under the highest moisture condi
tions. It also had the steepest gradient for its moisture reaction norm of -2.16, com
pared with values of -1.05 for A. vulgare and -0.19 for P. scaber (Table 1). Porcellio
scaber showed the least change in the proportion of time it spent sheltering because it
sheltered for more than 95% of the time under all substrate moisture conditions. In
contrast A. vulgare and Ph. muscorum sheltered for 74% and 51% respectively of the
time under the moistest conditions (Table 1). Ph. muscorum also had the steepest gradient for its moisture reaction norm for
feeding of 0.55 (Fig. 4b), which was significantly higher than those for either P. scaber
(0.05) or A. vulgare (0.03) (Table 1). Time spent feeding decreased from 11.5% when
moisture content of the sand was 20%, down to zero under the driest conditions when
none of the three species spent a significant amount of time feeding.i Overall Ph. muscorum showed significantly steeper moisture reaction norms than
did either of the other species just as O. asellus, which similarly lacks pleopodal lungs,
had a steeper gradient for its thermal reaction norm for aggregation than did either
P. scaber or A. vulgare. Life history responses to microclimate: thermal and moisture norms of
reaction for growth and survivorship Case study 3. Both thermal and moisture reaction norms for the key life history traits of
growth and mortality were compared for O. asellus and Porcellio dilatatus Brandt, 1833 by
Dixie et al. (2015) who give full details of the methodology and statistical analyses used.h The 2×2 factorial experimental design for investigating both relative growth rates and
mortality rates permits comparison of thermal and moisture reaction norms simultaneous
ly. Growth rates of O. asellus increased significantly at 5 °C higher temperatures under the
drier, 70% relative humidity conditions but did not grow significantly faster at the higher
temperatures in the moister, 90% humidity (Table 2). Growth of P. dilatatus increased with
temperature to a similar extent under both experimental relative humidity conditions. All
four of the moisture reaction norms for a 20% decrease in moisture are steeper than any Interspecific variation in responses to microclimate by terrestrial isopods... 13 13 Interspecific variation in responses to microclimate by terrestrial isopods... 13
Figure 4. Moisture reaction norms for a) sheltering and b) feeding behaviours with changing sand
moisture content (time spent in behaviour as percentages of total observed behaviours). Lines represent
linear regression models: A. vulgare (solid line) (sheltering: y = 95.24 – 1.05×; feeding: y = 0.32 + 0.03),
P. scaber (dashed line) (sheltering: y = 96.72 – 0.19×; feeding: y= 0.22 + 0.05×), Ph. muscorum (dotted
line) (sheltering: y = 93.14 – 2.16×; feeding: y = 0.36 + 0.55x). Further regression statistics and number
of observations (N), are given in Table 1. Figure 4. Moisture reaction norms for a) sheltering and b) feeding behaviours with changing sand
moisture content (time spent in behaviour as percentages of total observed behaviours). Lines represent
linear regression models: A. vulgare (solid line) (sheltering: y = 95.24 – 1.05×; feeding: y = 0.32 + 0.03),
P. scaber (dashed line) (sheltering: y = 96.72 – 0.19×; feeding: y= 0.22 + 0.05×), Ph. muscorum (dotted
line) (sheltering: y = 93.14 – 2.16×; feeding: y = 0.36 + 0.55x). Further regression statistics and number
of observations (N), are given in Table 1. of the thermal reaction norms for a 5 °C temperature difference for either species. This
indicates a greater sensitivity to a 20% change in relative humidity than to a rise of 5 °C in
temperature, suggesting that perhaps these species might respond more to predicted future
changes in rainfall than to predicted increases in temperature. Life history responses to microclimate: thermal and moisture norms of
reaction for growth and survivorship The difference in gradients of
reaction norms between species (Table 2) reflects the higher rates of growth for P. dilatatus
under all four combinations of temperature and relative humidity conditions. Mortality rates (Table 3) were very low at 90% relative humidity; none of the
O. asellus and very few of the P. dilatatus died at this relative humidity. The gradients
of thermal reaction norms at 90% relative humidity were therefore not significantly Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 14 Table 1. Regression statistics for moisture reaction norms for sheltering and feeding behaviours of three
species of isopods, Armadillidium vulgare, Porcellio scaber, and Philoscia muscorum, in laboratory arenas. Abbreviations: N: number of observations; a and b: parameters of regression equation; †: significant dif
ferences at P < 0.001(t test). Regression statistics
Sheltering
Feeding
N
a
b
t
a
b
t
A. vulgare
1648
95.24
-1.05
-21.84†
0.32
0.03
3.77†
P. muscorum
824
93.14
-2.16
-25.07†
0.36
0.55
15.32†
P. scaber
1648
96.72
-0.19
-8.39†
0.22
0.05
6.84†
Comparison of gradients
Sheltering
Feeding
A. vulgare vs. P. muscorum
18.99†
-30.19†
A. vulgare vs. P. scaber
-25.08†
-3.96†
P. muscorum vs. P. scaber
-44.43†
28.36† Table 2. Relative growth rates (mg g-1 day-1) of Oniscus asellus and Porcellio dilatatus. Three way ANOVA:
temperature F1,72 = 5.15, P = 0.026; humidity F1,72 = 88.62, P < 0.001; species F1,72 = 41.53, P < 0.001. Reac
tion norms are derived from the differences in response to the two temperature and two humidity conditions. Table 2. Relative growth rates (mg g-1 day-1) of Oniscus asellus and Porcellio dilatatus. Three way ANOVA:
temperature F1,72 = 5.15, P = 0.026; humidity F1,72 = 88.62, P < 0.001; species F1,72 = 41.53, P < 0.001. Reac
tion norms are derived from the differences in response to the two temperature and two humidity conditions. Temperature °C
Relative humidity %
Moisture reaction norms
90%
70%
O. asellus
13.5 °C
34 ± 6
2 ± 0.1
-32
18.5 °C
32 ± 2.5
17 ± 9
-15
Thermal reaction norms
-2
15
P. dilatatus
13.5 °C
66 ± 0.5
15 ± 4
-51
18.5 °C
77 ± 8
28 ± 4
-49
Thermal reaction norms
11
13 Table 3. Mortality rates (numbers dying container -1 7days-1) of O. asellus and P. dilatatus. Mann Whit
ney U test for O. Life history responses to microclimate: thermal and moisture norms of
reaction for growth and survivorship asellus: temperature NS, humidity U = 1851 P < 0.001; for P. dilatatus: temperature
U = 2754, P = 0.016, humidity U = 2277, P < 0.001. Reaction norms are derived from the differences in
response to the two temperature and two humidity conditions. Temperature °C
Relative humidity %
Moisture reaction norms
90%
70%
O. asellus
13.5 °C
0
0.73 ± 0.18
0.73
18.5 °C
0
0.58 ± 0.12
0.58
Thermal reaction norms
0
0.46
P. dilatatus
13.5 °C
0.09 ± 0.04
0.78 ± 0.16
0.69
18.5 °C
0.03 ± 0.03
0.18 ± 0.08
0.15
Thermal reaction norms
0.06
0.60 different to zero. Under drier conditions, at 70% relative humidity, more individuals
of both species died at 13.5 °C than at 18.5 °C, leading to steeper thermal reaction
norms at 70% relative humidity than at 90% relative humidity. different to zero. Under drier conditions, at 70% relative humidity, more individuals
of both species died at 13.5 °C than at 18.5 °C, leading to steeper thermal reaction
norms at 70% relative humidity than at 90% relative humidity. Overall, 20% lower relative humidity resulted in steeper moisture reaction norms
for mortality rates than for thermal reaction norms resulting from a 5 °C rise in tem
perature, with O. asellus being more susceptible to drier conditions than P. dilatatus. Interspecific variation in responses to microclimate by terrestrial isopods... 15 Physiological responses to microclimate: temperature and moisture
norms of reaction for respiratory rate and water loss One of the drivers of differences between species of isopods in their behavioural and
life history responses to differences in temperature and moisture is their different
physiological adaptations to the terrestrial environment. In this section we compare
responses of terrestrial isopod species, occurring in a wide range of biomes representing
a gradient of moisture conditions. Moisture reaction norms for differences in respiratory rate for O. asellus, P. scaber,
and A. vulgare are shown in Fig. 4 together with that for Ligia oceanica (Linnaeus,
1767). The pattern of differences between the physiological reaction norms closely
parallels that of behavioural reaction norms (Figs. 3, 4) with the gradient for O. asellus
over a range of a 50% difference in relative humidity being substantially higher than
for P. scaber and A. vulgare. However the gradient for L. oceanica, a littoral species, is
more than double that for O. asellus reflecting the higher stress that would result from
a reduction in relative humidity for a species that lives in a habitat with abundant avail
ability of crevices with saturated micro-climates. y
This interspecific comparison is extended for species from a wider range of habitats
for thermal reaction norms for the physiological process of water loss in Fig. 6. The
pattern for L. oceanica, O. asellus and P. scaber is similar to that for the moisture reac
tion norms for respiratory rate. Ph. muscorum has a higher rate than O. asellus, which,
together with P. scaber, has a slightly steeper moisture reaction norm than either A. vul
gare or Armadillidium nasatum (Budde-Lund, 1885). All five of these species are found
in mesic habitats. The three species of Buddelundia from semi-arid habitats in South
Australia all have lower gradients for their water loss thermal reaction norms than any
of the European species from mesic habitats. Least affected by differences in tempera
ture are species from xeric habitats including those found in the Negev and North Afri
can deserts such as Hemilepistus reaumurii (Milne-Edwards, 1840). The two Armadillo
species and Porcellio olivieri (Audouin, 1826) also occur in desert environments but
not so exclusively, as they are also found in other dry Mediterranean habitats. These
three species have gradients for their thermal reaction norms that are intermediate be
tween those of the desert specialists (including Venezillo arizonicus Mulaik & Mulaik,
1942) and the mesic species. Discussion The range of morphological adaptations in body shape, respiratory surfaces of the
pleopods and cuticle of isopods have been comprehensively described (Edney 1968,
Warburg 1992, Schmalfuss 1984, Hornung 2011, Csonka et al. 2013, Broly et al. 2014) and reflect a moisture gradient in the habitats in which they are found, from
littoral, through mesic to xeric, including desert. It can therefore be predicted that
there will be a parallel gradient in the magnitude of physiological responses to changes
in microclimate reflected in the gradients of thermal and moisture reaction norms. Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 16 Figure 5. Gradients of moisture reaction norms for respiration of isopods differing in their resistance to
desiccation. Reaction norms over the range 50–100% relative humidity for respiratory rates measured as rates
of oxygen uptake (mm3 mm-2 body surface h-1) (Edney 1968). Key to species: Lo Ligia oceanica, Oa Oniscus
asellus, Ps Porcellio scaber, Av Armadillidium vulgare. Key to habitats: littoral (blue), mesic (green). Figure 5. Gradients of moisture reaction norms for respiration of isopods differing in their resistance to
desiccation. Reaction norms over the range 50–100% relative humidity for respiratory rates measured as rates
of oxygen uptake (mm3 mm-2 body surface h-1) (Edney 1968). Key to species: Lo Ligia oceanica, Oa Oniscus
asellus, Ps Porcellio scaber, Av Armadillidium vulgare. Key to habitats: littoral (blue), mesic (green). This hypothesis was supported for respiratory rate moisture reaction norms (Fig. 5),
as L. oceanica, the species with the most primitive morphological adaptations to the
terrestrial environment, had the steepest reaction norm, while O. asellus, which does
not have pleopodal lungs, had a steeper moisture reaction norm than either P. scaber or
A. vulgare, both of which have well developed pleopodal lungs. Similarly, gradients of thermal reaction norms for water loss were substantially
higher for L. oceanica than for the more fully terrestrial species O. asellus and Ph. muscorum, both of which had steeper gradients for their thermal reaction norms than
members of either of the Porcellio species or A. vulgare (Fig. 6). There was a clear trend
of thermal reaction norms for species from semi-arid environments in South Australia
and xeric habitats in Europe, including specialist desert species, all having shallower
thermal reaction norms for rates of water loss. Discussion Thus it can be predicted that those spe
cies from mesic environments will be more sensitive to changes in rainfall patterns,
including longer periods of summer drought, than those from xeric habitats. Given that these patterns of inter-specific differences in physiological traits for
both thermal and moisture reaction norms reflect differences in morphology (Schmal
fuss 1984, Broly et al. 2015), it can be predicted that there would also be a parallel
pattern in thermal and moisture reaction norms for behavioural and life history traits. Aggregation in isopods is a behavioural tactic that reduces respiratory rates and water
loss (Allee 1926), further elaborated by Friedlander (1965), Cloudsey-Thompson and
Constaninou (1987), Caubet et al. (2008), and Broly et al. (2014). This morphologi
cally related hypothesis is supported by both the peak of the thermal response curve
and the gradient of the thermal reaction norm for aggregation being higher for O. asel
lus than for either P. scaber or A. vulgare. Interspecific variation in responses to microclimate by terrestrial isopods... 17 17 Dif
ferences in the gradients of reaction norms of life history traits could thus result in
differences in the way abundances of these species might change under future climates. f
An important conclusion of this overview is that, due to inter-specific differences
in morphological, physiological, behavioural and life history traits, different species of
isopods are likely to respond very differently to predicted changes in global climate. In
terspecific differences in response to changes in temperature (Hickling et al. 2006) are
now well documented for other taxa such as birds (Pearce-Higgins et al. 2015), Lepi
doptera (McDermott Long et al. 2017), Orthoptera (Buckley et al. 2015) and Odo
nata (Stoks et al. 2014). In contrast, much less is known about ecological responses to
predicted changes in rainfall patterns (Urban et al. 2016). Such responses are likely to
be particularly pronounced for soil animals due to their activity and ecology being so
strongly influenced by soil moisture, as shown by Blankinship et al. (2011) who found
that precipitation limited all taxa and trophic groups in forest floors. l
If the abundance of some species declines, it is not yet known whether other spe
cies, less sensitive to changes in microclimate, will respond by expanding their realised
niches. It is known that interspecific competition for high quality foods between dif
ferent species of isopods does occur in the field (Hassall and Dangerfield 1989) so it is
possible that total guild densities may change less than species composition of guilds
for macro-decomposers. It is known that differential impacts on basic biological pa
rameters of individual species can modify fundamental characteristics of community
structure and function (Jiguet et al. 2011). For isopods it is possible that the func
tional role of the whole isopod guild in stimulating microbial activity may alter less as
a result of future climate change than might be expected from effects on particularly
sensitive individual species. If other species in the guild, less sensitive to changes in
micro-climate, expanded their realised niches in response to the decline of a more
sensitive species, this could buffer consequences of climate change for overall decom
position rates. It is yet to be tested whether net community level responses, resulting
from differential impacts of climate change on individual species, will impact on the
extent to which future changes in rainfall patterns might mitigate potential increased
temperature/ induced carbon dioxide emission feedbacks. Figure 6. Thermal reaction norm gradients for evaporation rate (water loss) for isopods from biomes differ
ing in availability of moisture. Evaporation rate (g g-1 h-1× 10-2) standardised to a temperature range of 3.5 °C
(from Edney 1951; Warburg 1965, 1987, 1989). Key to species: Lo Ligia oceanica, Ph Philoscia muscorum, Oa
Oniscus asellus, Ps Porcellio scaber, Av Armadillidium vulgare, Ao Armadillo officinalis, Aa Armadillo albomargi
natus, Po Porcellio olivieri, An Armadillium nasatum, Ba Buddelundia albinogrisescens, Bf Buddelundia frontosa,
Va Veneziillo arizonicus, Bsp Buddelundia spp. probably lateralis, Hr Hemilepistus reaumurii, Sf Schizidium fes
tai. Key to habitats: littoral (blue), mesic (green), xeric (yellow), semi-arid (orange) habitats in South Australia. Sheltering behaviour is another tactic evolved in isopods to help reduce water loss
by taking refuge in more humid shelter sites. Again, Ph. muscorum had a significantly
higher gradient for its sheltering moisture reaction norm than either P. scaber or A. vul
gare. Similarly, feeding behaviour moisture reaction norms of Ph. muscorum decreased
more steeply than for either of the other two species. Thus within three species, all
typical in mesic environments, there was a consistent trend in all of these behavioural
traits, with species without pleopodal lungs being more sensitive to changes in micro
climate than species better adapted to resist desiccation. This again indicates that there
are significant inter-specific differences which suggest those species least adapted to
the terrestrial environment might be more susceptible to potential changes in micro-
climate resulting from changes in future patterns of rainfall. It would be logical to predict that such a trend might also apply to growth rates but
that was not supported by the data because thermal reaction norms were very similar
for O. asellus and P. dilatatus while the gradient for moisture reaction norms at both
temperatures were higher for P. dilatatus. P. dilatatus had higher growth rates under all
four combinations of temperature and relative humidity, possibly because the microcli
mate actually experienced by P. dilatatus was moister than that experienced by O. asel
lus due to P. dilatatus behavioural trait of burrowing into the sand substrates thereby
being subject to a higher relative humidity in its immediate microclimate. 18 Mark Hassall et al. / ZooKeys 801: 5–24 (2018) These life history traits of growth and survivorship are important correlates of
fitness and have a very strong influence on the population dynamics of isopods. It is widely agreed however,
more focus is required on biotic interactions to clarify the potential feedbacks between
climate change and soil carbon dynamics (Carrera et al. 2011). The importance of species-specific responses to climate change has been highlighted
for Collembola by Krab et al. (2010) and Makkonen et al. (2011), who found that in
terspecific faunal trait variation provided a valuable tool in predicting animal responses
to climate change. However, it is noteworthy that Crowther et al. (2015) conclude that
the regulatory effects of interspecific interactions are rarely considered in climate feed
back studies. Pelini et al. (2015) extend this conclusion further by arguing that without
isolating and including the significant impact of invertebrates, climate models will be
incomplete, hindering well-informed policy decisions. Because the adaptive value of
fundamental biological traits in isopods have been so comprehensively studied and Interspecific variation in responses to microclimate by terrestrial isopods... 19 because isopods form such a prominent component of so many soil macro-arthropod
communities, we propose that terrestrial isopods represent excellent model systems for
further investigation of species specific responses to predicted climate change and their
consequences for soil carbon dynamics. What are the most pressing needs for future research in this field? Our understanding of the global carbon cycle is predominantly encapsulated in mod
els. Our understanding of terrestrial isopod biology, in contrast, is mostly based on
results of empirical studies. There is a very important need to bridge the gap between
these two contrasting approaches and methodologies in these completely different dis
ciplines. Considering the whole series of symposium volumes on the Biology of Ter
restrial Isopods from 1984 until 2018, models may qualify as an endangered species. We know that carbon dynamics in the soil is the least well understood part of the
global carbon cycle. Whilst, it is extremely complicated, models by definition are sim
plifications of reality. Models can never represent the full complexity of the real world,
that is not their function, but what they have are both holistic and heuristic properties. Holistic in that it should be easier to appreciate emergent properties of a system as a
whole from a model of it, rather than from detailed studies of individual components. Heuristic in that models should generate testable predictions that cannot be made on
the basis of studying individual components in isolation. Terrestrial isopods are soil animals about which we have a wealth of knowledge based
on empirical studies on their anatomy, behaviour, physiology, life history, and ecology. What we do not have is integrative models of different aspects of their biology that can
then be interfaced with those of other soil animals and ultimately with those of micro-or
ganisms. This integrative approach would significantly contribute to our understanding
of how global climate change will affect the soil component of the global carbon cycle. Where can we start? Models do not necessarily have to be numerical, at least initially. A topographic map is
a model of a landscape. At a glance, that piece of paper only tens of cm wide, shows us
where hills, mountains, rivers and bridges are in landscapes at much larger scales. By us
ing a map we can predict how best to travel from A to B. What is very urgently needed
in soil biology is some comparably simplified maps of the interactions involving the soil
animals under our feet. A starting place could be for the context of many future studies
to be introduced using conceptual box and arrow models of the system under study
(Fig. 1). Our skills as scientists lie in how effectively we can simplify the real world we
are studying by being brave enough to make a whole array of untested assumptions.h g
g
g
The next stage could then be to parameterise these models using the extensive
empirical data available in the literature, making assumptions we know to be simplistic Mark Hassall et al. / ZooKeys 801: 5–24 (2018) 20 but which enable us to make further quantitative predictions. Progressively testing
these predictions experimentally to validate the models further could then increase our
understanding of the system. g
A problem will always be integrating models representing different levels of or
ganisation which, from previous studies, range from molecules to communities. These
may initially appear to be measured in different currencies but ultimately, while we are
working within the paradigm of neo-Darwinian evolutionary theory, the answer to this
problem may be in using fitness, or at least fitness correlates, to equate different traits at
the individual level. Moving from the individual level to the ecosystem function level
could then at least be based on a sound theoretical foundation.h The “functional traits” approach (Moretti et al. 2017) may be one way forward. Splitting whole trophic groups, treated as homogeneous entities in some large scale
models, into functional groups has been an important step in the right direction. Tak
ing account of interspecific differences in traits within functional groups and incor
porating the diversity of responses into systems models could be a further important
step. Where can we start? It would appear that the new generation of isopod biologists is now uniquely well
placed to do this, thus giving them the potential to fill an extremely important gap in
21st Century science in a way that is a meaningful contribution to our understanding
of the effects mankind is having by interfering with the global carbon cycle and thus
global climate systems. We wish younger colleagues all success with such a challenging
venture at such an exciting time and best wishes with carrying the science of terrestrial
isopod biology to new horizons. Acknowledgements MH is very grateful to Stephen Sutton for initiating, stimulating, and encouraging his
interest in the biology of terrestrial isopods. We also thank Martin Zimmer for suggest
ing an earlier version of this paper, NERC for a studentship for AM, and to the guest
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* 이 논문은 2019년 대한민국 교육부와 한국연구재단의 인문사회분야 신진연구자지원사업의 지원을 받아
수행된 연구임(NRF-2019S1A5A8038225).
†교신저자 : 정보영, 현대자동차 인재개발팀 책임매니저, viautumn@gmail.com 한국심리학회지: 산업및조직
Korean Journal of Industrial and Organizational Psychology
2021. Vol. 34, No. 1, 51-79
https://doi.org/10.24230/kjiop.v34i1.51-79 한국심리학회지: 산업및조직
Korean Journal of Industrial and Organizational Psychology
2021. Vol. 34, No. 1, 51-79
https://doi.org/10.24230/kjiop.v34i1.51-79 한국심리학회지: 산업및조직
Korean Journal of Industrial and Organizational Psychology
2021. Vol. 34, No. 1, 51-79
https://doi.org/10.24230/kjiop.v34i1.51-79 한국판 칼레이도스코프 경력태도 측정도구 타당화 연구* 정 보 영†
현대자동차 정 보 영†
현대자동차 이 재 은
상지대학교 이 재 은
상지대학교 이 연구의 목적은 Sullivan과 그의 동료들(2009)이 개발한 칼레이도스코프 경력태도 척도를 우
리나라의 기업근로자 대상으로 타당화하는 데 있다. 이를 위해 수정델파이법을 통해 내용타
당도를 검증하고, 남녀 기업근로자를 대상으로 2개의 표본자료를 수집하여(N=347, N=631)
구인타당도, 변별타당도, 준거타당도를 검증하였다. 연구결과는 다음과 같다. 첫째, 수정델파
이법을 통해 번안된 칼레이도스코프 경력태도 측정문항의 내용타당도를 확인하고 국내 기업
근로자에게 적용할 수 있는 3문항을 개발하였다. 둘째, 칼레이도스코프 경력태도 측정문항은
원척도와 같이 3개 요인으로 구성되었으며, 최종 13개 문항으로 구성된 측정도구의 적합도
가 적절하게 도출되어 이를 채택하였다. 셋째, 교차타당도 분석 결과 새로운 표본(N=631)에
서 13개 문항의 측정모형 적합도가 양호하였으며 남녀 성별 집단에서도 동일한 척도로 활용
될 수 있음을 확인하였다. 한편, 칼레이도스코프 경력태도 중 균형과 도전 인식에서 성별 집
단의 잠재평균 차이가 유의한 것으로 나타났다. 연구결과를 종합하여 한국판 칼레이도스코
프 경력태도가 타당하고 신뢰할 수 있는 척도임을 밝혔으며, 이를 토대로 연구의 의의와 한
계점, 향후 연구를 위한 제언 등이 논의되었다. 주요어: 칼레이도스코프경력, 만화경경력, 측정도구, 타당화, 잠재평균분석 * 이 논문은 2019년 대한민국 교육부와 한국연구재단의 인문사회분야 신진연구자지원사업의 지원을 받아
수행된 연구임(NRF-2019S1A5A8038225). †교신저자 : 정보영, 현대자동차 인재개발팀 책임매니저, viautumn@gmail.com - 51 - 한국심리학회지: 산업및조직 고 있다(McMahon, 2014; Savickas, 2012). 그러나
국내 뿐 아니라 국외에서도 칼레이도스코프
경력에 대한 연구는 아직까지 크게 확산되지
못하고 있는 실정이다. 이는 칼레이도스코프
경력이 특정 시점에서의 근로자 경력태도를
확인하고 이와 관련된 다양한 변인이 무엇인
지 밝히는 것에 초점을 두고 있기 보다는 근
로자의 생애에 걸친 경력패턴의 변화를 설명
하는 것을 목적으로 하고 있기 때문이라 짐작
된다. 최근 많은 근로자들이 경험하고 있는 급격
한 환경변화는 개인의 경력에 대한 태도와 경
력개발의 기준을 모두 바꾸어 놓고 있다. 조
직 중심의 경력개발에서 개인 주도의 경력개
발로 구심점이 이동한 경력 패러다임의 변화
는 더 이상 새로운 관점이 아닌 것으로 여겨
지고 있다. 조직 구성원들의 경력에 관한 태
도와 경력개발 의사결정 등과 관련하여 새로
운 개념 제시와 관점 채택이 필요한 시점이다. 이러한 경력 패러다임의 변화와 함께 국내외
경력 연구자들은 새로운 경력태도에 관심을
갖고 연구해 왔다. 새로운 경력태도와 관련하
여 프로티언 경력(protean career), 무경계 경력
(boundaryless career)과 함께 자주 거론되는 모
형이 Mainiero와 Sullivan(2005)이 제안한 칼레이
도스코프 경력모형(Kaleidoscope Career Model:
KCM)으로(Baruch, 2007; Sullivan & Baruch,
2009), 국내에서는 만화경 경력모형, 또는 만
화경 경력태도로 번안되기도 하였다. Mainiero
와 Sullivan(2005)은 환경변화에 따라 기존의
경력 개념에 변화가 필요하다고 설명하면서,
만화경이 반사에 의해 무한한 패턴을 만드는
것과 같이 개인은 경력과 관련하여 3개의 주
요한 경력 파라미터(career parameter: 진정성
(authenticity), 균형(balance), 도전(challenge))를 회
전시키며 다양한 경력 패턴을 만들어 간다고
보았다. 한국판 칼레이도스코프 경력태도 측정도구 타당화 연구* 이를 통해 향후 칼레이도스코프 경
력태도 관련 연구의 방향을 제시하고자 한다. 를 확인할 수 있을 것이다. 이 연구는 우리나라 기업근로자를 대상으로
하여 칼레이도스코프 경력태도 측정도구를
타당화하는 것을 목적으로 한다. 이를 위해
Sullivan, Forret, Carraher와 Mainiero(2009)가 개발
한 칼레이도스코프 경력태도 측정도구를 번안
하여 내용타당도를 확인하고, 탐색적 요인분
석과 확인적 요인분석을 통해 구성개념 타당
도를 검증하고자 한다. 또한 칼레이도스코프
경력태도의 구성개념과 관련된 일과 삶의 균
형, 프로티언 경력태도, 도전적 직무행동, 경
력만족, 고용가능성, 경력몰입 등의 변인과의
관계를 분석하여 준거관련 타당도를 확인하고
자 한다. 이를 통해 향후 칼레이도스코프 경
력태도 관련 연구의 방향을 제시하고자 한다. 이론적 배경 이론적 배경 한국판 칼레이도스코프 경력태도 측정도구 타당화 연구* 즉 개인은 자신의 생애 경력발달 단
계에서 나타날 수 있는 정체성 갈등을 해결해
나가면서 경력 발전의 중요한 축이 되는 경력
파라미터를 개인의 정체성에 맞게 조정하며
경력과 관련된 의사결정을 수행해간다는 것이
다(Kirk, 2016). 칼레이도스코프 경력모형은 삶
의 전체적인 맥락 안에서 구성주의적 관점을
갖고 개인의 경력개발에 초점을 두어야 한다
는 최근 경력 관련 연구자들의 주장을 수용하 선행연구에서 칼레이도스코프 경력태도는
주로 특정 집단의 경력에 대한 태도를 이해하
거나 경력 관련 행동에 차이가 있는 두 집단
의 특성을 비교할 목적으로 활용되어 왔다. 예를 들어, 여성(August, 2011; Cabrera, 2007;
Shaw & Leberman, 2015), 기업가(Sullivan, Forret,
Mainiero, & Terjesen, 2007), 교원(Carraher,
Crocitto & Sullivan, 2014)의 칼레이도스코프 경
력태도를 구명하고자 한 연구가 수행되었으며,
성차(Mainiero & Gibson, 2018)나 세대차(Sullivan,
Forret, Carraher, & Mainiero, 2009)를 밝히고자
한 연구들이 있었다. Sullivan과 Baruch(2009)는
개인의 경력 관련 의사결정에 있어 진정성,
균형, 도전의 속성이 점점 더 중요해지고 있
다고 설명하였으나, 서구 문화권에서 다양한
집단을 대상으로 칼레이도스코프 경력모형이
적용되어 온 것과 달리 국내 근로자를 대상으
로 칼레이도스코프 경력태도를 측정하거나
칼레이도스코프 경력모형을 적용한 연구는
매우 부족한 실정이다. 이에 우리나라 근로
자에게도 칼레이도스코프 경력을 구성하는
세 개의 경력 파라미터가 유효한지 검증할
필요가 있다. 경력 패러다임의 변화와 함께 나타난 근로
자의 새로운 경력태도를 대표하는 프로티언 - 52 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 경력태도는 주도적으로 자신의 가치에 기초하
여 경력을 개발하는 개인의 태도(Hall, 2004)로
자기주도적 경력관리와 개인가치 지향이라는
두 가지 하위요인으로 구성된다. 많은 연구에
서 프로티언 경력태도는 고용가능성, 주관적
경력성공, 경력만족, 직무만족, 조직몰입 등
개인의 긍정적인 경력, 조직 태도와 정적으로
관련되어 있어(Lee, Park, & Kim, 2019), 프로티
언 경력이 새로운 경력 패러다임 시대에 적응
적인 경력태도임을 밝힌 바 있다. 이 연구의
주요 관심인 칼레이도스코프 경력태도는 프로
티언 경력태도에서 강조하고 있는 ‘개인 가치
지향’의 실체를 보다 상세하게 설명해 준다는
측면에서 의의가 있다. 칼레이도스코프 경력
에서 강조하고 있는 세 가지 경력 파라미터가
개인이 지향하는 구체적인 가치가 될 수 있기
때문이다. 칼레이도스코프 경력은 ‘현대사회에
서 경력과 관련하여 개인이 추구하는 가치는
무엇인가?’, ‘이러한 가치는 전체 경력개발의
단계에서 어떻게 달라지는가?’, ‘개인의 경력
과 관련하여 변화된 가치는 경력관련 태도와
행동에 어떠한 영향을 미치는가?’ 등의 질문에
답변을 가능하게 한다. 를 확인할 수 있을 것이다. 이 연구는 우리나라 기업근로자를 대상으로
하여 칼레이도스코프 경력태도 측정도구를
타당화하는 것을 목적으로 한다. 이를 위해
Sullivan, Forret, Carraher와 Mainiero(2009)가 개발
한 칼레이도스코프 경력태도 측정도구를 번안
하여 내용타당도를 확인하고, 탐색적 요인분
석과 확인적 요인분석을 통해 구성개념 타당
도를 검증하고자 한다. 또한 칼레이도스코프
경력태도의 구성개념과 관련된 일과 삶의 균
형, 프로티언 경력태도, 도전적 직무행동, 경
력만족, 고용가능성, 경력몰입 등의 변인과의
관계를 분석하여 준거관련 타당도를 확인하고
자 한다. 칼레이도스코프경력모형 세 개의 파라미터는
진정성(authenticity), 균형(balance), 도전(challenge)
으로 구성되며, 진정성은 진정한 자기 자신이
되는 것으로, 균형은 일(work)과 일 외적인 요
구 사이의 균형으로, 도전은 개인을 자극하는
일과 경력에서의 발전으로 정의된다. 이러한
세 개의 파라미터는 생애주기 동안 모두 활성
화되어 있으나, 특정 시기에 어떤 파라미터가
강하게 나타나는가에 따라 개인의 경력 패턴
이 결정되게 된다. 보다 구체적으로 개인은
경력 파라미터에 기반하여 직무요구, 제한, 기
회와 개인이 얻게 될 가치, 이익, 관계 등을
비교하며 최적의 상태(best fit)를 추구한다는
것이다. 즉 개인의 생애에 걸친 경력 의사결
정과 그 결과로 나타나는 경력패턴을 경력 파
라미터를 중심으로 이해할 수 있다는 데 모형
의 의의가 있다. 칼레이도스코프 경력태도는 Sullivan과 그의
동료들(2009)이 개발한 측정도구를 통해 측정
할 수 있다. 칼레이도스코프 경력 측정도구는
세 개의 하위요인과 15문항으로 구성되어 있
으며, (1) 나를 전혀 표현하지 못하고 있다 부
터 (5) 나를 매우 잘 표현하고 있다 까지 자기
보고식 5점 LIkert 척도로 측정하게 된다. Sullivan과 그의 동료들(2009)의 연구는 칼레이
도스코프 경력태도의 세대 간 차이를 확인하
기 위한 목적으로 실시되었는데, 칼레이도스
코프 경력태도 측정도구의 구인타당도는 보고
되지 않았다. 다만 각 경력파라미터의 내적
일치도는 진정성 .76, 균형 .81, 도전 .84로 보
고되었다. 또한 경력 파라미터 간 상관관계의
범위는 .48-.56으로 중간 정도의 상관을 보였
다. 이 연구에서는 세대차 외에도 통제변인으
로 성별, 자녀 수, 혼인상태, 교육수준, 소득과
칼레이도스코프 경력의 상관관계도 분석하였
는데 남성에 비해 여성이 진정성(.14)과 균형
(.08)에서 유의한 정적 상관관계를 보였고, 자
녀 수와 진정성(.08), 균형(.27), 도전(.07) 간 상
관관계가 유의하였으며, 기혼일 때 균형(.18)과
의 상관관계가 유의하였다. 또한, 교육수준과 칼레이도스코프경력모형 칼레이도스코프 경력모형은 처음 제안된 당
시 여성의 경력경로 발달의 특징을 깊이 조명
하고자 하였다. 특히 기존의 전통적인 경력
경로가 아닌 새로운 경력 패러다임을 강조하
면서 경력과 관련된 남녀의 차이를 주요하게
다루었다. Mainiero와 Sullivan(2006)은 남성의 경
우 하나 혹은 두 개의 조직에서 일하는 경우
뿐만 아니라 다양한 조직을 거치더라도 상향
이동을 지향하는 전통적인 경력패턴을 갖는
반면, 대다수 여성들의 경력이동은 비선형적
이며 노동시장을 떠났다가 재진입하는 등 이
동을 반복하는 경향을 갖고 있다고 강조하면
서 칼레이도스코프 경력모형을 통해 여성의
경력 의사결정을 설명할 수 있다고 하였다. 칼레이도스코프 경력태도에서 강조하고 있
는 도전, 진정성, 일과 삶의 균형 등은 최근
일터에서의 경험, 리더십, 일-가정 균형 등 구
성원들의 일과 관련된 태도에서 중요한 가치
로 다루어지고 있다. 따라서 칼레이도스코프
경력태도가 높은 근로자일수록 보다 효과적인
경력 관련 태도 및 행동을 보일 가능성이 높
을 것으로 판단된다(Kim & Lee, 2020). 이 연구
에서는 칼레이도스코프 경력태도의 응답값이
경력 관련 긍정적인 태도 및 행동과 유의한
관계를 갖는지 검증하고자 한다. 이를 통해
칼레이도스코프 경력태도의 크기가 갖는 의미 - 53 - 한국심리학회지: 산업및조직 칼레이도스코프 경력모형(KCM)은 세 개의
평면거울이 반사에 의해 무한한 패턴을 만드
는 것과 같이 개인이 경력 관련 의사결정을
내릴 때, 세 개의 주요 경력 파라미터(career
parameter)를 회전시키며 다양한 경력 패턴을
만들어간다고 설명한다. 세 개의 파라미터는
진정성(authenticity), 균형(balance), 도전(challenge)
으로 구성되며, 진정성은 진정한 자기 자신이
되는 것으로, 균형은 일(work)과 일 외적인 요
구 사이의 균형으로, 도전은 개인을 자극하는
일과 경력에서의 발전으로 정의된다. 이러한
세 개의 파라미터는 생애주기 동안 모두 활성
화되어 있으나, 특정 시기에 어떤 파라미터가
강하게 나타나는가에 따라 개인의 경력 패턴
이 결정되게 된다. 보다 구체적으로 개인은
경력 파라미터에 기반하여 직무요구, 제한, 기
회와 개인이 얻게 될 가치, 이익, 관계 등을
비교하며 최적의 상태(best fit)를 추구한다는
것이다. 즉 개인의 생애에 걸친 경력 의사결
정과 그 결과로 나타나는 경력패턴을 경력 파
라미터를 중심으로 이해할 수 있다는 데 모형
의 의의가 있다. 환경, 맥락적 상황에 따라 변화한다. 경력태도
는 개인의 경력과 관련된 특정한 행동을 예측
하는 변인으로 다루어져 왔으며, 주요 연구도
경력태도의 결과변인에 초점을 맞추어 이루
어져왔다. 선행연구에서는 주로 전통적 경력
태도(traditional career attitude), 프로티언 경력
태도(protean career attitude), 무경계 경력태도
(boundaryless career attitude) 등 경력과 관련된
특정한 관점을 강조해 왔다. 이 연구에서는
칼레이도스코프 경력모형에서 제시된 칼레이
도스코프 경력태도에 초점을 맞추고 경력태도
를 설명하고자 한다. 칼레이도스코프 경력모형(KCM)은 세 개의
평면거울이 반사에 의해 무한한 패턴을 만드
는 것과 같이 개인이 경력 관련 의사결정을
내릴 때, 세 개의 주요 경력 파라미터(career
parameter)를 회전시키며 다양한 경력 패턴을
만들어간다고 설명한다. 칼레이도스코프경력태도의측정 경력태도(career attitude)는 경력에 대한 신념,
지식, 사고 등 인지적 요소와 선호되는 경력
인지 아닌지에 대한 평가적 요소 또는 정서적
요소, 특정 방향으로 행동하려는 행동 의도
및 경향에 해당하는 행동적 요소를 포괄하는
개념이다. 태도는 다양한 환경에 의해 학습되
고, 특정한 행동을 유발하며, 특정한 사물, 사
건에 대해 일관된 선호 또는 비선호를 갖게
하는 특징이 있다(Fishbein & Ajzen, 1975). 또한,
태도는 가변적 속성을 갖고 있어 개인의 경험, - 54 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 진정성(.07), 도전(.10) 간 상관관계가 유의하였
으며, 소득수준은 균형(.07)과 도전(.11) 간 상
관관계가 유의한 것으로 보고되었다. Shaw와 Leberman(2015)은 뉴질랜드의 여성 CEO
7명을 대상으로 한 심층 인터뷰를 통해 경력
파라미터의 하위 주제를 확인하여 칼레이도
스코프 경력모형을 확장하였는데, 구체적으로
진정성에는 열정과 관계구축이, 균형에는 자
기인식과 조직 내 영향이, 도전에는 기회수용
과 남성중심 환경에서의 업무수행이 도출되
었다. 한편, 최근 수행된 Kim과 Lee(2020)의 연구
에서 칼레이도스코프 경력태도를 측정하고 구
인타당도를 확인하였다. 이 연구는 Mainiero와
Gibson(2017)의 16문항으로 구성된 칼레이도스
코프 측정도구를 활용하였는데, 탐색적 요인
분석 과정에서 도전과 균형의 측정 문항이 부
적으로 할당되어 선행연구에서 제시하고 있는
칼레이도스코프 경력태도의 요인 구조를 확인
하는 데 한계가 있었다. 정량적 접근만 활용
했을 때 나타나는 이러한 한계점을 극복하기
위해, 이 연구는 델파이 조사를 실시하여 칼
레이도스코프 측정도구 번안 내용을 검토하고
국내 근로자들의 경력개발 환경에 적용할 수
있는 문항을 새롭게 추가하고자 하였다. 한편 칼레이도스코프 경력태도의 세 가지
경력 파라미터에 대한 태도를 각각 비교하여
성차나 세대차와 같이 집단 간 차이가 있음을
밝히는 데 초점을 둔 연구들도 수행되었다. 먼저 성차와 관련된 연구를 살펴보면, Sullivan
과 Mainiero(2007)는 응답자 중 대부분(84%)의
남성들은 초기-중기-말기의 경력단계에서 도전
-진정성-균형에 각각 순서대로 초점을 두는 알
파 칼레이도스코프 경력 패턴을 갖고 있는 반
면, 대부분의 여성(83%)들은 초기에는 도전,
중기에는 균형, 말기에는 진정성에 초점을 두
는 베타 칼레이도스코프 경력 패턴을 보였다
고 설명하였다. Cabrera(2009)도 25명의 여성을
심층 인터뷰한 질적 연구를 통해 베타 경력
패턴이 여성에게 있어서 가장 많이 나타나
는 패턴임을 밝혔다. 최근 수행된 Mainiero와
Gibson(2017)의 연구에서는 경력중기의 남녀
실업자를 대상으로 칼레이도스코프 경력태도
의 차이를 밝혔는데, 여성은 남성에 비해 균
형을 훨씬 중요하게 생각하고 있었으며, 경력
후기로 갈수록 남녀 모두 진정성을 더 강조
하는 것으로 나타났다. Elley-Brown, Pringle과
Harris(2018)가 교육분야 뉴질랜드 여성 직업인
을 대상으로 한 연구에서는 경력 중기 여성에
게서 알파 경력에 대한 선호가 관찰되었으며,
진정성은 여성에게 여전히 중요한 파라미터임
을밝혔다 칼레이도스코프경력태도관련선행연구 칼레이도스코프 경력태도를 활용하여
차이를 밝힌 연구는 제한적이나 직업가
경력성공에 대한 준거에 대한 연구를 통
레이도스코프 경력태도의 성차를 유추
있다. 예를 들어, Parker와 Chusmir(1992)
력성공의 준거를 성별에 따라 비교하였
여성은 남성에 비해 가족관계, 개인적
직업안정성 등을 중요하게 생각하고
에 대한 중요성은 낮게 평가하였다. D
Murphy(2006)의 연구에서는 여성은 남성
해 균형과 관계를 중요한 직업적 성공
여기고 있었다. 국내 연구에서는 Kim과
(2014)가 성별에 따라 경력성공 준거에
인식에 차이가 있음을 밝혔는데, 여성은 칼레이도스코프 경력모형을 적용한 세대차
연구도 수행되었다. Sullivan과 그의 동료들
(2009)은 X세대의 경우 베이비부머 세대에 비
해 진정성과 균형에 대한 높은 요구를 갖고
있는 것으로 나타났으나 도전에 대한 세대
차는 발견되지 않았다고 설명하였다. August
(2011)는 경력후기 여성을 대상으로 질적 연구
를 수행한 결과 여성은 상대적으로 젊을 때
도전을 더 강조하는 것으로 나타났으며, 경력
후기로 갈수록 진정성과 균형을 강조하고 있
다고 설명하였다. 프 경력태도의 성별, 세대별 집단 간 차이를
다룬 연구에서도 칼레이도스코프 경력태도의
각 경력 파라미터의 평균 차이를 분석하고 있
다. 이는 각 집단에서의 동일한 측정을 가정
한다. 그러나 실제로 이러한 집단 간에(예. 남
성 vs. 여성) 칼레이도스코프 경력에 대한 측
정동일성이 검증되지 않았다는 점에서 한계가
있다. 칼레이도스코프 경력태도와 성차
칼레이도스코프 경력모형은 남성과 다른 여
성의 경력태도를 제안하기 위한 목적으로 처
음 발표되었으므로 칼레이도스코프 경력태도
의 성차는 선행연구에서 중요하게 다루어져
왔다. 그러나 칼레이도스코프 경력태도의 차
이를 실증적으로 검증한 연구는 제한적으로
수행되었다. 예를 들어 Mainiero와 Gibson(2017)
은 미국의 남녀 경력 중기 실업자(N=744)를
대상으로 칼레이도스코프 경력태도의 성차를
검증하였는데, 경력중기 여성은 남성보다 균
형을 더 강조하는 것을 밝혔다. 칼레이도스코프 경력태도와 관련된 선행연
구를 종합해보면, 칼레이도스코프 경력모형이
제안된 이후 새로운 기업 환경에서의 경력을
설명하기에 칼레이도스코프 경력 모형이 적절
한 지에 대한 연구와 경력과 관련된 태도에서
의 성차, 세대차를 칼레이도스코프 경력모형
으로 설명할 수 있는지에 초점을 맞추고 연구
들이 수행되어 온 것을 알 수 있다. 특히 여
성의 생애사건과 경력단계에 따른 경력 파라
미터의 변화를 분석한 질적 연구들이 많이 수
행되었다. 그러나 다음과 같은 한계점이 관찰
된다. 첫째, 칼레이도스코프 경력태도의 측정
도구가 여러 문화에 걸쳐 타당한지 측정도구
를 타당화 한 연구는 찾아보기 어렵다. 둘째,
칼레이도스코프 경력모형이 현대인의 경력 태
도를 설명하는데 타당한 지 확인하기 위해 특
정 집단을 대상으로 한 질적 연구 방법에 초
점을 두고 있어 다양한 대상을 포괄한 양적
연구의 수행이 요구된다. 셋째, 칼레이도스코
프 경력태도의 성차와 관련하여 여성이 알파
경력패턴을 갖는다는 결과와 베타 경력패턴을
갖는다는 상반된 연구결과가 존재하므로 보다
다양한 맥락적 변인을 종합적으로 고려한 연
구설계가 필요하다. 마지막으로 칼레이도스코 칼레이도스코프 경력태도를 활용하여 성별
차이를 밝힌 연구는 제한적이나 직업가치나
경력성공에 대한 준거에 대한 연구를 통해 칼
레이도스코프 경력태도의 성차를 유추할 수
있다. 칼레이도스코프경력태도관련선행연구 Mainiero와 Sullivan(2005)이 처음으로 칼레이
도스코프 경력모형을 제안한 이후 이를 기반
으로 한 양적, 질적 연구와 이론 연구들이 다
수 수행되어 왔다. 특히 후속연구들은 세 개
의 경력 파라미터가 실제로 존재하는 지에 대
한 근거를 제시하였는데, Sullivan과 그의 동료
들(2009)은 928명을 대상으로 칼레이도스코프
경력태도가 타당하게 구분될 수 있음을 밝혔
으며, Cabrera(2007)는 497명의 전문직 여성을
대상으로 한 연구에서 62%의 응답자가 자신
의 경력에 대한 초점이 시간에 따라 변화되었
다고 기술하였음을 설명하였다. 또한 Baruch와
그의 동료들(2014)은 칼레이도스코프 경력모형
과 계획행동이론에 따라 교원의 연구년에 대
한 의사결정을 설명하는 모델을 제안하였다. - 55 - 한국심리학회지: 산업및조직 칼레이도스코프 경력모형을 적용한 세대차
연구도 수행되었다. Sullivan과 그의 동료들
(2009)은 X세대의 경우 베이비부머 세대에 비
해 진정성과 균형에 대한 높은 요구를 갖고
있는 것으로 나타났으나 도전에 대한 세대
차는 발견되지 않았다고 설명하였다. August
(2011)는 경력후기 여성을 대상으로 질적 연구
를 수행한 결과 여성은 상대적으로 젊을 때
도전을 더 강조하는 것으로 나타났으며, 경력
후기로 갈수록 진정성과 균형을 강조하고 있
다고 설명하였다. 칼레이도스코프 경력태도와 관련된 선행연
구를 종합해보면, 칼레이도스코프 경력모형이
제안된 이후 새로운 기업 환경에서의 경력을
설명하기에 칼레이도스코프 경력 모형이 적절
한 지에 대한 연구와 경력과 관련된 태도에서
의 성차, 세대차를 칼레이도스코프 경력모형
으로 설명할 수 있는지에 초점을 맞추고 연구
들이 수행되어 온 것을 알 수 있다. 특히 여
성의 생애사건과 경력단계에 따른 경력 파라
미터의 변화를 분석한 질적 연구들이 많이 수
행되었다. 그러나 다음과 같은 한계점이 관찰
된다. 첫째, 칼레이도스코프 경력태도의 측정
도구가 여러 문화에 걸쳐 타당한지 측정도구
를 타당화 한 연구는 찾아보기 어렵다. 둘째,
칼레이도스코프 경력모형이 현대인의 경력 태
도를 설명하는데 타당한 지 확인하기 위해 특
정 집단을 대상으로 한 질적 연구 방법에 초
점을 두고 있어 다양한 대상을 포괄한 양적
연구의 수행이 요구된다. 셋째, 칼레이도스코
프 경력태도의 성차와 관련하여 여성이 알파
경력패턴을 갖는다는 결과와 베타 경력패턴을
갖는다는 상반된 연구결과가 존재하므로 보다
다양한 맥락적 변인을 종합적으로 고려한 연
구설계가 필요하다. 마지막으로 칼레이도스코
프 경력태도의 성별, 세대별 집단 간 차
다룬 연구에서도 칼레이도스코프 경력태
각 경력 파라미터의 평균 차이를 분석하
다. 이는 각 집단에서의 동일한 측정을
한다. 그러나 실제로 이러한 집단 간에(
성 vs. 여성) 칼레이도스코프 경력에 대
정동일성이 검증되지 않았다는 점에서 한
있다. 칼레이도스코프 경력태도와 성차
칼레이도스코프 경력모형은 남성과 다
성의 경력태도를 제안하기 위한 목적으
음 발표되었으므로 칼레이도스코프 경력
의 성차는 선행연구에서 중요하게 다루
왔다. 그러나 칼레이도스코프 경력태도
이를 실증적으로 검증한 연구는 제한적
수행되었다. 예를 들어 Mainiero와 Gibson
은 미국의 남녀 경력 중기 실업자(N=
대상으로 칼레이도스코프 경력태도의 성
검증하였는데, 경력중기 여성은 남성보
형을 더 강조하는 것을 밝혔다. 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 에 비해 경제적 안정, 일과 삶의 균형, 평생
직장, 행복/즐거움 등을 더 중요하게 인식하
는 것으로 나타났다. 일반적으로 여성의 경
우 출산과 육아 뿐 아니라 생애단계에 따라
주어지는 돌봄 노동에 대한 기대로 균형에
대한 가치를 중요하게 생각할 것으로 가정할
수 있다. 에 비해 경제적 안정, 일과 삶의 균형, 평생
직장, 행복/즐거움 등을 더 중요하게 인식하
는 것으로 나타났다. 일반적으로 여성의 경
우 출산과 육아 뿐 아니라 생애단계에 따라
주어지는 돌봄 노동에 대한 기대로 균형에
대한 가치를 중요하게 생각할 것으로 가정할
수 있다. (true to oneself)’으로 일터에서의 삶, 또는 자기
자신을 진정한 나로 받아들일 수 있는지와 관
련된다(Mainiero & Sullivan, 2005). 이는 개인의
내적 가치와 연관되어 있을 것이라 가정할 수
있으며, 실제로 여성을 대상으로 한 질적 연
구에서 진정성을 유지하기 위해서는 개인의
가치와 조직의 가치를 정렬하는 것이 중요하
다는 것을 발견하였다(Shaw & Leberman, 2015). 따라서 칼레이도스코프 경력태도의 진정성 파
라미터는 프로티언 경력태도와 유의한 상관관
계를 가질 것이라 가정하였다. 가설 1. 여성은 남성에 비해 칼레이도스코
프 경력태도 중 균형에 대한 인식이 유의미하
게 높을 것이다. 가설 2. 칼레이도스코프 경력태도는 프로티
언 경력태도와 유의한 정적 상관관계를 가질
것이다. 칼레이도스코프 경력태도와 프로티언 경력
태도, 일-삶의 균형, 도전적 직무경험
프로티언 경력태도는 경력에 대한 새로운
사고방식으로 ‘개인적인 가치를 토대로 경력
을 선택하는 자기주도적이고 자유의지를 반영
한 경력 관련 태도’를 말한다(Briscoe & Hall,
2006, p.6). 즉 프로티언 경력태도는 환경의 변
화 뿐 아니라 개인이 갖고 있는 가치에 따라
자신의 경력을 주도적으로 개발하고 관리하는
것이다(Hall, 2004). 프로티언 경력태도는 크게
두 가지 하위요인으로 구성되는데, 자기주도
적 경력관리 태도(self-directed career management
attitude)와 가치기반 경력태도(value-driven career
attitude)로 나눌 수 있다(Briscoe, Hall, &
DeMuth, 2006). 자기주도적 경력관리 태도는
스스로 경력관리의 주체가 되어 주도적인 역
할을 수행하는 것을 의미하며, 가치기반 경력
태도는 외적 보상보다는 개인의 고유한 가치
에 기반하여 심리적 성공을 추구하는 정도를
의미한다(Briscoe & Hall, 2006). 프로티언 경력
태도의 두 가지 하위요인은 칼레이도스코프
경력태도의 하위요인 중 진정성 파라미터와
유사하다. 진정성은 ‘나 자신에게 진실한 것 일-삶의 균형(work-life balance: WLB)은 근로
자가 일 영역과 삶의 영역 간 균형을 이루기
위해 개인적 자원을 필요한 역할 영역에 적
절하게 배분하는 것을 의미한다(Lee & Choi,
2019). Kim과 Park(2008)은 개인적 자원에는 시
간, 심리적 에너지, 신체적 에너지 등이 있을
수 있으며, 적절한 분배를 통해 삶에 대한 통
제감을 갖고 삶에 만족한 상태를 의미한다고
설명하였다. 칼레이도스코프 경력태도의 하위
요인 중 균형 파라미터는 개인의 일과 삶에
있어서 균형을 유지하는 것이 얼마나 중요한
지에 대한 개인의 인식을 반영한다(Mainiero &
Sullivan, 2005). 칼레이도스코프경력태도관련선행연구 예를 들어, Parker와 Chusmir(1992)는 경
력성공의 준거를 성별에 따라 비교하였는데,
여성은 남성에 비해 가족관계, 개인적 성취,
직업안정성 등을 중요하게 생각하고 지위
에 대한 중요성은 낮게 평가하였다. Dyke와
Murphy(2006)의 연구에서는 여성은 남성에 비
해 균형과 관계를 중요한 직업적 성공으로
여기고 있었다. 국내 연구에서는 Kim과 Cha
(2014)가 성별에 따라 경력성공 준거에 대한
인식에 차이가 있음을 밝혔는데, 여성은 남성 - 56 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 실제로 선행연구에서 혼인여부
와 자녀의 수는 칼레이도스코프 경력태도 중
균형과 유의한 상관관계를 갖는 것으로 나타
났으며, 이러한 경력 파라미터를 중요하게 생
각하는 근로자는 실제 일-삶의 균형을 높게
유지할 것이라 가정할 수 있다. - 57 - 한국심리학회지: 산업및조직 한국심리학회지: 산업및조직 가설 3. 칼레이도스코프 경력태도는 일-삶의
균형과 유의한 정적 상관관계를 가질 것이다. 심리적 과정 또는 결과로 설명되기도 한다
(Aryee & Tan, 1992). 경력몰입은 몰입 대상에
대한 심리적 애착이라는 관점에서 직무몰입
및 조직몰입과 유사하지만, 몰입의 대상이 조
직이 아니라 개인의 내재적인 경력 목표라는
관점에서, 그리고 특정 시점이 아닌 장기적인
속성을 갖는다는 점에서 차이가 있다(Cho &
Lee, 2017). 개인의 경력몰입은 현재의 경력을
지속적으로 유지하고자 하는 동기를 나타내므
로 많은 연구에서 경력 적응의 지표로 활용되
어 왔다. 도전적 직무경험은 현재 일상적으로 사용
하고 있는 업무수행 전략이나 방식이 불충분
하여 과업을 수행하는 새로운 방법이 요구되
는 업무활동을 의미한다(Lee & Cho, 2019). 도
전적 직무경험은 개인으로 하여금 새로운 학
습의 기회나 동기를 제공해주는 직무 특성을
포함하고 있고(McCauley, Ruderman, Ohlott, &
Morrow, 1994), 이를 통해 개인은 지식, 기술,
능력, 통찰력, 가치 등을 개발하여 성장할 수
있게 된다. 칼레이도스코프의 도전 파라미터
는 개인이 지속적으로 학습할 수 있는 긍정적
이고 도전적인 환경을 구축하는 것과 관련된
다(Shaw & Leberman, 2015). 도전은 개인이 업
무로부터 오는 책임을 두려워하지 않고, 새로
운 일에서 성장의 기회를 발견하는 것을 의미
한다(Sullivan et al., 2009). 따라서 칼레이도스코
프 경력태도 중 도전 파라미터는 도전적 직무
경험과 높은 상관관계를 가질 것이라 가정할
수 있다. 주관적 경력성공은 개인의 경력과 관련된
성취나 만족감이라고 정의되며, 현재 직무를
포함한 개인의 모든 경력에서의 성공에 대
한 주관적 평가로 정의된다(Seibert, Crant, &
Kraimer, 1999). 주관적 경력성공을 연구한 많
은 선행연구들은 경력만족을 주관적 경력성공
의 지표로 측정해 왔으며(Gattiker & Larwood,
1986; Judge, Cable, Boudreau, & Bretz, 1995), 최
근에는 경력에 대한 주관적 만족감 외에 개인
의 지속적인 고용가능성에 대한 인식을 주관
적 경력성공의 하위요인으로 포함해 왔다(Lee
& Lee, 2016). 개인이 자신의 경력에 대해 긍정
적으로 평가하고 인식하는 것과 관련된 주관
적 경력성공은 개인의 경력 결과에 대한 주관
적인 판단으로 많은 선행연구에서 경력개발
노력에 대한 결과변인으로 다루어져 왔다(Kim
& Moon, 2018). 가설 4. 칼레이도스코프 경력태도는 도전적
직무경험과 유의한 정적 상관관계를 가질 것
이다. 칼레이도스코프 경력태도와 경력몰입, 주관
적 경력성공
경력몰입은 자신의 직업에 대한 일반적 태
도 중 하나로 개인이 선택한 직업에서 지속적
으로 일하고자 하는 동기이다(Blau, 1985). 연구방법 중 즐거움과 균형을 중요하게 생각하는 유형
과 도전과 기여를 중요하게 생각하는 유형이
있었는데, 두 가지 유형은 다른 유형(지위와
도전 중심 유형, 보상과 안정 중심 유형)에 비
해 경력만족이 더 높은 것으로 나타났다. 즐
거움, 균형, 도전과 같은 경력성공의 준거가
칼레이도스코프 경력태도의 진정성, 균형, 도
전과 유사한 개념이라고 가정하면, 칼레이도
스코프 경력태도의 각 하위요인의 값이 높은
것이 보다 적응적인 것을 의미하는 것으로 해
석할 수 있을 것이다. 따라서 이 연구에서는
칼레이도스코프 경력태도의 하위요인의 수준
이 높은 것은 주관적 경력성공 그리고 경력몰
입과 유의미한 정적 상관관계를 가질 것이라
가정하였다. 첫째, 번역된 도구의 심리측정적 등가성
(psychometric equivalence)을 확보하기 위해 번역
-역번역법(back-translation method)을 실시하였다
(Brislin, 1973). 번역-역번역법은 번역문의 정확
도를 교차검증하기 위해 목표어로 번역된 도
구를 다른 사람에 의해 원어(source language)로
다시 번역하여 원래의 도구와 역번역된 도구
를 비교하는 것이다. 이에 연구자는 측정도구
원문을 개발한 주저자 Sullivan의 동의를 얻어,
Sullivan과 그의 동료들(2009)이 제시한 칼레이
도스코프 경력태도 세 개 요인 15문항을 번역
-역번역 하였다. 최초 번역은 연구자 2인이
합의를 통해 실시하였다. 역번역 과정에서 미
국에서 10년 이상 거주하고 HRD 석사 학위
를 취득하여 한국어와 영어 모두 능숙한 이중
언어 구사자(bilingual) 1명에게 연구 도구에 대
한 배경설명 없이 도구를 역번역 하도록 하였
다. 대표적으로 진정성 파라미터의 5문항 중
하나로 ‘지금 당장 내 꿈을 따라갈 수 있다면,
나는 그렇게 할 것이다.’는 ‘If I can follow my
dream right now, I will’로 번역되었고, 균형 파라
미터 5문항 중 하나인 ‘일과 가정의 균형을 이루
는 것은 내 인생의 신성한 목표다.’는 ‘Achieving
work-life balance is the sacred goal of my life.’
로, 도전 파라미터 5문항 중 하나인 ‘나는
좌절을 극복해야 할 문제가 아니라 해결책
이 필요한 도전으로 본다.’는 ‘I see setbacks
as “challenges” that require solutions rather than
“problems” to be overcome.’으로 번역되었다. 둘째구인화및예비문항에대해내용타당 가설 5. 칼레이도스코프 경력태도는 경력몰
입과 유의한 정적 상관관계를 가질 것이다. 가설 6. 칼레이도스코프 경력태도는 주관적
경력성공과 유의한 정적 상관관계를 가질 것
이다. 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 경
력몰입은 자신의 직무에 대한 개인의 동일시
나 현재의 경력목표를 추구해 나가는 과정에
서 어려움을 감수할 수 있는 의지로써 정서적, 칼레이도스코프 경력태도와 경력몰입, 주관
적 경력성공 한편, Lee(2018)는 X세대와 Y세대의 경력성
공 준거에 대한 인식을 비교하면서 즐거움,
균형, 도전과 같은 내부 지향의 주관적인 준
거들을 포함시켰으며 Y세대의 경력성공 인식
프로파일을 분석하고 개인의 경력성공 중요도
및 성취도 인식 수준을 비교한 바 있다. 이
연구에서 Y세대의 경력성공 인식의 잠재유형 - 58 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 중 즐거움과 균형을 중요하게 생각하는 유형
과 도전과 기여를 중요하게 생각하는 유형이
있었는데, 두 가지 유형은 다른 유형(지위와
도전 중심 유형, 보상과 안정 중심 유형)에 비
해 경력만족이 더 높은 것으로 나타났다. 즐
거움, 균형, 도전과 같은 경력성공의 준거가
칼레이도스코프 경력태도의 진정성, 균형, 도
전과 유사한 개념이라고 가정하면, 칼레이도
스코프 경력태도의 각 하위요인의 값이 높은
것이 보다 적응적인 것을 의미하는 것으로 해
석할 수 있을 것이다. 따라서 이 연구에서는
칼레이도스코프 경력태도의 하위요인의 수준
이 높은 것은 주관적 경력성공 그리고 경력몰
입과 유의미한 정적 상관관계를 가질 것이라
가정하였다. 연구1. 내용타당도분석 선행연구 검토를 통해 한국형 칼레이도스코
프 경력태도를 측정하는 데 있어 Sullivan과 그
의 동료들(2009)의 도구가 가장 근접한 조사도
구라는 연구자 간 합의를 도출하고, 해당 도
구를 번역-역번역법을 통해 국문화한 뒤, 수
정델파이법을 활용하여 전문가들의 의견을 취
합함으로써 측정도구 초안을 개발하였다. 둘째, 구인화 및 예비문항에 대해 내용타당
도를 확인하기 위해 수정델파이법을 활용하였
다. 일반적으로 개발 완료된 측정도구는 하위
요인별 3~15 문항 정도로 구성할 것을 권고 - 59 - 한국심리학회지: 산업및조직 문으로 각 문항에 대한 수정 의견을 기재하
도록 하였다. 2라운드는 1라운드에서 수정된
문항과 타 전문가의 코멘트, 본인의 이전 응
답 등을 종합적으로 고려하여 타당성 정도를
Likert 5점 척도에 응하게 하고, 개방형 질문으
로 각 문항에 대한 최종 의견을 기재하도록
하였다. 문으로 각 문항에 대한 수정 의견을 기재하
도록 하였다. 2라운드는 1라운드에서 수정된
문항과 타 전문가의 코멘트, 본인의 이전 응
답 등을 종합적으로 고려하여 타당성 정도를
Likert 5점 척도에 응하게 하고, 개방형 질문으
로 각 문항에 대한 최종 의견을 기재하도록
하였다. 하고 있으므로(Walsh & Betz, 1995), 델파이 연
구 결과에 따른 문항 삭제 등 도구 수정을 고
려하여 파라미터별로 7개씩 총 21개로 구성된
칼레이도스코프 경력태도 측정도구 예비문항
을 선정하였다. 앞서 실시한 역번역 결과로
얻은 3개 요인 15문항 외에, 선행연구 분석을
토대로 연구자 2인이 상호 합의하여 요인별 2
개 문항을 추가하였다. 진정성(Authenticity) 파
라미터에 포함한 문항은 ‘나는 내 삶과 내 일
을 통해 진정한 나를 찾을 것이다.’와 ‘내가
나아갈 길에 대한 가치와 신념은 외부의 압박
에도 흔들리지 않는다.’ 등 2문항 이고, 균형
(Balance) 파라미터에는 ‘나에게는 가족이 가장
큰 우선순위이며, 그 다음이 일이다.’, ‘나는
가족으로부터 얻는 행복과 일에서 얻는 성취
를 동시에 추구한다.’ 등 2문항을 포함하였으
며, 도전(Challenge) 파라미터에는 ‘나에게 있어
새로운 변화란 나를 성장시킬 수 있는 기회이
다.’, ‘나는 도전적인 과제를 수행할 때 살아있
음을 느낀다.’ 등 2문항을 새로 개발하여 포함
하였다. * CVR: Content Validity Ratio 결 과 수정델파이 1라운드 결과, 진정성 1번 문항
은 번역이 어색하다는 의견 및 문장 수정 의
견에 따라 ‘나는 나에게 적합한 더 큰 삶의
의미를 찾길 원한다’로 수정하였다. 진정성 6
번 문항 ‘나는 내 삶과 내 일을 통해 진정한
나를 찾을 것이다’는 삶과 일이 개념적으로
‘과’로 묶이는 동일한 자격이 아니라고 사료된
다는 전문가 의견과 강한 어조를 사용한 경우
점수분포가 상대적으로 낮아질 수 있어 주의
를 요한다는 전문가 의견을 반영하여 ‘나는
내 일을 통해 내 삶 속에서 진정한 나를 찾길
바란다’로 수정하였다. 진정성 7번 문항 ‘내가
나아갈 길에 대한 가치와 신념은 외부의 압박
에도 흔들리지 않는다’는 ‘나아갈 길’ 및 ‘외
부 압박’이라는 개념이 모호하고 범위를 특정
할 수 없다는 지적에 근거하여 ‘내 진로에 대
한 가치와 신념은 어떠한 환경이나 조건에도
흔들리지 않는다’로 수정하였다. 한편, 전문가
가 제안한 ‘나에게는 마음, 정신, 신체 건강이
통합된 만족감이 중요하다’ 는 문항을 새롭게
추가하였다. 수정델파이 전문가 패널의 선정기준은 다음
과 같다. 먼저, 박사학위를 소지하고 있고 칼
레이도스코프 경력태도와 유관한 경력개발 또
는 진로발달 관련 변인 도구개발 경력이 있거
나 HRD 현업에 10년 이상 종사한 전문가 5인
을 선정하였으며, 측정 대상이 되는 근로자
중 박사학위를 소지하고 진정성, 균형, 도전을
경력발달 과정에서 경험했다고 응답한 경력
10년 이상의 여성과학기술인 5인을 포함하여
총 10인을 전문가 패널로 선정하였다. 수정델
파이는 총 2라운드로 구성하였다. 1라운드에
서는 전문가 패널로 하여금 파라미터에 각 문
항이 위치해 있는 것이 적절한지 타당성 정도
를 Likert 5점 척도에 응하게 하고, 개방형 질 균형 2번 문항 ‘나는 항상 가족의 요구에
우선하여 나의 일을 배치한다’는 가족의 요구
에 우선하기보다는 가족 요구와 나의 일을 지 - 60 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 * CVR: Content Validity Ratio 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 포함하였으나, 2라운드 결과 매우 낮은 내용
타당도비율(CVR)값을 보여 최종 삭제하였다. 이에 따라, 2라운드에 걸친 델파이 분석 결과
최종 진정성 파라미터 6문항, 균형 파라미터 6
문항, 도전 파라미터 6문항이 개발되었다. 도전 2번 문항에서 ‘좌절’은 전문가 의견을
바탕으로 ‘어려움’ 으로 순화하였다. 도전 3번
문항 ‘추가된 업무 책임은 나에게 걱정거리가
못 된다’ 는 번역투가 어색하다는 다수 전문
가의 의견에 따라 ‘나는 기존보다 업무가 추
가되어도 크게 걱정하지 않는다’로 수정하였
다. 도전 4번 문항 ‘내 주위 사람들은 나를 매
우 목표 지향적인 사람이라고 설명한다.’는 경
력태도 상의 도전이 지나치게 목표 지향성과
유사한 현상으로 이해될 수 있어 주의가 요구
된다는 전문가 의견에 따라 ‘내 주위 사람들
은 나를 목표 지향적인 사람이라고 얘기한다’
로 수정하였다, 도전 7번 문항 ‘나는 도전적인
과제를 수행할 때 살아있음을 느낀다’는 도전
적 직무 경험과 같은 외부 변수와 중복이 우
려된다는 전문가 의견에 따라 삭제하였다. 연구방법 조사대상 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 균형을 이루는 것은 내 인생의 신성한 목표
다’ 는 ‘신성한’이라는 단어의 국내 맥락에서
의 쓰임 및 정서를 감안하여야 한다는 의견을
반영하여 ‘일과 가족의 균형을 이루는 것은 속적으로 균형 있게 조정한다는 뜻이라고 판
단한 전문가 2인의 의견을 반영하여 ‘나는 가
족의 요구를 고려하여 나의 커리어를 계획한
다’ 로 수정하였다. 균형 4번 문항 ‘일과 삶의 items
First Round
Second Round
Mean
S.D. Median
CVR
Decision
Mean
S.D. Median
CVR
Decision
Authenticity
1
4.10
0.88
4
0.40
Modify
4.40
0.70
5
0.80
Accept
2
4.80
0.42
5
1.00
Maintain
4.70
0.48
5
1.00
Accept
3
4.60
0.70
5
0.80
Maintain
4.80
0.42
5
1.00
Accept
4
4.20
0.79
4
0.60
Modify
4.50
0.71
5
0.80
Accept
5
4.70
0.48
5
1.00
Maintain
4.70
0.48
5
1.00
Accept
6
3.90
0.99
4
0.40
Modify
4.40
0.84
5
0.60
Modify/
Accept
7
3.70
1.16
4
0.20
Modify
4.00
0.94
4
0.60
Delete
+
-
-
-
-
Generate
3.10
1.10
3
-0.40
Delete
Balance
1
4.20
0.63
4
0.80
Maintain
4.50
0.53
5
1.00
Accept
2
3.60
1.43
4
-0.20
Modify
4.40
0.70
5
0.80
Accept
3
4.10
1.20
4
0.80
Maintain
4.30
0.67
4
0.80
Accept
4
4.50
1.08
5
0.60
Modify
4.60
0.70
5
0.80
Accept
5
3.50
1.45
4
0.40
Modify
4.30
0.82
5
0.60
Accept
6
3.10
1.20
3
-0.20
Delete
-
-
-
-
-
7
4.00
0.82
4
0.40
Modify
4.60
0.52
5
1.00
Accept
Challenge
1
4.60
0.97
5
0.80
Maintain
4.90
0.32
5
1.00
Accept
2
3.90
1.12
5
0.40
Modify
4.70
0.48
5
1.00
Accept
3
4.10
0.99
4
0.60
Modify
4.20
0.63
4
0.80
Accept
4
3.90
1.10
4
0.20
Modify
4.60
0.70
5
0.80
Accept
5
4.50
0.71
5
0.80
Maintain
4.40
0.70
5
0.80
Accept
6
4.40
0.84
5
0.60
Modify
4.90
0.32
5
1.00
Accept
7
3.80
1.03
4
0.20
Delete
-
-
-
-
-
*
l d
Table 1
Summary of the result from modified delphi analysis Summary of the result from modified delphi analysis - 61 - 한국심리학회지: 산업및조직 포함하였으나, 2라운드 결과 매우 낮은 내용
타당도비율(CVR)값을 보여 최종 삭제하였다. 이에 따라, 2라운드에 걸친 델파이 분석 결과
최종 진정성 파라미터 6문항, 균형 파라미터 6
문항, 도전 파라미터 6문항이 개발되었다. 내 인생의 중요한 목표다’로 수정하였다. 균형
6번 문항은 CVR값이 낮고 균형 1번 문항과
사실상 같은 의미를 내포한다는 전문가 의견
에 따라 삭제하였다. 결 과 칼레이도스코프 경력태도
칼레이도스코프 경력태도는 연구1을 통해
내용타당도를 확보한 18개 문항(진정성, 균형,
도전 각 6개 문항)으로 구성되었다. Sullivan과
그의 동료들(2009)이 개발한 원척도는 5점
Likert로 응답하도록 하였으나 5점 Likert에 비
해 6점 Likert 응답을 활용하는 것이 신뢰도와
변별도를 유의미하게 향상시킨다는 Chomeya
(2010)의 연구결과에 따라 6점 Likert 척도(1=
전혀 동의하지 않는다. 2=별로 동의하지 않는
다. 3=동의하지 않는 편이다. 4=동의하는 편
이다. 5=동의한다. 6=매우 동의한다)를 활용
하여 측정하였다. 탐색적요인분석 요인분석에 필요한 기본가정을 표본이 충족
시키는 지 확인하기 위해 KMO(Kaiser-Meyer-
Olkin) 검정과 Bartlett이 제안한 구형성 검정을
실시하였다. KMO는 .921로 1에 근접하므로
적합하다고 보았고, Bartlett 검정값은 근사 카
이제곱 3209.433, p<.001 로 나타나 적절하므
로 요인분석의 선행조건을 만족한다고 판단하
였다. 탐색적 요인분석을 위해 고유값은 1보
다 큰 조건으로, 베리맥스 회전을 통해 최대
우도법으로 요인을 추출한 결과 3개의 요인
이 도출되었다(표 2 참조). 스크리도표를 함께
참고한 결과 3요인 모형이 가장 적합한 것으
로 나타났다. 도전 파라미터의 내적 일관성
(cronbach α)은 .882, 균형은 .869, 진정성은 .832
였으며, 전체적으로는 .911 로 나타났다. 탐색
적 요인분석 결과 원 도구와 마찬가지로 3개
의 요인 구조가 도출되었으나, 균형 1번과 진
정성 4번 등 일부 문항의 공통성이 .4 미만으
로 나타났으며, 진정성 3번, 5번, 6번은 도전
요인에도 요인부하량이 .4 이상으로 나타나
확인적 요인분석을 통해 요인구조를 재확인하
고자 하였다. 조사대상 연구 자료 수집을 위해 온라인 조사업체에
의뢰하여 국내의 다양한 기업에 근무하고 있
는 근로자를 대상으로 온라인 설문조사를 실
시하였다. 조사 대상을 선정함에 있어 성별,
연령을 고려하여 비율표집 하였고, 기타 고
려할 수 있는 인구통계학적 요인은 무선화
(randomize)함을 원칙으로 하였다. 전체 2,857명
을 대상으로 설문지가 발송되어 476명이 응답
을 시도하였으며(응답률 16.7%), 재직 근로자
가 아니거나 중간에 설문응답을 포기한 89명
을 제외한 396명의 응답자가 설문응답을 완료
하였다. 396명 중 응답이 불성실한 43명의 데
이터를 제외하고 최종적으로 347명의 응답자
료를 분석하였다. 347명의 인구통계학적 특성
은 다음과 같다. 성별은 남성이 193명(55.6%),
여성이 154명(44.4%)이고, 연령은 20~24세 24
명(6.9%), 25~29세 44명(12.7%), 30~34세 45명
(13.0%), 35~39세 48명(13.8%), 40~44세 43명
(12.4%), 45~49세 47명(13.5%), 50~54세 37명
(10.7%), 55~59세 32명(9.2%), 60세 이상 27명
(7.8%)이었다. 직급은 사원급 103명(29.7%), 대 수정델파이 2라운드 결과, 진정성 6번 문항
‘나는 내 일을 통해 내 삶 속에서 진정한 나
를 찾길 바란다’는 ‘찾는다’는 동사를 ‘발견한
다’ 는 행위로 변경하면 내적인 자각, 성찰을
의미할 것이라는 의견을 반영하여 최종 ‘나는
일을 통해 내 삶 속에서 진정한 나를 발견하
기를 원한다’로 수정 선택하였다. 진정성 7번
문항 ‘내 진로에 대한 가치와 신념은 어떠한
환경이나 조건에도 흔들리지 않는다’는 칼레
이도스코프 경력태도의 경우 진로에 대한 가
치와 신념이 변한다는 전제에 기반한 개념이
므로, 진정성 문항이라 하더라도 ‘불변’의 논
리를 사용하는 것은 무리라는 전문가 의견을
반영하여 최종 삭제하였다. 진정성 8번 문항
은 1라운드에서 전문가 의견에 따라 문항으로 - 62 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 인하였으며, 탐색적 요인분석을 통해 확인된
요인구조를 바탕으로 요인 및 문항의 구조적
관계를 검증하기 위해 확인적 요인분석을 실
시하였다. 적합도 지수는 TLI, CFI, RMSEA,
SRMR을 활용하였다. 리급 74명(21.3%), 과장급 62명(17.9%), 차장급
32명(9.2%), 부장급 43명(12.4%), 임원 이상 33
명(9.5%) 이었다. 학력은 박사 이상 4명(1.2%),
석사 24명(6.9%), 대졸 231명(66.6%), 전문대졸
46명(13.3%), 고졸이하 42명(12.1%) 이었다. 조사도구 분석방법 칼레이도스코프 경력태도의 이론적 구성개
념이 심리측정적으로 일치하는지 확인하기 위
해 통계 프로그램 IBM SPSS AMOS 25.0을 이
용하여 기술통계, 신뢰도 분석, 요인분석을 실
시하였다. 먼저 연구 1을 통해 확인한 18개
문항의 칼레이도스코프 경력태도 측정도구의
요인구조를 확인하기 위해 탐색적 요인분석을
실시하였다. 탐색적 요인분석은 최대우도법,
직교회전으로 추출하였으며 아이겐값을 기준
으로 요인의 수를 검토하였다. 요인부하량과
공통성을 검토하여 삭제할 문항이 있는지 확 - 63 - 한국심리학회지: 산업및조직 한국심리학회지: 산업및조직 Table 2
The result of exploratary factor analysis (N=347) Items
Factors
communality
1
2
3
Challenge 4
Most people would describe me as being very goal-directed. .763
.157
.167
.635
Challenge 5
I thrive on work challenges and turn work problems into
opportunities for change. .714
.266
.246
.642
Challenge 2
I view setbacks not as “problems” to be overcome but as
“challenges” that require solutions. .687
.201
.314
.611
Challenge 6
For me, a new change is an opportunity to grow. .661
.227
.389
.640
Challenge 3
Added work responsibilities don’t worry me. .644
.074
.097
.429
Challenge 1
I continually look for new challenges in everything I do. .610
.192
.342
.525
Balance 5
Nothing matters more to me right now than balancing
work with my family responsibilities. .096
.796
.176
.674
Balance 3
My work is meaningless if I cannot take the time to be
with my family. .102
.776
.111
.625
Balance 4
Achieving balance between work and family is life’s holy
grail. .067
.742
.322
.659
Balance 6
I pursue the happiness of my family and the achievement
of my work at the same time. .213
.682
.308
.605
Balance 1
If necessary, I would give up my work to settle
problematic family issues or concerns. .191
.589
.030
.384
Balance 2
I constantly arrange my work around my family needs. .352
.571
-.011
.450
Authenticity 2
I hunger for greater spiritual growth in my life. .205
.165
.750
.632
Authenticity 1
I hope to find a greater purpose to my life that suits who
I am. .115
.148
.709
.538
Authenticity 4
If I could follow my dream right now, I would. .280
.118
.531
.374
Authenticity 5
I want to have an impact and leave my signature on what
I accomplish in life. .421
.186
.514
.476
Authenticity 6
I want to find genuine me in my life through work. 분석방법 .458
.086
.513
.480
Authenticity 3
I have discovered that crises in life offer perspectives in
ways that daily living does not. .436
.185
.476
.451
Eigen Value
3.728
3.293
2.808
% of Variance Explained: 54.607
20.711
18.296
15.600
Cronbach α: .911
.882
.869
.832
Table 2
The result of exploratary factor analysis (N=347) - 64 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 연구방법 조사대상 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 삭제하였다. 문항 삭제 후 확인적 요인분석을
다시 실시한 결과 χ2 =161.788(df=62, p<.001),
CFI=.957,
TLI=.945,
RMSEA=.068
[하한값:
.055, 상한값: .081], SRMR=.049로 모형의 적합
도가 전반적으로 개선된 것으로 나타났다(표
3 참조). 삭제하였다. 문항 삭제 후 확인적 요인분석을
다시 실시한 결과 χ2 =161.788(df=62, p<.001),
CFI=.957,
TLI=.945,
RMSEA=.068
[하한값:
.055, 상한값: .081], SRMR=.049로 모형의 적합
도가 전반적으로 개선된 것으로 나타났다(표
3 참조). 확인적요인분석 확인적요인분석 탐색적 요인분석 결과를 통해 확인된 진정
성, 균형, 도전 파라미터를 잠재변인으로 설
정하고, 측정 문항을 측정변인으로 하여 잠
재변인과 측정변인 간 경로를 연결하고 확인
적 요인분석을 실시하였다. 그 결과, 모형적
합도 측면에서 χ2=415.826(p<.001), CFI=.880,
TLI=.911, RMSEA=.072[하한값: .064, 상한값:
.080], SRMR=.056로 나타나 χ2값과 CFI가 적
합도 기준에 부합하지 않는 것으로 나타났
다. 이에 문항의 요인부하량과 다중상관자승
(Squared Mutilple Correlations: SMC) 값을 검토하
였다. 모든 문항의 요인부하량은 .5 이상으로
적합하였으나, 진정성1(나는 나에게 적합한 더
큰 삶의 의미를 찾기를 원한다), 진정성4(지금
당장 내 꿈을 따라갈 수 있다면, 나는 그렇게
할 것이다), 균형1(필요하다면, 나는 가족과 관
련된 문제를 해결하기 위해 일을 포기할 것이
다), 균형2(나는 가족의 요구를 고려하여 나의
커리어를 계획한다), 도전3(나는 기존에 수행
하던 업무에 추가적인 과업이 주어져도 크게
걱정하지 않는다)의 SMC 값이 .4 미만으로 나
타났다. 이에 문항을 세부적으로 검토하여 연
구자 간 합의를 통해 5개 문항을 삭제하였다. 진정성1번 문항은 진정성 2번 문항과 오차 상
관이 높게 나타나 두 문항 간 측정내용이 유
사하여 이를 삭제하는 것이 적절하다고 판단
하였으며, 진정성 4번, 균형 1번, 균형 2번은
유연성이 낮은 우리나라 노동시장의 맥락적
특성을 고려할 때 국내 근로자의 경력 태도를
측정하기에 적합하지 않은 문항으로 판단하였
다. 마지막으로 도전 3번 문항은 추가적인 과
업에 대한 스트레스 감내성으로 도전에 대한
다른 측정문항들과 성격이 다르다고 판단하여 연구3: 교차타당도, 잠재평균차이,
준거관련타당도분석 조사대상 연구3을 위한 표본은 별도의 자료수집을 통
해 확보되었다. 연구2와 마찬가지로 2020년 7
월 기준 전체 근로자의 연령분포 비율을 확인
하여 연령별 비율 표집을 실시하였다. 온라인
전문조사업체에 의뢰하여 전체 4,456명을 대상
으로 설문지가 발송되었고 846명이 응답을 시
도하였으며(응답률 19.0%), 재직 근로자가 아
니거나 중간에 설문응답을 포기한 212명을 제
외한 634명의 응답자가 설문응답을 완료하였
다. 634명 중 응답이 불성실한 3명의 데이터
를 제외하고 최종적으로 631명의 응답자료를
분석하였다. 631명의 인구통계학적 특성은
다음과 같다. 성별은 남성이 347명(55.0%), 여
성이 284(45.0%)명이고, 연령은 20~24세 29명
(4.6%), 25~29세 75명(11.9%), 30~34세 82명
(13.0%), 35~39세 88명(13.9%), 40~44세 82명
(13.0%), 45~49세 83명(13.2%), 50~54세 73명
(11.6%), 55~59세 61명(9.7%), 60세 이상 58명
(9.2%) 이었다. 직급은 사원급 163명(25.8%), 대 - 65 - 한국심리학회지: 산업및조직 통해 번안된 바 있다. 예시문항은 ‘나는 일(직
장)과 삶 두 영역에 적절히 시간을 배분하고
사용하고 있다’이다. Valcour(2007)의 연구에서
신뢰도(cronbach’s α)는 .93으로 보고되었고,
Lee(2012)의 연구에서는 .836으로 나타났다. 이
연구에서 측정한 일-삶의 균형 5문항의 신뢰
도(cronbach’s α)는 .876이었다. 리급 135명(21.4%), 과장급 135명(21.4%), 차장
급 65명(10.3%), 부장급 84명(13.3%), 임원 이
상 49명(7.8%) 이었다. 학력은 박사 이상 14명
(2.2%), 석사 65명(10.3%), 대졸 401명(63.5%),
전문대졸 84명(13.3%), 고졸이하 67명(10.6%)
였다. 혼인 여부는 미혼이 234명(37.1%), 기혼
이 385명(61.0%) 이며, 12명(1.9%)은 기타(별거,
이혼, 사별)에 응답하였다. 리급 135명(21.4%), 과장급 135명(21.4%), 차장
급 65명(10.3%), 부장급 84명(13.3%), 임원 이
상 49명(7.8%) 이었다. 학력은 박사 이상 14명
(2.2%), 석사 65명(10.3%), 대졸 401명(63.5%),
전문대졸 84명(13.3%), 고졸이하 67명(10.6%)
였다. 혼인 여부는 미혼이 234명(37.1%), 기혼
이 385명(61.0%) 이며, 12명(1.9%)은 기타(별거,
이혼, 사별)에 응답하였다. 도전적 직무경험은 De Pater와 그의 동료들
(2009)이 개발한 7문항의 척도를 사용하였다. De Pater와 그의 동료들(2009)의 연구에서는 도
전적 직무경험이 하위요인이 없는 단일요인
구성개념으로 제시되었고 신뢰도는 .74로 양
호하였다. 이 연구에서는 De Pater와 그의 동
료들(2009)이 개발한 척도를 Lee와 Lee(2016)가
번역한 도전적 직무경험 7문항 척도를 활용하
였다. 예시문항은 ‘나는 우리 부서에서 전략적
변화를 시도하는 일을 담당한다’이며 신뢰도
(cronbach’s α)는 .912로 양호하였다. χ2=161.788(df=62, p<.001), CFI=.957, TLI=.945, RMSEA=.068 [하한값: .055, 상한값: .081], SRMR=.049 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 Path
B
β
S.E. t
AVE
CR
진정성 (Authenticity)
→
나는 내 인생에서 더 큰 정신적 성장을 갈망한다. (I hunger for greater spiritual growth in my life.)
1
.657
.515
.809
→
나는 인생의 위기를 통해 새로운 관점을 얻을 수 있다는 것을 알고 있다. (I have discovered that crises in life offer perspectives in ways that daily living
does not.)
1.190
.686
.112
10.664***
→
나는 스스로 성취한 것을 통해 영향력을 갖고 내 발자취를 남기고 싶다. (I want to have an impact and leave my signature on what I accomplish in
life.)
1.222
.712
.111
10.971***
→
나는 일을 통해 내 삶 속에서 진정한 나를 발견하기를 원한다. (I want to find genuine me in my life through work.)
1.213
.717
.110
11.032***
균형 (Balance)
→
가족과 함께 하는 시간을 낼 수 없다면, 내 일은 무의미하다. (My work is meaningless if I cannot take the time to be with my family.)
1
.745
.606
.860
→
일과 가족의 균형을 이루는 것은 내인생의 중요한 목표다. (Achieving balance between work and family is life’s holy grail.)
.959
.826
.065
14.783***
→
지금 당장 나의 일과 가족의 균형을 맞추는 것보다 더 중요한 것은 없다. (Nothing matters more to me right now than balancing work with my family
responsibilities.)
.951
.820
.065
14.682***
→
나는 가족으로부터 얻는 행복과 일에서 얻는 성취를 동시에 추구한다. (I pursue the happiness of my family and the achievement of my work at the
same time.)
.850
.771
.061
13.835***
도전 (Challenge)
→
나는 내가 하는 모든 일에서 끊임없이 새로운 도전을 추구한다. (I continually look for new challenges in everything I do.)
1
.720
.624
.892
→
나는 어려움을 극복해야 할 “문제”가 아니라 해결책이 필요한 “도전”으
로 본다. (I view setbacks not as “problems” to be overcome but as “challenges” that
require solutions.)
1.058
.790
.076
14.001***
→
내 주위 사람들은 나를 목표지향적인 사람이라고 얘기한다. (Most people would describe me as being very goal-directed.)
1.120
.744
.085
13.205***
→
나는 업무상 어려움을 잘 극복하고, 발생된 문제는 개선의 기회로 활용
한다. (I thrive on work challenges and turn work problems into opportunities for
change.)
1.002
.800
.071
14.179***
→
나에게 있어 새로운 변화란 나를 성장시킬 수 있는 기회이다. 조사도구 칼레이도스코프 경력태도는 연구 2를 통해
도출된 13문항을 활용하여 측정하였다. 진정
성 4문항, 균형 4문항, 도전 5문항으로 구성되
며, 문항은 표 3에 기술된 것과 같다. 프로티언 경력태도는 개인의 가치나 관심에
따라 자신의 경력을 자기주도적으로 개발하
고 관리하는 경력을 의미하며(Briscoe & Hall,
2006), 가치지향성과 자기주도성의 2가지 하위
요인으로 구성된다(Briscoe, Hall, & Demuth,
2006). 이 연구에서는 Briscoe, Hall과 DeMuth
(2006)의 측정도구를 번안한 Shin과 Jeong(2014)
의 도구를 활용하였다. 예시문항은 ‘내 경력의
성공과 실패에 대한 책임은 나에게 있다.’(자
기주도성), ‘나는 경력을 평가할 때, 임금이나
승진보다는 나의 고유한 가치를 더 추구한
다.’(가치지향성)이다. 이 연구에서 하위요인
의 신뢰도(cronbach’s α)는 자기주도성 .853,
가치지향성 .838였으며, 전체 문항의 신뢰도
(cronbach’s α)는 .890이었다. 주관적 경력성공은 경력만족과 고용가능성
을 각각 측정하여 합산한 값을 활용하였다. Greenhaus와 그의 동료들(1990)이 개발한 도구
를 바탕으로 Shim(2011)이 번안한 도구를 사용
하였다. 경력만족의 측정도구는 경력 전반에
대한 만족도와 목표달성, 기술과 능력, 보수,
직급 측면에서의 만족도를 묻는 5문항으로 구
성되었으며,
Shim(2011)의 연구에서 신뢰도
(cronbach’s
α)는 .88
이었다. 고용가능성은
Johnson(2001)이 개발한 도구를 바탕으로 Chang
과 Park(2004)이 번안한 도구를 사용하였다. 고
용가능성의 측정도구는 6문항의 5점 LIkert로
측정하도록 되어있으며 Chang과 Park(2004)의
연구에서 신뢰도는 .90였다. 이 연구에서는 경
력만족과 고용가능성의 신뢰도(cronbach’s α)는
각각 .919, .881이었으며, 전체 주관적 경력성
공의 신뢰도(cronbach’s α)는 .917였다. 일-삶의 균형을 측정하기 위해 Valcour(2007)
가 개발한 일-가정 균형에 대한 만족도를 측
정하는 5문항을 활용하였다. 이 측정도구는
일과 가정 균형에 대한 전반적 만족도를 측정
하고 있으며, 국내에서는 Lee(2012)의 연구를 - 66 - 교차타당도분석결과 교차타당도분석결과 교차타당도를 확인하기 위해 연구 2의 표본
1(N=347)을 추정표본으로 하고 연구 3의 표본
2(N=631)를 검증표본으로 하여 모형 적합도를
비교하였다. 교차타당화집단의 모형 적합도를
살펴보면 모수추정집단과 비교했을 때 상대적
으로 양호하지 않은 것을 알 수 있으나 CFI,
TLI가 .943, .928로 양호한 수준으로 나타났으
며, RMSEA도 .072로 양호한 범위로 판단할 수
있다. 또한 교차타당성 지수(ECVI: Expected
Cross-Validation Index)를 확인한 결과, 모수추정
집단은 .635, 교차타당화집단은 .514로 교차타
당화집단 모형이 더 안정적인 것으로 나타났
다. 즉, 표본이 달라졌음에도 불구하고 칼레이
도스코프의 3요인 측정모형은 타당한 것으로
확인되었다. 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 (For me, a new change is an opportunity to grow.)
1.012
.818
.070
14.488***
χ2=161.788(df=62, p<.001), CFI=.957, TLI=.945, RMSEA=.068 [하한값: .055, 상한값: .081], SRMR=.049
Table 3
The result of confirmative factor analysis (N=347) Table 3 The result of confirmative factor analysis (N=347) →
(For me, a new change is an opportunity to grow.) - 67 - 한국심리학회지: 산업및조직 경력몰입은 Blau(1985)가 개발하고 Chang
(1997)이 번안한 측정도구를 활용하였다. Chang
(1997)가 번안한 척도 중 ‘나는 내가 하고 있
는 직업분야에서 내 경력을 유지하기를 원한
다’ 등 4문항을 활용하였다. Chang(1997)의 연
구에서 보고한 신뢰도(cronbach’s α)는 .76이었
으며, 이 연구의 신뢰도(cronbach’s α)는 .824였
다. 모든 측정변인은 6점 Likert 척도(1=전혀
동의하지 않는다. 2=별로 동의하지 않는다. 3=동의하지 않는 편이다. 4=동의하는 편이다. 5=동의한다. 6=매우 동의한다)를 활용하여 측
정하였다. 분석을 실시하여 준거 변인들과의 관계를 확
인하였다. 연구결과 연구결과 교차타당도분석결과 분석방법 이 연구에서 설정한 구인 모형의 타당도를
다시 한 번 확인하기 위해 표본 2를 대상으로
확인적 요인분석을 실시하였다. 확인적 요인
분석 모형의 적합도와 더불어 기대교차타당성
지수(Expected Cross-Validation Index; ECVI)를 확
인하였다. 또한 다집단 분석을 실시하여 남녀
기업근로자의 측정동일성을 검증하였고, 측정
오차를 통제한 상태에서 칼레이도스코프 경력
태도 평균의 성별 차이를 확인하기 위해 잠재
평균분석을 실시하였다. 마지막으로 한국판
칼레이도스코프와 관련 변인들과의 상관관계 연구 2에서 사용하지 않았던 새로운 표본
2(N=631)을 추정표본으로 하여 확인적 요인
분석을 실시하였다. 연구 2의 결과로 제안된 Table 4 Model
χ2
df
χ2/df
CFI
TLI
RMSEA
ECVI
Parameter Estimation Group
(N=347)
161.788
62
2.609***
.957
.954
.068
(.055-.081)
.635
(.537-.755)
Cross-Validity Group
(N=631)
266.097
62
4.292***
.943
.928
.072
(.063-.081)
.514
(.440-.601)
***p <.001
Table 4
Model comparison for cross-validity analysis Model comparison for cross-validity analysis - 68 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 -
13문항 3요인 모형을 연구모형으로 설정하고
측정모형을 검증하였다. 확인적 요인분석의
적합도 지수는 χ2=266.097(df=62, p<.001),
CFI=.943, TLI=.928, RMSEA=.072 [하한값:
.063, 상한값: .081], SRMR=.045로 모형의 적
합도는 양호한 것으로 나타났다. 또한 각 요
인별 표준화계수가 모두 .6 이상으로 통계적
으로 유의한 적재량을 보이고 있었으며, 다
중상관자승(SMC) 값도 .4 이상으로 나타났다. 한편, 모든 잠재 변인의 개념 신뢰도(construct
reliability)가 .8 이상이었으며, 평균분산추출
(Average Variance Extracted)이 .5 이상으로 나타
나 내적 일관성 및 수렴 타당성을 확보하였다. 칼레이도스코프측정도구의성별에따른잠재
평균차이분석
칼레이도스코프 측정도구의 3요인 모형이
남녀 재직근로자 모두에게 적용할 수 있는 모
형인지 확인하기 위해 다집단분석을 실시하였
다. 먼저 칼레이도스코프 측정도구의 3요인
Model
χ2(df)
Δχ2(Δdf)
Model A
353.941(124)
-
Model B
372.422(134)
18.481* (10)
Model C
393.238(147)
20.816 (13)
Model D
414.703(153)
21.466** (6)
Note: Model A: unconstrained model; Model B:
measurement weights and intercepts; Model D: constrain
Table 5
The result of multi-group analysis between 13문항 3요인 모형을 연구모형으로 설정하고
측정모형을 검증하였다. 확인적 요인분석의
적합도 지수는 χ2=266.097(df=62, p<.001),
CFI=.943, TLI=.928, RMSEA=.072 [하한값:
.063, 상한값: .081], SRMR=.045로 모형의 적
합도는 양호한 것으로 나타났다. 또한 각 요
인별 표준화계수가 모두 .6 이상으로 통계적
으로 유의한 적재량을 보이고 있었으며, 다
중상관자승(SMC) 값도 .4 이상으로 나타났다. 한편, 모든 잠재 변인의 개념 신뢰도(construct
reliability)가 .8 이상이었으며, 평균분산추출
(Average Variance Extracted)이 .5 이상으로 나타
나 내적 일관성 및 수렴 타당성을 확보하였다. 모형이 남녀 재직근로자 모두에게 적합한 모
형인지 확인하기 위해 집단 간 형태동일성을
검증하였다. 그 결과 남녀 집단 모두에서 3
요인 모형이 적합한 것으로 나타났다(χ2=
353.941(df=124, p<.001), CFI=.936, TLI=.920,
RMSEA=.054 [하한값: .048, 상한값: .061]). 형
태동일성 모형을 기저모형으로 하고 잠재변인
과 측정변인 간 경로를 제약한 측정동일성 모
형을 비교하였다. 분석방법 측정요인 계수의 동일화 제
약을 가한 측정동일성 모형과 기저모형의 χ2
차이검증을 실시한 결과 기저모형(모형 A)과
측정동일성 모형(모형 B)간의 χ2값의 차이는
통계적으로 유의한 것으로 나타났으나(Δχ2(Δ
df)=18.481(10), p=.047), 모형A에 비해 적합도
가 근소한 차이로 감소하였으므로 측정동일성
모형을 수용하였다. 다음으로 집단 간 측정오
차의 잔차가 동일하다고 제약한 모형 C와 측
정동일성 모형을 비교하였다. 두 모형 간 χ2
값의 차이는 통계적으로 유의하지 않았다
(Δχ2(Δdf)=20.816(13), p=.077). 따라서 모형 C
를 수용하였다. 마지막으로 집단 간 요인계 칼레이도스코프측정도구의성별에따른잠재
평균차이분석 칼레이도스코프측정도구의성별에따른잠재
평균차이분석 칼레이도스코프측정도구의성별에따른잠재
평균차이분석 칼레이도스코프 측정도구의 3요인 모형이
남녀 재직근로자 모두에게 적용할 수 있는 모
형인지 확인하기 위해 다집단분석을 실시하였
다. 먼저 칼레이도스코프 측정도구의 3요인 Model
χ2(df)
Δχ2(Δdf)
χ2/df
CFI
TLI
RMSEA
Model A
353.941(124)
-
2.854***
.936
.920
.054
(.048-.061)
Model B
372.422(134)
18.481* (10)
2.779***
.934
.923
.053
(.047-.060)
Model C
393.238(147)
20.816 (13)
2.675***
.932
.928
.052
(.045-.058)
Model D
414.703(153)
21.466** (6)
2.710***
.927
.926
.052
(.046-.058)
Note: Model A: unconstrained model; Model B: constrained measurement weights; Model C: constrained
measurement weights and intercepts; Model D: constrained measurement weight, intercepts, and covariance
The result of multi-group analysis between gender The result of multi-group analysis between gender - 69 - 한국심리학회지: 산업및조직 집단 이상에서 요인의 평균을 동시에 추정할
수 없으므로 남성 근로자를 참조집단으로 하
여 하위 요인의 평균을 0으로 고정한 뒤 여성
근로자의 하위 요인 평균을 추정하였다. 표 6
의 남녀 근로자 집단 간 잠재평균 차이분석을
살펴보면 남성에 비해 여성의 잠재평균이 상
대적으로 낮은 것을 확인할 수 있으며, 진정
성 요인에서의 차이는 유의하지 않았으나, 균
형과 도전에서의 잠재평균차이는 유의한 것으
로 나타났다. Cohen’s d값을 살펴보면 균형의
경우 d값이 -.320, 도전의 경우 -.255로 둘 다
.20을 넘는 것으로 나타나 중간 수준의 효과
크기를 갖는 것(Cohen, 1988)으로 확인되었다. 즉, 남성 재직근로자가 칼레이도스코프 경력
태도 중 균형과 도전을 더 높게 인식하고 있
는 것을 확인할 수 있었다. 따라서 여성이 남
성에 비해 균형에 대한 인식이 더 높을 것이
라는 가설 1은 기각되었다. 수 뿐 아니라 각 잠재변인의 공분산도 동일
하다고 제약한 모형을 모형 C와 비교하였다. χ2차이검증을 실시한 결과 두 모형 간 χ2값
은 통계적으로 유의한 것으로 나타났다(Δχ2
(Δdf)=21.466(6), p=.002). 공분산까지 동일화
제약을 가한 모형 D는 모형 C보다 적합도가
다소 떨어지는 것을 알 수 있으나, χ2 차이검
증의 경우 사례 수에 민감하여 적합한 모형을
기각할 우려가 있으며, CFI가 .01보다 적게 감
소하거나 RMSEA가 적게 증가한 경우에는 적
합도에 차이가 없는 것으로 해석할 수 있다는
주장(Chen, 2007; Cheung & Rensvold, 2002)도
제기된 바 있다. 모형 D가 모형 C와 χ2차이
검증에서 유의한 차이를 보였으나, 모형 D의
적합도가 나쁘다고 해석하기에는 CFI, TLI 지
수가 .9 이상으로 양호하고, CFI의 감소폭이
0.1보다 적고, TLI와 RMSEA는 오히려 증가한
것을 고려할 때 적합도 차이가 없는 것으로
해석할 수도 있다. 따라서 성별에 따라 형태
동일성, 측정동일성, 측정오차의 잔차동일성,
공분산동일성이 모두 인정되므로 집단 간 측
정의 교차타당성이 확보되었다고 볼 수 있다. 집단 간 측정의 동일성 가정이 충족되어 두
집단 간 칼레이도스코프 하위요인의 잠재평균
비교분석을 실시하였다. 잠재평균 비교 시 두 칼레이도스코프
측정도구의
준거관련타당도
분석 A=Authenticity, B=Balance, C=Challenge
***p<.001 A=Authenticity, B=Balance, C=Challenge 칼레이도스코프
측정도구의
준거관련타당도
분석 칼레이도스코프의 구인타당도와 교차타당도
를 확인한 뒤 칼레이도스코프 경력태도의 총
점 및 하위요인과 관련변인 간 상관관계 분석 Table 6 Parameter
Male
Female
t
Effect size
(Cohen’s d)
Authenticity
.00
-.06
-1.030
-.015
Balance
.00
-.16
-2.734**
-.320
Challenge
.00
-.16
-2.386*
-.255
*p <.05, **p <.01
Table 6
The result of latent mean analysis between gender The result of latent mean analysis between gender - 70 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 을 통해 측정도구의 수렴타당도와 준거관련
타당도를 확인하였다. 먼저, 칼레이도스코프의
총점과 하위요인 간 상관관계 분석을 통해 수
렴타당도를 확인한 결과 칼레이도스코프 경력
태도 총점은 진정성과 .839, 균형과 .785, 도전
과 .866으로 높은 상관을 갖는 것으로 나타났
다. 또한 세 하위요인 간 상관관계를 살펴보
면 .444-.674의 범위를 갖고 있는 것으로 나타
나 칼레이도스코프 경력태도라는 구성개념을
측정하기 위한 내적 타당도를 갖추었다고 판
단할 수 있다. 과 유의미한 관계에 있을 것이라고 가정한 프
로티언 경력태도, 일-삶의 균형, 도전적 직무
경험, 주관적 경력성공, 경력몰입과의 상관분
석을 통해 준거관련 타당도를 확인하였다. 먼
저 자기주도성과 가치지향성의 하위요인으로
구성된 프로티언 경력태도와 칼레이도스코프
경력태도 총점과의 상관관계는 .708로 높은
수준으로 나타났으며, 특히 칼레이도스코프의
하위요인 중에서도 진정성은 자기주도성과 높
은 상관(r=.673, p<.001)을 갖는 것으로 확인
되었다. 칼레이도스코프 경력태도의 총점은
일-삶의 균형(r=.562, p<.001), 도전적 직무경 그 다음으로 칼레이도스코프의 각 하위요인 Table 7
Correlation between kaleidoscope career attitude and related variables (N=631) Variables
M
SD
Korean vesion of Kaleidoscope Career Attitude
(KKCA)
Result of
correlation
difference analysis
Total
KKCA
Authenticity
Balance
Challenge
Total KKCA
4.31
0.63
1
Authenticity
4.44
0.71
.839***
1
Balance
4.35
0.79
.785***
.444***
1
Challenge
4.15
0.77
.866***
.674***
.482***
1
Protean Career Attitude
4.22
0.60
.708***
.631***
.446***
.691***
B < A, C
Self-directed
4.37
0.62
.732***
.673***
.475***
.683***
B < A, C
Value-driven
4.07
0.72
.544***
.467***
.331***
.559***
B < A < C
Work-life Balance
4.20
0.76
.562***
.433***
.455***
.507***
A = B = C
Challenging job behavior
3.02
0.83
.481***
.353***
.298***
.546***
A = B < C
Subjective Career Success
3.69
0.81
.476***
.352***
.290***
.541***
A = B < C
Career satisfaction
3.61
0.98
.366***
.236***
.231***
.441***
A = B < C
Employability
3.78
0.84
.494***
.407***
.293***
.533***
B < A < C
Career commitment
3.62
1.00
.446***
.318***
.280***
.510***
A = B < C
Table 7
Correlation between kaleidoscope career attitude and related variables (N=631) Correlation between kaleidoscope career attitude and related variables (N=631) - 71 - 한국심리학회지: 산업및조직 하였다. 칼레이도스코프
측정도구의
준거관련타당도
분석 델파이조사를 통해 수정된 문항을 설
문조사 도구로 개발, 347명의 기업근로자를
대상으로 측정하고 탐색적 및 확인적 요인분
석을 실시하여 구성개념 타당도를 검증하였다. 마지막으로 새롭게 수집된 631명의 기업 근로
자 표본을 대상으로 칼레이도스코프 경력태도
와 관련 변인을 함께 측정하여 준거관련 타당
도를 확인하였다. 연구 결과를 통해 얻은 결
론 및 시사점은 다음과 같다. 험(r=.481, p<.001), 주관적 경력성공(r=.476,
p<.001), 경력몰입(r=.446, p<.001)과도 유의미
한 상관관계를 갖는 것으로 나타났다. 칼레이
도스코프 하위요인 중 도전은 도전적 직무경
험과의 상관관계가 다른 하위요인에 비해 가
장 큰 것으로 나타났으며(r=.546, p<.001), 균
형은 일-삶의 균형과 r=.455 (p.<.001)로 유의
한 상관관계를 가졌다. 따라서 가설 2, 3, 4, 5,
6은 모두 지지되었다. 한편, Fisher의 Z 변환점수를 통해 칼레이도
스코프 경력 파라미터와 관련 변인 간 상관관
계의 차이가 있는 지 분석하였다. 유의수준은
.05에서 양측검정을 통해 통계적 유의도를 판
단하였다. 일과 삶의 균형과 칼레이도스코프
경력 하위 파라미터 간 상관관계는 유의한 차
이를 보이지 않았으나, 프로티언 경력태도와
의 상관관계에서는 균형(.446)에 비해 진정성
(.631, z=-4.667, p<.000)과 도전(.691, z=-6.559,
p<.000)이 유의하게 높은 것으로 나타났으며,
일 삶의 균형, 도전적 직무행동, 주관적 경력
성공, 경력몰입에서는 진정성, 균형에 비해 도
전과의 상관관계가 유의하게 높은 것을 확인
할 수 있었다. 이는 도전 파라미터에 응답값
이 높은 사람들의 경우 자신의 경력 결과를
더 긍정적으로 평가하고 경력에 보다 몰입하
고 있음을 시사한다. 첫째, 문화적 배경에 따라 잠재변인을 측정
하는 구체적인 행동요소들이 달라질 수 있는
데, 이 연구는 델파이, 두 차례의 표집을 통해
반복 검증하여 타당도를 확보한 도구 개발 절
차를 거친 바, 칼레이도스코프의 한국판 측정
도구를 타당화했다는 데 의의가 있다. 최근
Kim과 Lee(2020)는 중소기업 초기 경력자를 대
상으로 칼레이도스코프 경력태도를 측정하여
탐색적 요인분석을 실시하고, 균형과 도전에
서 각각 한 문항씩 삭제된 도구를 활용했는데,
제거된 문항에는 이 연구에서 제거한 균형 1
번 (필요하다면, 나는 가족과 관련된 문제를
해결하기 위해 일을 포기할 것이다) 문항이
포함되어 있었다. 우리나라와 같이 노동시장
이 경직된 사회적 맥락에서는 퇴직 후 유사한
조건의 일을 구하는 것이 상대적으로 어려우
므로 해당 문항은 우리나라 문화에 적용하기
에 부적절한 것으로 판단된다. 이 연구는 정
성적 접근과 정량적 접근을 모두 활용한 통합
적 접근법(mixed method)을 바탕으로 수정델파
이법과 두 개의 표본을 통한 반복 검증을 통
해 원도구의 측정 문항 중 우리나라 기업근로
자에게 적용하기 어려운 문항을 가려냄과 동
시에 우리나라 기업근로자의 칼레이도스코프
경력태도를 측정하는 추가문항(진정성-나는 일
을 통해 내 삶 속에서 진정한 나를 발견하기 논 의 이 연구의 목적은 칼레이도스코프 경력태도
의 한국판 측정도구를 개발하는 데 있다. 칼레이도스코프
측정도구의
준거관련타당도
분석 이
를 위해 문헌고찰, 번역-역번역 방법을 통해
마련한 칼레이도스코프 경력태도 측정도구 초
안을 수정델파이법을 통해 내용타당도를 확인 - 72 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 를 원한다; 균형-나는 가족으로부터 얻는 행복
과 일에서 얻는 성취를 동시에 추구한다; 도
전-나에게 있어 새로운 변화란 나를 성장시킬
수 있는 기회이다)을 개발하고 타당도를 검증
했다는 점에서 의의가 있다. 이 연구를 통해
최종적으로 마련된 14문항은 후속연구에서
지속적으로 연구결과를 축적할 필요가 있다. 둘째, 한국판 칼레이도스코프 경력태도를
측정하는 3요인의 14문항은 서로 다른 두 표
본에서도 구인타당도가 확인되었으며, 남녀
성별 집단에서도 동일한 측정도구를 활용할
수 있는 것으로 확인되었다. 칼레이도스코프
경력태도를 구성하는 진정성, 균형, 도전의 3
요인 모형은 서로 다른 표본에서도 안정적으
로 도출되었다. 한편, 칼레이도스코프 경력태
도는 개인이 생애 단계에서 어떤 경력 파라미
터를 중요하게 생각하는지를 핵심으로 하고
있으며, 세 가지 파라미터의 절대 점수보다는
3요인 점수의 프로파일이 중요한 정보를 제공
한다. 선행연구도 이를 고려하여 알파 패턴,
베타 패턴 등으로 명명하면서 칼레이도스코프
경력 파라미터를 중심으로 판단해 왔다. 예를
들어, Sullivan과 Mainiero(2007)는 응답자 중 대
부분(84%)의 남성들은 초기-중기-말기의 경력
단계에서 도전-진정성-균형에 각각 초점을 주
는 알파 칼레이도스코프 경력 패턴을 갖고 있
는 반면, 대부분의 여성(83%)들은 초기에는 도
전, 중기에는 균형, 말기에는 진정성에 초점을
두는 베타 칼레이도스코프 경력 패턴을 보였
다고 보고하였다. 최근 수행된 Mainiero와
Gibson(2017)의 연구에서는 경력중기의 남녀
실업자를 대상으로 칼레이도스코프 경력태도
의 차이를 밝혔는데, 여성이 남성에 비해 균
형을 훨씬 중요하게 생각하고 있었으며, 경력
후기로 갈수록 남녀 모두 진정성을 더 강조하 는 것으로 나타났다. 이 연구에서는 진정성과
도전의 상관관계(r=.674)에 비해 진정성과 균
형(r=.444), 도전과 균형(r=.482)의 상관관계는
상대적으로 낮게 나타나 칼레이도스코프의 경
력 파라미터가 배타적인 양상으로 존재할 가
능성을 시사한다. 따라서 칼레이도스코프 경
력태도를 연구하고 해석하는 데 있어 칼레이
도스코프 경력태도의 프로파일을 분석하고 이
를 토대로 한 연구를 진행할 필요가 있다. 셋째, 이 연구는 칼레이도스코프 경력태도
중 균형 파라미터에서 여성이 남성에 비해 더
높은 점수를 가질 것이라 가정하였으나, 연구
결과 칼레이도스코프의 잠재평균은 남성이 여
성보다 높은 것을 밝혔다. 특히, 균형 및 도전
에서 잠재평균 차이가 유의미했으며, 균형에
서의 차이는 효과크기가 가장 큰 것으로 나타
났다. 이는 여성보다 남성들이 칼레이도스코
프의 전반적인 응답값이 상대적으로 높음을
의미하며, 여성들이 남성보다 균형을 더 강조
한다는 선행연구(Mainiero & Gibson, 2017)와는
상반된 결과이다. Mainiero와 Gibson(2017)의 선
행연구에서 칼레이도스코프 경력태도의 성별
차이를 밝히는 데 있어 경력단계를 함께 고려
했기 때문에 이러한 차이가 나타난 것으로 판
단된다. 단순히 남녀 간 성별의 차이가 아니
라 경력단계 또는 생애발달 단계에서 주어지
는 다양한 과업과 역할 변화에 따라 성차가
존재할 수 있다는 것이다. 한국 사회에서 일
반적으로 여성에 비해 남성의 평균적인 지위,
급여수준이 높고 고용형태가 보다 안정적이라
는 것은 잘 알려진 사실이다. 칼레이도스코프
측정도구의
준거관련타당도
분석 칼레이도스코프
경력태도의 차이가 이러한 사회맥락적 특성
및 보편적 인식에 따른 것인지, 또는 남성이
경력과 관련하여 변화된 가치를 여성보다 용
이하게 받아들인 결과인지 확인하는 연구가 는 것으로 나타났다. 이 연구에서는 진정성과
도전의 상관관계(r=.674)에 비해 진정성과 균
형(r=.444), 도전과 균형(r=.482)의 상관관계는
상대적으로 낮게 나타나 칼레이도스코프의 경
력 파라미터가 배타적인 양상으로 존재할 가
능성을 시사한다. 따라서 칼레이도스코프 경
력태도를 연구하고 해석하는 데 있어 칼레이
도스코프 경력태도의 프로파일을 분석하고 이
를 토대로 한 연구를 진행할 필요가 있다. 를 원한다; 균형-나는 가족으로부터 얻는 행복
과 일에서 얻는 성취를 동시에 추구한다; 도
전-나에게 있어 새로운 변화란 나를 성장시킬
수 있는 기회이다)을 개발하고 타당도를 검증
했다는 점에서 의의가 있다. 이 연구를 통해
최종적으로 마련된 14문항은 후속연구에서
지속적으로 연구결과를 축적할 필요가 있다. 둘째, 한국판 칼레이도스코프 경력태도를
측정하는 3요인의 14문항은 서로 다른 두 표
본에서도 구인타당도가 확인되었으며, 남녀
성별 집단에서도 동일한 측정도구를 활용할
수 있는 것으로 확인되었다. 칼레이도스코프
경력태도를 구성하는 진정성, 균형, 도전의 3
요인 모형은 서로 다른 표본에서도 안정적으
로 도출되었다. 한편, 칼레이도스코프 경력태
도는 개인이 생애 단계에서 어떤 경력 파라미
터를 중요하게 생각하는지를 핵심으로 하고
있으며, 세 가지 파라미터의 절대 점수보다는
3요인 점수의 프로파일이 중요한 정보를 제공
한다. 선행연구도 이를 고려하여 알파 패턴,
베타 패턴 등으로 명명하면서 칼레이도스코프
경력 파라미터를 중심으로 판단해 왔다. 예를
들어, Sullivan과 Mainiero(2007)는 응답자 중 대
부분(84%)의 남성들은 초기-중기-말기의 경력
단계에서 도전-진정성-균형에 각각 초점을 주
는 알파 칼레이도스코프 경력 패턴을 갖고 있
는 반면, 대부분의 여성(83%)들은 초기에는 도
전, 중기에는 균형, 말기에는 진정성에 초점을
두는 베타 칼레이도스코프 경력 패턴을 보였
다고 보고하였다. 최근 수행된 Mainiero와
Gibson(2017)의 연구에서는 경력중기의 남녀
실업자를 대상으로 칼레이도스코프 경력태도
의 차이를 밝혔는데, 여성이 남성에 비해 균
형을 훨씬 중요하게 생각하고 있었으며, 경력
후기로 갈수록 남녀 모두 진정성을 더 강조하 셋째, 이 연구는 칼레이도스코프 경력태도
중 균형 파라미터에서 여성이 남성에 비해 더
높은 점수를 가질 것이라 가정하였으나, 연구
결과 칼레이도스코프의 잠재평균은 남성이 여
성보다 높은 것을 밝혔다. 특히, 균형 및 도전
에서 잠재평균 차이가 유의미했으며, 균형에
서의 차이는 효과크기가 가장 큰 것으로 나타
났다. 이는 여성보다 남성들이 칼레이도스코
프의 전반적인 응답값이 상대적으로 높음을
의미하며, 여성들이 남성보다 균형을 더 강조
한다는 선행연구(Mainiero & Gibson, 2017)와는
상반된 결과이다. Mainiero와 Gibson(2017)의 선
행연구에서 칼레이도스코프 경력태도의 성별
차이를 밝히는 데 있어 경력단계를 함께 고려
했기 때문에 이러한 차이가 나타난 것으로 판
단된다. 단순히 남녀 간 성별의 차이가 아니
라 경력단계 또는 생애발달 단계에서 주어지
는 다양한 과업과 역할 변화에 따라 성차가
존재할 수 있다는 것이다. 한국 사회에서 일
반적으로 여성에 비해 남성의 평균적인 지위,
급여수준이 높고 고용형태가 보다 안정적이라
는 것은 잘 알려진 사실이다. 칼레이도스코프
측정도구의
준거관련타당도
분석 칼레이도스코프
경력태도의 차이가 이러한 사회맥락적 특성
및 보편적 인식에 따른 것인지, 또는 남성이
경력과 관련하여 변화된 가치를 여성보다 용
이하게 받아들인 결과인지 확인하는 연구가 - 73 - 한국심리학회지: 산업및조직 여성 CEO, 실업자, 교사, 은퇴기 여성 등을 대
상으로 연구가 수행된 바 있다. 개인의 주도
성이 강조되는 경력 패러다임 시대의 새로운
경력태도로 지칭되는 칼레이도스코프 경력태
도가 전체 직업군에 적용될 수 있는 개념인지
다양한 표본을 통한 반복 검증 연구를 실시하
여 확인할 필요가 있다. 이러한 연구들은 칼
레이도스코프 경력태도의 개념 뿐 아니라 경
력과 관련된 변인 간 관계와 주요 메커니즘에
대한 이해도 확장시킬 수 있을 것이다. 또한,
기업, 공공기관 등 조직의 인사부서에서 경력
태도 척도를 채택하여 활용할 경우, 보직임면
장면에서 경력태도 진단 및 피드백을 통해 긍
정적인 직원경험(employee experience)을 부여할
수 있을 것이다. 특히 세 가지 경력 파라미터
와 관련하여 경력개발 개입방안을 체계적으로
구조화하여 제공할 필요가 있다. 둘째, 칼레이
도스코프 경력태도의 세대차에 대한 연구도
수행될 필요가 있다. 이 연구에서는 표집의
한계로 인해 세대차이는 다루지 않았으나,
Sullivan과 그의 동료들(2009)은 X세대가 베이
비부머 세대에 비해 진정성과 균형에서 유의
하게 높은 것으로 나타났음을 보고하였다. 성
별 뿐 아니라 세대 간 집단에서 진정성, 균형,
도전과 같은 중요한 가치의 차이가 어떠한 지
확인할 필요가 있을 것이다. 마지막으로 경력
단계를 고려한 칼레이도스코프 경력태도에 대
한 연구가 이루어져야 할 것이다. 경력은 조
직 관점에서는 인적자원 차원에서 적시적소에
한정된 역량을 배치 및 활용하기 위한 매개물
로 받아들여지나, 개인 관점에서는 시간의 흐
름에 따른 역량의 발달뿐만 아니라 가족 돌봄
을 포함한 개인의 삶을 영위하는 태도 전반에
관계된 목적물이다. 일의 세계는 계속해서 변
화하고 있으며, 이에 따라 최근에 조직이 학 필요하다. 필요하다. 넷째, 이 연구는 칼레이도스코프 경력태도
의 합산 값을 어떻게 활용할 수 있을지에 대
한 자료를 제공하고 있다는 점에서 의의가 있
다. 선행연구는 주로 칼레이도스코프 경력태
도의 경력파라미터에 따른 패턴을 분석하는데
초점을 두고 있어 전체 칼레이도스코프 경력
태도 값을 어떻게 해석할 수 있을지에 대한
근거를 제시하는 데 한계가 있었다. 이 연구
에서는 칼레이도스코프 경력태도와 관련 변인
간 상관관계를 바탕으로 칼레이도스코프 경력
태도가 보다 적응적인 경력태도와 관련되어
있을 것이라는 가설을 확인하였다. 칼레이도
스코프 경력태도는 프로티언 경력태도, 일-삶
의 균형, 도전적 직무경험과 유의미한 상관관
계를 갖는 것으로 나타났는데, 그 중에서도
전반적으로 도전 파라미터가 준거 변인들과의
상관이 높게 나타났다. 이는 도전이라는 경력
파라미터를 중요하게 생각하는 기업근로자가
자신의 경력과 관련하여 보다 긍정적인 태도
를 갖는 것으로 해석할 수 있다. 최근 경력개
발에서의 개인의 주도성을 강조하는 새로운
경력태도는 고용환경 변화의 가속화와 무관하
지 않으며, 주어진 상황에서 새로운 도전을
추구하고 주도성을 발휘해 나가는 것이 개인
의 경력개발에서도 중요한 역할을 하고 있음
을 시사한다. 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 습하고, 다수의 개인이 채택하는 경력태도는
다양하다. 칼레이도스코프는 세 가지 경력파
라미터의 현재 수준보다는 이러한 경력 파라
미터가 어떻게 달라지는 지에 초점을 두고 있
는 개념이므로 향후 종단 연구나 다양한 경력
단계에서의 칼레이도스코프 경력태도 인식 차
이 연구 등이 이루어져야 할 것이다. Combinations and implications. Journal of
Vocational Behavior, 69(1), 4-18. doi:10.1016/j.jvb.2005.09.002 Briscoe, J. P., Hall, D. T., & DeMuth, R. L. F. (2006). Protean and boundaryless careers:
An empirical exploration. Journal of vocational
behavior, 69(1), 30-47. doi:10.1108/cdi-04-2013-0051 Brislin, R. (1973). Questionnaire wording and
translation. Cross-cultural research methods, 32-58. 칼레이도스코프
측정도구의
준거관련타당도
분석 연구결과를 바탕으로 칼레이도스코프 경력
태도와 관련하여 다음과 같은 연구들이 수행
될 필요가 있다. 첫째, 이 연구에서 확인한 한
국판 칼레이도스코프 경력태도를 활용하여 보
다 다양한 맥락의 근로자를 대상으로 연구할
필요가 있다. 이 연구는 우리나라의 20대에서
60대의 남녀 기업근로자들을 대상으로 비율
표집을 통해 표본을 구성하였다. 국외에서는 - 74 - 이재은․정보영/ 한국판칼레이도스코프경력태도측정도구타당화연구 참고문헌 Cabrera, E. F. (2007). Opting out and opting
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Resource Development, 48(1), 97-127. doi:10.23840/agehrd.2016.48.1.97 mediating effect of democratic characteristics. Journal of Agricultural Education and Human
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S.,
&
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Causal
Relationships
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Success,Job
Security,
Organizational
Career
Management Support, Protean Career Attitude,
and Protean Career Management Behavior of
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Career
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and
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A
Critical
Review and Agenda for Future Exploration. Journal of Management, 35(6), 1542-1571. †Corresponding Author: Boyoung Jung, Senior Manager, HRD Team, byjung@hyundai.com * This work was supported by the Ministry of Education of the Republic of Korea and the National Research
Foundation of Korea(NRF-2019S1A5A8038225). 한국심리학회지: 산업및조직 doi:10.1177/0149206309350082 Sullivan, S. E., & Baruch, Y. (2009). Advances in
Career
Theory
and
Research:
A
Critical
Review and Agenda for Future Exploration. Journal of Management, 35(6), 1542-1571. doi:10.1177/0149206309350082 - 78 - 한국심리학회지: 산업및조직
Korean Journal of Industrial and Organizational Psychology
2021. Vol. 34, No. 1, 51-79
https://doi.org/10.24230/kjiop.v34i1.51-79 한국심리학회지: 산업및조직
Korean Journal of Industrial and Organizational Psychology
2021. Vol. 34, No. 1, 51-79
https://doi.org/10.24230/kjiop.v34i1.51-79 Validation for Korean Version
of Kaleidoscope Career Attitude Measurement* Validation for Korean Version
of Kaleidoscope Career Attitude Measurement* BoYoung Jung† * This work was supported by the Ministry of Education of the Republic of Korea and the National Research
Foundation of Korea(NRF-2019S1A5A8038225).
†Corresponding Author: Boyoung Jung, Senior Manager, HRD Team, byjung@hyundai.com JaeEun Lee Sangji University Sangji University Hyundai Motor Company The purpose of this study is to validate the Kaleidoscope career attitude scale developed by Sullivan et
al. (2009) to measure the new career attitudes of Korean corporate employees. This study can be
summarized as follows: first, the Korean version of the Kaleidoscope career attitude measurement’s
psychometric equivalence and contents validity was verified. Second, the Kaleidoscope career attitude
measurement items consisted of three factors, equivalent to the original scale. Third, the fit of the
measurement model consisting of 13 items was confirmed with the new sample. And it was confirmed
that the same scale could be used in the male and female groups. Fourth, the difference in the latent
mean was significant in balance and challenge parameters. Comprehensive research results revealed that
the Korean version of the Kaleidoscope career attitude is valid and reliable. Based on this result, the
significance and limitations of the research and suggestions for future research were discussed. Key words : Kaleidoscope Career Model(KCM), Career Attitude, Measurement, Validation, Latent Mean Analysis - 79 -
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Hypersensitive-like response in Brassica plants is specifically induced by molecules from egg-associated secretions of cabbage white butterflies
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Frontiers in ecology and evolution
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cc-by
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Pieris, cabbage, cell death, plant-insect interaction, oviposition-induced response,
egg-associated molecular pattern TYPE Original Research
PUBLISHED 01 February 2023
DOI 10.3389/fevo.2022.1070859 TYPE Original Research
PUBLISHED 01 February 2023
DOI 10.3389/fevo.2022.1070859 TYPE Original Research
PUBLISHED 01 February 2023
DOI 10.3389/fevo.2022.1070859 COPYRIGHT
© 2023 Caarls, Bassetti, Verbaarschot, Mumm,
van Loon, Schranz and Fatouros. This is an
open-access article distributed under the terms
of the Creative Commons Attribution License
(CC BY). The use, distribution or reproduction
in other forums is permitted, provided the
original author(s) and the copyright owner(s)
are credited and that the original publication in
this journal is cited, in accordance with
accepted academic practice. No use,
distribution or reproduction is permitted which
does not comply with these terms. OPEN ACCESS EDITED BY
Maria L. Pappas,
Democritus University of Thrace, Greece
REVIEWED BY
Junji Takabayashi,
Kyoto University,
Japan
Islam S. Sobhy,
Cardiff University,
United Kingdom Lotte Caarls 1*†, Niccolò Bassetti 1, Patrick Verbaarschot 1,
Roland Mumm 2, Joop J. A. van Loon 3, M. Eric Schranz 1 and
Nina E. Fatouros 1* 1 Biosystematics Group, Wageningen University & Research, Wageningen, Netherlands, 2 Bioscience,
Wageningen University & Research, Wageningen, Netherlands, 3 Laboratory of Entomology, Wageningen
University & Research, Wageningen, Netherlands Plants perceive and respond to herbivore insect eggs. Upon egg deposition on leaves,
a strong hypersensitive response (HR)-like cell death can be activated leading to egg
desiccation and/or dropping. In Brassica spp., including many crops, the HR-like
mechanism against eggs of cabbage white butterflies (Pieris spp.) is poorly understood. Using two Brassica species, the crop B. rapa and its wild relative B. nigra, we studied the
cellular and molecular plant response to Pieris brassicae eggs and characterized potential
insect egg-associated molecular patterns (EAMPs) inducing HR-like cell death. We found
that eggs of P. brassicae induced typical hallmarks of early immune responses, such
as callose deposition, production of reactive oxygen species and cell death in B. nigra
and B. rapa leaf tissue, also in plants that did not express HR-like cell death. However,
elevated levels of ethylene production and upregulation of salicylic acid-responsive
genes were only detected in a B. nigra accession expressing HR-like cell death. Eggs and
egg wash from P. brassicae contains compounds that induced such responses, but the
eggs of the generalist moth Mamestra brassicae did not. Furthermore, wash made from
hatched Pieris eggs, egg glue, and accessory reproductive glands (ARG) that produce
this glue, induced HR-like cell death, whereas washes from unfertilized eggs dissected
from the ovaries or removal of the glue from eggs resulted in no or a reduced response. This suggests that there is one or multiple egg associated molecular pattern (EAMP)
located in the egg glue a that teresponse in B. nigra is specific to Pieris species. Lastly,
our results indicate that the EAMP is neither lipidic nor proteinaceous. Our study expands
the knowledge on the mechanism of Brassica-Pieris-egg interaction and is a step closer
toward identification of EAMPs in Pieris egg glue and corresponding receptor(s) in Brassica. OPEN ACCESS CITATION
Caarls L, Bassetti N, Verbaarschot P, Mumm R,
van Loon JJA, Schranz ME and
Fatouros NE (2023) Hypersensitive-like
response in Brassica plants is specifically
induced by molecules from egg-associated
secretions of cabbage white butterflies. Front. Ecol. Evol. 10:1070859. doi: 10.3389/fevo.2022.1070859 COPYRIGHT
© 2023 Caarls, Bassetti, Verbaarschot, Mumm,
van Loon, Schranz and Fatouros. This is an
open-access article distributed under the terms
of the Creative Commons Attribution License
(CC BY). The use, distribution or reproduction
in other forums is permitted, provided the
original author(s) and the copyright owner(s)
are credited and that the original publication in
this journal is cited, in accordance with
accepted academic practice. No use,
distribution or reproduction is permitted which
does not comply with these terms. frontiersin.org Hypersensitive-like response in
Brassica plants is specifically
induced by molecules from
egg-associated secretions of
cabbage white butterflies OPEN ACCESS
EDITED BY
Maria L. Pappas,
Democritus University of Thrace, Greece
REVIEWED BY
Junji Takabayashi,
Kyoto University,
Japan
Islam S. Sobhy,
Cardiff University,
United Kingdom
*CORRESPONDENCE
Lotte Caarls
lotte.caarls@wur.nl
Nina E. Fatouros
nina.fatouros@wur.nl
†PRESENT ADDRESS
Lotte Caarls,
Plant Breeding,
Wageningen University and Research,
Wageningen, Netherlands
SPECIALTY SECTION
This article was submitted to
Chemical Ecology,
a section of the journal
Frontiers in Ecology and Evolution
RECEIVED 15 October 2022
ACCEPTED 19 December 2022
PUBLISHED 01 February 2023
CITATION
Caarls L, Bassetti N, Verbaarschot P, Mumm R
van Loon JJA, Schranz ME and
Fatouros NE (2023) Hypersensitive-like
response in Brassica plants is specifically
induced by molecules from egg-associated
secretions of cabbage white butterflies. Front. Ecol. Evol. 10:1070859. doi: 10.3389/fevo.2022.1070859
COPYRIGHT
© 2023 Caarls, Bassetti, Verbaarschot, Mumm
van Loon, Schranz and Fatouros. This is an
open-access article distributed under the ter
of the Creative Commons Attribution Licens
(CC BY). The use, distribution or reproductio
in other forums is permitted, provided the
original author(s) and the copyright owner(s)
are credited and that the original publication 1. Introduction Plants rely on an immune system that regulates the perception of attackers and subsequent
activation of inducible defenses (Wilkinson et al., 2019). Perception involves detection of pathogen-
derived effector proteins and of molecular patterns, which can derive from different organisms, such as
microbes (MAMPs) or herbivores (HAMPs; Gust et al., 2017; Stahl et al., 2018; Van der Burgh and Frontiers in Ecology and Evolution 01 10.3389/fevo.2022.1070859 Caarls et al. prime plants for future larval attack (Fatouros et al., 2008, 2009, 2015;
Paniagua Voirol et al., 2020). Anti-aphrodisiacs transferred from the
butterfly male to the female during mating were shown to be present in
minute amounts in the ARG secretion and were suggested as potential
elicitors (Fatouros et al., 2008, 2009). However, glands from unmated
females induced HR-like, and therefore another, female-derived elicitor
is likely to play a role (Fatouros et al., 2015). In A. thaliana, egg-derived
phosphatidylcholines were found to induce production of H2O2, SA and
result in cell death as shown by trypan blue staining (Stahl et al., 2020). It is still unclear which compounds from eggs of Pieris spp. and
Anthocharis cardamines are detected by Brassica spp. and close relatives
that result in HR-like activation, and whether these reside in the eggs
themselves or in the secretions surrounding the eggs (Griese et al., 2021). Joosten, 2019). Perception is followed by an early signaling cascade
including rapid ion-flux changes and production of reactive oxygen species
(ROS; Couto and Zipfel, 2016). Induced defenses typically include
reinforcement of extracellular barriers, for example by callose deposition,
the production of antimicrobial or insecticidal metabolites and proteins,
and a localized, rapid cell death response, the hypersensitive response (Cui
et al., 2015; Couto and Zipfel, 2016; Campos et al., 2018; Balint-Kurti,
2019; Dalio et al., 2020). Most in-depth studies on activation of the plant
immune system have been performed on interactions with plant
pathogens. So far, 119 HAMPs and/or effectors mostly from oral secretions
of piercing sucking and chewing herbivores, have been identified, but
pattern recognition receptors for HAMPs are just starting to be discovered
(Erb and Reymond, 2019; Steinbrenner et al., 2020).h The detection by plants of herbivore eggs deposited on plant tissue is
remarkable, as eggs are immobile and seemingly harmless structures. However, insect eggs will turn into feeding larvae and thus pose a future
threat to the plant. 2.1. Plant material and rearing of butterflies Black mustard (B. nigra L.) accessions WUR-01 and WUR-02 used,
originated from plants that were collected near the river Rhine in
Wageningen, Netherlands (N51.96, E05.68). Brassica rapa L. genotypes
used in this study (L58, R-o-18, RC-144) were obtained from the
Laboratory of Plant Breeding (WUR). Plants were grown in a greenhouse
(18 ± 5°C, 50–70% RH, L16, D8) and were used when three to
five weeks old. Besides HR-like cell death, eggs of different insect species induce
immune responses similar to pattern-triggered immunity (PTI),
including callose deposition, accumulation of ROS and SA, and
transcriptome
changes
of
several
defense
genes,
including
PATHOGENESIS-RELATED PROTEIN 1 (PR1) in A. thaliana (Little
et al., 2007; Bruessow et al., 2010; Gouhier-Darimont et al., 2013;
Reymond, 2013; Lortzing et al., 2020). These early signaling defense
responses to eggs seem to be conserved across different plant species
(Lortzing et al., 2020). Expression of PR1 was also induced in leaves of
B. nigra underneath P. brassicae and P. rapae eggs (Fatouros et al., 2014,
2015). Whether brassicaceous species that are natural hosts of Pieris spp.,
including B. nigra and B. rapa, respond with a general immune response,
including ROS, callose and expression of different defense genes, to insect
eggs, and whether there is genetic variation for this response, is
largely unknown. i
Pieris brassicae L. (Lepidoptera: Pieridae) was reared on Brussels
sprouts plants (Brassica oleracea var. gemmifera cv. Cyrus) in a climate
room (21 ± 1°C, 50%–70% RH, L16: D8) at the Laboratory of
Entomology, Wageningen University. Virgin adult females were obtained
by isolating female butterflies immediately after eclosion. Otherwise,
twenty females and males could mate in a large cage (60 × 60 × 90 cm)
and females were used for oviposition in experiments or oviposition on
filter paper (grade 3 hw, Sartorius, Germany) for egg wash production
(see section below). The cabbage moth Mamestra brassicae
L. (Lepidoptera: Noctuidae) was reared on Brussels sprouts plants in a
climate room (21 ± 1°C, 50%–70% RH, L16: D8) at the Laboratory of
Entomology, Wageningen University. So far, few studies have identified EAMPs that activate defense
against insect eggs (Reymond, 2013; Hilker and Fatouros, 2015; Stahl
et al., 2018). In some studies, secretions surrounding eggs were sufficient
to elicit defense responses in plants and a few elicitors have been isolated
from these secretions (Hilker et al., 2005; Tamiru et al., 2011; Salerno
et al., 2013). 2.1. Plant material and rearing of butterflies Most EAMPs were found to be organic compounds of
low-molecular weight associated with the eggs, while recently a first
proteinaceous compound, diprionin, was found as an elicitor of pine
defense against sawfly eggs (Hundacker et al., 2021). In P. brassicae,
egg-enveloping secretions are produced by the females’ accessory
reproductive gland (ARG) and form a glue-like structure between the
eggs and leaves (Beament and Lal, 1957; Fatouros et al., 2012). Treatment
of Brassica plants with extracts from ARGs has shown to induce HR-like
cell death and plant chemical cues attracting egg parasitoids, and to 1. Introduction Upon detection, plants can mount defenses against
eggs that range from plant-mediated desiccation of eggs, egg dropping,
egg crushing and the production of ovicidal substances (Hilker and
Fatouros, 2015, 2016). Eggs of cabbage white butterflies (Pieris spp.)
trigger necrotic lesions in leaves of the black mustard, Brassica nigra that
can result in egg-killing by desiccating and/or dropping off singly laid
Pieris eggs (Shapiro and DeVay, 1987; Griese et al., 2017). As the
egg-induced plant phenotype resembles an HR, it has been referred to as
hypersensitive response-like (“HR-like”; Fatouros et al., 2012). Oviposition
by Pieris butterflies has been shown to induce HR-like cell death in several
other plants of the Brassicaceae family, although the severity of the
response varies between, and within species (Pashalidou et al., 2015;
Griese et al., 2020, 2021; Groux et al., 2021; Bassetti et al., 2022).f In this study, we developed and implemented a method to
characterize the compounds from Pieris spp. eggs and egg-enveloping
secretions. We specifically addressed: (1) the cellular and molecular
response of two Brassica species, (2) the specificity of these responses by
comparing eggs and egg washes of Pieris with the generalist moth
M. brassicae, (3) the origin of EAMP inducing HR-like cell death, and
(4) the chemical nature and stability of the elicitor. Frontiers in Ecology and Evolution 2.5. Dissection of butterfly reproductive
tract For all experiments testing the effects of treating plants with egg
washes, 10 μL egg wash (i.e., an equivalent of 25 eggs) was pipetted on
the abaxial side of the fourth or fifth emerged leaf of 3–4 weeks old
B. nigra plants, unless otherwise specified. Symptoms induced by egg
wash were scored 4 days after treatment. To quantify severity, a scoring
system from 0 to 4 was adapted from Griese et al. (2017). Severity score
0: no visual response; score 1: brown spots underneath eggs or egg wash
spot, only visible at abaxial side leaf; score 2: cell death also visible at
adaxial side of leaf, spot smaller than 2 mm diameter, score 3: cell death
the size of egg wash spot, and score 4: spreading lesion beyond spot of
treatment (Supplementary Figure S1). Score 0 and 1 are classified as
“non-HR,” score 2–4 are classified as “HR”. In female butterflies, eggs are produced in the ovaries, pass through
the common oviduct and are fertilized by sperm released from the bursa
copulatrix (BC) into the vagina. Before being expelled through the
ovipore, the eggs are covered by secretions released from the accessory
reproductive gland (ARG), a paired gland that contains egg-enveloping
secretions and glue-like material to attach eggs to leaves
(Supplementary Figure S2). For dissection of tissues from the female
reproductive tract, namely ARGs and BCs or eggs from the ovarian
tubules (hereafter termed “ovarian eggs”), mated P. brassicae females were
obtained by pairing a virgin female and virgin male 1 day after eclosion,
which were kept together until mating was observed. Virgin females were
obtained by keeping them separately shortly after eclosion. Three to four
days after eclosion, mated and virgin P. brassicae females were killed by
beheading using a scalpel and then dissected. ARGs, BC and ovarian eggs
were dissected from females under a stereomicroscope (optical
magnification 20×) in 20 mM Mes buffer. Dissected structures were
washed overnight (16 h) in 20 mM Mes pH 5.7 using 50 μL per each ARG,
100 μL buffer per each BC or 5 μL buffer per egg. Solution was pipetted
off the next morning and frozen until use (modified after Fatouros
et al., 2015). For experiments with oviposition on plants, one mated female
P. brassicae butterfly was placed in a cage with a B. nigra or B. rapa plant
and removed when ~10–25 eggs were laid. 2.4. Histochemical staining tested different buffers and found that using Mes buffer resulted in the
most consistent plant response (data not shown). The solution (egg
wash) was pipetted into a new tube the next morning. For histochemical staining of plant tissue underneath eggs,
P. brassicae egg-laden plants were sampled 24, 48, and 72 h after
oviposition by taking a 10 mm diameter leaf disk of the area
surrounding the eggs. Similarly this was done with plants treated
with egg wash. Eggs were carefully removed with a brush before
proceeding with histochemical staining. Pictures of the leaf disks
were taken with a Dino-Lite digital microscope (AnMo Electronics
Corporation, Taiwan) before staining (with eggs) and after staining
(without eggs). As cell death is generally preceded by production
of reactive oxygen species (ROS) such as hydrogen peroxide and
superoxide anion (Torres, 2010), we visualized ROS with different
histochemical stainings. 3,3′-diaminobenzidine (DAB; Sigma-
Aldrich, MO, United States) was used to stain hydrogen peroxide
(H2O2). Leaf disks (10 mm Ø) were submersed in 1 mg/mL DAB
solution and samples were incubated for 30–60 min in the dark. Nitroblue tetrazolium chloride (NBT; Sigma-Aldrich, MO,
United States) was used to stain superoxide radical O2
•−. For this
leaf disks (10 mm Ø) were submersed in 0.2% NBT with 50 mM
sodium phosphate buffer (pH 7.5) and samples were incubated
30–60 min in the dark. For visualization of cell death, leaves were
submersed in 0.08% trypan blue solution (Sigma-Aldrich, MO,
United States), overnight (16 h). For all three stainings, destaining
of leaves was performed with 96% ethanol. Pictures of leaf disks
before staining (with eggs) and after staining (without eggs) were
taken with a Dino-Lite digital microscope (AnMo Electronics
Corporation, Taiwan). For staining of callose, leaf disks were first
destained and then submersed in 0.01% aniline blue (Sigma-
Aldrich, MO, United States) with 150 mM K2HPO4 and imaged
after at least 2 h of incubation using a DAPI filter on a fluorescence
microscope equipped with a DS-5MC camera and NIS elements AR
2.30 software. To obtain egg-enveloping secretions for wash of only egg glue, 1-day
old P. brassicae eggs were collected as above, counted, and then carefully
removed from the paper using a brush. The spots of secretions
underneath eggs were then cut out and washed overnight (16 h) in 1 mL
20 mM Mes buffer (pH 5.7) per 400 eggs. 2.4. Histochemical staining A wash of the pieces of the
same filter paper without secretions was used as control. To remove egg-enveloping secretions from eggs, eggs on paper were
submersed in 1 mL per 400 eggs of either a solution of 1% bleach and 2%
Tween-20 or in 250 mM NaPO4 pH 9.0 for 30 min (two different treatments
that were shown to remove egg glue; Jacobs et al., 2013). After treatment,
eggs were rinsed once with 1 mL MILLI-Q® water (Sigma-Aldrich, MO,
United States), hereafter “MQ water,” and then 1 mL Mes buffer and then
washed overnight (16 h) in 20 mM Mes buffer pH 5.7. Eggs that were used
to compare where egg-enveloping secretions were not removed, were
submersed in Mes for 30 min, rinsed in MQ and Mes, and then washed. To study the induction by eggs of different ages, eggs were collected
as above on paper, and then kept at room temperature until they were
washed at the end of each day, until they hatched (6–7 days after
oviposition). After hatching, young caterpillars were carefully removed
using a brush. Empty eggshells and associated secretions (on paper)
were then washed overnight (16 h). To collect unfertilized eggs for egg wash, filter paper was pinned
underneath a B. nigra leaf of a plant placed in a cage with 10 virgin adult
females and then prepared in a similar way as fertilized eggs. Paper sheets with M. brassicae eggs were obtained from a rearing
population from the Laboratory of Entomology (WUR). To obtain egg
wash, eggs were counted, spots with eggs were cut out and washed in
1 mL 20 mM Mes buffer (pH 5.7) per 400 eggs. The wash was pipetted
off the next morning and frozen at −20°C until use. Frontiers in Ecology and Evolution 2.2. Preparation of egg washes The protocol for preparation of egg washes has been newly
developed to prepare egg wash free from leaf-surface related chemical
compounds which enables us to conduct chemical analysis and identify
possible EAMPs. Full procedure is described in Supplementary Data
(Supplementary Figure S1). Pieris brassicae eggs were collected on filter
paper pinned underneath a B. oleracea leaf in a cage containing 20
mated females. Egg clutches laid on the paper were cut out and
submersed in 1 mL 2-(N-morpholino)ethanesulfonic acid (Mes) buffer
per 400 eggs, overnight (16 h) without disturbance. Previously, we have 02 frontiersin.org Caarls et al. 10.3389/fevo.2022.1070859 2.7. Phospholipid treatments To compare gene expression between P. brassicae and M. brassicae
egg wash, B. nigra plants were treated with 10 μL of either egg wash (400
eggs/mL in Mes buffer) or a negative control (Mes buffer). Six leaf disks
(Ø 6 mm) were sampled directly next to the egg wash-treated spot 24 h
after treatment. Four plants were used for each treatment as
biological replicates. Phosphatidylcholines (PCs) were recently found to be the main
components of P. brassicae egg extracts that show a bioactivity in A. thaliana
Col-0 resulting in induction of SA and PR1 marker gene as well as ROS
accumulation and cell death formation (Stahl et al., 2020). Thus, we tested
whether the same PCs used in Stahl et al. (2020) may be associated with
development of HR-like cell death by B. nigra. Phospholipids were ordered
from Avanti Polar Lipids (Alabaster, Alabama, United States). Specifically,
PC(16:1/16:1), 1,2-dipalmitoleoyl-sn-glycero-3-phosphocholine, catalog
no. 850358 and PC(18:1/18:1), 1,2-dioleoyl-sn-glycero-3-phosphocholine,
catalog no. 850357 were used for experiments. Phospholipid stock solutions
were made in 100% MeOH. For phospholipid application, the MeOH was
evaporated under a nitrogen-flux and the phospholipids were dissolved in
MQ water with 1% DMSO, 0.5% glycerol, 0.1% Tween 20 by sonication. Phospholipids were applied on B. nigra leaves at different concentrations
(1, 5 and 10 μg/μL) that were previously shown to have biological activity
in A. thaliana (Stahl et al., 2020). An MQ water solution of 1% DMSO,
0.5% glycerol and 0.1% Tween 20 was used as control. To compare the gene expression of two B. nigra accessions with
contrasting ability to develop HR-like cell death, egg wash at higher
concentration (~1,000 eggs/mL in demineralised water) was used. Experimental design consisted of two treatments (egg wash,
control), two B. nigra accessions (WUR-01 without HR, and
WUR-02, developing HR), and three time points after treatment (6,
24, and 48 h after treatment). For each treatment combination, egg
wash or control were applied with two droplets of 5 μL on the
abaxial side of a single leaf. Leaf disks (Ø 6 mm) were harvested at
each time point on the treatment spots and disks from the same
treatment on a leaf were pooled. For each treatment combination,
a total of 15 plants were used and groups of 3 plants with similar
treatments were pooled to compose a total of 5 biological replicates. 2.5. Dissection of butterfly reproductive
tract HR-like cell death was scored
4 days after oviposition, using the same scoring system as with egg wash. For experiments with M. brassicae, five mated female moths were placed
together with a B. nigra plant in a cage to allow egg deposition overnight
(16 h) and removed in the morning. No visible cell death was observed
under M. brassicae eggs as was also reported in our previous studies
(Fatouros et al., 2012; Griese et al., 2021). 03 frontiersin.org Caarls et al. 10.3389/fevo.2022.1070859 2.6. Treatments of egg wash with heating,
proteinase K, freezing, methanol and
speedvac extract Netherlands). After sampling leaf disks for the ethylene assay, plants
were also treated with P. brassicae egg wash to determine their HR-like
cell death phenotype. Netherlands). After sampling leaf disks for the ethylene assay, plants
were also treated with P. brassicae egg wash to determine their HR-like
cell death phenotype. To characterize the chemical nature of EAMP in the egg wash
we exposed it to different treatments. Untreated egg wash was used as a
control. To study the stability of the elicitor, the egg wash was heated and
frozen. For heating, egg wash was made as described above, collected in
microcentifuge tubes and boiled in a water bath for 30 min at 95°C. For
freezing treatment, egg wash was frozen at −20°C and then thawed for
1 h at room temperature, then frozen again. Freezing–thawing cycles
were repeated up to four times, after which, all washes were used to treat
plants. To study if the elicitor was an intact protein, a proteinase
treatment was performed, a stock of 20 mg/mL Proteinase K (Qiagen,
Cat No./ID: 19133) was diluted 100× in 100 μL egg wash (10% volume,
pH 7.2) and incubated with egg wash for 30 min at 37°C. 2.8. Production of ethylene Ethylene production is regularly measured to characterize the
induction of early plant immune response by biotic stresses (Fan
et al., 2017). Ethylene was measured as previously published (Oome
et al., 2014). To measure the plant production of ethylene, in B. nigra,
ten plants for each accession (with and without HR-like phenotype)
were used. For B. rapa, three plants of each genotype were used. For
each plant, leaf disks (3 mm Ø) were sampled from mature leaves of
untreated plants and incubated overnight (16 h) in demineralised
water. Subsequently, three leaf disks for each biological replicate (a
single plant) were randomly chosen and incubated for 5 h in air tight
glass vials with in either 400 μL of 20 mM Mes pH 5.7 or 400 μL egg
wash (400 eggs/mL in same Mes buffer). After incubation, 1 mL of the
headspace of each sample was taken to measure ethylene
concentration on a Focus gas chromatograph (Thermo Electron
S.p.A., Milan, Italy) equipped with an FID detector and a RT-QPLOT
column, 15 m × 0.53 mm ID (Restek, Bellefonte, PA, United States). The system was calibrated with a certified gas of 1.01 μL L−1 (1 ppm)
ethylene in synthetic air (Linde Gas Benelux B.V., Schiedam, 2.9. Plant treatments for gene expression
experiments To compare gene expression induced by eggs and egg wash, B. nigra
plants were treated with either 10 μL of egg wash at lower concentration (400
eggs/mL in Mes buffer) or oviposited with an egg clutch of 10 eggs. Six leaf
disks (Ø 6 mm) were sampled with a leaf puncher directly next to the eggs
or the egg wash-treated spot at 0, 3, 6, 24, and 48 h, as done in previous
research (Fatouros et al., 2014; Griese et al., 2021). For each timepoint, four
plants were sampled individually and considered biological replicates. Brassica rapa plants were treated with three single eggs on a single leaf of
each plant. Leaf disks (Ø 6 mm) were then harvested next to the eggs at 0, 3,
6, 24, and 96 h after treatment. For each timepoint, three plants were
sampled individually and considered biological replicates. 2.7. Phospholipid treatments For each gene expression experiment, samples were snap frozen in
liquid nitrogen and stored at −80°C until use. Frontiers in Ecology and Evolution frontiersin.org 2.11. Data analysis All data analysis was carried out in R (R Core Team, 2020). Scoring of HR-like cell death in severity categories were analyzed
with a non-parametric method (Kruskal–Wallis test) on HR scores
(0, 1, 2, 3, and 4) and different treatments were included as categorical
fixed factors. For all other statistical analyses involving comparison
of mean values (gene expression, ethylene production), the choice of
parametric or non-parametric methods was made after checking the
assumptions of normality (Shapiro–Wilk normality test) and
homogeneity of variances (Fligner–Killeen test) on the raw data. As
parametric methods, Student’s T-test, Welch T-test and ANOVAs
followed by Tukey’s honestly significant difference test were used. As
non-parametric method, Kruskal–Wallis test followed by pairwise
Wilcoxon rank sum test was used. Gene expression data from
qRT-PCR were calculated with the ΔΔCt method (Livak and
Schmittgen, 2001) and were analyzed on log2-transformed data
(parametric test) or on raw data (non-parametric test) specifying
time points or treatments as factors. 3. Results
3.1. Pieris eggs triggered cellular responses
in Brassica spp. 3. Results
3.1. Pieris eggs triggered cellular responses
in Brassica spp. cDNA synthesis kit (Bioline, United Kingdom) according to the
manufacturer’s instructions. Real time qRT-PCR reactions were
performed using SensiFAST SYBR No-ROX Kit (Bioline,
United Kingdom) in 10 μL reaction volumes, containing 3 μL cDNA
and 500 nM of primers on a CFX96 Touch Real-Time PCR
Detection System (Bio-Rad, CA, United States). The following
qRT-PCR program was used: 95°C for 2 min followed by 40 cycles
of 95°C for 5 s; primer-specific annealing temperature for 5 s and
72°C for 10 s, with data collection at 72°C, followed by a melt curve
analysis. Relative gene expression was calculated with the ΔΔCq
method, using GAPDH as reference gene. Primers sequences are
available in Supplementary materials (Supplementary Table S1). We first investigated cellular responses against Pieris brassicae
oviposition in its natural host, the black mustard Brassica nigra. Reactive
oxygen species (ROS) such as superoxide anion (O2
•−) and hydrogen
peroxide (H2O2) accumulated underneath eggs at 24 h (Figure 1A). At
the same time point, aniline blue staining in B. nigra revealed also callose
deposition underneath eggs (Figure 1A). Further, the occurrence of cell
death in plant tissue under the eggs was investigated by staining with
trypan blue (TB), showing that cell death occurred 72 h after oviposition
(Figure 1B). Occasionally, a few TB-stained cells were visible at 48 h (not
shown). In B. nigra, we regularly observed a strong, macroscopically
visible HR-like cell death that spreads beyond the egg site and stops 96 h
after oviposition (Figure 1C). Cellular responses against P. brassicae
oviposition were also investigated in B. rapa. ROS accumulation and cell
death formation were also detected in B. rapa at 24 h and 72 h after
oviposition, respectively (Supplementary Figures S2A,B). Although these
early cellular responses appeared similar to what was observed in
B. nigra, the macroscopically visible HR-like cell death developed by
B. rapa appeared as black necrotic spots that never spread beyond the egg
site (Supplementary Figure S2C). 2.10. Expression of genes by real-time
qRT-PCR We investigated to which extent the difference in HR-like
phenotype between B. nigra accessions WUR-01 and WUR-02 was
related to activation of different phytohormones associated with
plant immunity. Thus, we quantified expression of genes induced
by SA, such as ICS1, PR1, and PR2 (Little et al., 2007), and genes
regulated by JA, i.e., MYC2, VSP1, and VSP2 (Reymond et al.,
2004). For experiments on gene expression induced by eggs or egg
wash in both B. nigra and B. rapa, RNA extraction was performed
according to Oñate-Sánchez and Vicente-Carbajosa (2008). For
experiments on gene expression induced by egg wash in two
B. nigra accessions, RNA extraction was performed with Direct-zol
RNA Miniprep Kit (Zymo Research, CA, United States) following
the manufacturer’s protocol. For preparation of cDNA of all
experiments, 1 μg of RNA was reverse-transcribed using SensiFAST 04 frontiersin.org Caarls et al. 10.3389/fevo.2022.1070859 3.2. Egg wash induced a plant response
similar to the response to eggs Expression at each time point was compared to the 0 h timepoint (Kruskal–Wallis, p < 0.05 for eggs and p < 0.05 for egg wash). (D) Ethylene production in
parts per million (ppm) by plants treated with egg wash. (Student’s T-test, p < 0.001). Each treatment consisted of 10 biological replicates. The height of the
boxes in C-D represents the first to the third quartile of the range; the horizontal line within the box is the median; the whiskers indicate the data minimum
and maximum; and dots represent data points. Asterisk indicates p < 0.05, *** <0.001. A B B A C D D D C FIGURE 2
Plant responses induced by Pieris brassicae eggs and egg wash in Brassica nigra. (A) Picture of responses induced by eggs and egg wash next to each other
on a leaf (microscopic image): both eggs and egg wash induce HR-like necrotic spots. (B) Quantification of severity of symptoms induced by egg wash and
eggs. Shown is the fraction of plants that were scored in a specific class for each treatment (N = 32 plants for both treatments; Kruskal–Wallis, p > 0.05). ns = no significant difference. (C) BnPR1 expression in plants after egg deposition (red box plots) or treatment with egg wash (blue box plots). Gene
expression was measured by qRT-PCR and normalized to the housekeeping gene BnGADPH. Each treatment consisted of 3–4 biological replicates. Expression at each time point was compared to the 0 h timepoint (Kruskal–Wallis, p < 0.05 for eggs and p < 0.05 for egg wash). (D) Ethylene production in
parts per million (ppm) by plants treated with egg wash. (Student’s T-test, p < 0.001). Each treatment consisted of 10 biological replicates. The height of the
boxes in C-D represents the first to the third quartile of the range; the horizontal line within the box is the median; the whiskers indicate the data minimum
and maximum; and dots represent data points. Asterisk indicates p < 0.05, *** <0.001. egg wash compared to incubation with control Mes buffer (Figure 2D;
Supplementary Table S4). In addition, there was a significant difference
in ethylene produced between plants with contrasting HR-like
phenotypes. 3.2. Egg wash induced a plant response
similar to the response to eggs To isolate potential EAMPs and to quickly screen plants for
HR-like cell death, we developed a method to dissolve egg-enveloping
secretions
from
P. brassicae
eggs
(Methods
section;
Supplementary Figure S1). We then compared visually whether the B
C A
B
C
FIGURE 1
Plant immunity responses induced in Brassica nigra by Pieris brassicae eggs (Bn). (A) B. nigra leaf 24 h after oviposition with accumulation of O2
•− (NBT
staining), H2O2 (DAB staining) and callose deposition (aniline blue staining). (B) B. nigra leaf 72 h after oviposition showing cell death (trypan blue staining). (C) Fully developed HR-like cell death that is macroscopically visible at 96 h after oviposition. A few eggs were removed from the clutch to show the cell
death underneath. Stainings were repeated multiple times with similar results. Scale bars = 1 mm. B
C B
C A A B C C FIGURE 1
Plant immunity responses induced in Brassica nigra by Pieris brassicae eggs (Bn). (A) B. nigra leaf 24 h after oviposition with accumulation of O2
•− (NBT
staining), H2O2 (DAB staining) and callose deposition (aniline blue staining). (B) B. nigra leaf 72 h after oviposition showing cell death (trypan blue staining). (C) Fully developed HR-like cell death that is macroscopically visible at 96 h after oviposition. A few eggs were removed from the clutch to show the cell
death underneath. Stainings were repeated multiple times with similar results. Scale bars = 1 mm. 05 Frontiers in Ecology and Evolution frontiersin.org 10.3389/fevo.2022.1070859 Caarls et al. A
B
C
D
FIGURE 2
Plant responses induced by Pieris brassicae eggs and egg wash in Brassica nigra. (A) Picture of responses induced by eggs and egg wash next to each other
on a leaf (microscopic image): both eggs and egg wash induce HR-like necrotic spots. (B) Quantification of severity of symptoms induced by egg wash and
eggs. Shown is the fraction of plants that were scored in a specific class for each treatment (N = 32 plants for both treatments; Kruskal–Wallis, p > 0.05). ns = no significant difference. (C) BnPR1 expression in plants after egg deposition (red box plots) or treatment with egg wash (blue box plots). Gene
expression was measured by qRT-PCR and normalized to the housekeeping gene BnGADPH. Each treatment consisted of 3–4 biological replicates. 3.2. Egg wash induced a plant response
similar to the response to eggs Plants responding with stronger HR-like cell death (score
2 or higher), produced a significantly higher amount of ethylene after
incubation with egg wash than plants with no HR-like cell death
(Student’s
T-test = −4.087,
df = 18,
p < 0.001;
Figure
2D;
Supplementary Table S4). Similarly, B. rapa plants developing HR-like
cell death, also showed a higher ethylene production upon incubation
of leaves with P. brassicae egg wash (Student’s T-test = −3.876, df = 4,
p = 0.018; Supplementary Figure S3E). These results suggest that there is
an early detection response in plants after contact with egg wash that
will ultimately lead to cell death. treatment of leaves with egg wash induced a similar HR-like cell
death as the butterfly eggs (Figure 2A). As a control, leaves were
treated with only the buffer (no eggs were washed). Symptoms
induced by eggs or egg wash were scored after 4 days, and HR-like
frequency (proportion of plants showing HR-like) and HR-like
severity (mean score of induced symptoms) were compared between
the two treatments in B. nigra. HR-like severity after oviposition by
eggs or treatment with egg wash did not differ (Kruskal–Wallis:
χ2 = 1.33, df = 1, p = 0.24; Figure 2B; Supplementary Table S2). We compared expression of SA-marker gene PR1 in B. nigra
oviposited on by P. brassicae butterflies or treated with egg wash. PR1
was significantly upregulated both after oviposition and after treatment
with egg wash. In B. nigra, PR1 expression increased at 6 h and was
significantly upregulated at 24 h and 48 h after egg deposition (Kruskal–
Wallis: χ2 = 12.23, df = 3, p = 0.015; Figure 2C). No significant differences
in PR1 expression were found between B. nigra plants treated with eggs
or egg wash (Figure 2C; Supplementary Table S3). Similarly, in B. rapa,
PR1 expression was significantly upregulated by egg deposition at 24 h
and showed further increase at 96 h (Kruskal–Wallis: χ2 = 11.18, df = 4,
p = 0.024; Supplementary Figure S2D). Frontiers in Ecology and Evolution 3.5. EAMP derived from female accessory
reproductive glands There was no significant difference in eliciting activity of eggs of
increasing age, from one-day-old, to five-day-old eggs, although HR-like
severity decreased slightly. Egg wash was also prepared from eggshells
and secretions that remained on the paper after caterpillars hatched, and
this still induced HR-like symptoms (Kruskal–Wallis: χ2 = 3.20, df = 5,
p > 0.1; Figure 5A; Supplementary Table S6). 3.3. Variation in HR-like cell death severity is
associated with SA-related defenses Asterisks indicate different p-values within the same time point: ** <0.01, *** <0.001, no asterisks indicate no significant
difference. followed by Tukey, p < 0.01), PR1 expression induced by M. brassicae
egg wash was not different from the control treatment (Figure 4C;
Supplementary Table S5). In addition, incubation with M. brassicae
egg wash induced lower ethylene production compared to P. brassicae
egg wash (Kruskal–Wallis: χ2 = 21.36, df = 2, p < 0.001; Figure 4D),
comparable to the ethylene levels previously observed in B. nigra
plants lacking a visible cell death (Figure 2D). Overall, eggs of
M. brassicae did not induce cell death and only a weak ROS and
ethylene response in B. nigra. relative expression of JA-related genes was generally very low and not
significantly different between two accessions. SA-marker BnICS1 was
significantly expressed at higher levels in plants that showed a visible
HR already at 6 h after treatment (Welch’s T-test = −4.901, df = 5.3,
p = 0.003). Both, BnPR1 and BnPR2 showed increased expression
across the time points, although with a different magnitude between
the two accessions. Both genes were already expressed at higher level
at 6 h in plants that showed a visible HR (Welch’s T-test = −2.932,
df = 5.3, p = 0.03 and Welch’s T-test = −4.850, df = 5.8, p = 0.003
respectively), and the difference with plants not expressing HR
increased up to 5-fold (24 h) and 20-fold (48 h) for BnPR1 and 4-fold
(24 h) and 5-fold (48 h) for BnPR2. The expression of JA-related genes
showed more stable profiles across time points, with a small peak at
24 h in all treatments, but with no significant differences between the
two accessions (MYC2, Welch’s T-test = −1.549, df = 5.8 p > 0.05; VSP1,
Welch’s T-test = −0.141, df = 3 p > 0.05; VSP2, Welch’s T-test = −0.301,
df = 3.5 p > 0.05). 3.3. Variation in HR-like cell death severity is
associated with SA-related defenses All SA marker genes were upregulated in B. nigra plants upon egg
wash treatment compared to control. However, the magnitude of
expression of SA-related genes was significantly different between the
two B. nigra accessions at different time points (Figure 3). In contrast, Next, we tested whether egg wash induced ethylene. Brassica nigra
leaves responded with higher ethylene production after incubation with 06 frontiersin.org Caarls et al. 10.3389/fevo.2022.1070859 FIGURE 3
Expression of SA- and JA-related defense genes upon P. brassicae egg wash treatment in two different B. nigra accessions. Gene expression was measured
with two treatments (egg wash, control), two accessions (no HR and HR) and at three time points (6, 24, and 48 h after treatment). HR phenotype of each
plant was scored after sampling for RNA isolation. Gene expression was measured by qPCR-RT and normalized to housekeeping gene BnGADPH. Expression levels were calculated at each time point relative to control treatment (water droplets). Each treatment consisted of 4–5 biological replicates
and is reported as boxplots. The height of the boxes in represents the first to the third quartile of the range; the horizontal line within the box is the median;
the whiskers indicate the data minimum and maximum; and dots represent data points. At each time point, gene expression between the two accessions
was compared (Welch’s T-test). Asterisks indicate different p-values within the same time point: ** <0.01, *** <0.001, no asterisks indicate no significant
difference. FIGURE 3
Expression of SA- and JA-related defense genes upon P. brassicae egg wash treatment in two different B. nigra accessions. Gene expression was measured
with two treatments (egg wash, control), two accessions (no HR and HR) and at three time points (6, 24, and 48 h after treatment). HR phenotype of each
plant was scored after sampling for RNA isolation. Gene expression was measured by qPCR-RT and normalized to housekeeping gene BnGADPH. Expression levels were calculated at each time point relative to control treatment (water droplets). Each treatment consisted of 4–5 biological replicates
and is reported as boxplots. The height of the boxes in represents the first to the third quartile of the range; the horizontal line within the box is the median;
the whiskers indicate the data minimum and maximum; and dots represent data points. At each time point, gene expression between the two accessions
was compared (Welch’s T-test). Frontiers in Ecology and Evolution frontiersin.org 4. Discussion In this study, we show that HR-like inducing EAMPs are in the
egg-enveloping secretions that are produced in female ARGs. Our
results indicate that the EAMP is unlikely of lipidic or proteinaceous
nature but an organic compound of low-molecular weight. When
plants are treated with egg wash, HR-like cell death is induced in
responsive plant genotypes, together with upregulation of
SA-responsive genes and elevated levels of ethylene suggesting that
the HR-like cell death is mediated by one or both phytohormones. In addition, deposition of eggs leads to ROS and callose deposition,
also in plant genotypes that do not show HR-like cell death. All
phenotypes observed were shared between the two Brassica species
that vary in cell death severity and are specific to eggs and egg wash
of the specialist P. brassicae. Finally, both egg wash and wash of glue alone induced a
severe HR-like cell death, and HR-like severity was significantly lower
when B. nigra was treated with a wash of eggs from which the glue was
removed (Kruskal–Wallis: H = 26.60, df = 3, p < 0.001; Figure 5D;
Supplementary Table S6). 3.4. Responses to eggs and egg wash from
Mamestra brassicae are dissimilar to Pieris
brassicae in Brassica nigra Different letters indicate significant differences in mean PR1 expression
(ANOVA followed by Tukey, p < 0.01). (D) Ethylene production in B. nigra leaf in response to egg washes. The height of the boxes in C,D represents the first
to the third quartile of the range; the horizontal line within the box is the median; the whiskers indicate the data minimum and maximum; and dots
represent data points. Values that have no accompanying letters in common differ significantly in production of ethylene (Kruskal–Wallis followed by
pairwise Wilcoxon test, p < 0.001). Scale bars = 1 mm. cell death upon treatment of B. nigra with neither PC16:1/PC16:1 nor
PC18:1/PC18:1 (Figure 7; Supplementary Table S8). A wash from unfertilized, deposited eggs (containing secretions
from the ARG) from virgin butterflies induced a similar response as
wash from fertilized, deposited eggs of mated females (Figure 5C). There was no significant effect of the mating status of the female
(mated or virgin) on the frequency of HR-like cell death elicited or on
HR severity (Kruskal–Wallis: χ2 = 2.61, df = 1, p > 0.1). In addition, the
wash of ARGs from virgin females induced strong symptoms similar
to those of mated females (Figure 5B). These results show that egg
fertilization is not necessary for the induction of the HR-like cell death
in B. nigra, and that an EAMP resides in the ARG, and is
female-derived. 3.4. Responses to eggs and egg wash from
Mamestra brassicae are dissimilar to Pieris
brassicae in Brassica nigra A wash of dissected accessory reproductive glands (ARGs) induced
HR-like cell death, similar to the positive control P. brassicae egg wash
(Figure 5B). On the contrary, neither a wash of mature but unfertilized
eggs dissected from the ovary (“ovarian eggs”) nor a wash of the sperm-
containing bursa copulatrix (BC) induced symptoms (Figure 5B). HR-like severity was significantly higher in plants treated with egg wash
or a wash of ARG, compared to wash of ovarian eggs or bursa copulatrix
(Kruskal–Wallis: χ2 = 19.833, df = 4, p < 0.001; Supplementary Table S6). Staining of leaves showed that plant cells did not die underneath
M. brassicae eggs as leaves did not stain with trypan blue
(Figures 4A,B). Further, M. brassicae eggs induced O2
•− production in
some plants, but weaker than Pieris eggs (Figures 4C,D). While
P. brassicae egg wash induced the expression of PR1 after 24 h (ANOVA 07 frontiersin.org 10.3389/fevo.2022.1070859 Caarls et al. A
B
C
D
FIGURE 4
Responses to eggs and egg wash of M. brassicae in B. nigra. (A) Leaf of B. nigra oviposited on by M. brassicae moth by NBT showing light O2•−
accumulation underneath M. brassicae eggs. (B) Leaf stained by trypan blue showing no cell death underneath M. brassicae eggs. (C) BnPR1 expression in
leaf tissue treated with control solution, P. brassicae wash or M. brassicae wash. Different letters indicate significant differences in mean PR1 expression
(ANOVA followed by Tukey, p < 0.01). (D) Ethylene production in B. nigra leaf in response to egg washes. The height of the boxes in C,D represents the first
to the third quartile of the range; the horizontal line within the box is the median; the whiskers indicate the data minimum and maximum; and dots
represent data points. Values that have no accompanying letters in common differ significantly in production of ethylene (Kruskal–Wallis followed by
pairwise Wilcoxon test, p < 0.001). Scale bars = 1 mm. A B B
D B A C D D D C FIGURE 4
Responses to eggs and egg wash of M. brassicae in B. nigra. (A) Leaf of B. nigra oviposited on by M. brassicae moth by NBT showing light O2•−
accumulation underneath M. brassicae eggs. (B) Leaf stained by trypan blue showing no cell death underneath M. brassicae eggs. (C) BnPR1 expression in
leaf tissue treated with control solution, P. brassicae wash or M. brassicae wash. Frontiers in Ecology and Evolution frontiersin.org 3.6. EAMP is neither a protein nor lipid The increasing knowledge on plant defense mechanisms to eggs
of herbivores suggests that plants can specifically recognize and
respond to egg deposition, presumably via the detection of EAMPs. However, very few EAMPs have been identified, and those that have
so far, are organic compounds of low-molecular weight associated
with the eggs (Reymond, 2013; Hilker and Fatouros, 2015; Stahl
et al., 2018). Only recently a novel type of EAMP, a proteinaceous
compound, diprionin, was found as an elicitor of pine defense None of the egg wash treatments conducted, i.e., freezing and
thawing, proteinase K, or boiling, had an effect on its bioactivity. That
the elicitor does not lose the capability to induce HR in B. nigra leaves
after these treatments, indicates that it is a small, stable, water-soluble
molecule that is likely not a protein (Figure 6; Supplementary Table S7). Lastly, we tested whether two phosphatidylcholines (PC) could also
induce a HR-like responses in B. nigra. We did not observe any visible 08 10.3389/fevo.2022.1070859 Caarls et al. A
B
D
C
FIGURE 5
Proportion of HR-like cell death induced by washes made of eggs, washes made of tissues of the reproductive tract of Pieris brassicae, and washes of eggs
with and without glue, in Brassica nigra. (A–D) (A) Egg washes made of eggs of different ages or eggshells and egg glue remaining on filter paper. N
plants = 13–26 (B) Egg wash of dissected structures of reproductive tract. N plants = 5. (C) Egg wash of eggs of mated females versus virgin females. N
plants = 8–24 (D) Wash of eggs, glue alone (in filter paper) or wash of eggs with egg-enveloping secretions removed. For glue removal, two treatments were
used, either a 1% bleach wash or wash with NaPO4. N plants = 8–16. Values that have no accompanying letters in common differ significantly (Kruskal–
Wallis). A C C A D B B D Proportion of HR-like cell death induced by washes made of eggs, washes made of tissues of the reproductive tract of Pieris brassicae, and washes of eggs
with and without glue, in Brassica nigra. (A–D) (A) Egg washes made of eggs of different ages or eggshells and egg glue remaining on filter paper. N
plants = 13–26 (B) Egg wash of dissected structures of reproductive tract. N plants = 5. (C) Egg wash of eggs of mated females versus virgin females. Frontiers in Ecology and Evolution FIGURE 7 FIGURE 7 FIGURE 7
Phosphatidylcholines (PC) does not induce HR-like cell death in B. nigra plants unlike P. brassicae egg wash. Brassica nigra (SF48-O1)
plants were treated with phosphatidylcholines PC(16:1/16:1) and
PC(18:1/18:1) or with egg wash (EW). PCs were solubilized in 1% DMSO,
0.5% Glycerol and 0.1% Tween and were applied at different
concentrations (1, 5, and 10 μg/μL). A solution of 1% DMSO, 0.5%
Glycerol and 0.1% Tween was used as control (ctl). Egg wash400 was
prepared with 400 eggs/mL, egg wash1000 was prepared with 1,000–
1,200 eggs/1 mL. All treatments were applied on the same leaf, with
two leaves per plant, on 15 biological replicates. Scale bar = 1. FIGURE 7
Phosphatidylcholines (PC) does not induce HR-like cell death in B. nigra plants unlike P. brassicae egg wash. Brassica nigra (SF48-O1)
plants were treated with phosphatidylcholines PC(16:1/16:1) and
PC(18:1/18:1) or with egg wash (EW). PCs were solubilized in 1% DMSO,
0.5% Glycerol and 0.1% Tween and were applied at different
concentrations (1, 5, and 10 μg/μL). A solution of 1% DMSO, 0.5%
Glycerol and 0.1% Tween was used as control (ctl). Egg wash400 was
prepared with 400 eggs/mL, egg wash1000 was prepared with 1,000–
1,200 eggs/1 mL. All treatments were applied on the same leaf, with
two leaves per plant, on 15 biological replicates. Scale bar = 1. Our study reveals that the EAMP associated with P. brassicae eggs likely
belongs to the first category of non-proteinaceous molecules. Neither
boiling nor treatment of the egg wash with proteinase K had an effect on
elicitation of HR-like. Recently, phosphatidylcholines (PCs) were identified
as EAMPs of the P. brassicae egg-induced response in A. thaliana. Active
PCs were mainly containing C16- and C18-fatty acyl chains and treatment
with the latter resulted in SA, H2O2, induction of PR1 and trypan blue
staining (Stahl et al., 2020). PCs are components of cell membranes. Earlier,
phospholipids of Sogatella furcifera Horváth were also found to induce an
ovicidal response in rice (Yang J. O. et al., 2014). Our results suggest that the
EAMPs in P. brassicae eggs that induce HR-like cell death in B. nigra are
other compounds than PCs. First, the B. nigra response is specific to Pieris
eggs and egg wash and is absent in response to other lepidopteran eggs. FIGURE 7 Second, given our method used of washing eggs in a water-like buffer, lipids
are not expected to be present in (high amounts in) the wash. Finally, as PCs
are present in membranes, ovarian eggs should also induce the response. Indeed, ovarian eggs alone induced PR1 expression in A. thaliana (Little
et al., 2007) while we showed that ovarian eggs did not induce HR-like cell
death in B. nigra. In A. thaliana, PR1 expression was also found in response to
crushed egg extracts of different insects (Bruessow et al., 2010; Stahl
et al., 2020). In B. nigra, there was no cell death underneath
M. brassicae eggs, and neither induction of PR1 nor ethylene
production in response to M. brassicae egg wash. Our results suggest
that cell death, ethylene production and gene expression, at least in
B. nigra, are specific to P. brassicae eggs, and we expect the response
to be activated after detection of a Pierinae-specific elicitor. This
hypothesis is strengthened by our previous work which showed
Pierinae-specific induction of PR1 in B. nigra (Griese et al., 2021). It
is possible that a mild cellular defense-like response is activated
against a general insect-derived EAMP, for example PCs, while the
strong HR-like is activated only by EAMPs specifically derived from
Pieris spp. in those plants that can detect these. The production of
ethylene after incubation with egg wash only in plants that show a
strong HR-like cell death (and not in non-HR plants), points to this
effect. Mapping efforts in B. nigra plants, can reveal whether the
genetic variation in plants is for the HR-like cell death and/or for the
detection of a Pieris-specific elicitor. g
When washing eggs, mainly compounds from outside of the eggs
and from egg-enveloping secretions are dissolved. We present
evidence that at least one Pieris-specific EAMP is in these secretions
that envelop the eggs: (i) wash of glue on filter paper is sufficient to
induce the HR-like cell death, (ii) when the secretions are removed
from eggs, and eggs are then washed, HR-like cell death in plants is
diminished, and (iii) a wash of ARGs, the organs where secretions
are produced, is also sufficient to induce HR-like cell death. As the
egg surface and egg-exterior associated secretions are in direct
contact with the leaves, it could be expected that plants evolve to
detect elicitors in egg-enveloping secretions. 3.6. EAMP is neither a protein nor lipid N
plants = 8–24 (D) Wash of eggs, glue alone (in filter paper) or wash of eggs with egg-enveloping secretions removed. For glue removal, two treatments were
used, either a 1% bleach wash or wash with NaPO4. N plants = 8–16. Values that have no accompanying letters in common differ significantly (Kruskal–
Wallis). A
B
FIGURE 6
Proportion of HR-like cell death induced by differently treated egg washes made of Pieris brassicae eggs in Brassica nigra. (A,B) Severity of HR-like
symptoms induced in plants by the different washes. (A) Egg washes that were subjected to increasing amount of freeze–thaw cycles. N plants/
treatment = 24–32. NS (B) Egg washes subjected to boiling (95°C for 30 min) or incubated with proteinase K. N plants/treatment = 9. NS, not significant,
Kruskal–Wallis test. A B B FIGURE 6
Proportion of HR-like cell death induced by differently treated egg washes made of Pieris brassicae eggs in Brassica nigra. (A,B) Severity of HR-like
symptoms induced in plants by the different washes. (A) Egg washes that were subjected to increasing amount of freeze–thaw cycles. N plants/
treatment = 24–32. NS (B) Egg washes subjected to boiling (95°C for 30 min) or incubated with proteinase K. N plants/treatment = 9. NS, not significant,
Kruskal–Wallis test. treated with diprionin they emitted similar quantities of the
sesquiterpene (E)-β-farnesene that is known to attract egg parasitoids
like when laden with sawfly eggs (Hundacker et al., 2021). against sawfly eggs (Hundacker et al., 2021). Diprionin is an annexin-
like protein which is released with the egg-associated secretions of
the sawfly Diprion pini into needles of pines. When pine twigs were Frontiers in Ecology and Evolution
0 09 frontiersin.org 10.3389/fevo.2022.1070859 Caarls et al. FIGURE 7
Phosphatidylcholines (PC) does not induce HR-like cell death in B. nigra plants unlike P. brassicae egg wash. Brassica nigra (SF48-O1)
plants were treated with phosphatidylcholines PC(16:1/16:1) and
PC(18:1/18:1) or with egg wash (EW). PCs were solubilized in 1% DMSO,
0.5% Glycerol and 0.1% Tween and were applied at different
concentrations (1, 5, and 10 μg/μL). A solution of 1% DMSO, 0.5%
Glycerol and 0.1% Tween was used as control (ctl). Egg wash400 was
prepared with 400 eggs/mL, egg wash1000 was prepared with 1,000–
1,200 eggs/1 mL. All treatments were applied on the same leaf, with
two leaves per plant, on 15 biological replicates. Scale bar = 1. and ARGs of virgin P. frontiersin.org Frontiers in Ecology and Evolution 3.6. EAMP is neither a protein nor lipid brassicae butterflies induced HR-like cell death
in an equal manner as mated butterflies. We thus hypothesize that at
least one HR-inducing EAMP is present in the egg-enveloping
secretions and ARGs and is female-derived. Chemical analysis of egg
wash and glands is currently carried out to identify this EAMP. g
y
y
Plant responses to insect eggs are similar to responses to
(microbial) pathogens, and include SA and ROS accumulation,
callose deposition, defense gene expression and cell death
(Reymond, 2013). For example, callose deposition was shown to
be associated with lesions following pathogen invasion or
autoimmune responses (Koga et al., 1988; Dietrich et al., 1994). We observed the deposition of callose in B. nigra associated with
lesions induced by P. brassicae eggs. Natural variation in strength of
egg-induced cell death was found in several brassicaceous species,
including B. nigra (Pashalidou et al., 2015; Griese et al., 2021; Groux
et al., 2021). Here, we show that plants that do not express a strong
HR-like cell death, still responded with ROS accumulation and cell
death as showed by trypan blue staining. Similarly, in S. dulcamara,
variation exists for egg-induced chlorosis, and a genotype that did
not respond with chlorosis and on which egg hatching rate of
S. exigua was not reduced, still accumulated SA after oviposition
(Geuss et al., 2017). We thus hypothesize that Pieris eggs generally
induce an immune response in all plants of B. rapa and B. nigra, that
is only in some plants accompanied by a stronger cell death
response. In pathogen-induced HR, cell death can often
be uncoupled from (preceding) biochemical and molecular changes
and the two processes can be genetically dissected (Künstler et al.,
2016). In that case, cell death is dispensable for resistance. However,
in egg-induced HR, previous studies show that the stronger the
HR-like cell death, the higher egg mortality (Fatouros et al., 2014;
Griese et al., 2017, 2020, 2021). References Erb, M., and Reymond, P. (2019). Molecular interactions between plants and insect herbivores. Annu. Rev. Plant Biol. 70, 527–557. doi: 10.1146/annurev-arplant-050718-095910 Balint-Kurti, P. (2019). The plant hypersensitive response: concepts, control and
consequences. Mol. Plant Pathol. 20, 1163–1178. doi: 10.1111/mpp.12821 Fan, L., Chae, E., Gust, A. A., and Nürnberger, T. (2017). Isolation of novel MAMP-like
activities and identification of cognate pattern recognition receptors in Arabidopsis
thaliana using next-generation sequencing (NGS)–based mapping. Current Protocols in
Plant Biology 2, 173–189. doi: 10.1002/cppb.20056 Bassetti, N., Caarls, L., Bukovinszkine’kiss, G., El-Soda, M., Van Veen, J.,
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pnas.0707809105 Bruessow, F., Gouhier-Darimont, C., Buchala, A., Metraux, J.-P., and Reymond, P. (2010). Insect eggs suppress plant defence against chewing herbivores. Plant J. 62, 876–885. doi:
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insects of different trophic levels. PLoS One 7:e43607. doi: 10.1371/journal.pone.0043607 Campos, M. L., De Souza, C. M., De Oliveira, K. B. S., Dias, S. C., and Franco, O. L. (2018). The role of antimicrobial peptides in plant immunity. J. Exp. Bot. 69, 4997–5011. doi: 10.1093/jxb/ery294 Fatouros, N. E., Paniagua Voirol, L. R., Drizou, F., Doan, Q. T., Pineda, A., Frago, E., et al. (2015). Role of large cabbage white butterfly male-derived compounds in elicitation of
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arplant-050213-040012 Fatouros, N. E., Pashalidou, F. FIGURE 7 It resembles the natural
situation of leaf-egg interaction, more so than, for example, crushing
of eggs (Little et al., 2007; Bruessow et al., 2010) or crushing of adults
(Doss et al., 2000; Yang Y. et al., 2014). Previously, a male-derived
anti-aphrodisiac compound transferred during mating to the female
ARG, benzyl cyanide, was suggested as potential elicitor (Fatouros
et al., 2008). We find no evidence for a male-derived elicitor: eggs Molecular patterns that are detected by plants are thought to
be structurally conserved molecules (Van der Burgh and Joosten, 2019). The presence of EAMPs in the glue suggests that their function could
be a structural component of the glue, or a compound with an essential
function to the fertilized eggs. Many proteins are found in glue of insect
eggs (Li et al., 2008), and egg glue of P. brassicae was described to consist
of proteins and unsaturated lipoids (Beament and Lal, 1957). In
addition, the ARG secretions could contain molecules that are produced
by the parents and/or microbial symbionts to protect the vulnerable egg, Frontiers in Ecology and Evolution 10 frontiersin.org Caarls et al. 10.3389/fevo.2022.1070859 Supplementary material This research was made possible by support of the Dutch Technology
Foundation TTW, which is part of Netherlands Organisation for
Scientific Research (NWO) (NWO/TTW VIDI grant 14854 to N.E.F.). The Supplementary material for this article can be found at:
https://www.frontiersin.org/articles/10.3389/fevo.2022.1070859/full#
Supplementary-material Acknowledgments for example compounds with antimicrobial activity (Flórez et al., 2018). Further identification of EAMPs can lead to new research in
this direction. We are grateful to the employees of Unifarm (WUR) for rearing and
caring of the plants used in the experiment. We thank Pieter Rouweler,
André Gidding, and the late Frans van Aggelen for rearing of Pieris
brassicae and Mamestra brassicae. We thank Martijn Flipsen, Gabriella
Bukovinszkine’Kiss, Klaas Bouwmeester, Femke van Doesburg, Ewan
van Eijden and Arjen van der Peppel for assistance with some of the
experiments. Guusje Bonnema and Erik Poelman are acknowledged for
seeds of B. rapa and B. nigra, respectively. In summary, we present a method to obtain EAMPs from insect
eggs, and using this method, show that the EAMP inducing the HR-like
cell death response is in the egg glue, derived from the female
ARG. We furthermore assess the chemical nature and specificity of
these elicitors and the molecular response of Brassica plants to Pieris
and other eggs. The obtained knowledge paves the way for future
studies on identification of EAMPs in Pieris egg glue, and the
corresponding receptor genes in Brassica plants. Data availability statement The authors declare that the research was conducted in the absence
of any commercial or financial relationships that could be construed as
a potential conflict of interest. The datasets presented in this study can be found in the data
repository Zenodo at doi.org/10.5281/zenodo.7194716. Publisher’s note LC, NB, JL, RM, and NF planned and designed the research. LC,
NB, and PV performed the experiments and/or analyzed the data. LC,
NB, and NF wrote the manuscript with contributions by PV, RM, JL, and
MS. All authors contributed to the article and approved the
submitted version. All claims expressed in this article are solely those of the authors and
do not necessarily represent those of their affiliated organizations, or
those of the publisher, the editors and the reviewers. Any product that
may be evaluated in this article, or claim that may be made by its
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A systematic approach for risk assessment and mitigation of wind integrated electricity markets
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*Corresponding author: naresh.2021ree07@mnnit.ac.in E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 A systematic approach for risk assessment
and mitigation of wind integrated electricity
markets Naresh Boda1,*, Prashant Kumar Tiwari2, and Rajesh Panda3
1,2Motilal Nehru National Institute of Technology Allahabad, UP, India, 211014
3SSN College of Engineering Chennai, India Abstract. A risk-based double auction mechanism with dispatchable loads
is one of the most cost-efficient trading mechanisms in the modern real-time
electricity markets. Risks in the system due to various uncertainties, such as
line failure, generator failure, load changes and other severe contingencies
causes non-systematic execution of real-time electricity markets. This paper
presents a simple and systematic procedure for reducing system risks in
deregulated power markets considering wind power integrations. For
maximizing the considerable system social welfare and minimizing the
system's economic risk, the optimal placement and operation of wind power
producers (WPP) have been suggested. The critical buses have been
identified by accessing the different assessment tools e.g., Value at Risk
(VaR), Conditional Value at Risk (CVaR), and locational marginal pricing
(LMP) based Marginal Pricing Index (MPI). The presented approach has
been evaluated and validated on modified IEEE 30-bus system. Keywords: Risk Assessment, Locational Marginal Price, Wind Power
Producers, Real Time Electricity Market. Keywords: Risk Assessment, Locational Marginal Price, Wind Power
Producers, Real Time Electricity Market. © The Authors, published by EDP Sciences. This is an open access article distributed under the terms of the Creative Commons
Attribution License 4.0 (https://creativecommons.org/licenses/by/4.0/). E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 In this context, a strategic bidding model is used for different contingencies and
uncertainties with reference to WPP, the market price, demand bids and competitor offers
[3]. Determine the best offer price for WPP by using multistage risk restrained stochastic
interrelated model is used in electricity market[4]. The stability and security of the
restructured power system can be improved by reducing the system risk and to assess the risk
using risk hedging tools are proposed[5]. To trade the future value of the wind power the
WPP may take a part in the prediction market and by accepting a different stance from the
electricity market, the imbalance costs will be compensated for by rewards in the prediction
market [6]. The energy output of wind power producers could be increased, and the cost of producing
energy might be decreased, with the use of wind power bidding techniques. When wind
power penetration is minimal, it is fair to view WPP as price-takers, as stated in [7]. The
producers of wind energy are strategic participants in DA and RT markets, and impact of
WPP penetration on energy prices cannot be ignored. By using the EPEC method, market
equilibrium was discovered, which maximized the sum of expected profits from all producers
[8]. While maximize the profit of the DA and ancillary services a strategic bidding is used to
reduce the generation cost, and improving the revenue of GenCos & DisCos in electricity
market [9]. In order to increase the profit, a bidding technique is proposed for WPP and
energy storage systems in DA and Real-time electricity markets [10–11]. The proper risk
measure approach is suggested to find the best scheduling for various risk levels [12–13]. At
GenCo’s bids are generated by using best bidding strategy, profit of the system is increased
while reducing risk [14–15]. The risk is considered an impediment to the bid that would
increase the profit while reducing the uneven prices of the optimal placement of WPP in spot
market. A risk-averse trading mechanism is proposed and assessed against several risk
indicators, and an investigation of systemic risk of the power market is assessed by using two
indices, i.e., contagious VaR (CoVaR) and marginal contagious VaR (ΔCoVaR) [16]. By considering uncertainty and coupling of risk in restructured power system, the accuracy
of risk evaluation to be increased while reducing financial losses by applying a proper
hierarchical risk evaluation method[17]. 1 Introduction Renewable energy becomes optimistic in a deregulated power system in response to rising
environmental perturbations. The most popular renewable energy source today is wind
power, which is a great addition to the current energy grid. Because of its benefits, including
quick building timelines, cost competitiveness, and environmental consequences, wind
power has emerged as a strong contender in long-term system planning. As a result, the
world's installed wind power capacity has rapidly increased, with China, the United States,
Brazil, Germany, and Sweden leading the way. Currently, there has been 906 GW of installed
wind capacity worldwide. This reflects 9% year-over-year growth. The addition of 78 GW
globally in 2022 made it the third-biggest year ever for new capacity [1]. In India, wind
capacity increased two times from 21 GW to 42.6 GW since 2014, and India is globally in
the fourth position in renewable energy [2]. Looking towards the increased wind potential in the power system networks, there are
several research findings have been reported by the researchers in the past to handle the
integration issues, intermittency etc. related to wind power in conventional and competitive
power markets. https://doi.org/10.1051/e3sconf/202447201010 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 2.1 Electricity Market Mechanism A market is a system that facilitates the exchange of related goods and services for economic
benefit. Sellers and buyers are getting profit in an auction market. This is defined as a place
to trade with the auctioneers, and it is classified into two types, i.e., single-sided, and double-
sided auction markets. New uncertainties are introduced as a result of the deregulation of the power sector. Recently,
deregulation has given generation firms (GenCo’s) and distribution entities (DisCo’s) a
platform to investigate price potential in the universally competitive electricity markets. In
power system deregulation, DA and RT are the two notable time periods for market
settlement. The RT market utilization is calculated according to real time balance of supply
and demand. Market clearance occurs in the DA market before 24 hours of the following
trading day, where offers and bids are decided on one DA. Sellers in electrical market
typically enter the DA for the following day’s trade to submit offers to the system operator. In double auction market both buyers and sellers submit their offers and bids to the ISO in
real time[27]. To avoid market power during the multi-round auction procedure, both
generation company and distribution company transmit offers and bids to the ISO at the same
time, and ISO clear the market based on MCP. The MCP is determine the offers price and
demand bids are interact at one point as submitted to the ISO. using specific protocols
selected by the ISO based on their transmission grid restrictions and pricing methods. Uniform and discriminatory are the two different ways pricing mechanisms are classified. Seller and buyer pairs are matched individually in a discriminatory pricing scheme, and every
time in a double auction market, buyer and seller are matched equally in a uniform pricing
scheme. The bid and offer price-quantity are iteratively connected through buyers and sellers in a
double auction model to be submitted to the ISO. The MCP is the interaction between the bid
curve and offer curve and the nodal price given by the LMP at each node of the deregulated
power system [28]. LMP can be maximized at the voltage controlled node by maximizing
consumer excess, while LMP can be reduced at P-Q bus by reducing produced excess. Flexible loads and fixed loads are the two load models in a power system. Under uncertain
conditions in the power system, fixed loads do not change. E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 The primary contribution of this article is to assess the risk associated with the
dispatchable loads are used in double auction strategy under various uncertainties, including
line fault, generator fault, and load changing in the RT market using various risk assessment
tools. By locating wind power producers, the risk-averse solution is attained, and the GenCo’s
profits can be maximized in RT market. The remainder of this article is structured as follows: Section II initially describes the
overview of the methodology. Section III indicates problem formulation of presented
strategy. Numerical results and discussion of presented strategy describes in Section IV. Finally, Conclusion of the article describe in Section V. E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 A bidding strategy to minimize projected cost under
risk constraint optimum rigid and malleable bids in Day ahead and Real time markets are
evaluated in [18]. By considering sensitivity factor, the system profit increase while reducing
the generation cost of an electricity market[19]. A regular scheduled market auction bidding
method has been used to increase the profit of the GenCo by setting bid price as payas-bid
[20-21]. To improve prediction models and represent various preferences on WPP models of
a real-world system to propose a risk assessment based adaptive USTWPP frame work is
evaluated[22]. While considering the uncertainty surrounding renewable generation, real-
time price, and power consumption method ensures the robustness of the day-ahead (DA)
bidding strategy are used to reduce predicted regret value over a group of worst-case
situations whose combined probability is no greater than a certain threshold[23]. For optimal
placement of WPP in DA and RT electricity market by using an optimum bidding strategy
of various market players are considered[24]. Find optimum strategy to place WPP in day
ahead and balancing market by using a multi-stage risk-constrained stochastic approach is
proposed in [25]. A Systemic risk investigation of future electricity market to propose a
theoretical model for systemic analysis of the power market[26]. 2 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 2.2 Risk Assessment The existence or severity of system risks in the evaluated networks in accordance with
LMP is investigated using the concept of risk evaluation techniques (i.e., VaR and CVaR). Fig.1. shows that VaR & CVaR are risk evaluation indices, with VaR designating a small
loss of (1-α) the loss distribution of the system [30]. loss of (1 α) the loss distribution of the system [30]. Fig.1. Illustration of VaR and CVaR[29] Fig.1. Illustration of VaR and CVaR[29] 2.1 Electricity Market Mechanism A relatively easy technique is to
model price-sensitive loads accompanying negative charges with uncooperative real power 3 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 injections. The negative output of the generator is described with injection levels, ranging
from zero to highest negative conceivable loads[29]. injections. The negative output of the generator is described with injection levels, ranging
from zero to highest negative conceivable loads[29]. 2.3 Marginal Pricing Index(MPI) The presented strategic index is used to determine which nodes are most price-sensitive
after line interruptions and other contingency situations. The level of risk in the system is
identified by using price sensitive buses.
,
b a se
A O
i
M a x A O
a b s
M P I
(1) (1) Where is MPIi at bus i. λBas and λAO base case LMP values at bus ‘i’ after line interruptions. After various contingency conditions, λMax,AO has the highest LMP value among all buses in
the system [31]. 2.4 Proposed Strategy To find the optimal placement of WPP in deregulated power system by using a strategic risk
assessment of the system and to assess the risk by using risk indices i.e., VaR and CVaR and
to find most critical bus by using marginal pricing index(MPI). The application of these risk
assessment methods and MPI has greatly reduced the area of search for the optimal spot, and
it has been demonstrated that placement locations are important for managing economic risk. 4 4 4 https://doi.org/10.1051/e3sconf/202447201010 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 Fig.2. Flow Chart of presented strategy Fig.2. Flow Chart of presented strategy 3 Problem Formulation A method consists of a risk-constrained double auction strategy in RT electricity market
under deregulated power system has been presented that evaluate cost minimization, profit
maximization, and social welfare maximization for each GenCo’s and DisCo’s with ideal
placement of WPP. Mathematically, objective function is defined as below: Objective Function-I Objective Function-I
2
1
2
1
gi
dj
j
dj
i
gi
ng
i
gi
i
i
gi
i
i
nd
j
dj
j
j
dj
j
j
MaxF
B
P
C
P
C
P
a P
b P
c
B
P
a P
b P
c
(2) j
2
1
2
1
gi
dj
j
dj
i
gi
ng
i
gi
i
i
gi
i
i
nd
j
dj
j
j
dj
j
j
MaxF
B
P
C
P
C
P
a P
b P
c
B
P
a P
b P
c
(2) Here, Ci(Pgi) is Power system generation cost and Bj(Pdj) is the profit of the pool users to
pay from demand. Social welfare will be increased while reducing the generation costs of the
system. Here, Ci(Pgi) is Power system generation cost and Bj(Pdj) is the profit of the pool users to
pay from demand. Social welfare will be increased while reducing the generation costs of the
system. 5 5 https://doi.org/10.1051/e3sconf/202447201010 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 Objective Function-II Risk of the system measured with the risk evaluation tools i.e., VaR and CVaR, with VaR
designating a small loss of (1- α) the loss distribution of the system for a certain level [0,1],
and VaR is less successful than CVaR in a scenario of continuous loss distribution. To
conclude that CVaR is the average loss at the lower back end of the loss dispensation.
(
,
)
,
min
:
,
g A Y
a
A
x Y dY
A
R
A
(3)
1
1
1
Ns
s
s
s
MinimizeCVaR
(4)
(
,
)
,
min
:
,
g A Y
a
A
x Y dY
A
R
A
(3)
1
1
1
Ns
s
s
s
MinimizeCVaR
(4)
subjected to
,
1,2,3,....,
S
s
EP
s
Ns
(5)
0,
1, 2,3,....,
s
s
Ns
(6) (3) (4) subjected to j
,
1,2,3,....,
S
s
EP
s
Ns
(5)
0,
1, 2,3,....,
s
s
Ns
(6) ,
1,2,3,....,
S
s
EP
s
Ns
0,
1, 2,3,....,
s
s
Ns
(6) Here, β indicate the aide positive coefficient and distinguish between VaR(ξ) and scenario
payoff. Equations (3) and (4) show the equations for VaR and CVaR, respectively. The
optimal configuration of wind power producers (WPP) is used to fulfil the objective function
concurrently. Here, β indicate the aide positive coefficient and distinguish between VaR(ξ) and scenario
payoff. Equations (3) and (4) show the equations for VaR and CVaR, respectively. The
optimal configuration of wind power producers (WPP) is used to fulfil the objective function
concurrently. In this article, VaR, CVaR, and MPI are the risk assessment tools. Fig. 2 shows an example
of VaR and CVaR with profit maximization. When the values of VaR and CVaR are negative
maximums on left side of the chart tail end, then the loss is increased and profit of the system
is reduced, respectively. Equality Constraints: 1
g
N
g
W
d
Loss
a
P
P
P
P
(11)
2
2
1
2
cos
d
N
Loss
j
i
j
i
j
i
j
a
P
G
V
V
V V
(12)
1
cos
B
N
i
i
k ik
ik
i
k
k
P
VV Y
(13)
1
sin
B
N
i
i
k
ik
ik
i
k
k
Q
VV Y
(14) (12) (14) From equations (11) and (12), power balance equations are power from wind, Ploss is
transmission loss, and Gj is conductance between node a and k. Vi and Vj are the magnitudes
of voltages at node i and j. Equation (13) and (14), shows that active and reactive power of
the system. Fig.1 illustrates that, for the negative greatest value of VaR and CVaR, the system loss is
high and the profit of the system is decreased, while for the positive greatest value of VaR &
CVaR, the system loss decreases and the profit of the system increases at the right-back end
of the diagram. At higher system-risk conditions, maintaining the stability and security of the
system required some further arrangements. So, at most severe risk conditions, the cost of
generation is improved in addition to the actual cost of production and ancillary costs. According to the above explanation, it is clear that the system's risk has an inverse
relationship with VaR and CVaR. It means that the system's risk will be higher for the lower
(highest negative) values of VaR and CVaR. So, it is essential to reduce the risk of the system
and shift the graph (Fig.1.) from the left back-end to the right back-end, i.e., the value of VaR
and CVaR is at its low value in the positive direction. The main aim of the presented strategy
is to reduce the generation cost of the system. Consequentially, the values of VaR and CVaR
have an inverse relation to the objective function (system generation cost) of the presented
strategy. Therefore, the generation cost of the system decreases while increasing the value of
social welfare. So, VaR and CVaR are proportional to the social welfare of the system. Objective Function-II At the tail end of the histogram, the values of generation cost and
social welfare are minimized and maximized, for the maximum(positive) value of VaR and
CVaR. To maintain the system's stability and security, some additional arrangements are required
for higher system risks. The actual generation cost increases with higher risk and additional
costs. Therefore, in a higher-risk scenario, social welfare is minimized. Constraints: Ensuring equality & inequality constraints are contemplate while resolving OPF
issues. Inequality Constraints:
min
max
i
i
i
V
V
V
1,2,3,.....,
B
i
N
(7)
min
max
i
i
i
1,2,3,.....,
B
i
N
(8)
min
max
gi
gi
gi
P
P
P
1,2,3,.....,
B
i
N
(9)
min
max
gi
gi
gi
Q
Q
Q
1,2,3,.....,
B
i
N
(10) Inequality Constraints:
min
max
i
i
i
V
V
V
1,2,3,.....,
B
i
N
(7)
min
max
i
i
i
1,2,3,.....,
B
i
N
(8)
min
max
gi
gi
gi
P
P
P
1,2,3,.....,
B
i
N
(9)
min
max
gi
gi
gi
Q
Q
Q
1,2,3,.....,
B
i
N
(10) (10) (10) Here
min
iV
and
max
iV
lower limit and upper limit of the voltage at node i. min
i
and
max
i
are
lower and upper limits of phase angle at node i. min
gi
P
and
min
gi
Q
are lower limit of real and 6 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 reactive power at node i. max
gi
P
, and
max
gi
Q
upper limit of real and reactive power at node i. Pd
& Ploss are the power demand and power loss of the system. reactive power at node i. max
gi
P
, and
max
gi
Q
upper limit of real and reactive power at node i. Pd
& Ploss are the power demand and power loss of the system. 4 Simulation Results and Discussion The customized IEEE 30-bus data system has been evaluated to demonstrate the proposed
strategy. The customized data test system contains 41 transmission lines, 30 node, 6 thermal
generators, and four dispatchable loads. A complete proposed strategy has been developed in
MATPOWER software [29], and MILP based AC optimal power flow has been employed
for optimization. Three separate case studies on the system were studied in the entire execution and evaluation
of the presented strategy in three cases as shown below. 7 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 1. Generate Scenarios by considering different contingencies and study the VaR, CVaR
by using scenarios based LMP and calculated most critical buses by using marginal
pricing index(MPI). 1. Generate Scenarios by considering different contingencies and study the VaR, CVaR
by using scenarios based LMP and calculated most critical buses by using marginal
pricing index(MPI). p
g
(
)
2. Finest of the most endangered buses for placement of wind power producers(WPP). 2. Finest of the most endangered buses for placement of wind power producers(WPP). 3. After the placement of WPP, to calculate the profit of the system before and after
placement of WPP. 3. After the placement of WPP, to calculate the profit of the system before and after
placement of WPP. Case 1: Given this, 42 different contingency scenarios have been created based on generator
outages, line outages, and changes in loads. To determine the value of VaR & CVaR based
on LMP of the system at different contingency conditions are used to determine the risk level
of the selected scenarios. Fig. 3 represents risk(VaR and CVaR) according to confidence level
95%. From Fig. 3, The VaR and CVaR that the system is inversely correlated to risk of the system. So, social welfare of the deregulated system is reduced when the VaR and CVaR values are
maximised, and vice versa. maximised, and vice versa. (a)
(b)
Fig.3. a) VaR and Social welfare ($/hr) at 95% before WPP and
b) CVaR and Social welfare($/hr) at 95 % before WPP
To find the riskiest nodes in the deregulated power system by using marginal pricing
index(MPI) for optimal placement of the WPP. The calculation of MPI is based on LMP of
the system. The values of MPI are subsequently organised into descending order to create a
list of things to prioritise (Table 1 illustrates this). 4 Simulation Results and Discussion The system's most sensitive bus is
represented by MPI's highest value. For the installation of wind farms, the top 6 buses (aside
from generator buses) with the highest MPI values (bus numbers 29, 30, 27, 26, and 25) have
been selected. 500
550
600
-0.25
-0.15
-0.05
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welfare ($/hr)
VaR
Scenario's
VaR at 95%
Social Welfare
500
550
600
650
-0.40
-0.20
0.00
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welafare ($/hr)
CVaR
Scenario's
CVaR at 95%
Social Welfare (a)
(b)
Fig.3. a) VaR and Social welfare ($/hr) at 95% before WPP and
b) CVaR and Social welfare($/hr) at 95 % before WPP
500
550
600
-0.25
-0.15
-0.05
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welfare ($/hr)
VaR
Scenario's
VaR at 95%
Social Welfare
500
550
600
650
-0.40
-0.20
0.00
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welafare ($/hr)
CVaR
Scenario's
CVaR at 95%
Social Welfare (b)
Fig.3. a) VaR and Social welfare ($/hr) at 95% before WPP and
b) CVaR and Social welfare($/hr) at 95 % before WPP
500
550
600
650
-0.40
-0.20
0.00
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welafare ($/hr)
CVaR
Scenario's
CVaR at 95%
Social Welfare ( )
Fig.3. a) VaR and Social welfare ($/hr) at 95% before WPP and
b) CVaR and Social welfare($/hr) at 95 % before WPP To find the riskiest nodes in the deregulated power system by using marginal pricing
index(MPI) for optimal placement of the WPP. The calculation of MPI is based on LMP of
the system. The values of MPI are subsequently organised into descending order to create a
list of things to prioritise (Table 1 illustrates this). The system's most sensitive bus is
represented by MPI's highest value. For the installation of wind farms, the top 6 buses (aside
from generator buses) with the highest MPI values (bus numbers 29, 30, 27, 26, and 25) have
been selected. 4 Simulation Results and Discussion To find the riskiest nodes in the deregulated power system by using marginal pricing
index(MPI) for optimal placement of the WPP. The calculation of MPI is based on LMP of
the system. The values of MPI are subsequently organised into descending order to create a
list of things to prioritise (Table 1 illustrates this). The system's most sensitive bus is
represented by MPI's highest value. For the installation of wind farms, the top 6 buses (aside
from generator buses) with the highest MPI values (bus numbers 29, 30, 27, 26, and 25) have
been selected. 8 8 8 https://doi.org/10.1051/e3sconf/202447201010 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 Table 1: shows the MPI-based priority list. Bus
No. Locational Marging Pricing
Marginal
Pricing
Index(MPI)
Rank
Base
Case
(λbase)
After Line
ouatge
(λAO)
Difference
(λbase-λAO)
29
4.075
8.573
4.498
0.52467
1
30
4.155
7.174
3.019
0.35215
2
27
3.960
4.456
0.496
0.05786
3
26
4.064
4.518
0.454
0.05296
4
25
3.988
4.386
0.398
0.04642
5
24
3.981
4.144
0.163
0.01901
6
23
3.994
4.094
0.100
0.01166
7
28
3.939
4.045
0.106
0.01236
8
22
3.966
4.041
0.075
0.00875
9
21
3.970
4.042
0.072
0.00840
10
19
4.029
4.101
0.072
0.00840
12
20
4.008
4.075
0.067
0.00782
11 Table 1: shows the MPI-based priority list. Case 2: From Fig.4&5, VaR and CVaR at each bus of the system shows before and after
placement of WPP in existing system at different confidence levels i.e., 95% and
99%.According to the below shown figure before WPP, the most critical buses are 30, 29,
18, 20, and 25. Compared to 18, 20, and 25, the critical buses are 30 and 29. The optimal placement of wind power producers is between 30 and 29 buses. After the
placement of WPP, the risk of the system is reduced due to the reduced generation cost and
increase in social welfare. (a)
-0.60
-0.30
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
VaR
Bus No. VaR at 99% before WPP
VaR at 99% after WPP (a)
-0.60
-0.30
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
VaR
Bus No. VaR at 99% before WPP
VaR at 99% after WPP (a) 9 9 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 (b)
Fig.4. 4 Simulation Results and Discussion VaR at each bus of the system before and after placement of WPP
at 95% confidence level b) at 99% confidence level
(a)
(b)
Fig.5. VaR at each bus represents before and after placement of WPP
a)
at 95% confidence level b) at 99% confidence level
ase 3: The social welfare of every seller and buyer under various scenarios is computed
fore and after the placement of wind power producers. The system's social welfare with
d without wind energy producers as compared to 42 different scenarios, as illustrated in
g. 6. -0.40
-0.20
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
VaR
Bus No. VaR at 95% before WPP
VaR at 95% after WPP
-1.60
-0.80
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
CVaR
Bus No. CVaR at 99% before WPP
CVaR at 99% after WPP
-0.50
-0.25
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
CVaR
Bus No. CVaR at 95% before WPP
CVaR at 95% after WPP
SD 2023 (b)
Fig.4. VaR at each bus of the system before and after placement of WPP
95%
fid
l
l b)
99%
fid
l
l
-0.40
-0.20
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
VaR
Bus No. VaR at 95% before WPP
VaR at 95% after WPP (b)
Fig.4. VaR at each bus of the system before and after placement of WPP
at 95% confidence level b) at 99% confidence level (b)
ig.4. VaR at each bus of the system before and after placement of WPP
at 95% confidence level b) at 99% confidence level (b)
Fig.5. VaR at each bus represents before and after placement of WPP
a)
at 95% confidence level b) at 99% confidence level
-0.50
-0.25
0.00
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
CVaR
Bus No. CVaR at 95% before WPP
CVaR at 95% after WPP Case 3: The social welfare of every seller and buyer under various scenarios is computed
before and after the placement of wind power producers. The system's social welfare with
and without wind energy producers as compared to 42 different scenarios, as illustrated in
Fig. 6. 5 Conclusion Presented strategy is effective for minimizing system financial risks according to the LMP
of the system and recognizing the riskiest bus for optimal placement of WPP in centralized
double-auctioned restructured power system. A MPI also used to evolve and find out the
more sensitive nodes with the help of LMP at different contingencies in deregulated system. To minimize economic risk, priority based MPI and risk assessment tools are used to position
wind generators in a deregulated system. The test results illustrate the success of the
recommended strategy with reference to risk mitigation, system social welfare enhancement,
and total generation cost reduction. As a result, the suggested approach is demonstrating its robustness in terms of reducing
economic risk in competitive power markets. This generalised technique can be relating any
small-scale, medium-scale, and large scale integrated system. 4 Simulation Results and Discussion Case 3: The social welfare of every seller and buyer under various scenarios is computed
before and after the placement of wind power producers. The system's social welfare with
and without wind energy producers as compared to 42 different scenarios, as illustrated in
Fig. 6. 10 E3S Web of Conferences 472, 01010 (2024)
ICREGCSD 2023 https://doi.org/10.1051/e3sconf/202447201010 Fig.6. Represent Social Welfare of the system afore and later placement of WPP
0
400
800
1
3
5
7
9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welfare ($/hr)
Scenario's
Social Welfare Before WPP
Social Welfare after WPP Fig.6. Represent Social Welfare of the system afore and later placement of WPP
0
400
800
1
3
5
7
9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
Social Welfare ($/hr)
Scenario's
Social Welfare Before WPP
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https://genomebiology.biomedcentral.com/counter/pdf/10.1186/s13059-022-02692-0
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English
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De novo reconstruction of cell interaction landscapes from single-cell spatial transcriptome data with DeepLinc
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Genome biology
| 2,022
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cc-by
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© The Author(s). 2022 Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to
the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The
images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise
in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright
holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. The Creative Commons Public Domain
Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless
otherwise stated in a credit line to the data. METHOD Open Access * Correspondence: yangxuerui@
tsinghua.edu.cn * Correspondence: yangxuerui@
tsinghua.edu.cn Abstract Based on a deep generative model of variational graph autoencoder (VGAE), we
develop a new method, DeepLinc (deep learning framework for Landscapes of
Interacting Cells), for the de novo reconstruction of cell interaction networks from
single-cell spatial transcriptomic data. DeepLinc demonstrates high efficiency in
learning from imperfect and incomplete spatial transcriptome data, filtering false
interactions, and imputing missing distal and proximal interactions. The latent
representations learned by DeepLinc are also used for inferring the signature genes
contributing to the cell interaction landscapes, and for reclustering the cells based
on the spatially coded cell heterogeneity in complex tissues at single-cell resolution. MOE Key Laboratory of
Bioinformatics, Center for Synthetic
& Systems Biology, School of Life
Sciences, Tsinghua University,
Beijing 100084, China Keywords: Single-cell spatial transcriptome, Cell interaction, Variational graph
autoencoder, VGAE, Deep learning De novo reconstruction of cell interaction
landscapes from single-cell spatial
transcriptome data with DeepLinc Runze Li and Xuerui Yang* Background The physiological functions of multicellular tissues are not only defined by heteroge-
neous cells forming these tissues but are also highly dependent on complicated local
and distal cell-cell interactions [1, 2]. Furthermore, it is increasingly recognized that
many of the intracellular activities of each single cell are closely related to its interac-
tions with the multicellular context [3]. The intrinsic gene expression profile of each
single cell is both a consequence and a defining factor of the complicated cell inter-
action network in physiological contexts [4, 5]. Recent advances in various spatially resolved transcriptome profiling techniques have
made it possible to measure gene expression profiles at single-cell or subcellular reso-
lution while simultaneously retaining information on the spatial locations of cells. These techniques include in situ sequencing methods, such as FISSEQ [6] and STAR-
map [7]; imaging methods based on fluorescence in situ hybridization (FISH), such as
MERFISH [8], seqFISH [9], SPOTs [10], and osmFISH [11]; spatial barcoding tech-
niques, such as Slide-seq [12], HDST [13], and DBiT-seq [14]; and laser capture micro-
dissection (LCM) combined with flow cytometry methods, such as GEO-seq [15] and Li and Yang Genome Biology (2022) 23:124
https://doi.org/10.1186/s13059-022-02692-0 Li and Yang Genome Biology (2022) 23:124
https://doi.org/10.1186/s13059-022-02692-0 Li and Yang Genome Biology (2022) 23:124 Page 2 of 24 Page 2 of 24 Li and Yang Genome Biology TSCS [16]. Due to different experimental designs, the coverage of these transcriptome
profiles ranges from tens to thousands of genes. Spatially resolved transcriptome profiles provide insightful resources for understand-
ing the cellular organization patterns in multiple types of tissues and organs, such as
the central nervous system [13, 17, 18], developing human heart [19], and tumors [13,
20]. However, these snapshots of tissue sections are still incomplete observations of the
multicellular organization and the potential local interactions between geometrically
adjacent cells. It remains a major challenge to infer full cell interaction networks, in-
cluding distal cell-cell interactions, which are potentially mediated by a broad range of
different mechanisms. Various methods are currently available for spatially resolving
predefined cell types with bulk RNA-seq or scRNA-seq data and for inferring cell-cell
interactions based on known ligand-receptor pairs or other predefined features and ref-
erences [21–25]. However, methods for the de novo reconstruction of cell interaction
networks in an unbiased and more comprehensive manner by taking full advantage of
spatial transcriptome profiles at single-cell resolution are still lacking. In addition, due to various technical limitations, spatially resolved single-cell tran-
scriptome profiles suffer to different extents from data imperfections such as too many
missing values, batch effects, biased and low coverage, and high noise levels [26, 27],
which necessitate methods with a sophisticated design for mining cell-cell interactions
from such imperfect and incomplete snapshots. Ideally, such methods should be cap-
able of reducing the noise of spatial transcriptome data, reconstructing existing interac-
tions, restoring missing interactions (including distal interconnections), and mining
latent features related to cell-cell interaction landscapes. The nonlinear, high-dimensional, sparse and multimodal features of single-cell spatial
transcriptome data make it a proper and feasible target of the deep learning strategy. Multiple deep learning models have been applied for various tasks with single-cell
spatial transcriptome data, for example, integration of histology to define spatial do-
mains and predict local gene expressions with convolutional networks [28, 29], imput-
ation of spatial transcriptome profiles by graph-regularized tensor completion [30],
inference of gene-gene interactions with convolutional networks [31], and integration
of sc/snRNA-seq data with nonconvex optimization to resolve in situ cell clusters [32]. These methods serve as powerful frameworks for learning from multimodal data and
resolving the spatial cell and gene organizations in tissues [33]. However, none of them
were designed to directly uncover the cell-cell interactions that shape the tissue
organization and define tissue physiological functions. Deep generative models such as generative adversarial networks (GANs) and vari-
ational autoencoders (VAEs) have been proven to be powerful tools for leveraging la-
tent features and modeling high-dimensional scRNA-seq data for various tasks, such as
denoising [34], clustering [35], dimension reduction [36], and missing value imputation
[37]. In the present study, we adapted another type of deep generative model, vari-
ational graph autoencoders (VGAEs) [38], for encoding cell-cell interaction features
from spatial single-cell transcriptome data and eventually regenerating full cell-cell
interaction landscapes. VGAE is a recently developed state-of-the-art machine learning
algorithm in which the core of variational autoencoders has been adapted [39] and ex-
tended for graph representation learning [40]. Our method, referred to as DeepLinc
(deep learning framework for landscapes of interacting cells), uses the VGAE model to Li and Yang Genome Biology (2022) 23:124 Page 3 of 24 Page 3 of 24 integrate and learn from the two dimensions of information (i.e., cell interactions and
gene expression profiles) during the encoding phase. Furthermore, the adversarial strat-
egy was applied to force the encoder to explicitly approximate the Gaussian distribution
[41]. The decoding phase then uses the latent representation learned during encoding
to reconstruct a cell-cell interaction graph. Specifically, DeepLinc assumes that the neighboring cells should be much more likely
to have some types of interactions than randomly picked non-neighboring cells that are
far away from each other. Therefore, for a particular tissue region, the neighboring cell
pairs comprise an incomplete and potentially noisy observation of a subset of the full
cell-cell interaction network. The main task of DeepLinc is to learn from this subset of
cell-cell interactions, extract the underlying features of single-cell transcriptome pro-
files, and finally, regenerate a more unbiased and complete landscape of cell-cell inter-
actions, which would include both proximal and distal interactions. To the best of our
knowledge, DeepLinc is the first framework of its kind to apply a deep generative
model for recovering cell interactions from spatial single-cell transcriptomic data. Based on a wide array of tests with real and simulated data, DeepLinc demonstrated
its high efficiency in learning from imperfect and incomplete spatial transcriptome data,
filtering false interactions, and inferring missing distal and proximal interactions. The
reconstructed full networks of cell interactions exhibited high physiological relevance. Such de novo reconstructions of cell interaction networks do not depend on prior
knowledge of cell types, ligand-receptor pairs, or cell interaction mechanisms. Further-
more, the interrogation of the pipeline revealed signature genes that are potentially in-
volved in shaping cell interaction landscapes. Finally, the reconstructed cell interaction
landscapes, which are presumably more complete than the original snapshots of cell
spatial organization, categorized hub cells and partitioned cells into further subclusters
based on both the features of cell interaction landscapes and transcriptome profiles. These new insights could greatly aid in elucidating the biological relevance and poten-
tial machinery underlying cell organization patterns in physiological contexts. In sum-
mary, as a specially designed method based on deep learning, DeepLinc is of unique
value for mining the latent features of cell interaction landscapes and, thus, for taking
full advantage of previous and future spatial transcriptome profiling data at single-cell
resolution. The DeepLinc model and the datasets used for testing The DeepLinc model and the datasets used for testing We assume that in a solid tissue the neighboring cells with direct contacts should be
much more likely to have some types of interactions than randomly picked non-
neighboring cells that are far away from each other. It is well recognized that single-cell
transcriptome profiles represent both the driving force and consequences of cell-cell
interaction landscapes [4, 5]. DeepLinc combines the VGAE and an adversarial network
to learn from single-cell spatial transcriptome profiles and generate a latent distribution
capturing the intrinsic associations between cell-cell interactions and the gene expres-
sion patterns of single cells (Fig. 1A). Specifically, adjacent cell pairs defined simply by the geometric closeness between
single cells were summarized as an undirected adjacency network in which the nodes Page 4 of 24 Page 4 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology Fig. 1 Overview of DeepLinc and the datasets. A Schematic description of the DeepLinc pipeline. DeepLinc
consists of an encoder, a decoder, and an adversarial regularization module in the VGAE framework. The
encoder is a two-layer graph convolutional network (GCN) and the decoder is a Sigmoid function for the
dot product of latent variables. The single-cell transcriptome profile and the cell adjacency matrix derived
from a single-cell spatial transcriptome dataset serve as inputs of DeepLinc. The output of DeepLinc is a
new matrix (A’) presenting the reconstructed cell-cell interaction network. In addition, DeepLinc also uses
the latent information of cell interaction landscapes and gene expression profiles for the visualization and
clustering of single cells. B Statistics of the 4 datasets used for reconstruction of cell interaction networks
by DeepLinc represent the cells and the edges indicate the neighboring cell pairs. This network is
represented by cell adjacency matrix A. The single-cell gene expression profiles, as fea-
tures of the nodes in A, are presented as the matrix X (Fig. 1A). The graph-structured
data (A) and the node features (X) are fed into the VGAE consisting of two graph con-
volutional layers. As the output of this variational graph convolutional network
(VGCN) encoder, the latent representation (H) captures the characteristics of a single
cell itself and its neighboring cells. In addition, H is further constrained by an adversar-
ial regularization module from a prior Gaussian distribution (Fig. 1A). Modeling of cell interactions by DeepLinc and the effects of denoising Modeling of cell interactions by DeepLinc and the effects of denoising
Single-cell spatial transcriptome profiles provide snapshots of cell-cell interaction land-
scapes, which are presumably just small subsets of interactions from complete cell-cell
interaction networks and usually contaminated by high levels of noise. Therefore, the
ability of DeepLinc to efficiently learn from the provided information on cell spatial
organization and single-cell transcriptome profiles, filter out noise, correctly infer latent
interactions, and eventually recover complete cell interaction landscapes is a critical
benchmark. DeepLinc was applied to the 4 seqFISH, MERFISH, HDST olfactory bulb, and HDST
breast cancer datasets (Fig. 1B). First, cell adjacency maps were defined by assembling
the neighboring cells in the tissue sections. Specifically, the distances between each pair
of cells (Additional file 1: Fig. S1A) or between each cell and its 3 closest neighbors
(Additional file 1: Fig. S1B) were used to assess the overall distance distributions of ad-
jacent cell pairs with potential direct interactions. The distance threshold for defining
direct contacts in each tissue section was then determined based on the above distribu-
tions (Additional file 1: Fig. S1B). For each cell, only the 3 closest neighbors falling
under the distance threshold were defined as direct contacts. The union of the direct
contacts of all the cells then constituted the full adjacency matrix. DeepLinc uses neigh-
boring cells with direct contacts as the positive set for learning the transcriptome fea-
tures related to cell-cell interactions. From a general biological point of view, we think
it is reasonable to assume that in a solid tissue, most of the cells in 2-D could directly
contact with 3 or more other cells. The edges of direct contacts were randomly divided into two groups for training
(90%) and testing (10%). The negative set for testing, which was 100 times larger than
the positive set, was composed of fake edges between two randomly picked non-
neighboring cells on the tissue section. For all of the 4 datasets, DeepLinc showed high
sensitivity and accuracy in recovering the originally annotated cell-cell interactions,
with area under the receiver operating characteristic (AUROC) ranging from 0.8 to lar-
ger than 0.9 (Fig. 2A) and false positive rate (FPR) below 5% (Additional file 1: Fig. S2). The DeepLinc model and the datasets used for testing Next, with the
information learned above, the decoder performs a dot product operation on H to gen-
erate a new adjacency matrix (A’) presenting the reconstructed cell-cell interaction net-
work (Fig. 1A). On the other hand, the vectors of H, which represent the latent
information of cell interaction landscapes and gene expression profiles, could be ex-
tracted for the visualization and clustering of single cells. DeepLinc was applied to 4 published spatial transcriptomic datasets obtained from
the mouse visual cortex [17], the preoptic region of the mouse hypothalamus [18], the
mouse olfactory bulb [13], and human breast cancer [13] (Fig. 1B). Note that the spatial Li and Yang Genome Biology (2022) 23:124 Page 5 of 24 Page 5 of 24 Li and Yang Genome Biology transcriptome profiling techniques and the sparseness of the gene expression profiles
vary greatly across these studies (Fig. 1B). In brief, the seqFISH and MERFISH datasets
show much lower gene coverage (fewer than 200 genes) than the two HDST datasets
(approximately 10,000 genes). However, the latter 2 high-throughput datasets are very
sparse, and large numbers of genes were not detected in all the single cells in these
datasets. Modeling of cell interactions by DeepLinc and the effects of denoising This process was repeated
30 times for each proportion of the missing edges to draw a boxplot. C The AUROCs between mixed-in fake
edges and pre-existing real edges. Different numbers (1- to 10-fold of the size of the original cell adjacency
map) of randomly generated fake edges were added into the original cell adjacency map, which was then fed
to DeepLinc. This process was repeated 30 times for each level of fake edges to draw a boxplot Fig. 2 Performance of DeepLinc in reconstructing cell interaction networks. A The levels of AUROC at different
training epochs of DeepLinc. The original data or data with different levels of random noise in the gene
expression profiles were used. Larger σ values would result in higher levels of random noise in general (see
“Methods” for details). B The AUROCs between the randomly removed real edges and originally nonexistent
edges. Different proportions of the edges in the original cell adjacency maps were randomly picked and
discarded. Then, the remaining edges were used for training the DeepLinc model. This process was repeated
30 times for each proportion of the missing edges to draw a boxplot. C The AUROCs between mixed-in fake
edges and pre-existing real edges. Different numbers (1- to 10-fold of the size of the original cell adjacency
map) of randomly generated fake edges were added into the original cell adjacency map, which was then fed
to DeepLinc. This process was repeated 30 times for each level of fake edges to draw a boxplot Methods for details). B The AUROCs between the randomly removed real edges and originally nonexistent
edges. Different proportions of the edges in the original cell adjacency maps were randomly picked and
discarded. Then, the remaining edges were used for training the DeepLinc model. This process was repeated
30 times for each proportion of the missing edges to draw a boxplot. C The AUROCs between mixed-in fake
edges and pre-existing real edges. Different numbers (1- to 10-fold of the size of the original cell adjacency
map) of randomly generated fake edges were added into the original cell adjacency map, which was then fed
to DeepLinc. This process was repeated 30 times for each level of fake edges to draw a boxplot transcriptome data. Modeling of cell interactions by DeepLinc and the effects of denoising Notably, DeepLinc modeled the cell interaction networks almost equally well with ei-
ther the highly sparse high-throughput datasets (HDST olfactory bulb and HDST
breast cancer) or the low-throughput datasets (seqFISH and MERFISH). Next, we tested the tolerance of DeepLinc to artificial noise in the single-cell gene ex-
pression data. Specifically, different levels of random noise with Gaussian distributions
were introduced on top of the original gene expression profiles of all the cells
(Methods). As shown in Fig. 2A, performances of DeepLinc were not compromised by
fairly high levels of noise, indicating the robustness of DeepLinc to noise in gene ex-
pression data, which is critical when dealing with highly noisy single-cell spatial Page 6 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology Fig. 2 Performance of DeepLinc in reconstructing cell interaction networks. A The levels of AUROC at different
training epochs of DeepLinc. The original data or data with different levels of random noise in the gene
expression profiles were used. Larger σ values would result in higher levels of random noise in general (see
“Methods” for details). B The AUROCs between the randomly removed real edges and originally nonexistent
edges. Different proportions of the edges in the original cell adjacency maps were randomly picked and
discarded. Then, the remaining edges were used for training the DeepLinc model. This process was repeated
30 times for each proportion of the missing edges to draw a boxplot. C The AUROCs between mixed-in fake
edges and pre-existing real edges. Different numbers (1- to 10-fold of the size of the original cell adjacency
map) of randomly generated fake edges were added into the original cell adjacency map, which was then fed
to DeepLinc. This process was repeated 30 times for each level of fake edges to draw a boxplot Fig. 2 Performance of DeepLinc in reconstructing cell interaction networks. A The levels of AUROC at different
training epochs of DeepLinc. The original data or data with different levels of random noise in the gene
expression profiles were used. Larger σ values would result in higher levels of random noise in general (see
“Methods” for details). B The AUROCs between the randomly removed real edges and originally nonexistent
edges. Different proportions of the edges in the original cell adjacency maps were randomly picked and
discarded. Then, the remaining edges were used for training the DeepLinc model. Modeling of cell interactions by DeepLinc and the effects of denoising However, further corruption of the data dramatically impaired the
performance of DeepLinc, illustrating the minimal gene expression profile information
needed for DeepLinc to correctly rebuild the cell interaction landscapes. Single-cell spatial transcriptome data is largely suffering from high sparsity due to
large numbers of dropouts, especially for the high-throughput data such as HDST. Therefore, we also generated two more noise models to simulate different types of
dropouts (Additional file 1: Fig. S3). In the first model, different percentages of genes
were randomly removed from the expression dataset to mimic dropouts of genes, and
in the second model, different percentages of the non-zero values were randomly
picked and forced to be zero, which mimics dropouts of individual datapoints in single
cells. DeepLinc showed high tolerance to the noise from both types of dropouts (Add-
itional file 1: Fig. S3). In general, the performances of DeepLinc were still fairly good
with the dropout ratio of 50%. As well expected, more severe dropouts would strongly
reduce the accuracy of DeepLinc. In summary, these tests confirm the advantage of
DeepLinc in extracting the latent information of cell-cell interactions from the highly
sparse and noisy spatial transcriptome profiles. Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology (2022) 23:124 Page 7 of 24 Page 7 of 24 DeepLinc also demonstrated relatively stable and high levels of fitting (AUC 85~95%)
with different sizes of tissue sections that were randomly picked from the original
frame of the tissue sections and therefore included fewer cells than the original data
(Additional file 1: Fig. S4). The performance of DeepLinc was still fairly good with small
tissue sections containing as few as 500 cells, indicating the efficient learning capability
of DeepLinc. This is a useful feature for dissecting the cell organization of ultrafine tis-
sue sections. We also tested DeepLinc for its capability to recover cell-cell interaction landscapes
with different proportions of missing interactions. Specifically, certain percentages of
the existing edges were randomly removed from the original cell-cell interaction net-
work. The remaining network and the single-cell transcriptome data were then used
for training the DeepLinc model. Ultimately, AUROC values were calculated to evalu-
ate the performance of DeepLinc in correctly distinguishing the arbitrarily removed in-
teractions (positives) and the originally nonexistent interactions (negatives). As shown
in Fig. 2B, DeepLinc exhibited high precision in imputing the missing positive interac-
tions. Modeling of cell interactions by DeepLinc and the effects of denoising For example, even when half of the interactions were lost in the input, DeepLinc
still managed to recover these missing edges with 65–75% precision (Fig. 2B). This indi-
cates that DeepLinc can indeed efficiently learn from limited and incomplete networks
of cell interactions to reinstate missing edges. Such an imputation and prediction cap-
ability is critical for applications of DeepLinc for the inference of full cell interaction
landscapes from the incomplete snapshots generated from spatially resolved single-cell
transcriptome profiles. In addition to the missing interactions (i.e., false negatives), false annotations of cell-
cell interactions (i.e., false positives) that are fed to DeepLinc are another major issue
to address. To test the robustness of DeepLinc in correctly recovering cell interactions
with random false interactions (i.e., the denoising capability), we added fake edges be-
tween randomly picked cell pairs into the original cell interaction network. Taking such
highly noisy data as the input, DeepLinc can still nicely distinguish the originally non-
existent edges and the real pre-existing edges (Fig. 2C). Even when fake edges were arti-
ficially added in numbers as high as several-fold the original number of cell
interactions, DeepLinc still managed to filter out the extensive noise and recover the
predefined cell interactions with fairly good precision (Fig. 2C). This indicates the high
tolerance of DeepLinc to random false interactions and its efficient denoising effects Reconstructed cell-cell interaction landscapes The landscapes of the reconstructed and the original cell-cell interaction networks were
interrogated with a permutation test as described previously [21, 24, 42] to evaluate the
enrichment or depletion of certain types of interactions in a network by comparison
with randomly assembled networks with similar scales. As shown in Fig. 3A–D, the
cell-cell interaction landscapes reconstructed by DeepLinc exhibited substantial differ-
ences from the original cell adjacency networks composed of proximal cell pairs with
direct contacts. For the visual cortex data of seqFISH, relative to the original network, the recon-
structed network reinforced the interactions between the astrocytes and the two types
of neurons (glutamatergic and GABAergic) (Fig. 3A). Interactions between epithelial Page 8 of 24 Page 8 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology Fig. 3 Landscapes of cell interactions reconstructed by DeepLinc. A–D Evaluation of the over- or under-
representation of particular types of interactions in the original cell adjacency networks (left) and in the
reconstructed cell interaction networks (right) with the 4 datasets. The results are represented as dot
heatmaps showing the interaction relationships between cell types. Dot size represents the connectivity
between two cell types, calculated by the number of connections divided by the product of the cell
numbers. The color of the dots represents significance (−log10P) determined by comparing with the
permutation test, in which positive values indicate enrichment and negative values indicate depletion. E
Original and reconstructed cell interaction networks in a representative tissue section from MERFISH dataset
of hypothalamic preoptic area. The red box indicates a microglia cell with distal interactions with many
other cells in the reconstructed networks Fi
3 L
d
f
ll i
i
d b D
Li
A D E
l
i
f h
d Fig. 3 Landscapes of cell interactions reconstructed by DeepLinc. A–D Evaluation of the over- or under-
representation of particular types of interactions in the original cell adjacency networks (left) and in the
reconstructed cell interaction networks (right) with the 4 datasets. The results are represented as dot
heatmaps showing the interaction relationships between cell types. Dot size represents the connectivity
between two cell types, calculated by the number of connections divided by the product of the cell
numbers. The color of the dots represents significance (−log10P) determined by comparing with the
permutation test, in which positive values indicate enrichment and negative values indicate depletion. Reconstructed cell-cell interaction landscapes E
Original and reconstructed cell interaction networks in a representative tissue section from MERFISH dataset
of hypothalamic preoptic area. The red box indicates a microglia cell with distal interactions with many
other cells in the reconstructed networks Fig. 3 Landscapes of cell interactions reconstructed by DeepLinc. A–D Evaluation of the over- or under-
representation of particular types of interactions in the original cell adjacency networks (left) and in the
reconstructed cell interaction networks (right) with the 4 datasets. The results are represented as dot
heatmaps showing the interaction relationships between cell types. Dot size represents the connectivity
between two cell types, calculated by the number of connections divided by the product of the cell
numbers. The color of the dots represents significance (−log10P) determined by comparing with the
permutation test, in which positive values indicate enrichment and negative values indicate depletion. E
Original and reconstructed cell interaction networks in a representative tissue section from MERFISH dataset
of hypothalamic preoptic area. The red box indicates a microglia cell with distal interactions with many
other cells in the reconstructed networks cells and almost all the other cell types were significantly enriched in the reconstructed
network. On the other hand, despite the high count, the interactions among the gluta-
matergic neurons appeared depleted when compared to random networks (Fig. 3A),
whereas such interactions were highly enriched in the original cell adjacency network. Therefore, the reconstructed network indicates high selectivity of the interactions
among the excitatory glutamatergic neurons. For the MERFISH data of the mouse hypothalamic preoptic region, the reconstructed
network strengthened the interactions within endothelial cells and between endothelial
cells and astrocytes (Fig. 3B), similar to the observations based on the mouse visual cor-
tex data discussed above. However, unlike the visual cortex, the mouse hypothalamic
preoptic region showed enriched interactions between excitatory (mainly glutamatergic)
and inhibitory (GABAergic) neurons in the reconstructed network. In addition, the in-
teractions within excitatory neurons were strongly enriched (Fig. 3B), whereas the op-
posite observation was obtained from the visual cortex data (Fig. 3A). Therefore, the
above results revealed both common and tissue-specific cell interaction patterns in the
two types of CNS systems. Reconstructed cell-cell interaction landscapes Finally, unlike the original cell adjacency network, the re-
constructed network of the mouse hypothalamic preoptic region showed strong Li and Yang Genome Biology (2022) 23:124 Page 9 of 24 Page 9 of 24 enrichment according to the interactions among the mature oligodendrocytes, the in-
teractions between microglia and many other cells, and the interactions between peri-
cytes and other cells (Fig. 3B). The cell interaction network reconstructed from HDST data of the olfactory bulb
showed more enriched interactions between oligodendrocytes and most of the other
cell types, whereas the interactions between immune cells and other cells were reduced
relative to the random networks (Fig. 3C). In addition, the interactions among vascular
cells and glial cells (peripheral glia, astrocytes, and ependymal cells) were specifically in-
tensified in the reconstructed network (Fig. 3C), indicating strong internal interactions
within the complex blood–brain barrier composed of these cells [43–45]. Finally, for the breast cancer tissue profile data obtained with HDST, the recon-
structed network exhibited strong interactions between lymphocytes (T cells and B
cells) and almost all the other cell types (Fig. 3D). In contrast, the interactions between
stromal cells and endothelial cells were enriched in the original network but depleted
in the reconstructed network, suggesting high specificity of such interactions (Fig. 3D). In summary, by learning from the cell adjacency maps defined simply by the spatial
organization of cells, which are incomplete, sparse, and noisy, DeepLinc performs the
imputation of missing interactions and the filtration of potentially false connections. Ultimately, the reconstructed interaction networks are significantly enlarged and
reshaped, which should recapitulate the cell interaction landscapes in a more compre-
hensive and precise manner. As shown by the local regions from different tissues in Fig. 3E and Additional file 1:
Fig. S5 as examples, small numbers of the originally defined direct cell interactions
were removed, and new interactions were added by DeepLinc. The reconstructed net-
works had more hierarchical and modularized structures. Interestingly, in the two ex-
amples of visual cortex and hypothalamic preoptic area, microglia showed dense
connections with many other cells, including neurons, astrocytes, endothelial cells, and
oligodendrocytes, thereby serving as hub cells (Fig. 3E, Additional file 1: Fig. S5C),
which nicely recapitulated the wide-ranging functions of microglia and their well-
acknowledged interactions with other cell types in the CNS [46]. Such insight was miss-
ing in the original cell-cell interaction network. Distal interactions in the reconstructed cell networks The probability of cell-cell interactions predicted by DeepLinc showed slightly negative
correlations with geometric distances in general, and significant numbers of distal cell
pairs presented high probabilities of interactions (Additional file 1: Fig. S6). Such distal
interactions were missing in the original cell adjacency map, and DeepLinc demon-
strated its capability to recover cell-cell interactions that are not restricted by spatial
proximity. The newly recovered distal interactions showed distinct patterns of participating cell
types. For example, in the seqFISH study of the mouse visual cortex, distal interactions
were highly enriched between neurons (Fig. 4A). An example is provided in Fig. 4E to
show the interactions between a glutamatergic neuron and other GABAergic neurons. By contrast, distal interactions were depleted between endothelial cells (Fig. 4A). This
is in line with the notion that endothelial cells form a tight one-cell-thick interior Page 10 of 24 Page 10 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology Fig. 4 Landscapes of distal interactions in the reconstructed cell networks. A–D Evaluation of the over- or
under-representation of particular types of distal interactions in the reconstructed cell interaction networks
with the 4 datasets. The results are represented as dot heatmaps showing the enrichment or depletion of
the distal interactions between certain cell types. The color of the dots represents significance (−log10P)
determined by comparing with the permutation test, in which positive values indicate enrichment, and
negative values indicate depletion. E–G Examples of distal interactions between a glutamatergic neuron
and GABAergic neurons (E), between an epithelial cell and stromal cells (F), and between a T cell and other
T cells (G). The proximal interactions between neighboring cells are shown as backgrounds Fig. 4 Landscapes of distal interactions in the reconstructed cell networks. A–D Evaluation of the over- or
under-representation of particular types of distal interactions in the reconstructed cell interaction networks
with the 4 datasets. The results are represented as dot heatmaps showing the enrichment or depletion of
the distal interactions between certain cell types. The color of the dots represents significance (−log10P)
determined by comparing with the permutation test, in which positive values indicate enrichment, and
negative values indicate depletion. E–G Examples of distal interactions between a glutamatergic neuron
and GABAergic neurons (E), between an epithelial cell and stromal cells (F), and between a T cell and other
T cells (G). Distal interactions in the reconstructed cell networks The proximal interactions between neighboring cells are shown as backgrounds Fig. 4 Landscapes of distal interactions in the reconstructed cell networks. A–D Evaluation of the over- or
under-representation of particular types of distal interactions in the reconstructed cell interaction networks
with the 4 datasets. The results are represented as dot heatmaps showing the enrichment or depletion of
the distal interactions between certain cell types. The color of the dots represents significance (−log10P)
determined by comparing with the permutation test, in which positive values indicate enrichment, and
negative values indicate depletion. E–G Examples of distal interactions between a glutamatergic neuron
and GABAergic neurons (E), between an epithelial cell and stromal cells (F), and between a T cell and other
T cells (G). The proximal interactions between neighboring cells are shown as backgrounds interface in blood vessels, and it is not surprising to see a lack of distal interactions be-
tween endothelial cells. In addition, a similar depletion of distal interactions between
endothelial cells was observed in the HDST breast cancer data (Fig. 4D). In the hypothalamic preoptic region dataset obtained with MERFISH, the newly re-
constructed distal interactions were depleted between pericytes and other cell types
(Fig. 4B) relative to the expectation based simply on chance. Indeed, embedded in the
basement membrane of blood capillaries, pericytes directly interlock and communicate
with endothelial cells [47]. Long-distance interactions between pericytes and other cell
types are indeed not expected. Being similar to the seqFISH study of the mouse visual cortex, an enrichment of the
distal interactions between neurons was also observed in the MERFISH data of hypo-
thalamic preoptic region and HDST olfactory bulb dataset (Fig. 4B, C). In addition, in
both the hypothalamic preoptic region and the olfactory bulb, the newly reconstructed
distal interactions were depleted between neurons and glial cells (Neu-Glial in Fig. 4B,
and Neu-Epen/Astro/Per/Oligo in Fig. 4C) relative to the expectation based simply on
chance. By contrast, in the olfactory bulb, immune cells showed a broad spectrum of
long-range interactions with almost all cell types, especially with neurons, indicating
abundant nerve-immune circuits (Fig. 4C). In the HDST breast cancer data, the newly recovered distal interactions were
enriched with interactions between epithelial and stromal cells (Fig. 4D). An example is Li and Yang Genome Biology (2022) 23:124 Page 11 of 24 provided in Fig. 4F to show the interactions between an epithelial cell and multiple
stromal cells. Distal interactions in the reconstructed cell networks This was not surprising given the frequent reports of stromal-epithelial
cell interactions mediated by cytokines and extracellular vesicles [48–50], which confer
high potential for remote interactions [51, 52]. Similarly, T cells or B cells heavily rely
on long-range communication via cytokines, a notion reflected by the distal interac-
tions enriched among T or B cells in the reconstructed network (Fig. 4D, and an ex-
ample illustrating interactions between T cells is given in Fig. 4G). In contrast, the
long-distance interactions were depleted in interactions that are highly dependent on
direct cell contacts (for example, antigen presentation), such as the interactions be-
tween T and B cells, T cells and epithelial cells, T cells and macrophages, B cells and
epithelial cells, and B cells and macrophages (Fig. 4D). In the HDST breast cancer data, the epithelial cells were mostly tumor cells. Taking 4 randomly picked T cells and epithelial cells as examples, Additional file
1: Fig. S7 shows the distances between the interacting cell pairs, which were gener-
ally much shorter than those from the randomly shuffled networks. Therefore,
DeepLinc recovered the enriched interactions between T cells and tumor cells (Fig. 3D), which were strongly limited to proximal and not distal interactions (Fig. 4D,
Additional file 1: Fig. S7). Interestingly, in contrast to epithelial cells, endothelial cells were more involved in
distal interactions with immune cells (Fig. 4D). Indeed, it has been frequently reported
that endothelial cells secrete chemokines to facilitate the transmigration and homing of
immune cells into tumor tissues [53, 54]. DeepLinc identifies signature genes shaping the cell interaction landscapes Next, we further interrogated the DeepLinc model to infer the signature genes that play
important roles in shaping the comprehensive landscapes of cell-cell interactions. In
brief, the expression profile of a particular gene was shuffled across all the single cells,
and the new transcriptome data and the cell adjacency network were then fed to Dee-
pLinc. The AUPRC calculated from the cell network reconstructed by DeepLinc with
these modified data was then compared to the original AUPRC with the unmodified
gene expression data. The reduction, defined as the sensitivity score, indicates the sen-
sitivity of the DeepLinc model to the particular gene being shuffled and therefore serves
as an assessment of the potential contribution of the gene to the cell-cell interaction
landscape. The same procedure was performed for each gene one at a time to obtain
the sensitivity scores of all the genes (Additional file 1: Fig. S8). The top-rated genes with the highest sensitivity scores were indeed enriched by mul-
tiple molecular and cellular processes related to cell-cell interactions (Fig. 5A–D). For
example, 205 genes (Additional file 2: Table S1) showed outstanding sensitivity scores
with the HDST breast cancer data, and these genes were involved in the processes of
cell-substrate adhesion, junction formation, and protein targeting to the membrane
(Fig. 5A, and the full list of functional categories supplied in Additional file 1: Fig. S9A). SPATA2, which showed the highest sensitivity score (Additional file 1: Fig. S8B),
mediates the recruitment of CYLD to the LUBAC complex [55, 56], thereby playing a
crucial role in the TNFR1- and NOD2-signaling pathways. The rest of the top 10 genes
listed in Additional file 1: Fig. S8B include PKHD1 (transmembrane or secreted protein, Page 12 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology Fig. 5 Biological processes enriched by the signature genes in shaping the cell interaction networks. A, B
For the two high-dimensional datasets of HDST, the biological processes involving the top-ranked genes
were obtained through Gene Ontology Enrichment Analysis. C, D For two low-throughput datasets
(seqFISH and MERFISH), due to the limitation of gene numbers, all the genes were sorted by their sensitivity
scores, and gene set enrichment analysis was performed. E Prediction of cell interactions with genes
annotated as ligands or receptors and the same number of top-ranked signature genes identified by
DeepLinc in the seqFISH (left) and MERFISH (right) datasets. DeepLinc identifies signature genes shaping the cell interaction landscapes Simple machine learning models, including
support vector classification (SVC), K-nearest neighbor (KNN), and gradient boosting decision tree (GBDT)
models, were used for this test. The originally defined cell adjacency map was used as a positive set of cell
interactions, and randomly selected nonadjacent cell pairs with the same numbers were used as a negative
set. This process was repeated 30 times with different training and testing set assignments. P-value were
obtained with Student’s t test Fig. 5 Biological processes enriched by the signature genes in shaping the cell interaction networks. A, B
For the two high-dimensional datasets of HDST, the biological processes involving the top-ranked genes
were obtained through Gene Ontology Enrichment Analysis. C, D For two low-throughput datasets
(seqFISH and MERFISH), due to the limitation of gene numbers, all the genes were sorted by their sensitivity
scores, and gene set enrichment analysis was performed. E Prediction of cell interactions with genes
annotated as ligands or receptors and the same number of top-ranked signature genes identified by
DeepLinc in the seqFISH (left) and MERFISH (right) datasets. Simple machine learning models, including
support vector classification (SVC), K-nearest neighbor (KNN), and gradient boosting decision tree (GBDT)
models, were used for this test. The originally defined cell adjacency map was used as a positive set of cell
interactions, and randomly selected nonadjacent cell pairs with the same numbers were used as a negative
set. This process was repeated 30 times with different training and testing set assignments. P-value were
obtained with Student’s t test involved in ciliogenesis, cell-cell and cell-matrix adhesion, and calcium ion homeosta-
sis), S100PBP (binding partner of the calcium sensor S100P, involved in cell adhesion)
[57], PSD4 (GEF for ARF6, involved in endocytosis), TRIM5 (capsid-specific restriction
factor, involved in innate immune signaling and autophagy), ADAP2 (GTPase-activat-
ing protein for ARFs, localizing to the plasma membrane, showing phospholipid bind-
ing activity and mediating TCR signaling), and CCL26 (C-C motif chemokine ligand
for CCR3). For the other HDST high-throughput spatial transcriptome dataset of the mouse ol-
factory bulb, the top-rated genes (Additional file 3: Table S2) were more specifically Li and Yang Genome Biology (2022) 23:124 Page 13 of 24 Page 13 of 24 involved in the cell-cell interactions of the nervous system, such as synaptic transmis-
sion, ion transmembrane transport, axonogenesis, synaptic vesicle activities, and trans-
synaptic signaling (Fig. DeepLinc identifies signature genes shaping the cell interaction landscapes 5B, and the full list of functional categories supplied in
Additional file 1: Fig. S9B). For the two low-throughput datasets of the mouse visual cortex (seqFISH) and mouse
hypothalamic preoptic region (MERFISH), we performed gene set enrichment analysis
(GSEA) to identify the processes involving more of the top-rated signature genes for
the cell interaction landscapes (Fig. 5C, D, and the full lists in Additional file 1: Fig. S10). In the mouse visual cortex, these processes included the inflammatory response,
response to external stimulus, defense response, extracellular structure organization,
signal release, secretory vesicle processes, and peptide secretion (Fig. 5C). In the hypo-
thalamic preoptic region, the identified processes included actin-based cell projection,
axon, membrane organization, cell-cell adhesion, secretory granule, and peptide secre-
tion (Fig. 5D). Together, the above results indicate the capability of the DeepLinc model to mine
transcriptome signatures that are directly or indirectly associated with cell interaction
landscapes. The high-impact genes revealed by the interrogation of the DeepLinc pipe-
line provide insights into the key processes defining different types of cell interactions
in specific tissue contexts. Interestingly, compared to the annotated ligand and receptor
genes, equal numbers of the top-rated signature genes were much more efficient in dis-
tinguishing interactive and noninteractive cell pairs with simple classifiers in both the
mouse visual cortex and mouse hypothalamic preoptic region datasets (Fig. 5E). This
once again suggests that the information on cell-cell interactions is largely coded in the
transcriptome and not limited to specific ligand and receptor genes, a notion that lays
the foundation of DeepLinc for the inference of cell interaction landscapes with spatial
transcriptome data. DeepLinc identifies multicellular domains informing organizational tissue structures The dot size represents the significance of the one-versus-
rest test (Mann–Whitney U test). The color indicates the median value of the normalized z-scores of gene
expression in each cluster Fig. 6 Spatial domains defined by reclustering of cells based on latent vectors recovered by DeepLinc. A, B
The UMAP graph and spatial locations of different spatial domain clusters and cell types. The spatial domain
clusters were acquired by the k-means method with latent vectors of the single cells. C, D Spatially specific
gene expression signatures of homologous cells reclustered by DeepLinc. The heatmaps show the
differential expression of domain-specific genes. The dot size represents the significance of the one-versus-
rest test (Mann–Whitney U test). The color indicates the median value of the normalized z-scores of gene
expression in each cluster g
p
y
g
y
p
,
The UMAP graph and spatial locations of different spatial domain clusters and cell types. The spatial domain
clusters were acquired by the k-means method with latent vectors of the single cells. C, D Spatially specific
gene expression signatures of homologous cells reclustered by DeepLinc. The heatmaps show the
differential expression of domain-specific genes. The dot size represents the significance of the one-versus-
rest test (Mann–Whitney U test). The color indicates the median value of the normalized z-scores of gene
expression in each cluster Taking astrocytes and GABAergic and glutamatergic neurons in the mouse visual cor-
tex as examples, the same types of cells allocated in the 6 domains showed significantly
differential expression patterns of some signature genes (Fig. 6C). The details of these
patterns were well in line with the inner-outer structure of the mouse visual cortex. However, the same types of cells located in the same layer were also partitioned into
different cell clusters (e.g., Clusters D1 and D3) (Fig. 6A). Although the astrocytes,
GABAergic neurons, and glutamatergic neurons in D1 and D3 showed similar patterns Taking astrocytes and GABAergic and glutamatergic neurons in the mouse visual cor-
tex as examples, the same types of cells allocated in the 6 domains showed significantly
differential expression patterns of some signature genes (Fig. 6C). The details of these
patterns were well in line with the inner-outer structure of the mouse visual cortex. However, the same types of cells located in the same layer were also partitioned into
different cell clusters (e.g., Clusters D1 and D3) (Fig. 6A). DeepLinc identifies multicellular domains informing organizational tissue structures As a deep data-generative model, DeepLinc generates a latent representation (H), which
captures both the characteristics of the single-cell transcriptome profile and its position
in the cell-cell interaction landscape. Therefore, we used this latent information of the
cells learned by DeepLinc to redefine the multicellular spatial domains in heteroge-
neous tissues. For example, in the seqFISH data of the mouse visual cortex, the embedding vectors in
H for each cell were used for unsupervised K-means clustering, resulting in 6 subgroups
of cells (Fig. 6A). The optimal cluster number was defined by Calinski-Harabasz scores
[58] (see methods for details). These cell clusters were largely different from the previ-
ously annotated cell types based only on known marker genes (Fig. 6A). Instead, the new
cell groups resembled the multilayered organization of the heterogeneous cells in the
mouse visual cortex [17] (Fig. 6A), which reflected the spatially restricted developmental
trajectory from the inner to outer regions of the tissue. Similarly, for the MERFISH data-
set of the hypothalamic preoptic area, the same analysis classified the tissue section into 7
subregions (Fig. 6B), which were again highly correlated with the original anatomy. Next, for the same types of cells that are allocated to these different spatially coded
cell clusters, differential expression analysis identified domain-specific signature genes. Page 14 of 24 Li and Yang Genome Biology (2022) 23:1 Li and Yang Genome Biology Fig. 6 Spatial domains defined by reclustering of cells based on latent vectors recovered by DeepLinc. A, B
The UMAP graph and spatial locations of different spatial domain clusters and cell types. The spatial domain
clusters were acquired by the k-means method with latent vectors of the single cells. C, D Spatially specific
gene expression signatures of homologous cells reclustered by DeepLinc. The heatmaps show the
differential expression of domain-specific genes. The dot size represents the significance of the one-versus-
rest test (Mann–Whitney U test). The color indicates the median value of the normalized z-scores of gene
expression in each cluster Fig. 6 Spatial domains defined by reclustering of cells based on latent vectors recovered by DeepLinc. A, B
The UMAP graph and spatial locations of different spatial domain clusters and cell types. The spatial domain
clusters were acquired by the k-means method with latent vectors of the single cells. C, D Spatially specific
gene expression signatures of homologous cells reclustered by DeepLinc. The heatmaps show the
differential expression of domain-specific genes. DeepLinc identifies multicellular domains informing organizational tissue structures Although the astrocytes,
GABAergic neurons, and glutamatergic neurons in D1 and D3 showed similar patterns Li and Yang Genome Biology (2022) 23:124 Page 15 of 24 Page 15 of 24 of gene expression in general, some genes were differentially expressed, indicating the
differences between D1 and D3 (Fig. 6C). This again indicates that DeepLinc captures
features from both gene expression profiles and spatially coded cell interaction land-
scapes. Similarly, for the MERFISH dataset of the hypothalamic preoptic area, neurons
allocated into different domains also showed differentially expressed signature genes
(Fig. 6D). In summary, DeepLinc rebuilt more comprehensive cell interaction landscapes by
learning from spatially resolved single-cell transcriptome profiles. The latent informa-
tion on spatially dependent cellular features redefined the geometric domains of tissues
composed of heterogeneous cells. These results therefore provide unique insights into
the spatially coded cellular organization underlying the potential physiological relevance
of cell interaction landscapes. Discussion As a major methodological breakthrough, technologies for profiling spatially resolved
transcriptomes have provided unprecedented opportunities for the dissection of highly
heterogeneous cellular organizations in physiological and pathophysiological contexts. However, it has been a major challenge to comprehensively and precisely infer cell-cell
interactions based on spatial transcriptome profiles at single-cell resolution. First,
spatial transcriptome profiles are random snapshots of very limited numbers of tissue
sections. The cellular organization patterns contain information on cell-cell interactions
and presumably represent only small collections sampled from the full landscape. Therefore, methods for correctly inferring the complete landscapes of cell interactions
by taking advantage of the observed cell geometric distributions and gene expression
profiles are urgently needed. Second, current spatial transcriptome data are limited by a
series of technical issues, including biased and low gene coverage, low signal-to-noise
ratios, high data sparsity under high-throughput techniques, and/or batch effects,
thereby necessitating data-mining methods that are highly robust to these technical
limitations. Powered by the strong data representation capacity of the VGAE model, DeepLinc
was specifically designed and optimized to denoise and infer missing interactions by ef-
ficiently learning from the incomplete sets of cell organization revealed by different
methods of single-cell spatial transcriptome profiling. Further analysis of the features
mined by DeepLinc helps identify the signature genes shaping cell interaction land-
scapes and cells with special features, such as hub cells and cells in specific domains of
the interaction landscape. Various methods have been available for inferring the cell-cell interaction landscapes
utilizing bulk RNA-seq, scRNA-seq, or spatially resolved single-cell data. These
methods are mostly based on predefined or previously annotated ligands, receptors,
and downstream target genes [23, 25, 59–61]. These methods have mostly been applied
to scRNA-seq data and are not well suited for single-cell spatial transcriptome data,
which show very limited and uneven coverage of the ligand and receptor genes due to
high sparsity and noise. This necessitates de novo methods, such as DeepLinc, for re-
construction of cell-cell interaction networks in a more unbiased manner independent
of prior knowledge such as ligand-receptor pairs. Instead, by assuming that cell inter-
action potentials have been coded in single-cell transcriptome profiles, DeepLinc is Li and Yang Genome Biology (2022) 23:124 Page 16 of 24 trained by the observed cell-cell connections and then utilized to infer missing interac-
tions. As shown in Fig. 4 and Additional file 1: Fig. Discussion S5, DeepLinc recovered great num-
bers of distal interactions, and meanwhile, DeepLinc removed some of the predefined
proximal interactions between neighboring cells. Therefore, DeepLinc is positioned as a
data-mining and processing strategy tailored for single-cell spatial transcriptome profil-
ing data, with the goal of recovering the full landscapes of cell interactions from limited
observations of tissue sections. Specifically, DeepLinc uses VGAE to encode the intrinsic characteristics of spatial
transcriptome data and decode cell interactions. The dual modal data (i.e., the graph-
like cell adjacency map and the high- or mid-throughput transcriptome profile) are in-
tegrated at the same time. Here, the design of VGAE constrains the model to put more
weight on the key genes by force-fitting the global cell interaction graph signatures. To
the best of our knowledge, DeepLinc is the first deep generative learning model for in-
ferring cell interactions from spatial transcriptomic data. Based on a wide array of tests
with real and simulated data, DeepLinc demonstrated high efficiency in learning from
imperfect and incomplete spatial transcriptome data, filtering false interactions, and in-
ferring missing distal and proximal interactions, thereby suggesting the reliability and
effectiveness of the deep learning strategy. With the DeepLinc model, our sensitivity test of transcriptome profiles suggested sig-
nature genes with potentially high impacts on the cell interaction landscape. Import-
antly, compared to known ligand and receptor genes, these genes showing high
sensitivities performed better in simple classification models of cell interactions. This
supports our presumption that cell-cell interactions are coded in single-cell transcrip-
tome profiles, rather than simply being defined by ligand and receptor genes. The iden-
tification of these signature genes could facilitate the understanding of the various
machineries underlying heterogeneous cell interactions and further suggests the possi-
bility of directly predicting cell interactions simply from single-cell transcriptome pro-
files. Further refined models would be needed for this purpose. Due to the lack of more detailed information in spatial transcriptome data, DeepLinc
does not provide the directionality or strength of the cell interactions. Potentially different
modes of cell interactions also cannot be differentiated by DeepLinc in its current form. Finally, similar to most of the deep learning models based on GAE, it is difficult to effi-
ciently and directly backtrack the important features encoded by the graph convolutional
networks used in DeepLinc. Conclusions DeepLinc is a novel computational method, with a strategy of deep learning, for de
novo reconstruction of cell interaction landscapes from single-cell spatial transcriptome
profiles. With simulated and real data, DeepLinc demonstrates high efficiency in learn-
ing from imperfect and incomplete spatial transcriptome data, filtering false interac-
tions, and inferring missing distal and proximal interactions. Interrogations of the
DeepLinc pipeline reveal signature genes that are potentially involved in shaping cell
interaction landscapes. In addition, clustering of cells based on the latent features of
cell interactions and transcriptome profiles learned by DeepLinc indicates multicellular
domains informing organizational tissue structures. Discussion Technical advances in spatial transcriptome profiling and
breakthroughs in graph convolutional models would help further improve DeepLinc in
the future. In addition, if more information becomes available due to future technological
advances, DeepLinc could be further expanded to incorporate features such as cell
morphology, surface markers, and metabolic profiles. Finally, although this is not the main
target of DeepLinc, the framework can be used for inferring single-cell gene expression
profiles by using information on cell interactions based on the assumption that the gene
expression profiles are partly correlated with local and distal cell interaction patterns [62]. In summary, as a tool for deep data mining, DeepLinc further empowers rapidly
emerging spatial transcriptome profiles for the de novo reconstruction of cell inter-
action maps. This new strategy, facilitated by deep graph convolutional networks,
does not rely on any prior knowledge of cells or gene functions. We anticipate that
the combination of state-of-the-art spatial transcriptome profiling techniques and Li and Yang Genome Biology (2022) 23:124 Page 17 of 24 an efficient data deep mining framework will greatly facilitate the identification of
the biological mechanisms underlying complex cell communication networks with
physiological relevance. Datasets of single-cell spatial transcriptome profiles Four published datasets were used for the current study [13, 17, 18], including mouse
visual cortex profiled by seqFISH [17], preoptic region of the mouse hypothalamus by
MERFISH [18], mouse olfactory bulb, and human breast cancer by HDST [13] (Fig. 1B). Fewer than 200 genes were profiled by seqFISH and MERFISH, whereas HDST
generates high-throughput transcriptome profiles coving around 10,000 genes (Fig. 1B). Four published datasets were used for the current study [13, 17, 18], including mouse
visual cortex profiled by seqFISH [17], preoptic region of the mouse hypothalamus by
MERFISH [18], mouse olfactory bulb, and human breast cancer by HDST [13] (Fig. 1B). Fewer than 200 genes were profiled by seqFISH and MERFISH, whereas HDST
generates high-throughput transcriptome profiles coving around 10,000 genes (Fig. 1B). The seqFISH dataset was downloaded from the data portal at https://bitbucket.org/qzhu/
smfish-hmrf/src/master/. The data was processed according to the procedure described in
the original study [17]. The final dataset covers 125 genes measured in 1597 cells. The seqFISH dataset was downloaded from the data portal at https://bitbucket.org/qzhu/
smfish-hmrf/src/master/. The data was processed according to the procedure described in
the original study [17]. The final dataset covers 125 genes measured in 1597 cells. The MERFISH dataset of the mouse hypothalamic preoptic region was downloaded
from https://datadryad.org/stash/dataset/10.5061/dryad.8t8s24818. We used the slice re-
gion at Bregma+0.11 mm of the animal No. 18, as it contains the largest number of sin-
gle cells. After removing the ambiguous cells, the final dataset contains 4975 cells,
covering 160 genes, five of which were blank controls. The datasets of mouse olfactory bulb and human breast cancer profiled by HDST
were downloaded from the data portal at https://portals.broadinstitute.org/single_cell/
study/SCP420 [13]. The olfactory bulb dataset contains 3 sections (CN13_D2, CN24_
D1, and CN24_E1) from two mice. We selected a field of CN13_D2 (x:8447~11447, y:
5447.5~6447.5), which consists of 1981 nuclei. In total, 14,909 genes were measured
among these cells, although a majority of them were not detected in each cell. The
breast cancer dataset also includes 3 tissue sections (CN21_E2, CN21_C1, and CN21_
D1), which were obtained from a histological grade 3 HER2+ patient. We selected a
field of CN21_E2 (x:8464~9664, y:5000~6500) consisting of 1765 nuclei. This data, be-
ing very sparse as well, covers 8336 genes. The algorithm of DeepLinc DeepLinc assumes that the neighboring cells in a solid tissue should be much more
likely to have some types of interactions than randomly picked non-neighboring cells Li and Yang Genome Biology (2022) 23:124 Page 18 of 24 that are far away from each other. Therefore, for a particular tissue region, the neigh-
boring cell pairs comprise an incomplete and potentially noisy observation of cell-cell
interactions sampled from the comprehensive cell interaction landscape. In other
words, neighboring cell pairs in tissue sections reveal some cell-cell interactions,
whereas large numbers of other interactions are not explicitly shown. We then assume
that the features related to cell interactions are encoded in the gene expression profile
of each singe cell. DeepLinc uses neighboring cells with direct contacts as the positive set for learning
the transcriptome features related to cell-cell interactions. From a general biological
point of view, it is reasonable to assume that in a solid tissue, most of the cells in 2-D
could directly contact with 3 or more other cells. From a technical point of view, for
the strategies of machine learning, it is critical to minimize the potential false positives
in the predefined positive sets for training. Therefore, for each cell, we only used the 3
nearest neighbors to define direct contacts, which we believe is a balanced choice gen-
erating enough number of direct contacts for training the DeepLinc pipeline, and at the
same time, ensuring few false positives to contaminate the positive training set. DeepLinc leverages a variational graph autoencoder (VGAE) with an adversarial net-
work for regularization, to infer the unobserved interactions between cells. The entire
architecture includes three parts: a variational graph convolutional network (VGCN)
encoder, an inner product decoder, and an adversarial module. Given a cell adjacency matrix A and a gene expression matrix X, the VGAE learns
the embedded features H of the cell graph. A two-layer graph convolutional network
(GCN) (250-125) is used as the encoder module. q HjX; A
ð
Þ ¼
Yn
i¼1q hijX; A
ð
Þ
q hijX; A
ð
Þ ¼ Ν hijμi; diag σ2
Similar to variational autoencoder, μ = GCNμ(X, A) is the matrix of mean vectors and
logσ = GCNσ(X, A). Implementation of DeepLinc DeepLinc is implemented as a Python package, which is available at https://github. com/xryanglab/DeepLinc. Two files are needed as inputs of the pipeline, one for the
cells-by-genes matrix of gene expression data and the other for the predefined cell adja-
cency matrix. If an enrichment analysis of the interactions between cell populations, an
annotation file defining the cell types would be needed. DeepLinc allows users to specify the node numbers. For datasets with more cells, lar-
ger numbers of nodes are recommended to capture the complexity of data better. Al-
though the number of hidden layers for the encoder is also adjustable, extra caution
has to be taken, since deeper graph convolutional networks are prone to over-
smoothing [64]. In general, the two-layer graph convolutional network has been widely
used and proven suitable for most of the tasks. The Adam optimizer [65] with 4e-4
learning rate is used for optimization. The dropout method is also used to ease the
overfitting effect. A new cell interaction matrix is the main output of DeepLinc. The latent representa-
tion output by DeepLinc can also be used for unsupervised clustering of the single cells. Implementation of DeepLinc requires the following packages: Numpy v1.16.4, Scipy
v1.2.1, Pandas v0.20.3, Matplotlib v3.0.2, Seaborn v0.8.1, Networkx v2.1, Scikit-learn
v0.21.2, Tensorflow v1.4.0, Python 3.5.2 (Anaconda3-4.2.0). The DeepLinc model was
trained on a single GPU of GTX 1050 or Tesla K80. The algorithm of DeepLinc As a typical structure, the GCN is defined as GCNðX; AÞ ¼ ~AReLUð
~AXW 0ÞW 1 with weight Wi and normalized adjacency matrix ~A ¼ D−1
2ðA þ IÞD−1
2 to
realize layer-to-layer information transmission. We use ReLU as the activation function
of the first layer instead of the second layer. The purpose of this step is to learn mean-
ingful embedded features supporting the interaction formation by aggregating informa-
tion of each cell itself and its interacting neighbor cells. The inner product decoder p(A| H) = σ(H · HT) is employed to reconstruct a new cell
interaction matrix, which reports probability scores of the interactions between each
pair of single cells. VGAE has exhibited outstanding performance in learning the embedded features of
graph elements [38]. However, for single-cell spatial transcriptome data, a major chal-
lenge is its high sparsity and data noise. For such non-ideal situations, the standard
variational encoder-decoder structure adopted by VGAE does not make more con-
straints on latent variables, often resulting in underfitting of the sparse data [63]. Here
we implement strong regularization on the latent distribution with the adversarial
network. Specifically, the adversarial module is built on a multilayer perceptron (MLP). The
architecture is similar to the discriminator of generative adversarial networks (GAN). Li and Yang Genome Biology (2022) 23:124 Page 19 of 24 Page 19 of 24 The adversarial network has two fully-connected hidden layers with ReLU activation
(125–150). The output layer uses sigmoid function to judge whether a latent sample hi
is from a prior Gaussian distribution or from VGAE. Eventually, the DeepLinc pipeline was optimized to maximize the log-likelihood of
cell-cell interaction network. The objective of VGAE is to minimize its cost function by
optimizing the evidence lower bound (ELBO): ζELBO ¼ Eq HjX;A
ð
Þ logp AjH
ð
Þ
½
−DKL q HjX; A
ð
Þ p H
ð
Þ
k
ð
Þ For the adversarial module, the loss function is binary cross-entropy. For the adversarial module, the loss function is binary cross-entropy. Reconstruction of the cell interaction networks As introduced above, DeepLinc inferred interaction probability scores for all the cell
pairs. Next, cell interaction networks were assembled with the cell pairs with high
probability scores. Specifically, different thresholds of the probability scores were tested
for identifications of cell interactions. The optimal threshold was then determined so
that the newly assembled cell interaction network on the testing set has the highest ac-
curacy for recovering the predefined cell adjacency maps. Evaluating the over- or under-representation of particular types of interactions A previously described strategy [21, 24, 42] was used to evaluate the enrichment or de-
pletion of the interactions between different cell types or within a cell type. In brief, we
randomly shuffled the reconstructed networks for 1000 times. From these 1000 net-
works, the numbers of cell interactions of a certain type, e.g., interactions between two
types of cells, were used to generate a null distribution. The observed interaction num-
ber was then compared to this null distribution, generating a P-value indicating how
often the random values were higher or lower than the observed ones, which corre-
sponds to either enrichment or depletion of the particular type of interactions in the
network. Two-tail test was done for such analysis. Evaluation of the enrichment or depletion of the distal interactions between certain
cell types was done with a similar strategy. For each dataset, we first generate a distance
distribution with all the edges in the reconstructed cell interaction network, from which
a threshold was determined to classify the distal interactions (Additional file 1: Fig. S6). The same number of interactions was established between randomly picked distal cell
pairs. Such procedure was repeated for 1000 times, generating a null background of the
distal cell interaction landscapes. Next, the observed number of distal interactions was
compared to this null distribution, generating a P-value indicating how often the ran-
dom values were higher or lower than the observed one, which corresponds to either
enrichment or depletion of the particular type of distal interactions in the network. Two-tail test was done for such analysis. Benchmark of DeepLinc for recovering missing interactions Different proportions of the edges in the original cell adjacency maps were randomly
picked and discarded. The remaining edges were used for training the DeepLinc model. The area under the receiver operating characteristic (AUROC) was calculated to assess
the accuracy of the reconstructed interaction networks for recovering the discarded
edges. Here equal numbers of the originally non-existing edges were used as true nega-
tive sets. Page 20 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology Benchmark of DeepLinc for differentiating real and fake interactions Different numbers (1 to 10 folds of the size of the original cell adjacency map) of ran-
domly generated fake edges were added into the original cell adjacency map, which was
then fed to DeepLinc. AUROC of the reconstructed network was calculated by taking
the pre-existing real edges as positives and the mixed-in fake edges as negatives. Benchmark of DeepLinc for the tolerance to random noise in gene expression profiles Random noise was independently generated and implemented to each gene in each sin-
gle cell. Specifically, for each gene, the new expression level was regenerated by adding
random noise as follows: Expnoiseþ i; j
ð
Þ
Expnormal i; j
ð
Þ ¼ 2rand;
rand Ν 0; σ
ð
Þ Expnoiseþ i; j
ð
Þ
Expnormal i; j
ð
Þ ¼ 2rand; Among them, Expnoise+(i, j) represents the regenerated expression value of gene j in
sample i, and Expnormal(i, j) represents the original expression level. The fold change
used to simulate noise, after log2 transformation, follows the normal distribution N(0,
σ), whereas different σ values would result in different levels of random noise in
general. Sensitivity analysis for each gene Sensitivity analysis was performed to evaluate the potential contribution of a gene to
shaping the reconstructed cell interaction landscape. In brief, for a particular gene in Page 21 of 24 Li and Yang Genome Biology (2022) 23:124 Page 21 of 24 the transcriptome profile, the expression profile was shuffled among all the single cells. This new data, with the expression of one gene shuffled at a time, was then fed to Dee-
pLinc, which then generates a new cell interaction network. ΔAUPRC was then calcu-
lated by comparing the AUPRC values of the two cell interaction networks built by
DeepLinc with the original unperturbed gene expression data and the data with the ex-
pression of one gene perturbed. For each gene, the process above was repeated for 30
times, and the average ΔAUPRC was used to quantitatively evaluate the sensitivity of
the reconstructed cell interaction network to the gene, thereby defined as the sensitivity
score. For the two high-throughput HDST datasets, distributions of the sensitivity scores of
all the genes were prepared, and the biological processes involving the top-ranked
genes were obtained with the Gene Ontology enrichment analysis [66]. For two low-
throughput datasets of seqFISH and MERFISH, all the genes were sorted by their sensi-
tivity scores, and the Gene Set Enrichment Analysis [67] was performed to acquire the
biological processes enriched by the top-ranked genes. The biological processes were
filtered by gene numbers (min size=3 and max size=500), resulting in 1240 and 1691
gene sets for the GSEA analysis with the seqFISH and MERFISH data, respectively. Prediction of cell interactions with simple machine learning models For the seqFISH and MERFISH datasets, simple machine learning models including
Support Vector Classification (SVC), K-nearest neighbor (KNN), and Gradient Boosting
Decision Tree (GBDT) were used to test the capability of specified genes for classifica-
tions of the interacting and non-interacting cell pairs. The sklearn package in Python
was used to build these models. The originally defined cell adjacency map was used as
positive sets of cell interactions, and randomly selected unadjacent cell pairs, with the
same numbers, were used as negatives. Sixteen genes annotated as ligands or receptors and the same number of top-ranked
signature genes identified by DeepLinc were used for the classification models. Their
expression vectors of a pair of cells were concatenated and used as the feature vectors. The combined set of positive and negative cell interactions were randomly divided into
4:1, serving as the training and testing sets, respectively. The three classification models
were then applied on the data to predict the interacting cell pairs with the testing set. This process was repeated for 30 times with different training and testing set assign-
ments. The AUROC values were calculated based on the predictions. Clustering of the single cells based on the latent representation learned by DeepLinc Dimension of the latent representation H was set as 10 for clustering of the single cells
with the k-means method. The Calinski-Harabasz score was used to determine the op-
timal cluster number. The “umap” package in Python was used for dimension reduction
and visualization of the latent vectors of the single cells. Review history Review history is available as additional file 4. Supplementary Information pp
y
The online version contains supplementary material available at https://doi.org/10.1186/s13059-022-02692-0. The online version contains supplementary material available at https://doi.org/10.1186/s13059 022 02692 0. Additional file 1: Figures S1-S10
Additional file 2: Table S1. Signature genes of the HDST breast cancer data. Additional file 3: Table S2. Signature genes of the HDST mouse olfactory bulb data. Additional file 4: Availability of data and materials y
The DeepLinc pipeline has been deposited in github (https://github.com/xryanglab/DeepLinc) [68] and Zenodo
(https://doi.org/10.5281/zenodo.6564143) [69]. The source code is released under MIT License (http://opensource.org/
licenses). Four published datasets were used for the current study [13, 17, 18], including mouse visual cortex profiled
by seqFISH [17, 70], preoptic region of the mouse hypothalamus by MERFISH [18, 71], mouse olfactory bulb and
human breast cancer by HDST [13, 72]. Funding
h
k g
This work was funded by the National key research and development program, Precision Medicine Project
(2016YFC0906001), the Tsinghua University Spring Breeze Fund, the Tsinghua University Initiative Scientific Research
Program (2019Z06QCX01), and the National Natural Science Foundation of China (81972912 and 31671381). Peer review information Stephanie McClelland was the primary editor of this article and managed its editorial process and peer review in
collaboration with the rest of the editorial team. Authors’ contributions
R L
d X Y
d R.L. and X.Y. conceived and designed the project. R.L. developed the DeepLinc pipeline and conducted the
bioinformatics analyses. X.Y. supervised the whole project. R.L. and X.Y. wrote the manuscript. All author(s) read and
approved the final manuscript. Acknowledgements The authors would like to thank the supports from the Tsinghua University Branch of China National Center for Protein
Sciences (Beijing) and Tsinghua University Technology Center for Protein Research, including the core facilities of
Biocomputing, Genome Sequencing and Analysis at Tsinghua University. Competing interests Competing interests
The authors declare that they have no competing interests. The authors declare that they have no competing interests. Received: 7 November 2021 Accepted: 20 May 2022 Received: 7 November 2021 Accepted: 20 May 2022 Received: 7 November 2021 Accepted: 20 May 2022 Ethics approval and consent to participate
Ethical approval is not needed for this study Ethical approval is not needed for this study. Differential gene expression analysis between single-cell groups For the cells of the same type but distributed in different spatial domains, differential
expression analysis was performed to identify the upregulated genes specifically in each
spatial domain. For each domain, the genes that are significantly upregulated in both Page 22 of 24 Li and Yang Genome Biology (2022) 23:124 Li and Yang Genome Biology one-versus-one and one-versus-rest tests (Mann–Whitney U test, P < 0.01) were
deemed as domain-specific high-expression genes. one-versus-one and one-versus-rest tests (Mann–Whitney U test, P < 0.01) were
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https://pubs.rsc.org/en/content/articlepdf/2021/md/d1md00138h
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English
| null |
Will the chemical probes please stand up?
|
RSC medicinal chemistry
| 2,021
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cc-by
| 12,613
|
a CZ-OPENSCREEN, National Infrastructure for Chemical Biology, Institute of
Molecular Genetics of the Czech Academy of Sciences, Vídeňská 1083, 142 20,
Prague 4, Czech Republic. E-mail: ctibor.skuta@img.cas.cz
b Deanery of Biomedical Sciences, University of Edinburgh, Edinburgh EH8 9XD, UK
† Electronic supplementary information (ESI) available: Full data set used for
the study. See DOI: 10.1039/d1md00138h
‡ Current address: Medicines Discovery Catapult, Alderley Park, Macclesfield
SK10 4TG, UK. Introduction contemporary efforts. These include considerations of the
MLSMR
fitness-for-purpose
as
a
library
accrued
in
an
academic context (compared to arguably better-resourced
pharmaceutical company screening collections), persistent
probe quality issues and remaining confusion on exactly how
many probe compounds the program generated.6–8 In
2005,
the
NIH
Molecular
Libraries
Program
(MLP)
undertook
the
first
large-scale
identification
of
tool
compounds to expand biological insights, now termed small-
molecule chemical probes.1,2 Their systematic generation
against a range of molecular targets was a key driver for the
establishment of the PubChem database in order to collate
structures and data from the initial ten funded screening
centres.3 The concomitant screening compound collection
was established as the Molecular Libraries Small Molecule
Repository (MLSMR) for which PubChem had hosted 255 000
compounds by the end of 2005 and since expanded to 406 000
by 2015 (but updates have ceased). Although 25 of the 64 early
compounds were judged to be of equivocal quality by a
crowdsourcing assessment in 2009 (ref. 4) the program
progressed to 375 probes (see data section below) before
ending in 2014. As conceived from the outset, the availability
of
these
compounds
and,
crucially,
their
associated
characterisation data, have facilitated the exploration of new
targets,
pathways
and
therapeutic
hypotheses.5
Notwithstanding these success stories, the MLP undertaking
has been subject to criticisms that remain relevant to A less tangible but equally important success of the MLP
is that, from approximately 2010 onwards, it inspired other
organisations to also initiate probe discovery with open
dissemination. The three most recent announcements (but
not yet surfacing data) are EU-OPENSCREEN9 with probe
development as one of their main objectives, the EUbOPEN10
consortium aiming to synthesize at least 100 new chemical
probes and the Target 2035 initiative to accrue probes for all
human targets by 2035.11 As of June 2021, we were able to
collect probe data from the sources listed below. • MLP probes (NIH screening initiative) • MLP probes (NIH screening initiative) • Structural Genomics Consortium (SGC, 3D structure-
based)12 • Nathanael Gray Laboratory (cancer research focused)13 • Chemical Probes Portal (literature curation, expert
opinion)14 • Pharmaceutical companies (offering in-house compounds)15 • Probe Miner (data filtration for putative probes)16 • Probes & Drugs (comprehensive collation of probe data)17 Detailed descriptions of these sources and their individual
approaches to probe development are available from their
websites and related publications. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54
This article is licensed under a Creative Commons Attribution 3.0 U Received 23rd April 2021,
Accepted 28th June 2021 Received 23rd April 2021,
Accepted 28th June 2021 DOI: 10.1039/d1md00138h rsc.li/medchem This journal is © The Royal Society of Chemistry 2021 Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Ctibor Škuta,
*a Christopher Southan
‡b and Petr Bartůněk
a In 2005, the NIH Molecular Libraries Program (MLP) undertook the identification of tool compounds to
expand biological insights, now termed small-molecule chemical probes. This inspired other organisations
to initiate similar efforts from 2010 onwards. As a central focus of the Probes & Drugs portal (P&D), we
have standardised, integrated and compared sets of declared probe compounds harvested from 12
different sources. This turned out to be challenging and revealed unexpected anomalies. Results in this
work address key questions including; a) individual and total structure counts, b) overlaps between sources,
c) comparisons with selected PubChem sources and d) investigating the probe coverage of druggable
targets. In addition, we developed new high-level scoring schemes to filter collections down to probes of
higher quality. This generated 548 high-quality chemical probes (HQCP) covering 447 distinct protein
targets. This HQCP collection has been added to the P&D portal and will be regularly updated as
established sources expand and new ones release data. Experimental vs. calculated We chose to internally partition P&D probes into two main
categories; experimental and calculated. Experimental denotes compounds from published probe
characterisation experiments. These papers include profiling
data for target modulation potency, selectivity, and possible
secondary
targets. Also
important
to
note
is
that
the
experimental data largely originate from a single laboratory
and are thus likely to have more consistent and reproducible
data (e.g. where intra-laboratory assay variation is controlled
via sufficient replicates and internal standards). Examples of
such
experimental
probe
sets
include;
bromodomains
chemical
toolbox,37
Chemical
Probes
Portal,14
Gray
Laboratory Probes,13 Nature Chemical Biology Probes, Open
Science Probes,15 opnMe Portal,38 Protein methyltransferases
chemical toolbox,39 and SGC Probes.12 We use the term calculated here to denote in silico
evaluation using the combination of public data with a custom
scoring function (we intentionally avoided the term predicted
in this context because machine learning methods were not
used).16,40 The evaluation of probes at scale involves comparing
public data at different stringencies according to availability. The seven key criteria are: 1) <100 nM target potency in vitro,
2) <1 uM for cell-based assays with evidence of direct target
engagement, 3) target selectivity >100-fold, 4) absence of
structural alerts indicating chemical liabilities 5) identification
of an inactive analogue as control with significantly lower
potency
or
inactive
against
the
primary
target,41
6) an
orthogonal probe with a different chemotype against the same
primary target, and 7) SAR data to increases confidence in
specific target modulation. The simplest approach to assigning
a compound as a probe is to use these criteria (or a subset
thereof). However, this can be nuanced by weightings (e.g. for
potency and selectivity) as well as more complex scoring (e.g. functions favouring compounds with a wider range of target
profiling data). In contrast to experimental probes, the data for
the evaluation of the calculated probes from different sources
can be less consistent and may not be acquired with the
objective of developing and validating a probe per se. The
calculated sources included here are from Probe Miner16 and
the tool compound set.40 P&D
has
additional
advantages
for
this
study. An
important one is that chemical structures from all sources
are standardised after importation. This means that our
internal comparisons are as rigorous as we can make them
(notwithstanding cheminformatic nuances that preclude this
from being perfect). Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Addressing these questions is specifically enabled by the
Probes & Drugs portal (P&D, https://probes-drugs.org).17 P&D
was designed as a hub for the integration of high-quality
bioactive
compound
sets
enabling
their
analysis
and
comparison. As the name indicates, the main focus is on
probes and drugs but includes additional relevant sets
extracted from, or supplied by, recently published specialist
databases (e.g. BiasDB19), vendor sets, ESI† from papers (e.g. kinase inhibitors) or harvested from publication out-links
(e.g. the British Journal of Pharmacology “Concise Guide”
series20–26). Other high-quality curated bioactive chemistry
sources,
including
ChEMBL,27
BindingDB,28
Guide
To
Pharmacology (GtoPdb),29 DrugCentral30 and DrugBank31 are
utilized for compound biological annotation (the latter 3 are
also P&D compound sets). The P&D compound database is
currently composed of 69 sources including 12 probe-related,
7 drug compilations, 36 academic (non-commercial) sets and
14
precompiled
sets
from
bioactive
compound
vendors
(suggestions for expansion are welcome). Experimental vs. calculated We have used the standardiser python
package32
for
salt-striping,
charge
neutralization,
standardizing
common
functional
groups,
preserving
stereochemistry
and
identification
of
the
main
active
pharmaceutical
ingredients
(API)
in
mixtures. Structural
uniqueness is based on the InChIKey, a hashed version of
the International Chemical Identifier, InChI.33 Within the
text of this article, we have used InChIKeys to designate
mentioned compounds. These can be searched not only
against P&D and all major databases but also in Google.34
On our website, users can choose to browse and compare
the structures in three different forms: 1) standardized; 2)
original (i.e. as imported from the source) and 3) non-
isomeric (i.e. core connectivity without stereochemistry). Importantly,
we
made
the
considered
decision
not
to
submit
our
entire
content
to
PubChem
for
two
main
reasons. The first is to reduce non-obvious circularity which
can confound database users.35,36 The second is that this
enables
informative
Boolean
query
combinations
to
be
made between P&D
sets
and
essentially
any
PubChem
source or filtration selects. This journal is © The Royal Society of Chemistry 2021 Introduction These are expanded in a
recent review18 as well as articles in this special issue. This work was conceived to answer the following questions
that can not be answered via the individual sources: • How many declared chemical probe structures are there? 1428 | RSC Med. Chem., 2021, 12, 1428–1441 This journal is © The Royal Society of Chemistry 2021 View Article Online RSC Medicinal Chemistry RSC Medicinal Chemistry Research Article Research Article • What is the distribution of their physicochemical
properties? • What is the distribution of their physicochemical
properties? • What is the distribution of their physicochemical
properties? considered equivalent in a canonical sense because they
have been generated by divergent approaches. By analysing
the structures and associated metadata, we have tried to
establish a high-quality subset based on the probe origin
and by the application of scoring schemes. In addition,
where
the
data
allow
plausible
assignments,
we
have
compiled target coverage. • What are the differences between experimental and
calculated probes? • What is their representation in PubChem sources? • What are their intersections with each other? • What are their individual targets? • What is their combined human proteome coverage? • What is their combined human prot Historical compounds The
use
of
these
obsolete
compounds
is
no
longer
recommended by the CP portal because new data indicates
promiscuity or displacement by better tool compounds.42 These
structures include the well-known and notorious medicinal
chemistry time-wasters of staurosporine (HKSZLNNOFSGOKW-
FYTWVXJKSA-N), quercetin (REFJWTPEDVJJIY-UHFFFAOYSA-N),
resveratrol
(LUKBXSAWLPMMSZ-OWOJBTEDSA-N),
and
curcumin (VFLDPWHFBUODDF-FCXRPNKRSA-N). Set comparisons The sets are compared by exact matches in Table 2. The
resulting matrix is unique in that no individual source has
published a comparable analysis. However, some results were
unexpected. The first surprise was the low intersection
between MLP and the calculated sets. We attribute this to the
80 compounds from the MLP set without bioactivity data on
P&D. In addition, 203 have a primary target potency above
the
100
nM
threshold. The
unexpectedly
high
overlap
between the Chemical Probes Portal and tool compound sets
is a consequence of the (already mentioned) inclusion of the
former in the latter but without data-supported evaluation. Also surprising is the low overlap between the two calculated
sets, even though the data sources and selection criteria were
conceptually similar. However, the main goal of the tool
compound set was to select effective agonists or antagonists
with a high stringency for target selection and cell potency. Bromodomains and protein methyltransferases chemical
toolboxes These compounds were extracted from publications focused
on the study of bromodomains and methyltransferases. Except for four bromodomain probes developed elsewhere,
these also belong to the SGC set. MLP and Nature Chemical Biology Probes MLP and Nature Chemical Biology Probes The former has been outlined in the introduction. The latter
was extracted from articles published in Nature Chemical
Biology (although since 2018 this dedicated section of the
Journal is no longer available) As legacy collections, probes
from these two sources may lack the stringent characterisation
of maintained collections. This is reflected in some cases by a)
the lack of controls or orthogonal probes, b) unclear potency
and selectivity criteria or c) no target annotation. SGC Probes, Open Science Probes, opnMe Portal, and Gray
Laboratory Probes These organisations apply the currently accepted probe
quality criteria and are maintained sets in that P&D has
picked up at least some new compounds since their initial
release (even if at a variable frequency). Tool compound set and Probe Miner These
are
calculated
selections,
mainly
from
ChEMBL,
selected via probe-likeness criteria. While the tool compound
set is a one-off extraction from the publication, Probe Miner
is regularly updated. The tool compound set independently
includes more than 100 compounds available from the
Chemical Probes Portal at the time of its publication. Another surprising observation was that, while overlap
with historical compounds is reassuringly low, Table 2
indicates there are still nine of these undesirables in Probe
Miner, four in the MLP, and two in the high-quality SGC
Probes. The
first
of
these,
bromosporine43
(UYBRROMMFMPJAN-UHFFFAOYSA-N), was designed to be a
pan-bromodomain inhibitor and could usefully be family-
selective. The second, GSK-J1 (ref. 44) (AVZCPICCWKMZDT-
UHFFFAOYSA-N), an inhibitor of the KDM protein family is This journal is © The Royal Society of Chemistry 2021 Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:5
This article is licensed under a Creative Commons Attribution 3.0 U lists historical probes (these are also captured as a P&D set,
see next section). n Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. RSC Medicinal Chemistry
View Article Online View Article Online Research Article RSC Medicinal Chemistry Table 1
Sources with their compound numbers and probe type. “E” refers to experimental and “C” to calculated probe type. The targets column counts
distinct probe-target annotations. The total number of compounds/targets represents a distinct number of compounds/targets for all sets combined Table 1
Sources with their compound numbers and probe type. E refers to experimental and C to calculated probe type. The targets column counts
distinct probe-target annotations. The total number of compounds/targets represents a distinct number of compounds/targets for all sets combined
Set
Probe type
Set class
Compounds
Targets
1
Bromodomains toolbox
E
High-quality
25
26
2
Chemical Probes.org
E
High-quality
362
322
3
Gray Laboratory
E
High-quality
53
56
4
MLP
E
Legacy
375
156
5
Nature Chemical Biology
E
Legacy
58
51
6
Open Science Probes
E
High-quality
83
95
7
opnMe Portal
E
High-quality
55
57
8
Probe Miner
C
Calculated
3187
326
9
Methyltransferases toolbox
E
High-quality
19
20
10
SGC Probes
E
High-quality
81
97
11
Tool compound set
C
Calculated
515
392
12
Historical compounds
E
Historical/obsolete
239
—
Total
(Not historical)
4466
819 Source descriptions and counts As of version 02.2021 (March 2021), P&D contains 77 130
compounds with 4466 labelled as chemical probes by their
sources used for this study. However, these should not be Those compared in this study are listed in Table 1 with brief
descriptions below. This journal is © The Royal Society of Chemistry 2021 RSC Med. Chem., 2021, 12, 1428–1441 | 1429 RSC Medicinal Chemistry
Research Article
View Article Online Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54
This article is licensed under a Creative Commons Attribution 3.0 Un not cell-permeable. The SGC Probes resource has noted this
and consequently now recommends a pro-drug of GSK-J1,
GSK-J4
(WBKCKEHGXNWYMO-UHFFFAOYSA-N)
for
cell-
based assays. Fig. 1
Physico-chemical
properties
distribution
(top:
molecular
weight, middle: calculated log P, bottom: TPSA) for experimental
probes
(blue),
calculated
probes
(orange)
and
small-molecule
approved drugs set from ChEMBL (green) (x: property range, y:
compounds percentage). The properties were calculated by RDKit. Dataset compilation Merging individual sets resulted in 4466 structurally distinct
probe compounds (i.e. as unique InChIKeys). This includes
940 (21%) experimental plus 3670 (82.2%) calculated probes
including 3178 from Probe Miner. The overlap of 143 (3.2%)
compounds is mainly due to the inclusion of the Chemical
Probes Portal in the tool compound set. The full 4466 set
includes 275 labelled as drugs (reported to be in clinical
phases) with 103 labelled as approved by FDA, EMA and
other agencies. The full set also includes 29 PROTACs
(Proteolysis Targeting Chimeras) from PROTAC-DB45 and
Chemical
Probes
Portal,
60
covalent
binders
from
CovalentInDB,46 and 21 biased GPCR ligands from BiasDB.19 Our analysis also established that 132 compounds were
flagged with one or more structural alerts from either a)
PAINS filters,47,48 b) aggregators,49 c) cellular assay nuisance
compounds50
or
d)
historical
compounds. Of
the
60
stereoisomers,
54
originate
from
the
Probe
Miner
set. Compared to small-molecule approved drugs extracted from
ChEMBL (as a set in P&D) probes are generally larger and
more complex (Fig. 1). This correlates with higher target selectivity that, in turn,
is reflected in the number of associated targets (Fig. 2). However, these values could be biased by approved drugs
accumulating more cross-screening data and hence a wider
range of secondary targets. Fig. 1
Physico-chemical
properties
distribution
(top:
molecular
weight, middle: calculated log P, bottom: TPSA) for experimental
probes
(blue),
calculated
probes
(orange)
and
small-molecule
approved drugs set from ChEMBL (green) (x: property range, y:
compounds percentage). The properties were calculated by RDKit. This journal is © The Royal Society of Chemistry 2021 Chemical Probes Portal (CP portal) This
provides
expert
usage recommendations
based
on
publication evaluations from a Scientific Advisory Board
(SAB, of which one of us, CS, is a member). While content
had languished below 200 compounds for some time, this
has recently expanded to 362. Importantly, this portal also This journal is © The Royal Society of Chemistry 2021 1430 | RSC Med. Chem., 2021, 12, 1428–1441 1430 View Article Online RSC Medicinal Chemistry Table 2
A matrix showing the intersections between 12 sources. This was computed using the InChIKey exact match for the standardised structures
from the P&D portal. The diagonal figures in white represent the source counts in Table 1 Table 2
A matrix showing the intersections between 12 sources. This was computed using the InChIKey exact match for the standardised structures
from the P&D portal. The diagonal figures in white represent the source counts in Table 1 Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. RSC Medicinal Chemistry Ki, IC50 or Kd). Some may also have
secondary targets with a data-supported potency below that
against the primary target (we have avoided using the term
“off-target” since there are few cases where secondary targets
have been mechanistically assigned as a side-effect or toxicity
liabilities). The 132 probes without primary target annotation
were, in most cases, directed against viruses, bacteria, cell
lines or pathways. They also predominantly belonged to the
MLP and Nature Chemical Biology legacy sets. For the
remainder, we collated 819 single and multi-component
protein targets with 549 for both experimental and calculated
with an overlap of 279. In total these constituted 807 distinct
single
protein
identifiers
(i.e. UniProt
IDs51),
544
for
experimental and 535 for calculated probes with 272 in-
common. The human Swiss-Prot target count was 796. The differences in Fig. 3 reflect inherent biases. For
example, predicted probes have almost double the number of
GPCRs52 but, compared to predicted probes, proportionally
fewer kinases and epigenetic regulators. This is likely to be
due to the challenges of optimising single-target selective
ligands within these target families.37,53 However, probes
with intra-family selectivity can also be experimentally useful
but run the risk of being rejected by quality scoring weighted
towards single-target selectivity. RSC Medicinal Chemistry RSC Medicinal Chemistry there are several length forms in TrEMBL but not Swiss-Prot)
or ABL1 (P00519) or, in the case of ∼20 probes from the
Probe Miner set, BCR (P11274). Other complicating examples
are the cyclin-dependent kinases (CDKs), where the probes
may be annotated by sources with one of the human CDKs, a
CDK in complex with a specific cyclin, or both. The highest probe target numbers (5 experimental and
225 calculated) have been assigned against mTOR (MTOR,
P42345). Next in rank are histone deacetylase 1 (HDAC1,
Q13547)
with
2/161
experimental/calculated,
epidermal
growth
factor
receptor
(EGFR,
P00533)
with
7/103
and
estrogen receptor (ESR1, P03372) with 1/92. The larger
number for calculated probes reflects their more frequent
origin from panel screening papers and consequent higher
average compounds: target ratio of 9.8 for the Probe Miner
set compared to ∼1.0 for experimental probes. For these, the
highest assigned target numbers are BRD4 (O60885) and the
BRD3/BRD2 (Q15059/P25440) subfamily pair with 18 and 15
probes, respectively. This is a consequence of the declared
SGC Probes focus on epigenetic regulators. The family
distribution of all annotated targets is shown in Fig. 3. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Fig. 2
The number of associated targets for probes (blue) and the
small-molecule approved drugs set from ChEMBL (green) (x: axis
number of targets, y: compounds percentage). Only compounds with
at least one associated target were included encompassing 4257
probes and 2005 approved drugs. in vitro binding data (e.g. Ki, IC50 or Kd). Some may also have
secondary targets with a data-supported potency below that
against the primary target (we have avoided using the term
“off-target” since there are few cases where secondary targets
have been mechanistically assigned as a side-effect or toxicity
liabilities). The 132 probes without primary target annotation
were, in most cases, directed against viruses, bacteria, cell
lines or pathways. They also predominantly belonged to the
MLP and Nature Chemical Biology legacy sets. For the
remainder, we collated 819 single and multi-component
protein targets with 549 for both experimental and calculated
with an overlap of 279. In total these constituted 807 distinct
single
protein
identifiers
(i.e. UniProt
IDs51),
544
for
experimental and 535 for calculated probes with 272 in-
common. The human Swiss-Prot target count was 796. in vitro binding data (e.g. RSC Medicinal Chemistry
View Article Online View Article Online View Article Online Fig. 2
The number of associated targets for probes (blue) and the
small-molecule approved drugs set from ChEMBL (green) (x: axis
number of targets, y: compounds percentage). Only compounds with
at least one associated target were included encompassing 4257
probes and 2005 approved drugs. Research Article RSC Medicinal Chemistry Target mapping The majority of probes have primary targets specified in their
sources. In most cases these are supported by quantitative RSC Med. Chem., 2021, 12, 1428–1441 | 1431 This journal is © The Royal Society of Chemistry 2021 Target intersections in UniProt In practice, the number of protein targets is below 819,
since multi-component targets may be variably annotated
against either a protein subunit, the complex target, or both. For example, probes directed against the BCR–ABL1 fusion
protein may be annotated with the fusion protein (of which Having assigned target IDs to both probe sets we compared
these with informative cross-references in UniProt.51 We
selected 4213 protein IDs (as human Swiss-Prot entries)
based on the union (OR operator) of the four high-quality Fig. 3
A bar chart showing the target families distribution separately for all (green), experimental (blue) and calculated (orange) probes (x: target
family, y: number of targets). The assignments are based on the ChEMBL and Guide to Pharmacology target classification. Fig. 3
A bar chart showing the target families distribution separately for all (green), experimental (blue) and calculated (orange) probes (x: target
family, y: number of targets). The assignments are based on the ChEMBL and Guide to Pharmacology target classification. 1432 | RSC Med. Chem., 2021, 12, 1428–1441 This journal is © The Royal Society of Chemistry 2021 This journal is © The Royal Society of Chemistry 2021 Research Article
View Article Online View Article Online RSC Medicinal Chemistry Research Article Research Article curated chemistry-to-target databases (already mentioned) of
ChEMBL, BindingDB, GtoPdb and DrugCentral. These IDs
represent liganded targets for 21% of the UniProt proteome
of 20 395 (although this drops to 19 205 for HUGO Gene
Nomenclature
Committee
annotation). The
comparative
protein sets we also selected were from the four target
development
levels
(TDLs)
of
the
Pharos
resource
for
Illuminating the Druggable Genome (IDG).54 This facilitates
exploration of both the characterised and the understudied
(or “dark”) regions of the human proteome with a view to
expanding functional insights and finding new drug targets. We initially selected the combination of the Tchem54 (1593
proteins)
known
to
bind
small
molecules
(other
than
approved drugs) with target-class specific potency thresholds
plus the Tclin54 (659 proteins) as targets of approved drugs. The union of these two is 2221 proteins. The intersections of
these four lists are shown in Fig. 4. • Concentration unit errors for secondary target activity
(i.e. the compound was thus not selective). • Erroneous target annotations that falsely indicate potent
secondary target activity. • Potency values assigned to only a subunit in a multi-
component target (i.e. thus technically without supporting
bioactivity data). ess Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. As an operating principle P&D fixes any unequivocal errors
we spot. At the same time, we notify the originating sources
about
these
errors
that
could
otherwise
persist
and
proliferate between databases. However, we have found the
speed with which these are fixed at source has been variable
(although this is clearly dependent on build cycle times and
release versions). The two notable features are: The two notable features are: Probe scoring schemes 1. Probes have activity against 53 proteins not in Tclin
or Tchem. We optimised four different scoring schemes to support users
for probe triage and selection. As explained, the Probe Miner
(PMIS)
and
P&D
probe-likeness
scores
(PDPS)
are
data-
supported. The other two scores are expert opinion-based and
thus more subjective. These are abstracted from the Chemical
Probes Portal rating for use in cells (CPOC) and in organisms
(CPOO). These
represent
a
summed
rating
of
chemical
properties, primary targets, secondary targets and in most
cases, expert judgments. The conceptual difference with data-
supported scores is that these are calculated for compound-
target pairs. Thus, a single compound can have multiple scores
against each of its assigned targets. Users may thus select the
most suitable of these pairings but the probes can also be used
for intra- or inter-family selectivity. For comparability, all scores
are normalized to between 0 and 100%. 2. The Swiss-Prot liganded proteome includes 743 probe
targets. Analysis with other TDL sets established that of the 659
approved targets 265 were also covered by probes. However,
there were no intersections between probes and the 6368
Tdark proteins. This implies that the current probes may
have already expanded Tchem but there is no overlap with
dark targets. This journal is © The Royal Society of Chemistry 2021 RSC Med. Chem., 2021, 12, 1428–1441 | 1433 Target intersections in UniProt • Some sources had incorrectly assigned human TrEMBL
partial sequence entries as targets rather than the human
Swiss-Prot IDs (although only three cases were found). Source errors Despite curatorial diligence, low levels of annotation errors,
including
the
transitive
inheritance
of
author
mistakes
extracted from papers, inevitably creep into the bioactivity
databases we have used as sources.55 During manual cross-
checking we identified the following error types affecting
approximately 40 probe entries: The PMIS, ranging from 0 to 1, combines partial scores for
1) potency, 2) selectivity 3) activity in cells, 4) SAR data 5)
availability of an inactive analogue and 6) a PAINS score. Nevertheless, probe suitability is not prescribed by the score
value but by so-called minimum quality criteria. These include
100 nM potency in biochemical assays, 10-fold target selectivity,
and 10 μM potency in cell-based assays (but not necessarily
evidence of intra-cellular primary target engagement). The
Probe Miner set of 3187 compounds meeting these criteria thus
have
a
PMIS
between
0.38 and
0.85. A
more detailed
description is given in the Probe Miner publication.16 • Substitution of a biochemical for a cell-based assay as
well as vice versa cases (the most common problem). Fig. 4
Venn diagram of P&D targets against selected human UniProt
cross-references and two Pharos TDLs. The PDPS, scaled from 0 to 1, incorporates partial scores
in common with PMIS but adds in orthogonal probes. The
comparison of both scoring schemes is shown in Table 3. Unlike the Probe Miner selectivity score, P&D also highlights
target sub-family selectivity beyond just single proteins. The
probe-likeness of a compound is closely related to the PDPS
value. Each compound with a score above 0.7 is labelled as
P&D-approved based on the available data. The score is
capped to not exceed 0.7 unless it passes all three core
criteria (i.e., in vitro potency, cell potency and selectivity). Fig. 4
Venn diagram of P&D targets against selected human UniProt
cross-references and two Pharos TDLs. RSC Med. Chem., 2021, 12, 1428–1441 | 1433 This journal is © The Royal Society of Chemistry 2021 Table 3
Comparison of PMIS and PDPS. For parameters with a defined range, the score is 0% for values below the minimum and 100% for values
greater than the maximum. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. —
Binary (1 and −3 for historical compounds) PAINS,
aggregators + other nuisance compounds in cellular assays,
historical compounds
Compounds labelled as P&D approved for PDPS >70% 1109 probe with low ratings from the Chemical Probes Portal (not
scored
by
Probe
Miner),
AGI-5198
(ref. 59)
(FNYGWXSATBUBER-UHFFFAOYSA-N), was proposed as a
prototypical
IDH-1
R132H
inhibitor. However,
this
was
surpassed in potency and characterization details by the
more
recent
GSK864
(ref. 60)
(DUCNNEYLFOQFSW-
PMERELPUSA-N) which also has an inactive control for proof
of
target
involvement. However,
as
a
second,
distinct
chemotype, this probe could be used for corroborative
phenotypic assays. Compounds labelled as historical are down-weighted by
subtracting 0.3 and thus cannot be labelled as P&D-approved. Currently, there are 1109 probes labelled as P&D approved. More details on this are given on the P&D FAQ page.56 The
CPOC and CPOO scores (from 0 to 4 stars) are based on
Scientific
Advisory
Board
(SAB)
reviews. These
may
be
accompanied by comments and usage recommendations. However, there is currently a review backlog in that out of
362 compounds, 274 have been rated for use in cells and 225
for use in model organisms. Compounds labelled as historical are down-weighted by
subtracting 0.3 and thus cannot be labelled as P&D-approved. Currently, there are 1109 probes labelled as P&D approved. More details on this are given on the P&D FAQ page.56 The
CPOC and CPOO scores (from 0 to 4 stars) are based on
Scientific
Advisory
Board
(SAB)
reviews. These
may
be
accompanied by comments and usage recommendations. However, there is currently a review backlog in that out of
362 compounds, 274 have been rated for use in cells and 225
for use in model organisms. In Table 4, we review three examples of compounds with
assigned probe scores. The first is a selective RIPK1 inhibitor,
GSK2982772 (ref. 57) (LYPAFUINURXJSG-AWEZNQCLSA-N),
highly scored by all three probe sources. The second is BET
family bromodomain inhibitor from SGC Probes, JQ-1 (ref. 58) (DNVXATUJJDPFDM-KRWDZBQOSA-N), scored highly by
P&D and Chemical Probes Portal, but as a family-selective
probe with lower PMIS. The latter is the only P&D approved Source errors Within this range, there is a linear relationship between the value and the score
Parameter
PMIS value range (weight) note
PDPS value range (weight) note
Potency
(biochemical)
5–10 [−log(M)] (4)
6.5–7 [−log(M)] (2)
Selectivity
Complex selectivity score normalized per target (8)
10–30-Fold (2)
Potency
(cell-based)
5 [−log(M)] (2) without the evidence of primary target engagement
5.5–6 [−log(M)] (2) with the evidence of primary target
engagement
Inactive
analogue
Binary (1)
Binary (1)
Orthogonal
probe
—
Binary (1)
SAR
Binary (1)
—
Structural alert Binary (1) PAINS
Binary (1 and −3 for historical compounds) PAINS,
aggregators + other nuisance compounds in cellular assays,
historical compounds
Probe-likeness
determination
Independent of the score value, compounds labelled as possible
suitable probes if they meet minimum quality criteria (100 nM
potency, 10 μM cell potency, 10-fold selectivity)
Compounds labelled as P&D approved for PDPS >70%
Probe-like
compounds
count
3187
1109
RSC Medicinal Chemistry
Research Article
View Article Online Research Article RSC Medicinal Chemistry Table 3
Comparison of PMIS and PDPS. For parameters with a defined range, the score is 0% for values below the minimum and 100% for values
greater than the maximum. Within this range, there is a linear relationship between the value and the score High-quality chemical probes set Table 5
Matrix showing the intersections between six different probe
scores and probe types. This was computed using the InChIKey exact
match for the standardised structures from the P&D portal As an outcome of this study, we have compiled a high-quality
chemical probes subset (HQCP). We have used the PDPS for
the addition of P&D approved experimental probes plus those
P&D approved calculated probes that are in at least one
established tool compound set. We have thus partitioned
four compound sets from P&D: 1. Concise Guide to Pharmacology 2019/20 is a set extracted
from a biennial series of publications providing concise
overviews of the key properties of ∼1800 human drug targets
with an emphasis on selective pharmacology.20–26 Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. 2. Kinase chemogenomics set is a collection of narrow-
spectrum small molecule kinase inhibitors assembled by
the SGC-UNC to study the biology of dark kinases. This is
the most diverse and highly annotated public collection of
kinase inhibitors.61 3. Kinase
inhibitors
were
extracted from
a series
of
Molecular Cell papers by Wang and Gray summarising
recently-reported kinase inhibitors.62,63 since the highest value is 85%, a setting of 75% would leave
only 54 compounds from more than 3000. We thus chose to
set the Probe Miner threshold at 60%, thus leaving 1282
compounds, a similar number to P&D approved probes. 3. Kinase
inhibitors
were
extracted from
a series
of
Molecular Cell papers by Wang and Gray summarising
recently-reported kinase inhibitors.62,63 4. Novartis Chemogenomic Library – NIBR MoA Box was
compiled via data mining and institutional crowdsourcing. It
is regularly updated and used widely both within Novartis
and by their external collaborators.64 4. Novartis Chemogenomic Library – NIBR MoA Box was
compiled via data mining and institutional crowdsourcing. It
is regularly updated and used widely both within Novartis
and by their external collaborators.64 The
comparison
between
different
scoring
schemes
(Table 5) highlights differences between judgment-based and
calculated scores. One of the reasons for these differences is
data incompleteness, for example where affinity data is only
for
the
presumed
primary
target
thereby
precluding
selectivity assessment. We used the quality criteria in Table 6 to select 548 probes
for HQCP (451 experimental, 208 calculated with 114 in
common). Score comparisons To
extend
our
systematic
comparison
of
scoring,
we
introduced quality thresholds. PDPS was set at 70% as used
for P&D approved probes. For CPOC and CPOO, we raised
this to 75% (equivalent to 3 out of 4 stars in the original
rating system). For the PMIS, there is no clear threshold and Table 4
Three selected chemical probes with assigned probe scores from P&D, Probe Miner and Chemical Probes Portal Three selected chemical probes with assigned probe scores from P&D, Probe Miner and Chemical Probes Portal Table 4
Three selected chemical probes with assigned probe scores from P&D, Probe Miner and Chemical Probes P probes with assigned probe scores from P&D, Probe Miner and Chemical Probes Portal Table 4
Three selected chemical probes with assigned probe scores from P&D, Probe Miner and Chemical Probes Portal
Name
GSK2982772
JQ-1
AGI-5198
PDPS
100%
100%
86%
PMIS
70% (in Probe Miner set)
48% (not in Probe Miner set)
—
CPOC
100%
100%
50%
CPOO
83%
75%
42% 1434 | RSC Med Chem 2021 12 1428–1441
This journal is © The Royal Society of Chemistry 2021
Table 4
Three selected chemical probes with assigned probe scores from P&D, Probe Miner and Chemical Probes Portal
Name
GSK2982772
JQ-1
AGI-5198
PDPS
100%
100%
86%
PMIS
70% (in Probe Miner set)
48% (not in Probe Miner set)
—
CPOC
100%
100%
50%
CPOO
83%
75%
42% This journal is © The Royal Society of Chemistry 2021 Research Article
View Article Online Research Article
View Article Online View Article Online RSC Medicinal Chemistry Research Article Table 6
The criteria used for the selection of HQCP. The count column contains the number of compounds matched by the criterion. The total
number represents the union of all criteria number represents the union of all criteria
Criterion
Count
1 Belong to one of the high-quality probe sets (except Chemical Probes Portal)
256
2 CPOC or CPOO score at least 75% (i.e. three out of four stars in the original Chemical Probes Portal rating system)
208
3 P&D approved experimental probes
150
4 P&D approved probes belonging to one of the non-commercial high-quality sets (Concise Guide To Pharmacology, Kinase
Chemogenomic Set, Kinase Inhibitors, and Novartis Chemogenetic Library)
177
5 Not labelled as a historical compound
−2
Total
548 This journal is © The Royal Society of Chemistry 2021 High-quality chemical probes set The intersections are shown in Table 7 including
the EU-OPENSCREEN Bioactive Compound Library and Drug
Repurposing Hub65 set. As non-commercial bioactive libraries,
these are included in P&D as relevant for probe research. Even the experimental probes included 195 compounds
without
bioactivity
data. In
addition,
we
found
142
compounds
annotated
against
single
targets
without
selectivity data. The union of these represents 36% of the
experimental probes that cannot thus be properly scored. Another reason for differences arises between stringent
criteria-based evaluation and expert judgment. The HQCP set contains 42 approved drugs with 102
clinical candidates, 27 PROTACs, 15 covalent binders and 10
compounds
tagged
with
a
structural
alert
(four
for
aggregation and six for PAINS). The overlap between HQCP
and the calculated sets is largest for the P&D approved
probes with 244 out of 1109 compounds, but these were also
partly used for the HQCP selection. From the complete Probe
Miner set (i.e. without the PMIS threshold applied), there are
66 compounds from nearly 3200 meeting the Probe Miner
minimum quality criteria. On the other hand, there are 153
compounds from 515 in the tool compound set, mainly from
the inclusion of the Chemical Probes Portal compounds in
the tool compound set. We also found differences between calculated scores for
the 265 compounds-in-common between the 1109 (P&D) and
1282 (Probe Miner) sets. This could be attributed to the
differences
in
the
scoring
methodology
but
also
the
associated data (i.e. not all experimental probes have PMIS). While Probe Miner currently employs bioactivity data from
ChEMBL and BindingDB, P&D uses more recent versions and
complements these with smaller data sources such GtoPdb. This is reflected in a high P&D score for 150 experimental
probes (with 55 highly-rated by CPOC) while Probe Miner
detects 20 (including 10 based on the CPOC). The intersections between the three bioactive screening
libraries (Novartis Chemogenetic Library, EU-OPENSCREEN RSC Med. Chem., 2021, 12, 1428–1441 | 1435 This journal is © The Royal Society of Chemistry 2021 View Article Online Research Article RSC Medicinal Chemistry Table 7
Matrix showing the intersections between HQCP and other selected sets. This was computed using the InChIKey exact match for the
standardised structures from the P&D portal Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. High-quality chemical probes set first part of this necessitated the mapping of all P&D probe
structures to PubChem CIDs via InChIKey matches and
SMILES strings for cross-corroboration. This was done using
the PubChem Identifier Exchange Service.67 We expected
high coverage from the probe sources which we knew to have
entered
PubChem
by
various
routes. From
the
940
experimental probes, we recorded 915 CID matches (910 from
IKs
plus
five
more
from
SMILES). Inspection
of
the
unmapped probes confirmed that most from SGC Probes,
opnMe Portal and Gray Laboratory had no submission path
into PubChem (directly, or via other source). In addition, we
found the three unmatched MLP probes had different or
flattened stereochemistry in PubChem (i.e. matched different
non-isomeric
CIDs). The
corresponding
3670
calculated
probes matched 3557 CIDs. While the mismatches were still
only 3%, the reasons behind these are (again) differences in
the handling of stereochemistry between PubChem and P&D
(the
latter
uses
RDKit
as
its
main
cheminformatics
framework68). We also discovered that the links to some
compounds are missing from ChEMBL because of InChIKey
differences (ChEMBL is also using the RDKit framework69). Bioactive Compound Library, Drug Repurposing Hub) are
5.0%, 7.5% and 2.8%, respectively. For target assessment, the HQCP covers 447 distinct
proteins. The
distribution
of
target
families
for
all,
experimental, calculated and HQCP probes is shown in
Table 8. The HQCP was added as a separate compound set to
the P&D portal and will be updated regularly. As new
bioactivity data and new versions of compound sets are
integrated we expect the number to increase. This journal is © The Royal Society of Chemistry 2021 1436 | RSC Med. Chem., 2021, 12, 1428–1441 Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. The 67% inclusion in the MLSMR means these particular
probes may have expanded profiling data from unpublished
assays not captured by ChEMBL, including testing against
malaria, other disease parasites and cancer cell lines (this is
particularly the case for the older MLP compounds). The
extensive data overlap between ChEMBL and BindingDB arises
from their mirroring collaboration but the latter has unique
content from patent SAR extractions. For some years GtoPdb
has included probe curation from papers selected for their
pharmacological relevance and this is reflected in the capture
of 305 probes.29 The availability of a PDB ligand structure for
242 probes is clearly enabling for many reasons but note these
may not all be for the probe-primary target pair or species. The
explanation for the low hits to the Chemical Probes Portal was
the inclusion of historical probes in their 2017 PubChem
submission (we suggest the separate submission of this
cautionary subset in the future). The last row in the table
presents two anomalies. As discussed above, at least 100
additional nominal MLP probes can be found in various lists
beyond the 223 in the PubChem CID select for “BioAssay,
Probes”.7 While reasons for the low match in P&D are being
investigated the historical confusion associated with legacy
MLP compounds may confound explanation. PubChem (note also the opnMe probes are free upon
application). The 770 matches in ChEMBL indicate high
levels of probe-target activity data extracted from papers. However, there is an unexpected shortfall of 115 probes
without any active results in BioAssay. The explanations are
either the probe generators have not published in their assay
results
or
these
were
not
in
journals
that
ChEMBL,
BindingDB
or
the
Guide
to
Pharmacology
would
have
extracted and then submitted to PubChem. PubChem (note also the opnMe probes are free upon
application). The 770 matches in ChEMBL indicate high
levels of probe-target activity data extracted from papers. However, there is an unexpected shortfall of 115 probes
without any active results in BioAssay. The explanations are
either the probe generators have not published in their assay
results
or
these
were
not
in
journals
that
ChEMBL,
BindingDB
or
the
Guide
to
Pharmacology
would
have
extracted and then submitted to PubChem. The fact that 71% of the experimental probes have patent
matches
was
a
surprise
since
the
impact
of
potential
Intellectual Property (IP) issues on probe usage has not been
widely discussed. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. While this high proportion seems at odds
with the Open Science context that probe development teams
espouse, the matches only mean the structures are specified
in patent documents rather than necessarily being within the
scope of allowed claims. Many of the automated extractions
may merely represent prior-art mentions including where
applicants have exploited analogue expansions from existing
probe
structures
as
drug
discovery
starting
points. Notwithstanding, some probe structures may be explicitly
claimed
in
maintained
and
granted
patents
(although
precisely how many is difficult to assess). However, open
patent information has become increasingly available and
compound-to-patent document mappings are now indexed
for
nearly
40
million
PubChem
CIDs.70
An
interesting
example is the Boehringer opnMe GPR142 agonist BI-1046
(MLOGCHDCTRINMU-UHFFFAOYSA-N). Two
sources
in
PubChem have extracted the structure from Boehringer's
WO2020007729 “Triazole benzamide derivatives as GPR142
agonists”. From
CID
146293963
(via
SureChEMBL
SID
405725530), we can map the structure to example 2 and a
table of low nM IC50 SAR values for 20 analogues (with
synthesis details) that can also be found in the PubChem
“Similar Compounds” section. The explanations for the calculated probes are the same
as for the experimental but show a different pattern in the 11
rows of Table 9. Since these are predominantly derived from
ChEMBL, the matches against this source, BioAssay active
and BindingDB are all high. In contrast, vendor matches are
proportionally much lower. While the patent intersection
drops to 61% this still impacts 2227 CIDs. The explanation
lies in the fact that many of the organisations (academic or
commercial) generating the medicinal chemistry papers that
ChEMBL curates (and Probe Miner selects) also file patents
on their characterised compounds in advance of publication. Notwithstanding potential IP complications, it is important
to note that patent matches are potentially advantageous for
probe evaluation because they may well contain unpublished
selectivity and SAR data not captured in probe sources.70 This journal is © The Royal Society of Chemistry 2021 PubChem intersections As the de facto global hub for chemical structures, associated
bioactivity
data
and
a
massive
range
of
informatic
connectivity it was of considerable interest to profile probe
sets against the 110 million compounds in PubChem.66 The 1436 | RSC Med. Chem., 2021, 12, 1428–1441
Table 8
The target families distribution separately for all, experimental,
calculated and HQCP probes
Target family
All
Experimental
Calculated
HQCP
Kinase
201
179
146
152
GPCR
146
65
115
70
Hydrolase
98
59
54
36
Epigenetic regulator
75
73
44
68
Transferase
39
28
23
26
Ion channel
34
22
21
16
Oxidoreductase
33
10
27
8
Transporter
29
11
22
12
Nuclear receptor
26
14
20
15
Cytochrome P450
9
2
9
3
Isomerase
8
6
3
2
Ligase
5
3
5
2
Lyase
5
0
5
0
Other
111
77
55
37
Total
819
549
549
447 Table 8
The target families distribution separately for all, experimental,
calculated and HQCP probes The second part of this analysis compared the two probe
sets with selected PubChem sources to give additional
insights. The numbers, shown in Table 9, are, again, a
mixture of the expected and unexpected. We can propose
explanations, starting with the experimental probes. The high
level of BioAssay positive results is expected but does not
establish whether those are the same probe-target pairs
annotated in P&D. From the 915 CID matches in PubChem, 784 (85%)
include vendors submissions, indicating a high availability
for purchase. However, this expands slightly since additional
vendor matches internal to P&D may not be indexed in This journal is © The Royal Society of Chemistry 2021 View Article Online RSC Medicinal Chemistry Research Article Table 9
CID intersections between experimental and calculated probes
for selected PubChem sources, ranked by the number of experimental
probe matches. Note that most of these results can alternatively be read
off directly from the P&D portal and give the same or close numbers. The
total column represents a number of distinct compounds in the
respective sources patent holders can only be addressed on a case-by-case basis. The assumption of Research Use Exemption should apply to US
academics but the position of commercial institutions is less
clear.71 Note, however, despite the detailed data package in the
opnMe portal, the absence of a publication (BI-1046 is PubMed-
negative but has over 40 false positives in Google Scholar
because of an HIV clinical trial designation BI 1046) means that
CID 146293963 has neither ChEMBL nor BioAssay links. Source
Total
Experimental
Calculated
PubChem
109 818 005
915
3557
BioAssays – active
1 457 929
800
3487
Vendors
59 867 622
784
810
ChEMBL
2 067 192
770
3519
Patents
39 401 959
652
2227
MLSMR
406 097
622
416
BindingDB
975 228
608
3331
GtoPdb
8705
305
335
PDBe
33 543
242
287
Chemical Probes Portal
467
186
131
BioAssays Probes
223
152
2 Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Discussion This
work
provides
a
uniquely
comprehensive
and
comparative overview of probe sources and targets. This will
be
maintained
and
expanded
for
experts
and
non-
informaticians seeking probes to use in their work. Although
our results are presented in good faith, we understand the
causes of fuzziness (some of which have been discussed) that
caution against these numbers being taken as ground truth. While the opnMe portal magnanimously declares that
results generated with their molecules belong to the ordering
scientists, the IP situation regarding other probe structure RSC Med. Chem., 2021, 12, 1428–1441 | 1437 This journal is © The Royal Society of Chemistry 2021 RSC Medicinal Chemistry
View Article Online View Article Online View Article Online Author contributions CŠ collated the data with CŠ and CS being responsible for
the analysis. All authors edited and approved the final
manuscript. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM.
This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. Open Access Article. Published on 16 July 2021. Downloaded on 10/24/2024 4:54:43 AM. This article is licensed under a Creative Commons Attribution 3.0 Unported Licence. 5. There
are
many
selective
and
potent
compounds
appearing in the recent medicinal chemistry and chemical
biology literature that, while not officially yet declared in probe
sources, include sufficient characterisation for useful probe
criteria scoring. However, the rate of data extraction from these
publications and flow into databases remains slow.72 6. We are considering how to address the mismatched
and missing probes in PubChem but we need to iterate with
the originating sources in the first instance. FAIR and reproducible This situation
can
obviously
be
ameliorated by generating more data but, going forward, it is
not clear how the existing data gaps can best be backfilled. 2. Comprehensive characterisation of a probe, including
the
necessary
broad
cross-screening,
requires
extensive
experimental work. In addition, the results need to be
accessible
(ideally
in
an
open-access
text-minable
publication), reproducible and easily captured for transfer
into database
records. This situation
can
obviously
be
ameliorated by generating more data but, going forward, it is
not clear how the existing data gaps can best be backfilled. FAIR and reproducible We have endeavoured to make this work findable, accessible,
interoperable
and
reusable,
according
to
Open
Science
principles.73 As for P&D itself, the licence is CC BY-SA 4.0. We
have submitted a ESI† sheet that includes compound names,
SMILES, InChIKeys, target identifiers, source assignments as
well as other data used for the study. For interoperability, this
will be deposited into Figshare in .xlsx format. No proprietary
software has been used in this work and we thus expect any
analysis reported here to be reproducible (if users encounter
difficulties, they are welcome to contact us). As mentioned, all
the intersections between sources inside the P&D database can
simply be read off, combined, downloaded and users' own sets
uploaded for further intersection analysis. Also as described,
we recommend the PubChem Identifier Exchange Service for
casting SMILES or InChIKeys at a medium scale against
PubChem in total or selected sources within it. Many of the
data sources used in this work (and consequently P&D) will
expand with new releases so we expect numbers to change
within a few months of these compilations made in April 2021. During the course of this work and the preceding years of
P&D
operation,
the
team
has
encountered
a
range
of
technical challenges most of which have been alluded to
above. In this regard, while most stand-alone probe sources
are designed with the needs of their users in mind, it is
important for scientists to be able to navigate across multiple
sources to obtain an overview of all potential probes in
advance of experimental planning. This presents a particular
challenge for non-informaticians and for which we needed
much data-wrangling effort to complete the overview that
P&D now offers. During
this
work,
we
also
detected
problematic
anomalies, some of which are listed below. These are not
presented as criticisms but more as pointers towards what
could be improved. 1. The current probe data landscape is particularly patchy. This means for many compounds their associated data falls
short of the well-publicised criteria and thus compromises
the utility of scoring. 2. Comprehensive characterisation of a probe, including
the
necessary
broad
cross-screening,
requires
extensive
experimental work. In addition, the results need to be
accessible
(ideally
in
an
open-access
text-minable
publication), reproducible and easily captured for transfer
into database
records. Conflicts of interest 3. The bias towards known targets seems counter-intuitive. Given that mTOR has 38 020 PubMed hits and ChEMBL has
4557
compounds
aligned
against
P42345
(including
20
clinical candidates), the need for 5 experimental and 225
calculated probes is not obvious (although new highly
selective and potent allosteric modulators of old targets could
provide new insights). As the Pharos TLD categories indicate,
probe development that would broaden Tchem and make There are no conflicts to declare. 3. The bias towards known targets seems counter-intuitive. Given that mTOR has 38 020 PubMed hits and ChEMBL has
4557
compounds
aligned
against
P42345
(including
20
clinical candidates), the need for 5 experimental and 225
calculated probes is not obvious (although new highly
selective and potent allosteric modulators of old targets could
provide new insights). As the Pharos TLD categories indicate,
probe development that would broaden Tchem and make This journal is © The Royal Society of Chemistry 2021 Research Article RSC Medicinal Chemistry Notwithstanding, we have analysed 940 experimental and
3670 calculated probe candidates. Together these provide
evidence of specific binding for 796 human proteins across
the target classes. We have flagged unsuitable (i.e. potentially
misleading and resource-wasting) compounds from both
probe groups. Compared to ChEMBL approved drugs, probes
tend to be larger and more complex structures. inroads into Tdark could lead to functional illumination (but
with
the
caveat
of
the
obvious
paucity
of
assays
for
understudied proteins). 4. The identification and provision of the crucial control
compounds lag behind probe availability. Notably, a recent
analysis of negative controls extols the council of perfection in
that
a
quartet
of
compounds
is
needed
to
maximise
interpretation (i.e. two probes of different chemotypes and two
negative controls, also matched as different chemotypes).41 Although calculated probes are in a large majority, we
established that their scoring is influenced by methodology
and biases in data sources. Consequently, the application of
PMIS and PDPS scoring retrieves different numbers of
quality-rated probes from the Chemical Probes Portal set (i.e. 6 : 1 in favour of PDPS). We thus support scoring as a
pragmatically useful means of compound prioritisation. By
combining established criteria, we developed this further to
delineate 548 high-quality chemical probes (HQCP) covering
just under 450 targets. As we shown above, the Swiss-Prot
bioactive
chemistry
cross-references
indicate
a
data-
supported druggable proteome of 20%. The current “probe
proteome” targets would reach only 4% dropping to half of
that for the HQCP set. Notes and references Bunnage, C. Buser-Doepner, R. M. Campbell, A. J. Carter, P. Cohen, R. A. Copeland, B. Cravatt, J. L. Dahlin, D. Dhanak,
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PHAGOCYTOSIS ACTIVITY OF BINAHONG (Anredera cordifolia (Tenore.) Steenis) FROM SECANG, MAGELANG, CENTRAL JAVA, INDONESIA
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Jurnal farmasi sains dan komunitas
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*Corresponding author: Nunung Yuniarti
Email: nunung@ugm.ac.id, nunzyuniarti@gmail.com Vol. 16 No. 1 Vol. 16 No. 1 JURNAL FARMASI SAINS DAN KOMUNITAS, May 2019, 7-13
p-ISSN 1693-5683; e-ISSN 2527-7146
doi: http://dx.doi.org/10.24071/jpsc.001693 PHAGOCYTOSIS ACTIVITY OF BINAHONG (Anredera cordifolia (Tenore.) Steenis)
FROM SECANG, MAGELANG, CENTRAL JAVA, INDONESIA
Dika Sotya Sakti, Perdana Priya Haresmita, Nunung Yuniarti*), Subagus Wahyuono
Faculty of Pharmacy, Universitas Gadjah Mada, Bulaksumur, Yogyakarta, Indonesia Received January 14, 2019; Accepted March 18, 2019 ABSTRACT The use of medicinal plants is increasing due to the search for alternative resources to treat
diseases such as hypertensions and infection. Along with the development in science, preventive
action should take place to prevent our body from suffering from these diseases. This can be done
by increasing the human immune status with immunomodulatory agents. Binahong empirically
have benefits for wound healing. The purpose of this research was to investigate the
immunomodulatory effect of ethanolic extract of binahong leaves. The non-specific modulatory
effects of the ethanolic extracts of binahong leaves on the immune systems were measured based
on phagocytosis index and phagocytosis capacity. Tests were conducted on strain Balb/C male
mice at the age of 6-8 weeks. Mice were administered orally with the extract of binahong leaves
(doses of 25, 50, 75 mg/kgBW) for 14 days. The test results with the index parameters and
macrophages phagocytosis capacity at doses of 50 and 75 mg/kgBW did not significantly
increased when compared with the controls. From these results, we concluded that the ethanolic
extract of binahong leaves with a dose of 25, 50 and 75 mg/kgBW cannot significantly the activity
of macrophages by phagocytosis index parameters and phagocytosis capacity significantly. Keywords: Anredera cordifolia; immunomodulatory agent; macrophage; phagocytosis capacity;
phagocytosis index Preparation of macrophage cells CMC-Na, Distilled water, alcohol 70%,
ethanol 96% (General Labora), methanol,
Giemsa 20%, latex beads (Sigma™), RPMI
medium 1640 (Gibco™), FBS (Sigma™),
Fungizone (Gibco™), pen-strep (Gibco™),
PBS
(Sigma™),
hepatitis
B
vaccine
(Euvax™). Macrophage cells were isolated from
BALB/c mice (6–8 weeks old). Mice were
euthanized with neck dislocation and 10 mL of
cold RPMI medium were then injected inside
the stomach. After 3-5 minutes, the RPMI was
withdrawn from the stomach using syringe
and put into a conical flask, centrifuged at
1,500 rpm (4oC, for 10 minutes). The
supernatant was removed and the residue
resuspended with RPMI (80% FBS). The
numbers
of
cells
was
calculated with
hemocytometer, diluted with RPMI (80%
FBS) till 2.5 x 106/mL cells density was
obtained. The cells suspension that had been
cultured on 24 wells (200 µL/well, 5x105
cells/well) plate for 24 hours was put in round
coverslips, incubated in a CO2 (5%) incubator,
at 37oC for 30 minutes and then complete
medium, containing 10% FBS, fungizone
0.5% and penicillin streptomycin 2% (1.0 mL)
was added to each well and then incubated for
another 2 hours. The cells were washed twice
with RPMI, and then complete medium (1.0
mL) was added to each well and incubated for
the next 24 hours (Hartini et al., 2013). Preparation of animal test Male mice, strain Balb/c aged 6-7 weeks
weighing 25-35 g were obtained from the
Animal Cage Test Faculty of Pharmacy,
Universitas Gadjah Mada. The protocol of the
study was approved by the Ethics Commission
for Preclinical Trials of Integrated Research
and Testing Laboratory (LPPT), Universitas
Gadjah
Mada
with
certificate
number
00123/04/LPPT/X/2017. Extraction of binahong leaves infected
by
Pseudomonas
aeruginosa
(Sukrama et al., 2017). Binahong leaves were obtained from
Secang, Magelang Regency, Central Java
Province, Indonesia. Powdered leaves material
of Anredera cordifolia was macerated with
ethanol 96% (2x) for 24 hours each, filtered
and the filtrates obtained were combined and
evaporated in vacuum to give thick liquid
material of ethanol extract. The ethanol extract
with the dose of 25, 50 and 75 mg/kgBW were
given orally to assess its immunomodulatory
effects by measuring macrophage activity
using phagocytosis index and phagocytosis
capacity and compared with normal and
CMC-Na groups. We therefore designed this study to
determine the immunomodulatory effects of
binahong
leaves
extract
by
measuring
phagocytosis activity and phagocytosis index. The antimicrobial-antioxidant activity was
correlated with immunomodulatory effects
(Umar et al., 2012; Yuniarti and Lukiswanto,
2017). Phagocytosis
ratio
indicates
the
percentage of active macrophage to 100
macrophages,
and
phagocytosis
index
indicates the number of latex able to be
consumed by active macrophages. These
number are compared to the controls. INTRODUCTION medicinal plant in Indonesia. Binahong leaves
are used for treatment of wounds, refreshing
the body, headache and lowering blood
pressure. Ether fraction of binahong leaves
extract exhibited antioxidant activity measured
by
DPPH
(1,1-diphehyl-2-picrylhydrazyl)
(Ardianti and Guntarti, 2014). Binahong
leaves extract accelerated wound healing
infected by Staphylococcus aureus in mice. Binahong leaves extract also inhibited the
growth of Staphylococcus aureus and as a
result, the healing process the wounds is
faster than wound healing without binahong
leaves extract (Umar et al., 2012). Topical
application of binahong leaves extract makes
wound healing process faster, IL-6 level
higher and increases vascular endothelial
growth factor (VEGF) production in burns Medicinal plants have been used as an
alternative to synthetic drugs, for a long time. Medicinal plants are potential sources of drugs
and are widely used to gain health benefit
empirically
(Rodríguez
et
al.,
2018). Medicinal plants are used because they are
easier to obtain, cheaper than synthetic drugs
and have less side effect (Wardhani and
Sulistyani, 2013). Various diseases such as
hypertension, diabetes mellitus, headache,
inflammation and modulation of the immune
system have been widely treated with
medicinal plants (Leliqia et al., 2017; Putri et
al., 2017). Binahong (Anredera cordifolia (Ten.)
Steenis) is one of the species from the family
of Basellaceae, which has fleshy leaves and
thick aerial tubers. It is widely used as a Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 Preparation of sample Binahong plant leaves were obtained from
the District Secang, Magelang, Central Java. The identify was authenticated by the
Department
of
Pharmaceutical
Biology,
Faculty of Pharmacy, Universitas Gadjah
Mada
in
certificate
number
UGM/FA/2413/M/03/02. Based on the results,
the plants studied is binahong (Anredera
cordifolia (Tenore) Steenis). Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 added into each well containing peritoneum
macrophages, then incubated for another 60
minutes in a CO2 incubator. The cells
suspension was washed 3 times with PBS in
order to remove particles. The cells suspension
was dried at room temperature and then fixed
with methanol. Coverslips were dyed with
Giemsa 20% for 20 minutes, washed with
aquadest, lifted up from the wells and re-dried
at room temperature. The macrophages
activity was calculated as the number (%) of
consumed latex (substrates), visualized by
light microscope (magnified 400x) as seen as
Figure 1. Phagocytosis ratio was indicated by
the percentage of active macrophage in 100
macrophages, and phagocytosis index was
indicated by the number of latex able to be
consumed by active macrophages (Hartini et
al., 2014). These data were compared to the
controls. obtained by calculating the percentage of the
number of active macrophages phagocyted
latex beads per 100 macrophages observed,
whereas macrophage phagocytosis index was
obtained by counting the number of latex
beads phagocyted per 100 macrophages
(Jensch-Junior et al., 2006). These parameters
would be able to show immunostimulatory
effects of ethanolic extract of binahong leaves. Macrophages were isolated from the
peritoneal cavity of mice because the number
of macrophages in the peritoneal cavity is
larger than the other organs and easy to obtain
from the peritoneal fluid. The medium used
was RPMI because this medium can attract
macrophages in the peritoneal cavity and
provide nutrition such as vitamins, amino
acids, and essential materials required for
macrophage cell culture processes. Liquids
that have been isolated from the peritoneal
cavity contains not only macrophage cells but
also granulocytes and lymphocytes. Peritoneal
fluid that was place into a conical flasks was
centrifuged to separate macrophage cells from
other
cells
such
as
lymphocytes
and
granulocytes (Hay and Westwood, 2002). Macrophages activity measurement Macrophages activity measurement was
done involving latex (2 µm in diameter) as
substrates (suspended in PBS, at 2.5 x
107/mL). The cells suspension (200 µL) was 8 Nunung Yuniarti et al. Nunung Yuniarti et al. Phagocytosis Activity of Binahong … Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 Hepatitis B vaccine was administered
intraperitoneally on day 5 and 12 as an
immune booster. The first vaccination was
done to activate the non-specific and specific
immune system and the second vaccination
was done to increase the expression of the
immune system (as a booster) so it is easier to
analyze. Surgery was performed on the 15th
day because the immune system will be
activated optimally up to 3 days after
induction of the vaccine and gradually decline
thereafter (Abbas et al., 2014). macrophages. Other
studies
found
that
binahong leaves extract can also increase the
production of interleukin-6 in the blood
plasma of mice during the healing of burns. Interleukin-6 is a cytokine produced by T-cells
and macrophages to stimulate the immune
response during an infection or during the
healing process (Sukrama et al., 2017). From
the previous research of Sukrama et al., (2017)
and Wijayanti et al., (2018), ethanolic extract
of binahong leaves has the effect to increase
the response of the immune system, but in the
present study, the increasing phagocytosis
capacity and phagocytosis index were not
significant when compared to the control
group. Quantification
of
macrophages
was
performed using latex beads with a size of 3
μm which were resuspended in the serum of
test animals. The size of the latex beads that
resemble the size of bacteria could trigger
macrophage phagocytosis by being perceived
as foreign particles. The addition of serum of
the test animals into the latex suspension
would help the macrophage phagocytosis
process
as
serum
would
facilitate
the
introduction of the antigen by macrophages
(Harvath and Terle, 1999). Research by Wijayanti et al., (2018) was
using a guinea pig test animals while this
study used Balb/c strain mice. Previous
research was conducted to see the effect of
binahong leaves extract on blood leukocytes
profile and the average number of descendants
of the test animals. The increase in the number
of monocytes as one of the parameters of
blood leukocytes profile in these studies was
not always followed by an increase in activity
of macrophages in the tissue. Monocytes from
the blood can differentiate into multiple cell
types of the immune system and in such a
network of dendritic cells, osteoclasts and
macrophages depends on the existing stimulus
inside the body. Monocyte activity in the
network tends to be more specific and
different in each tissue. Phagocytosis Activity of Binahong … RESULTS AND DISCUSSION The parameters of the observed activity of
macrophages were phagocytosis capacity of
macrophages and macrophage phagocytosis
index. Macrophage phagocytosis capacity was Figure 1. Macrophage observation under microscope magnification 400x. A: active macrophage; B: inactive
macrophage. B
A Figure 1. Macrophage observation under microscope magnification 400x. A: active macrophage; B: inactive
macrophage. 9 Phagocytosis Activity of Binahong … Phagocytosis Activity of Binahong … Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 A Figure 2. Results of statistical analysis of phagocytosis index (A) and phagocytosis capacity (B). B
A B
A A B Figure 2. Results of statistical analysis of phagocytosis index (A) and phagocytosis capacity (B). Table I. Phagocytosis Capacity and Phagocytosis Index of Binahong Leaves Ethanolic Extracts
Phagocytosis Capacity
Phagocytosis Index
Results
Mean ±SD
Results
Mean ±SD
Normal Group
44.00%
46.20%
± 0.0414
1.97
2.07
± 0.1521
46.67%
1.92
40.33%
2.32
50.33%
2.04
49.67%
2.08
CMC-Na + Vaccine Group
48.67%
49.87%
± 0.0899
242
2.70
± 0.4076
65.33%
3.02
42.00%
3.08
47.33%
284
46.00%
2.13
Dose 25 mg/kgBW
+ Vaccine
63.33%
49.33%
± 0.1269
2.88
2.59
± 0.2267
33.67%
2.65
54.67%
2.35
45.67%
2.48
Dose 50 mg/kgBW
+ Vaccine
40.00%
58.86%
± 0.1324
2.81
2.55
± 0.3002
59.67%
2.88
53.00%
2.76
74.33%
2.25
67.33%
3.06
Dose 75 mg/kgBW
+ Vaccine
61.00%
62.33%
± 0.0610
3.39
3.12
± 0.6526
61.67%
2.20
56.00%
3.72
70.67%
3.17 Table I. Phagocytosis Capacity and Phagocytosis Index of Binahong Leaves Ethanolic Extra Table I. Phagocytosis Capacity and Phagocytosis Index of Binahong Leaves Ethanolic Extracts
Phagocytosis Capacity
Phagocytosis Index Table I. Phagocytosis Capacity and Phagocytosis Index of Binahong Leaves Ethanolic Extracts
Phagocytosis Capacity
Phagocytosis Index 10 Nunung Yuniarti et al. Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 (33), 1582–1614. (33), 1582–1614. (33), 1582–1614. has a lower bioavailability compared with
administration by injection and the drug can
be metabolized by gastrointestinal fluids or
often called first-past experience effect
(Atanasov et al., 2015). Hartini, Y.S., Wahyuono, S., Widyarini, S.,
and Yuswanto, A., 2013. Phagocytic
Macrophage Activity of Fractions from
Methanolic Leaf Extract of Red Betel
(Piper crocatum Ruiz & Pav.) In Vitro. Jurnal Ilmu Kefarmasian Indonesia,
(11), 108–115. Another factor that could affect the results
when compared with previous studies was the
source of the sample. Wahyukundari and
Praharani (2016) collected the binahong leaves
from Jember, East Java whereas in this study
the binahong leaves were collected from
Secang, district of Magelang, Central Java. Different areas could cause differences in the
content of secondary metabolites in plants
because each area has different soil nutrients. Differences in soil nutrients would affect the
availability of nutrients and plant precursor to
form secondary metabolites (Salim et al.,
2017). Hartini, Y.S., Wahyuono, S., Widyarini, S.,
and Yuswanto, A., 2014. In vivo
Immunomodulatory
Effect
and
Histopathological Features of Mouse
Liver
and
Kidney
Treated
With
Neolignans Isolated from Red Betel
(Piper crocatum Ruiz & Pav) Leaf. Tropical Journal of Pharmaceutical
Research. (13), 1609–1614. Harvath, L. and Terle, D.A., 1999. Assay for
Phagocytosis,
In:
Javois,
L.C.,
Immunocytochemical
Methods
and
Protocols. New York: Humana Press,
281-290. CONCLUSION Ethanolic extract of binahong leaves at
dose of 25, 50 and 75 mg/kgBW cannot
significantly
increase
the
activity
of
macrophages
by
phagocytosis
index
parameters and phagocytic capacity. Ethanolic extract of binahong leaves at
dose of 25, 50 and 75 mg/kgBW cannot
significantly
increase
the
activity
of
macrophages
by
phagocytosis
index
parameters and phagocytic capacity. Hay, F.C. dan Westwood, O.M.R., 2002. Practical Immunology, Fourth edition,
Blackwell Science, United Kingdom. Hulin, I., Jakubovsky, J., and Viera, S., 1995. Inflammation and Fever, in: Hulin, I.,
Jakubovsky,
J.,
and
Viera,
S.,
Pathophysiology
Principles
of
Diseases. Bratislava:
Comenius
University, 65-72. Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 The difference is
caused by different stimuli derived from the
macrophage microenvironment (Hulin et al.,
1995). Based on our results which was presented
in Table I, after examining macrophage
phagocytic
capacity
and
macrophage
phagocytosis index, the ethanolic extract of
binahong leaves at dose of 25, 50 and 75
mg/kgBW did not significantly enhance the
ability of macrophage through an increase of
both parameters compared with the control
group 0.5% CMC-Na. A significant difference
was found only in the phagocytosis index data
for the dose group 75 mg/kgBW when
compared with the normal group, as seen as in
Figure 2. Research conducted by Wahyukundari et
al. (2016) was performed in vitro by taking
blood monocytes. Experiments in vitro have
less variables that cannot be controlled as
compared to experiments in vivo. In vivo
experiments
involve
factors
including
pharmacokinetics (absorption, distribution,
metabolism and excretion) and the first-past
effect that could affect the availability of the
extract in the body. In addition, the
physiological state of the test animals such as
hormones could also influence the effect of the
test sample. Absorption through the oral drug According to previous research, 70%
ethanolic extract of binahong leaves dose of
50 mg/ kgBW could raise the profile of
leukocytes as an increase in total leukocytes,
neutrophils and total monocytes in guinea pigs
(Wijayanti et al., 2018). In addition, the
ethanolic extract of binahong leaves at
concentrations of 50% and 100% could
increase the phagocytic monocytes with in
vitro method (Wahyukundari and Praharani,
2016). Monocytes are produced by the bone
marrow which would circulate in the blood
before it becomes differentiated in the tissue to 11 Jurnal Farmasi Sains dan Komunitas, 2019, 16(1), 7-13 Phagocytosis Activity of Binahong … REFERENCES Abbas, A.K., Lichtman, A.H., and Pillai, S.,
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Pemberian 13 Phagocytosis Activity of Binahong …
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https://openalex.org/W4372295311
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https://www.brazilianjournalofscience.com.br/revista/article/download/385/223
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English
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Histopathology of broiler chickens fed diets supplemented with Prosopis africana (African mesquite) essential oil
|
Brazilian Journal of Science
| 2,023
|
cc-by
| 6,348
|
Abstract This study was carried out to investigate the hepatic histopathology of broiler chickens fed diets supplemented
with Prosopis africana (African mesquite) essential oil (PRSO). 300, 1-day old broiler chicks were randomly
distributed into six treatments with 5 replicates consisting of 10 birds each. Birds in diet 1 (D1) was fed basal
diet supplemented with 0 % PRSO, D2: basal diet with 1.0 g/kg-1 oxytetracycline, D3, D4, D5 and D6 were fed
basal diet supplemented with PRSO at 200 mg, 400 mg, 600 mg and 800 mg/kg-1 respectively. The experiment
lasted for 56 days and a completely randomized design was adopted. Phyto-constituents of PRSO revealed the
presence of tannins (201.21 mg/g-1), flavonoids (677.83 mg/g-1), alkaloids (405.90 mg/g-1), saponins (75.62
mg/g-1), phenols (508.28 mg/g-1), phytates (20.10 mg/g-1) and oxalates (5.26 mg/g-1). Hepatic histopathology
showed that tissues from D1, D3, D4, D5 and D6 showed no signs of any damage; the central vein, blood portal
vessels showed no congestion. Mild hepatocellular damage was recorded among liver of birds in D2. It can be
concluded that Prosopis africana essential oil are among the key alternative solutions to antibiotics because they
contain bioactive compounds that are safe, effective and performs multiple biological functions in the body of
animals. Keywords: Prosopis africana oil, phytochemicals, broilers, liver, histopathology. Resumo Este estudo foi realizado para investigar a histopatologia hepática de frangos de corte alimentados com dietas
suplementadas com óleo essencial de Prosopis africana (mesquita africana) (PRSO). 300 pintos de corte de 1 dia
de idade foram distribuídos aleatoriamente em seis tratamentos com 5 repetições de 10 aves cada. Aves na dieta
1 (D1) receberam dieta basal suplementada com 0 % de PRSO, D2: dieta basal com 1,0 g/kg-1 de oxitetraciclina,
D3, D4, D5 e D6 receberam dieta basal suplementada com PRSO a 200 mg, 400 mg, 600 mg e 800 mg/kg-1,
respectivamente. O experimento teve duração de 56 dias e foi adotado o delineamento inteiramente casualizado. Os fitoconstituintes do PRSO revelaram a presença de taninos (201,21 mg/g-1), flavonoides (677,83 mg/g-1),
alcaloides (405,90 mg/g-1), saponinas (75,62 mg/g-1), fenóis (508,28 mg/g-1), fitatos (20,10 mg/g-1) e oxalatos
(5,26 mg/g-1). A histopatologia hepática mostrou que os tecidos de D1, D3, D4, D5 e D6 não apresentavam sinais
de danos; a veia central, os vasos sanguíneos portais não apresentaram congestão. Danos hepatocelulares leves
foram registrados entre fígados de aves em D2. Pode-se concluir que o óleo essencial de Prosopis africana está
entre as principais soluções alternativas aos antibióticos porque contém compostos bioativos que são seguros,
eficazes e desempenham múltiplas funções biológicas no corpo dos animais. Palavras-chave: óleo de Prosopis africana, fitoquímicos, frangos de corte, fígado, histopatologia. Histopathology of broiler chickens fed diets supplemented with
Prosopis africana (African mesquite) essential oil Olujimi John Alagbe1, Oluchi C. P. Agubosi1 & Rufus Adebisi Oluwafemi1
1 Department of Animal Science, University of Abuja, Gwagwalada, Nigeria
Correspondence: Olujimi John Alagbe, Department of Animal Science, University of Abuja, Gwagwalada,
Nigeria. E-mail: dralagbe@outlook.com Accepted: April 25, 2023
Publi
URL: https://doi.org/10.14295/bjs.v2i9.385 Published: September 01, 2023
9.385 Received: April 05, 2023
DOI: 10.14295/bjs.v2i9.385 Received: April 05, 2023
DOI: 10.14295/bjs.v2i9.385 Published: September 01, 2023 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 Histopathology of broiler chickens fed diets supplemented with
Prosopis africana (African mesquite) essential oil
Olujimi John Alagbe1, Oluchi C. P. Agubosi1 & Rufus Adebisi Oluwafemi1
1 Department of Animal Science, University of Abuja, Gwagwalada, Nigeria
Correspondence: Olujimi John Alagbe, Department of Animal Science, University of Abuja, Gwagwalada,
Nigeria. E-mail: dralagbe@outlook.com 1. Introduction Nowadays, there is an increasing awareness on the need to maintain good health via the use of natural products
from plants because they are efficient, cheap, easily metabolized with minimum health hazards and do not pose 49 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 any environmental pollution (Alagbe, 2022; Wallace et al., 2011). Medicinal plants are also known as phytogenic
feed additives are regarded as complex mixtures of bioactive components or phytochemicals found in varying
amount depending on geographical location, stage of growth or harvesting, parts used (leaf, seeds, bark, buds,
flowers and twig) as well as method of processing (Shittu; Alagbe, 2020; Simitzis, 2017). any environmental pollution (Alagbe, 2022; Wallace et al., 2011). Medicinal plants are also known as phytogenic
feed additives are regarded as complex mixtures of bioactive components or phytochemicals found in varying
amount depending on geographical location, stage of growth or harvesting, parts used (leaf, seeds, bark, buds,
flowers and twig) as well as method of processing (Shittu; Alagbe, 2020; Simitzis, 2017). Bioactive compounds or phytochemicals from plants includes: phenolics, flavonoids, terpenoids, carotenoids,
saponins, alkaloids, flavours and fragrances. Plant extracts (essential oils) with these bioactive molecules are
widely used in the food, pharmaceutical and cosmetic industries (Lewis et al., 2003; Alagbe, 2017). They are
also capable of enhancing livestock productivity through the improvement of digestibility, nutrient absorption
and elimination of pathogens residents in the animal gut (Denli et al., 2004). Medicinal plants had possessed
antimicrobial (Agubosi et al., 2022; Nursoy et al., 2011), antioxidants (Adewale et al., 2021), antibacterial
(Dhama et al., 2015), antiviral (Mehr et al., 2014), immune-modulatory (Krishan; Narang, 2014; Olorunmaiye et
al., 2019), hepato-protective, antifungal and cytotoxic (Rostagno et al., 2011; Souza et al., 2015), antifungal,
anti-proliferative, anti-helmithic, antiviral and anti-tumor (Souza et al., 2015), anti-cancer (Assis et al., 2017)
and anti-protozoal, anti-proliferative, anticoagulant, anticonvulsant, antipsychotic (Brenes; Roura, 2010; Cho et
al., 2014). Essential oils (EOs) has shown positive effect in livestock by promoting growth, relieving the host animals from
immune defense stress during critical situations, scavenging free radicals capable of causing diseases and
increase the intestinal availability of essential nutrients for absorption, thereby helping animals to grow better
within the framework of their genetic potential (Lee et al., 2003; Bozkurt et al., 2016). 1. Introduction EOs have been
demonstrated to work through the antioxidative mechanisms serving as free radical scavengers promoted by
polyphenols, flavonoids and terpenoids which possess the ability to protect the cell organelles from damage
caused by free radicals induced oxidative stress either by inhibiting the initiation or propagation of oxidative
chain reactions (Sevim et al., 2017; Zeng et al., 2015). Studies have shown that the plants active principle agents
in the herbs have a strong capability for scavenging superoxide radicals, hydrogen peroxide and nitric oxide from
activated macrophages, reducing iron complex and inhibiting lipid peroxidation (Stef et al., 2009; Umashanker et
al., 2011). Prosopis africana (African mesquite) is a leguminous multipurpose tree that belongs to the family Fabaceae and
subfamily Mimosoideae (Ogunshe et al., 2017). It is known as Iron wood in English, Kiriya in Hausa, Ubwa in
Igbo and it consist of 14 species and widely grown in tropical, arid and semi arid regions of the world
(Olorunmaiye et al., 2019). Essential oils can be extracted mostly from the seed which contains compounds with
therapeutic properties (alkaloids, saponins, tannins, phenols, cardiac glycosides, flavonoids and terpenoids)
(Olorunmaiye et al., 2019; Gberikon et al., 2015; Ugwoke et al., 2019). Traditionally, the roots and stem bark extracts are used for the treatment and prevention of fever, skin diseases,
diarrhea, dysentery, respiratory disorders, rheumatism, pneumonia and ulcer (Gberikon et al., 2015; Oguntoyinbo
et al., 2017). Prosopis africana oil can also inhibit the growth of pathogenic microorganisms; Staphyllococcus
spp, Bacillus spp, Escherichia coli, Enterobacter spp, Enterococcus spp, Pseudomonas aeruginosa, Shigella spp,
Klebsiella spp and Proteus spp (Alagbe, 2023; Oguntoyinbo et al., 2007; Gordon et al., 1973). There is no justification for assuming that just because essential oils are natural, they must therefore be safe;
certain phytochemicals or bioactive compounds can interfere with the body’s system for metabolizing
medications in the liver, which can have a chain reaction of negative effects. Therefore, this experiment was
designed to investigate the histopathology of broiler chickens fed diets supplemented with Prosopis africana
(African mesquite) essential oil. 2. Materials and Methods 2.1 Ethical approval, collection and processing of Prosopis africana oil (PRSO) 2.4 Determination of phytochemical components of Prosopis africana oil 2.4 Determination of phytochemical components of Prosopis africana oil Phenols and total tannins in PSO were evaluated using Folin-Ciocalteau method outlined by Otles and Yacin
(2012). Total flavonoids, alkaloids, phytates and oxalates were determined using the methods described by
Harborne (1973); Odebiyi and Sofowora (1978). Saponins was quantified using the protocols used by Brunner
(1984). Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 Prosopis africana oil was extracted employing the steam-distillation method, which also calls for the use of a
digital scale, a round bottom flask, distilled water, a heating mantle made of glass yarn, a measuring cylinder,
and a separatory funnel. 250 mL of distilled water was added to a round-bottom flask along with 50 grams of
ground Prosopis seed. The mixture was added to a glass yarn heating mantle, warmed to a temperature of 80 °C,
and then the condenser was positioned above the flask with a round bottom. The combination is forcefully boiled
for 15 min, the distillate is collected in a beaker until no more oil droplets flow over, 5 grams of sodium chloride
are added, and the mixture is constantly agitated. To get Prosopis africana oil, the distillate was then put into the
separatory funnel and evaporated in a steam bath. 2.2 Management of experimental animals and design 2.2 Management of experimental animals and design 300, 1-day old broiler chicks of mixed sex (Arbo Acres) were purchased from a reputable commercial hatchery
in Oyo State, Nigeria and randomly distributed into six treatments with 5 replicates consisting of 10 birds each. A galvanized battery cage measuring (100 cm × 75 cm × 50 cm) (length × depth × height) was equipped with
automatic nipple drinkers and galvanized manual feeder in a semi open pens. Chicks were weighed on arrival to
determine their average initial body weight and thereafter on weekly basis and given anti-stress (glucose + water)
at the rate of 2 grams of glucose to 10 liters of water. Completely randomized experimental design was used in
this study. 2.3 Experimental diets Diets were compounded based on the nutrient requirements for broilers according to Olomu (2021) as presented
in (Table 1). Experimental diet at the starter phase (1-28 days) and finisher phase (29-56 days) contains 23.06%
crude protein (CP), 21.40% CP and metabolizable energy content (ME) of 2991.5 Kcal/kg-1 and 3108.1 Kcal/kg-1
dry matter respectively. All birds had unrestricted access to feed and water throughout the entire experimental
period (8 weeks). Experimental set-up is shown below: Experimental set-up is shown below: Diet 1 (D1): Basal diet with no Prosopis africana oil (Negative control)
Diet 2 (D2): Basal diet + Oxytetracycline 1.00 g/kg-1 (Positive control)
Diet 3 (D3): Basal diet + 200 mg/kg-1 Prosopis africana oil (PRSO)
Diet 4 (D4): Basal diet + 400 mg/kg-1 PRSO
Diet 5 (D5): Basal diet + 600 mg/kg-1 PRSO
Diet 6 (D6): Basal diet + 800 mg/kg-1 PRSO Diet 1 (D1): Basal diet with no Prosopis africana oil (Negative control) 2.1 Ethical approval, collection and processing of Prosopis africana oil (PRSO) The study was carried out at the University of Abuja Teaching and Research Farm in the month of April - June,
2022 according to the guidelines of animal protocol agreed by the Research and Ethic committee (ANSJ/004) of
the Department of Animal Science, University of Abuja, Gwagwalada, Nigeria. Fresh seeds of Prosopis africana were purchased from a local market in Gwagwalada, authenticated at the
Department of Biological Sciences, University of Abuja by a certified taxonomist where a Voucher specimen
number PRO/ANS/004 was deposited at the herbarium of the same department. Air dried on a clean metallic
plate for 11 days and grinded into powder using a blender thereafter poured into a well labeled zip log before it
was transported to a laboratory for further laboratory examination. 50 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 2.5 Histopathological evaluation **Premix supplied per kg diet: -
vit A, 8,500 I.U; vit E, 10 mg; vit D3, 1500 I.U, vit K, 3.8 mg; vit B2, 10 mg; Niacin, 15 mg; vit B12, 10 mg;
choline chloride, 250 mg; Mn, 5.0 mg; Zn, 56 mg; Cu, 1.6 g; folic acid, 2.8 mg; Fe, 5.1 g; pantothenic acid, 10
mg; biotin, 30.5 g; antioxidant, 56 mg (finisher’s mash). Source: Autors, 2023. 2.5 Histopathological evaluation Liver specimens (1 cubic cm) were taken on the 56th day of the experiment and placed in a well-sealed leak
proof labelled container containing 10% formalin. The 10% formalin solution prevents the tissues from
degeneration. Thereafter, specimens were transported at room temperature to the laboratory for further
examination. All tissues were stained with Hematoxylin and Eosin (H & E) stain and the hsitopathological
changes were observed under a microscope. 51 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 Table 1. Ingredient composition of the experimental diets. Materials
Starters mash (1-28 days)
Finishers mash (29-56 days)
Maize
52.55
57.55
Wheat offal
2.00
5.00
Soya bean meal
29.00
22.45
Groundnut cake
8.00
5.00
Fish meal (72%)
3.00
3.00
Limestone
1.50
2.00
Bone meal
3.00
4.00
Lysine
0.20
0.20
Methionine
0.20
0.20
**Premix
0.25
0.25
Salt
0.30
0.35
Total
100.0
100.0
Determined analysis (% DM)
Crude protein
23.06
21.40
Crude fibre
4.00
5.30
Ether extract
4.03
4.47
Calcium
1.66
1.83
Phosphorus
0.78
0.99
Energy (Kcal/kg-1)
2991.5
3108.1
Note: *Premix supplied per kg diet: - vit A, 10,000 I.U; vit E, 5 mg; vit D3, 3000 I.U, vit K, 3 mg; vit B2, 5.5 mg;
Niacin, 25 mg; vit B12, 16 mg; choline chloride, 120 mg; Mn, 5.2 mg; Zn, 25 mg; Cu, 2.6 g; folic acid, 2 mg; Fe,
5g; pantothenic acid, 10 mg; biotin, 30.5 g; antioxidant, 56 mg (starter’s mash). **Premix supplied per kg diet: -
vit A, 8,500 I.U; vit E, 10 mg; vit D3, 1500 I.U, vit K, 3.8 mg; vit B2, 10 mg; Niacin, 15 mg; vit B12, 10 mg;
choline chloride, 250 mg; Mn, 5.0 mg; Zn, 56 mg; Cu, 1.6 g; folic acid, 2.8 mg; Fe, 5.1 g; pantothenic acid, 10
mg; biotin, 30.5 g; antioxidant, 56 mg (finisher’s mash). Source: Autors, 2023. Note: *Premix supplied per kg diet: - vit A, 10,000 I.U; vit E, 5 mg; vit D3, 3000 I.U, vit K, 3 mg; vit B2, 5.5 mg;
Niacin, 25 mg; vit B12, 16 mg; choline chloride, 120 mg; Mn, 5.2 mg; Zn, 25 mg; Cu, 2.6 g; folic acid, 2 mg; Fe,
5g; pantothenic acid, 10 mg; biotin, 30.5 g; antioxidant, 56 mg (starter’s mash). 3.1 Phytochemical constituents of Prosopis africana oil Constituents
Concentrations (mg/g-1)
Tannins
201.21
Flavonoids
677.83
Alkaloids
405.90
Saponins
72.62
Phenols
508.28
Phytates
20.10
Oxalates
5.26
Source: Authors, 2023. Figure 1. Phyto-constituents in Prosopis africana oil. Source: Authors, 2023. 3.2 Histopathology examination of birds fed Prosopis africana oil (PRSO)
The examined histopathogical liver sections is presented in (Board 1 (Figure 1 at 6) Histopathology) The liver is Table 2. Phytochemical constituents of Prosopis africana oil. Constituents
Concentrations (mg/g-1)
Tannins
201.21
Flavonoids
677.83
Alkaloids
405.90
Saponins
72.62
Phenols
508.28
Phytates
20.10
Oxalates
5.26
Source: Authors, 2023. Table 2. Phytochemical constituents of Prosopis africana oil. Figure 1. Phyto-constituents in Prosopis africana oil. Source: Authors, 2023. Figure 1. Phyto-constituents in Prosopis africana oil. Source: Authors, 2023. Figure 1. Phyto-constituents in Prosopis africana oil. Source: Authors, 2023. 3.1 Phytochemical constituents of Prosopis africana oil Phytochemical constituents of Prosopis africana oil is summarized in (Table 2 and Figure 1). Flavonoids had the
highest concentration (677.83 mg/g-1) followed by phenols (508.28 mg/g-1), alkaloids (405.90 mg/g-1), tannins
(201.21 mg/g-1), saponins (72.62 mg/g-1), phytates (20.10 mg/g-1) and oxalates (5.26 mg/g-1) respectively. All the
phytochemicals have stated above have different medicinal and pharmacological importance. For instance,
flavonoids therapeutically function as strong antimicrobial against pathogenic organisms (Alagbe, 2020),
antioxidants against reactive oxygen species which are generated from free radicals (Oluwafemi et al., 2021). Alkaloids have several pharmacological properties on human and livestock health including anti-inflammatory,
analgesics, antifungal, antimalarial, anti-hypertensive, neuro-protective, anti-parasitic and anti-tumor (Xie et al.,
2021; Roy, 2017). High concentrations of alkaloids in the body system can lead to chronic toxicity and other
health complications such as abdominal pains, nausea, increased cardiac rates, neuro-toxicity, genotoxicity and
liver diseases (Ma et al., 2007; Adibah et al., 2019). Tannins exhibits strong biological activities (anti-bacterial,
antimicrobial, hepato-protective, antiarrhythmic and immune-modulatory) (Adewale et al., 2021; Singh et al.,
2021). Phenols have strong antioxidant properties and are capable of preventing oxidative damage to biomolecules such
as DNA, lipids and proteins (Hassan et al., 2012; Singh et al., 2023). Saponins are immune stimulators (Hassan
et al., 2012), antimicrobials (Soetan et al., 2006). Researchers have reported its hypocholesterolemic properties
(Cheeke, 2000). Phytates have been reported to be effective in the treatment of wounds and haemorrage 52 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 (Oluwafemi et al., 2021). They are antioxidants capable of reducing the risk of cardiovascular disease, kidney
stones, osteoporosis and iron absorption (Shittu; Alagbe, 2021). Excessive ingestion of oxalates can interfere
with the absorption of calcium in the body (Varadharajan et al., 2012); Sombié et al. (2011). The results obtained
in this research is in consonance with previous findings of Ogbeba et al. (2017); Singh & Bhat (2013). Tobias
(2010) reported that the composition of essential oils is influenced by cultivar within the plant variety, climatic
conditions and time of harvest. The presence of multiple phyto-constituents in Prosopis africana shows that it
has notable beneficial effects in maintaining animal’s performance, antioxidant and osmotic balance, palatability,
retention time of feed, modulating fatty acid in the muscle as well as supporting the intestinal integrity (Singh et
al., 2023). Table 2. Phytochemical constituents of Prosopis africana oil. 3.2 Histopathology examination of birds fed Prosopis africana oil (PRSO) The examined histopathogical liver sections is presented in (Board 1 (Figure 1 at 6) Histopathology). The liver is
the largest gland in the body. It is made up of hepatic lobules which are classically described as been hexagonal
in structure (Dooley et al., 2002). Birds in Figure 1 (fed 0% PRSO) showed no histopathological signs of any
damage; the central vein, blood portal vessels showed no congestion. Figure 2 (fed oxytetracycline) showed mild
perivascular lymphocytic aggregation and moderate hepatic sinusoids. Figure 3, 4, 5 and 6 (fed PRSO) revealed
normal tissue achietecture with normal hepatic lobules, central vein and hepatic sinusoids. 53 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 Board 1. Figure 1. Basal diet with no Prosopis africana oil (Negative control). Figure 2. Basal diet + Neomycin
1.00 g/kg-1 (Positive control). Figure 3. Basal diet + 200 mg/kg-1 Prosopis africana oil (PRSO). Figure 4. Basal
diet + 400 mg/kg-1 PRSO. Figure 5. Basal diet + 600 mg/kg-1 PRSO. Figure 6. Basal diet + 800 mg/kg-1 PRSO. Source: Authors, 2023. Board 1. Figure 1. Basal diet with no Prosopis africana oil (Negative control). Figure 2. Basal diet + Neomycin
1.00 g/kg-1 (Positive control). Figure 3. Basal diet + 200 mg/kg-1 Prosopis africana oil (PRSO). Figure 4. Basal
diet + 400 mg/kg-1 PRSO. Figure 5. Basal diet + 600 mg/kg-1 PRSO. Figure 6. Basal diet + 800 mg/kg-1 PRSO. Source: Authors, 2023. According to Harold et al. (2008); Plinio & Broglia (2000), the liver help to detoxify toxic substances,
metabolism of micro-nutrients, excretion of bilirubin and synthesis of bile salts. The healthy tissue architecture
of birds given Prosopis africana oil is evident that both their bioactive components and inclusion levels are safe. PRSO are well known for their beneficial effects on animals, from anti-inflammatory, antioxidant and
antimicrobial properties (Agubosi et al., 2022; Oluwafemi et al., 2020). The outcome of this investigation is
consistent with the findings of Oloruntola (2019) when phytogenics (pawpaw leaf and seed meal) were
supplemented in the diet of broiler chickens. Similar findings were made by Samantha et al. (2019) when essential oils were given to broilers that were
coccidiosis-challenged. Omokore & Alagbe (2019); Alagbe & Oluwafemi (2019) claimed that bioactive
substances found in essential oils may inhibit the pathogenicity features in bacteria, such as flagella, which are
important for bacterial adhesion or adhesion to any mammalian preference sites. 5. Acknowledgments We will like to acknowledge the efforts of Mrs. Mercy Cincinsoko Kadiri in sourcing Prosopis seeds. 3.2 Histopathology examination of birds fed Prosopis africana oil (PRSO) Alagbe (2022) asserts that the
liver is crucial for the creation of bile, the metabolism of carbohydrates, and the removal of toxins from an
animal's system. The abundance of flavonoids, a significant phytogeenic compound could significantly decrease
liver tissue degenerative conditions particularly blockages in the liver of animals when compared to the other
treatments. Antibiotics like oxytetracycline are widely used to prevent, control, and treat disease in animals; however, due to
their uncontrolled usage and insufficient sensitivity testing before treatment, there are a rising number of
incidences of resistance (Adewale et al., 2021). Alagbe et al. (2019) claim that bioactive substances found in
essential oils may inhibit the pathogenicity features in bacteria, such as flagella, which are important for bacterial
adhesion or adhesion to any mammalian preference sites. Alagbe (2022) asserts that the liver is crucial for the
creation of bile, the metabolism of carbohydrates, and the removal of toxins from an animal's system. The abundance of flavonoids, a significant botanical compound could significantly decrease liver tissue
degenerative conditions particularly blockages in the liver of animals. Degeneration in liver histopathology has
also been observed to be enhanced by contaminants (Soetan; Oyewole, 2009). Infections with bacteria may have
caused the severe and mild hepatic damage seen in birds in T2. Despite the fact that antibiotics were
administered to the birds in T2, this finding suggests that there may be an antimicrobial resistance issue. 4. Conclusions 4. Conclusions 54 Brazilian Journal of Science, 2(9), 49-59, 2023. ISSN: 2764-3417 It can be concluded that the more antibiotics (oxytetracycline) fed to animals the greater the selective pressure
that favours resistant strains. However, Prosopis africana essential oil possess several properties – antioxidant,
antiviral, anti-inflammatory, anti-depressant, immune-stimulatory, antimicrobial, hepato-protective amongst
others, all of which are vital for the performance enhancing effects in livestock’s. 8. Ethics Approval Yes applicable. 6. Auhors’ Contributions Olujimi John Alagbe: field work and compilation of results, subimission, corrections and publication. O. C. P
Agubosi: field work and compilation of results. R. A. Oluwafemi: field work and compilation of results. 7. Conflicts of Interest No conflicts of interest. http://dx.doi.org/10.32861/ajls.511.107.111 http://dx.doi.org/10.32861/ajls.511.107.111 Alagbe, J. O. (2020). Caecal microbial population of growing grass cutters fed Phyllantus amarus and
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Copyright for this article is retained by the author(s), with first publication rights granted to the journal. Copyright for this article is retained by the author(s), with first publication rights granted to the journal This is an open-access article distributed under the terms and conditions of the Creative Commons Attribution
license (http://creativecommons.org/licenses/by/4.0/). This is an open-access article distributed under the terms and conditions of the Creative Commons Attribution
license (http://creativecommons.org/licenses/by/4.0/). 59
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German
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Ueber gemischte, Aethylendiamin und Ammoniak enthaltende Triamminkobaltsalze
|
Berichte der Deutschen Chemischen Gesellschaft
| 1,905
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public-domain
| 4,031
|
das komtnt bei Gleicbgewicbtastudien, wie diejenigen von S c h e n c k
iind H e l l e r , vallkommen auf da-selbe hinaos. das komtnt bei Gleicbgewicbtastudien, wie diejenigen von S c h e n c k
iind H e l l e r , vallkommen auf da-selbe hinaos. studirt, man bei Renutzurig von
amorpher Kolrle resp. Diamant
co*+c+2co . . . . . . (1)
P a w
PCC),
fur dae Verhaltniss
Werihe bekomrnen muss. welche, z. B. bei
64 I " , 5.4 resp. 4.2 hlal so gross sind als bei Graphit. Wir durfen
hier aber nicht vergessen, dass wir zu diesem Rtwltate gelangt sind
ntiter der Annahme, dass der Quotient
bei dem Versuclir mit den
verscbiedenen Kohlenstotrmodificationen denselben Werth bat, und dieses
iet sicher nicht der Fall. Jedenfalls wird also dns Studium des
(;leichgewichtes
( I )
ons mit grnsserer Sicherheit etwas iiber die
Ddntpfspannungen der Kohlettstoffmoclificatiouert aussagen. als es die
Untersuchung von S c h e n c k und H e l l e r vermag. Das Gleichgewicht
( 1 ) w i d von mir in dieser Richtung untersucht. Ptw I
P 1.' Q verscbiedenen Kohlenstotrmodificationen denselben Werth bat, und dieses
iet sicher nicht der Fall. Jedenfalls wird also dns Studium des
(;leichgewichtes
( I )
ons mit grnsserer Sicherheit etwas iiber die
Ddntpfspannungen der Kohlettstoffmoclificatiouert aussagen. als es die
Untersuchung von S c h e n c k und H e l l e r vermag. Das Gleichgewicht
( 1 ) w i d von mir in dieser Richtung untersucht. A. rns t e r d a m . Aoorg.-chem. Laboratarium der Univereitat. A. rns t e r d a m . Aoorg.-chem. Laboratarium der Univereitat. A. rns t e r d a m . Aoorg.-chem. Laboratarium der Univereitat. 703. A. Werner und A. Griin: Ueber gemischte, Aethylen-
diamin und Ammoniak enthaltende TI-iamminkobaltsalae. (Eingegangen am 30. Novemb1.r 1905) Zur Klarung bestimmter structureller und stereachemischer Fragen
bei Coordiriationsrerbindungen erschien es wunschenswerth , an Stelle
der nicht sehr bestfindigen, nur Ammonia% enthaltenden. reinen Tri-
:imminsalze analoge. virlleicht bestandigere, aminsubstititirte Verbin-
dungen zu gewinnen. Wir hahen deshalb einige athylendiatninbaltige
Vertreter der Triamminkobaltreihe dargestellt, die wir im Folgenden
beschreibcn wollen. Die Unterruchung hat aber gezeigt, dass der Ein-
tritt von Aetbylendiarnin den Chemismus und die physikalischen Eigen-
schaften van Triamminsalzen mehr andpi t, als es bei Tetraamminsalzen
der Fall ist, nnd dass, inr Gegensatz zum Verhalten der Tetrammin-
salze, die Hestandigkeit des Kobaltiakrndica!s der Triamminsalze dnrch
den Eintritt vou hethylendiamin nicht erbiiht, soodern erniedrigt wird. das komtnt bei Gleicbgewicbtastudien, wie diejenigen von S c h e n c k
iind H e l l e r , vallkommen auf da-selbe hinaos. I) Zeitscbr. fiir anorg. Chem. 13, 1SO [189G]. 2, Ztsitscbr. fiir anorg. Chem. 8, 171 [1S95]. das komtnt bei Gleicbgewicbtastudien, wie diejenigen von S c h e n c k
iind H e l l e r , vallkommen auf da-selbe hinaos. Als Ausgsngsmnterial zur Dnrstellung iinserer Verbindungen diente
ein Tr i n i tri t o - a t h y I e n d i am in-a mrni t i - ko b al t , das sic+, neben
anderen Producten, bei der Einwirkung ron Aethylendi:iinin auf tetra- 1034 iiitritodiamminkobaltsaure Salze: [(HJN)~CO(SC~)~]R
( E r d m a n n -
sches Salz) bildet. Es liornmt ihm die Forniel zu, uud seine Bilduug aus dern Erdmanii’schen Salz lasst sich fol-
eendermanssen forniuliren: Dab ‘L’rinitritoatliylet diarninamminkobalt ahnelt in seinen Eigen-
schaften dem ’~ririitritotii;~mmi~iknbalt,
deniles m c h constitutionell an
die Seite zu srrllen ist: Triammin kobalt. l’rinitritoathylendiamin:rulminkobalt. Wie das Trianiminkobalt zeigt e~ die Eigenschaft. in zwei so sehr
yon einander verschiedenen und bestgudigen Krystallgestalteri aufzu-
trrten, dass man :LU isomere Formen denken kiinnte. Da nber ein-
schlagige Versuche geieigt hat~en, dass die Ansiclit \-on J i i r g e n s e n l ) ,
die von dem Eiiien von uns [We r n e r 2)] a18 isornrr bezeichneteo Formen
des Trinitritotriamriiinko~~:ilts seien auf Lhnorphie zurlckzufuhren, wahr-
sctieinlich riclitig ist, s o sollen nrich die beiden Pormen des Trinitri-
toathylendiaminamniinkob:~lts vord+i hand niir als dirnnrphe Modifica-
tionen au fgefasst w erd en. Durch Einwirkung von kochender Salzsiiure verwandelt sich die
neue Trioitritoverbindung in riii schwarzes Salz, welches nach Kry-
atallisation aus s:ilzsiiurehalti~erii Wasser grlnschwaree Nadeln bildet
cind sich durch seiue Zusarnrnmsetzung uiid sein Verhalten als athylen-
diaminheltiges Dichrosalz erweist. g
Es konimt ihm folgende Constitutii,nJforineI zu: Nur e i n Chloratorn dieses Salies zeigt den gewiihnlichen doppelten
Umsatz, wie die Einwit kung von Salpetersaure letlrt, welche folgendes
Nitrat lielei t : I) Zeitscbr. fiir anorg. Chem. 13, 1SO [189G]. I) Zeitscbr. fiir anorg. Chem. 13, 1SO [189G]. 2, Ztsitscbr. fiir anorg. Chem. 8, 171 [1S95]. 2, Ztsitscbr. fiir anorg. Chem. 8, 171 [1S95]. In wassriger Losung rrfahrt e i ti intraradicales Cbloratorn, infolge
VOII Hydratation, sehr rasch rlektrolptische Dissociation, sodass die
anfanglich griine Losung blao wird. I) Dieser Niederschlag besteht aus : [(Oa N)g Co :en)a]. [(OS
N)dCo (K tj;>j],
wic in einer spLteren Abhatodlung gezoigt wird das komtnt bei Gleicbgewicbtastudien, wie diejenigen von S c h e n c k
iind H e l l e r , vallkommen auf da-selbe hinaos. Ein Salz des dann in der Lo-
sung
vorhandenen
C h lor o- d i a q uo -a t h y I end i a m i n-a m rn i n - k o -
b a l t r a d i c a l e , namlich das Oxalat: C1 CO (NHj) & 0 4 ,
[
(O;::;] ,
C1 CO (NHj) & 0 4 ,
[
(O;::;] [
in welchem sich Oxalsiiure durch analytische Reagentien sofort, Chlor
dagegen erst uach IangeIeiii Kochen nachweisen lasst, ist schwer 16s-
lich und kann deshalb leicht isolirt werden. Wird die blaue Liisung des Chlorids [nit RrornwasserstoffsZur~
versetzt, so scheiden sich olivengriine h'adehl
aus. Die so gebildete
Verbindung zeigt iu ihreni complexen positiven Radical die gr6sste. bis jrtzt i n complexen Ioneri brobacbtete Variation von Componmten,
denn sie ist als Chloro-bromo-aquo-Bthylendiamin-ammin-
k o b ; i l ~ i b r o n i i d zu bezeichnen, d. h. in directer Bindung mit dem
Rob;, ltatoni befinden sich: eiu Chloratorn, ein Bromatoin, ein Wasser-,
ein Animoniak- und ein Aethylendiamin-Molekul. Die Forrnel der
Verbindung ist folgendermaassen 711 schreibm: W irkt KromwasseratoHbIure auf eine ill der WIrme bereitete Lo-
rung von Dichloroaqu~ithylendiaminarnminkobaltchlorid ein, so werdeii
siimmtliche Chloratonie durch Brom ersetzt, und man gelangt zu dem
stark dicbroytischen D i brorno-:tcl uo - a t h y le ndiamin - a m m in- ko-
bnlt tlromid: 1)ie Erystalle dieser Verbinduilg erscheiueu, je nach der Krystall-
axe, blauschwarz oder braun; eerdruckt man sie, so grben sie eii~
griines Pulver. Sowohl die beschriebene leichte Ersetzbarkeit der Chloratome als
anch die leichte Bildung von Diaquooxalat u. s. w. zeigm, dass der
Eintritt ron Aethylendiamin in das Triamminkobaliradical die Beweg-
lichkeit der an Kobalt gebundenen Saurereste begiinstigt, sodas8 die
innere Stabilitiit der iii Form von Salzen nachgewieseiien complexen'
Kobaltradicale wesentlicli grringer ibt 11s bei den gawiihnlicheii Tri-
arnminsalzeii. Infolgedessen sind auch die eiiizelneii VerLinduugstypen
bei den ithylei~diaminhaltigen Verbindungen weniger charakteristisch
:rls bei den gew6hnlichen Triamminsalzen. Es wird aber erst durch 4036 eine eingehendere Untersucbung der einzeluen Salze , specie11 in phy-
sikalisch chernischer Richtung, rniiglich sein, den Unterwhied in den
Htabilitatsoerhaltniesen genauer festzustrllen. eine eingehendere Untersucbung der einzeluen Salze , specie11 in phy-
sikalisch chernischer Richtung, rniiglich sein, den Unterwhied in den
Htabilitatsoerhaltniesen genauer festzustrllen. T ri n i t r i to-ath y 1 end i am i n- a n i niin- ko b a It , [(OrN)sCo (NH3)
(en) I . hei der Modification in Blattchen: Co 21.94, N 29.27. D
))
,)
n
D Nadeln:
))
21.83,
)) 29 15. Di c h 1 o r o- aq u 0-9 t ti y 1 e n d i a m in - a m ni i 11 - li o b a1 t c h 1 o r i d , Die Darstellung dieses Salzes aus dem Nitritos.ilz ert'olgt :iiialog
der von S. M. J o r g e n s e n ' ) aogegebenen Darstelluug von Dichloro-
aquotriamminkobaltchlorid aus Tiinitritotriamminkobalt. 10 g Nitrit werden mit 80 ccm 1,oncentrirter Salzsiiure verrieben
and kurze Zeit (3 Minuteii) au1' dein Wavserbade erhiczt, wobei leb-
hafte Entwickelurrg von salpetriger Same eintritt. Sofort beginnt die
Ansscheidung eiiies schwarzen Salzes, weiche bcim Stehen des Re-
actionsgemisches in der Kiilte nach einigen Stunden beendet ist. Deri abgesaugten, durch Waschen mit Alkohol von Saurr be-
freiten Niederschlag 16st man in so wenig Wasser, dasa ein kleiner
Theil ungeliist bleibt, und filtrirt. Das Filtrat versetzt man sofort mit
25 cc'm concentrirter Salzsaure und fugt nach Ablauf einer Stunde
ooch einaial 25 ccm und nach 2 Stunden noch 50 ccrn hinzu. Das
Salz scheidet sic11 dabei in Form kleiner, griinschwarzer Nadeln ans,
welcbe scliwachen Dichroismus zeigen. Gewinnt man das Salz durch
Einleiten von Salzsauregas in die wassrige Losung, so bildet es ein
graues Pulver, aber die Ausbeuten sind grBsser. Die auf einem Saugfilter gesammelten und mit saurehaltigeni
Wasser und Alkohol gewaschenen Krystalle ergaben bei der Analyse: 0.1774 g Sbst.: 0.0826 g HsO. - 0.2305 g Shst.: 03845 g AgCl. - 0.1274 p
Sbst.: 18.7 ccm N (15O, 720mm). 0.0886 g Sbst.: 0.0532 g COSO~. - 0.1774 g Sbst.: 0.0505 g CO,. - 0.1774 g Sbst.: 0.0826 g HsO. - 0.2305 g Shst.: 03845 g AgCl. - 0.1274 p
Sbst.: 18.7 ccm N (15O, 720mm). 0.0886 g Sbst.: 0.0532 g COSO~. - 0.1774 g Sbst.: 0.0505 g CO,. - C O N ~ G C I ~ H I S O . Ber. Co 22.64, N I G . l G , C 9.20, H 5.04, C1 40.81. Gef. )) 22.84,
)) 16.25,
7.76, )) 5.17,
)) 41.17. C O N ~ G C I ~ H I S O . Ber. Co 22.64, N I G . l G , C 9.20, H 5.04, C1 40.81. Gef. )) 22.84,
)) 16.25,
7.76, )) 5.17,
)) 41.17. I ) Zeitsclir. f. anorg. Chem. 14, 418-419. T ri n i t r i to-ath y 1 end i am i n- a n i niin- ko b a It , [(OrN)sCo (NH3)
(en) I . Fiigt man zu einer Liisung YOU Tetranitritodiarnminkobaltnatrium
bei 60'' Aethylendiamin (Monohydrat) und zwar 1.4 g auf 5 g des
Yalzes, so scheidet sich ein gelber, k rystallinisrher Niederschlag ') ails,
wHhrend in der braungefarbten Mutterlauge eine Anzahl leichter 16s-
licher Reactionsproducte enthalten ist. Dampft man die Muiterlauge auf ungefahr die Halfte itirrs ur-
spriinglichen Volumens ein , so scheidet sich nach dem Erkalteri Tri-
~iitritoathylendiarninarnmiokobalt als gelber Kryetallbrei aue. dem aber
iioch andere Salze heigemisclit sind. Nach drei- bis vier-maligem
Umkrystallisiren ails essigaaurehaltigrrn Wabser erhalt man ea rein,
in Form hellbrauner Blattchen, welche in Wasser schwer liislich sind. 0.0578 g Sbst.: 0.0508 g CoSO,. - 0.07i3 g Sbst.: 0.0417 g CoSOh. -
0.0787 g Sbst.: 22.2 ccm N (l3.So, 720 mrn). - 0.07f3 g Sbst.: 20 2 CCIU N
# 1 3 O , 722 mm). - 0.1518 g Sbs:.: 0.04s; g COT. 0.0765 g Sbst.: 0.0442 a CO SO&. - 0.0974 R Sbst.: 0.0562 g C O S O ~ . - ,
)
g
; g
(JoNsC.rOaH11. Ber. Co B l . i S ,
N 3O.ti5,
c 8.84. Gcf. 21.94, 21.98, 21.99, 22.00,
n 31.45, 29.95,
D 8.66, , ,
g
; g
(JoNsC.rOaH11. Ber. Co B l . i S ,
N 3O.ti5,
c 8.84. ,
Gcf. 21.94, 21.98, 21.99, 22.00,
n 31.45, 29.95,
D 8.66, , Dieser Zusarnmrnsetzung entepricht folgende Formel: Dieser Zusarnmrnsetzung entepricht folgende Formel: Die Nitritgruppen verhalten sich nicht als Iorieu, iind dement
eprechend kann man das Salz mit kalter Salzeiiure iiberschichten,
( h e dass vine Entwickelung von salpetriger Saure bemerkbar wire. Natiirlich wirkt die Salzsiiure ill der Warrne daranf ein. Hierbei
entsteht ein schwarzes Salz, welches, nach allen seinen Eigenschaften
zu schliessen, dae Analogon des Dichloroaquotriarnminkobaltchlorids
(Dichroaalz) ist. Damit ist bewiesen, dass nnsere Verbindung einer
Triarnminreihe angehort. Dae Trinitritoathylendiaminarnminkobalt wurde gelegentlich auch
i n einer anderen Krystallform erhalten, obwohl die Daretellungaweise
i n keiner Weiee von der bereits angegebenen abwich. 4037 Es krystallisirte in ungefiihr 2 cm langen, breiten, federartig
gestreiften Nadeln. Nach Analyse und Verhalten hat dieae Modifi-
cation dieselbe Zusammensetzung wie die Rlattchenform. g
0.0807 g Shst.: 0.0464 g CoSO,. - 0.0978 g Shst.: 28.4 ccm N (14O, 713mm). Gef. hei der Modification in Blattchen: Co 21.94, N 29.27. D
))
,)
n
D Nadeln:
))
21.83,
)) 29 15. g
0.0807 g Shst.: 0.0464 g CoSO,. - 0.0978 g Shst.: 28.4 ccm N (14O, 713mm). Gef. T ri n i t r i to-ath y 1 end i am i n- a n i niin- ko b a It , [(OrN)sCo (NH3)
(en) I . In eiskaltem Wasser 16st pith das Salz mit griiner Farbe, welche
bald in blan umschlagt; bei weiterem Verdiinnen wird die LBsuog
roth, und dann tallt auf Zusatz von Silbernitrat sammtliches Chlor als
Ghlorsilber nus. Erhitzt man die neutrale Losung des Salzes, so tritt
Zersvtzung unter Abscheidung von Kobalti hydroxyd ein. Wird mit I ) Zeitsclir. f. anorg. Chem. 14, 418-419. 4038 Salzsaure erhitzt, so erhalt man als Zersetzungsproduct blaue blattrige
Krystalle, welcbe sich in Wasser mit rother Farbe 18sen. Diese LB-
sung wird bei Zusatz eines Tropfens Salzsaure blau und ecbeidet
heirn Umriihren salzsaures Aethylendiarnin aus, wiihrend Kobaltchloriir
i n Lijsung hleibt. Die hlauen Krystallc sind dernnnch das bpkannte
Chlorokobaltoat des Aethylendiamins. Auf Ziisatz der berechneten Menge Natriumnitrii wird die schwach
saure L8sung des Cblorids gelb gefiirlit und scheidet nach einiger
Zeit Trinitritoathylendiaminamrninkobxlt in den
charakteristischeir
Formen aus. Nitrat,
CIzCo(NH3) NO?. i
“I Wird fein gepulvertes, angefeuchtetes Chlorid rnit Salpetersaure
vorn spec. Gewicht 1.4 in solcher Menge verrieben, dass ein diinn-
Hiissiger Brei entstebt, so wird die Maese nach einiger Zeit ziernlich
zahfliissig. Man wascht sie dann rnit Wasser, um die Salpetersanre
zn entfernen, liierauf rnit Alkotiol und trocknet. Der Process wird
dreimal wiederholt, worauf das Nitrat rein ist. Es wird in wenig
kaltern Wasser geltist und die filtrirte Lasung mit einigen Tropfen
concentrirter Salpetersaure versetzt. Rei laugsamem Verdonste‘n der
LBsung krystallisirt das Salz in g r h e n , metallisch glaozenden Flittern. In Wasser ist es ausser~t leicht ldslich. In Wasser ist es ausser~t leicht ldslich. 01021
Sb t
0 0553
CoSOa
0 2045
Sh t
0 2015 g A CI - 0.1021 g Sbst.: 0.0553 g CoSOa. - 0.2045 g Shst.: 0.2015 g AgCI. -
0.1485 g Sbbt.: 26.2 ccrn N (1.50, 723 mm). CoClNdC12HtoO. Ber. Co 20.58, K 19.59, C1 24.73. Gef. n 20.61,
I) 19.61,
D 24.36. CoClNdC12HtoO. Ber. Co 20.58, K 19.59, C1 24.73. Gef. n 20.61,
I) 19.61,
D 24.36. C b lo ro-d i aq uo-a t h y 1 e n d i a rn i n- a m m i II - k o b a 1 to x a1 a t . C b lo ro-d i aq uo-a t h y 1 e n d i a rn i n- a m m i II - k o b a 1 to x a1 a t . ClCO (NHB) God. T ri n i t r i to-ath y 1 end i am i n- a n i niin- ko b a It , [(OrN)sCo (NH3)
(en) I . Erhitzcn auf 105O nurmehr O.lti17 g. g
p
Ber. fur obige Formel: 1 Mol. HsO -= 6.05 pCt. Ch I o r n - b r o m 0-
a q u o-at 11 y 1 end i a min- a m m i 11 - k o b a 1 t b r o m id , Ch I o r n - b r o m 0-
a q u o-at 11 y 1 end i a min- a m m i 11 - k o b a 1 t b r o m id , Lost u~aii das Chlorid in der eben nothwendigen Menge Wasser
auf iind setzt das gleiche Volurnen Bromwasserstoffsaure vom spec. Gewicht 1.49 zii, so scheiden sich sofort kleine, olivengriin gefarbte
Nadeln am, die rnit Alkohol gewaschen und neben Natronkalk ge-
trocknet werden. Die Zusammensetzuog entspricht dem aiich in der
reineo Triarnminreihe sich bildenden Chlorobromobromid. Chlor und Brom wurden in dcr Weiw heetimmt, dass ein Theil des
Halogensilbrrs eine Stunde lang im Chlorstrorn erhitzt nnd die in Folge der
Ueberfiihrung des Bromsilbers in Chlorsilber entstandene Gewichtsverminde-
rung auf die ganzc! Halogensilbermenge umgerechnet wurde. g
g
g
g
g
0.2033 g Sbst.: 0.3021 g Halogensilber. - 0.2034 g Sbst.: 0.3022 g Ha-
logensilber. - 0.2845 g Halogensilber crgabeii, im Chlorstrom erhitzt, eine
Gewichtsabnahme von 0.0479 8; den, entspricht fur 0.3022 g eine Abnabme
von 0.0513 g, woraus sicb 0.21584 g Bg Br und 0.08636 g AgCl berecbnen. - 0.106 g Sbst.: 0.0468 g COSOJ. - 0.115 g Sbst.: 12.4 ccm N (13q 703 mm). CoBr2ClC?NgH130. Ber. Co 16.88, N 12.05, Br 45.74, C1 10.14. Gef. n 16.80,
D 11.75,
))
45.14, * 10.49. g
g
g
g
g
0.2033 g Sbst.: 0.3021 g Halogensilber. - 0.2034 g Sbst.: 0.3022 g Ha-
logensilber. - 0.2845 g Halogensilber crgabeii, im Chlorstrom erhitzt, eine
Gewichtsabnahme von 0.0479 8; den, entspricht fur 0.3022 g eine Abnabme
von 0.0513 g, woraus sicb 0.21584 g Bg Br und 0.08636 g AgCl berecbnen. - 0.106 g Sbst.: 0.0468 g COSOJ. - 0.115 g Sbst.: 12.4 ccm N (13q 703 mm). CoBr2ClC?NgH130. Ber. Co 16.88, N 12.05, Br 45.74, C1 10.14. Gef. n 16.80,
D 11.75,
))
45.14, * 10.49. Eine noch nicht naher untersuchte Verhderung erleidet die Ver-
binduug durch Erhitzen: g
0.1518 g Sbst. wogen nach einstiindigem Erhitzen iin Trockenschrd
0.1437 8 ; Abnahmc = 0.0081 g = 5.34 pCt. Bw. T ri n i t r i to-ath y 1 end i am i n- a n i niin- ko b a It , [(OrN)sCo (NH3)
(en) I . [
(:e:)al [
5 g Chlorid werden mit eiuer Loaung von 5 g Oxalsaure iii 50 ccm,
Wasser; sowie rnit 50 ccrn Alkohol versetzt und eine Minute erhitzt. Reim Erkalteu scheidet sich das Salz nus. Es bildet blaue, silber-
gliinzende Krystallblattcben, die auf dern Filter durch Waschen rnit
salzsaurehaltigern Wasser und Alkohol gereinigt werden. In kaltem
Wasser ist es sehr schwer, in heissem leichter Ioslicb; in Alkohol ist
es fast uulrislich. Die wassrige Losung giebt rnit Salzsaure wieder
Dichlorochlorid. Aus arnrnouiakhaltiger Liisung entsteht auf Zusatz
\-on Chiorcalcium ao fort ein weisser Niederschlag von Calciurnoxalat. Mit Silbwnitrat giebt die wassrige Lijsung anfanglich nor pine schwacbe Triibung, und erst nacb Iangerem Steheu fallt d:is Cblor rollstindig
aus -- ein Bewris dafiir. dass das Chlor nicht als Ion wirkt. Triibung, und erst nacb Iangerem Steheu fallt d:is Cblor rollstindig 0.1063 g Sbst.: 13.7 ccm N (170, 723 mm). - 0.2834 g Sbst.: 0.13YY g
AgCI. - 0.1260 g Sbst.: 0.0585 g H20. - 0.1260 g Sbst.: 0.0766 g Con. -
0.08li4 p Sbst.: 00150 g CoSOi. 0.1063 g Sbst.: 13.7 ccm N (170, 723 mm). - 0.2834 g Sbst.: 0.13YY g
AgCI. - 0.1260 g Sbst.: 0.0585 g H20. - 0.1260 g Sbst.: 0.0766 g Con. -
0.08li4 p Sbst.: 00150 g CoSOi. 0.1063 g Sbst.: 13.7 ccm N (170, 723 mm). - 0.2834 g Sbst.: 0.13YY g
AgCI. - 0.1260 g Sbst.: 0.0585 g H20. - 0.1260 g Sbst.: 0.0766 g Con. -
0.08li4 p Sbst.: 00150 g CoSOi. p
g
C O C ~ N B C I O ~ H ~ ~ , . Uer Co l!l.Y6, I\i 14.02, Cl 11.97, H 5.11, C 16.23. Gef. ))
19.81,
I) 14.30, 8 12.19,
)) 4.63. 16.56. p
g
C O C ~ N B C I O ~ H ~ ~ , . Uer Co l!l.Y6, I\i 14.02, Cl 11.97, H 5.11, C 16.23. Gef. ))
19.81,
I) 14.30, 8 12.19,
)) 4.63. 16.56. stanz eineii Gewichtsverlust, der eiriem MolekLil Wasser entspricht. Gewichtsvcrliist = 0.01 g = 5.82 pct. RrO. Heini Erhitzeri erleidet die iiber Schwefelsaure getrocknete Sub-
0.1717 g Shst. wogen nach I-stdg. Erhitzcn auf 105O nurmehr O.lti17 g. Ber. fur obige Formel:
1 Mol. HsO -=6.05 pCt. stanz eineii Gewichtsverlust, der eiriem MolekLil Wasser entspricht. Heini Erhitzeri erleidet die iiber Schwefelsaure getrocknete Sub- ,
p
Gewichtsvcrliist = 0.01 g = 5.82 pct. RrO. 0.1717 g Shst. wogen nach I-stdg. D i b r o m o - a q u o - 6: I I , y I end i a rn in- a m m in- k o b a 1 t b r o m i d , D i b r o m o - a q u o - 6: I I , y I end i a rn in- a m m in- k o b a 1 t b r o m i d , D i b r o m o - a q u o - 6: I I , y I end i a rn in- a m m in- k o b a 1 t b r o m i d , D i b r o m o - a q u o - 6: I I , y I end i a rn in- a m m in- k o b a 1 t b r o m i d , Zur Darstelluug dieses Salzes liist m:in das soeben beschriebenc
Chlorobromobromid in etwas mehr als der nothigen Menge IV'asser
:tiif,
erwarmt die Losung schwech
:tiif dem Wasserbade und setzt
nach erfolgter Abkiihlung daa gleiche Volumen Kromwasserstoffsiiure
(spec. Grwicht 1.49) hiinu. Nach 12 Stdn. ist das Salz augkrystalli-
sirt. Es bildet kleine. aber wohlansgebildete, sternformige Nadel-
:iggregate und zeigt einen Y lei ausgepriigteren Dichroismus als das
entsprechende Dichloroch!orid. Die Krystalle erscheinen blauscbwarz
oder braun gefiirbt uud geben beini Zerreiben ein griines Pulver. In Wasser lost sich das Salz mit brauner Farbe. 0.1102 g Sbst.: 0.0443 g CoSO,. - 0.1060 g Sbst.: I0 ccm N (ll*,#. 711 mm). - 0.1466 g Sbst.: 14.2 ccm N (141/2", 724 mm). - 0.1003 g Sbst.:
0.2872 g AgBr. CoBr3N3C2H130. Ber. Co 14.92, N 10.69,
Br 60.91. Gef. D
15.'?Y,
10.47, 10.80, D FI.01. Ziirich, Unirersitatslaboi atorium, December 1905. Ziirich, Unirersitatslaboi atorium, December 1905. I) Diese Berichte 82, 2197 [l899].
2, Arch. f. Pharm. 234, 487 [1896l. 704. H. K i l i a n i : Ueber Digitoxose.
[Aus der medic. Abth. des Univer3.-Laboratoriums Freiburg i. B.;
(Eingegsngen am 5. December 1905.) (Eingegsngen am 5. December 1905 Meine friihereli Beobaclitungen hatten es zweifelhaft gelassen, ob die
Digitoxose, CsH1201, ale Aldeliyd oder als Keton zu betrachten sei I). Jetzt habe ich sicher festgestellt, dn53 der Zucker eine Aldose ist. Diese enthalt ein CHS (Y. 1. c.), folglicli muss, w e n n eine normale
Kohlenstoff-Kette vorliegt, geniiiss obiger Formcl ein CH2 vorhanden
sein, das w a h r s c h e i n l i c h die u-Stellung zur Aldehydgruppe ein-
nimmt, weil es nicht gelungeu war, ein Osazon ZII gewinnen2). Da-
mit ergab sich aber sofort Eiir die drei Hydroxyle die Stellung ani
$-, y- und 8-Kohlenstoff, d. h. die C o n s t i t u t i o n d e r D i g i t o x o s r
rnusste sein: Ziirich, Unirersitatslaboi atorium, December 1905. T ri n i t r i to-ath y 1 end i am i n- a n i niin- ko b a It , [(OrN)sCo (NH3)
(en) I . fiir 1 Mol. HnO = 5.15 pCt 0.1518 g Sbst. wogen nach einstiindigem Erhitzen iin Trockenschrd
0.1437 8 ; Abnahmc = 0.0081 g = 5.34 pCt. Bw. fiir 1 Mol. HnO = 5.15 pCt Dabei war die Substanz zusarnmengrsintert
und bildete e k e
gelbe Kriistp. die sich ill Wasser leicht mit gelber Farbe liiste. Dabei war die Substanz zusarnmengrsintert
und bildete e k e
gelbe Kriistp. die sich ill Wasser leicht mit gelber Farbe liiste. A 118 der Liisung des Clilorobroniobromids fiillt Jodkaliumliisung
ein scliwer IiislicheR Jodid, und diirch Vcrreiben des trocknen Salzes
rnit conceriti irter Salpetereaure bildet sich ein sehr leicht lijslicheq
hellgrCines Sitrat. 4040 CHx.CH(OH) .CH (OH) .CH(OH) .CX2 CHO. CHx.CH(OH) .CH (OH) .CH(OH) .CX2 CHO. CHx.CH(OH) .CH (OH) .CH(OH) .CX2 CHO.
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https://openalex.org/W4230099459
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|
English
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Association of Intradialytic Hypotension Events With All-Cause Mortality in Hemodialysis Patients
|
American journal of kidney diseases
| 2,016
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cc-by
| 347
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UC Irvine
UC Irvine Previously Published Works
Title
Association of Intradialytic Hypotension Events With All-Cause Mortality in Hemodialysis
Patients
Permalink
https://escholarship.org/uc/item/4cr9d8d3
Journal
American Journal of Kidney Diseases, 67(5)
ISSN
0272-6386
Authors
Chou, Jason
Obi, Yoshitsugu
Kim, Tae Hee
et al.
Publication Date
2016-05-01
DOI
10.1053/j.ajkd.2016.03.075
Copyright Information
This work is made available under the terms of a Creative Commons Attribution License,
availalbe at https://creativecommons.org/licenses/by/4.0/
Peer reviewed UC Irvine
UC Irvine Previously Published Works
Title
Association of Intradialytic Hypotension Events With All-Cause Mortality in Hemodialysis
Patients
Permalink
https://escholarship.org/uc/item/4cr9d8d3
Journal
American Journal of Kidney Diseases, 67(5)
ISSN
0272-6386
Authors
Chou, Jason
Obi, Yoshitsugu
Kim, Tae Hee
et al. Publication Date
2016-05-01
DOI
10.1053/j.ajkd.2016.03.075
Copyright Information
This work is made available under the terms of a Creative Commons Attribution License,
availalbe at https://creativecommons.org/licenses/by/4.0/
Peer reviewed UC Irvine Copyright Information
This work is made available under the terms of a Creative Commons Attribution License,
availalbe at https://creativecommons.org/licenses/by/4.0/ Am J Kidney Dis. 2016;67(5):A1-A118 Am J Kidney Dis. 2016;67(5):A1-A118 A35 NKF 2016 Spring Clinical Meetings Abstracts NKF 2016 Spring Clinical Meetings Abstracts Powered by the California Digital Library
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https://openalex.org/W4241539654
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https://journals.iium.edu.my/ejournal/index.php/iiumej/article/download/1706/818
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English
| null |
MODELLING A 1-DOF FINGER EXTENSOR MACHINE FOR HAND REHABILITATION
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IIUM engineering journal
| 2,021
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cc-by
| 5,742
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KEYWORDS: modelling; simulation; experimental validation; hand rehabilitation;
hardware-in-the-loop IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. 1. INTRODUCTION Every year, up to 15 million people suffer from stroke, making it one of the leading
causes of severe disability in the world [1]. After a neurological injury such as stroke,
patients suffer from several impairments, of which the most common and impeding is the
impairment to the hand sensorimotor function. The finger extensor muscles develop
weakness during voluntary movements as a result of activity deficit. This leads to
spasticity which is a velocity-dependent increase in muscle tone. Spasticity causes intense
muscle contractions and stiffness that manifest in the form of a clenched fist, tensed
fingers, a curled wrist and other deformities. These impairments lead to a decreased ability
to open and close the hand, control individual finger movements and perform force
coordination, thereby severely affecting the quality of life of stroke patients [2]. Patients undergo intense physical therapy under the guidance of an occupational
therapist to regain functionality in the affected limb. Activity-based exercises are known to
be effective in helping overcome the motor deficits induced by post-stroke spasticity [3]. However, occupational therapy is expensive and as a result, a vast majority of the patients
do not have continuous access to it. Besides, conventional therapy is monotonous,
repetitive and difficult to be sustained for prolonged periods of time, leading to high drop-
out rates. Furthermore, with conventional therapy, it is not possible to measure the exact
progress of patients over time and provide them with well-regulated and a wide range of
rehabilitative forces. Due to these factors, robot-assisted rehabilitation has seen a growing
interest in the last few decades. Robotic devices, due to their features of high repeatability,
interactivity, ability to provide a wide range of accurate forces and automatic measurement
of patient progress, are becoming increasingly popular. The interactive nature of robot-
aided therapy not only increases physical engagement but also encourages cognitive
engagement of the patient through immersive and challenging exercises. Patient
engagement in rehabilitation is vital because encouraging human users to perform self-
initiated movements is an essential requirement to achieve motor learning and
neuroplasticity [4]. Furthermore, robot-aided rehabilitation is capable of producing assistive as well as
resistive forces that can be tailored to the specific impairment of the patient. When patients
gain some amount of motor function in the impaired part, they are subjected to active-
resistive therapy whereby, forces are applied on the patient in a direction opposite to the
one they are tracing. *Corresponding author: sinnira@iium.edu.my (Received: 24th November 2020; Accepted: 22nd December 2020; Published on-line: 4th July 2021) ABSTRACT: It is essential to have an accurate representation of a robotic rehabilitation
device in the form of a system model in order to design a robust controller for it. This
paper presents mathematical modelling and validation through simulation and
experimentation of the 1-DOF Finger Extensor rehabilitation machine. The machine’s
design is based on an iris mechanism, built specifically for training open and close
movements of the hand. The goal of this research is to provide an accurate model for the
Finger Extensor by taking into consideration various factors affecting its dynamics and
to present an experimental validation of the devised model. Dynamic system modelling
of the machine is performed using Lagrangian formulation and the involved physical
parameters are obtained experimentally. To validate the developed model and
demonstrate its effectiveness, hardware-in-the-loop experiments are conducted in the
Simulink-MATLAB environment. Mean absolute error between the simulated and
experimental response is 1.38° and the relative error is 1.13%. The results obtained are
found to be within the human motion resolution limits of 5 mm or 5º and exhibit
suitability of the model for application in robotic rehabilitation systems. The model
accurately replicates the actual behavior of the machine and is suitable for use in
controller design. ABSTRAK: Gambaran tepat mengenai model sistem peranti rehabilitasi robotik adalah
sangat penting bagi pembangunan sesebuah reka bentuk alat kawalan tahan lasak. Kajian
mengenai model matematik dan pengesahan melalui simulasi dan eksperimentasi mesin
pemulihan 1-DOF ‘Finger Extensor’. Mesin ini direka bentuk berdasarkan mekanisme
iris, dibangunkan khusus bagi melatih gerakan buka dan tutup tangan. Tujuan kajian ini
adalah bagi menyediakan model Finger Extensor yang tepat dengan mengambil kira
faktor mempengaruhi dinamik dan pengesahan model eksperimen yang dirancang. Model sistem dinamik mesin ini diuji menggunakan formula Lagrangian dan parameter
fizikal yang terlibat diperoleh melalui eksperimen. Model ini disahkan dan diuji
keberkesanannya
menggunakan
eksperimen
Perkakasan-dalam-gelung
melalui
MATLAB-Simulink. Purata ralat mutlak antara dapatan simulasi dan respon eksperimen
adalah 1.38° dan ralat relatif 1.13%. Dapatan kajian adalah dalam had resolusi gerakan
tangan manusia iaitu 5 mm atau 5º dan didapati model ini sesuai bagi aplikasi sistem
rehabilitasi robotik. Model ini tepat dalam mereplikasi kelakuan sebenar mesin dan
sesuai digunakan bagi reka bentuk kawalan. 384 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. Before the implementation of these control strategies, dynamic and kinematic
modelling of the rehabilitation machine is necessary. Modelling is the mathematical
representation of a physical system [14]. A system model is developed from mathematical
equations and then represented graphically in software such as MATLAB, Simulink and
StateFlow, etc. The developed model is then validated through simulations, reducing the
cost associated with experimental validation. During the process of model development,
trade-offs have to be made between the fidelity and simplicity of the model. Fidelity
implies the extent to which the system model mimics the behavior of the real system and
simplicity ensures that the model remains simple enough for smooth controller
development in later stages. Thus, there are some deviations between the behavior of the
actual system and its model. After balancing simplicity and fidelity, the model is validated
using a Hardware-in-the-Loop (HiL) approach. In this approach, one or more real
subsystems interact in a closed loop with the virtual subsystems of the model. A
comparison between the response of such a closed-loop interaction with the simulation
output reveals performance of the developed model [15]. In [16], dynamic and kinematic modelling of the multipurpose rehabilitation robot,
the Universal Haptic Pantograph, was performed. Validation of the model was carried out
through experiments with healthy subjects reflecting its performance. Dynamic modelling
and experimental validation of a parallel robot for upper limb rehabilitation was presented
in [17]. Approximately zero steady-state errors points to a high degree of accuracy and
suitability of the developed model. Dynamic modelling of a haptic finger actuated by a
McKibben artificial muscle built for tele-operated rehabilitation was performed in [18]. Upon experimental validation, the model demonstrated a satisfactory performance. In this work, mathematical modelling and system simulation of a 1 degree-of-freedom
(DOF) Finger Extensor rehabilitation machine was performed. A Model-Based
Development (MBD) [15] approach to controller design was taken. Such an approach to
design enabled verification and validation at each stage. Based on simulation results,
modifications to the system could be made easily without incurring high costs. As a first
step, mathematical modelling was done, followed by simulation of the system behavior. Simulation results were then compared to hardware measurements which demonstrated the
accuracy of the realized model. Hardware-in-the-Loop testing of the model carried out in
this study ensures minimization of risks and costs associated with experimental validation. IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 The developed model aids in the design and implementation of a control algorithm for the
Finger Extensor machine. 1. INTRODUCTION Resistive strategies enhance patient performance as they require
additional effort from the patient to resist opposite forces. Such efforts induce
neuroplasticity and help in development of neural pathways from the impaired muscles to
the brain which aids in regaining of motor function [5]. However, all these features of robotic rehabilitation devices are possible only when
efficient controllers are developed [6]. A variety of control strategies have been proposed
for rehabilitation robots over the past few decades. These control schemes are designed
around the central idea of modulating assistance/resistance provided to the patient based
on various factors such as force exerted by the patient, limb velocity, tracking error,
interaction force/torque, EEG/EMG activity [7]. In recent literature, approaches such as force/stiffness control [8] Assist-as-Needed
control (AAN) [9-11] and performance-based impedance control [7,12,13] have been
implemented on rehabilitation robots. AAN control aims at providing minimal
intervention in patient recovery. Performance-based impedance control modulates the
assistance provided, based on real-time measurements of the patient's biomechanical
characteristics. 385 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 3.1 Mechanical Model Newton-Euler and the Lagrange-Euler methods are the most commonly used
approaches for deriving dynamic equations of mechanical parts of robots [20]. The former
is ideal for on-line control applications due to reduced computational time. However, it
falls short in providing an adequate insight into control design due to recursive
computation. The Lagrange-Euler method is simple and systematic, providing dynamic
equations in a matrix form and was chosen for deriving the mechanical model for the
Finger Extensor. Dynamic equation describing motion of the Finger Extensor was written
as 𝑀(𝜃(𝑡), 𝜉)𝜃̈(𝑡) + 𝐷(𝜃(𝑡), 𝜃̇(𝑡), 𝜉) + 𝐺(𝜃(𝑡), 𝜉) = 𝑇(𝑡)
(1)
where, 𝑀(𝜃(𝑡), 𝜉)𝜃̈(𝑡) + 𝐷(𝜃(𝑡), 𝜃̇(𝑡), 𝜉) + 𝐺(𝜃(𝑡), 𝜉) = 𝑇(𝑡)
(1)
where, (1) 𝑀(θ(𝑡), ξ) is the N x N inertia matrix. 𝑀(θ(𝑡), ξ) is the N x N inertia matrix. 𝐷(𝜃(𝑡), 𝜃̇(𝑡), 𝜉) is the N x 1 vector of coriolis and centrifugal forces. 𝐺(𝜃(𝑡), 𝜉) is the N x 1 vector of gravitational forces. 𝑇(𝑡) is the N x 1 vector of torques applied by the actuators. 𝐷(𝜃(𝑡), 𝜃̇(𝑡), 𝜉) is the N x 1 vector of coriolis and centrifugal forces. 𝐺(𝜃(𝑡) 𝜉) is the N x 1 vector of gravitational forces 𝐷(𝜃(𝑡), 𝜃̇(𝑡), 𝜉) is the N x 1 vector of coriolis and centrifugal forces. 𝐺(𝜃(𝑡), 𝜉) is the N x 1 vector of gravitational forces. 𝐺(𝜃(𝑡), 𝜉) is the N x 1 vector of gravitational forces. 𝑇(𝑡) is the N x 1 vector of torques applied by the actuators. 𝜃(𝑡), 𝜃̇(𝑡), 𝜃̈(𝑡) are the N x 1 vectors of joint displacements, velocities and
accelerations respectively. 𝜉 is a vector (with appropriate dimension) of parameters of the mechanism such as
payload. For the Finger Extensor, 𝑀(𝜃(𝑡), 𝜉) is a 1x1 matrix, with one element equal to the
moment of inertia of a solid cylinder with iris radius of 0.055m. 𝐷(𝜃(𝑡), 𝜃̇(𝑡), 𝜉) and
𝐺(θ(𝑡), ξ) for a 1-DOF machine were null matrices. 𝑇(𝑡) for the Finger Extensor is
calculated in the subsequent sections. For developing the mechanical model of the Finger Extensor, the next step was to
derive equations of motion of the moving parts of the mechanism. These were represented
by flywheel and spring mechanical elements as shown in Fig. 4. 𝐽1, 𝐽2 represent inertia
elements of the motor and iris mechanisms respectively, 𝑘1, 𝑏1 represent the spring
stiffness effect and viscous and coulomb friction. Fig. 4: Block diagram representation. Fig. 4: Block diagram representation. 2. THE FINGER EXTENSOR MACHINE The Finger Extensor rehabilitation machine is a 1-DOF device with an iris mechanism
lying at the heart of its dynamics. The design of the iris or diaphragm is inspired by the
constriction and dilation of the iris of the human eye, which varies size of the pupil,
adjusting the aperture of the eye. The iris mechanism built into the Finger Extensor
constricts and dilates the poles at the top of its surface, to allow openings of variable sizes. These poles, as shown in Fig. 1 and Fig. 2 are grasped by the patient during therapy and
enable him/her to practice finger flexion and extension [19]. There are six poles in the current Finger Extensor design and each pole has a blade
associated with it. On the upper end, each blade is connected to a slider and at the bottom,
each blade goes into a slot in a disc using a bearing, as illustrated in Fig. 2. The disc with
slots is in turn connected to the motor actuation system through a chain and sprocket
mechanism. As the motor turns, the sprocket induces motion into the slotted disc. This 386 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. https://doi.org/10.31436/iiumej.v22i2.1706 causes the blades to slide and move the poles in their own respective slots at the top of the
machine. Fig. 1: 1-DOF Finger Extensor machine. Fig. 1: 1-DOF Finger Extensor machine. Fig. 2: CAD drawing of the side view of the iris mechanism. Fig. 1: 1-DOF Finger Extensor machine. Fig. 1: 1-DOF Finger Extensor machine. Fig. 2: CAD drawing of the side view of the iris mechanism. Fig. 2: CAD drawing of the side view of the iris mechanism. Fig. 3: Inner view of the machine. Fig. 3: Inner view of the machine. Fig. 3: Inner view of the machine. Figure 3 shows the actuation system of the machine comprising of a brushed planetary
geared DC motor (Gear Ratio- 32.5:1). Rotation of the motor is recorded by a rotary
encoder by ESB electronics, Japan (Counts per Revolution- 500) connected to the sprocket
using a flexible shaft coupling. A torque sensor, connected close to the chain-sprocket
mechanism provides the torque applied by the machine on to the patient. Actuation system
and sensors are controlled by Arduino microcontroller board which is programmed in the
Simulink environment. 387 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. 3.1 Mechanical Model (5), 𝑇𝑔=
1
𝑁𝐽2𝜃2̈ +
1
𝑁𝑇𝑒+ 𝑁𝑘1𝜃2 + 𝑉𝑐(𝜃2̇ ) + 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ )
(6 𝑇𝑔=
1
𝑁𝐽2𝜃2̈ +
1
𝑁𝑇𝑒+ 𝑁𝑘1𝜃2 + 𝑉𝑐(𝜃2̇ ) + 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ )
(6)
where, 𝑉𝑐(𝜃2̇ ) and 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ ) are the viscous and coulomb friction terms. Therefore,
dynamic equation of the 1 DOF Finger Extensor was represented as (6) 𝑇𝑔= 𝑁𝐽2𝜃2 + 𝑁𝑇𝑒+ 𝑁𝑘1𝜃2 + 𝑉𝑐(𝜃2) + 𝐹𝑐𝑆𝑔𝑛(𝜃2)
(6)
where, 𝑉𝑐(𝜃2̇ ) and 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ ) are the viscous and coulomb friction terms. Therefore,
dynamic equation of the 1-DOF Finger Extensor was represented as where, 𝑉𝑐(𝜃2̇ ) and 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ ) are the viscous and coulomb friction terms. Therefore,
dynamic equation of the 1-DOF Finger Extensor was represented as where, 𝑉𝑐(𝜃2̇ ) and 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ ) are the viscous and coulomb friction terms. Therefore,
dynamic equation of the 1-DOF Finger Extensor was represented as 𝑇𝑔(𝑡) = 1
𝑁𝐽2(𝜃2)𝜃2̈ (𝑡) + 1
𝑁𝑇𝑒(𝑡) + 𝑁𝑘1𝜃2(𝑡) + 𝑉𝑐(𝜃2̇ )(𝑡) + 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ )
(7) 𝑇𝑔(𝑡) = 1
𝑁𝐽2(𝜃2)𝜃2̈ (𝑡) + 1
𝑁𝑇𝑒(𝑡) + 𝑁𝑘1𝜃2(𝑡) + 𝑉𝑐(𝜃2̇ )(𝑡) + 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ )
(7 𝑇𝑔(𝑡) = 1
𝑁𝐽2(𝜃2)𝜃2̈ (𝑡) + 1
𝑁𝑇𝑒(𝑡) + 𝑁𝑘1𝜃2(𝑡) + 𝑉𝑐(𝜃2̇ )(𝑡) + 𝐹𝑐𝑆𝑔𝑛(𝜃2̇ )
(7) (7) 3.1 Mechanical Model Forces acting on the motor and iris sprockets are shown in Fig. 5, where 𝜃1 is the
angle of rotation of the motor sprocket, 𝜃2 is the angle of rotation of the iris mechanism,
𝑇𝑔 is the torque on the driver sprocket, 𝑇𝑒 is the torque exerted by the user, 𝑓𝑐𝑟1 and 𝑓𝑐𝑟2 388 eering Journal, Vol. 22, No. 2, 2021
Shahdad et al. g/10.31436/iiumej.v22i2.1706 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. are the contact force torques. Figures 4 and 5, hence represent Eq. (1) in terms of sum of
forces (inertial, gravitational, frictional) acting on the machine. are the contact force torques. Figures 4 and 5, hence represent Eq. (1) in terms of sum of
forces (inertial, gravitational, frictional) acting on the machine. As per D’Alembert’s law, sum of torques on both flywheels was given by the
equations 𝑇𝑔= 𝐽1𝜃1̈ + 𝑘1𝜃1 + 𝑏1(𝜃1̇ −𝜃2̇ ) −𝑓𝑐𝑟1
(2)
−𝑓𝑐𝑟2 −𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒= 0
(3)
From Eq. (3), value of 𝑓𝑐 is calculated as
𝑓𝑐=
1
𝑟2 [−𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒]
(4) 𝑇𝑔= 𝐽1𝜃1̈ + 𝑘1𝜃1 + 𝑏1(𝜃1̇ −𝜃2̇ ) −𝑓𝑐𝑟1
(2)
−𝑓𝑐𝑟2 −𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒= 0
(3)
From Eq. (3), value of 𝑓𝑐 is calculated as
𝑓𝑐=
1
𝑟2 [−𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒]
(4) 𝑇𝑔= 𝐽1𝜃1̈ + 𝑘1𝜃1 + 𝑏1(𝜃1̇ −𝜃2̇ ) −𝑓𝑐𝑟1
−𝑓𝑐𝑟2 −𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒= 0
From Eq. (3), value of 𝑓𝑐 is calculated as
𝑓𝑐=
1
𝑟2 [−𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒] (2) (3) 𝑓𝑐=
1
𝑟2 [−𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒] (4) Fig. 5: Forces acting on mechanism. Fig. 5: Forces acting on mechanism. Substituting value of 𝑓𝑐 in Eq. (2) Substituting value of 𝑓𝑐 in Eq. (2) Substituting value of 𝑓𝑐 in Eq. (2)
𝑇𝑔= 𝑘1𝜃1 + 𝑏1(𝜃1̇ −𝜃2̇ ) −
𝑟1
𝑟2 [−𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒]
(5) 𝑇𝑔= 𝑘1𝜃1 + 𝑏1(𝜃1̇ −𝜃2̇ ) −
𝑟1
𝑟2 [−𝐽2𝜃2̈ −𝑏1(𝜃2̇ −𝜃1̇ ) −𝑇𝑒]
(5) (5) Using the inverse gear ratio,
𝑟2
𝑟1 = N and the equality of arc lengths, 𝑟1𝜃1 = 𝑟2𝜃2, it follows
that 𝜃1 =
𝑟2
𝑟1 𝜃2 = 𝑁𝜃2. Substituting in Eq. (5), Using the inverse gear ratio,
𝑟2
𝑟1 = N and the equality of arc lengths, 𝑟1𝜃1 = 𝑟2𝜃2, it follows
that 𝜃1 =
𝑟2
𝑟1 𝜃2 = 𝑁𝜃2. Substituting in Eq. 3.2 Actuator Model The Finger Extensor is powered by a planetary-geared, 24 V brushed DC motor. A
schematic of the motor is shown in Fig. 6. Using Kirchhoff's voltage law and Newton's
law of motion, dynamic equations of the actuator were written as 𝐽𝑚𝜃𝑚̈ (𝑡) = −𝐵𝑣𝜃𝑚̇ (𝑡) + 𝑘𝑡𝑖𝑎(𝑡) −𝑇𝐿(𝑡)
(8)
𝐿𝑖𝑎(𝑡) = −𝑘𝑣𝜃𝑚̇ (𝑡) −𝑅𝑖𝑎(𝑡) + 𝑉(𝑡)
(9) (8) (9) 𝐿𝑖𝑎(𝑡) = −𝑘𝑣𝜃𝑚(𝑡) −𝑅𝑖𝑎(𝑡) + 𝑉(𝑡)
(9)
where, 𝐽𝑚 is the moment of inertia of the motor (kg. m2) 𝐽𝑚 is the moment of inertia of the motor (kg. m2) 389 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. 𝜃𝑚(𝑡) is the angular displacement of the motor (rad)
𝐵𝑣 is the viscous friction coefficient (Nm/rad/s)
𝑘𝑡, 𝑘𝑣 are the torque and back emf constants (Nm/A, V/rad/s)
L, R are the armature inductance and resistance (H, Ω)
𝑖𝑎(𝑡) is the armature current for the motor (A)
𝑇𝐿(𝑡) is the load torque for the motor (Nm)
𝑉(𝑡) is the voltage input (V)
Fig. 6: Motor schematic. Fig. 6: Motor schematic. Fig. 6: Motor schematic. Utilizing the relationship between loading torque 𝑇(𝑡) (from the motor) and load
torque 𝑇𝐿(𝑡) (on the motor shaft) and the relationship between the motor's angular position
and the machine's angular position, we formulated 𝑇𝐿(𝑡) =
𝑇(𝑡)
𝑁, 𝑁≥1
𝜃𝑚(𝑡) = 𝑁𝜃(𝑡) 𝑇𝐿(𝑡) =
𝑇(𝑡)
𝑁, 𝑁≥1 𝑇𝐿(𝑡) =
𝑇(𝑡)
𝑁, 𝑁≥1
(10)
𝜃𝑚(𝑡) = 𝑁𝜃(𝑡)
(11) 𝑇𝐿(𝑡) =
𝑇(𝑡)
𝑁, 𝑁≥1
(10)
𝜃𝑚(𝑡) = 𝑁𝜃(𝑡)
(11) (10) (11) where N is the inverse gear ratio. 3.3 Integrated Model For developing the integrated model of the machine, a time derivative of the dynamic
equation of the machine was found. Differentiating Eq. (7) gave 𝑇̇𝑔(𝑡) = {
𝐽2(𝜃2)
𝑁} 𝜃2⃛(𝑡) + {
𝑇̇𝑒(𝑡)
𝑁} + 𝑁𝑘𝜃2̇ (𝑡) + 𝐹𝑐{2𝛿(𝜃2̇ )𝜃2̈ } + 𝑉𝑐𝜃2̈ (𝑡)
(15)
Substituting Eq. (15) in Eq. (12), (15) 𝜃2⃛(𝑡) {𝐽𝑚𝑁3 + 𝐽2(𝜃2)
𝐽𝑚𝑁3
} + 𝜃2̈ (𝑡) {𝑁2(𝐵𝑣𝐿+ 𝐽𝑚𝑅) + 2𝐹𝑐𝐿𝛿(𝜃2̇ ) + 𝑉𝑐𝐿
𝐽𝑚𝑁2𝐿
}
+ 𝜃2̇ (𝑡) {𝑁(𝑘𝑣𝑘𝑡+ 𝐵𝑣𝑅) + 𝐿𝑘1
𝐽𝑚𝑁𝐿
} + 𝑇𝑔(𝑡) {
𝑅
𝑁2𝐽𝑚𝐿} + 𝑇̇𝑒(𝑡) {
1
𝑁3𝐽𝑚
}
= { 𝑘𝑡
𝑁𝐽𝑚𝐿} 𝑉(𝑡)
(16) (16) Equation (16) was solved for 𝜃 and put in state-space form, the integrated model of the
Finger Extensor was obtained as Equation (16) was solved for 𝜃 and put in state-space form, the integrated model of the
Finger Extensor was obtained as 𝑋̇𝐵(𝑡) = 𝐴𝐵(𝑋𝐵, 𝑡)𝑋𝐵(𝑡) + 𝐵𝐵(𝑋𝐵, 𝑡)𝑈(𝑡) + 𝐹𝐵𝑇𝑔(𝑡) + 𝑄𝐵𝑇̇𝑒(𝑡)
(17) (17) where 𝑋𝐵= [𝜃2 𝜃2̇ 𝜃2̈ ]
𝑇 where 𝑋𝐵= [𝜃2 𝜃2̇ 𝜃2̈ ]
𝑇 where 𝑋𝐵= [𝜃2 𝜃2̇ 𝜃2̈ ]
𝑇 𝐴𝐵, 𝐵𝐵, 𝐹𝐵, 𝑄𝐵 are the system, input, load distribution and rate of load distribution
respectively. 𝑈(𝑡), 𝑇𝑔(𝑡), 𝑇𝑒(𝑡) are the input vector, mechanical link torque and toque
exerted by the patient. 𝐴𝐵(𝑋𝐵, 𝑡) = [
0
0
1
0
1
0
0
𝑎32
𝑎33
] , 𝐵𝐵(𝑋𝐵, 𝑡) = [
0
0
𝑏31
] , 𝐹𝐵(𝑋𝐵, 𝑡) = [
0
0
𝑓31
] , 𝑄𝐵(𝑋𝐵, 𝑡) = [
0
0
𝑞31
] Equation (17) was put in the state-space form and solved for 𝑋̇𝐵. Equation (17) was put in the state-space form and solved for 𝑋̇𝐵. 𝑎32 = −{𝑁2(𝑁𝑘𝑣𝑘𝑡+ 𝑁𝐵𝑣𝑅+ 𝐿𝑘1)
𝐿(𝐽𝑚𝑁3 + 𝐽2(𝜃2))
} , 𝑎33 = −{𝑁(𝑁2𝐵𝑣𝐿+ 𝑁2𝐽𝑚𝑅+ 2𝐹𝑐𝐿𝛿(𝜃2
˙
) + 𝑉𝑐𝐿
𝐿(𝐽𝑚𝑁3 + 𝐽2(𝜃2))
}
𝑏31 = {
𝑘𝑡𝑁2
𝐿(𝐽𝑚𝑁3 + 𝐽2(𝜃2))} , 𝑓31 = −{
𝑅𝑁
𝐿(𝐽𝑚𝑁3 + 𝐽2(𝜃2))} , 𝑞31 = −{
1
(𝐽𝑚𝑁3 + 𝐽2(𝜃2))} The state-space model derived in Eq. (17) represents the integrated dynamics of the Finger
Extensor and was utilized in the simulation of the developed system. The state-space model derived in Eq. (17) represents the integrated dynamics of the Finger
Extensor and was utilized in the simulation of the developed system. where N is the inverse gear ratio. where N is the inverse gear ratio. Substituting Eq. (10) and Eq. (11) in Eq. (8) and Eq. (9), the otherwise linear, time-
invariant actuator dynamic was converted into a non-linear time-variant one. The actuator
model in the form of a third-order differential equation was represented as: 𝜃2⃛(𝑡) + {
𝐵𝑣𝐿+𝐽𝑚𝑅
𝐽𝑚𝐿
} 𝜃2̈ (𝑡) + {
𝑘𝑣𝑘𝑡+𝐵𝑣𝑅
𝐽𝑚𝐿
} 𝜃2̇ (𝑡) + {
𝑅
𝑁2𝐽𝑚𝐿} 𝑇𝑔(𝑡) + {
1
𝑁2𝐽𝑚} 𝑇̇𝑔= {
𝑘𝑡
𝑁𝐽𝑚𝐿} 𝑉(𝑡) (12) To represent the actuator dynamic model in the state variable form, 𝜃, 𝜃̇, 𝜃̈ were
chosen as state variables and the state vector for the actuator was given by To represent the actuator dynamic model in the state variable form, 𝜃, 𝜃̇, 𝜃̈ were
chosen as state variables and the state vector for the actuator was given by 𝑋𝐵(𝑡) = [𝜃(𝑡) 𝜃̇(𝑡) 𝜃̈(𝑡)]
𝑇
(13) 𝑋𝐵(𝑡) = [𝜃(𝑡) 𝜃̇(𝑡) 𝜃̈(𝑡)]
𝑇
(13 (13) The state equation was formulated as The state equation was formulated as 𝑋𝐵̇ (𝑡) = 𝐴𝐵𝑋𝐵(𝑡) + 𝐵𝐵𝑈(𝑡) + 𝐹𝐵𝑇𝑔(𝑡) + 𝑄𝐵𝑇̇𝑒(𝑡)
(14) 𝑋𝐵̇ (𝑡) = 𝐴𝐵𝑋𝐵(𝑡) + 𝐵𝐵𝑈(𝑡) + 𝐹𝐵𝑇𝑔(𝑡) + 𝑄𝐵𝑇̇𝑒(𝑡)
(14)
where (14) 𝐴𝐵(𝑋𝐵, 𝑡) = [
0
0
1
0
1
0
0
𝑎32
𝑎33
] , 𝐵𝐵(𝑋𝐵, 𝑡) = [
0
0
𝑏31
] , 𝐹𝐵(𝑋𝐵, 𝑡) = [
0
0
𝑓31
] , 𝑄𝐵(𝑋𝐵, 𝑡) = [
0
0
𝑞31
] 390 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. Here, 𝐴𝐵, 𝐵𝐵, 𝐹𝐵, 𝑄𝐵 are the system, input, load distribution and rate of load distribution
respectively. 𝑈(𝑡), 𝑇𝑔(𝑡), 𝑇𝑒(𝑡) are the input vector, mechanical link torque and toque
exerted by the patient. The values of elements 𝑎32, 𝑎33, 𝑏31, 𝑓31, and 𝑞31 are 𝑎32 = {−𝑘𝑣𝑘𝑡+ 𝐵𝑣𝑅
𝐽𝑚𝐿
}; 𝑎33 = {−𝐵𝑣𝐿+ 𝐽𝑚𝑅
𝐽𝑚𝐿
}; 𝑏31 = { 𝑘𝑡
𝑁𝐽𝑚𝐿} ; 𝑓31 = −{
𝑅
𝑁2𝐽𝑚𝐿} ;
𝑞31 = −{
1
𝑁2𝐽𝑚
} 4. MODEL VALIDATION Using the MBD approach for designing a controller for the Finger Extensor, dynamic
system modelling was followed by graphical representation of the mathematical
formulation in the MATLAB-Simulink environment. Simulation results were validated
through hardware-in-the-loop experimentation. The same input was provided to both the
developed model as well as the machine and comparisons were drawn between resultant
outputs. 3.4 Mechanical and Electrical Parameter Determination The development of the integrated model for the Finger Extensor was followed by
determination of mechanical and electrical parameters involved. This step aids in
enhancing the fidelity of the developed model. Mechanical parameters such as radius of 391 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. the iris and moment of inertia of the inner mechanism were adapted from [20]. Electrical
parameters associated with the actuator model were experimentally found from the motor
characteristic curve as shown in Fig. 7. Torque and back EMF constants 𝑘𝑡, 𝑘𝑣 were
found from the torque-current and torque-speed curves with similar values of 0.791 and
0.886, respectively. Other parameters such as the viscous friction constant 𝐵𝑣, armature
resistance R and inductance L were adapted from [20]. the iris and moment of inertia of the inner mechanism were adapted from [20]. Electrical
parameters associated with the actuator model were experimentally found from the motor
characteristic curve as shown in Fig. 7. Torque and back EMF constants 𝑘𝑡, 𝑘𝑣 were
found from the torque-current and torque-speed curves with similar values of 0.791 and
0.886, respectively. Other parameters such as the viscous friction constant 𝐵𝑣, armature
resistance R and inductance L were adapted from [20]. Fig. 7: Motor characteristic curves. Fig. 7: Motor characteristic curves. 4.1 Simulation For carrying out simulation of the mathematical model of the Finger Extensor, the
voltage and torques from the motor and user acted as inputs to the Finger Extensor plant
while the angle of rotation of the iris was the output. A PID controller was used for
optimal position control. Rotational position output and its derivatives were used to
calculate velocity and acceleration. These values helped in calculating the force acting on
the patient. A desired angle of 𝜃 = 121° (2.11 radians) was fed into the plant and the
simulated position of the iris was plotted as shown in Fig. 8. Fig. 8: Comparison between target and simulated angular position of the iris. Fig. 8: Comparison between target and simulated angular position of the iris. 392 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. The figure shows a complete open and close cycle of the Finger Extensor for 30
seconds. The poles of the Extensor open to their maximum rotation angle of 121° followed
by closing motion to 7.89°. From the figure, it is clear that the hardware had a slower
response than the simulation and took about 11.20 seconds to sync with the simulation
results. However, beyond that time, error was negligible and actual behavior of the Finger
Extensor overlapped with the simulated, modeled behavior. From experimental data, it
was observed that the mean absolute error was 1.38° and the relative error was 1.13%. However, for rehabilitation applications, movement accuracy is less critical as compared
to surgical applications. Also, position resolution of human arm movement is 5 mm or 5º
[21], which indicates that a mean absolute error of 1.38° is likely to be acceptable for
rehabilitation purposes. Figure 11 shows the comparison between the simulated result and the hardware-in-
the-loop response, when the Finger Extensor was programmed to open to an angle of
59.58° (1.04 radians). Simulation and experimental curves took 0.3 seconds to reach the
desired angle. The Root Mean Squared Error (RMSE) was calculated as 0.9875. There
was a slight overshoot of 2.86º (0.05 radians) which caused the poles of the Finger
Extensor to open to an angle 2.86º greater than the target. However, this negligible
overshoot did not inflict any discomfort on the subject. Also, the response settled down to
its desired value within 0.08 seconds. Fig. 11: Simulated and experimental responses for target angle experiment. Fig. 11: Simulated and experimental responses for target angle experiment. 4.2 Hardware-in-the-Loop Experiments To validate the developed dynamic model and demonstrate its effectiveness,
experiments were conducted making the Finger Extensor a part of the simulation loop. These experiments were conducted with a healthy subject, a female of 29 years of age. A
simple open-close movement of the machine was performed. Results from the simulations
run in MATLAB were compared with the experimental tests with the Finger Extensor. The experimental set-up as shown in Fig. 9 consisted of the Finger Extensor connected to
an Intel Core i5 PC by means of an Arduino Uno microprocessor board. This board
received sensory data from the encoder as well as the torque sensor. Communication
between the Arduino and MATLAB 2018b software, running on the PC was established
using a MATLAB s-function. This function read data from the encoder i.e. the position
sensor of the machine and converted the encoder counts to angular displacement of the
iris. Fig. 9: Experimental setup for hardware-in-the-loop simulation. Fig. 9: Experimental setup for hardware-in-the-loop simulation. The movement commands to the motor and consequently to the encoder were
provided using Simulink's support package for Arduino which allowed Simulink blocks to
control hardware connected to the Arduino using PWM signals. In this case, hardware
controlled was the motor driver connected to the motor. The desired input angle was fed
into the hardware and the Finger Extensor was run for about 30 seconds, performing open-
close movements. The comparison between the simulated and the experimental angular
displacement of the machine is shown in Fig. 10. Fig. 10: Simulated and experimental responses during open-close movements. Fig. 10: Simulated and experimental responses during open-close movements. 393 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 5. CONCLUSION This paper presented a mathematical model for the 1-DOF Finger Extensor
rehabilitation machine, taking into consideration the dynamics of the mechanism as well
as that of the actuation system. Simulation results show that the developed model tracks
the desired position efficiently with minimal error. Hardware-in-the-Loop experiments
carried out with the machine demonstrate the validity of the model. System response using
the developed model was compared with that obtained through hardware experimentation. The results illustrate the accuracy of the model to the actual behavior of the rehabilitation
system and make it suitable for use in the design of a control scheme for it. Obtained
results with mean absolute error of only 1.38° are within the human motion resolution
limits and exhibit suitability of the model for application in robotic rehabilitation systems. Future research with the Finger Extensor and the developed model will be directed
towards development of a control scheme for the machine. This will be aimed at
modulating assistance/resistance provided to the patient based on his/her performance
using various metrics such as limb velocity, applied force, position error etc. Considering
patient's strength and residual ability, primary goal of the controller would be to increase 394 IIUM Engineering Journal, Vol. 22, No. 2, 2021
https://doi.org/10.31436/iiumej.v22i2.1706 Shahdad et al. https://doi.org/10.31436/iiumej.v22i2.1706 self-initiated movements and patient engagement in training exercises which would lead to
an increase in neuroplasticity. As part of future study, clinical testing of the Finger
Extensor will also be carried out with stroke patients. This will include a detailed study of
the level of comfort of the device for use with such patients. self-initiated movements and patient engagement in training exercises which would lead to
an increase in neuroplasticity. As part of future study, clinical testing of the Finger
Extensor will also be carried out with stroke patients. This will include a detailed study of
the level of comfort of the device for use with such patients. REFERENCES [1]
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Efectos de la realización de un megaevento deportivo sobre una economía local. El caso de los Juegos Mundiales 2013 Cali
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Article
Efectos De La Realización De Un Megaevento Deportivo
Sobre Una Economía Local. El Caso De Los Juegos
Mundiales 2013 Cali Journal of Economics, Finance and Administrative Science Journal of Economics, Finance and Administrative Science Provided in Cooperation with:
Universidad ESAN, Lima Suggested Citation: Aguado, Luis F.; Osorio Mejía, Ana M.; Arbona, Alexei; Peña-Vinces, Jesús C. (2017) : Efectos De La Realización De Un Megaevento Deportivo Sobre Una Economía Local. El Caso
De Los Juegos Mundiales 2013 Cali, Journal of Economics, Finance and Administrative Science, ISSN
2218-0648, Emerald Publishing Limited, Bingley, Vol. 22, Iss. 43, pp. 131-153,
https://doi.org/10.1108/JEFAS-01-2017-0012 Aguado, Luis F.; Osorio Mejía, Ana M.; Arbona, Alexei; Peña-Vinces, Jesús C. Terms of use: Die Dokumente auf EconStor dürfen zu eigenen wissenschaftlichen
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www.emeraldinsight.com/2077-1886.htm © Luis F. Aguado, Ana M. Osorio, Alexei Arbona and Jesús C. Peña-Vinces. Published in
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Limited. This article is published under the Creative Commons Attribution (CC BY 4.0)
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article (for both commercial & non-commercial purposes), subject to full attribution to the
original publication and authors. The full terms of this licence may be seen at http://
creativecommons.org/licenses/by/4.0/legalcode Resumen Propósito – En el artículo se presenta una síntesis del impacto sobre la producción y el empleo generado por
la realización de un megaevento deportivo en la ciudad de Cali, Colombia. Diseño/metodología/enfoque – El impacto se estima a través de los efectos sistemáticos producidos por
las actividades de inversión/gasto de los agentes identificados y los consecuentes efectos multiplicadores
capturados a partir de un modelo insumo-producto. Hallazgos – La realización de los Juegos Mundiales significó un nuevo flujo de ingreso monetario a la
ciudad de Cali, de $101.036 millones de pesos (=53,4 millones de dólares) y se generaron 9.598 puestos de
trabajo (=7.711 empleos de tiempo completo). Adicionalmente, se atrajo a la ciudad a 2.174 turistas
internacionales y 11.250 turistas nacionales. Originalidad/valor – El artículo aporta aspectos innovadores metodológicamente útiles para futuro
estudios de impacto económico en ciudades poco turísticas situadas en países en vías desarrollo. Palabras clave – Cali, Colombia, Impacto económico, Los Juegos Mundiales, Megaeventos deportivos Tipo de artículo – Artículo de investigación Efectos de la realización de un
megaevento deportivo sobre una
economía local. El caso de los
Juegos Mundiales 2013 Cali
Luis F. Aguado, Ana M. Osorio and Alexei Arbona
Departamento de Economía, Pontificia Universidad Javeriana, Cali, Colombia, y Efectos de la
realización de
un megaevento
deportivo 131 Received 25 January 2017
Accepted 8 February 2017 Jesús C. Peña-Vinces
Business Management, Universidad de Sevilla, Sevilla, España JEL classification – L83, C67 Journal of Economics, Finance and
Administrative Science
Vol. 22 No. 43, 2017
pp. 131-153
EmeraldPublishingLimited
2077-1886
DOI 10.1108/JEFAS-01-2017-0012 1. Introducción En el artículo se presentan los resultados del impacto económico a corto plazo, sobre la
ciudad de Santiago de Cali[1] y su área metropolitana (en adelante AM), asociado a la
celebración de la IX edición de los Juegos Mundiales (World Games). Los Juegos Mundiales
están constituidos por las federaciones de disciplinas deportivas que no participan en el
programa de los Juegos Olímpicos. La IX edición de los Juegos Mundiales se llevó a cabo
entre el 25 de julio y el 4 de agosto de 2013 y ha sido la primera organizada por una ciudad de
habla hispana y situada en un país de América del Sur[2]. Los Juegos Mundiales, como
megaevento deportivo, involucraron la competición simultánea en 31 disciplinas
deportivas[3], el desplazamiento de 2.929 deportistas y de 1.903 delegados de 97 países de los
cinco continentes. Adicionalmente, la novedad y la diversidad que implica la presencia de
atletas de élite en deportes poco conocidos y practicados como el sumo, el wushu y el
korfball, por mencionar algunos en el contexto de América Latina. El artículo se centra en los impactos producidos por las actividades de inversión/gasto de
los agentes identificados en la celebración de los Juegos Mundiales: Comité Local
Organizador de Cali (inversión en infraestructura, gasto en logística) y espectadores (gasto
en consumo), y los consecuentes efectos multiplicadores capturados a partir de un modelo
insumo-producto. Estos impactos se sintetizan en los efectos sobre la producción y el empleo
en el área metropolitana de la ciudad, definida como área de impacto. p
,
p
La inversión para garantizar la infraestructura física deportiva y la organización y
logística de los Juegos Mundiales más el gasto de los espectadores ascendió a $140.357
millones de pesos (:74,2 mill. US$)[4]. De este total, $101.036 millones de pesos (:53,4
mill. US$) significaron un nuevo flujo de ingreso monetario derivado de agentes de
fuera de Cali y A.M. (Gobierno Nacional; Turistas Extranjeros y Nacionales) que de no
haberse realizado Los Juegos Mundiales no hubiese llegado a la ciudad. Adicionalmente,
los Juegos Mundiales atrajeron a 2,174 turistas internacionales y 11,250 turistas
nacionales. Asimismo, generaron 9.598 puestos de trabajo (:7.711 empleos de tiempo
completo) en sectores vinculados con la logística de la organización de los Juegos y con
la “turbina de gasto” de los turistas. g
Los resultados lucen interesantes para la literatura empírica sobre los estudios de
impacto económico de megaeventos deportivos. Abstract Purpose – A summary of the economic impact resulting from the celebration of a sporting mega-event in
the city of Cali, Colombia, is carried out in this article. Design/methodology/approach – The impacts are estimated by means of the systematic effects
produced by the activities of investment/expenditure of the identified agents and the consequent multiplying
effects captured from an input-output model. Findings – The World Games represented a new income flow to the city of Cali of $101.036 million pesos (:
US$53.4 mill.) and 9.598 jobs were created (:7,711 full-time jobs). Additionally, 2,174 foreign tourists and
11,250 national tourists were attracted to the city. Journal of Economics, Finance and
Administrative Science
Vol. 22 No. 43, 2017
pp. 131-153
EmeraldPublishingLimited
2077-1886
DOI 10.1108/JEFAS-01-2017-0012 Originality/value – The article contributes with innovative aspects methodologically speaking, useful for
future studies of economic impact in cities with low tourism located in developing countries. Keywords Economic impact, Sporting mega-events, World Games, Cali
Paper type Research paper Originality/value – The article contributes with innovative aspects methodologically speaking, useful for
future studies of economic impact in cities with low tourism located in developing countries. Keywords Economic impact, Sporting mega-events, World Games, Cali Originality/value – The article contributes with innovative aspects methodologically speaking, useful for
uture studies of economic impact in cities with low tourism located in developing countries. Keywords Economic impact, Sporting mega-events, World Games, Cali Keywords Economic impact, Sporting mega-events, World Games, Cali
Paper type Research paper Keywords Economic impact, Sporting mega-events, World Games, Cali
Paper type Research paper 132 Efectos de la
realización de
un megaevento
deportivo 2. Los Juegos Mundiales como megaevento deportivo
D
d
l li
d l
í d l d
(R
h De acuerdo con la literatura de la economía del deporte (Roche, 1994; Llopis-Goig, 2012), los
megaeventos deportivos se caracterizan por ser eventos a gran escala con un profundo
significado internacional; tener una duración breve y perfectamente acotada; estar
organizados por un país o ciudad que generalmente compite por ser sede; atraer un
importante número de participantes (deportistas) y espectadores, tanto locales como
extranjeros; ser cubiertos por un amplio número de medios internacionales de comunicación;
requerir alta inversión en infraestructura, logística y seguridad; y ser regentados por un
Comité Local Organizador integrado por autoridades de gobiernos nacionales y locales y
organizaciones y federaciones deportivas internacionales. Los Juegos Mundiales encajan dentro de la tipología de megaevento deportivo (ver Tabla I). En efecto, estos tienen carácter multideportivo, un gran número de deportistas y países
participantes, se celebran cada cuatro años, son regentados por la Asociación
Internacional de los Juegos Mundiales (IWGA)[5]. La literatura de la economía del
deporte identifica como megaeventos deportivos a los Juegos Olímpicos, la Copa
Mundial de Fútbol (FIFA) y los Juegos Olímpicos de Verano e Invierno y en menor escala
los Juegos Panamericanos. Los Juegos Mundiales constituyen un megaevento
impulsado por los resultados deportivos —“competitor driven”— que alcancen los
deportistas de élite en competencia, frente a aquellos megaeventos impulsados por su
alto número de seguidores y espectadores —“spectator driven”— (Gratton et al., 2006,
p. 51). 1. Introducción Por un lado, en el contexto latinoamericano
es muy limitada la información de los estudios de esta naturaleza, exceptuando la
investigación ex-post para los Juegos Panamericanos de Río de Janeiro 2007, el estudio ex-
ante de los Juegos Panamericanos de Guadalajara 2011 y el más reciente para el pasado
mundial de fútbol de Brasil. Por otro lado, se presentan elementos innovadores en la
metodología seguida. Así, la metodología para determinar los turistas internacionales por
vía administrativa ofrece un cálculo directo y, por tanto, realista del flujo de visitantes
extranjeros, usualmente sobreestimados en los estudios de impacto económico. En el caso de
la estimación de los turistas nacionales, se combina información de una encuesta cara a cara
aplicada a los asistentes con información de las entradas vendidas por el Comité
Organizador Local. En ambos casos (turistas nacionales e internacionales), se tiene en cuenta
el tamaño de los escenarios, la frecuencia de asistencia y la rotación de entradas, lo que hace
más precisas las estimaciones. Adicionalmente, se plantea el limitado alcance de los efectos “expulsión” y de
“cancelación de reservas” en ciudades poco turísticas situadas en países en vías desarrollo,
lo que incrementa para este tipo de ciudades los beneficios económicos de realizar este tipo
de eventos internacionales. El artículo se organiza en siete secciones, la primera es esta
introducción. En la segunda y tercera secciones, se caracterizan los Juegos Mundiales como
megaevento deportivo y se detalla la metodología empleada en la estimación del impacto
económico. En la cuarta sección se presenta la estimación del flujo de espectadores, tanto
nacionales como extranjeros. En quinto lugar se estima el impacto económico de los Juegos
Mundiales a través de un modelo insumo-producto. En la sexta sección se discute el legado
de los Juegos Mundiales para Cali, es decir los efectos de largo plazo esperados por la
celebración del megaevento deportivo, los cuales no necesariamente se capturan por las
relaciones monetarias implícitas en los modelos insumo-producto. Por último, se presentan
las conclusiones. 133 3. Metodología
l
l En el artículo se utiliza la metodología de efectos, propia de los Estudios de Impacto
Económico —en adelante EIE—. La literatura empírica de estudios de impacto económico
aplicada al sector deportivo está concentrada en países desarrollados situados en Europa,
Estados Unidos y Asia. Una aproximación detallada de la metodología empleada, sus
limitaciones y aplicaciones se encuentra en: (Baade and Matheson, 2016; Zimbalist, 2015;
Barajas et al., 2012; Matheson, 2012; Porter and Chin, 2012; Barget and Gouguet, 2008;
Gratton et al., 2006; Matheson, 2006). Para el contexto de América Latina (Devesa et al., 2012;
Ávalos, 2011). El análisis del impacto económico resulta útil para ofrecer una imagen cuantitativa de la
actividad económica adicional generada en la ciudad por la celebración de los Juegos
Mundiales. Por un lado, referida a la focalización de las inversiones —públicas y privadas—
en infraestructura deportiva y urbana y el gasto en gestión y logística para llevar a cabo el Tabla I. Perfil de los Juegos
Mundiales 2013 -
Cali
Características
Descripción
Gobierno del evento
International World Games Association (IWGA)
Comité Local Organizador de Cali (CLOC)
Comité Nacional Organizador
Ámbito geográfico
Internacional
97 países: 40 (Europa) þ 23 (América) þ 23 (Asia) þ 2 (Oceanía) þ 9 (África)
Frecuencia de
realización
Cada 4 años con cambio de sede
Duración
11 días (10 de competiciones þ 1 de inauguración)
Competidores
2.929 deportistas: 1.511 (Europa) þ 823 (América) þ 407 (Asia) þ 110 (Oceanía) þ
78 (África)
Delegaciones
1.903 delegados de las federaciones deportivas, jueces internacionales, delegaciones
nacionales —entrenadores, médicos, ayudantes—, incluidos los representantes de
los medios internacionales
Deportes
31 (26 oficiales þ 5 invitados)
Interés mediático
Cali compitió por ser sede con las ciudades de Johannesburgo (Sudáfrica) y San
Petersburgo (Rusia)
96 periodistas de 34 países
Escenarios
25 (20 cerrados þ 5 al aire libre)
Voluntariado
2.539 voluntarios
Novedad en América
Latina
Primera edición organizada por una ciudad de habla hispana y situada en un país de
América del Sur
Nota: Elaboración propia con información del Comité Organizador Local Cali
JEFAS
22,43
134 Nota: Elaboración propia con información del Comité Organizador Local Cali megaevento deportivo. De otro lado, referida a la generación de gasto de los asistentes —
turistas nacionales y extranjeros— en bienes y servicios relacionados con su estadía en la
ciudad. 3. Metodología
l
l Desde esta perspectiva, el gasto de los residentes de Cali y AM no
constituye una entrada neta de dinero a la economía local, sino una reasignación de su gasto. Este supuesto señala que los residentes en Cali y AM disponen de un presupuesto fijo para
sus gastos de recreación y ocio. Por tanto, incluir este gasto puede conducir a sobrestimar los
efectos indirectos, error muy común en los estudios de impacto económico (Siegfried and
Zimbalist, 2006). De igual forma, se debe tener presente el posible efecto expulsión provocado por el
desplazamiento de público local residente, y/o la cancelación de visitas a la ciudad de
turistas (extranjeros/nacionales), para evitar la congestión y los altos precios que supone el
exceso de demanda provocado por los espectadores atraídos por la celebración del
megaevento deportivo. En el primer caso, la celebración de los Juegos Mundiales representó
Figura 1. Metodología impacto
económico Juegos
Mundiales 2013 Cali
IMPACTOS
Fuentes de demanda
Corto plazo
Comité Local Organizador
Inversión en infraestructura deportiva
Construcción y remodelación de escenarios
deportivos. Inversión en gestión y logística
Preparación y gestión por parte del equipo que
integra el CLOC. Equipamiento de las
instalaciones deportivas; transporte y
alimentación de deportistas, jueces y
delegados, uniformes de los voluntarios, etc. Gasto visitantes/espectadores
Gasto en bienes y servicios por parte de los
visitantes/espectadores no locales
Encuesta cara/cara
asistentes
Efectos directos
Efectos indirectos
Producción y empleo
adicionales generados por
los Juegos Mundiales 2013
Cali
Comité Local
Organizador de Cali,
CLOC
[Presupuesto/Gastos]
Fuentes de información
Efectos inducidos : Matriz insumo producto adaptada par
el Valle del Cauca
FLUJO MONETARIO DE GASTO
Elaboración propia
Efectos de la
realización de
un megaevento
deportivo
135 Figura 1. Metodología impacto
económico Juegos
Mundiales 2013 Cali
IMPACTOS
Fuentes de demanda
Corto plazo
Comité Local Organizador
Inversión en infraestructura deportiva
Construcción y remodelación de escenarios
deportivos. Inversión en gestión y logística
Preparación y gestión por parte del equipo que
integra el CLOC. Equipamiento de las
instalaciones deportivas; transporte y
alimentación de deportistas, jueces y
delegados, uniformes de los voluntarios, etc. 3. Metodología
l
l Ambas fuentes de nueva demanda generan un efecto multiplicador sobre la
economía de la ciudad a través de la interrelación sectorial capturada a partir de un modelo
insumo-producto (Schuschny, 2005). La metodología se centra en los efectos sistemáticos
producidos por las actividades de inversión/gasto de los agentes identificados (ver Figura 1). megaevento deportivo. De otro lado, referida a la generación de gasto de los asistentes —
turistas nacionales y extranjeros— en bienes y servicios relacionados con su estadía en la
ciudad. Ambas fuentes de nueva demanda generan un efecto multiplicador sobre la
economía de la ciudad a través de la interrelación sectorial capturada a partir de un modelo
insumo-producto (Schuschny, 2005). La metodología se centra en los efectos sistemáticos
producidos por las actividades de inversión/gasto de los agentes identificados (ver Figura 1). Es así como la estimación del impacto económico de los Juegos Mundiales 2013 Cali se
lleva a cabo a partir de la suma de tres efectos: directo, indirecto e inducido. 3.1 Efectos directos
Los efectos directos miden los gastos/inversión totales en que incurre el Comité Local
Organizador de Cali, CLOC, nómina de sueldos, bienes y servicios comprados, construcción
y adecuación de infraestructura física deportiva, logística, alquileres, etc. Son obtenidos a
partir de la información suministrada por el Comité Local Organizador discriminada por la
fuente de financiación, gobierno local o entidades del gobierno regional/nacional. Estos
últimos constituyen la inyección de gasto nuevo en el área metropolitana de Santiago de Cali
y no una recomposición del gasto del municipio. Asimismo, un aspecto a considerar en el
análisis del impacto económico de los megaeventos deportivos es el tipo de financiación
usado (Zimbalist, 2010). La financiación vía deuda pública deja compromisos en el mediano
y largo plazo asumibles mediante mayores impuestos y/o recortes del gasto en otros
servicios prestados por el gobierno (local/nacional). Si bien este estudio se enfoca en el corto
plazo, es importante señalar que la financiación de los Juegos Mundiales provino en mayor
medida de recursos del gobierno nacional y no requirió contraer deuda pública por parte del
gobierno local[6]. 3.2 Efectos indirectos
Los efectos indirectos miden los gastos que realizan los espectadores en bienes y servicios —
compra de entradas, alojamiento, alimentación, transporte local, recuerdos—. Los gastos
relevantes son los realizados por los espectadores de fuera del área metropolitana de
Santiago de Cali. 3. Metodología
l
l Gasto visitantes/espectadores
Gasto en bienes y servicios por parte de los
visitantes/espectadores no locales
Encuesta cara/cara
asistentes
Efectos directos
Efectos indirectos
Producción y empleo
adicionales generados por
los Juegos Mundiales 2013
Cali
Comité Local
Organizador de Cali,
CLOC
[Presupuesto/Gastos]
Fuentes de información
Efectos inducidos : Matriz insumo producto adaptada par
el Valle del Cauca
FLUJO MONETARIO DE GASTO
Elaboración propia
Efectos de la
realización de
un megaevento
deportivo
135 3.2 Efectos indirectos 3.2 Efectos indirectos f
Los efectos indirectos miden los gastos que realizan los espectadores en bienes y servicios —
compra de entradas, alojamiento, alimentación, transporte local, recuerdos—. Los gastos
relevantes son los realizados por los espectadores de fuera del área metropolitana de
Santiago de Cali. Desde esta perspectiva, el gasto de los residentes de Cali y AM no
constituye una entrada neta de dinero a la economía local, sino una reasignación de su gasto. Este supuesto señala que los residentes en Cali y AM disponen de un presupuesto fijo para
sus gastos de recreación y ocio. Por tanto, incluir este gasto puede conducir a sobrestimar los
efectos indirectos, error muy común en los estudios de impacto económico (Siegfried and
Zimbalist, 2006). ,
)
De igual forma, se debe tener presente el posible efecto expulsión provocado por el
desplazamiento de público local residente, y/o la cancelación de visitas a la ciudad de
turistas (extranjeros/nacionales), para evitar la congestión y los altos precios que supone el
exceso de demanda provocado por los espectadores atraídos por la celebración del
megaevento deportivo. En el primer caso, la celebración de los Juegos Mundiales representó
un evento extraordinario que motivó a los locales a participar como asistentes a las
competiciones y a demostrar su hospitalidad a los visitantes internacionales. El segundo
caso parece ajustarse más a la realidad de grandes ciudades turísticas en países
desarrollados. En la Figura 2 se observa que la demanda adicional de hospedaje provocada
por la celebración de los Juegos Mundiales fue perfectamente absorbida por la capacidad
hotelera disponible en la ciudad, para los mismos meses de 2014 la tasa de ocupación estuvo
alrededor del 50 per cent, por tanto no sería válido concluir que se presentó un efecto
desplazamiento y de “cancelación de reservas”. Para la estimación del número de espectadores y la descripción de su perfil
sociodemográfico y de gasto se requirió el uso de varias fuentes de información, entre ellas la
colaboración de Migración Colombia[7] y la realización de una encuesta cara a cara aplicada
a los espectadores en la entrada/salida y los alrededores de los escenarios deportivos. Figura 2. Tasa de ocupación
hotelera en Cali
2010-2013
43.8%
41.0%
44.0%
92.0%
47.8%
42.0%
49.0%
78.0%
0%
20%
40%
60%
80%
100%
2010
2011
2012
2013*
Agosto
Julio
Nota: *Corresponde al período del 1 al 4 de agosto y al
período del 24 al 31 de julio. . Estimación del número de espectadores y de su gasto 4. Estimación del número de espectadores y de su gasto
Los Juegos Mundiales se desarrollaron durante 10 días, los seis últimos del mes de julio y los
primeros cuatro del mes de agosto de 2013. Las competiciones tuvieron lugar en 25 escenarios
con capacidad para albergar a 108.259 espectadores[9]. Los tres escenarios de mayor capacidad
son el Estadio Olímpico Pascual Guerrero (25.486)[10], la Plaza de Toros de Cañaveralejo (19.000) y
el Coliseo El Pueblo (18.700) que representan el 58 per cent del total de la capacidad. El 25 de julio
se llevó a cabo la ceremonia de inauguración (no hubo competencias) y el 4 de agosto se realizó la
ceremonia de clausura. El cálculo de los espectadores requirió usar distintas fuentes de
información, a través de un conjunto de matices y especificaciones que procedemos a explicar. 4.1 La encuesta Asistentes a los Juegos Mundiales 2013 Cali 3.2 Efectos indirectos Fuente: Asociación Hotelera del Valle del Cauca,
COTELVALLE
JEFAS
22,43
136 136 Figura 2. Tasa de ocupación
hotelera en Cali
2010-2013 Nota: *Corresponde al período del 1 al 4 de agosto y al
período del 24 al 31 de julio. Fuente: Asociación Hotelera del Valle del Cauca,
COTELVALLE 3.3 Efecto inducido El efecto inducido es el efecto combinado de los dos gastos anteriores (Comité Organizador
Local y espectadores) sobre el conjunto de la economía local, a través de dos canales, el canal de
relaciones intersectoriales (por ejemplo, la construcción de un nuevo escenario deportivo
significa un aumento en la demanda de insumos —ladrillo, aluminio, acero— a otros sectores)
y el canal de consumo asociado a los nuevos ingresos que la actividad económica genera (por
ejemplo, la contratación de mano de obra —ingenieros, obreros de construcción—, utilidades
del sector de obras civiles). Los efectos inducidos son calculados a través de multiplicadores
derivados de la Matriz insumo-producto (MIP) para el Valle del Cauca, elaborada por Duque
et al. (2013). Esta matriz es la más reciente disponible en la región para este tipo de estudios a
falta de una MIP para Cali y AM[8]. p
y
[ ]
La matriz parte de información del valor bruto de la producción, del valor agregado y de
la demanda final, y utiliza el método RAS para calcular el cuadrante de consumo intermedio
(Duque et al., 2013). La matriz calculada es de tipo producto por producto, simétrica, a
valores básicos y presentados en pesos constantes de 2005. Se encuentra desagregada en 37
sectores de la economía del Valle del Cauca; sin embargo, para los fines de este artículo se ha
agregado en 18 sectores de acuerdo con las dos grandes fuentes de demanda identificadas. 4.2 El cálculo de los turistas extranjeros Gracias al apoyo de Migración Colombia se habilitó temporalmente en el sistema de registro
de los diferentes puestos de control migratorio en el país la categoría “Juegos Mundiales”
entre el 18 de julio y el 4 de agosto de 2013[13]. Los resultados muestran que en el período
ingresaron 10.481 extranjeros al país con destino a la ciudad de Cali, de los cuales 7.006
manifestaron que la principal razón para visitar Cali era participar de las actividades
asociadas a los Juegos Mundiales, distribuidos así: 4.832 visitantes en calidad de deportistas,
jueces, miembros de federaciones deportivas, prensa, invitados especiales, etc. y 2.174 como
espectadores/turistas extranjeros. Esta estrategia para determinar el número de turistas extranjeros resulta novedosa en
este tipo de estudios, en la medida en que implica un cálculo directo y preciso del grupo de
interés. Usualmente, en la literatura de evaluación de impacto económico se utilizan
estimaciones indirectas basadas en encuestas al público asistente y en la ocupación hotelera. Otra ventaja de esta estrategia es que elimina el sesgo de los visitantes casuales, es decir, los
espectadores extranjeros que asisten a los Juegos pero su visita a la ciudad es explicada por
otra razón distinta a la celebración de los Juegos (visita a familiares, razones de estudio/
trabajo). 4.1 La encuesta Asistentes a los Juegos Mundiales 2013 Cali J
g
Se diseñó una encuesta cara a cara[11] aplicada a los espectadores en los sitios de entrada/
salida y alrededores de los escenarios de competición, para ello se requirió el apoyo de 56 voluntarios/as de los Juegos que a través de tres reuniones recibieron capacitación en el
desarrollo de encuestas cara a cara, un conocimiento profundo de los cuestionarios y de los
objetivos de la investigación. Los voluntarios/as se distribuyeron en función de la
programación de las competiciones y la capacidad de los escenarios. Asimismo, se diseñaron
dos modelos de cuestionario: uno dirigido a los turistas extranjeros (en inglés) y otro para los
locales, tanto para residentes como para no residentes de Cali y AM. En total, se aplicaron
1.299 encuestas: 508 a turistas extranjeros, 282 a turistas nacionales y 509 a residentes del
área metropolitana de Santiago de Cali[12]. voluntarios/as de los Juegos que a través de tres reuniones recibieron capacitación en el
desarrollo de encuestas cara a cara, un conocimiento profundo de los cuestionarios y de los
objetivos de la investigación. Los voluntarios/as se distribuyeron en función de la
programación de las competiciones y la capacidad de los escenarios. Asimismo, se diseñaron
dos modelos de cuestionario: uno dirigido a los turistas extranjeros (en inglés) y otro para los
locales, tanto para residentes como para no residentes de Cali y AM. En total, se aplicaron
1.299 encuestas: 508 a turistas extranjeros, 282 a turistas nacionales y 509 a residentes del
área metropolitana de Santiago de Cali[12]. 137 La encuesta permitió capturar:
las
características
generales
de
los
asistentes,
tanto
demográficas
com
socioeconómicas;
la frecuencia de asistencia a las competiciones; es decir, para un mismo individuo, el
número medio de competiciones y el número medio de días que asiste;
la estructura del gasto derivado de la asistencia a las competiciones; y
el motivo de asistencia y grado de satisfacción. Para cada uno de los anteriores apartados la información distingue entre los tres tipos de
espectadores identificados: locales residentes en Cali y AM, turistas nacionales y turistas
extranjeros. Efectos de la
realización de
un megaevento
deportivo Figura 3.
Número de
espectadores* que
presenciaron los
Juegos Mundiales
2013 Cali 4.3 Otros aspectos metodológicos clave La empresa encargada por el Comité Local Organizador de Cali para la venta de las entradas
proporcionó el número total de entradas disponibles, las efectivamente vendidas y las de
cortesía por día, escenario y disciplina deportiva, incluyendo las ceremonias de inauguración
y clausura. Si bien en principio resulta claro que incluir el gasto del público local residente en
Cali y AM representa una recomposición de su gasto en el tiempo y en la misma área
geográfica, se hace necesario incorporar a aquellos espectadores locales residentes en el área
de impacto que pudiendo salir de la ciudad (fuga de gasto) se quedaron, producto de una
elección racional, para participar como espectadores por la novedad que significaban los
Juegos Mundiales. Este público asistente es capturado a través de la siguiente pregunta de la encuesta: >Ha asistido a otras competiciones internacionales en los últimos tres años? Asimismo, se tiene en cuenta el fenómeno observado de rotación de entradas entre
familiares/amigos de la población residente en Santiago de Cali. Este fenómeno es explicado
en parte por las largas jornadas de las competiciones y porque la entrada permitía el acceso
durante toda la jornada. Así, por ejemplo, una persona usaba la misma entrada en la mañana
y otra la usaba en la tarde, a lo largo del mismo día. 138 4.4. >Cuántos espectadores presenciaron los juegos mundiales 2013 cali? La anterior información y refinamientos metodológicos permitieron convertir el número
total de entradas (vendidas þ cortesía) en número total de asistentes por tipo de residencia
que se muestran en el Figura 3. Esta metodología de estimación del número de espectadores
también se puede calificar de novedosa en los estudios de impacto económico. En efecto,
conociendo el número de espectadores internacionales, a partir de la encuesta se puede
estimar el número de turistas nacionales y de espectadores residentes en Cali y AM, a través
de la pregunta sobre el “motivo de asistencia”. Además, la información de la encuesta
permite corregir el número de entradas vendidas por la frecuencia de asistencia, es decir, por
el número medio de competiciones vista por cada tipo de espectador. De esta forma, se
convierte el número de entradas en número de espectadores, lo que asegura evitar cometer
un error muy común en los estudios de impacto económico de sobreestimar el número de
asistentes, que en muchos estudios rebasa la capacidad de los escenarios de competencia y/o
la capacidad hotelera local. JEFAS
22,43 4.3 Otros aspectos metodológicos clave 2,174
11,250
90,249
50,540
0
20,000 40,000 60,000 80,000100,000
Turistas Internacionales
Turistas Nacionales
Residentes en Cali y área
metropolitana**
Ceremonia Inaugural y Clausura***
Notas: *Corresponde al número de espectadores corregido por la
frecuencia de asistencia. Es decir, teniendo presente que un mismo
espectador asiste a distintas competiciones a lo largo de los días de
celebración de los Juegos; **incluye corrección por el número
de residentes de Cali que compartieron la entrada, con ello se
aumentó el número de personas que disfrutaron de las
competiciones; ***incluye corrección para evitar la duplicación
de asistentes que presenciaron ambas ceremonias
Fuente: Elaboración propia sobre la base de la Encuesta Asistentes
a los Juegos Mundiales 2013 Cali e información de Migración
Colombia 2,174
11,250
90,249
50,540
0
20,000 40,000 60,000 80,000100,000
Turistas Internacionales
Turistas Nacionales
Residentes en Cali y área
metropolitana**
Ceremonia Inaugural y Clausura*** Figura 3. Número de
espectadores* que
presenciaron los
Juegos Mundiales
2013 Cali
2,174
11,250
90,249
50,540
0
20,000 40,000 60,000 80,000100,000
Turistas Internacionales
Turistas Nacionales
Residentes en Cali y área
metropolitana**
Ceremonia Inaugural y Clausura***
Notas: *Corresponde al número de espectadores corregido por la
frecuencia de asistencia. Es decir, teniendo presente que un mismo
espectador asiste a distintas competiciones a lo largo de los días de
celebración de los Juegos; **incluye corrección por el número
de residentes de Cali que compartieron la entrada, con ello se
aumentó el número de personas que disfrutaron de las
competiciones; ***incluye corrección para evitar la duplicación
de asistentes que presenciaron ambas ceremonias
Fuente: Elaboración propia sobre la base de la Encuesta Asistentes
a los Juegos Mundiales 2013 Cali e información de Migración
Colombia Notas: *Corresponde al número de espectadores corregido por la
frecuencia de asistencia. Es decir, teniendo presente que un mismo
espectador asiste a distintas competiciones a lo largo de los días de
celebración de los Juegos; **incluye corrección por el número
de residentes de Cali que compartieron la entrada, con ello se
aumentó el número de personas que disfrutaron de las
competiciones; ***incluye corrección para evitar la duplicación
de asistentes que presenciaron ambas ceremonias
Fuente: Elaboración propia sobre la base de la Encuesta Asistentes
a los Juegos Mundiales 2013 Cali e información de Migración
Colombia Figura 3. Efectos de la
realización de
un megaevento
deportivo 4.5 Estimación del gasto de los espectadores La estructura de gasto de los espectadores, en especial de aquellos que vienen de fuera de la
ciudad atraídos por la celebración de los Juegos, depende de un conjunto de factores, entre
ellos: la duración de la estancia en la ciudad y del número de competiciones a las que asista,
del tipo de alojamiento y el sistema de transporte que utilicen en la ciudad. La duración de la
estadía en la ciudad por parte de los turistas extranjeros es en promedio de 4,8 días y asisten
en promedio a 4,4 competiciones en los diez días de duración de los Juegos (Figura 4). En el
caso de los turistas nacionales la estancia media es de 4,1 días y una media de asistencia a
4,0 competiciones. La participación como asistente a eventos deportivos dinamiza la
interacción social que se refleja en estilos de vida compartidos. En efecto, los turistas
extranjeros asisten con un promedio de 4.5 acompañantes frente a 3,9 y 3,6 de los turistas
nacionales y asistentes locales, respectivamente. El 58,3 per cent de los turistas extranjeros se hospeda en hoteles, cerca del 13 per cent en
pensiones/hostales y el 23,3 per cent en casa de familiares/amigos (Figura 5). Por su parte, el
58,4 per cent de los turistas nacionales usó como principal forma de alojamiento la casa de
familiares/amigos y el 30,6 per cent se aloja bien sea en hoteles o pensiones/hostales. Las
anteriores cifras permiten ilustrar cómo cada subgrupo de espectadores tiene una estructura
de gasto diferente. Para los efectos de la estimación tomamos como criterio de clasificación
del gasto el lugar de residencia. La Tabla III muestra el gasto total por tipo de espectador, excluyendo el gasto en
entradas, que fue suministrado por la empresa encargada de su venta ($4.137 mill. de pesos),
lo que permite identificar las actividades económicas más beneficiadas por la “turbina de
gasto” asociada a los Juegos. Este gasto constituye la inyección total de recursos a través del
consumo que dinamiza los sectores vinculados al turismo como la alimentación, la hotelería,
el transporte y el comercio local. De igual forma se beneficia el Estado a través de una mayor
recaudación de impuestos derivados de las distintas transacciones. 4.3 Otros aspectos metodológicos clave Número de
espectadores* que
presenciaron los
Juegos Mundiales
2013 Cali En resumen, los Juegos Mundiales 2013 Cali fueron vistos en vivo en los distintos escenarios
deportivos por 103.673 personas (ver Figura 3), de las cuales 2.174 fueron turistas
internacionales, 11.250 turistas nacionales y 90.249 residentes en Cali y AM. Asimismo,
50.540 espectadores presenciaron las ceremonias de inauguración y clausura. p
p
g
y
El perfil de los asistentes a los Juegos Mundiales 2013 Cali indica que se trata de un
espectador de mediana edad, en su mayoría hombres, casados o en unión libre y con
educación universitaria (ver Tabla II). En el caso de los turistas extranjeros, el rango de edad
que más asistió fue entre los 30 y 39 años (29,6 per cent), el 64,8 per cent son hombres, el 51,7
per cent de estado civil con algún vínculo formal (casado/unión libre) y el 80 per cent cuenta
con titulación universitaria. Asimismo se trata, en el caso del turista internacional, de un
espectador especializado y con experiencia. En efecto, para el caso del 41,3 per cent de los
turistas extranjeros su trabajo/estudio está vinculado con actividades deportivas y el 71,1
per cent ha asistido a eventos deportivos internacionales en los últimos tres años. Para los
turistas nacionales y los residentes en Cali estos porcentajes caen fuertemente, indicando con
ello, en parte, que se trata de espectadores más motivados por la novedad de las disciplinas
deportivas y la experiencia de asistir a un evento deportivo de escala mundial. Por nivel de
ingresos mensuales, la mayoría de los espectadores tiene ingresos de entre $1.200.000 [US
$635] y $3.600.000 [US$1,904]. 139 Tabla II. Tabla II. Figure 5.
Tipo de hospedaje de
los asistentes a los
Juegos Mundiales
2013 Cali 5. Impacto económico p
El objetivo de este apartado es presentar los resultados de la estimación del impacto
económico de los Juegos Mundiales sobre la economía de Cali y AM. El impacto Variables
Turista Extranjero
Turista Nacional
Residente en Cali y área metropolitana
Rango de Edad (años)
18-24
25-29
30-39
40-49
>50
18-24
25-29
30-39
40-49
>50
18-24
25-29
30-39
40-49
>50
15.2%
17.2%
29.6%
17.8%
20.2%
13.9%
13.1%
25.3%
22.0%
25.7%
12.5%
9.7%
26.3%
27.9%
23.6%
Sexo
Hombre
64.8
61.6
56.2
Mujer
35.2
38.4
43.8
Estado civil
Soltero
44.1
44.1
37.0
Casado/unión libre
51.7
49.0
56.3
Viudo/divorciado
4.1
6.9
6.7
Nivel educativo
Ninguno
0.2
0.0
0.4
Primaria
0.4
1.2
0.8
Secundaria
10.2
6.9
11.5
Técnico
9.1
11.8
12.7
Universitaria/Postgrado
80.0
80.0
74.6
Su trabajo/estudios están relacionados con la actividad deportiva
Si
41.3
22.9
12.7
Ha asistidio a otras competiciones internacionales en los últimos 3 años
Si
71.1
46.1
40.2
Actividad principal
Estudiar
9.6
13.5
12.1
Trabajar
81.1
77.1
74.3
Oficios del hogar
0.7
4.1
6.5
Buscar trabajo
1.1
0.0
1.0
Jubilado
3.7
3.3
4.4
Otra
3.9
2.0
1.6
(continued)
Tabla II. Perfil
sociodemográfico de
los asistentes a los
Juegos Mundiales
2013 Cali
JEFAS
22,43
140 140 Variables
Turista Extranjero
Turista Nacional
Residente en Cali y área metropolitana
Nivel de ingresos mensuales (pesos colombianos)
<$600.000
10.3
8.2
11.4
$601.000-$1.200.000
15.7
15.5
17.7
$1.201.000-$2.400.000
19.0
23.7
23.3
$2.401.000-$3.600.000
19.7
21.6
16.8
$3.601.000-$4.800.000
16.2
13.5
11.0
$4.801.000-$6.000.000
7.9
8.6
8.9
>$6.001.000
11.4
9.0
11.0
Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales 2013 Cali. Elaboración propia
Tabla II. Efectos de la
realización de
un megaevento
deportivo
141 Variables
Turista Extranjero
Turista Nacional
Residente en Cali y área metropolitana
Nivel de ingresos mensuales (pesos colombianos)
<$600.000
10.3
8.2
11.4
$601.000-$1.200.000
15.7
15.5
17.7
$1.201.000-$2.400.000
19.0
23.7
23.3
$2.401.000-$3.600.000
19.7
21.6
16.8
$3.601.000-$4.800.000
16.2
13.5
11.0
$4.801.000-$6.000.000
7.9
8.6
8.9
>$6.001.000
11.4
9.0
11.0
Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales 2013 Cali. Elaboración propia Efectos de la
realización de
un megaevento
deportivo 141 141 económico se mide como la suma de tres efectos. En primer lugar, los efectos directos
derivados del gasto/inversión en que incurrió el Comité Local Organizador de Cali para
garantizar la celebración de los Juegos. 5. Impacto económico Tipo de hospedaje de
los asistentes a los
Juegos Mundiales
2013 Cali Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos
Mundiales 2013 Cali Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos
Mundiales 2013 Cali p
p
Mundiales 2013 Cali
la inversión en construcción de nuevos escenarios deportivos y la adecuación y
remodelación de los ya existentes;
la inversión en construcción de nuevos escenarios deportivos y la adecuación y
remodelación de los ya existentes;
el gasto en preparación, gestión y logística. La inversión en construcción y adecuación de infraestructura deportiva para llevar a cabo
los Juegos Mundiales fue un esfuerzo tripartito entre los gobiernos local, departamental y
nacional. El esfuerzo del gobierno local corresponde al 44,2 per cent del total de la inversión
(Tabla IV), el gobierno nacional apoyó con el 48 per cent y el gobierno departamental con
cerca del 8 per cent. 143 Siguiendo la metodología adoptada en este estudio, para evitar cometer un error
ampliamente documentado en los estudios de impacto económico, la inversión de la Alcaldía de
Santiago de Cali en escenarios deportivos se excluye del efecto directo, dado que no puede ser
considerada como una nueva inyección de recursos en el área de impacto de los Juegos. Esta
constituye una transferencia de los mismos residentes locales en forma de gasto público local,
frente a otros usos potenciales de estos recursos. Finalmente, como inyección de nuevos
recursos financieros al área de impacto, en este caso Cali y AM, ingresaron $24.175 millones de
pesos correspondientes al apoyo de los gobiernos nacional y departamental. p
p
p y
g
y
p
El gasto en operación y logística para la realización de los Juegos se muestra en el Tabla
V. La información se presenta en las partidas que utiliza el Comité Olímpico Colombiano
(COC). En total los recursos ascienden a $66.255 millones de pesos financiados en su
totalidad por el gobierno nacional en cabeza de Coldeportes[15]. El gasto en operación y
logística de acuerdo con la metodología adoptada, para tener en cuenta solo la inyección de
recursos provenientes de fuera del área de impacto y que beneficien a los agentes locales
(sectores productivos locales), requiere ajustes en tres partidas:
contratos internacionales, puesto que corresponden a equipos importados alquilados
para los procesos de medición y cronometraje en las competencias; Tabla III. Efectos de la
realización de
un megaevento
deportivo 5. Impacto económico En segundo lugar, los efectos indirectos,
derivados del gasto que los espectadores de los Juegos realizan en la ciudad, en la
medida en que son inherentes al hecho de su celebración. Por último, los efectos
inducidos derivados de la dinámica que genera el impulso de las dos demandas
anteriores, vía relaciones intersectoriales (compras a otros sectores) y de consumo
(nuevos ingresos), sobre toda la economía local. 142 5.1 Efecto directo
El efecto directo incluye todos los gastos/inversiones necesarios para poner en marcha los
Juegos Mundiales. En este caso se consideran dos grandes fuentes de gasto/inversión por
parte del Comité Local Organizador de Cali[14]: 5.1 Efecto directo
El efecto directo incluye todos los gastos/inversiones necesarios para poner en marcha los
Juegos Mundiales. En este caso se consideran dos grandes fuentes de gasto/inversión por
parte del Comité Local Organizador de Cali[14]: 4.8
4.1
4.3
4.4
4.0
4.5
4.5
3.9
3.6
0.0
1.0
2.0
3.0
4.0
5.0
6.0
Turista Extranjero
Turista Nacional
Residente en Cali y A.M. Número medio de acompañantes
Número medio de competiciones vistas
Número medio de días asistencia
Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales
2013 Cali 4.8
4.1
4.3
4.4
4.0
4.5
4.5
3.9
3.6
0.0
1.0
2.0
3.0
4.0
5.0
6.0
Turista Extranjero
Turista Nacional
Residente en Cali y A.M. Número medio de acompañantes
Número medio de competiciones vistas
Número medio de días asistencia
Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales
2013 Cali Figure 4. Duración media de la
estancia, número
medio de
competiciones y
acompañantes de los
asistentes a los Juegos
Mundiales 2013 Cali Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales
2013 Cali 58.3%
22.0%
12.8%
8.6%
23.3%
58.4%
5.7%
11.0%
0%
10%
20%
30%
40%
50%
60%
70%
Turista Extranjero
Turista Nacional
Otro
En casa de familiar/amigo
Pensión/Hostal
Hotel
Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos
Mundiales 2013 Cali Turista Nacional Turista Extranjero Figure 5. 5. Impacto económico Gasto total de los
espectadores [cifras
en pesos 2013]
Tipo de Consumo
Turista
Extranjero
Turista Nacional
Residente
Cali y A.M. Total
Alojamiento
$ 1,248,745,461
$ 3,154,258,391
$ 0
$ 4,403,003,852
14.3%
Alimentación
$ 713,253,218
$ 1,900,533,906
$ 7,738,784,649
$ 10,352,571,773
33.6%
Transporte Local
$ 534,951,751
$ 1,252,874,373
$ 5,101,579,580
$ 6,889,405,703
22.4%
Compras/Recuerdos
$ 574,170,087
$ 1,015,911,937
$ 4,136,692,156
$ 5,726,774,180
18.6%
Ocio/Diversión
$ 852,810,232
$ 509,430,261
$ 2,074,349,251
$ 3,436,589,745
11.2%
Total
$ 3,923,930,749
$ 7,833,008,867
$ 19,051,405,636
$ 30,808,345,252
100%
Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales 2013 Cali Elaboración propia sobre la base de la Encuesta Asistentes a los Juegos Mundiales 2013 Cali Tabla IV. Inversión en
infraestructura
deportiva (cifras en
pesos de 2013)
Entidad
Obra civil
Interventoria
Total
(%)
Alcaldía Santiago de Cali
$ 18,163,169,691
$ 955,956,300
$ 19,119,125,990
44.2
Gobernación del Valle del Cauca
$ 3,230,000,000
$ 170,000,000
$ 3,400,000,000
7.9
Gobierno Nacional [Coldeportes]
$ 19,735,863,587
$ 1,038,729,662
$ 20,774,593,249
48.0
Total
$ 41,129,033,277
$ 2,164,685,962
$ 43,293,719,239
100
Fuente: Comité Local Organizador de Cali - Secretaria del Deporte, Alcaldía de Santiago de Cali Tabla V. Gasto en gestión,
logística y
organización (cifras
en pesos de 2013)
Concepto
Valor
(%)
Implementación
$ 11.962.898.607
18.1
Contratos internacionales
$ 8.376.422.862
12.6
Tecnología
$ 5.862.516.211
8.8
Hoteles
$ 5.422.557.519
8.2
Gastos administrativos
$ 4.573.791.131
6.9
Alimentación
$ 4.461.985.231
6.7
Promoción
$ 3.866.146.939
5.8
Recurso humano
$ 3.820.796.721
5.8
Montaje
$ 3.329.738.754
5.0
Manejo administrativo
$ 3.286.534.087
5.0
Transporte
$ 2.594.802.981
3.9
Plan de medios
$ 2.025.572.397
3.1
Inauguración/clausura
$ 2.000.000.000
3.0
Otras \bacute\aéreas logísticas
$ 1.892.106.901
2.9
Protocolo
$ 672.045.813
1.0
Voluntariado
$ 551.180.928
0.8
Deportes
$ 541.774.066
0.8
Dopaje
$ 430.857.684
0.7
Aérea médica
$ 363.569.969
0.5
Medio ambiente
$ 219.828.700
0.3
Total
$ 66.255.127.503
100
Fuente: Comité Local Organizador de Cali
JEFAS
22,43
144
tecnología y servicios de transmisión de televisión, en ambos casos solo se incluye el
gasto realizado en el área de impacto (gastos locales por parte de los técnicos), y
tecnología y servicios de transmisión de televisión, en ambos casos solo se incluye el
gasto realizado en el área de impacto (gastos locales por parte de los técnicos), y
manejo administrativo, en lo correspondiente a la devolución de los derechos de
inscripción del equipo de Gimnasia de Israel que no se presentó. 5. Impacto económico
manejo administrativo, en lo correspondiente a la devolución de los derechos de
inscripción del equipo de Gimnasia de Israel que no se presentó. Asimismo, se excluyeron los convenios —canje— con el Masivo Integrado de Occidente –
MIO– para el transporte de los voluntarios y con EMCALI para suministro de servicios
públicos y comunicaciones. Asimismo, se excluyeron los convenios —canje— con el Masivo Integrado de Occidente –
MIO– para el transporte de los voluntarios y con EMCALI para suministro de servicios
públicos y comunicaciones. Una vez realizados los ajustes, como inyección de nuevos recursos financieros al área de
impacto ingresaron $57.445 millones de pesos. En las Tablas 8, 9 y 10 se presenta la
información homologada con los sectores de la Matriz insumo-producto (MIP) para el Valle
del Cauca. En resumen, el efecto directo asociado a la celebración de los Juegos Mundiales
2013 Cali significó una inyección de $81.620 millones de pesos a la economía de Cali y AM,
representados por $24.175 millones de pesos de inversión en infraestructura deportiva y
$57.445 millones de pesos en gastos de operación, gestión y logística[16]. 5.2 Efecto indirecto
i
i
d l La estimación del número de espectadores efectuada en el apartado 4.5. se refina al incluir
residentes locales que no se desplazaron a otro lugar debido precisamente a la celebración de
los Juegos (ver la Figura 6). Se trata de una equivalencia entre individuos que decidieron
quedarse en su casa y asistir a los eventos programados en lugar de desplazarse de
vacaciones a otro lugar, producto de una elección racional. El cálculo se ha realizado con
ayuda de la pregunta consignada en la Encuesta a Asistentes sobre si ha asistido a otras
competiciones internacionales en los últimos tres años: Figure 6. Espectadores de fuera
del área
metropolitana de
Santiago de Cali
Atracción de visitantes de fuera
de Cali y área metropolitana
Turistas extranjeros
11.250
Turistas nacionales
2.174
36.280
Residentes locales
Gasto en: entradas, alojamiento,
alimentación, transporte local,
compras/recuerdos, ocio/diversión
Impacto en la producción de bienes y
servicios y sobre el empleo en Cali y
AM
Elaboración propia
Efectos de la
realización de
un megaevento
deportivo
145 Efectos de la
realización de
un megaevento
deportivo Atracción de visitantes de fuera
de Cali y área metropolitana Gasto en: entradas, alojamiento,
alimentación, transporte local,
/
d
i /di
ió Figure 6. Espectadores de fuera
del área
metropolitana de
Santiago de Cali Figure 6. Espectadores de fuera
del área
metropolitana de
Santiago de Cali Impacto en la producción de bienes y
servicios y sobre el empleo en Cali y
AM Elaboración propia Elaboración propia
El supuesto subyacente indica que quienes responden afirmativamente siguen la
actividad deportiva, disponen de los recursos financieros para desplazarse pero
deciden quedarse en la ciudad para seguir las competiciones del deporte de su
interés y/o por la novedad que implican los Juegos Mundiales.
El supuesto subyacente indica que quienes responden afirmativamente siguen la
actividad deportiva, disponen de los recursos financieros para desplazarse pero
deciden quedarse en la ciudad para seguir las competiciones del deporte de su
interés y/o por la novedad que implican los Juegos Mundiales.
Adicionalmente, el 97 per cent de los espectadores residentes en Cali y AM
considera que la celebración de los Juegos Mundiales 2013 Cali generó un impacto
positivo sobre el “orgullo y sentido de pertenencia de los caleños por su ciudad”. Figure 6.
Espectadores de fuera
del área
metropolitana de
Santiago de Cali 5.2 Efecto indirecto
i
i
d l Tal
como lo ha puesto de manifiesto Zimbalist (2010) para los residentes locales este tipo
de celebraciones es un momento “corto pero intenso” a través del cual “la atención
del mundo se centrará en sus hogares” lo que a corto plazo genera “una gran
sensación de logro”.
Adicionalmente, el 97 per cent de los espectadores residentes en Cali y AM
considera que la celebración de los Juegos Mundiales 2013 Cali generó un impacto
positivo sobre el “orgullo y sentido de pertenencia de los caleños por su ciudad”. Tal
como lo ha puesto de manifiesto Zimbalist (2010) para los residentes locales este tipo
de celebraciones es un momento “corto pero intenso” a través del cual “la atención
del mundo se centrará en sus hogares” lo que a corto plazo genera “una gran
sensación de logro”. La Tabla VI muestra el efecto indirecto asociado a la inyección de gasto adicional en
Cali y AM derivado de los espectadores de los Juegos Mundiales. Un total de $19.416
millones de pesos que resultan de la suma del gasto de los turistas extranjeros y
nacionales, y el gasto de los residentes locales que se quedaron en la ciudad para
presenciar las competiciones. Tabla VI. Estimación del efecto
indirecto (cifras en
pesos de 2013)
Tipo de espectadores
Número de espectadores
Gasto medio por persona
Gasto total
Turistas extranjeros
2.174
$ 1.805.237
$ 3.924.586.128.93
Turistas nacionales
11.250
$ 696.242
$ 7.832.724.858.48
Residentes locales
36.280
$ 211.099
$ 7.658.665.065.52
Elaboración propia 5.3 Efecto total sobre la producción Una vez descritos y estimados los efectos directos e indirectos, resulta clave analizar cómo
estas fuentes de demanda inicial se irrigan hacia los distintos sectores productivos para
conocer y determinar el efecto o impacto total sobre el sistema económico local. Una parte de
esa demanda directa que perciben las empresas locales constituye fuente de nueva demanda
en forma de compras a otras empresas e individuos, que les proveen los insumos necesarios
para realizar su producción (efecto indirecto), este aumento de la producción a su vez genera
nuevos ingresos, que inducen nuevas rondas de gasto y aumento de la producción y así
sucesivamente (efecto inducido), hasta llegar al efecto total sobre el sistema económico. JEFAS
22,43 5.2 Efecto indirecto
i
i
d l 146 El resultado generado en la simulación, al introducir el cambio en la demanda final por
$101.036 millones de pesos derivados de las dos fuentes de demanda identificadas: $81.620
millones de pesos (infraestructura y logística) más $19.416 millones de pesos (gasto de
espectadores) generó un impacto total acumulado de $569.493 millones de pesos sobre la
economía de la ciudad de Cali y AM medido a través del valor bruto de la producción y
utilizando los multiplicadores de producción tipo II (ver Tabla VII). Como puede apreciarse
en la Tabla VII, el impacto sobre la producción recae sobre tres sectores (72,9 per cent) donde
se concentra el gasto en logística de los Juegos (servicios a las empresas, 28,01
per cent), la inversión en infraestructura deportiva (construcción, 23,38 per cent) y la
“turbina de gasto” de los espectadores (hostelería, 21,55 per cent). s
Sector
Descripción
Inyección de
nuevos recursos
[Mill. $]
Impacto sobre la
producción
Mill. $
(%)
Servicios a las empresas
[p.e. gastos en gestión del
CLOC; montaje y adecuación
de escenarios; interventoria
obras civiles; alquiler de
equipos de medición y
cronometraje; transport e de
equipos; servicios de
vigilancia]
$27.803
$159.497
28.01
Construcción y reparación
de edificaciones
[p.e. inversión en
infraestructura de escenarios]
$22.966
$133.122
23.38
Alojamiento, suministro
de comidas y bebidas
[p.e. gasto de los espectadores
en alojamiento y alimentación]
$20.110
$122.739
21.55
Otra industria
manufacturera
[p.e. gasto en uniformes de
voluntarios]
$11.748
$46.899
8.24
Servicios de transporte
[p.e. gasto en transporte local
de los turistas y delegaciones]
$6.563
$38.807
6.81
Servicios de
esparcimiento
[p.e. gasto de los espectadores
en ocio/diversión, boletería]
$4.249
$25.839
4.54
Comercio y servicios de
reparación
[p.e. gasto de los espectadores
en compras de recuerdos]
$3.253
$19.536
3.43
Servicios financieros e
inmobiliarios
[p.e. gasto en arrendamiento
sedes, bodegas]
$1.808
$10.873
1.91
Correo y
telecomuncaciones
[p.e. gasto en promoción y
medios de comunicación]
$1.115
$6.755
1.19
Maquinaria y equipo de
transporte
[p.e. gasto en compra PC,
impresoras, celulares, etc.]
$1.421
$5.428
0.95
Total
$101.036
$569.493
100
Elaboración propia Sector Tabla VIII. Impacto sobre el
empleo por sectores
económicos
Número de empleos
Sector
Descripción
Ocupados
Equivalente de tiempo
completo
Comercio y servicios de reparación [p.e. gasto de los espectadores en
compras de recuerdos]
3.085
2.479
32.14%
Servicios de transporte
[p.e. gasto en transporte local de los
turistas y delegaciones]
1.355
1.089
14.12%
Producción agropecuaria,
silvicultura y pesca
[p.e. demanda derivada]
935
751
9.74%
Alojamiento, suministro de
comidas y bebidas
[p.e. gasto de los espectadores en
alojamiento y alimentación]
772
620
8.04%
Construcción y reparación de
edificaciones
[p.e. gastos en infraestructura de
escenarios]
636
511
6.63%
Servicios de enseñanza y salud de
mercado
[p.e. gasto de logistica en el \bacute
\aérea médica y manejo de dopaje]
558
448
5.82%
Servicios de esparcimiento
[p.e. gasto de los espectadores en ocio/
diversión, boletería]
482
387
5.02%
Servicios a las empresas
[p.e. gastos en gestión del CLOC;
montaje y adecuación de escenarios;
interventoria obras civiles; alquiler de
equipos de medición y cronometraje;
transporte de equipos; servicios de
vigilancia]
481
387
5.01%
Otra industria manufacturera
[p.e. gasto en uniformes de voluntarios] 383
307
3.99%
Servicios del gobierno
[p. ej. demanda derivada]
240
193
2.50%
Correo y telecomuncaciones
[p.e. gasto en promoción y medios de
comunicación]
234
188
2.44%
Industria alimenticia
[p. ej. demanda derivada]
200
161
2.09%
Servicios financieros e
inmobiliarios
[p.e. gasto en arrendamiento sedes,
bodegas]
143
115
1.49%
Maquinaria y equipo de transporte [p.e. gasto en compra PC, impresoras,
celulares, etc.]
42
34
0.44%
Trabajos de construcción,
construcción de obras civiles y
servicios de arrendamiento de
equipo con operario
[p.e. demanda derivada]
28
22
0.29%
Electricidad, gas, agua y
saneamiento b\bacute\aésico
[p.e. demanda derivada]
23
19
0.24%
Productos de minería
[p.e. demanda derivada]
1
1
0.01%
Total
9.598
7.711
100%
Elaboración propia
Efectos de la
realización de
un megaevento
deportivo
147 Elaboración propia 5.4 Efecto sobre el empleo 5.4 Efecto sobre el empleo
En cuanto al empleo, el cambio en la demanda final por $101.036 millones de pesos creó un
total de 9.598 puestos de trabajo (ocupados), equivalentes a 7.711 empleos de tiempo
completo (ver Figura 7)[17]. De estos últimos, 1.775 son causa directa del impulso inicial de
demanda (empleos directos), 1.013 son empleos indirectos, derivados de los incrementos en
la demanda de otros sectores productivos y 4.923 inducidos asociados a los nuevos efectos
sobre los propios proveedores vía ingresos y consumo. La Tabla VIII muestra el empleo por sectores económicos. Figure 7.
Impacto agregado
sobre el empleo 6. El legado de los Juegos Mundiales para Cali
D
d
R
F
hli h
id
d
I 6. El legado de los Juegos Mundiales para Cali
De acuerdo con Ron Froehlich, presidente de International World Games Association
(IWGA) los pasados Juegos Mundiales 2013 Cali son “la mejor de las nueve versiones de los
Juegos, por eso la medalla de oro es para Cali y Colombia[18]”. Este éxito inicial de los Juegos
Mundiales 2013 Cali no garantiza que en el largo plazo los efectos positivos se mantengan. Lo que podemos denominar “el legado de los Juegos”, depende de un conjunto complejo de
factores e interacciones que no son capturados por las relaciones monetarias entre agentes y
sectores que se describen en la Figura 4. Con el objetivo de que perdure la ventaja
competitiva generada por la realización del megaevento deportivo, se requiere de una
adecuada gestión por parte de los actores clave (sector público —local, regional, nacional—,
sector deportivo, las empresas privadas y la comunidad) que mitigue los riesgos
identificados (ver la Tabla IX). (
)
La Figura 8 propone una interacción estratégica entre los actores clave (stakeholders) que
pueden dinamizar “el legado de los Juegos”. Los actores clave arriba identificados a través de
una adecuada interacción y adopción de un conjunto de acciones son quienes garantizan el
sostenimiento en el largo plazo de los impactos positivos de los Juegos. En la Figura 4 se
presentan en forma sucinta algunas de las acciones relevantes, que implican algo más allá de
una actitud de colaboración entre los actores locales clave, significa que realmente se
comparten experiencias, recursos y capacidades para adquirir, desarrollar y consolidar
ventajas competitivas en la organización de megaeventos deportivos dentro de una visión
estratégica global de gestión del territorio; al respecto, ver Radicchi (2012, pp. 38-39). Sector Los sectores que más empleo
generaron, en términos relativos, fueron los vinculados con la logística de la organización de los Juegos y la “turbina de gasto” de los turistas. Es así como en el sector comercio y
servicios de reparación se generaron 2.479 empleos equivalentes de tiempo completo,
seguidos del sector de servicios de transporte con 1.089, el sector de producción
agropecuaria, silvicultura y pesca con 751 y el sector de alojamiento, suministro de comidas
y bebidas con 620. 148 6. El legado de los Juegos Mundiales para Cali JEFAS
22,43 RIESGOS Capital Físico/Infraestructura: infraestructura física
para la realización de eventos deportivos de escala
mundial Los escenarios deportivos se deterioran
físicamente; las adecuaciones y remodelaciones
sin mantenimiento y actualización permanente
se vuelven obsoletas. Adem\bacute\aés de que
representan una carga para el presupuesto
público local/nacional 149 Capital humano/social: profesionales capacitados en
áreas relacionadas con la realización de grandes
eventos deportivos. Beneficios sociales del Programa
de Voluntarios*. Beneficios sociales de la nueva
infraestructura deportiva y la diversidad en la
práctica deportiva que posibilitan**
“Marca de Ciudad”: mejora en la percepción y
posicionamiento de la ciudad en el contexto nacional
e internacional como ciudad “deportiva, moderna y
competitiva” Capital humano/social: profesionales capacitados en
áreas relacionadas con la realización de grandes
eventos deportivos. Beneficios sociales del Programa
de Voluntarios*. Beneficios sociales de la nueva
infraestructura deportiva y la diversidad en la
práctica deportiva que posibilitan** La experiencia de los profesionales vinculados
con la organización y logística, asimismo de los
voluntarios, se puede subutilizar en la medida en
que no puedan aplicar su experiencia con la
organización y oferta de otros eventos de gran
escala “Marca de Ciudad”: mejora en la percepción y
posicionamiento de la ciudad en el contexto nacional
e internacional como ciudad “deportiva, moderna y
competitiva” Sin promoción y apoyo a la práctica del deporte
recreativo y competitivo el uso de las
instalaciones deportivas se puede desviar para
otros fines. Sin que ello implique que la nueva
infraestructura deportiva no se incorpore dentro
del equipamiento cultural y social de la ciudad
para el uso y disfrute del tiempo libre de los
residentes y visitantes de Cali
Sin una gestión adecuada de la Marca de Ciudad
se pueden perder los beneficios positivos
asociados con la buena imagen para la ciudad
que deja el éxito deportivo y de asistencia a los
Juegos Mundiales como potencial destino
turístico de futuros visitantes Intangible: Aumento del sentido de identidad,
pertenencia, autoestima y cohesión social Notas: *La convocatoria para voluntarios recibió la inscripción de 5.872 personas, se capacitaron 5.366 y
finalmente se seleccionaron 2.539; **Por ejemplo, como legado de los Juegos Mundiales quedan nuevos
deportes incluidos en el sistema nacional, entre ellos: batalla de fuerza; baile deportivo y escalada. Elaboración propia conocimiento de los autores, este sería el primer estudio de esta naturaleza aplicado a un
megaevento deportivo en Colombia. Tabla IX.
El legado de los
Juegos Mundiales
2013 Cali 7. Conclusiones
l En este artículo se estimó el impacto económico a corto plazo asociado a la celebración de la
IX edición de los Juegos Mundiales sobre la ciudad de Cali y su área metropolitana. En Figure 7. Impacto agregado
sobre el empleo
2,209
1,261
6,128
9,598
1,775
1,013
4,923
7,711
Directo
Indirecto
Inducido
Total
Directo
Indirecto
Inducido
Total
Ocupados
Empleo Tiempo
Completo
Elaboración propia 2,209
1,261
6,128
9,598
1,775
1,013
4,923
7,711
Directo
Indirecto
Inducido
Total
Directo
Indirecto
Inducido
Total
Ocupados
Empleo Tiempo
Completo
Elaboración propia Tabla IX. El legado de los
Juegos Mundiales
2013 Cali
RIESGOS
Los escenarios deportivos se deterioran
físicamente; las adecuaciones y remodelaciones
sin mantenimiento y actualización permanente
se vuelven obsoletas. Adem\bacute\aés de que
representan una carga para el presupuesto
público local/nacional
La experiencia de los profesionales vinculados
con la organización y logística, asimismo de los
voluntarios, se puede subutilizar en la medida en
que no puedan aplicar su experiencia con la
organización y oferta de otros eventos de gran
escala
Sin promoción y apoyo a la práctica del deporte
recreativo y competitivo el uso de las
instalaciones deportivas se puede desviar para
otros fines. Sin que ello implique que la nueva
infraestructura deportiva no se incorpore dentro
del equipamiento cultural y social de la ciudad
para el uso y disfrute del tiempo libre de los
residentes y visitantes de Cali
Sin una gestión adecuada de la Marca de Ciudad
se pueden perder los beneficios positivos
asociados con la buena imagen para la ciudad
que deja el éxito deportivo y de asistencia a los
Juegos Mundiales como potencial destino
turístico de futuros visitantes
ripción de 5.872 personas, se capacitaron 5.366 y
legado de los Juegos Mundiales quedan nuevos
batalla de fuerza; baile deportivo y escalada. Efectos de la
realización de
un megaevento
deportivo
149 Efectos de la
realización de
un megaevento
deportivo RIESGOS En resumen, la celebración de los Juegos Mundiales
significó para la ciudad de Cali:
En lo económico, la movilización de recursos que dinamizó diversos sectores de
la economía local: 2.174 turistas internacionales, 11.250 turistas nacionales, 9.598
puestos de trabajo equivalentes a 7.711 empleos de tiempo completo.
En lo deportivo, una oportunidad para continuar consolidando y explotando las
ventajas competitivas que ha adquirido en la realización de eventos deportivos de
escala mundial.
En lo deportivo, una oportunidad para continuar consolidando y explotando las
ventajas competitivas que ha adquirido en la realización de eventos deportivos de
escala mundial.
En lo social, exaltar el sentido de pertenencia y el orgullo de los residentes locales
por su ciudad, a través de la celebración de un megaevento deportivo
excepcional,
inédito en América
Latina. Noventa y
siete
de
cada
100
espectadores residentes en Cali y AM consideraron que la celebración de los
Juegos Mundiales tuvo un efecto positivo sobre el “orgullo y sentido de
pertenencia de los caleños por su ciudad”. Figure 8. Actores y acciones
para sostener “el
legado de los Juegos”
JEFAS
22,43
150 150 150 Figure 8.
Actores y acciones
para sostener “el
legado de los Juegos” Efectos de la
realización de
un megaevento
deportivo Figure 8. El principal aporte del artículo se centra en el desarrollo de algunos puntos que resultan
metodológicamente útiles en la elaboración de estudios de impacto económico en ciudades
de países en desarrollo. En efecto, en el cálculo del flujo de espectadores (nacionales y
extranjeros) se refina la estimación teniendo presente: El principal aporte del artículo se centra en el desarrollo de algunos puntos que resultan
metodológicamente útiles en la elaboración de estudios de impacto económico en ciudades
de países en desarrollo. En efecto, en el cálculo del flujo de espectadores (nacionales y
extranjeros) se refina la estimación teniendo presente: El principal aporte del artículo se centra en el desarrollo de algunos puntos que resultan
metodológicamente útiles en la elaboración de estudios de impacto económico en ciudades
de países en desarrollo. En efecto, en el cálculo del flujo de espectadores (nacionales y
extranjeros) se refina la estimación teniendo presente:
El cálculo directo vía administrativa de los turistas extranjeros, a través de un
acuerdo con la autoridad migratoria colombiana, lo que implica un cálculo preciso
del grupo de interés.
Se corrige el posible “efecto expulsión” y de “cancelación de reservas”.
Se incluyeron aquellos espectadores locales, residentes en el área de impacto, que
ante el costo de oportunidad de salir de la ciudad (fuga de gasto) optó por quedarse
y presenciar los Juegos.
Se corrige el número de espectadores al incluir el fenómeno observado de rotación de
entradas entre familiares/amigos de la población residente en Santiago de Cali. Por último, desde una perspectiva de largo plazo, el principal legado de los Juegos se dirige a
consolidar las ventajas competitivas que ha adquirido la ciudad en la organización de
megaeventos deportivos. En el marco de una visión estratégica global de gestión del
territorio, resulta necesaria una fluida interrelación entre los actores locales clave (sector
público, sector privado, sector deportivo y comunidad). Notas El aporte del gobierno de Cali se
estima en un 44,2 per cent para la infraestructura construida. 7. La Unidad Administrativa Especial Migración Colombia es la entidad del Estado Colombiano
encargada de “ejercer control como autoridad migratoria a ciudadanos nacionales y extranjeros
en el territorio colombiano”, ver www.migracioncolombia.gov.co/index.php/entidad/la–unidad/
mision–vision.html 8. La elaboración de una MIP para Cali y AM constituiría un esfuerzo que sobrepasa los alcances de
este artículo por los altos requerimientos de información y tiempo necesarios para su
construcción, y revela la falta de una contabilidad regional y local en Colombia. La utilización de
la MIP actualizada para el Valle del Cauca constituye la mejor aproximación disponible para el
cálculo de los multiplicadores, puesto que la estructura económica de Cali refleja la estructura
económica del Valle del Cauca. Lo anterior se observa en el peso relativo de Cali y AM en el PIB
del Valle del Cauca, que asciende al 67,9 per cent (DANE, 2013). 9. Los escenarios se utilizaron en forma simultánea, aunque no todos los días. Por ejemplo, el
Estadio Olímpico Pascual Guerrero se utilizó del 28 al 30 de julio para las competencias de disco
volador y del 1 al 2 de agosto para rugby 7. 10. Corresponde a las tribunas habilitadas para presenciar las competiciones y no a la capacidad
total del estadio. 11. El sistema de encuesta elegido fue la entrevista personal cara a cara. Este sistema tiene sus
ventajas; con un entrevistador entrenado, una vez que se aborda al entrevistado y este acepta la
realización de la misma, es difícil que la encuesta no sea válida ya que el número de preguntas
contestadas es prácticamente del 100 per cent. No obstante, tiene el inconveniente de realizarse en
un momento próximo al disfrute del evento por parte del entrevistado, lo que la hace difícil y
costosa. 12. El diseño muestral siguió el procedimiento sugerido por Levy and Lameshow (1991) aceptando un
error muestral del 5 per cent y un nivel de confianza del 95 per cent. Las 1.299 encuestas constituyen
una muestra representativa del total de espectadores de los Juegos Mundiales 2013 Cali, estratificada
según el origen (turistas extranjeros, turistas nacionales y residentes en Cali y AM). 12. El diseño muestral siguió el procedimiento sugerido por Levy and Lameshow (1991) aceptando un
error muestral del 5 per cent y un nivel de confianza del 95 per cent. Notas 1. La ciudad de Santiago de Cali tiene amplia tradición en la celebración de eventos deportivos de
gran escala, por citar algunos: VI Juegos Panamericanos en 1971, I Juegos del Océano Pacífico en
1995, Campeonato del Mundo de Patinaje de Velocidad en 2007, Copa Mundo de Ciclismo de Pista
en 2008, XVIII Campeonato Mundial de Futbol Sub-20 en 2011 (sede grupo B). 1. La ciudad de Santiago de Cali tiene amplia tradición en la celebración de eventos deportivos de
gran escala, por citar algunos: VI Juegos Panamericanos en 1971, I Juegos del Océano Pacífico en
1995, Campeonato del Mundo de Patinaje de Velocidad en 2007, Copa Mundo de Ciclismo de Pista
en 2008, XVIII Campeonato Mundial de Futbol Sub-20 en 2011 (sede grupo B). 2. Los primeros Juegos Mundiales se realizaron en 1981 en la ciudad de Santa Clara (Estados
Unidos). Desde entonces ocho ciudades han sido sede: Londres (Reino Unido), Karlsruhe
(Alemania), La Haya (Países Bajos), Lahti (Finlandia), Akita (Japón), Duisbur (Alemania) y
Kaohsiung (China). La próxima edición en 2017 se llevará a cabo en la ciudad de Wroclaw
(Polonia). 151 3. Adicionalmente, cinco disciplinas deportivas participaron como invitadas. 4. Tasa de cambio representativa del mercado (TCRM) $/US$1,890.33, fin de mes (julio de 2013). Fuente: Banco de la República/Superintendencia Financiera de Colombia. (www.banrep.gov.co/
es/series–estadisticas/see_ts_trm.htm). 4. Tasa de cambio representativa del mercado (TCRM) $/US$1,890.33, fin de mes (julio de 2013). Fuente: Banco de la República/Superintendencia Financiera de Colombia. (www.banrep.gov.co/
es/series–estadisticas/see_ts_trm.htm). 5. Por sus siglas en inglés: International World Games Association, IWGA (www.theworldgames. org/). 5. Por sus siglas en inglés: International World Games Association, IWGA (www.theworldgames. org/). 5. Por sus siglas en inglés: International World Games Association, IWGA (www.theworldgames
org/). 6. La estrategia de financiación de los Juegos Mundiales implicó la construcción de cinco nuevos
escenarios en terrenos del municipio y/o públicos; los demás escenarios fueron adaptados y
remodelados para las competiciones. La financiación recayó sobre el gobierno nacional
(Coldeportes) tanto para infraestructura como para la logística. El aporte del gobierno de Cali se
estima en un 44,2 per cent para la infraestructura construida. 6. La estrategia de financiación de los Juegos Mundiales implicó la construcción de cinco nuevos
escenarios en terrenos del municipio y/o públicos; los demás escenarios fueron adaptados y
remodelados para las competiciones. La financiación recayó sobre el gobierno nacional
(Coldeportes) tanto para infraestructura como para la logística. Notas De acuerdo con el DANE, los empleos (u ocupados) hacen referencia al contrato
explícito o implícito entre una persona y una unidad institucional para realizar un trabajo a
cambio de remuneración, durante un período definido o hasta nuevo aviso, y se dividen en dos
grandes grupos: los asalariados y los independientes. Estos trabajos pueden ser de tiempo
completo, medio tiempo o empleos ocasionales. 17. Los empleos de tiempo completo equivalente se calculan como el total de horas trabajadas
dividido el promedio anual de horas trabajadas en puestos de trabajo considerados de tiempo
completo. De acuerdo con el DANE, los empleos (u ocupados) hacen referencia al contrato
explícito o implícito entre una persona y una unidad institucional para realizar un trabajo a
cambio de remuneración, durante un período definido o hasta nuevo aviso, y se dividen en dos
grandes grupos: los asalariados y los independientes. Estos trabajos pueden ser de tiempo
completo, medio tiempo o empleos ocasionales. 17. Los empleos de tiempo completo equivalente se calculan como el total de horas trabajadas
dividido el promedio anual de horas trabajadas en puestos de trabajo considerados de tiempo
completo. De acuerdo con el DANE, los empleos (u ocupados) hacen referencia al contrato
explícito o implícito entre una persona y una unidad institucional para realizar un trabajo a
cambio de remuneración, durante un período definido o hasta nuevo aviso, y se dividen en dos
grandes grupos: los asalariados y los independientes. Estos trabajos pueden ser de tiempo
completo, medio tiempo o empleos ocasionales. 18. Ver:
www.ntn24.com/noticias/italia–se–consagra–campeon–de–los–juegos–mundiales–en–la–
ciudad–colombiana–de–cali–101438 18. Ver:
www.ntn24.com/noticias/italia–se–consagra–campeon–de–los–juegos–mundiales–en–la–
ciudad–colombiana–de–cali–101438 Notas Las 1.299 encuestas constituyen
una muestra representativa del total de espectadores de los Juegos Mundiales 2013 Cali, estratificada
según el origen (turistas extranjeros, turistas nacionales y residentes en Cali y AM). 13. Por razones de deficiente infraestructura de transporte terrestre, ferroviario y fluvial en el país y
la topografía del territorio se estima como mínima y no significativa la llegada de turistas
extranjeros por otra vía distinta del trasporte aéreo. 14. Las diferentes partidas de la inversión en infraestructura, los gastos de organización y logística y
el gasto de los espectadores se homologa en los sectores económicos en los que se desagrega la
matriz insumo producto del Valle del Cauca, utilizada en la simulación. Se emplean deflactores
implícitos calculados a partir de las Cuentas Departamentales y de las Cuentas Trimestrales del
DANE. 15. El Departamento Administrativo del Deporte, la Recreación, la Actividad Física y el
Aprovechamiento del Tiempo Libre –Coldeportes– es el ente oficial del Estado colombiano
“encargado de formular, coordinar y vigilar la práctica del deporte, la recreación, la educación
física, el aprovechamiento del tiempo libre y la actividad física, orientados al mejoramiento de la
calidad de vida de la sociedad colombiana”, ver [www.coldeportes.gov.co/]. 16. En este total se incluye el ingreso por venta de entradas que de acuerdo con la contabilidad del
CLOC se incorporó en el presupuesto general de los Juegos. 17. Los empleos de tiempo completo equivalente se calculan como el total de horas trabajadas
dividido el promedio anual de horas trabajadas en puestos de trabajo considerados de tiempo
completo. De acuerdo con el DANE, los empleos (u ocupados) hacen referencia al contrato
explícito o implícito entre una persona y una unidad institucional para realizar un trabajo a
cambio de remuneración, durante un período definido o hasta nuevo aviso, y se dividen en dos
grandes grupos: los asalariados y los independientes. Estos trabajos pueden ser de tiempo
completo, medio tiempo o empleos ocasionales. 18. Ver:
www.ntn24.com/noticias/italia–se–consagra–campeon–de–los–juegos–mundiales–en–la–
ciudad–colombiana–de–cali–101438
JEFAS
22,43
152 13. Por razones de deficiente infraestructura de transporte terrestre, ferroviario y fluvial en el país y
la topografía del territorio se estima como mínima y no significativa la llegada de turistas
extranjeros por otra vía distinta del trasporte aéreo. 13. Notas Por razones de deficiente infraestructura de transporte terrestre, ferroviario y fluvial en el país y
la topografía del territorio se estima como mínima y no significativa la llegada de turistas
extranjeros por otra vía distinta del trasporte aéreo. 14. Las diferentes partidas de la inversión en infraestructura, los gastos de organización y logística y
el gasto de los espectadores se homologa en los sectores económicos en los que se desagrega la
matriz insumo producto del Valle del Cauca, utilizada en la simulación. Se emplean deflactores
implícitos calculados a partir de las Cuentas Departamentales y de las Cuentas Trimestrales del
DANE. 14. Las diferentes partidas de la inversión en infraestructura, los gastos de organización y logística y
el gasto de los espectadores se homologa en los sectores económicos en los que se desagrega la
matriz insumo producto del Valle del Cauca, utilizada en la simulación. Se emplean deflactores
implícitos calculados a partir de las Cuentas Departamentales y de las Cuentas Trimestrales del
DANE. 152 15. El Departamento Administrativo del Deporte, la Recreación, la Actividad Física y el
Aprovechamiento del Tiempo Libre –Coldeportes– es el ente oficial del Estado colombiano
“encargado de formular, coordinar y vigilar la práctica del deporte, la recreación, la educación
física, el aprovechamiento del tiempo libre y la actividad física, orientados al mejoramiento de la
calidad de vida de la sociedad colombiana”, ver [www.coldeportes.gov.co/]. 15. El Departamento Administrativo del Deporte, la Recreación, la Actividad Física y el
Aprovechamiento del Tiempo Libre –Coldeportes– es el ente oficial del Estado colombiano
“encargado de formular, coordinar y vigilar la práctica del deporte, la recreación, la educación
física, el aprovechamiento del tiempo libre y la actividad física, orientados al mejoramiento de la
calidad de vida de la sociedad colombiana”, ver [www.coldeportes.gov.co/]. 16. En este total se incluye el ingreso por venta de entradas que de acuerdo con la contabilidad del
CLOC se incorporó en el presupuesto general de los Juegos. 16. En este total se incluye el ingreso por venta de entradas que de acuerdo con la contabilidad del
CLOC se incorporó en el presupuesto general de los Juegos. 17. Los empleos de tiempo completo equivalente se calculan como el total de horas trabajadas
dividido el promedio anual de horas trabajadas en puestos de trabajo considerados de tiempo
completo. Referencias Ávalos, A. (2011), “Estimación del impacto económico de los Juegos Panamericanos Guadalajara 2011:
análisis de insumo-producto”, Revista Econoquantum, Vol. 8 No. 1, pp. 35-60. aade, R. y Matheson, V. (2016), “Going for the gold: the economics of the Olympics”, Journal of
Economic Perspectives, Vol. 30 No. 2, pp. 201-218. arajas, A., Salgado, J. y Sánchez, P. (2012), “Problemática de los estudios de impacto económico de
eventos deportivos”, Estudios De Economía Aplicada, Vol. 30, pp. 441-462. arget, E. y Gouguet, J. (2008), “Un análisis de la copa del mundo de rugby 2007 en Francia”, RAE:
Revista Asturiana De Economía, Vol. 42, pp. 103-127. DANE (2013), Metodología Para Calcular el Indicador de Importancia Económica Municipal, DANE,
Bogotá. Devesa, M., Báez, A., Figueroa, V. y Herrero, L.C. (2012), “Repercusiones económicas y sociales de los
festivales culturales. El caso del festival internacional de Cine de Valdivia”, EURE–Revista
Latinoamericana de Estudios Urbano Regionales, Vol. 38 No. 115, pp. 95-115. Duque, H., Escobar, J., Aponte, E., Flórez, J. y Aguirre, A. (2013), Impacto Del TLC Con Estados Unidos
en la producción y el empleo del Valle del Cauca. Gratton, C., Shibli, S. y Coleman, R. (2006), “The economic impact of major sports events: a review of ten
events in the UK”, The Sociological Review, Vol. 54 No. s2, pp. 41-58. Levy, P. y Lameshow, S. (1991), Sampling of Populations: Methods and Applications, Wiley, New York,
NY. Llopis-Goig, R. (2012), Megaeventos Deportivos: Perspectivas Científicas y Estudios de Caso, Editorial
UOC, Barcelona. Matheson, V. (2006), “Economic impact analysis”, en Andreff, W. and Szymanski, S. (Edits), Handbook
on the Economics of Sport, Edward Elgar, Cheltenham. Matheson, V. (2012), “Efectos de los principales megaeventos deportivos en las economías locales”, en
Llopis-Goig, R. (Ed.), Megaeventos Deportivos. Perspectivas Científicas y Estudios de Caso, UOC,
Barcelona, pp. 53-74. Porter, P. y Chin, D. (2012), “Economic impact of sports events”, en Maennig, W. and Zimbalist, R. (Eds),
International Handbook on the Economics of Mega Sporting Events, Edward Elgar Publishing,
pp. 246-268. 153 Radicchi, E. (2012), “Megaeventos deportivos y creación de valor para las economías anfitrionas”, en
Llopis-Goig, R. (Ed.), Megaeventos deportivos. Perspectivas científicas y Estudios de Caso, UOC,
Barcelona, pp. 25-51. Roche, M. (1994), “Mega-events and urban policy”, Annals of Tourism Research, Vol. 21 No. 21, pp. 1-19 Schuschny, A. (2005), Tópicos Sobre el Modelo de Insumo-producto: Teoría y Aplicaciones, Serie
Estudios Estadísticos y Prospectivos No. 37. Referencias División de Estadística y Proyecciones Económicas
de la CEPAL, Santiago de Chile. Siegfried, J. y Zimbalist, A. (2006), “The economic impact of sports facilities, teams and mega-events”,
The Australian Economic Review, Vol. 39 No. 4, pp. 420-427. Zimbalist, A. (2010), “>Merece la pena?”, Revista Finanzas y Desarrollo, marzo, pp. 8-11. Zimbalist, A. (2015), “The illusory economic gains from hosting the Olympics and World Cup”, World
Economics, Vol. 16 No. 1, pp. 35-42. Efectos de la
realización de
un megaevento
deportivo For instructions on how to order reprints of this article, please visit our website:
www.emeraldgrouppublishing.com/licensing/reprints.htm
Or contact us for further details: permissions@emeraldinsight.com Corresponding author
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A
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b Luis F. Aguado can be contacted at: lfaguado@javerianacali.edu.co For instructions on how to order reprints of this article, please visit our website:
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Frozen elephant trunk reconstruction for right-sided aortic arch with aberrant left subclavian artery and aneurysm of the descending aorta: a case report
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Journal of cardiothoracic surgery
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cc-by
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© 2016 Kokotsakis et al. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver
(http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Background A right-sided aortic arch is an uncommon congenital lesion
occurring in 0.05–0.10 % of the population. It is often associ-
ated with an aberrant left subclavian artery, and patients are
particularly prone to developing aneurysm and dissection [1]. The optimal approach to this condition requires meticulous
planning and depends on the extent of aneurysm formation. We report the successful single stage repair of this vascular
anomaly using the frozen elephant trunk technique. Abstract Background: A 59-year old man being investigated for back pain was found to have aneurysmal dilatation of a
right-sided aortic arch and descending thoracic aorta together with an aberrant left subclavian artery. Case Presentation: He underwent repair of this utilising the frozen elephant trunk technique, which dealt with
all three pathologies in one-stage. He made an unremarkable recovery and was discharged home on the 8th
post-operative day. Conclusions: This case report further demonstrates the flexibility and safety of the frozen elephant trunk in
dealing with complex aortic pathology as a single-stage procedure. Keywords: Aortic arch, Endovascular procedures thoracic aorta (DTA, 6.8 cm at its largest point) located in
the right hemi-thorax, which returned to normal size
below the diaphragm. The root and ascending aorta were
not aneurysmal. An aberrant left subclavian artery (ALSA)
arose from a Kommerell’s diverticulum (3.5 cm) at the
junction between the distal arch and the proximal DTA. The head and neck vessels arose individually from the
arch in the following order: left common carotid artery
(LCCA), right common carotid artery (RCCA), right sub-
clavian artery (RSA) and finally the ALSA as mentioned
above (Fig. 1). Coronary angiography, echocardiography
and carotid doppler scan were unremarkable. After multi-
disciplinary team discussion it was decided to proceed
with the frozen elephant trunk technique. Frozen elephant trunk reconstruction for
right-sided aortic arch with aberrant left
subclavian artery and aneurysm of the
descending aorta: a case report Frozen elephant trunk reconstruction for
right-sided aortic arch with aberrant left
subclavian artery and aneurysm of the
descending aorta: a case report hn Kokotsakis1, Omar A. Jarral2*, Leanne Harling2, Panteleimon Tsipas1 and Thanos Athanasiou2 CASE REPORT
Open Access
Frozen elephant trunk reconstruction for
right-sided aortic arch with aberrant left
subclavian artery and aneurysm of the
descending aorta: a case report
John Kokotsakis1, Omar A. Jarral2*, Leanne Harling2, Panteleimon Tsipas1 and Thanos Athanasiou2
Abstract
Background: A 59-year old man being investigated for back pain was found to have aneurysmal dilatation of a
right-sided aortic arch and descending thoracic aorta together with an aberrant left subclavian artery.
Case Presentation: He underwent repair of this utilising the frozen elephant trunk technique, which dealt with
all three pathologies in one-stage. He made an unremarkable recovery and was discharged home on the 8th
post-operative day.
Conclusions: This case report further demonstrates the flexibility and safety of the frozen elephant trunk in
dealing with complex aortic pathology as a single-stage procedure.
Keywords: Aortic arch, Endovascular procedures
Kokotsakis et al. Journal of Cardiothoracic Surgery (2016) 11:77
DOI 10.1186/s13019-016-0479-y CASE REPORT
Open Access
Frozen elephant trunk reconstruction for
right-sided aortic arch with aberrant left
subclavian artery and aneurysm of the
descending aorta: a case report
John Kokotsakis1, Omar A. Jarral2*, Leanne Harling2, Panteleimon Tsipas1 and Thanos Athanasiou2
Abstract
Background: A 59-year old man being investigated for back pain was found to have aneurysmal dilatation of a
right-sided aortic arch and descending thoracic aorta together with an aberrant left subclavian artery. Case Presentation: He underwent repair of this utilising the frozen elephant trunk technique, which dealt with
all three pathologies in one-stage. He made an unremarkable recovery and was discharged home on the 8th
post-operative day. Conclusions: This case report further demonstrates the flexibility and safety of the frozen elephant trunk in
dealing with complex aortic pathology as a single-stage procedure. Keywords: Aortic arch, Endovascular procedures
Kokotsakis et al. Journal of Cardiothoracic Surgery (2016) 11:77
DOI 10.1186/s13019-016-0479-y Kokotsakis et al. Journal of Cardiothoracic Surgery (2016) 11:77
DOI 10.1186/s13019-016-0479-y CASE REPORT
Open Access
Frozen elephant trunk reconstruction for
right-sided aortic arch with aberrant left
subclavian artery and aneurysm of the
descending aorta: a case report
John Kokotsakis1, Omar A. Jarral2*, Leanne Harling2, Panteleimon Tsipas1 and Thanos Athanasiou2
Abstract
B
k
d A 59
ld
b i
i
i
d f
b
k
i
f
d
h
l dil
i
f
DOI 10.1186/s13019 016 0479 y CASE REPORT
Open Access
Frozen elephant trunk reconstruction for
right-sided aortic arch with aberrant left
subclavian artery and aneurysm of the
descending aorta: a case report
John Kokotsakis1, Omar A. Jarral2*, Leanne Harling2, Panteleimon Tsipas1 and Thanos Athanasiou2 * Correspondence: omarjarral@gmail.com
2Department of Surgery and Cancer, Imperial College London, London W2
1NY, UK
Full list of author information is available at the end of the article Case presentation A 59-year old male was referred to our clinic with a six-
month history of back pain. His medical history included
hypertension and a 60-pack year history of smoking. Phys-
ical examination including assessment of peripheral pulses
was unremarkable. Chest radiograph revealed a laterally
displaced trachea and a right-sided aortic arch. Computer-
ized tomographic angiography (CTA) confirmed a right-
sided aortic arch (4.1 cm) and an aneurysmal descending After induction of anaesthesia, arterial lines were
placed in the left femoral and both radial arteries. A
lumbar cerebrospinal fluid drain was placed in order to
manipulate cord perfusion pressure. A soft guidewire
was placed in the right femoral artery (RFA) and ad-
vanced in to the arch. A median sternotomy was per-
formed with left supraclavicular extension in order to
expose the distal LSA. The right axillary artery (RAA)
was exposed with an infraclavicular incision. After Page 2 of 4 Kokotsakis et al. Journal of Cardiothoracic Surgery (2016) 11:77 Fig. 1 Computer generated images of aorta Fig. 1 Computer generated images of aorta were transected 1 cm distal to their origin and the RSA was
ligated at its origin from the aortic arch. Perfusion catheters
were inserted in to the LCCA, the RCCA, and together with
the RAA cannula already in place, continuous bilateral se-
lective antegrade cerebral perfusion (SACP) was started. heparinization, a 8 mm graft was anastomosed to the
RAA through this incision on the underside of the artery. The other end of the graft was tunnelled through the right
pleural space in to the pericardial cavity and the arterial
line (bifurcated) of the cardiopulmonary bypass (CPB) cir-
cuit was connected to this end. Examination of the aorta
revealed no evidence of dissection or rupture. At this point the soft guidewire in the RFA was
exchanged for a stiff one. A hybrid stent-graft (E-vita
OPEN PLUS 40 × 160 mm, Jotec Inc., Hechingen,
Germany) was advanced in to the DTA in an antegrade
fashion. The proximal landmark for stent deployment
was just distal to the origin of the RSA. The incorpo-
rated polyester graft was cut back and sutured to the
transected distal arch with interrupted horizontal mattress
sutures (3-0 Prolene, Ethicon, Somerville, NJ) reinforced
with a strip of Teflon. A sterilized bronchoscope enabled
the distal part of the stent graft to be checked under direct
vision. Case presentation After going on to CPB a right superior pulmonary vein
vent was placed and the patient was cooled to 25 °C. During
cooling the head and neck vessels were dissected out, and
upon fibrillation the ascending aorta was clamped and trans-
ected just above the sinotubular junction. The heart was
arrested using direct cold crystalloid cardioplegia. Subsequent
shots were delivered retrogradely every 20 min. Once the
target temperature was reached, CPB was arrested and the
ascending aorta and part of the arch (up to 2 cm before the
origin of the RSA) were resected. The LCCA and RCCA Page 3 of 4 Page 3 of 4 Kokotsakis et al. Journal of Cardiothoracic Surgery (2016) 11:77 Attention was then turned to the ascending aorta where
a vascular prosthesis with four side branches (30 mm
Tetrabranch, Jotec Inc.) was anastomosed distally to the
arch cuff (comprising the native aorta and the E-vita pros-
thesis/Teflon strip as described above) using a continuous
suture (3-0 Prolene, Ethicon). Systemic perfusion was re-
established by connecting the arterial line of the CPB
circuit to the 4th side branch of the vascular prosthesis. The 3rd side branch (8 mm) was passed under the innom-
inate vein and was anastomosed end-to-end to the RCCA
(after removal of the perfusion catheter) with a continuous
suture (5-0 Prolene, Ethicon). The proximal anastomosis
of the main graft to the sinotubular junction was then per-
formed using a continuous suture (3-0 Prolene, Ethicon)
with an external strip of Teflon. Rewarming was com-
menced; a de-airing drill completed and the cross-clamp
was removed. The 1st side branch (10 mm) was passed
under the innominate vein and anastomosed end-to-end
to the LCCA (after removal of the perfusion catheter) with
a continuous suture (5-0 Prolene, Ethicon). Arterial inflow
to the RAA graft, which had been tunnelled to sit in the pericardial cavity, was then stopped and this end of the
graft was anastomosed end-to-end to the 2nd side
branch(8 mm) of the prosthesis using a continuous su-
ture (5-0 Prolene, Ethicon). A separate 8 mm spiral
graft was then anastomosed end-to-side to the ALSA in
the left supraclavicular area and connected proximally
end-to-side to the 1st branch of the main vascular pros-
thesis (Fig. 2). The ALSA was ligated proximal to the
origin of the left vertebral artery. After complete rewarming the patient was separated
from CPB uneventfully. Fig. 2 Intraoperative photo References Submit your next manuscript to BioMed Central
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and we will help you at every step: Case presentation The bypass times were as
follows: CPB (160 mins), SACP (90 mins) and, distal
ischaemic time (50mins). Following haemostasis, the
synthetic grafts were covered with a strip of pericardium
to exclude them from the sternal wound prior to rou-
tine closure. The patient was extubated 6 h following
the operation and discharged on the 8th post-operative
day at which point he was mobile and neurologically in-
tact. Twelve-month follow-up with CTA confirmed sat-
isfactory appearance of the reconstruction with absence
of endoleak. Fig. 2 Intraoperative photo Fig. 2 Intraoperative photo Page 4 of 4 Kokotsakis et al. Journal of Cardiothoracic Surgery (2016) 11:77 Page 4 of 4 Received: 17 November 2015 Accepted: 2 May 2016 Received: 17 November 2015 Accepted: 2 May 2016 Received: 17 November 2015 Accepted: 2 May 2016 Competing interests Competing interests
The authors declare that they have no competing interests. Competing interests
The authors declare that they have no competing interests. Authors’ contributions
JK PT KV
d TA
f JK, PT, KV and TA performed the operation. TA and JK conceived the report. OJ and LH wrote the first draft with input from all authors. All authors read
and approved the final manuscript. Acknowledgements
None. Author details
1E
l
H 1Evaggelismos Hospital, Athens, Greece. 2Department of Surgery and Cancer,
Imperial College London, London W2 1NY, UK. References 1. Cina CS, Althani H, Pasenau J, Abouzahr L. Kommerell’s diverticulum and
right-sided aortic arch:a cohort study and review of the literature. J Vasc
Surg. 2004;39(1):131–9. 1. Cina CS, Althani H, Pasenau J, Abouzahr L. Kommerell’s diverticulum and
right-sided aortic arch:a cohort study and review of the literature. J Vasc
Surg. 2004;39(1):131–9. 2. Easo J, Holzl P, Dikov V, Chavan A, Dapunt O. Frozen elephant trunk
procedure in a dextropositioned arch and descending aorta. J Endovasc
Ther. 2010;17(6):751–4. In this case we utilised the frozen elephant trunk
technique in one-stage to deal with three pathologies:
arch and descending aortic aneurysm together with an
ALSA originating in a Kommerell’s diverticulum. To
our knowledge, this is only the third such report in
the literature [2, 3]. We consider it to be a safe and
effective technique given the excellent outcome this
patient achieved, especially as some of the above tech-
niques are associated with very significant morbidity. There are now a number of other devices on the mar-
ket (e.g., Thoraflex Hybrid), which would allow a simi-
lar or more advanced outcome to be achieved. Whilst
it could be suggested that TEVAR(after debranching)
is a less invasive option, this was not considered due
to the acute angle between the distal arch and the
DTA, which may have led to difficulty in securing a
good proximal landing. Proceeding with TEVAR in
such a situation may have led to incomplete treatment
of the arch aneurysm and perhaps unwanted compli-
cations such as endoleak. A further disadvantage of
this approach is that of embolic related complications
in patients with extensive atheroma of the distal aorta. In retrospect, we could have simplified our operation
further by establishing CPB via arch cannulation and
reanastomosing the RSA to the new arch graft directly
via the median sternotomy. 3. Munakata M, Itaya H, Fukui K, Ono Y. One-stage repair for aortic
regurgitation and kommerell diverticulum with aneurysmal right aortic arch. J Thorac Cardiovasc Surg. 2007;133(3):798–9. Consent Right-sided aortic arch with ALSA is a rare congenital
abnormality frequently associated with aneurysm and
dissection. Whilst appropriate surgical indications are
not yet clearly established, intervention in the asymp-
tomatic is usually recommended for aneurysms over
3 cm [1]. Meticulous pre-operative imaging and plan-
ning are essential. Current treatment options are tai-
lored to the anatomical features (e.g., type I or II right-
sided arch and the degree of aneurysm formation) of
each case and include: right (or left) thoracotomy with
replacement of the descending thoracic aorta and/or
reconstruction of the ALSA, sternotomy and deb-
ranching of the arch vessels with subsequent thoracic
endovascular aneurysm repair (TEVAR), sternotomy
and total arch replacement with insertion of elephant
trunk and subsequent TEVAR, total arch replacement
with subsequent thoracotomy for DTA replacement,
arch replacement and DTA replacement through the
clamshell approach, and left carotid to left subclavian
artery bypass with ligation or coiling at the origin of
the ALSA [1]. Written informed consent was obtained from the patient
for publication of this case report and any accompanying
images. Conclusion This case report demonstrates the flexibility and safety
of the frozen elephant trunk in dealing with right-sided
aortic arch aneurysms as a single-stage procedure.
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https://openalex.org/W1606192030
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https://revistas.pucp.edu.pe/index.php/boletindearqueologia/article/download/1854/1789, https://revistas.pucp.edu.pe/index.php/boletindearqueologia/article/download/1854/1789/
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El dominio del Inka, identidad local y complejidad social en las tierras altas del desierto de Atacama, Norte Grande de Chile (1450-1541 d.C.)
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Boletín de arqueología PUCP
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301
DEL INKA,
IDENTIDAD
LOCAL
Y COMPLEJIDAD SOCIAL EN ATACAMA...
BOLETÍN EL
DEDOMINIO
ARQUEOLOGÍA
PUCP,
N.° 6, 2002,
301-336
EL DOMINIO DEL INKA, IDENTIDAD LOCAL Y
COMPLEJIDAD SOCIAL EN LAS TIERRAS
ALTAS DEL DESIERTO DE ATACAMA,
NORTE GRANDE DE CHILE (1450-1541 D.C.)
1
Mauricio Uribe,* Leonor Adán** y Carolina Agüero***
Resumen
En este artículo se presenta una aproximación a la expansión del Tawantinsuyo en las tierras altas
del desierto de Atacama (norte de Chile), proponiendo un enfoque alternativo a la tradicional concepción de
una presencia débil o indirecta del Inka en este territorio. Por ello, se exponen aquellas evidencias y argumentos
que parecen representar mejor la complejidad social, económica, política y étnica que se encontraba en pleno
desarrollo cuando el imperio tomó contacto con las poblaciones de este desierto. En particular, los autores se
centran en el caso de San Pedro de Atacama, porque aún cuando se tratán de una de las localidades que ha
servido como punto de referencia para la comprensión de los grupos que habitaron sus tierras altas, todavía
se conoce muy poco de aquellos momentos. Pero, sobre todo, porque el estudio de la convergencia de las
poblaciones atacameñas e «incaicas» dentro de San Pedro, de acuerdo a experiencias en otras localidades de
este territorio, permite vislumbrar en ese encuentro los elementos que particularizan la complejidad social del
desarrollo local.
Abstract
THE INKA’S DOMINION, LOCAL IDENTITY AND SOCIAL COMPLEXITY IN THE HIGHLAND OF
ATACAMA’S DESERT, CHILE’S NORTE GRANDE (1450-1541 AD)
This paper presents our approach to the Tawantinsuyo expansionism in the highlands of Atacama
Desert (Northern Chile), in which we suggest an alternative view to the traditional idea of a weak or indirect
Inka presence in the Atacamenian territory. Therefore, we show the evidence and arguments that represent the
politics, economic and ethnic complexity development when the empire had contact with the populations from
the chilean desert. We concentrate in the case of San Pedro de Atacama, because it had been a point of reference
to the archaeological understanding of groups that lived in its highlands, but the history is poorly known. In
particular the convergence of the study of Atacamenian and Inca people in San Pedro, like in others parts of
this space, allows to appreciate best the elements that characterize the social complexity of this local development.
«La existencia de pucaras [...] hace suponer el deseo de defender un patrimonio territorial e
indica también el establecimiento en esa zona del núcleo fuerte y posiblemente solitario de los
atacameños, tomados como unidad étnica, política, religiosa y social» (Montandón 1950: 15).
1. Introducción
Este trabajo sintetiza la aproximación de los autores acerca de la expansión del Tawantinsuyo
en las tierras altas del desierto de Atacama, norte de Chile (Fig. 1). Sobre la base de las investigaciones
* Universidad de Chile, Departamento de Antropología. E-mail: mur_cl@yahoo.com
** Universidad Austral de Valdivia, Museo Histórico Mauricio van de Maele. E-mail: ladan@uach.cl
*** Universidad Católica del Norte, Instituto de Investigaciones Arqueológicas. E-mail: maguero@ucn.cl
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
realizadas por los autores en la zona desde 1994, se propone un enfoque alternativo a la tradicional
concepción de una presencia débil o indirecta del Inka en este territorio (Llagostera 1976). Por el
contrario, se perfila un conjunto de estrategias económicas, políticas y simbólicas implicadas en el
proceso de anexión de su población al estado, sugeridas por el comportamiento y naturaleza de la
cultura material de este periodo, distribuida entre el río Loa y San Pedro de Atacama (Uribe 19992000).
Esto no concuerda con las interpretaciones que enfatizan el carácter marginal de estos
territorios para los intereses del Tawantinsuyo, lo cual se ha discutido con datos arqueológicos,
etnohistóricos y etnográficos, teniendo en cuenta las particularidades de la historia de la investigación
sobre la temática y las actuales perspectivas de la arqueología regional (Uribe et al. 1999; Adán y
Uribe e.p.). En este sentido, los antecedentes sugieren desde ya un dominio o intervención más
directa del Estado Incaico en Atacama, el que habría privilegiado la interacción con grupos dirigentes
locales, potenciando las contradicciones al interior de su sociedad, entre aquellos y sus bases
(Uribe y Adán e.p.).
Tales contradicciones, consecuentemente, comprometerían la identidad local como la del
Inka. Elementos incaicos y locales fusionados, observados en segmentos específicos de la población
atacameña (Ayala et al. 1999), representan a esos grupos dirigentes que asumieron el funcionamiento
estatal en la región, explotando su propia fuerza de trabajo, sus excedentes agrícolas y riquezas
minerales.
La tutela del Inka, entonces, se expresaría materialmente pura en otros sectores, como los
periféricos respecto al núcleo local, manifestándose con mayor claridad en la vialidad y los
asentamientos asociados al capac ñam (Cornejo 1995).
Estas ideas toman como referente trabajos previos (Castro y Cornejo 1990; Cornejo 1995;
Gallardo et al. 1995) y se fundamentan en análisis tipológicos, tecnológicos y funcionales en cuanto
a la arquitectura, alfarería, industria lítica, textilería y otros materiales de estos periodos, llevados a
cabo por los autores. Los resultados obtenidos permiten, en esta oportunidad, acceder a la complejidad
sociopolítica del desierto de Atacama y, en particular ,de San Pedro (Fig. 1), a través de una evaluación
del substrato local frente al «cinturón» creado por más de una decena de instalaciones del
Tawantinsuyo que se conocen a lo largo de la vertiente occidental circunpuneña de este segmento
de los Andes centro-sur (v.g., Raffino 1981; Castro 1992; Lynch y Núñez 1994).
2. Poblaciones de Atacama en los siglos XVI y XVII
Las poblaciones indígenas que habitaron el área de estudio durante los siglos XVI y XVII
representan identidades locales distinguibles a través de fuentes etnohistóricas, ya sea por discursos externos recogidos por los hispanos, o por la propia documentación colonial local generada a
través del contacto directo con las poblaciones originales de este territorio (v.g., Hidalgo 1983;
Martínez 1985; Castro 1997; Manríquez 2002).
Las informaciones para los siglos en cuestión mencionan, de manera general, a «indios»,
«gentes» o «naturales» de determinados lugares de Atacama «la Grande» o «la Chica» (San Pedro y
el Loa Superior respectivamente), de sus ayllus o pueblos, diferenciando claramente entre éstos y
los indios asentados en la costa.
En el siglo XVI, Oviedo grafica a estos «naturales» como de «tierra alzada é de guerra, y la
gente por los montes, fuera de sus casas é simientes, y puestos en montañas y sierras muy asperas»
(Núñez 1992: 85), relacionando la actitud de los «naturales» con las determinadas características del
entorno que habitaban, es decir, del desierto.
EL DOMINIO DEL INKA, IDENTIDAD LOCAL Y COMPLEJIDAD SOCIAL EN ATACAMA...
303
Fig. 1. Mapa de la región con las principales localidades y sitios mencionados en el texto (Cortesía: V. Castro).
304
MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
Paralelamente, en la provincia de Atacama crónicas y documentos hispanos locales señalan
la existencia de indígenas cuya designación coincide con la de la «provincia». Esta manera de
nombrar subsiste durante el periodo colonial, en el cual la denominación otorgada a un espacio
geográfico (valle), un hito (río), o una división jurídica territorial (provincia, partido, pueblo, etc.),
comienzan a ser apelativos de ciertas poblaciones específicas (Manríquez 2002).
En estos primeros siglos de ocupación hispana tales poblaciones son caracterizadas de
manera genérica como «yndios», «gente que sirvió al inga» y «jente dispuesta y bien bestidos como
los del Piru» (Vivar 1979 [1558]: 20), los cuales, desde ya, son adscritos al espacio territorial nombrados como indios de la provincia de Atacama. Y, en 1557, cuando se inicia el proceso de «pacificación» de la provincia, se los menciona como «yndios de Atacama que estaban de guerra».2
La segunda mitad del siglo XVI marca el inicio de la consolidación hispana en la zona. El
virrey del Perú y el aparato administrativo colonial deciden tomar el control directo de la provincia de
Atacama, por tratarse de uno de los lugares claves en la comunicación entre La Plata y Potosí,
Tucumán y el reino de Chile (Hidalgo 1982). El control se concretó aún más con el traspaso de una
parte de esta extensa encomienda a Juan Velázquez Altamirano, en porción de las distintas
«provincias» que constituían la audiencia de Charcas en Bolivia (Manríquez 2002). Este hecho
indica que los indígenas de Atacama y su territorio son integrados por los españoles dentro de dicha
audiencia, idea que la arqueología y la etnohistoria también comparten al integrar a esta unidad
como parte de la subárea circunpuneña (Lumbreras 1981). De acuerdo a ello, es muy probable que la
administración europea estuviera utilizando, aunque con ciertos reacomodos, un ordenamiento
territorial prehispánico, basado en dinámicas históricas particulares y en un profundo conocimiento
del medioambiente local.
Es entonces cuando estas poblaciones comienzan a ser denominadas en propiedad como
«atacamas». Al respecto, la investigación etnohistórica no reconoce antecedente alguno que oriente
hacia la mención de una autodenominación, puesto que, salvo por los hispanos, ningún indígena se
define como «atacama» o «indio atacama». Por ello se reafirma la idea de una exodenominación que
bien podría tener su origen en las categorizaciones realizadas por otras poblaciones originarias o
por el Tawantinsuyo mismo (Manríquez 2002). El término «Atacamas», por lo tanto, denominaría
genéricamente a indígenas adscritos a un determinado territorio, pero también distingue a los
indígenas que siendo señalados como «originarios» o «naturales» de Atacama se desplazaban de
manera permanente o por periodos variables de tiempo, de preferencia hacia lugares situados en el
Noroeste argentino, Chichas, Tarapacá y el extremo norte del reino de Chile (v.g., Copiapó),
considerados lugares al exterior de los límites de la provincia (Martínez 1998).
Frente a esta representación externa, también se desarrollan otras identidades desde una
escala local o «microscópica» (Manríquez 2002). Así, poblaciones de Atacama la Alta o San Pedro
compartirían códigos de identidad cultural con aquellos que reconocen su origen en espacios y
ayllus situados en Atacama la Baja o el río Loa, en tanto las mismas u otras reconocerían membresía
en el Noroeste argentino, Chichas o Tarapacá (v.g., Odone 1995).
Dentro de esta lógica, los indígenas parecieran relacionar su pertenencia con el origen y
éste con el lugar donde se encuentran sus parientes. En toda la región y en cada uno de los ayllus
serían las relaciones de parentesco y de un origen reconocido como común, piezas importantes en
la articulación de sus relaciones sociales, económicas, políticas y religiosas. Se establece así un
nexo significativo entre parentesco, territorio de origen e identidad que se define por el lugar
donde se ubican los familiares muertos y sus expresiones sagradas (Manríquez 2002). Estas relaciones estarían documentadas ya en el siglo XVI por descripciones como las que realiza indirectamente Vivar (1979 [1558]), sobre el «pueblo o valle de Atacama» cuando se refiere a las «costumbres
EL DOMINIO DEL INKA, IDENTIDAD LOCAL Y COMPLEJIDAD SOCIAL EN ATACAMA...
305
de yndios» y describe que en un apartado de las casas, cerca de donde duermen: «...qu’ es el más
prinçipal, está hecho de bobeda alta fasta el entresuelo y cuadrada. Aqueste es su enterramiento
y sepulcro. Y allí dentro tienen a sus bisaguelos y aguelos y padres y toda su generaçion» (op.
cit.: 20).
Esta situación se advierte igualmente en prácticas religiosas del siglo XVII en las que cada
especialista ofrendaba a un «ídolo», correspondiente a un determinado pueblo, por ejemplo, Qumaquma o Qumaquina de Chiuchiu, Socomba de Aiquina, Sintalacna de Caspana y otros (Castro 1997).
Preliminarmente, por lo tanto, se desprende de estas descripciones y las inferencias hechas que, al
momento del contacto, el territorio atacameño se encontraba constituido por distintas comunidades
adscritas a un extenso espacio desértico, pero, al mismo tiempo, a los espacios de otros a los cuales
podían acceder, de acuerdo a los nexos establecidos por el parentesco, los ancestros y las deidades
propias como compartidas por cada una de esas unidades.
3. San Pedro de Atacama antes y durante el Inka
De manera coherente con lo anterior, las investigaciones arqueológicas llevadas a cabo han
permitido comenzar a definir en Atacama la existencia de un sistema social articulado por poblaciones segmentadas en el espacio, pero fuertemente vinculadas en el plano territorial por aspectos
sagrados y, seguramente, parentales, alcanzando un importante grado de integración macroregional,
al menos entre las poblaciones del río Loa y San Pedro.
A continuación se presentan las evidencias y argumentos arqueológicos que soportan este
planteamiento, particularmente aquellos de San Pedro de Atacama donde se centran actualmente los
estudios de los autores.
3.1. Los asentamientos
El estudio de la arquitectura de los periodos tardíos de San Pedro de Atacama señala un
patrón temprano de construcción en barro representado por la ocupación de los Campos de Sólor
(Fig. 2). La vigencia de esta técnica constructiva tradicional apunta a una cierta continuidad con los
tiempos previos, particularmente con los periodos Formativo y Medio (Le Paige 1958), que se expresa a través de asentamientos-núcleos construidos de barro, articulando a los oasis con los principales cursos de agua del Salar como los ríos San Pedro y el Vilama (Núñez 1992; Llagostera 1996; Uribe
et al. e.p.).
La ocurrencia de estos asentamientos, fuertemente asociados al aprovechamiento económico de los recursos cruciales de la cuenca, los ríos San Pedro y Vilama, plantea la existencia de un
sistema social y económico complejo, aunque, probablemente, poco competitivo en momentos
tempranos del periodo (v.g., dispersión de los asentamientos, ausencia de fortalezas, etc.), pero con
una creciente presión hacia el bien básico y crítico para la vida de las poblaciones atacameñas: el
agua. Este sistema debió generar una dinámica social a lo largo del recurso hídrico, por la cual las
poblaciones fueron acercándose al norte del salar y hacia las quebradas del oriente, resultando
favorecidas aquellas comunidades que podían controlar las cabeceras de dichos cursos (cf. Núñez
1992). Esta situación podría haber generado una importante tensión social durante la evolución del
sistema, lo que en algún momento se expresa en la construcción de pequeños sitios en lugares
estratégicos como los pukara de Quítor y Vilama, en los sectores más septentrionales de la localidad
(Fig. 3 a). Del mismo modo, se aprecia en la experimentación con técnicas agrícolas y de irrigación en
sistemas estancieros de altura como los de Guatin (Fig. 3 b), inicialmente destinados al pastoreo
(Serracino y Stehberg 1974). Por último, esta tensión también se identifica por la introducción de
elementos de apariencia foránea, en especial altiplánica, correspondiente a las estructuras conoci-
a
b
Fig. 2. Campos de Sólor. a. Planta de estructura habitacional de barro; b. Cántaro o tinaja usada para
almacenamiento de líquidos y/o como urna funeraria.
a
b
Fig. 3. a. Pucaras de Vilama. Reductos de cumbre asociados al río Vilama; b. Estancia de Guatin. Se observan
campos o canchas de cultivo.
308
MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
a
b
Fig. 4. a. Aldea de Zápar. Estructura tipo chullpa; b. Aldea de Peine. Estructura tipo chullpa.
das como chullpa (Figs. 4 a, b), entre otras, que aparecen en poblados que se instalan a partir de
aquellos pukara, como Quítor, o en otros completamente nuevos, por ejemplo, Zápar y Peine.
Es casi un consenso, en consecuencia, que durante los primeros momentos de los Desarrollos Regionales se ocuparon pueblos «sin defensa», como Sólor y Guatin, para luego comenzar a
gestarse los pukara típicos del periodo (Le Paige 1958; Bittman et al. 1978; Schiappacasse et al.
1989; Núñez 1992). Al comienzo del Periodo Intermedio Tardío se nota, por lo tanto, la existencia de
un sistema de asentamiento centrado en la explotación de los recursos de los oasis regados por los
principales ríos que drenan el Salar, articulándose los asentamientos-núcleos con otros de pisos
más altos, con gran probabilidad, ocupados para la ganadería de camélidos. Esta arquitectura fue la
que caracterizó a la región después del 950 d.C. hasta cerca del 1350 d.C. (Berenguer et al. 1986;
Tarragó 1989; Uribe e.p.), surgiendo luego otro patrón constructivo más competitivo y directamente
relacionado con el control de los recursos hídricos que se manifestó en la aparición de los pukara,
sean o no amurallados, a partir de los cuales se constituyeron las comunidades donde justamente el
Inka dejó rastros materiales notables.
EL DOMINIO DEL INKA, IDENTIDAD LOCAL Y COMPLEJIDAD SOCIAL EN ATACAMA...
309
Catarpe, Quítor, Zápar y Peine. El trabajo realizado se concentró en ocho sitios habitacionales de
San Pedro, los que han sido registrados en su totalidad a través de la aplicación sistemática de una
ficha arquitectónica (Castro et al. 1991). De éstos, fueron seleccionados los asentamientos Catarpe
Oeste (W), Quítor, Zápar y Peine (Fig. 1), los cuales proveen de información significativa para
entender la época del contacto entre el nuevo desarrollo atacameño y el Inka.
En estos yacimientos (Tabla 1), los rasgos arquitectónicos más distintivos son el gran número de recintos (entre 141 y 210 estructuras), el trazado ortogonal de los poblados, su orientación
cardinal O-E, con sus muros transversales 60º al Norte, así como la conformación de conjuntos
aglutinados y densos constituyendo conglomerados que en algunos casos presentan una evidente
distribución espacial en mitades o bipartita. Los sitios exhiben espacios domésticos con diferentes
tipos de estructuras, algunos con notables concentraciones de molienda, los cuales comprometen
cocinas-dormitorios, patios interiores, recintos sin techar a modo de ramadas y otros grandes a
manera de plazas, corrales u otros espacios de uso comunal.3 Afuera, junto y dentro de ellos se
disponen sepulturas y depósitos —silos o trojas—, formalmente correspondientes a «cistas» y
estructuras tipo chullpa; del mismo modo que tumbas «...adosadas a bloque rocoso», «adosadas y
bajo bloque rocoso» y «bajo bloque rocoso»4 (Fig. 5), acercándose a la composición constructiva
de los asentamientos del Loa superior (Aldunate y Castro 1981; Castro et al. 1991; Adán 1996; Ayala
2000).
Por más que el emplazamiento sea más o menos plano, los sitios se ubican siempre en
quebradas o asociadas a éstas, lo que exigió la aplicación de técnicas constructivas de terrazas y
muros de contención, hasta el día de hoy comunes en las quebradas (Fig. 6 a), lo que equivale a la
arquitectura habitacional con los sistemas de cultivo (Adán 1999). Los muros son principalmente
simples, rústicos y a plomo, pero también aparecen los dobles, sedimentarios y desaplomados. Las
piedras con las que se edificaron las viviendas fueron unidas con argamasa de barro y, en su
mayoría, se encuentran en estado natural, aunque con un alto porcentaje de piezas desbastadas y
toscamente canteadas.
Además de estar asociados a extensos campos de cultivo y elaborados sistemas de regadío
en las quebradas o valles del río, otra característica relevante de estos poblados es su conexión con
rutas de tránsito, seguramente de origen preincaico, que conectan los oasis con la región del Loa
superior y, más lejos aún, con las regiones de Bolivia y Argentina, hacia el norte y oriente. Estas
rutas son reforzadas luego por la política expansiva del Tawantinsuyo, como lo evidencian los
tambos y camino incaicos que se ubican en ambos extremos del Salar, como Catarpe Este y Peine, los
dos anexados a los poblados locales (Lynch 1977; Niemeyer y Rivera 1983; Lynch y Núñez 1994).
En definitiva, la localización, el emplazamiento, la visibilidad y relación de los poblados con
los recursos hídricos, agrícolas y viales sugieren una idea de control y defensa, razón por la que
tradicionalmente estas construcciones han sido llamadas pukara. Algunos de ellos como Catarpe y
Quítor, por su ubicación que aprovecha y controla las mejores aguas y tierras del Salar en el sector
más septentrional de San Pedro, debieron generar notables relaciones de dependencia respecto a las
comunidades o ayllus más meridionales. Sugerentemente, Latcham (1938: 96-98) propone que, adosado
a uno de los riscos del Pukara de Quítor, se levantó el edificio «más grande y más perfecto» que, en
opinión del autor, habría correspondido a un «cacique» o autoridad, lo cual sugiere la existencia de
una población internamente diferenciada, muy poco igualitaria, que pudo estar jerarquizada de acuerdo
a esta dependencia de los ayllus según su acceso al agua.
A lo anterior se suma que la disposición de estos asentamientos en zonas de quebradas y la
significativa ocurrencia de estructuras tipo chullpa en ellos, señalan la configuración de una tradición
de «tierras altas» equivalente a la del curso superior del río Loa (Uribe y Adán 1995). Por lo mismo,
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
Catarpe E
Catarpe W
Quítor
Zápar
Peine
163
177
201
141
210
Superficie (m )
12.000
10.080
17.400
8.400
16.000
Densidad (n.º rec./hec.)
135,83
163,89
115,52
167
131,25
Número de recintos
2
1a. Sitios estudiados, número de recintos, superficie total y densidad ocupacional
Catarpe E
Forma de recintos
Circular
Cuadrangular
Elipsoidal
Irregular
Oval
Rectangular
Subcircular
Subcuadrangular
Subrectangular
Subtriangular
Trapezoidal
n
0
16
0
13
0
56
3
10
52
0
2
%
0
9,82
0
7,98
0
34,36
1,84
6,13
31,90
0
1,23
Catarpe W
Quítor
%
n
%
n
%
n
%
7
3,48
2
1,42
1
0,48
18
8,96
10
7,09
8
3,81
3
1,49
0
0,00
1
0,48
45
22,39
46
32,62
57
27,14
6
2,99
3
2,13
13
6,19
17
8,46
18
12,77
30
14,29
24
11,94
13
9,22
28
13,33
1
0,50
3
2,13
8
3,81
53
26,37
32
22,70
48
22,86
5
2,49
1
0,71
8
3,81
0
0,00
1
0,71
0
0,00
12
8,51
210
3,81
141
100
100
100
n
1
13
0
39
1
31
8
9
54
0
1
0,56
7,34
0
22,03
0,56
17,51
4,52
5,08
30,51
0,00
0,56
No observable
11
6,75
20
11,30
22
10,95
Total
163
100
177
100
201
100
Zápar
Peine
1b. Forma de planta de los recintos y su frecuencia en los sitios.
Catarpe E
Catarpe W
Quítor
Zápar
Peine
Rango de tamaño de los
recintos (m 2)
n
%
n
%
n
%
n
%
n
%
hasta 5
49
30,06
51
28,81
38
18,91
58
41,13
102
48,6
entre 5-10
40
24,54
33
18,64
25
12,44
23
16,31
33
15,7
entre 10-20
38
23,31
37
20,90
47
23,38
13
9,22
26
12,4
entre 20-40
21
12,88
34
19,21
54
26,87
14
9,93
22
10,5
entre 40-60
1
0,61
3
1,69
17
8,46
6
4,26
10
4,76
más de 60
8
4,91
7
3,95
7
3,48
19
13,48
7
3,33
Sin registro
6
3,68
12
6,78
13
6,47
8
5,67
10
4,76
Total
163
100
177
100
201
100
141
100
210
100
1c. Frecuencia del tamaño de los recintos y su frecuencia en los sitios.
Tabla 1. Principales características arquitectónicas de sitios habitacionales del Periodo Tardío de San Pedro
de Atacama
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311
a
b
Fig. 5. a. Aldea de Zápar. Cistas funerarias asociadas a estructuras habitacionales; b. Pucara de Catarpe
Oeste. Sepulturas adosadas y bajo bloque rocoso.
la ocupación de estos espacios ecológicos, las quebradas, no sólo resalta por la mera incorporación
de nuevos rasgos arquitectónicos, sino porque redunda en la adopción de un sistema de tecnologías,
conocimientos y poder asociados que hacen habitables y productivos los territorios altos previamente
desestimados o subutilizados por las poblaciones de los oasis (Adán y Uribe 1995). En este sentido,
una singularidad en la ocupación «quebradeña» de San Pedro es la coexistencia de estructuras tipo
chullpa con entierros en cistas al interior de las viviendas, lo que indica una combinación de
elementos culturales propios del Salar, en tanto una ancestral tradición del desierto y otros rasgos
de la llamada tradición Altiplánica (Schiappacasse et al. 1989). Tal situación expresaría una
convivencia de tradiciones culturales que caracterizaría a las poblaciones del Salar a partir de esos
momentos que, de acuerdo a los fechados absolutos con que se cuenta hoy día para San Pedro, se
iniciaría hacia el 1340 d.C. y perdura hasta el contacto con el Inka (Uribe e.p.).
Al mismo tiempo, estas clases de tumbas y estructuras, cistas y tipo chullpa, al ser aéreas,
exhiben dos cualidades que las antiguas fosas y cementerios atacameños no poseían (cf. Costa y
Llagostera 1992). En primer lugar son visibles, por lo tanto, opera una presencia física determinada
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
en parte por la percepción o contacto visual constante que puede establecerse entre vivos y muertos. Por otra parte, este tipo de sepulturas permiten una eventual manipulación de los restos mortales (Montt 2002). La inclusión de estas prácticas innovadoras es de suma relevancia social, porque
los rituales relativos a la muerte ofrecen a los vivos la eficacia simbólica de percibir visualmente y
movilizar a sus difuntos (Isbell 1997). Por consiguiente, los muertos debieron constituir un significante
explícito, sobre todo porque se ha visto que en esos tiempos la posesión y usufructo de la tierra y el
agua parece haber sido crucial, lo que se encontraría legitimado ante las fuerzas productivas que
componen la población local y ante otros, a través de la noción de ancestro, cuyo valor pudo radicar
en su origen altiplánico. Se integra así un segundo elemento de distinción entre las poblaciones de
San Pedro, las cuales no sólo se diferencian por su ubicación respecto a los recursos críticos locales
—y, de acuerdo a ello, a un territorio específico— sino también por su vínculo con lo externo, en
especial, lo altiplánico (Uribe 1996; Ayala 2000).
Frente a todo esto, Catarpe Este y los tambos de Licancábur, Peine y otros manifiestan la
imposición de la arquitectura del Tawantinsuyo en la región de San Pedro de Atacama (Niemeyer y
Schiappacasse 1988). Sin embargo, Catarpe Este es paradigmático en términos de las manifestaciones
de un dominio e incorporación efectiva de las poblaciones del Salar al funcionamiento estatal. El
asentamiento, si bien comparte varias características con el resto de los sitios habitacionales de la
región (v.g., el trazado ortogonal), evidencia la presencia de una importante plaza doble en una
ubicación central (Fig. 6 b), una frecuencia similar de unidades domésticas y de silos o collcas, así
como una abundancia relativa de muros dobles. La arquitectura de Catarpe Este indica, por lo tanto,
la presencia de abundantes actividades de almacenaje, la disposición de espacios habitacionales sin
sepulturas y, sobre todo, la existencia de grandes espacios de utilidad para funciones administrativas
como ceremoniales. Su construcción se remontaría según los fechados radiocarbónicos a 1510 d.C.5
(Uribe e.p.), indicando una lógica de ocupación que obviamente se aleja de la que se ha señalado
para los poblados locales y que parece incidir en el funcionamiento de la estructura económica y
social de sus habitantes.
Cerámica. Cerca de 53.000 fragmentos de cerámica, recolectados sistemáticamente a partir de un
muestreo dirigido del 20% de cada sitio considerado en la investigación (Tabla 2), permiten detectar
variaciones tipológicas y funcionales en los recintos analizados. Estas diferencias tienen dos lecturas,
una temporal y otra funcional, las que se articulan conformando un panorama complejo y dinámico
de los periodos Intermedio Tardío y Tardío.
A partir de un análisis tipológico y funcional de la cerámica, para lo cual se usó con fines
comparativos y cuantitativos, la cantidad y el peso de los fragmentos provenientes de cada recinto
y asentamiento, se pudo establecer una secuencia ocupacional de los yacimientos estudiados. De
acuerdo a esto, Catarpe Oeste y Quítor aparecen relacionados por una tradición alfarera local (Figs.
7, 8), la cual se remonta a tiempos previos y a los antiguos espacios de los oasis del Salar como los
campos de Sólor (v.g., a través de la presencia predominante de tipos cerámicos conocidos como
Dupont y Aiquina, o de otros más esporádicos como el San Pedro rojo violáceo [Uribe 1997]). Sin
embargo, estos mismos sitios presentarían un momento posterior, más «clásico» del desarrollo
regional y, justamente, serían los que reciben el impacto incaico, elocuente —como se ha visto— en
Catarpe Este. Lo anterior significa que no se reconocen los ejemplares más tempranos o éstos
pierden relevancia frente a una diversidad tipológica mayor (Uribe 1996), en la cual es posible
reconocer exponentes «exóticos» como la cerámica altiplánica Hedionda (Fig. 9). De este modo, los
tipos foráneos se incrementan desde estos momentos y remiten a vínculos con el Altiplano meridional, la vertiente oriental circunpuneña e incluso los Valles Occidentales, expresándose especialmente con el Inka, el cual exhibe un carácter marcadamente trasandino (Fig. 10 d, e).
Debido a que presentan esta misma situación y porque, en la práctica, no existen evidencias
cerámicas más tempranas en ellos, se incluyen Zápar y Peine como asentamientos ocupados en
forma contemporánea a Quítor y Catarpe Oeste. No obstante, éstos exhiben la introducción de una
alfarería cuyas características se relacionarían con el mayor impacto del Tawantinsuyo en la zona, lo
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313
a
b
Fig. 6. a. Pucara de Quítor. Aterrazamientos para construcción de estructuras habitacionales en ladera; b.
Tambo de Catarpe E. Plaza rectangular doble en el centro del asentamiento (Cortesía: F. Maldonado).
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
Tipos cerámicos (%)
Sitios
Pre Intermedio
Tardío
Intermedio
Tardío
Tardío
Foráneos
Hispano-indígena
Guatin
10,96
86,73
1,82
0,20
0,00
Sólor
10,45
88,78
0,31
0,22
0,00
Vilama Norte
13,69
82,89
2,69
0,00
0,00
Vilama Sur
2,33
95,35
1,55
0,00
0,00
Catarpe Este
0,93
93,39
4,23
1,32
0,00
Catarpe Oeste
0,64
96,62
1,27
0,80
0,56
Quítor
0,06
98,50
0,36
0,90
0,02
Zápar
1,19
86,71
11,12
0,49
0,22
Peine
1,97
62,47
34,29
0,52
0,30
n=26.692 fragmentos
Tabla 2. Resumen de la frecuencia de tipos cerámicos de sitios habitacionales tardíos de San Pedro de Atacama
de acuerdo a su adscripción cultural y cronológica.
cual redundaría en la transformación de la cerámica local (Fig. 10 a). Además, aquí la ocupación de
estos asentamientos se extendería hasta épocas coloniales tempranas, marcando un tercer momento
de la secuencia regional coincidente con el Periodo Tardío.
Ahora bien, la cerámica local que se vuelve significativa inmediatamente antes del Inka,
indica, por una parte, el incremento de grandes contenedores para almacenamiento (Uribe y Carrasco
1999). En este sentido, destacan aquellos para líquidos que no sólo pudieron ser utilizados para el
agua, sino también para chicha, lo cual es especialmente relevante en situaciones ceremoniales o
festivas (Fig. 8).
Por otro lado, aumentan los contenedores asociados a la manipulación y preparación de
alimentos, incluida su cocción, lo que otra vez se vincula con actividades festivas que habrían
involucrado la producción de alimentos, generalmente para muchas personas (Fig. 8 c). Un tercer
aspecto interesante es la aparición de cerámica local revestida y roja de uso ceremonial en recintos habitacionales y basurales junto con el resto del material doméstico (Fig. 8 b). Lo anterior
indica que esas prácticas ceremoniales y festivas tendrían lugar en áreas tanto públicas como en
los espacios particulares de las unidades domésticas, generando correlatos materiales sumamente
similares.
En suma, el tipo de asentamiento inferido a partir de la anterior asignación funcional de sus
construcciones indica que todos los sitios inmediatamente preincaicos como Catarpe, Quítor, Zápar
y Peine tendrían un carácter eminentemente doméstico, con recintos habitacionales donde se preparan
y sirven alimentos, siendo sumamente importante el almacenaje. No obstante, también ocurren
actividades ceremoniales, las cuales difieren tanto en escala como en intensidad, pues en dichos
momentos esta práctica pareciera incentivarse en cada uno de los yacimientos mencionados y
realizarse de manera independiente de los otros. Tal situación es coherente con la inclusión en todos
ellos de prácticas funerarias manifestadas en la arquitectura que se incorpora dentro, junto o cerca
de los asentamientos, sugiriendo una dinámica de comunidad o ayllu para las sociedades que
habitaron estos poblados.
Así, dentro de la escala temporal se observa un primer momento del Periodo Intermedio
Tardío caracterizado por sitios habitacionales en los que se encuentran representadas una amplia
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315
Fig. 7. Tipos cerámicos locales de inicios del periodo Intermedio Tardío. a. Tipo Dupont (escudilla); b. tipo
San Pedro rojo violáceo (cántaro doble cuerpo); c. Tipo Turi rojo burdo (cántaro o tinaja); d. Tipo Turi gris
alisado (olla con protúberos).
gama de actividades, aunque no todos los aspectos del ciclo de vida de las personas. Lo anterior se
restringió a muy pocos lugares, los que actuarían como asentamiento-núcleo, tal cual se ha descrito
más arriba para los Campos de Sólor. En cambio, un segundo momento, representado en los sitios
Catarpe, Quítor, Zápar y Peine, corresponde a asentamientos mayores y aglutinados que integran
todo un espectro de actividades: habitacionales, de almacenamiento, ceremoniales y mortuorias en
particular, que los convierte en unidades independientes y autónomas, sin distinguirse uno o unos
pocos núcleos como antes.
Seguramente, este cambio se habría gestado en función de los trastornos económicos,
sociales, políticos y demográficos experimentados por las poblaciones locales después del colapso
tiwanaku, mostrando un énfasis en la segmentación social, extensión territorial, en las conductas de
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
Fig. 8. Tipos cerámicos locales clásicos del Periodo Intermedio Tardío. a. Tipo Aiquina (escudilla); b.
Tipo Turi rojo revestido alisado (cántaro); c. Tipo Turi gris alisado (olla con asa); d. Turi rojo alisado
(cántaro).
almacenamiento, las relaciones con sus fronteras y un cambio en la naturaleza y escala de lo ceremonial y/o público (cf. Albarracín-Jordán 1996). En este ámbito, el desarrollo de una organización
jerarquizada, la generación de excedentes y el rol de los líderes o autoridades en festividades
redistributivas debieron jugar un papel protagónico al interior de los grupos de San Pedro de Atacama,
muy ligado a la competencia sobre el acceso a los recursos de agua y tierra como a los contactos a
larga distancia para la complementariedad de recursos.
Material lítico. Siguiendo el mismo procedimiento de recuperación y la perspectiva analítica con
que se abordó la cerámica, se estudiaron 4080 piezas líticas y 1216 artefactos de molienda, producto
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317
Fig. 9. Tipos cerámicos altiplánicos del Periodo Intermedio Tardío. a, b. Tipo Hedionda negro sobre ante
(escudilla y cántaro miniatura); c, d. Tipo Yavi-Chichas (cántaro y balde).
de la recolección superficial en los yacimientos como del registro in situ de los mismos (Carrasco
2002). Considerando esta división en talla y molienda del material lítico, se llegó a establecer un
comportamiento común o patrón dominado por desechos secundarios e implementos para moler
(Tabla 3). En este contexto, la talla siempre se presenta más abundante en momentos tempranos y de
manera muy escasa en los sitios tardíos, a pesar incluso de evidencias de loza y vidrio empleados en
la elaboración de chaquiras en épocas indígena-coloniales.
Se observa que los sitios con menor material tallado exhiben una alta proporción de instrumentos de molienda (Fig. 11 a), con la excepción de Sólor que habría sido un importante centro
aglutinante de actividades en momentos tempranos como anteriores al desarrollo regional, tal cual lo
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
Fig. 10. Tipos cerámicos inkas locales y foráneos del Periodo Tardío. a. Tipo Lasana café-rojizo revestido
pulido (aríbalo pasta con mica), b. Tipo Turi rojo revestido, exterior negro, alisado interior (jarro «aribaloide»),
c. Tipo Turi rojo revestido pulido ambas caras (escudilla ornitomorfa); d, e. Tipo Yavi-La Paya (aríbalos).
afirma la arquitectura y la cerámica. En la misma época, otros sitios, como Vilama, muestran una talla
más expeditiva y relacionada con situaciones defensivas, quizás también aplicables a Quítor, vinculadas a la competencia por las cabeceras de los principales cursos que desaguan en el salar y que se
manifiesta en concentraciones de piedras «huevillo» para usar como proyectiles (Fig. 11 b).
EL DOMINIO DEL INKA, IDENTIDAD LOCAL Y COMPLEJIDAD SOCIAL EN ATACAMA...
Categorías
Guatin
Sólor Vilama Vilama Catarpe Catarpe Quítor
Norte
Sur
Este
Oeste
319
Zápar
Peine
Total
Adornos
0
18
0
0
2
0
2
3
14
39
Contenedores
0
1
0
0
0
0
0
0
0
0
Herramientas
30
30
2
0
2
6
4
2
9
5
Molienda
35
251
0
3
66
105
0
423
157
1.040
Talla
1.381
1.041
197
9
254
277
92
74
567
3.892
Polifuncionales
0
81
0
1
1
6
0
57
7
153
Indeterminados
3
13
0
0
0
4
0
9
0
29
Material temprano
6
44
0
0
1
0
1
0
5
57
Total
1.455
1.479
199
13
326
398
99
568
759
5215
Tabla 3: Resumen de las principales categorías líticas morfofuncionales de sitios habitacionales tardíos de
San Pedro de Atacama.
Junto con este carácter expeditivo adquirido por la tecnología lítica, es importante destacar
que durante la parte tardía del periodo los productos líticos enfatizan la molienda, tanto agrícola
como con otros fines (v.g., ceremoniales, especialmente ritos funerarios o minería), desplegando una
gran diversidad morfológica y polifuncionalidad de los artefactos referidos a dicha actividad. De
este modo, es posible apreciar que junto con lo funcional, el uso de los artefactos líticos o de la
piedra en general comienzan a adquirir un carácter versátil y simbólico que se expresa en la challa de
materiales molidos en tumbas y chullpas, así como también en la reutilización y polifuncionalidad de
muchas herramientas asociadas a esos materiales, en especial la molienda. Así, se configura, una
conducta común para el resto del área, es decir, en conjunto con el río Loa, donde es bastante equivalente la manera en que se producen los artefactos líticos, con un énfasis en la molienda y el procesamiento de minerales con connotaciones simbólicas, como la malaquita (Uribe y Carrasco 1999).
De acuerdo con lo mencionado, esto es coherente con la concentración de estructuras tipo
chullpa y sepulturas al interior de los poblados en Catarpe, Quítor y sobre todo en Zápar y Peine, lo
cual es a su vez coincidente con las apreciaciones sobre el material cerámico y su énfasis en el
almacenamiento y la actividad ceremonial. Al respecto, el material lítico confirma la necesidad de
procesar y acopiar ciertos alimentos, los que seguramente se emplearon en las festividades comunales, pero, asimismo, en el mantenimiento de otras labores como una minería asociada a la extracción de recursos de valor simbólico (Núñez 1999). Dentro de esta lógica, resulta considerable la
producción de cuentas de malaquita y calcedonia, aparte de las de concha, que se suman al ámbito
de la lítica y que se registran de manera generalizada en todos los asentamientos estudiados,
aunque en proporciones modestas.
Esto, sin duda, potenciaría el intercambio y caravaneo característico de la región con el
propósito de ampliar la complementariedad de recursos a través de contactos a larga distancia. Para
ello, en esos momentos las autoridades de cada comunidad debieron liderar su promoción y organización, ya que debido a la segmentación territorial predominante cada unidad lo llevaría a cabo
independientemente y no de forma tan centralizada en San Pedro, como parece haber ocurrido con
actividades semejantes durante el Periodo Medio (Llagostera 1996).
Restos de fauna y vegetales. Bajo la misma óptica con que se trabajaron los materiales cerámicos y
líticos se recolectaron y analizaron restos óseos y vegetales de los asentamientos descritos (Tabla
4), los cuales fueron identificados por especies e inicialmente asignados a labores vinculadas con la
preparación, manipulación y almacenamiento de alimentos, además de actividades funerarias
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MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
a
b
Fig. 11. a. Aldea de Zápar. Concentración de instrumentos de molienda asociados a estructuras habitacionales;
b. Pucara de Vilama Norte. Concentraciones de piedras asociadas a muro perimetral, posiblemente usadas
como proyectiles.
(González 2002). No obstante, muchos también pudieron ser el resultado de otras actividades, como
las pastoriles, y de esa manera haber ingresado al contexto arqueológico perdurando, más o menos,
gracias a las condiciones de conservación específicas de cada uno de los yacimientos.
De este modo, se recuperaron restos vegetales, entre los cuales predominan el chañar, maíz
y algarrobo, aparte de ramas y maderas no consideradas dentro de las labores domésticas señaladas. Por su parte, el material óseo se encuentra constituido principalmente por astillas de hueso,
siendo preponderantes los restos de camélidos (Lama sp.), además de fauna local, silvestre e introducida. Comparando cada uno de los sitios estudiados, desde Sólor a Peine, en todos se observa
que la presencia de fauna se mantiene uniforme, mientras que el material vegetal adquiere un notorio
aumento en Catarpe, Quítor, Zápar y Peine.
Al respecto, comparando con la cerámica y el lítico, se sostiene que la segregación observada entre las proporciones de las evidencias óseas y vegetales refleja la existencia de dos modos de
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Materiales orgánicos (%)
Oseo
Vegetal
Sólor
34,9
0,40
Guatin
2,4
0,1
Vilama Norte
8,4
0,2
Vilama Sur
1,2
0,6
Catarpe Este
2,8
3,4
Catarpe Oeste
1,3
8,9
Quítor
4,9
13,8
Zápar
26,0
31,4
Peine
17,6
40,8
n=2275,5g
n=2571g
Sitios
Tabla 4: Resumen de la frecuencia de materiales orgánicos de sitios habitacionales tardíos de San Pedro de
Atacama.
producción distintos en el tiempo. En los momentos más tempranos se trataría de una economía con
énfasis en la ganadería, que sustentaba principalmente actividades de tráfico o caravaneo, como lo
sugiere la clara configuración de un transecto pastoril entre Sólor, Vilama y Guatin, del mismo modo
que la existencia de una industria para fabricación de cuentas en el primero, especialmente a partir de
moluscos, e incluso de metales (cf. Llagostera 1996). Este sistema económico estaría apoyado en un
sistema agrícola comparativamente poco relevante, quizás de autosubsistencia, con mayor énfasis
en la recolección de productos de fácil manejo, si es que no silvestres, como son los frutos de chañar
y algarrobo abundantes en todos los oasis de San Pedro, coherente con el patrón de asentamiento
señalado entre 950 y 1350 d.C.
En cambio, los sitios del segundo grupo, más tardíos, representarían una transformación de
ese sistema ganadero y caravanero, el cual estaría apoyado en una producción agrícola con otra
importancia, más estable y de mayor escala, capaz de sustentar una cantidad de población más
grande y segmentada. Lo anterior confirma y es del todo coherente con las particularidades que
adquieren los asentamientos y las actividades desarrolladas en ellos en términos de sus desechos
cerámicos y líticos, coincidentes con una nueva realidad demográfica y cultural, evidentes en prácticas de almacenamiento y ceremoniales incluidas en los ámbitos habitacionales que constituyen
unidades independientes, aunque relacionadas.
Ajuares y ofrendas. Para acercarse a la población que habría protagonizado estos cambios, se
estudiaron 90 cráneos de los contextos funerarios de Yaye, Sólor y Catarpe con el propósito de
caracterizar biológicamente a los habitantes tardíos del Salar de Atacama, evaluándolos según su
condición de salud (edad, sexo, patologías, etc.), a partir de atributos cráneo-faciales, mandibulares,
dentales y deformación cefálica (Reyes 2002; Uribe et al. e.p.). De acuerdo a este estudio preliminar
—y a pesar del grave problema de la muestra, debido al trabajo exclusivo con el esqueleto craneal,
incluso incompleto a veces— el conjunto correspondería a una misma población, bastante relacionada
con las tierras altas del río Loa (cf. Ayala et al. 1999), afectada toda ella por las mismas eventualidades
durante su existencia (v.g., alta esperanza de vida, pero trabajo duro, muchas enfermedades dentales
y a veces etapas de carencia de alimentos o de baja calidad nutricional que afectan bastante la salud,
entre otros). Sobre esta misma base poblacional, sin embargo, se desarrollarían diferencias internas
importantes, cronológicas o sociales que se manifiestan en ajuares y ofrendas. Pero, además, ciertos
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rasgos óseos —como la deformación craneana que se registra en la mitad de la muestra de manera
indistinta entre hombres y mujeres, bastante diversa, aunque casi siempre bajo el predominio de la
deformación tabular erecta, seguida de la oblicua y las circulares— también podrían vincularse a la
complejidad sociopolítica de este grupo (cf. Torres-Rouff 2002), aunque todavía se necesita más
estudios para afirmarlo. Por lo mismo, se enfocará el comportamiento de los textiles y los objetos
misceláneos de otras materias primas que componen sus ajuares y ofrendas.
Material textil. Estudios textiles desarrollados con anterioridad distinguen, a partir del Periodo
Intermedio Tardío, un comportamiento estilístico particular del territorio comprendido por el desierto
de Atacama, el cual se desenvuelve en la configuración de una identidad que transforma los parámetros
del periodo anterior e incorpora aquellos de sus zonas de frontera, fortaleciendo su propio carácter
y, quizá con ello, su acceso a otros espacios y recursos (Agüero et al. 1997, 1999; Agüero 1998).
Tomando en cuenta esos avances, el trabajo realizado consideró el análisis de tejidos funerarios de San Pedro de Atacama como de cementerios del río Loa y del Noroeste argentino
(específicamente del río Doncellas), a través de un registro sistemático de sus atributos tecnológicos, formales y decorativos (Agüero 2000). De todo este territorio se estudió un total de 946 prendas
de vestir, tocados y bolsas, distinguiéndose dos versiones textiles propias del Salar. Por un lado,
aquella representada en los cementerios que continúan ocupados después del Periodo Medio (Fig.
13), los cuales comparten otros elementos mucho más populares en el río Loa e incluso de origen
foráneo (v.g., Tarapacá). Y, paralela a la anterior, otra versión que consiste en tejidos estructuralmente
idénticos, pero propios de cementerios sin ocupaciones previas y con una mayor continuidad hacia
los momentos tardíos (Fig. 12). Esto significa que este segundo conjunto —formado por túnicas
cuadradas o rectangulares tejidas en tramas múltiples, con bordados laterales en puntada satín,
además de gorros tipo «corona», «saquitos-amuletos» y bolsas hechas en sprang o torzal oblicuo
doble— coexiste con la textilería de mayor tradición en el oasis de San Pedro y que comienza a
desarrollarse en el Loa y sus bordes, junto con aquella que se describe a continuación.
Hacia el 1350 d.C. se aprecia en el Loa (Agüero 1998) y las tierras altas del Noroeste argentino (Boman 1908; Rolandi de Perrot 1973, 1979; Pelissero et al. 1997), la popularización de un grupo
de prendas rectangulares correspondientes a túnicas y bolsas, que utilizan técnicas decorativas de
urdimbres transpuestas con uso de tramas múltiples y tapicería dentada (dovetailed), los cuales
reproducen motivos con forma de «V» o rombos en sucesión vertical (en el caso de la transposición
de urdimbres), así como cuadrados y una característica figura zoomorfa de tres dedos (en el caso de
la tapicería), todos ellos confeccionados con hilados de colores blanco, azul y rojo, al igual que en
las numerosas bolsas elaboradas en faz de urdimbre (Fig. 14). Su distribución geográfica y la utilización de tramas múltiples permiten insertar estas prendas dentro del componente «atacameño» o,
mejor dicho, circunpuneño. Al mismo tiempo, la amplitud geográfica que abarca este conjunto,
sumada a la abundancia de bolsas, sugiere su asociación con la actividad caravanera, lo que abre la
posibilidad de que puedan ser situadas hacia finales del Periodo Intermedio Tardío y que hayan
tenido contacto con el Inka, extendiéndose su circulación hasta Calingasta por el sur (Michieli 1994)
y por el norte hasta Camarones y Arica (Ulloa, comunicación personal 1998).
Coincide con esta situación que ciertos sitios propiamente tardíos de San Pedro de Atacama
exhiben algunas túnicas, bolsas y gorros tipo «corona» como los de dicho conjunto textil, en momentos que parecieran ingresar los atuendos incaicos pertenecientes a figurillas de santuarios de
altura (CIADAM 1978; Agüero 2000). Por lo tanto, aunque el tamaño de la muestra es bastante
exiguo, se ha logrado distinguir un nuevo conjunto textil —el incaico—, que presenta novedosas
combinaciones de técnicas y colores (Fig. 15 a), aún cuando está reservado a un contexto sumamente específico, diferente al del resto de los textiles considerados. De acuerdo a ello, es evidente que
el estudio de los tejidos tardíos se ha enfocado en prendas elitistas, lo cual impide tener información
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Fig. 12. Textiles de Atacama del Periodo Intermedio Tardío, presentes en cementerios sin ocupaciones del
periodo Medio. a. Gorros de piel tipo «corona»; b. «Saquitos-amuletos»; c. Bolsa en torzal oblicuo doble;
d. Túnicas decoradas por bordados laterales en puntada satín.
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Fig. 13. Túnicas de Atacama del Periodo Intermedio Tardío, presentes en cementerios con ocupaciones previas, así como en zonas de frontera, mostrando bordados en puntada satín y listas laterales que incluyen
decoración por urdimbres flotantes y/o complementarias.
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acerca de su comportamiento en otros contextos con los que eventualmente podrían ser comparadas
las prendas locales de San Pedro (Rowe 1978, 1995-1996; Michieli 1990; Dransart 1995). Sin embargo,
la situación anterior se ve compensada en parte por hallazgos en Peine (gorro tipo corona y bolsa),
los cuales se vinculan con la textilería previa y local. En particular, una bolsa presenta urdimbres
transpuestas, decoración y colores que la insertan dentro del estilo tardío propuesto para la cuenca
del Loa y el Noroeste argentino (Agüero et al. 1997, 1999; Agüero 2000). Pero, al mismo tiempo,
muestran una gran estandarización en el uso de los colores y en los patrones decorativos, en cierto
sentido la misma de los tejidos de estatuillas y santuarios de altura, por lo que casi con certeza
pueden ser atribuidos a la presencia del Inka en la zona.
Esto, por otra parte, no sugiere una simple adscripción a los patrones estilísticos incaicos,
porque de hecho estéticamente no parece haber sido así (no es el mismo estilo de los santuarios de
altura), sino más bien se trataría de una consecuencia de la intervención de la industria local por
parte del Tawantinsuyo. Esta intervención significaría una reorganización de la producción textil,
cuyo propósito finalmente se vincularía con los cambios ocurridos por la integración a un aparato
estatal. Al mismo tiempo, en el ámbito del intercambio caravanero implicaría un tráfico distinto,
totalmente controlado por el Inka, perfilando una manufactura de bolsas bien evidenciada en el
Noroeste argentino (Fig. 14 c), casi en serie para un movimiento de recuas a una escala mucho mayor,
observándose a través de este estilo un indudable vínculo entre ambos territorios. Sin duda, el
manejo de los líderes locales debió ser clave para la implementación de esta clase de producción en
cada una de las comunidades comprometidas o reuniendo a sus propios grupos de especialistas.
Tabletas, calabazas y otro objetos misceláneos. Siguiendo un estudio y comportamiento muy similar
al de los tejidos, la observación de 3432 objetos misceláneos de contextos funerarios consistentes
en piezas enteras permite distinguir como primera característica importante: la existencia de una
sólida tradición o continuidad cultural desde los inicios del Periodo Intermedio Tardío en los oasis
de San Pedro de Atacama. Lo anterior se observa principalmente en los tipos de tabletas, tubos,
morteros y espátulas de madera y hueso que componen el característico complejo sicotrópico de la
región. Junto a ellos, además, se encuentran calabazas con y sin decoración pirograbada, cucharas
e instrumentos relacionados con el trabajo agrícola, textil, caravaneo y caza, entre otros. Dicha
tradición habría perpetuado un estilo o, más bien, una «estética» respecto a las clases de objetos
ofrendados, los modos de fabricación de los mismos y, con certeza, los criterios para la selección
como adquisición de productos de intercambio (Catalán 2002).
A inicios del periodo, en los contextos funerarios de San Pedro los ajuares y ofrendas
tienden a mantener ciertas características de la época previa (Costa y Llagostera 1992), pero ya no
tienen el despliegue iconográfico y estético de antes, apreciándose más bien una especie de «simplificación» iconográfica y formal de los objetos, que ha sido mal interpretada como un «empobrecimiento» de la sociedad atacameña (Le Paige 1964; Tarragó 1989). Al contrario, esto significa un
mayor acercamiento a lo cotidiano y una potenciación de la producción de excedentes basado en
una agricultura y ganadería intensivas (Adán y Uribe 1995). Esta característica se ve reforzada con
la incorporación de implementos para labores cotidianas asociadas al trabajo agrícola, textil y la
subsistencia en general, los cuales empiezan a ocupar un importante lugar en el rito mortuorio (cf.
Ayala et al. 1999). Tal simplificación, entonces, radicaría posiblemente en la concepción más cercana
de la convivencia entre los vivos y los muertos dentro de un espacio simbólico que se ha hecho
explícito en la arquitectura ceremonial y para el almacenamiento de los asentamientos más clásicos
del desarrollo regional.
De este modo, los elementos del complejo sicotrópico que se habrían incrementado
profusamente en esta región con la influencia tiwanaku persisten durante los periodos tardíos, pero
no de forma masiva. Las tabletas, especialmente como las de Catarpe, se caracterizan por tener un
recipiente rectangular y mango con extensión plana que puede o no llevar decoración, correspon-
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diente a figuras antropomorfas y zoomorfas, en cuyo caso tienden a representar sólo la cabeza y el
cuello del animal (Fig. 16 g, h). Por su parte, las cucharas son objetos que muestran una alta sensibilidad al cambio morfológico más que al decorativo, ya que en el Periodo Tardío no suelen llevar
manifestaciones de esta clase, la que se restringe a motivos zoomorfos, al combinar trazos naturalistas con otros más esquemáticos o, bien, aserrados o «recortes» a lo largo de los mangos (Fig. 16 f).
Pero el rasgo formal más diagnóstico es que la pala se ubica sobre el nivel del mango, el cual puede
ser recto o levemente divergente hacia el extremo terminal y de pala oval (Figs. 16 e, f). Por último, las
calabazas presentan una evolución clara con relación a sus temáticas iconográficas, distinguiéndose un primer momento en los cementerios de Yaye, donde se observa una decoración simple representada por triángulos dispuestos por fuera alrededor del borde que, al parecer, se trataría de un
diseño local (Fig. 16 a). No obstante, luego se produciría una situación diferente vista en Catarpe,
donde se distinguen motivos directamente importados desde el Noroeste argentino, representados
por triángulos con espirales o volutas que se asemejan a los de la cerámica puneña de Humahuaca
(Fig. 16 b, d).
Todas estas evidencias materiales señalan contactos culturales con los cursos medio e
inferior del río Loa, sobre todo a través de los objetos de madera (Ayala et al. 1999), al mismo tiempo
que, como los textiles, aluden a una relación más profunda con el Noroeste argentino. Además, entre
todas estas regiones se desarrollaron temáticas iconográficas similares, existiendo una tendencia
hacia la humanización de las figuras en los objetos relacionados con el complejo del rapé (Fig. 15 b),
donde los personajes muestran un aspecto humano pero sin los rasgos del felino que se observan
en momentos de la influencia tiwanaku (cf. Torres 1984a, b). Tal situación sugiere que la organización
social y el poder político no estarían basados en un conocimiento y manejo de lo sobrenatural, sino
en el dominio estratégico de prácticas asociadas a las actividades cotidianas mencionadas. Consecuentemente se integraría al río Loa dentro de esta dinámica debido a lazos más estrechos que se
establecen con su curso medio, en particular con las localidades de Lasana y Chiuchiu, ricas en
tierras agrícolas y ganaderas, distinguiéndose motivos decorativos compartidos entre las tabletas y
tubos, a los que se unen los de las calabazas. No obstante, también se perciben ciertas distancias
estilísticas con el Loa superior (cf. Alliende 1981), mientras que en el Loa inferior se aprecian elementos que representan un espacio de contactos con las tradiciones de los Valles Occidentales de Arica
y Tarapacá (cf. Cervellino y Téllez 1980). Por su parte, en estos mismos momentos del Periodo
Intermedio Tardío se integraría una iconografía foránea y más abstracta que la observada en las
calabazas y que incluso llegarían a ser más populares que el complejo sicotrópico. Estos, quizás,
constituirían objetos de importación desde el Noroeste argentino donde son mucho más comunes
(Durán 1976), siendo prácticamente idénticos los diseños y motivos que aparecen en ese territorio,
particularmente los del río Doncellas (Gentile 1990).
En suma, se confirma una situación cultural diferenciada en el tiempo, la cual se vincularía
con la existencia de poblaciones que en una etapa inicial se mantienen dentro de las antiguas
tradiciones materiales, frente a otras más tardías que remiten a elementos nuevos, especialmente
compartidos con el Noroeste argentino y el Altiplano meridional de Bolivia, volviéndose predominantes. Esto implica la presencia de una importante interacción circunpuneña que podría
extenderse, al menos, hasta Tarapacá. Sin duda, una importante complejidad social y política debió
verse representada por esta situación, involucrando diversas negociaciones entre las autoridades
que dirigían estas «empresas», dentro de una ardua competencia por el mejor posicionamiento de
las distintas comunidades. Por lo mismo, las concepciones acerca del poder habrían cambiado como
Fig. 14. (Desplegable en la página siguiente). Textiles de finales del Periodo Intermedio Tardío y del Periodo
Tardío circunpuneño. a. Túnica con decoración por urdimbres transpuestas; b. Túnica con decoración por
tapicería enlazada o dovetailed; c. Bolsas.
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a
b
Fig. 15. a. Textil de estilo foráneo del Periodo Tardío; b. Tableta del complejo sicotrópico con mango de tallado
volumétrico con figura antropomorfa de los periodos Intermedio Tardío y Tardío (Museo Caspana).
parece expresarse en las nuevas manifestaciones iconográficas, en las cuales destacan las figuras
humanas por sobre las imágenes de seres sobrenaturales. Esta complejidad «atacameña» debió
implicar, por otro lado, un impulso de gran escala para el desarrollo experimentado en toda la región
por la producción agrícola y caravanera (cf. Núñez 1992), igualmente explícito en los materiales
funerarios, lo que el Inka supo anexar para su propio provecho, en particular empleando el vínculo
con el Noroeste argentino.
4. Conclusiones: Atacama en tiempos del Inka
A lo largo de este trabajo se han expuesto, en la medida de lo posible, aquellas evidencias y
argumentos que parecen representar mejor la complejidad social, económica, política y étnica que se
MAURICIO URIBE, LEONOR ADÁN Y CAROLINA AGÜERO
328
Fig. 16. Objetos funerarios misceláneos. a. Calabaza pirograba de inicios del periodo Intermedio Tardío; b, c, d. Calabazas pirograbadas de los periodo
Intermedio Tardío y Tardío; e, f. Cucharas con pala sobre el mango de los periodos Intermedio Tardío y Tardío; g, h. Tabletas del complejo sicotrópico con
mango plano y tallado volumétrico de los periodos Intermedio Tardío y Tardío.
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encontraba en pleno desarrollo cuando el Inka tomó contacto con las poblaciones del desierto de
Atacama. En particular, los autores se concentraron en el caso de San Pedro de Atacama, porque
tratándose de una de las localidades que ha servido como punto de referencia para la comprensión
arqueológica de los grupos que habitaron este desierto y sus tierras altas (v.g., Berenguer et al.
1986; Tarragó 1989; Núñez 1992), todavía se conoce muy poco de aquellos momentos. Es un hecho
que la investigación se ha centrado en otros periodos de la prehistoria regional, en la que la
esplendorosa época de relaciones con Tiwanaku ha cegado la mirada hacia los momentos posteriores
y se ha asumido, sin mayor cuestionamiento, una continuidad cultural entre las poblaciones
prehispánicas, coloniales y actuales, la cual casi no ha sido sometida a evaluación arqueológica (cf.
Orellana 1963; Castro et al. 1979; Adán et al. 1995). Por último, se hace hincapié en el estudio de este
momento de convergencia de las poblaciones atacameñas e «incaicas» dentro de San Pedro, porque
la experiencia obtenida en otras localidades de este territorio —específicamente Caspana— ha
permitido vislumbrar en ese encuentro los elementos que particularizaron a la complejidad social
local durante el Periodo Intermedio Tardío.
Retomando ese conocimiento (Uribe et al. 1999; Adán y Uribe e.p.; Uribe y Adán e.p.), se
sabe que después de 1450 d.C., en el Loa superior las poblaciones locales y el Inka dialogan a través
de la arquitectura, probablemente porque se trata de un dominio significativo para ambos actores
sociales, demostrando cierta «tensión social» (Uribe y Adán e.p.). Esto, dado que, por un lado el
Inka —o lo foráneo— construyen para dominar a través de su monumentalidad, estandarización y
visibilidad, donde se exhibe y sobrerepresenta (Adán 1999), conectado por una notable red vial
vinculada a recursos mineros y agrícolas, con importantes manifestaciones de arte rupestre. Sin
duda, se trata de una jerarquía administrativa, funcional y productiva que, en este caso, relaciona
sitios de distinta envergadura que se denominaron instalaciones, instaurando la soberanía de agentes
externos.
En este contexto, lo local, por su parte, está libre de la monumentalidad y notoriedad del
dominio inkaico (ibid.); no obstante, las labores productivas muestran una importante intervención
a través de una expresión inka local (cerámica) en todas las clases de contextos funcionales que
abarcan ampliamente cada uno de los asentamientos como los estudiados —consumo de alimentos
y líquidos, molienda, almacenamiento y elaboración de artesanías— al mismo tiempo que en ellos se
incrementan los elementos foráneos y exóticos, en especial del Noroeste argentino (Uribe 1999;
Uribe y Carrasco 1999). El Inka, con relación a la producción lítica, concentra en sus instalaciones las
evidencias de la explotación de mineral y su procesamiento a través de martillos y cobre triturado
como la malaquita, pero dentro de una práctica ya conocida localmente, implementada de manera
ceremonial y, tal cual se vio, realizada en los mismos poblados.
En su expansión, por lo tanto, el Tawantinsuyo habría impulsado el desarrollo de las fuerzas
productivas a través de una fuerza de trabajo numerosa previa, apoyada en el crecimiento demográfico
local y de sus medios de producción, sobre todo agrícolas, cuya mayor fuerza de trabajo requirió de
una organización social más compleja. Eso mismo obligaría al control de esas fuerzas, a modo de una
burocracia administrativa o un grupo encargado de dirigirla desde la actividad doméstica a la estatal,
modificando las relaciones de propiedad objetiva y, consecuentemente, la división social del trabajo.
Las poblaciones locales de Atacama, entonces, desarrollaron aún más sus fuerzas productivas y
potenciaron la generación de excedentes impulsando una jerarquía social que conllevaba a un sistema
políticamente cada vez menos igualitario (Uribe y Adán e.p.).
De este modo, tal cual se aprecia en Catarpe Este, con el Inka son más claros los espacios
públicos como expresión del aparato administrativo, que demuestran el control a través del cual se
dirige la fuerza de trabajo hacia la producción estatal, distinguiéndose un grupo productor de bienes
y alimentos, de otro especializado en las actividades político-ideológicas y administrativas expuestos
en ciertos cementerios (Ayala et al. 1999), y las cuales tampoco faltan en el Salar (v.g., Hostería de
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San Pedro). En definitiva, el Tawantinsuyo se apropia de la fuerza de trabajo y la producción de las
unidades domésticas como un excedente que es manejado por los encargados de los segmentos ya
diferenciados de la misma población local, que ahora se convierten en administradores, incluso
políticos e ideológicos. Así, paralelamente, se genera un amplio control de la conciencia social a
través de la participación ideológica en la formación económica-social del imperio (Uribe y Adán
e.p.). De esta manera, lo inka local mantendría la cohesión y el dominio sin violencia real. Ello por
medio de referentes sociales significativos como las fiestas de redistribución dirigidas por jefes
locales, las que jugaron un papel crucial como lo evidencia el incremento de contenedores de líquidos
y contextos de preparación de alimentos y almacenamiento que se venía dando de antes,
convirtiéndose en la trama religiosa, social y estética que sustenta y reproduce al sistema inkaico.
En suma, antes de la aparición del Inka, en el desierto de Atacama existían sociedades
complejas, segmentadas e internamente jerarquizadas; con el Tawantinsuyo se agudiza la división
social y la propiedad particular, dando como resultado sistemas sociales andinos originales, pero
desiguales, que emplean prácticas sociales arraigadas como la reciprocidad y la redistribución,
generando estructuras verticales al interior de las sociedades locales, entre éstas como las del Loa
y San Pedro, y con las poblaciones del núcleo estatal más cercano como el Noroeste argentino. En
este contexto, el carácter «sagrado» del aparato imperial acompañaría y encubriría las contradicciones
anteriores, dejando que cada entidad actuara dentro de una aparente autonomía (segmentación),
bajo la necesidad de complementariedad económica y en una armonía social, tal cual lo sugiere la
presencia del Inka que articula el sistema en los bordes o periferias más que en los centros. En este
marco, el Inka pareciera imponer un nuevo orden en la geografía y paisaje de las poblaciones locales,
lo cual pareciera ser explícito en otras materialidades no abordadas en esta oportunidad, como la
vialidad y el arte rupestre. Se sabe que instauran lugares especiales, quizás a modo de huacas, que
al estar cerca de las áreas productivas como campos de cultivo, pastoreo y minas las incluyen
dentro de su propiedad (Vilches y Uribe 1999; Sepúlveda 2002). Una clara evidencia material de este
comportamiento lo constituye un ushnu entre el Loa y San Pedro —Cerro Verde—, que pareciera
articular la unidad «atacameña» preexistente y consolidada por el Inka, la misma que en tiempos
coloniales se arrogaría a una misma deidad y, en este sentido, a un solo orden: «...el idolo Llamado
sotar condi a quien todos los yndios de estas Prouincias teníamos Por dios el qual idolo estaua
bestido de cumbe con supillo y plumas en el de oro y Pajaro flamenco...» (Castro 1997: A-30, fol. 40r).
A estas deidades los «...indios cassados y cassas de familias por uia de mita [daban] carneros de la
tierra cuyes chicha plumas de pajaros flamencos cossas yerbas de olor Para saumerios del demonio
y los que no eran cassados ni cabeças de cassas yban a asistir a los dichos sacrificios» (ibid.: A-30,
fol. 39r).
Agradecimientos
Agradecemos en primer lugar a los doctores Peter Kaulicke, Ian Farrington y Gary Urton por
permitirnos participar en el IV Simposio de Arqueología PUCP; al Fondo Nacional de Ciencia y
Tecnología (FONDECYT) que financió esta investigación; al Instituto de Investigaciones Arqueológicas y al Museo R. P. Gustavo Le Paige S. J. de San Pedro de Atacama, por su apoyo constante y,
asimismo, a las comunidades indígenas «atacameñas» de San Pedro, por su comprensión con este
trabajo. Este trabajo no habría sido posible sin la colaboración del equipo conformado por amigos y
colegas: Patricia Ayala, Carlos Carrasco, Bárbara Cases, Claudia del Fierro, Josefina González, Indira
Montt, Viviana Manríquez, Sergio Morales, Omar Reyes y Lorena Sanhueza. Además, les damos
gracias a todos aquellos que participaron responsable y desinteresadamente en las distintas campañas
de campo. Asimismo, nuestro reconocimiento a Paulina Chávez, por sus excelentes ilustraciones.
EL DOMINIO DEL INKA, IDENTIDAD LOCAL Y COMPLEJIDAD SOCIAL EN ATACAMA...
331
Notas
1
Este artículo es resultado del Proyecto FONDECYT 1000148, denominado Historia cultural y
materialidad en la arqueología de los periodos Intermedio Tardío y Tardío de San Pedro de Atacama
y su relación con la cuenca del río Loa.
2
Archivo General de Indias (1557), Patronato, legajo 188, n.° 4.
3
Categorías funcionales para arquitectura tardía han sido propuestas por Adán (1995, 1996).
4
Tipología propuesta por Ayala (1998 y 2000).
5
UCTL-1432 (1590 ± 40 d.C.), UCTL-1433 (1540 ± 30 d.C.) y UCTL-1434 (1570 ± 40 d.C.).
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Direct serogrouping of Dichelobacter nodosus from Victorian farms using conventional multiplex polymerase chain reaction
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Best et al. BMC Res Notes (2018) 11:108
https://doi.org/10.1186/s13104-018-3229-5 Best et al. BMC Res Notes (2018) 11:108
https://doi.org/10.1186/s13104-018-3229-5 BMC Research Notes © The Author(s) 2018. This article is distributed under the terms of the Creative Commons Attribution 4.0 International License
(http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium,
provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license,
and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/
publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Abstract Objective: Dichelobacter nodosus is the causative agent of footrot in sheep. Ovine footrot is a major problem in
Australia that results in large economic losses and a represents a very significant animal welfare issue. D. nodosus is
divided into 10 serogroups (A–I, M), based on sequence variation in the type IV fimbriae gene, fimA. Control of the
bacteria is possible through use of serogroup-specific vaccination, however traditional identification of the sero-
groups of D. nodosus on infected sheep is time-consuming and costly. With the aim of reducing time and cost, a PCR
assay was used to identify serogroups of D. nodosus directly from foot swabs of infected sheep in Victoria. Results: It was shown that serogroup B was most common (10 locations), followed by A, G and H (4 locations), I and
C (2 locations), D, E and F (1 location). Infections with multiple serotypes were observed in 50% of farms, with the
remaining 50% having only a single serogroup detected. The ability to identify serogroups quickly and cheaply direct
from foot swabs will aid the understanding of the epidemiology of D. nodosus and support control programs. Keywords: Footrot Serogroup PCR Keywords: Footrot, Serogroup, PCR from the host. Vaccination with serogroup-specific FimA
antigen provides a protective response, however, cross
protection between serogroups does not occur, and mul-
tiple serogroup infections have been reported to be com-
mon [4]. Understanding what D. nodosus serogroups
are on a particular farm can support better epidemio-
logical investigations, control and eradication programs
[5]. Currently, the testing for serotype of D. nodosus is
time-consuming and labour intensive. It requires bacte-
rial isolation, culturing and slide agglutination testing to
determine the serogroup. Slide agglutination requires the
production antisera specific for each serogroup, which is
beyond most diagnostic laboratories capacity. Direct serogrouping of Dichelobacter
nodosus from Victorian farms using
conventional multiplex polymerase chain
reaction Nickala Best1, Jacek Gwozdz2, Robert Suter3, Grant Rawlin2 and Travis Beddoe1* *Correspondence: t.beddoe@latrobe.edu.au
1 Department of Animal, Plant and Soil Science and Centre
for AgriBioscience (AgriBio), La Trobe University, Bundoora, Melbourne,
VIC, Australia
Full list of author information is available at the end of the article Introduction Ovine footrot is a bacterial infection in sheep, with viru-
lent infections showing severe lesions in the hoof and
resulting in lameness [1]. Footrot infections cost the
sheep meat and wool industry $45 million per year in
Australia [2]. The causative agent, Dichelobacter nodo-
sus, is a genetically diverse, gram-negative anaerobe. D. nodosus is classified into different serogroups based on
the sequence of the fimbrial A gene (fimA). In Australia,
the bacteria are classed into 10 serogroups (A–I, M), with
serogroups A–I being fully sequenced, while the Austral-
ian serogroup M has been partially sequenced [3]. Each
serogroup elicits a specific and strong immune response Recently, a multiplex PCR was developed to deter-
mine the serogroup of D. nodosus from bacterial colo-
nies grown on media [6]. This molecular assay has been
shown to be highly specific and reduces time in identify-
ing the serogroup of D. nodosus. The ability to determine
D. nodosus serogroup directly from foot swabs would aid Best et al. BMC Res Notes (2018) 11:108 Page 2 of 6 particle handling system (Thermo Fisher Scientific, Fin-
land), as per manufacturer’s instructions. epidemiological investigations of footrot in Australia. Here, we have identified the serogroups of D. nodosus
from foot swabs collected on Victorian farms using this
multiplex PCR assay. qPCR for detection of D. nodosus
Detection of D. nodosus was performed by quantita-
tive polymerase chain reaction (qPCR) as described by
Stäuble et al. [7]. Briefly, the qPCR program consisted of
an initial denaturation step of 2 min at 50 °C, a hold of
10 min at 95 °C, followed by 40 cycles of 15 s at 95 °C,
1 min at 60 °C. The AgPath-IDTM One-Step RT-PCR Kit
(Ambion, Austin, USA) was used as master mix accord-
ing to manufacturer’s instructions, with final concentra-
tions of 300 nM primers, 100 nM DnAprTM- vMGB,
250 nM DnAprTM-bMGB and 5 μL of template DNA
used. Primers and probes were synthesised and supplied
by Applied Biosystems (California, USA). Reactions were
carried out and analysed using the 7500 Fast Real-Time
PCR System (Life Technologies), with a set threshold of
0.05. Results are reported as cycling threshold (Ct) val-
ues, the point at which the sample signal exceeds the
threshold of 0.05. Materials and methods
Field collectionh The clinical samples were obtained retrospectively from
the Victorian Government Veterinary Diagnostic unit
collection. The original samples were submitted by Victo-
rian District Veterinary Officers, Animal Health Officers
and private veterinary practitioners from the interdigital
skin of lame sheep for routine diagnostic testing. Samples
(n = 137) were collected from 16 farms across 15 Victo-
rian locations. These farms were diagnosed as clinically
virulent, benign or healthy during a disease surveillance
survey from November 2014 to August 2015 (Fig. 1). The
interdigital skin was swabbed and the cotton head of the
swab transported at 4 °C in phosphate buffer saline (PBS)
(8.1 mM Na2HPO4, 137 mM NaCl, 1.4 mM KH2HPO4
and 2.6 mM KCl) with an addition of 20 mM ethylen-
ediaminetetraacetic acid (EDTA). Total nucleic acid was
extracted using the MagMAX™ − 96 Viral RNA isolation
kit (Ambion, Austin, USA) and Kingfisher-96 magnetic Results Foot swab samples were collected from farms that have
been known previously to have clinical forms of footrot,
or were clinically healthy yet had D. nodosus detected
on animals. These samples and the collection method
were used due to availability, as they were concurrently
used for additional studies. Genomic DNA was extracted
from all samples and the presence of D. nodosus in all 137
sheep were confirmed by qPCR as described by Stäuble
et al. [7]. Initial experiments to determine the serotype
of D. nodosus by PCR from DNA extracted from foot
swabs was unsuccessful, however the addition of extra
magnesium to the PCR reaction was able to achieve
amplification of PCR products for the determination of
serogroups present on different farms (Fig. 2). The repre-
sentative PCR products were verified by DNA sequenc-
ing as shown in Additional file 1: Figure S1. These results give a basic overview of the epidemiology
of D. nodosus on farms throughout Victoria, which previ-
ously has not been known. The ability to serotype directly
from a swab greatly reduces the time and cost tradition-
ally required to serotype field samples. This information
and additional virulence testing may help in both under-
standing and control of the disease in the future. i
Overall the result revealed that 87 (63%) of the sheep
tested resulted in a serogroup being determined by mul-
tiplex PCR using DNA directly extracted from a foot
swab sample (Table 1). From the animals that showed a
serogroup band, 74% of the infected sheep had a single
serogroup detected, while 26% had infections from mul-
tiple serogroups, ranging from 2 to 4 serogroups detected
(Table 1). Serogroup B was the most commonly detected
serogroup as shown in Additional file 2: Figure S2), found
in 48% of infections, on 56% of farms and in 60% of loca-
tions in Victoria. Serogroup B was also involved in 91% of Discussion We have modified an existing protocol by increasing the
concentration of MgCl2, for faster and more cost-effec-
tive identification of D. nodosus serogroup determination
by PCR, by analysing DNA extracted from foot swabs. This has allowed us to investigate the epidemiology of D. nodosus infection on farms throughout Victoria. It was
found that on the sampled farms, there was a high level
of detection of serogroup B. This is in agreement with a
previous study that showed serogroup B was most com-
monly detected [8]. Additionally, it was also found that
serogroup B was most commonly found in multiple
infections. Additional sampling across more locations is needed
to gain an overall insight into the prevalence of differ-
ent serogroups in Victoria, with this method providing
a starting point for a higher throughput study. Currently,
Victorian diagnostics laboratories are not equipped to
isolate or grow D. nodosus or produce the antisera that
is required for the traditional slide agglutination tests to
determine serogroup. The further improvement of the
methods here would allow for molecular diagnostics
based laboratories to access this information. A more
affordable and accessible method of serotyping would aid
in epidemiological studies in Victoria and contribute to
the improved control of the disease.h Serotyping by PCR The following amplification con-
ditions consisted of an initial denaturation step of 4 min
at 95 °C, followed by 5 cycles of 30 s at 94 °C, 30 s at 60 °C,
and 30 s at 72 °C, followed by 25 cycles of 30 s at 94 °C,
30 s at 58 °C, 30 s at 72 °C for 30 s and then a final elon-
gation of 4 min at 72 °C. Amplicon size was checked by
agarose electrophoresis migration by loading directly on
2% (w/v) MetaPhor® Agarose (Lonza, USA) gel, prepared
as per manufacturer’s instructions, with the addition of
0.5 µL of Sybr® Safe DNA Gel Stain (Life Technologies). The gel was run at 100 V for 55 min and imaged using the
GelDoc™ XR+ (BioRad) instrument and software. Sanger
DNA sequencing of representative amplicons was per-
formed at Australian Genomics Research Facility (AGRF)
to verify amplified serogroups. the farms with multiple serogroup infections. Serogroups
A, G and H were the next most common serogroups,
found in 15, 25 and 12% of infections respectively. All
were found in 4 locations in both single and multiple
infections. Serogroup C and I were found in 2 farms in 2
separate locations, with D, E and F present in 1 location
only. Serotyping by PCR Multiplex PCR for determination of serogroup was per-
formed as described by Dhungyel et al. with the following Fig. 1 Victorian farms sampled. The locations of the 16 Victorian farms sampled and number of serogroups detected at each location. The figure
was generated in Tableau 6.1 Fig. 1 Victorian farms sampled. The locations of the 16 Victorian farms sampled and number of serogroups detected at each location. The figure
was generated in Tableau 6.1 Best et al. BMC Res Notes (2018) 11:108 Page 3 of 6 modifications [6]. GoTaq® Green Master Mix 2X (Pro-
mega, Madison, USA) was used for amplification as per
manufacturer’s instructions, with the addition of 1 µL
100 mM MgSO4 to increase the Mg+ concentration to
5.5 mM. Reactions were carried out in a final volume of
25 µL and amplified in a C-Master GT thermal cycler
(Dynamica, Australia). The following amplification con-
ditions consisted of an initial denaturation step of 4 min
at 95 °C, followed by 5 cycles of 30 s at 94 °C, 30 s at 60 °C,
and 30 s at 72 °C, followed by 25 cycles of 30 s at 94 °C,
30 s at 58 °C, 30 s at 72 °C for 30 s and then a final elon-
gation of 4 min at 72 °C. Amplicon size was checked by
agarose electrophoresis migration by loading directly on
2% (w/v) MetaPhor® Agarose (Lonza, USA) gel, prepared
as per manufacturer’s instructions, with the addition of
0.5 µL of Sybr® Safe DNA Gel Stain (Life Technologies). The gel was run at 100 V for 55 min and imaged using the
GelDoc™ XR+ (BioRad) instrument and software. Sanger
DNA sequencing of representative amplicons was per-
formed at Australian Genomics Research Facility (AGRF)
to verify amplified serogroups. modifications [6]. GoTaq® Green Master Mix 2X (Pro-
mega, Madison, USA) was used for amplification as per
manufacturer’s instructions, with the addition of 1 µL
100 mM MgSO4 to increase the Mg+ concentration to
5.5 mM. Reactions were carried out in a final volume of
25 µL and amplified in a C-Master GT thermal cycler
(Dynamica, Australia). Limitations It should be noted that not all samples tested returned a
visible serogroup PCR band, and these will require further
investigation. Experimentally, it may be possible bands
were present yet were not detected using the current
method. Additional detection methods were not inves-
tigated. There is also the possibility that undetermined Best et al. BMC Res Notes (2018) 11:108 Page 4 of 6 l
b l
t
M
thi
t i
l d d
i
l ti
lt
i
d lid
l ti
ti
d
t th l
k
Fig. 2 Identification of serogroups from foot swab samples by PCR. Samples from various farms were amplified in serogroups mixtures (ABC, DEF,
and GHI) and analyzed on a 2% MetaPhor agarose gel. Banding is shown that corresponds to serogroup I (a and b), serogroups A, B, C and H (c, d
and e), serogroups B, D and E (f, g and h) and serogroup F (i). The serogroup A and C positive control with no amplification seen in the negative, no
template (NT), control Fig. 2 Identification of serogroups from foot swab samples by PCR. Samples from various farms were amplified in serogroups mixtures (ABC, DEF,
and GHI) and analyzed on a 2% MetaPhor agarose gel. Banding is shown that corresponds to serogroup I (a and b), serogroups A, B, C and H (c, d
and e), serogroups B, D and E (f, g and h) and serogroup F (i). The serogroup A and C positive control with no amplification seen in the negative, no
template (NT), control isolation, culturing and slide agglutination due to the lack
of suitable primers for identification of serogroup M by
PCR. Therefore, it is imperative to develop primers suit-
able for detection of serogroup M by PCR to understand
the prevalence of serogroup M in Victorian sheep flocks. samples belong to serogroup M as this was not included
in the PCR. There is limited information on the prevalence
of serogroup M in Australian isolates, however, in two
studies the numbers ranged from 5 to 40% of flocks [3, 9]. Currently, serogroup M identification requires bacterial Best et al. BMC Res Notes (2018) 11:108 Page 5 of 6 Table 1 Summary of serogroups found on farms across different locations in Victoria
The animals tested were all confirmed to be D. Publisher’s Note Publisher s Note
Springer Nature remains neutral with regard to jurisdictional claims in pub-
lished maps and institutional affiliations. Springer Nature remains neutral with regard to jurisdictional claims in pub-
lished maps and institutional affiliations. Funding NB was responsible for the experimental work, data analysis, and writing of
the manuscript. JG contributed to the writing of the manuscript and reviewed
the drafts. RS, GR and TB coordinated the project, contributed to the experi-
mental design, and reviewed the drafts. All authors read and approved the
final manuscript. Funding
The Victorian Sheep and Goat Compensation Fund provided funding for this
project. The Victorian Sheep and Goat Compensation Fund provided funding for this
project. Limitations nodosus positive via qPCR
Location
Farms
Animals tested
(n)
Animals with serogroup
detected (n)
Serogroup (times detected)
Animals
with multiple
serogroups (n)
Avenel
1
10
6
G (6)
0
Bendoc
1
10
4
G (4), H (2)
2
Carboor
1
10
9
A (7), B (3), C (4), H (5)
8
Cavendish
1
10
9
B (1), G (8)
5
Giffard
1
1
1
C (1)
0
Hamilton
2
20
15
B (8), G (3), F (7)
3
Kernot
1
2
2
A (2), H (1)
1
Kialla East
1
10
5
I (5)
0
Kilmore East
1
10
4
A (1), B (2), H (2)
1
Lindenow South
1
10
1
I (1)
0
Murroon
1
9
6
B (1), D (4), E (4)
3
Newstead
1
10
8
A (3), B (5)
0
Rutherglen
1
9
6
B (6)
0
Strathbogie
1
6
1
B (1)
0
Tooborac
1
10
10
B (10)
0
Total
16
137
87
23 Table 1 Summary of serogroups found on farms across different locations in Victoria The animals tested were all confirmed to be D. nodosus positive via qPCR VIC, Australia. 3 Department of Economic Development, Jobs, Transport
and Resources, Victorian Government, Attwood, Melbourne, VIC, Australia. Acknowledgements Additional file 1: Figure S1. PCR amplicons of different serogroups
subjected to Sanger sequencing. Blastn alignment of serogroup G (panel
A), I (panel B), B (panel C, E, F and G) and F (panel D) amplicon sequence
obtained from Sanger sequencing from various farms. Accession number
is shown in brackets. g
We would like to thank the participating sheep owners for their time and
assistance during sample collection, and the Victorian District Veterinary Offic-
ers and Animal Health Officers for sample collection. Abbreviations
PCR
i
i qPCR: quantitative polymerase chain reaction; EDTA: ethylenediaminetet-
raacetic acid; DNA: deoxyribonucleic acid; PBS: phosphate buffer saline. Availability of data and materials The information supporting the conclusions of this article is included in the
article. Competing interests p
g
The author(s) declared no potential competing interest with respect to the
research, authorship, and/or publication of this article. Additional file 2: Figure S2. PCR amplicons of different serogroups
subjected to Sanger sequencing. Blastn alignment of serogroup G (panel
A), I (panel B), B (panel C, E, F and G) and F (panel D) amplicon sequence
obtained from Sanger sequencing from various farms. Accession number
is shown in brackets. 6.
Dhungyel OP, Whittington RJ, Egerton JR. Serogroup specific single and
multiplex PCR with pre-enrichment culture and immuno-magnetic bead
capture for identifying strains of D. nodosus in sheep with footrot prior to
vaccination. Mol Cell Probes. 2002;16(4):285–96.
7.
Stauble A, Steiner A, Frey J, Kuhnert P. Simultaneous detection and
discrimination of virulent and benign Dichelobacter nodosus in sheep of
flocks affected by foot rot and in clinically healthy flocks by competitive
real-time PCR. J Clin Microbiol. 2014;52(4):1228–31.
8.
Claxton PD, Ribeiro LA, Egerton JR. Classification of Bacteroides nodosus
by agglutination tests. Aust Vet J. 1983;60(11):331–4.
9.
Chetwin DH, Whitehead LC, Thorley SE. The recognition and prevalence
of Bacteroides nodosus serotype M in Australia and New Zealand. Aust
Vet J. 1991;68(4):154–5. Author details
1 1 Department of Animal, Plant and Soil Science and Centre for AgriBioscience
(AgriBio), La Trobe University, Bundoora, Melbourne, VIC, Australia. 2 Depart-
ment of Economic Development, Jobs, Transport and Resources Centre
for AgriBioscience (AgriBio), Victorian Government, Bundoora, Melbourne, Received: 8 December 2017 Accepted: 2 February 2018 Best et al. BMC Res Notes (2018) 11:108 Page 6 of 6 References
1. Kennan RM, Han X, Porter CJ, Rood JI. The pathogenesis of ovine footrot. Vet Microbiol. 2011;153(1–2):59–66. 2. Lane J, Jubb T, Shephard R, Webb-Ware J, Fordyce G: Final Report
B.AHE.0010: Priority list of endemic diseases for the red meat industries. In: Edited by Australia. MaL; 2015. 3. Dhungyel O, Schiller N, Whittington R. Identification and characterization
of serogroup M Dichelobacter nodosus from sheep with virulent footrot. Vet Microbiol. 2015;176(3–4):378–81. 4. Dhungyel OP, Lehmann DR, Whittington RJ. Pilot trials in Australia
on eradication of footrot by flock specific vaccination. Vet Microbiol. 2008;132(3–4):364–71. 5. Dhungyel O, Hunter J, Whittington R. Footrot vaccines and vaccination. Vaccine. 2014;32(26):3139–46. 6.
Dhungyel OP, Whittington RJ, Egerton JR. Serogroup specific single and
multiplex PCR with pre-enrichment culture and immuno-magnetic bead
capture for identifying strains of D. nodosus in sheep with footrot prior to
vaccination. Mol Cell Probes. 2002;16(4):285–96. References References
1. Kennan RM, Han X, Porter CJ, Rood JI. The pathogenesis of ovine footrot. Vet Microbiol. 2011;153(1–2):59–66. 2. Lane J, Jubb T, Shephard R, Webb-Ware J, Fordyce G: Final Report
B.AHE.0010: Priority list of endemic diseases for the red meat industries. In: Edited by Australia. MaL; 2015. 3. Dhungyel O, Schiller N, Whittington R. Identification and characterization
of serogroup M Dichelobacter nodosus from sheep with virulent footrot. Vet Microbiol. 2015;176(3–4):378–81. 4. Dhungyel OP, Lehmann DR, Whittington RJ. Pilot trials in Australia
on eradication of footrot by flock specific vaccination. Vet Microbiol. 2008;132(3–4):364–71. 5. Dhungyel O, Hunter J, Whittington R. Footrot vaccines and vaccination. Vaccine. 2014;32(26):3139–46. • We accept pre-submission inquiries
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Stauble A, Steiner A, Frey J, Kuhnert P. Simultaneous detection and
discrimination of virulent and benign Dichelobacter nodosus in sheep of
flocks affected by foot rot and in clinically healthy flocks by competitive
real-time PCR. J Clin Microbiol. 2014;52(4):1228–31. y
9.
Chetwin DH, Whitehead LC, Thorley SE. The recognition and prevalence
of Bacteroides nodosus serotype M in Australia and New Zealand. Aust
Vet J. 1991;68(4):154–5. 8.
Claxton PD, Ribeiro LA, Egerton JR. Classification of Bacteroides nodosus
by agglutination tests. Aust Vet J. 1983;60(11):331–4. 5.
Dhungyel O, Hunter J, Whittington R. Footrot vaccines and vaccination.
Vaccine. 2014;32(26):3139–46. Submit your next manuscript to BioMed Central
and we will help you at every step: • We accept pre-submission inquiries
• Our selector tool helps you to find the most relevant journal
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https://openalex.org/W4362676277
|
https://eprints.soton.ac.uk/476760/2/1_s2.0_S2590136223000402_main.pdf
|
English
| null |
Trust and vaccine hesitancy during the COVID-19 pandemic: A cross-national analysis
|
Vaccine. X
| 2,023
|
cc-by
| 8,016
| ERROR: type should be string, got "https://doi.org/10.1016/j.jvacx.2023.100299\n2590-1362/\u0001 2023 The Author(s). Published by Elsevier Ltd.\nThis is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). ⇑Corresponding author.\nE-mail address: w.j.jennings@soton.ac.uk (W. Jennings). a r t i c l e\ni n f o Article history:\nReceived 21 November 2022\nReceived in revised form 31 March 2023\nAccepted 1 April 2023\nAvailable online 6 April 2023\nKeywords:\nCOVID-19\nVaccination\nTrust\nMisinformation Article history:\nReceived 21 November 2022\nReceived in revised form 31 March 2023\nAccepted 1 April 2023\nAvailable online 6 April 2023 Previous studies of vaccine hesitancy in the context of COVID-19 have reported mixed results in terms of\nthe role played by political and institutional trust. This study addresses this ambiguity with a global anal-\nysis of the relationship between trust and vaccine hesitancy, disentangling the effects of generalized trust\norientations, trust in specific institutions and conspiracy mentality. It first draws upon a cross-national\nsurvey of 113 countries to demonstrate that trust in government is a predictor of vaccine hesitancy across\nglobal regions. It further draws on original surveys fielded in seven countries (France, Germany, Spain,\nArgentina, Croatia, Brazil, India), which deploy a diverse range of measures, to disentangle the\nindividual-level predictors of vaccine hesitancy. Our findings confirm the robust effects of trust in gov-\nernment across countries, but when including other trust measures in the same models, the most robust\neffects are those of trust in health institutions and conspiracy mentality. Weaker associations are\nobserved for right-wing ideology and online political engagement, while the consumption of traditional\nmedia tends to predict the willingness of individuals to be vaccinated. Keywords:\nCOVID-19\nVaccination\nTrust\nMisinformation Keywords:\nCOVID-19\nVaccination\nTrust\nMisinformation \u0001 2023 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY license\n(http://creativecommons.org/licenses/by/4.0/). The global battle against COVID-19 has drawn attention to the\nperceived critical importance of political trust for public health\ninterventions designed to mitigate the spread of infectious disease\n[3,4,8,9]. This perception seems well-founded: citizens’ trust in\nauthorities and social institutions speaks to whether they believe\nthat they are actually looking out for their interests and has thus\nlong been understood as fundamental to cooperation between gov-\nernment and the public [8,13]. Indeed, trust has been found to be\nimportantly related to countries’ general success in combating\nthe pandemic on the aggregate level [3,30] and with citizens’ com-\npliance with public health guidelines on the individual level\n[12,29,27,29]. they report non-significant effects for trust in government in all\nbut one of their samples. On the other hand, Lazarus et al. Trust and vaccine hesitancy during the COVID-19 pandemic: A cross-\nnational analysis Will Jennings a,⇑, Viktor Valgarðsson a, Lawrence McKay a, Gerry Stoker a, Eduardo Mello b,\nHasan Muhammad Baniamin c a University of Southampton, UK\nb Fundação Getulio Vargas, Brazil\nc North South University, Bangladesh Vaccine: X 14 (2023) 100299 Vaccine: X 14 (2023) 100299 https://doi.org/10.1016/j.jvacx.2023.100299\n2590-1362/\u0001 2023 The Author(s). Published by Elsevier Ltd.\nThis is an open access article under the CC BY license (http://c Original survey data For the second part of the analysis, we use data from online\nnationally representative surveys in seven major countries –\nFrance, Germany, Spain, Croatia, Argentina, Brazil and India –\nwhich we conducted between September and December 2021. Each of our surveys apply quotas on multiple demographic factors,\nwhich include age, gender, and region for all surveys (and addition-\nally social class for NetQuest surveys and education and past vote\nfor YouGov surveys). Full details of the survey fieldwork, sample\nsize, survey company and mode/design are reported in Table 1. The dependent variable is vaccine willingness, the inverse of vac-\ncine hesitancy. At the time of fieldwork, the distribution of free\nvaccines was becoming a reality rather than (as with the data in\nour previous analysis) a theoretical future possibility, even in\ncountries where the rate of doses had been stalled by poor avail-\nability (such as in the case of India, which may be reflected in its\nlow level of first doses and full vaccinations received), so the mea-\nsures reflect actual reported prior behaviour, rather than reported\nvaccine intentions. In the survey, we asked respondents the follow-\ning pair of questions, coding as vaccine ‘willing’ (1) those respon-\ndents who answered that they had received a dose or that they\nwould if offered. Those who had declined the dose offered, or\nwould not get a future dose if offered, are coded as unwilling\n(0).2 The questions were formulated as follows: p\nOur empirical findings confirm that political trust is a consistent\npredictor of vaccine hesitancy across the world, one that is robust\nto the inclusion of multiple controls and holds in over 90 percent of\nthe more than 100 countries covered here. In our more detailed\nanalysis of the role of trust, using original survey data, we find that\ngeneralised distrust in government is positively related to vaccine\nhesitancy and generalised trust is negatively related to it, whereas\nthe effects of political mistrust and social trust are non-significant,\nbut conspiracy mentality has the strongest effect by some margin. When separating political trust into trust in different institutions,\nwe find, in line with previous studies, that trust in health institu-\ntions stands out as by far the most robust, but this is fairly strongly\nrelated to confidence in political institutions, which has a signifi-\ncant effect when the former is not included in the same model. Methods lumped together in statistical models, without great clarity about\nthe reasons behind the choice of variables, the correlations\nbetween them or how the measures overlap. In this study, we\nmove\nbeyond\nthese\nprevious\napproaches\nby\ndistinguishing\nbetween generalized trust orientations and more specific confi-\ndence in particular institutions, by distinguishing between political\ntrust, mistrust, distrust and conspiracy mentality as distinct types of\nattitudes within the ‘‘trust” family, and by explicitly interrogating\nthe inter-relationships between different types of trust and vaccine\nacceptance. Finally, our original surveys contain a broader range of\nrelevant measures of personal, attitudinal and behavioural factors\nthan most others, allowing us to compare the effects of trust with\nfactors such as media use, personality traits and demographics to a\ngreater extent than prior studies. Wellcome global monitor data For the global analysis, we use data from The Wellcome Global\nMonitor (WGM) included in the Gallup World Poll. Between Octo-\nber 2020 and January 2021 inclusive, telephone surveys were com-\npleted by nearly 120,000 respondents in 113 countries, which\ncollectively represent 90 % of the global population aged 15 +. The surveys concentrated on themes related to COVID-19, includ-\ning its economic impacts, and crucially featured questions on both\nvaccine uptake intentions and on trust. The measures used here\nasked respondents: (a) how much they trusted the national gov-\nernment in their country, and (b) whether they would agree to\nbe vaccinated if a vaccine to prevent coronavirus ‘‘was available\nright now at no cost”.1 Our approach in this paper thus seeks to advance our under-\nstanding of the role(s) of trust in vaccine acceptance by addressing\nthese limitations of prior studies. Firstly, we undertake a global\nanalysis of the degree to which trust predicts vaccine hesitancy,\nbased on high quality survey data from over 100 countries. This\nenables us to determine how general the link between trust and\nvaccine hesitancy is, covering a broader scope of countries than\nany prior study on the individual level. Secondly, we report results\nfrom our original surveys conducted in seven countries, which\nincluded an extensive set of measures which allow us to disentan-\ngle the relationship between different types of trust and vaccine\nacceptance and compare with the role of related factors such as\npersonality traits, cognition and political ideology. 1 The full wording for these questions is included in Online Appendix A.\n2 Those who give ‘prefer not to say’ responses at Q13, or respond ‘don’t know’ at\nQ14, are dropped from the analysis. Methods In terms of\ntrust, we include measures of generalized political trust, mistrust\nand distrust, reflecting insights from recent studies that suggest\nthat lack of trust may either denote a type of cautious scepticism\n(mistrust) or settled dissatisfaction (distrust) – and that the two\nmay have different implications for vaccine hesitancy, in that we\nwould expect distrust but not necessarily mistrust to drive it\n([2]; [28]; [12]). In addition, we include a scale measuring ‘conspir-\nacy mentality’ (developed by 5 and more conventional measures of\ntrust in representative institutions, health institutions and the\nmedia as well as a measure of social trust, separating these in our\nanalysis and comparing their relative effects. This offers a more\ncomprehensive test of the precise ways that trust influences beha-\nviour and its relative importance. The WGM also included demographic information on respon-\ndents’ age, sex, education, income and employment status. In\nterms of sample size, 98,166 respondents in 105 countries are valid\non the vaccine and trust measures, with 94,620 in 102 countries\nvalid once control variables are added in a regression model. a r t i c l e\ni n f o In this study, we\nmove\nbeyond\nthese\nprevious\napproaches\nby\ndistinguishing\nbetween generalized trust orientations and more specific confi-\ndence in particular institutions, by distinguishing between political\ntrust, mistrust, distrust and conspiracy mentality as distinct types of\nattitudes within the ‘‘trust” family, and by explicitly interrogating\nthe inter-relationships between different types of trust and vaccine\nacceptance. Finally, our original surveys contain a broader range of\nrelevant measures of personal, attitudinal and behavioural factors\nthan most others, allowing us to compare the effects of trust with\nfactors such as media use, personality traits and demographics to a\ngreater extent than prior studies. a r t i c l e\ni n f o [16] find\na significant effect for trust in government in their 19-country\nstudy and Lindholt et al. [17] find in their 8-country study that\ntrust in government has an effect in addition to trust in scientists\nand health authorities, although that effect is only marginally sig-\nnificant in their full model. Finally, Allington et al. [1] find in their\nstudy of the UK that trust in government had a significant effect\nbut a much smaller one than trust in scientists and medical profes-\nsionals. Other studies on the topic have been limited by their use of\nconvenience samples, although these tend to find significant\neffects for political trust [13,25,25]. However, when it comes to the particular topic of the role of\npolitical trust in shaping vaccine acceptance, studies have reported\nmixed results. There appears to be no doubt that some kind of trust\nmatters, but while Rozek et al. [21] report in their 17-country\nstudy that trust in health institutions matter, they report no effects\nfor trust in political leaders. Similarly, Kerr et al. [14] find in their\n25-sample study that trust in experts plays an important role, but In summary, of the four large, cross-national studies published\nto date on the topic, two report significant effects for political trust\non vaccine acceptance, whereas two report no significant effects. In\nthis study, we directly address this ambiguity in the existing liter-\nature, using secondary survey data from over 100 countries and\noriginal survey data from seven countries, with the latter explicitly\ndesigned to disentangle the role of different types of trust in seven\ncountries. The broad concept of political trust is typically not mea-\nsured in a systematic or theoretically driven way in prior studies;\ninstead, different measures of trust in various entities tend to be https://doi.org/10.1016/j.jvacx.2023.100299\n2590-1362/\u0001 2023 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). Vaccine: X 14 (2023) 100299 Vaccine: X 14 (2023) 100299 W. Jennings, V. Valgarðsson, L. McKay et al. lumped together in statistical models, without great clarity about\nthe reasons behind the choice of variables, the correlations\nbetween them or how the measures overlap. Original survey data Indeed, trust in health institutions, conspiracy mentality and age\nare the only predictors whose significant effects persist even after\nincluding all 29 variables in our analysis in the same model. Con-\nsuming news in traditional media is also a strong (negative) pre-\ndictor of hesitancy and those who identify on the right in politics\nare more likely to be hesitant, whereas the effects of other factors\nare weaker and less consistent. By combining breadth and depth of\nanalysis, we highlight the consistency of trust as a predictor of vac-\ncine hesitancy and shed light on the dynamic role that different\ntypes of trust play, with conspiracy mentality and trust in health\ninstitutions standing out as the most potent predictors. Q13. Which of these applies to you? (1) I have received one or two doses of a COVID-19 vaccine\n(2) I have been offered a vaccine against COVID-19, but declined the\noffer to be vaccinated\n(3) I have not yet been offered a vaccine against COVID-19\n(4) Prefer not to say\nQ14. To what extent, if at all, do you agree, or disagree, with the\nfollowing statements? If I was offered a vaccine for COVID-19, I would get it [ASK IF\nQ13 = 3]\n(1) Strongly agree\n(2) Tend to agree\n(3) Neither agree nor disagree\n(4) Tend to disagree (2) I have been offered a vaccine against COVID-19, but declined the\noffer to be vaccinated (3) Neither agree nor disagree (4) Tend to disagree 2 2 Vaccine: X 14 (2023) 100299 W. Jennings, V. Valgarðsson, L. McKay et al. Table 1\nOriginal survey details. Source: Our World In Data [18]. Country\nFieldwork\nSample\nCompany\nDesign\nVaccination rate*\nFrance\n23.09.2021 – 20.11.2021\n1,548\nYouGov\nOnline, nationally representative\n65 % (75 %)\nGermany\n14.10.2021 – 23.11.2021\n1,558\nYouGov\nOnline, nationally representative\n65 % (70 %)\nSpain\n23.09.2021 – 26.09.2021\n1,022\nYouGov\nOnline, nationally representative\n77 % (79 %)\nCroatia\n29.09.2021 – 06.10.2021\n1,017\nYouGov\nOnline, nationally representative\n40 % (46 %)\nArgentina\n29.09.2021 – 08.10.2021\n1,113\nYouGov\nOnline, nationally representative\n50 % (66 %)\nBrazil\n06.12.2021 – 28.12.2021\n2,123\nNetQuest\nOnline, nationally representative\n64 % (77 %)\nIndia\n22.09.2021 – 30.09.2021\n1,040\nNetQuest\nOnline, nationally representative (of the literate population)\n15 % (44 %)\nTotal\n22.09.2021 – 28.11.2021\n9,421\nNote: France includes a top-up sample of 498 respondents 19.11.2021–20.11.2021; Germany includes top-up sample of 534 respondents 19.11.2021–20.11.2021. 3 In Brazil, this item specifically referred to the publicly funded healthcare system,\n‘‘Sistema Único de Saúde (SUS)”. Global analysis: Wellcome Global Monitor data We begin with an analysis of the relationship between political\ntrust and vaccine hesitancy across the globe. Is this robust, and\ndoes it consistently emerge globally? To answer this, we turn to\nthe Wellcome Global Monitor included in the Gallup World Poll,\nwhich includes measures of trust in government and willingness\nto be vaccinated, along with demographic measures. Further infor-\nmation about this data source and the variables used is presented\nin the methods section below and in Online Appendix B. We first\npresent the country-level aggregate relationships between trust\nin\ngovernment\nand\nvaccine\nintention\non\nthe\ncountry-level\n(Fig. 1).4 The y-axis represents the percentage of respondents in each\ncountry who said that they would agree to be vaccinated against the\ncoronavirus if a vaccine was available at no cost,5 while the x-axis\nrepresents the percentage of respondents who said they trusted\nthe national government in their country ‘a lot’ or had ‘some’ trust\nin it.6 The aggregate correlation between these two variables is pos- Following previous work, we took the first three statements (1–\n3) to indicate trust, the next two (4–5) to reflect mistrust, and the\nlast three (6–8) to measure distrust. For each set of statements, we\ncreated scales for each respondent taking the mean level of agree-\nment with the statements (on the Likert scale) before rescaling the\nvalues to 0–1. Cronbach’s alpha scores vary between 0.60 (mis-\ntrust) to 0.84 (distrust) and 0.90 (trust), all of which pass the 0.6\nthreshold established as acceptable by Nunally and Bernstein [20]. Following previous work, we took the first three statements (1–\n3) to indicate trust, the next two (4–5) to reflect mistrust, and the\nlast three (6–8) to measure distrust. For each set of statements, we\ncreated scales for each respondent taking the mean level of agree-\nment with the statements (on the Likert scale) before rescaling the\nvalues to 0–1. Cronbach’s alpha scores vary between 0.60 (mis-\ntrust) to 0.84 (distrust) and 0.90 (trust), all of which pass the 0.6\nthreshold established as acceptable by Nunally and Bernstein [20]. Turning to our other independent variables, we provide more\ndetail in Online Appendix B, and only summarise our measures\nbriefly here. In terms of demographics, we include age, gender,\nand a dummy for university level education. Our models include\nleft–right ideological self-placement, measured on a 0–10 scale. (5) Strongly disagree\n(6) Don’t know Turning to our independent variables, the survey includes mul-\ntiple measures of trust. First, it includes conventional measures of\nconfidence in institutions, drawn from the World Values Survey,\nfrom which we derive three separate measures of institutional\ntrust: 1) trust in health institutions, which is calculated for each\nrespondent as the mean of trust in the World Health Organisation\n(WHO) and ‘‘the health service in your country”,3 2) trust in the\nmedia, which is the mean of trust in ‘‘the press” and ‘‘TV”, and 3)\ntrust in political institutions, which is calculated as the mean of trust\nin ‘‘parliament”, ‘‘political parties” and ‘‘the government in your\ncountry”. We also include a module developed in previous studies\nto measure generalized orientations of trust, mistrust and distrust\ntowards government [6,14]. The following set of questions was\nasked in all seven countries: To what extent do you agree, or disagree, with the following\nstatements? statements? (1) The government is honest and truthful\n(2) In general, the government usually does the right thing\n(3) The government usually has good intentions\n(4) I am usually cautious about trusting the government\n(5) I am unsure whether to believe the government\n(6) The government acts unfairly towards people like me\n(7) The government usually ignores my community\n(8) The government doesn’t respect people like me\n(1)\nStrongly agree\n(2)\nTend to agree\n(3)\nNeither agree nor disagree\n(4)\nTend to disagree\n(5)\nStrongly disagree\n(6)\nDon’t know 4 Readers should bear in mind the timing of data collection (October 2020 through\nJanuary 2021) and that the level of trust in each country may have increased or\ndecreased since (for instance in Myanmar – ‘MM’ in Fig. 1 – where trust may have\nfallen significantly since the 2021 coup d’etat).\n5 The response options were binary: ‘‘Yes, would agree” and ‘‘No, would not agree”.\nThose who responded ‘‘don‘t know” or refused to respond are coded as missing for the\npurposes of this analysis.\n6 The other response options were ‘‘not much” and ‘‘not at all”. Don’t knows and\nrefusals are once again treated as missing. Results Global analysis: Wellcome Global Monitor data Original survey data *Share of\npeople vaccinated against COVID-19 at start of survey fieldwork: complete initial vaccine protocol (partially vaccinated). Table 1\nOriginal survey details. Source: Our World In Data [18]. 4 Readers should bear in mind the timing of data collection (October 2020 through\nJanuary 2021) and that the level of trust in each country may have increased or\ndecreased since (for instance in Myanmar – ‘MM’ in Fig. 1 – where trust may have\nfallen significantly since the 2021 coup d’etat). Table 1 We further incorporate measures of the ‘Big Five’ personality traits,\nbased on the ‘Short 15-item Big Five Inventory (BFI-S)’ devised and\nvalidated by Lang et al. [15], creating scales for the mean (dis)\nagreement with the statements measuring each of the Big Five. We also include a scale of ‘conspiracy mentality’, designed for\nuse across cultures and languages, based on the 5-item scale by\nBruder et al. [5], albeit with a slightly modified question and\nresponse scale. We take the simple mean of these items as our\nmeasure of conspiracy mentality. Finally, we include a measure\nof cognitive reflection based on Frederick (2005; see also [25]),\nwhich principally measures the ability or disposition of respon-\ndents to reflect on a question and resist reporting the first response\nthat comes to mind (using classic games/experiments from beha-\nvioural economics). We take the sum of correct answers (0–3) to\nbe the respondent’s cognitive reflection score. Finally, we include\nmeasures of media consumption and activity: how much respon-\ndents follow news in different types of media and whether they\nhave recently ‘fact-checked’ articles or posted political content\nonline. All these scales and all other variables used in our analysis\nare rescaled to 0–1 to enable direct comparison of effects. After\nremoving all cases with missingness in the final model, 6,210\nrespondents are retained for the multivariate analysis. (5) Strongly disagree\n(6) Don’t know 6 The other response options were ‘‘not much” and ‘‘not at all”. Don’t knows and\nrefusals are once again treated as missing. 5 The response options were binary: ‘‘Yes, would agree” and ‘‘No, would not agree”.\nThose who responded ‘‘don‘t know” or refused to respond are coded as missing for the\npurposes of this analysis. Global analysis: Wellcome Global Monitor data Turning to our other independent variables, we provide more\ndetail in Online Appendix B, and only summarise our measures\nbriefly here. In terms of demographics, we include age, gender,\nand a dummy for university level education. Our models include\nleft–right ideological self-placement, measured on a 0–10 scale. 3 W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 100299 Fig. 1. Trust in government vs self-reported vaccine intentions (aggregate-level), Wellcome Global Monitor. Countries denoted by Alpha-2 ISO codes, see: https://www. iban.com/country-codes. Fig. 1. Trust in government vs self-reported vaccine intentions (aggregate-level), Wellcome Global Monitor. Countries denoted by Alpha-2 ISO codes, see: https://www. iban com/country-codes Fig. 1. Trust in government vs self-reported vaccine intentions (aggregate-level), Wellcome Global Monitor. Countries denoted by Alpha-2 ISO codes, see: https://www. iban.com/country-codes. orted vaccine intentions (aggregate-level), Wellcome Global Monitor. Countries denoted by Alpha-2 ISO codes, see: https://www levels, which may drive both trust and vaccine intention). The full\nresults from these regression models are presented in Online Appen-\ndix, Table A1, and they show that moving through each stage from\nthe lowest to the highest level of trust in government is associated\nwith a significant increase in vaccine willingness, when controlling\nfor individual and contextual factors. Fig. 3 presents the marginal\neffects of each category of trust from these models, illustrating that\nthe global prediction (holding all control variables constant) for vac-\ncine intention is about 54 % for those who say they trust their\nnational government ‘‘not at all”, whereas it is about 71 % for those\nwho say they trust it ‘‘a lot”. Thus, the global relationship between\npolitical trust and vaccine intention appears to be significant and\nsubstantial even when accounting for demographic and contextual\nfactors. itive and moderate (0.49, p < 0.001). The simple bivariate regression\nline shows that at the lowest observed levels of trust (20 %), fewer\nthan 50 % are expected to take vaccines, while at the highest levels\n(near to 100 %), nearly 80 % would accept a no-cost vaccine. To explore how far this relationship is universal, we run bivari-\nate logistic regression models of vaccine intention on trust within\neach country on the individual level and present the odds ratios for\neach country in Fig. 2 (where trust in government is an ordinal\nvariable that takes the values 1 ‘not at all’, 2 ‘not much’, 3 ‘some’\nand 4 ‘a lot’). 7 Here, we include the full categorical variable with dummies for each category and\n‘‘not at all” as the baseline. Global analysis: Wellcome Global Monitor data While there is significant cross-national heterogene-\nity, it is striking that there is statistically significant (at the 95 per\ncent confidence level) positive relationship between trust in gov-\nernment and vaccine willingness in 75 out of 105 countries. In just\na single country (Bosnia Herzegovina), the relationship between\ntrust and vaccine willingness is significantly negative (i.e. confi-\ndence intervals of the estimated odds ratio do not cross at 1.0). In another 29 countries, the relationship was not statistically sig-\nnificant, interestingly including both high trust countries, such as\nNorway and Sweden, and low trust countries, such as Slovakia,\nSenegal and Serbia. The average odds ratio across countries is equal\nto 1.4, meaning that each one unit increase in the ordinal scale of\ntrust in government typically leads to a 40 % increase in the likeli-\nhood of being willing to be vaccinated. Detailed analysis: original surveys in seven countries While the Wellcome Global Monitor provides impressive global\nscope in terms of the total number of countries and proportion of\nthe global population covered, it is understandably limited in\nhow far it explores alternative measures of trust, as well as other\nrelevant values, beliefs and behaviours which may account for or\novershadow the relationship. We build upon this through fielding\nour own bespoke survey that specifically explored multiple dimen-\nsions of trust and a range of related factors in multiple countries. For our study, we are therefore able to draw on an array of mea-\nsures that distinguish between categories of generalized trust, mis-\ntrust and distrust in government, as well as more conventional\nmeasures of trust in specific institutions, where we differentiate\nbetween political institutions (parliament, political parties and\nthe national government), health institutions (the national health\nservice and WHO) and the media (the press and TV). We also\ninclude various measures of conspiratorial mentality, political ide-\nology, cognitive reflection, personality traits and traditional and\nonline media use, as well as demographic information. This\napproach enables us to explore the degree to which a much wider Turning to a more sophisticated analysis of this relationship, we\nnext run multi-level regression models on the entire dataset (with\ncountry as the second level). These again predict vaccine intent as a\nfunction of trust in government,7 and add several individual-level\ncontrols: age (in four categories), sex, education (a binary variable\nwhere those with university education are coded as equal to 1),\nemployment status and income on the individual level (in quintiles). At the country-level, we include the Human Development Index\n(HDI) score from the UN Human Development Reports (to partly\naccount for the potential confounding effect of societal welfare 4 Fig. 2. Bivariate regression of trust in government on self-reported vaccine willingness, odds ratios (individual-level, by country). Countries denoted by Alpha-3 ISO\nsee: https://www.iban.com/country-codes. W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 1 W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 100299 Fig. 2. Bivariate regression of trust in government on self-reported vaccine willingness, odds ratios (individual-level, by country). Countries denoted by Alpha-3 ISO codes,\nsee: https://www.iban.com/country-codes. Fig. 3. Marginal predictions of vaccine intention by levels of trust in government in the 2020 Wellcome Global Monitor survey, from multi-level regression models controlling\nfor individuals’ demographics and countries’ HDI. W. Jennings, V. Valgarðsson, L. Detailed analysis: original surveys in seven countries However, these measures are moderately correlated (the\npairwise Pearson’s r coefficients are between 0.45 and 0.5) and\nwhen the other institutional trust measures are included without\nthe others but with controls, both are significant predictors, so this\nshould not be interpreted to mean that trust in political or media\ninstitutions does not matter for vaccine willingness. In other\nwords, people’s general trust orientations towards government\nand their trust in political institutions matter for vaccine accep-\ntance, but when accounting for their trust in different specific state\ninstitutions, it turns out that health institutions are the most clo-\nsely relevant institution. We begin by considering the degree to which each of our mea-\nsures in isolation is associated with willingness to be vaccinated\nagainst COVID-19. Bivariate logistic regression models are calcu-\nlated for all predictors for the full, pooled seven-country sample,\nwith the odds ratios plotted in Fig. 4. In the figure, the dark circles\nindicate the coefficient value, while the whiskers indicate the 95\nper cent confidence intervals. The red line indicates the point at\nwhich there is no significant relationship between the variables if\nthose confidence intervals cross. Several measures of trust have statistically significant bivariate\nrelationships with vaccine willingness. The strongest positive rela-\ntionship\nis\nobserved\nfor\ntrust\nin\nhealth\ninstitutions\n(odds\nratio = 16.32, 95 % CI 12.40 to 21.48): in other words, those most\ntrusting were between 12 and 21 times more likely to accept the\nvaccine than those lowest in trust. Trust in the media (OR 4.96,\n95 % CI 3.70 to 6.64) and political institutions (OR 3.72, 95 % CI\n2.78 to 5.01) also have significant, but weaker, bivariate relation-\nships with vaccination willingness. p\ng\nAs expected, generalized trust orientations towards govern-\nment also have a positive effect on willingness (OR: 3.61, 95 % CI\n2.83 to 4.62), whereas distrusting and mistrusting orientations\nhave negative effects with mistrusters (OR: 0.49, 95 % CI 0.38 to\n0.62) and distrusters (OR: 0.52, 95 % CI 0.40 to 0.68) each about half\nas likely. Perhaps surprisingly, the difference between the effects of\ndistrust and mistrust are small, and social trust has weak and non-\nsignificant effects. Besides trust, we observe numerous other sig-\nnificant relationships. Detailed analysis: original surveys in seven countries McKay et al. Vaccine: X 14 (2023) 100299\n5 Fig. 2. Bivariate regression of trust in government on self-reported vaccine willingness, odds ratios (individual-level, by country). Countries denoted by Alpha-3 ISO codes,\nsee: https://www.iban.com/country-codes. Fig. 3. Marginal predictions of vaccine intention by levels of trust in government in the 2020 Wellcome Global Monitor survey, from multi-level regression models controlling\nfor individuals’ demographics and countries’ HDI. Fig. 3. Marginal predictions of vaccine intention by levels of trust in government in the 2020 Wellcome Global Monitor survey, from multi-level regression models controlling\nfor individuals’ demographics and countries’ HDI. 5 W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 100299 set of measures of political attitudes are predictive of vaccine will-\ningness/hesitancy, and to situate different types of trust in that\ncontext. We provide further details on these data and measures\nin the methods section and the full questionnaire in Online Appen-\ndix A. age, and consumption of traditional media. However, we run all\nour block models on the same sample of respondents with non-\nmissing values on all variables (N = 6,235), to enable comparison\nof effects between models. The block models are displayed in Fig. 5, with effects again\nexpressed in terms of odds ratios. The first model includes gener-\nalized trust orientations, in both government and other people,\nand shows that generalized trust retains its positive effect\n(OR = 3.08, 95 % CI 2.08 to 4.56) on vaccine willingness and distrust\nits negative effect (OR = 0.58, 95 % CI 0.38 to 0.87), whereas mis-\ntrust becomes a non-significant predictor. Here, social trust\nbecomes a weakly negative predictor of willingness, albeit not sig-\nnificant at the 95 % level (p = 0.08), when accounting for political\ntrust orientations. In model 2, we include measures of institutional\ntrust, with intriguing results. Here, trust in political institutions\nbecomes non-significant and trust in the media only significant\nat the 90 % level (p = 0.08), but trust in health institutions remains\na strong predictor (OR = 15.37, 95 % CI 10.06 to 23.47) – suggesting\nagain that the latter is the most robust driver of vaccine willing-\nness. Detailed analysis: original surveys in seven countries Most notably, those with most ‘conspiracy\nmentality’ are about 10 % as likely as the least conspiratorial to\nbe willing to be vaccinated (OR = 0.10, 95 % CI 0.07 to 0.14) and\nthe oldest respondents are four times as likely as the youngest\n(OR = 3.98, 95 % CI 2.94 to 5.40). Engagement with different infor-\nmation sources also appears related to vaccine hesitancy: online\ninformation consumption, sharing and fact-checking material\nonline are associated with slightly lower willingness to get vacci-\nnated, though the positive association with consumption of tradi-\ntional media is slightly larger in magnitude. People who identify\nas on the right of the political spectrum are also somewhat less\nlikely to get vaccinated, as are those who score higher on some\nof the Big Five personality trait scales; especially on ‘openness’,\nwhereas cognitive reflection shows no association. Finally, as in\nother studies, university education is related to greater willingness\nto get vaccinated for COVID-19. Finally, the third model shows that conspiracy mentality is an\neven stronger negative predictor (OR = 0.07, 95 % CI 0.04 to 0.10)\nwhen including demographic controls. Because political trust and\nconspiracy mentality are closely related attitudes [10],8 we explore\nwhether trust is a confounder for the effects of conspiracy mentality\nor vice versa, by presenting the results of models including all of\nthese variables together in Online Appendix Fig. A2: these show that\nthe effects of conspiracy mentality survive controlling for all of the\ntrust measures, whereas the only trust measure that still has a sig-\nnificant effect is that of trust in health institutions, suggesting that\nboth shape vaccine willingness above and beyond political trust.9 Returning to other factors, we find that the effects of most per-\nsonality traits lose their significance in the block model (neuroti-\ncism is significant at the 90 % level, p = 0.06) but here, cognitive\nreflection\nshows\na\npositive\neffect\non\nvaccine\nwillingness\n(p < 0.01) and information engagement still has significant effects. In particular, the frequency of consuming traditional media (news-\npapers, television, radio) remains a strong predictor of vaccine\nwillingness (OR = 4.27, 95 % CI 2.89 to 6.32), while getting informa-\ntion from ‘people’ and online has negative effects, although rela-\ntively weak ones (and non-significant for the former). 8 The correlation between the conspiracy mentality scale and the generalized trust\nscales is between 0.22 and 0.28 Pearson’s r in the pooled dataset, with stronger\ncorrelations in France, Germany, and Spain.\n9 Generalized trust in government is still a significant predictor when including\nthose and conspiracy mentality together in a model, but its effects become\ninsignificant when controlling for trust in institutions as well. 8 The correlation between the conspiracy mentality scale and the generalized trust\nscales is between 0.22 and 0.28 Pearson’s r in the pooled dataset, with stronger\ncorrelations in France, Germany, and Spain. 9 Generalized trust in government is still a significant predictor when including\nthose and conspiracy mentality together in a model, but its effects become\ninsignificant when controlling for trust in institutions as well. Detailed analysis: original surveys in seven countries We find\nthat more ‘active’ internet users – those who post political content\n– are more vaccine hesitant (OR = 0.51, 95 % CI 0.37 to 0.69),\nalthough the effect of fact-checking becomes non-significant. We\nalso find that moving from left to right on a left/right scale is asso-\nciated with lower levels of vaccine willingness in all models. The\neffect of university education becomes non-significant in most\nmodels, but age retains a strong positive effect on willingness\n(while the effects of gender are still non-significant). The latter is While informative in a descriptive sense, these bivariate rela-\ntionships may well be confounded by various related demographic\nand attitudinal factors. To get closer to a causal relationship, we\nnext run a series of multivariate regression models, to isolate and\ndisentangle the effects of specific factors. We estimate six separate\n‘block’ models of related variables here, each retaining a selection\nof closely related variables. Each of the block models controls for\ndemographic factors (gender, age, education), left–right ideology\nand country dummies (because the dependent and independent\nvariables are likely to vary by country and the other control vari-\nables, in ways which might result in spurious effects reported in\nthe absence of these controls). In Online Appendix A, we also report\nresults of a single model including all 29 variables. We focus on\nblock models in this discussion, because including so many corre-\nlated variables in one model creates problems of multicollinearity,\nwide confidence intervals and very unintuitive interpretations of\nindividual coefficients 29. We see this model primarily as a conser-\nvative test of which predictors are the most robust of all: these\nturn out to be trust in health institutions, conspiracy mentality, 6 W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 100299 Fig. 4. Bivariate logistic regression of vaccine willingness, odds ratios. Fig. 4. Bivariate logistic regression of vaccine willingness, odds ratios. traits interestingly appear to play a bigger role in those countries\nthan in the others. perhaps unsurprising given the sharp age gradient of the severity\nof COVID-19. Finally, we explore whether the determinants of vaccine hesi-\ntancy (or willingness) differ importantly between the countries\nin our dataset by conducting the bivariate and block models sepa-\nrately for each country. Coefficient plots from these models can be\nfound in the Online Appendix, Figs. A3-A23. Detailed analysis: original surveys in seven countries Most of the central\nfindings remain the same across countries, but generalized trust\norientations have by some margin the strongest relationship with\nvaccine hesitancy in France and Germany, whereas in Brazil polit-\nical trust negatively predicts vaccine willingness. This finding for\nBrazil is consistent with other studies suggesting that partisan cues\ncan reverse expected relationships between political trust and\nCOVID-related attitudes and behaviours (e.g., [12]). President Bol-\nsonaro was notoriously vaccine- and COVID-sceptic, so those\nrespondents expressing trust in his government were more likely\nto also hold anti-vax attitudes. Generally, most variables have\nweaker and often non-significant effects in India and particularly\nArgentina than in the other countries, but conversely, personality Discussion Our analysis here has served to demonstrate the critical link\nbetween political trust and vaccine hesitancy across the globe,\ndemonstrating how trust in government is predictive of willing-\nness to get vaccinated against COVID-19 for much of the world’s\npopulation. Generalized trust orientations towards government\nand trust in political institutions are consistent predictors of vac-\ncine hesitancy, although when including multiple interrelated\npolitical trust measures in the same models, trust in health institu-\ntions and conspiracy mentality stand out as the most robust pre-\ndictors. The design of this study has offered unprecedented\nglobal scope in its consideration of the degree to which trust in\ngovernment predicts vaccine hesitancy for COVID-19 (113 coun-\ntries) and systematic comparison of alternative measures of differ-\nent types of trust as well as other beliefs, opinions and traits, again\nin a substantial cross-national design (7 countries). This allows us 7 7 W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 100299 onclude with increased confidence that trust is an essential\nmedia) are less willing. There is little evidence to support the clai\nFig. 5. Multivariate block models of vaccine willingness, odds ratios. Fig. 5. Multivariate block models of vaccine willingness, odds ratios. Fig. 5. Multivariate block models of vaccine willingness, odds ratios. to conclude with increased confidence that trust is an essential\nresource for societies in their fight against COVID-19, and that this\nis true both relative to other attitudes and controlling for the\ndemographics of individuals. Our findings confirm that willingness\nto be vaccinated against COVID-19 is positively linked to trust in\ngovernment, and negatively linked to distrust of government. Prior\ncross-national studies have reported mixed results in terms of the\neffects of political trust on vaccine hesitancy, but our findings sug-\ngest that this is likely because those that report non-significant\neffects of political trust do so via regression models which simulta-\nneously include various closely related measure of trust [14,24]. In\nterms of other factors, we find that consumers of traditional media\n(i.e. television, newspapers) are more willing to get vaccinated,\nwhereas the politically online (i.e. people who post on social media) are less willing. There is little evidence to support the claim\nthat personality traits are drivers of vaccine hesitancy, even in\nmodels that exclude strong predictors such as conspiracy beliefs\nand political trust. Discussion We find that cognitive reflection is associated\nwith greater vaccine willingness, but this is a small effect relative\nto other attitudinal and demographic predictors. Of the 29 predic-\ntors included in our regression models, political trust orientations\nand conspiracy mentality emerge as the strongest predictors of\nvaccine willingness, along with respondents’ age. to conclude with increased confidence that trust is an essential\nresource for societies in their fight against COVID-19, and that this\nis true both relative to other attitudes and controlling for the\ndemographics of individuals. Our findings confirm that willingness\nto be vaccinated against COVID-19 is positively linked to trust in\ngovernment, and negatively linked to distrust of government. Prior\ncross-national studies have reported mixed results in terms of the\neffects of political trust on vaccine hesitancy, but our findings sug-\ngest that this is likely because those that report non-significant\neffects of political trust do so via regression models which simulta-\nneously include various closely related measure of trust [14,24]. In\nterms of other factors, we find that consumers of traditional media\n(i.e. television, newspapers) are more willing to get vaccinated,\nwhereas the politically online (i.e. people who post on social We have here built on a still growing literature examining the\nlink between trust and vaccine hesitancy in the context of\nCOVID-19. Our study draws upon high quality survey data from\nthe Wellcome Global Monitor, enabling us to cover a substantially\ngreater number of countries than most empirical studies con- 8 W. Jennings, V. Valgarðsson, L. McKay et al. Vaccine: X 14 (2023) 100299 [7] Devine D, Gaskell J, Jennings W, Stoker G. Exploring Trust, Mistrust and\nDistrust (1st Digital Workshop of the ESRC ‘‘TrustGov” Project); 2020. https://\ntrustgov.net/s/TrustGov-Trust-mistrust-distrust-20042020.pdf. ducted during the pandemic, and through our own cross-national\nsurvey programme, allowing us to disentangle the roles of different\ntypes of trust and to directly compare their effects with other com-\nmonly cited factors. While the COVID-19 pandemic has eased in\nseverity by now, these results point to the importance of rebuilding\npolitical trust for the potential success of future mass vaccination\nprogrammes. [8] Devine D, Gaskell J, Jennings W, Stoker G. Trust and the Coronavirus Pandemic:\nWhat are the Consequences of and for Trust? An Early Review of the Literature. Polit\nStud\nRev\n2021;19(2):274–85. https://doi.org/10.1177/\n1478929920948684. [9] Devine D, Valgarðsson V, Smith J, Jennings W, di Vettimo MS, Bunting H,\nMcKay L. References [23] Nunnally JC, Bernstein IH. Psychometric Theory. McGraw-Hill; 1994. [24] Rozek LS, Jones P, Menon A, Hicken A, Apsley S, King EJ. Understanding Vaccine\nHesitancy in the Context of COVID-19: The Role of Trust and Confidence in a\nSeventeen-Country Survey. Int J Public Health 2021;66(May):. https://doi.org/\n10.3389/ijph.2021.636255636255. [1] Allington D, McAndrew S, Moxham-Hall V, Duffy B. Coronavirus conspiracy\nsuspicions, general vaccine attitudes, trust, and coronavirus information\nsource as predictors of vaccine hesitancy among UK residents during the\nCOVID-19\npandemic. Psychol\nMed\n2021. https://doi.org/10.1017/\nS0033291721001434. [25] Schernhammer E, Weitzer J, Laubichler MD, Birmann BM, Bertau M, Zenk L,\net al. Correlates of COVID-19 vaccine hesitancy in Austria: Trust and the\ngovernment. J Public Health 2021;44(1):E106–16. https://doi.org/10.1093/\npubmed/fdab122. [2] Bertsou E. Rethinking political distrust. Eur Polit Sci Rev 2019;11(2):213–30. https://doi.org/10.1017/S1755773919000080. [26] Schmeisser Y, Renström E, Bäck H. Who follows the rules during a crisis? Personality traits and trust as predictors of compliance with containment\nrecommendations during the COVID-19 pandemic in Sweden. Front. Polit. Sci. 2021;3(November):1–13. https://doi.org/10.3389/fpos.2021.739616. [3] Bollyky TJ, Hulland EN, Barber RM, Collins JK, Kiernan S, Moses M, et al. Pandemic preparedness and COVID-19: an exploratory analysis of infection\nand fatality rates, and contextual factors associated with preparedness in 177\ncountries, from Jan 1, 2020, to Sept 30, 2021. Lancet (London, England)\n2022;399(10334):1489–512. https://doi.org/10.1016/S0140-6736(22)00172-\n6. (\n)\np\n//\ng/\n/ p\n[27] Seyd B, Bu F. Perceived risk crowds out trust? Trust and public compliance\nwith coronavirus restrictions over the course of the pandemic. Eur Polit Sci Rev\n2022;14(2):155–70. https://doi.org/10.1017/S1755773922000078. [4] Brezzi M, Gonzalez S, Nguyen D, Prats M. An Updated OECD Framework on\nDrivers of Trust in Public Institutions to Meet Current (OECD Working Papers\non\nPublic\nGovernance,\nIssue\n48);\n2021. https://www.oecd-ilibrary. org/governance/an-updated-oecd-framework-on-drivers-of-trust-in-public-\ninstitutions-to-meet-current-and-future-challenges_b6c5478c-en. ( )\np\n//\ng/\n/\n[28] Stecula DA, Pickup M. Social Media, Cognitive Reflection, and Conspiracy\nBeliefs. Front. Polit. Sci. 2021;3(June):1–8. https://doi.org/10.3389/\nfpos.2021.647957. [29] Weinberg J. Can Political Trust Help to Explain Elite Policy Support and Public\nBehaviour in Times of Crisis? Evidence from the United Kingdom at the Height\nof the 2020 Coronavirus Pandemic. Polit. Stud. 2020. https://doi.org/10.1177/\n0032321720980900. [5] Bruder M, Haffke P, Neave N, Nouripanah N, Imhoff R. Measuring Individual\nDifferences in\nGeneric Beliefs\nin Conspiracy Theories Across Cultures:\nConspiracy Mentality Questionnaire. Front Psychol 2013;4(April). https://doi. org/10.3389/fpsyg.2013.00225. [30] Yuan H, Long Q, Huang G, Huang L, Luo S. Different roles of interpersonal trust\nand institutional trust in COVID-19 pandemic control. Ethical review [10] Easton D. A Re-Assessment of the Concept of Political Support. Br J Polit Sci\n1975;5(4):435–57. https://doi.org/10.1017/S0007123400008309. [11] Frederick S. Cognitive reflection and decision making. J Econ Perspect 2005;19\n(4):25–42. https://doi.org/10.1257/089533005775196732. This study was approved by the Faculty of Social Sciences\nResearch Ethics Committee at the University of Southampton\n(Ethics/ERGO Number: 61735). [12] Garry J, Ford R, Johns R. Coronavirus conspiracy beliefs, mistrust, and\ncompliance:\nTaking\nmeasurement\nseriously. Psychol\nMed\n2021;1–11. https://doi.org/10.1017/S0033291720005164. p\n//\ng/\n/\n[13] Hetherington MJ. Why Trust Matters: Declining Political Trust and the Demise\nof American Liberalism (Kindle Edi). Princeton University Press; 2005. Discussion Political trust in the first year of the COVID-19 pandemic: a meta-\nanalysis of 67 studies. J Eur Publ Policy 2023. https://doi.org/10.1080/\n13501763.2023.2169741. Declaration of Competing Interest 10.1007/s11135 017 0584 6. [16] Karic´ T, Mededovic´ J. Covid-19 conspiracy beliefs and containment-related\nbehaviour: The role of political trust. Pers Individ Differ 2021;175. https://doi. /10 1016/j\nid 2021 11069 /\n[16] Karic´ T, Mededovic´ J. Covid-19 conspiracy beliefs and containment-related\nbehaviour: The role of political trust. Pers Individ Differ 2021;175. https://doi. org/10.1016/j.paid.2021.110697. The authors declare the following financial interests/personal\nrelationships which may be considered as potential competing\ninterests: Will Jennings reports financial support was provided by\nthe UK Economic and Social Research Council (ESRC). Will Jennings\nreports financial support was provided by Research England QR\nGlobal Challenges Research Fund (GCRF). [17] Kerr JR, Schneider CR, Recchia G, Dryhurst S, Sahlin U, Dufouil C, et al. Correlates\nof\nintended\nCOVID-19\nvaccine\nacceptance\nacross\ntime\nand\ncountries: Results from a series of cross-sectional surveys. BMJ Open\n2021;11(8). https://doi.org/10.1136/bmjopen-2020-048025. [18] Lang FR, John D, Lüdtke O, Schupp J, Wagner GG. Short assessment of the Big\nFive: Robust across survey methods except telephone interviewing. Behav Res\nMethods 2011;43(2):548–67. https://doi.org/10.3758/s13428-011-0066-z. Acknowledgments [19] Lazarus JV, Ratzan SC, Palayew A, Gostin LO, Larson HJ, Rabin K, et al. A global\nsurvey of potential acceptance of a COVID-19 vaccine. Nat Med 2021;27\n(2):225–8. https://doi.org/10.1038/s41591-020-1124-9. Financial support was provided by the ESRC (ES/S009809/1) and\nResearch\nEngland\nQR\nGlobal\nChallenges\nResearch\nFund\n(519016146). [20] Lindholt MF, Jørgensen F, Bor A, Petersen MB. Public acceptance of COVID-19\nvaccines: Cross-national evidence on levels and individual-level predictors\nusing observational data. BMJ Open 2021;11(6):1–12. https://doi.org/10.1136/\nbmjopen-2020-048172. [21] Mathieu E, Ritchie H, Rodés-Guirao L, Appel C, Giattino C, Hasell J, et al. ‘‘Coronavirus Pandemic (COVID-19)”. Published online at OurWorldInData. org; 2022. Retrieved from: ’https://ourworldindata.org/coronavirus’ [Online\nResource]. Data will be made available on request. Data will be made available on request. [15] Johnston R, Jones K, Manley D. Confounding and collinearity in regression\nanalysis: a cautionary tale and an alternative procedure, illustrated by studies\nof British voting behaviour. Qual Quant 2018;52(4):1957–76. https://doi.org/\n10.1007/s11135-017-0584-6. Appendix A. Supplementary material Supplementary data to this article can be found online at\nhttps://doi.org/10.1016/j.jvacx.2023.100299. ]\n[22] McCarthy M, Murphy K, Sargeant E, Williamson H. Examining the relationship\nbetween conspiracy theories and COVID-19 vaccine hesitancy: A mediating\nrole for perceived health threats, trust, and anomie? Anal Soc Issues Public\nPolicy 2022;22(1):106–29. https://doi.org/10.1111/asap.12291. Data availability [14] Jennings W, Stoker G, Valgarðsson V, Devine D, Gaskell J. How trust, mistrust\nand distrust shape the governance of the COVID-19 crisis. J Eur Publ Policy\n2021;28(8):1174–96. https://doi.org/10.1080/13501763.2021.1942151. References Soc Sci Med 2022; 293\n(August 2021): 114677. doi:10.1016/j.socscimed.2021.114677. [6] Bunting H., Gaskell J., Stoker G. Trust, Mistrust and Distrust: A Gendered\nPerspective\non\nMeanings\nand\nMeasurements. Front. Polit. Sci. 2021:3\n(November):1-15. doi:10.3389/fpos.2021.642129. 9 9"
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Historicising Entanglements: Science, Technology and Socio-Ecological Change in the Postcolonial Anthropocene
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de Hoop, E., Sridhar, A., da Silva, C. M., & van der Vleuten, E. B. A. (2022). Historicising Entanglements:
Science, Technology and Socio-Ecological Change in the Postcolonial Anthropocene . Global Environment,
15(2), 195-208. https://doi.org/10.3197/ge.2022.150201 Citation for published version (APA):
de Hoop, E., Sridhar, A., da Silva, C. M., & van der Vleuten, E. B. A. (2022). Historicising Entanglements:
Science, Technology and Socio-Ecological Change in the Postcolonial Anthropocene . Global Environment,
15(2), 195-208. https://doi.org/10.3197/ge.2022.150201 Citation for published version (APA):
de Hoop, E., Sridhar, A., da Silva, C. M., & van der Vleuten, E. B. A. (2022). Historicising Entanglements:
Science, Technology and Socio-Ecological Change in the Postcolonial Anthropocene . Global Environment,
15(2), 195-208. https://doi.org/10.3197/ge.2022.150201 DOI:
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Science, Technology and Socio-
Ecological Change in the
Postcolonial Anthropocene Evelien de Hoop, Aarthi Sridhar, Claiton
Marcio da Silva and Erik van der Vleuten Evelien de Hoop, Aarthi Sridhar, Claiton
Marcio da Silva and Erik van der Vleuten 1 P.J. Crutzen and E.F. Stoermer, ‘The “Anthropocene”’, IGPB Newsletter 41
(2000): 17–18; P.J. Crutzen, ‘Geology of mankind’, Nature 415 (51) (2002): 23.
For a history, see H. Trischler, ‘The Anthropocene: A challenge for the history of
science, technology, and the environment’, NTM Journal of the History of Science,
Technology and Medicine 24 (3) (2016): 309–35. gy
2 J. Lorimer, ‘Multinatural geographies for the Anthropocene’, Progress in Hu-
man Geography 36 (5): 593–612, at 606. Also, D. Chakrabarty, The Climate of
History in a Planetary Age (Chicago and London: The University of Chicago Press,
2021); G. Caluya, ‘Fragments for a postcolonial critique of the Anthropocene: in-
vasion biology and environmental security’, in J. Frawley and I. McCalman (eds),
Rethinking Invasion Ecologies from the Environmental Humanities (London: Rout-
ledge, 2014); A.S. Mathews, ‘Anthropology and the Anthropocene: criticisms, ex-
periments, and collaborations’, Annual Review of Anthropology 49 (2020): 67–82;
B. Latour, ‘Anthropology at the time of the Anthropocene: A personal view of
what is to be studied’, in M. Brightman and J. Lewis (eds), The Anthropology
of Sustainability (New York: Palgrave Macmillan, 2017), pp. 35–49; J. Lorimer,
‘The Anthropo-scene: A guide for the perplexed’, Social Studies of Science 47 (1)
(2017):117–42. T.J. LeCain, ‘Against the Anthropocene: a Neo-materialist per-
spective’, International Journal for History, Culture and Modernity 3 (2015): 1–28. Evelien de Hoop, Aarthi Sridhar, Claiton
Marcio da Silva and Erik van der Vleuten his special issue focuses on connected histo-
ries of science, technology and socio-ecolog-
ical change in what we call the ‘postcolonial
Anthropocene’. We used this term to guide
the papers in this issue towards research
questions that interrogate both human-na-
ture relations and postcolonial relations, as
entangled components of each inquiry. Of
T
course, both constituent terms ‘postcolonial’ and ‘Anthropocene’
are contentious and much-debated, with diverse connotations and
perceived implications to diverse readerships; using both together
can be a risky business. We nevertheless chose to do so, because we
felt that, both in historiography and in wider academic discourses,
Anthropocene and postcolonial research perspectives have too often
remained poorly connected – despite the growing number of schol-
ars arguing that Anthropocene research must include postcolonial
perspectives and vice versa. In this introductory essay, we begin by
discussing specific lines of (postcolonial) Anthropocene scholarship
we chose to connect to, before we specify this issue’s overarching
research questions and introduce the context and content of the in-
dividual papers. GE
195 GE
195 The ‘Anthropocene’ and its critiques Caluya, ‘Fragments for a postcolonial critique of the Anthropocene: in-
vasion biology and environmental security’, in J. Frawley and I. McCalman (eds),
Rethinking Invasion Ecologies from the Environmental Humanities (London: Rout-
ledge, 2014); A.S. Mathews, ‘Anthropology and the Anthropocene: criticisms, ex-
periments, and collaborations’, Annual Review of Anthropology 49 (2020): 67–82;
B. Latour, ‘Anthropology at the time of the Anthropocene: A personal view of
what is to be studied’, in M. Brightman and J. Lewis (eds), The Anthropology
of Sustainability (New York: Palgrave Macmillan, 2017), pp. 35–49; J. Lorimer,
‘The Anthropo-scene: A guide for the perplexed’, Social Studies of Science 47 (1)
(2017):117–42. T.J. LeCain, ‘Against the Anthropocene: a Neo-materialist per-
spective’, International Journal for History, Culture and Modernity 3 (2015): 1–28. The ‘Anthropocene’ and its critiques It has been over two decades since Paul J. Crutzen and Eugene F. Stoermer proposed the term ‘Anthropocene’ for the current geological
epoch in the International Geosphere-Biosphere Programme IGBP
newsletter (and soon after in a famous article in Nature). The term,
informally in use since at least the 1980s, was intended to emphasise
the ‘central role of mankind’ as ‘a powerful geological force’ changing
the ‘systemic properties’ of the planet and the atmosphere.1 Today (at
the time of writing) geologists are still debating whether to accept the
Anthropocene as an official subdivision of geological time. Meanwhile
humanities and social science scholars of many stripes have also mas-
sively engaged with the Human Epoch as a geological as well as a cul-
tural notion, based on their own (inter)disciplinary perspectives and
concerns. Amongst those perspectives, we would like to highlight that
many have lauded ‘the diagnosis of the Anthropocene’ as a decisive
acknowledgement of ‘the public death of the modern understanding
of Nature’, whereafter humans can no longer be regarded as operat-
ing in isolation from the ecologies in which they live.2 This insight gy
2 J. Lorimer, ‘Multinatural geographies for the Anthropocene’, Progress in Hu-
man Geography 36 (5): 593–612, at 606. Also, D. Chakrabarty, The Climate of
History in a Planetary Age (Chicago and London: The University of Chicago Press,
2021); G. Caluya, ‘Fragments for a postcolonial critique of the Anthropocene: in-
vasion biology and environmental security’, in J. Frawley and I. McCalman (eds),
Rethinking Invasion Ecologies from the Environmental Humanities (London: Rout-
ledge, 2014); A.S. Mathews, ‘Anthropology and the Anthropocene: criticisms, ex-
periments, and collaborations’, Annual Review of Anthropology 49 (2020): 67–82;
B. Latour, ‘Anthropology at the time of the Anthropocene: A personal view of
what is to be studied’, in M. Brightman and J. Lewis (eds), The Anthropology
of Sustainability (New York: Palgrave Macmillan, 2017), pp. 35–49; J. Lorimer,
‘The Anthropo-scene: A guide for the perplexed’, Social Studies of Science 47 (1)
(2017):117–42. T.J. LeCain, ‘Against the Anthropocene: a Neo-materialist per-
spective’, International Journal for History, Culture and Modernity 3 (2015): 1–28. gy
2 J. Lorimer, ‘Multinatural geographies for the Anthropocene’, Progress in Hu-
man Geography 36 (5): 593–612, at 606. Also, D. Chakrabarty, The Climate of
History in a Planetary Age (Chicago and London: The University of Chicago Press,
2021); G. 6 E.W. Krauss, ‘Anthropology in the Anthropocene: sustainable development,
climate change and interdisciplinary research’, in H. Greschke and J. Tischler
(eds), Grounding Global Climate Change (New York: Springer 2015), pp. 59–76;
L. Ogden et al., ‘The politics of Earth stewardship in the uneven Anthropocene’,
in R. Rozzi et al. (eds), Earth Stewardship. Linking Ecology and Ethics in Theory
and Practice (New York: Springer 2015), pp. 137–57; J. Hope, ‘The anti‐politics
of sustainable development: Environmental critique from assemblage thinking in
Bolivia’, Transactions of the Institute of British Geographers 46 (1) (2021): 208–22. h
p
4 R. Costanza et al., ‘Sustainability or collapse: what can we learn from in-
tegrating the history of humans and the rest of nature?’, AMBIO: A Journal of
the Human Environment 36 (7) (2007): 522–27; E. Russell, Evolutionary History:
Uniting History and Biology to Understand Life on Earth (Cambridge: Cambridge
University Press, 2011); J.L. Caradonna (ed.), Routledge Handbook of the History of
Sustainability (New York, NY: Routledge, 2018); S. Sörlin, ‘Reform and respon-
sibility – the climate of history in times of transformation’, Historisk tidsskrift 97
(1) (2018): 7–23.hh 5 C. Bonneuil and J.-B. Fressoz, The Shock of the Anthropocene. The Earth, His-
tory and Us (London and New York: Verso, 2016), p. 13. 3 Trischler, ‘The Anthropocene’, 312. 7 J. Guldi and D. Armitage, The History Manifesto (Cambridge, Ma: Cam-
bridge University Press, 2014), p. 69; Z.B. Simon, ‘History manifested: mak-
ing sense of unprecedented change’, European Review of History 22 (5) (2015):
819–34; Z.B. Simon, The Epochal Event: Transformations in the Entangled Human,
Technological, and Natural Worlds (London: Palgrave MacMillan, 2020). g
g
8 D. Haraway et al., ‘Anthropologists are talking – about the Anthropocene’,
Ethnos. Journal of Anthropology 81 (3) (2016): 535–64, p. 539. Also: D. Chandler,
‘The transvaluation of critique in the Anthropocene’, Global Society 33 (1): 26-44;
Chakrabarty, The Climate of History in a Planetary Age; D. Haraway, ‘Anthropo-
cene, Capitalocene, Plantationocene, Chthulucene: Making kin’, Environmental
Humanities 6 (2015): 159–65; B. Latour, Down to Earth: Politics in the New Cli-
matic Regime (Cambridge: Polity Press, 2018); A. Malm and A. Hornborg, ‘The
geology of mankind? A critique of the Anthropocene narrative’, The Anthropocene
Review 1 (1) (2014): 62–69; A. Blok and G.B. Jensen, ‘The Anthropocene event
in social theory: On ways of problematizing nonhuman materiality differently’,
The Sociological Review 67 (6) (2019): 1195–211. The classic is J. Ferguson, The Anti-politics Machine: ‘Development’, Depoliticiza-
tion and Bureaucratic Power in Lesotho (Chicago: Chicago University Press, 1990).h INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN Ferguson, The Anti-politics Machine: ‘Development’, Depoliticiza-
tion and Bureaucratic Power in Lesotho (Chicago: Chicago University Press, 1990).h g
g
7 J. Guldi and D. Armitage, The History Manifesto (Cambridge, Ma: Cam-
bridge University Press, 2014), p. 69; Z.B. Simon, ‘History manifested: mak-
ing sense of unprecedented change’, European Review of History 22 (5) (2015):
819–34; Z.B. Simon, The Epochal Event: Transformations in the Entangled Human,
Technological, and Natural Worlds (London: Palgrave MacMillan, 2020). INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 196 inspired scholars to rethink interrelations between histories of science,
technology and the environment.3 It also fostered, or resonated with,
diverse initiatives towards collaboration across the natural sciences,
social sciences and humanities to develop new knowledges informing
the mitigation of the Anthropocene’s ecological challenges – including
new historiographical knowledges.4 g p
g
However, and crucially, the proliferating use of the term An-
thropocene has also met with substantial critique. We would like to
emphasise two related critiques that are widely shared among histo-
rians, anthropologists and geographers. First, concerning human re-
lations, natural science-initiated accounts of the Anthropocene have
been much criticised for depoliticising history by presenting ‘history
as a contest between the human species as a whole and the planet,
with societies as ignorant and passive masses who can only be guided
by scientists and saved by green technologies’, as Bonneuil and Fres-
soz bluntly phrased it.5 Research that universalises human agency
tends to obscure differential human experiences, responsibilities and
politics – some scholars therefore call such research an ‘anti-politics
machine’.6 Second, concerning human-nature relations, the critique GE
197 G
197 G
197 is that omnipresent modernist uses of the term Anthropocene do
not transcend the nature-society dualism at all, but instead enact,
reproduce and amplify human exceptionalism. Implicitly and often
quite explicitly, such modernist Anthropocene scholarship attributes
to ‘humankind’ the position of primal Earth-changer and, as a con-
sequence, Earth stewardship to repair environmental harm done in
the ‘Human Epoch’. Such unabashed anthropocentrism also echoes
in historiographical debate of the Anthropocene; for example, the
History Manifesto endorsed calls for (long-term) histories to assists
humans in their role as responsible Earth managers – and was fierce-
ly criticised by others arguing that such human exceptionalist think-
ing is part of the problem rather than the solution.7 Either way, the
combination of over-emphasising and universalising human agency
obscures how change on Earth emerges from complex, connected
and situated interactions between a wide variety of human and non-
human agents, and that such change may be distributed unequally
across the earth. The world is not a ‘human species act’, as the term
Anthropocene may portray it to be.8 h These critiques elicited many responses. One prominent line
has been to develop a range of alternative terms to the notion of The classic is J. 9 J.W. Moore, Anthropocene or Capitalocene? Nature, History, and the Crisis of
Capitalism (Oakland: PM Press, 2016).
10 D. Haraway, Staying with the Trouble: Making Kin in the Chthulucene (Dur-
ham, NC: Duke University Press, 2016); M. Armiero, Wasteocene: Stories from
the Global Dump (Cambridge: Cambridge University Press, 2021); C.M. da Silva
and C. de Majo ‘Towards the soyacene: Narratives for an environmental history
of soy in Latin America’s Southern Cone’, Historia Ambiental Latinoamericana y
Caribeña 11 (1) (2021): 329–56. 11 Haraway, ‘Anthropocene, Capitalocene, Plantationocene, Chthulucene’; J.
Davis et al., ‘Anthropocene, Capitalocene, … Plantationocene? A manifesto for
ecological justice in an age of global crises’, Geography Compass 13 (5) (2019):
e12438; B. Latour et al., ‘Anthropologists are talking – about capitalism, ecology,
and apocalypse’, Ethnos 83 (3) (2018): 587–606. N. Myers, ‘From the Anthro-
pocene to the Planthroposcene: Designing gardens for plant/people involution’,
History and Anthropology 28 (3) (2017): 297–301. 9 J.W. Moore, Anthropocene or Capitalocene? Nature, History, and the Crisis of
Capitalism (Oakland: PM Press, 2016).
10 D. Haraway, Staying with the Trouble: Making Kin in the Chthulucene (Dur-
ham, NC: Duke University Press, 2016); M. Armiero, Wasteocene: Stories from
the Global Dump (Cambridge: Cambridge University Press, 2021); C.M. da Silva
and C. de Majo ‘Towards the soyacene: Narratives for an environmental history
of soy in Latin America’s Southern Cone’, Historia Ambiental Latinoamericana y
Caribeña 11 (1) (2021): 329–56.
11 Haraway, ‘Anthropocene, Capitalocene, Plantationocene, Chthulucene’; J.
Davis et al., ‘Anthropocene, Capitalocene, … Plantationocene? A manifesto for
ecological justice in an age of global crises’, Geography Compass 13 (5) (2019):
e12438; B. Latour et al., ‘Anthropologists are talking – about capitalism, ecology,
and apocalypse’, Ethnos 83 (3) (2018): 587–606. N. Myers, ‘From the Anthro-
pocene to the Planthroposcene: Designing gardens for plant/people involution’,
History and Anthropology 28 (3) (2017): 297–301. INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 198 the Anthropocene, which explicitly invite scrutiny of human and
multispecies differential politics, including postcolonial relations. To mention but a few: the notion of Capitalocene presents, in the
words of Moore, the world as a ‘multi-species assemblage, a world-
ecology of capital, power, and nature’.9 The term Plantationocene
takes inspiration from the study of colonial plantation systems and
refers to the radical simplification of previously diverse living sys-
tems and their relocation elsewhere, and thereby producing life for
value extraction. The Wasteocene concept invites historians to scru-
tinise the embodied stratigraphy of power and toxicity with special
attention to subaltern human and more-than-human communities;
the notion of the Soyacene invites study of how soybean has medi-
ated human and non-human lives especially in the last fifty years.10
Notions such as Chthulucene (derived from the Greek word chthon
meaning earth) and Planthroposcene refer not to critical analysis of
historical epochs, but to an ongoing attempt to deliberately shift the
terms of encounter with Anthropocene-type-of-concerns for future
modes of multispecies engagement.11 Note that these are only some
of the alternative ‘Big Words’ that have been proposed recently to
guide research on today’s major social and environmental challenges
in more critical and sensitive directions. Still, as observed by Hara-
way, the use of such terms comes with the risk of creating a Theory GE
199 GE
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199 of Everything at the expense of empirical study that takes historical
contingency, specificity and situatedness seriously.12 12 Haraway et al., ‘Anthropologists are talking – about the Anthropocene’,
550 and 651; C. Bos et al, ‘Steering with big words: Articulating ideographs in
research programs’, Journal of Responsible Innovation 1 (2) (2014): 151–70.h p
g
f
p
13 S. Harding (ed.), The Postcolonial Science and Technology Studies Reader
(Durham, NC: Duke University Press, 2011); J. Law and W. Y. Lin, ‘Provincial-
izing STS: Postcoloniality, symmetry, and method’, East Asian Science, Technology
and Society 11 (2) (2017): 211–27; A. Kumar et al. (eds), Dilemmas of Energy
Transitions in the Global South (New York: Routledge, 2021); Ute Hasenörl, ‘His-
tories of technology and the environment in post/colonial Africa: Reflections on
the field’, Histories 1 (3)(2021): 122–44; J. van der Straeten, ‘The rhythms behind
change. Historiography and the temporality of non-Western technological land-
scapes’, Technikgeschichte 88 (2) (2021): 191–96. Understanding socio-ecological
entanglements in the postcolonial
Anthropocene This latter observation leads us to literatures highlighting and
studying the historical, spatial, social and material situatedness of
specific socio-ecological entanglements and changes, most notably
historical and postcolonial Science and Technology Studies (STS).13
As such, this special issue approaches humans as diverse beings who
are co-constituted by and act together with the (social, material,
ecological) environments in which they live.14 This entanglement g
f
p
13 S. Harding (ed.), The Postcolonial Science and Technology Studies Reader
(Durham, NC: Duke University Press, 2011); J. Law and W. Y. Lin, ‘Provincial-
izing STS: Postcoloniality, symmetry, and method’, East Asian Science, Technology
and Society 11 (2) (2017): 211–27; A. Kumar et al. (eds), Dilemmas of Energy
Transitions in the Global South (New York: Routledge, 2021); Ute Hasenörl, ‘His-
tories of technology and the environment in post/colonial Africa: Reflections on
the field’, Histories 1 (3)(2021): 122–44; J. van der Straeten, ‘The rhythms behind
change. Historiography and the temporality of non-Western technological land-
scapes’, Technikgeschichte 88 (2) (2021): 191–96. p
g
14 Note also that in his ‘Postcolonial Studies and the Challenge of Climate
Change’, Chakrabarty speaks of the postcolonial ‘view of the human as the same
everywhere ... endowed with … anthropological difference’. This has received criti-
cism from within postcolonial scholarship, and we follow the latter – work which
recognises the emergence of difference without making the assumption of univer-
sality being hidden behind difference (e.g. D. Boscov-Ellen, ‘Whose universalism? Dipesh Chakrabarty and the Anthropocene’, Capitalism Nature Socialism 31 (1):
70–83; G. Caluya, ‘Fragments for a postcolonial critique of the Anthropocene: inva-
sion biology and environmental security’, in J. Frawley and I. McCalman (eds), Re-
thinking Invasion Ecologies from the Environmental Humanities (London: Routledge,
2014); G. Jack, ‘Advancing postcolonial approaches in critical diversity studies’, in
R. Bendi et al. (eds), The Oxford Handbook of Diversity in Organizations (Oxford: INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 200 has implications not only for our understanding of humans, their
agency and power imbalances in the Anthropocene, but also for
our understanding of what constitutes these humans’ material and
ecological ‘environments’. It means we do not approach such en-
vironments as singular, not a ‘One-World world’ as geological and
other natural sciences studying the Anthropocene would have it,15 Oxford University Press, 2015)). Postcolonial theorists’ conceptualisation of human
beings and their agency as being co-constituted by the environments in which they
live draws upon both (older) STS work, such as actor-network theory (B. Latour,
Reassembling the Social: An Introduction to Actor-Network-Theory (New York: Oxford
University Press, 2005)) and on more recent new materialist work which has been
spurred in part by the challenge of the Anthropocene (A.F. Conty, ‘The Politics of
nature: New materialist responses to the Anthropocene’, Theory, Culture and So-
ciety 35 (7–8) (2018): 73–96). See, for example, the notion of trans-corporeality
proposed by feminist critical theorist Alaimo, denoting that ‘the human is always
intermeshed with the more-than-human world, ... ultimately inseparable from “the
environment”’ (S. Alaimo, Bodily Natures: Science, Environment, and the Material
Self (Bloomington, IN: Indiana University Press, 2010), p. 2). f
g
y
p
15 M. De La Cadena and M. Blaser, A World of Many Worlds (Durham, NC:
Duke University Press, 2018). This One-World world has been critiqued for creat-
ing a colonial divide between ‘those who function within the OWW from those
who insist on other ways of bringing a world into being, or “worlding”’ (A. Es-
cobar, ‘Thinking-feeling with the Earth: Territorial struggles and the ontological
dimension of the epistemologies of the South’, Revista de Antropología Iberoameri-
cana 11 (1) (2016): 11–32, at 21). Indeed, scholarship on indigenous protests
against extractivism, such as Australian Aboriginal resistance to drilling (D. Dan-
owski and E. de Castro, The Ends of the World (Cambridge, UK and Malden,
Ma: Polity Press, 2016)) or Meswaki and Sioux resistance to the Dakota Access
Pipeline (G. Giuliani, Monsters, Catastrophes and the Anthropocene; A Postcolonial
Critique (Abingdon, Oxon and New York, NY: Routledge, 2021)) has power-
fully brought the power and relevance of a plurality of locally-situated and (often)
non-anthropocentric cosmologies to the fore, and contrasted these with those en-
acted through extractivist approaches. 16 A. Blunt and C. McEwan, Postcolonial Geographies (London and New York:
Continuum, 2002).
17 R. Wenzlhuemer, Doing Global History: An Introduction in 6 Concepts (Lon-
don and New York: Bloomsbury, 2020). On infrastructure, supply chain, and
commodity circulation histories see e.g., A. Johnson, ‘Europe without borders:
Environmental and global history in a world after continents’, Contemporary Eu-
ropean History 31 (1) (2022): 129–41; L. Marques, ‘Commodity chains and the
global environmental history of the colonial Americas’, Esboços: histórias em con-
textos globais 28 (49) (2021): 640–97; P. Högselius et al., Europe’s Infrastructure
Transition: Economy, War, Nature (London: Palgrave MacMillan, 2016). INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN Other ways of worlding have not only been
identified in scholarship on resistance against extractivism, but of course also in,
amongst others, work on Indigenous worldviews on ecology and American Indian
cosmologies, and on non-modern philosophies of life such as Buen Vivier. See,
amongst many others, J.M. Beijer, ‘Beyond hegemonic state(ment)s of Nature. Indigenous knowledge and non-state possibilities in international relations’, in C. Geeta and S. Nair (eds), Power, Postcolonialism and International Relations. Read-
ing Race, Gender and Class (London: Routledge, 2002). GE
201 GE
201 G
201 G
201 but as multiple and possibly inequitable, arising together with the
imaginations and practices of the humans who live in them. Indeed,
and critically, understanding both humans and their environments
as co-constituting each other helps to bring (post)colonial power re-
lations and inequalities in the shaping of sited human-environment
relationships to the fore. This resulted in the following question that
guided the investigations of the papers published in this special is-
sue: which human-environment constellations emerged at the site(s)
studied, how did they relate to one another, and what inequalities were
embedded therein? Second, we approach this process of co-constitution in (post)
colonial worlds as transnationally entangled.16 In particular, the pa-
pers in this special issue take the lead from two bodies of historical
work in which such transnational entanglements feature particu-
larly prominently, namely histories of relations and relational histo-
ries. The former constitute historical investigations of connections
and relations between different sites and the ways in which these
relations and sited socio-ecological dynamics co-constituted each
other.17 These histories of relations predominantly approach diverse
sites’ histories from the perspective of a common logic (and, despite
claims to the contrary, not seldom a European or global North-in-
spired one), and their attentiveness to multi-sited historical diver-
sity has therefore been limited. In contrast, relational and connected
histories investigate how regional histories may be at once highly
geographically distant, diverse, specific and situated, yet neverthe- INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 18 See S. Subrahmaniam, Europe’s India. Words, People, Empires, 1500–1800
(Harvard: Harvard University Press, 2017).
19 A.L. Stoler, Along the Archival Grain: Epistemic Anxieties and Colonial Com-
mon Sense (Princeton and Oxford: Princeton University Press, 2009). We consider
this important, amongst others, in view of providing room to diverse ways of
understanding and living in today’s world(s), cf. A. Escobar, Designs for the Pluriv-
erse Radical Interdependence, Autonomy, and the Making of Worlds (Durham: Duke
University Press, 2018).h INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 202 less develop in mutual interaction.18 Such work may explicitly move
against the archival grain, deconstructing the dominant narratives of
archives and contemporary historical ‘sources’ that scholars use and
contrasting these with the narratives of a wide diversity of voices.19
In both types of histories, relations are established through the trav-
els of a wide variety of entities; in line with this issue’s grounding in
postcolonial STS we particularly focus on (expert and lay) knowl-
edges and technologies as phenomena that connect and co-construct
geographically dispersed and locally situated socio-ecological histo-
ries. This resulted in the second question that guided the investiga-
tions of the papers published in this special issue: what roles did
science and technology, which often travelled between diverse locations
across the globe, play in the emergence of and inequalities embedded in
the diverse human-environment constellations identified in response to
the first question? 20 Crutzen and Stoermer, ‘The “Anthropocene”’, 18. 21 The initiative sprang from a workshop in Lisbon in Autumn 2019, organ-
ised in the context of the Tensions of Europe programme Technology and Societal
Challenges 1850–2050 and its ‘Technology, environment and resources’ working
group. For the broader programme: E. van der Vleuten, ‘Technology, societal
challenges, and global sustainability history’, Icon 24 (2018): 34–52; E. van der
Vleuten, ‘History and technology in an age of grand challenges: Raising ques-
tions’, Technology and Culture 61(1): 260–71. For the ‘Technology, environment
and resources’ working group: M. Heymann et al. ‘Challenging Europe: Tech-
nology, environment, and the quest for resource security’, Technology and Cul-
ture 61 (1) (2020): 282–94; O. Sparenberg and M. Heymann (eds), ‘Resource
challenges and constructions of scarcity in the 19th and 20th centuries’, Special
Issue in European Review of History 27 (1–2) (2020): 243–369; C. Kehrt and J.
Martin (eds), ‘Reconfiguring nature: Resource security and the limits of expert
knowledge’, Global Environment 13 (3) (2020): 512–658; A. Åberg and F. Ve-
raart (eds), ‘Creating, capturing and circulating commodities: The technology and
politics of material resource flows, from the 19th century to the present’, Special
section in The Extractive Industries and Society 7 (1) (2020): 1–67; J. Daheur (ed.),
‘Extractive Peripheries in Europe: Quest for Resources and Changing Environ-
ments (Fifteenth-Twentieth Centuries)’, Special issue of Global Environment 15
(1) (2022): 7–147. Contributions to this special issue Crutzen and Stoermer offered the term ‘Anthropocene’ in order
to inspire and empower the development of ‘a world-wide accepted
strategy leading to sustainability of ecosystems against human in-
duced stresses’; such ‘great future tasks of mankind’ in their words
required intensive research efforts and the ‘wise application of the
knowledge thus acquired’.20 Critiquing such calls and research for
their universalisation of humanity and the obscuring of differential
experiences, responsibilities and politics, this special issue has sought
to study situated and diverse engagements with socio-ecological GE
203 GE
203 G
203 G
203 change, guided by the two questions raised in the foregoing para-
graphs.21 This effort has resulted in the following papers. hf
First, de Hoop and van der Vleuten study how human-environ-
ment constellations were variously articulated in scientific knowl-
edge on palm oil sustainability in Southeast Asia and Europe since
1980. They ask how such research enacted a postcolonial politics
of difference between Southeast Asia and Europe with regard to
defining ‘sustainability’ problems and solutions. They observe that
palm oil sustainability research originating from Southeast Asia fore-
grounded problems experienced and to be redressed within the re-
gion itself. By contrast, diverse strands of research lead by scholars
from Europe variously framed migrant, smallholder and large-scale
palm oil farmers in Southeast Asia as primarily responsible for caus-
ing and solving ‘global sustainability problems’, notably global de-
forestation and climate change. This ‘global sustainability’ discourse
by and large acquitted European actors and markets from such re-
sponsibilities – even though these had long deforested their own ter-
ritories, had played a major role in establishing palm oil cultivation, INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 204 trade and use, and had constituted prime markets for most of the
twentieth century. Next, da Silva and de Majo’s study of Southern Brazilian tradi-
tional populations known as Caboclos shows how a relationship of
mutual dependence between people and their forests had been built
up over centuries. This ‘bio-anthropophagic’ relationship changed
dramatically when a new wave of settlers advanced westward in the
early twentieth century, creating new human-environment constel-
lations based on governments’ and settlers’ ideas about both the for-
est and its traditional populations. These ideas were prominently
informed by the travels of both racial and eugenic theories and as-
sociated sanitation policies. Thus followed a rearrangement of the
exploitation of natural materials – such as yerba mate or herbal
medicines – based on new hierarchies. Local populations survived
through astute strategies of incorporating exogenous values, giving
rise to deep inequalities with respect to the ability to both materially
and ontologically define one’s one way of life. g
yi
y
Also located in South America, França de Oliveira and Zarilli’s
contribution analyses how the transformation of the Argentine
pampas into an area of large-scale cattle farming went hand in hand
with profound environmental, economic and demographic transfor-
mations. This included the incorporation of over forty million hec-
tares of pampas into market-based agriculture and cattle-farming;
the introduction of new animal and plant species as well as a range
of modern agricultural technologies; new economic models of farm-
ing; and a substantial increase in the number of people living in the
area. This paper demonstrates how these changes were mediated not
only by the introduction of cattle and the early establishment of
farms in the area in the sixteenth century, but also by transportation
infrastructures whereby railroads connected the pampas to Argen-
tina’s ports, reconfiguring the landscape. i
Next, Abazeed and Hafez investigate how cultural discourse co-
constructed the Nile as a ‘modern river’ through an analysis of imag-
inaries and knowledges of rivers as represented in two different sets
of writing: Egyptian renaissance writing by modernist Alī Mubārak,
who hybridised a European education into his nationalist develop- GE
205 GE
205 ment visions, and a set of travelogue diaries of orientalist European
travelers writing for European audiences. INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN Both cultural traditions
harnessed modernist ideas, but did so in situated ways, using dif-
ferent vantage points, to remake the free-flowing River Nile and its
people into a modern ‘hybrid entity’ drawing in multiple cultural
and natural attributes. Fotopoulos and Araposthatis highlight how explorations of hy-
drocarbons, regional visions and governing practices were historical-
ly co-constructed in the Eastern Mediterranean, with implications
for who has access to Greece’s hydrocarbons. They argue that Euro-
pean and North American expertise on hydrocarbon explorations as
well as various foreign exploration technologies and infrastructures
were critical to encoding these power constellations in regional hy-
drocarbon development. p
The final paper of the issue represents the tradition of investi-
gating histories of relations and connections rather than relational
histories: Veraart investigates how the development and travels of
catalysis technologies, which enabled the production of margarine
from a diversity of oils and fats, gave rise to a variety of new hu-
man-environment constellations at different sites across the world,
through putative supply chain relations forged in colonisation con-
texts across Congo, the Dutch Indies and the Antarctic, between
1910 and 1940. Acknowledgements This special issue springs from the inspiring workshop on ‘Re-
sources, Infrastructures and the Anthropocene: Dialogues between
the Global-North and the Global-South’, organised at the Univer-
sidade NOVA de Lisboa in 2019. We are particularly indebted to
the workshop participants for vivid, critical and inspiring contribu-
tions and debate. We also thank the organizers and funders of the
workshop: the Centro Interuniversitário de História das Ciências e da
Tecnologia, the Dutch research council NWO’s internationalising
humanities program GREASE (nr. IG.18.036), the Danish Council INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN 206 for Independent Research’s research network programme EurReS,
and the Foundation for the History of Technology SHT. for Independent Research’s research network programme EurReS,
and the Foundation for the History of Technology SHT. Erik van der Vleuten teaches at Eindhoven University of Technology and
researches the connected histories of technology and socioecological change in
an infrastructured world. Earlier works include: Europe’s Infrastructure Transition:
Economy, War, Nature (2015, with Per Högselius and Arne Kaijser) and Engineer-
ing the Future, Understanding the Past: A Social History of Technology (2017, with
Ruth Oldenziel and Mila Davids). His current research and education focus on
diverse and contested sustainability histories and futures of distant-yet-connected
regions across the globe. Email: e.b.a.v.d.vleuten@tue.nl INTRODUCTION / DE HOOP, SRIDHAR, DA SILVA, V. D. VLEUTEN for Independent Research’s research network programme EurReS,
and the Foundation for the History of Technology SHT. Evelien de Hoop is based at the transdisciplinary Athena Institute (Vrije
Universiteit Amsterdam, NL). With a background in Science and Technology
Studies, she works with scholars from diverse disciplines and societal stakehold-
ers to address contemporary health and sustainability challenges. Empirical foci
include Indian farmers’ engagements with biofuels, sustainable Dutch country-
sides, and inclusive food systems in deprived urban areas. She mobilises research
on the historical and transnational entanglements of contemporary challenges,
with a particular focus on knowledge politics, to render her contemporary work
historically-sensitive and pluralise ways of thinking about the future. Aarthi Sridhar is one of the founder trustees of Dakshin Foundation and
is pursuing a Ph.D. from the University of Amsterdam on a historical sociology
of fisheries science in India. Her academic interests centre on historical and con-
temporary socio-legal studies, Science and Technology Studies, and sociological
approaches to knowledge and expertise. Her empirical focal areas are coastal and
marine environments, resource politics, maritime infrastructures and practices
of environmental norms and justice. She enjoys collaborative projects and has
worked with diverse disciplinary teams on writing projects, documentary films,
technical websites, field manuals and other learning material on the subject of
marine environments and people. Email: aarthi77@gmail.com Email: aarthi77@gmail.com Claiton Marcio da Silva is Associate Professor at the Federal University
of the Southern Frontier (UFFS), Santa Catarina, Brazil and CNPq Productiv-
ity Researcher. He began studying the modernisation of Brazilian agriculture in
2002, completing his MA in history at the Federal University of Santa Catarina
(UFSC) with a thesis on rural youth in Southern Brazil. At Casa de Oswaldo Cruz
(COC/Fiocruz), his doctoral dissertation explored the role of Nelson Rockefeller’s
American International Association for Economic and Social Development (AIA)
in introducing the US extension service model in Latin America. Recently, he has
been exploring the global environmental history of soybeans and its impacts for
traditional populations in the Global South. Email: claiton@uffs.edu.br GE
207 GE
207 Erik van der Vleuten teaches at Eindhoven University of Technology and
researches the connected histories of technology and socioecological change in
an infrastructured world. Earlier works include: Europe’s Infrastructure Transition:
Economy, War, Nature (2015, with Per Högselius and Arne Kaijser) and Engineer-
ing the Future, Understanding the Past: A Social History of Technology (2017, with
Ruth Oldenziel and Mila Davids). His current research and education focus on
diverse and contested sustainability histories and futures of distant-yet-connected
regions across the globe.
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Symmetry Extensions and Their Physical Reasons in the Kinetic and Hydrodynamic Plasma Models
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Symmetry, integrability and geometry: methods and applications
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Symmetry Extensions and Their Physical Reasons
in the Kinetic and Hydrodynamic Plasma Models⋆ Volodymyr B. TARANOV Institute for Nuclear Research, 47 Nauky Ave., 03028 Kyiv, Ukraine
E-mail: wlatar@inet.ua URL: http://www.geocities.com/vebete/mypage.html p
g
yp g
Received October 31, 2007, in final form January 14, 2008; Published online January 17, 2008
Original article is available at http://www.emis.de/journals/SIGMA/2008/006/ Received October 31, 2007, in final form January 14, 2008; Published online January 17
O igi al a ticle is a ailable at htt
//
i
d /j
l /SIGMA/2008/006/ Received October 31, 2007, in final form January 14, 2008; Published online January 17, 2008
Original article is available at http://www.emis.de/journals/SIGMA/2008/006/ Original article is available at http://www.emis.de/journals/SIGMA/2008/006/ Abstract. Characteristic examples of continuous symmetries in hydrodynamic plasma
theory (partial differential equations) and in kinetic Vlasov–Maxwell models (integro-
differential equations) are considered. Possible symmetry extensions conditional and ex-
tended symmetries are discussed. Physical reasons for these symmetry extensions are
clarified. Key words: symmetry; plasma; hydrodynamic; kinetic Key words: symmetry; plasma; hydrodynamic; kinetic 2000 Mathematics Subject Classification: 35A30; 35Q60; 45K05; 58J70; 22E70 2000 Mathematics Subject Classification: 35A30; 35Q60; 45K05; 58J70; 22E70 ⋆This paper is a contribution to the Proceedings of the Seventh International Conference “Symmetry in
Nonlinear Mathematical Physics” (June 24–30, 2007, Kyiv, Ukraine).
The full collection is available at
http://www.emis.de/journals/SIGMA/symmetry2007.html Symmetry, Integrability and Geometry: Methods and Applications Symmetry, Integrability and Geometry: Methods and Applications SIGMA 4 (2008), 006, 7 pages 1
Introduction Symmetry considerations essentially help us to solve nonlinear problems of plasma physics. Among the recent publications, a survey of methods which allow us to find symmetries of integro-
differential equations of kinetic plasma theory [1] and a review of applications of symmetry
methods to the hydrodynamic plasma models based on the partial differential equations [2] can
be cited. In the present paper, some characteristic examples of the symmetries are presented for var-
ious plasma theory models: electron magnetohydrodynamics, collisionless electron plasma os-
cillations, drift waves in plasma and multi-component collisionless plasma containing particles
with equal charge to mass ratios. In Section 2, Lie point symmetries are obtained for the partial differential equations of the
electron magnetohydrodynamics by means of the standard Maple 11 program. A simple invariant
solution is presented, which is in fact a background solution of a perturbation theory formalism. An additional condition is underlined which can lead to the symmetry extension. In Section 3, continuous symmetries of the integro-differential kinetic equations of the colli-
sionless electron plasma model are discussed. The symmetries were found in [3] by an indirect
algorithm which allows us to obtain symmetries of the kinetic equations from the symmetries of
an infinite set of partial differential equations for the moments of distribution functions. In Section 4, some conditional symmetries are presented for the Hasegawa–Mima hydrody-
namic model describing drift waves in a plasma. Due to these symmetries, invariant solutions
are possible describing the asymptotic structure of the nonlinear waves. In Section 5, additional symmetries for integro-differential equations of the kinetic theory
of collisionless plasma containing particles with equal charge to mass ratio are considered. For
example, alpha particles and deuterium ions participating in a thermonuclear reaction D+ +
T+ →He++ + n + 17.6 MeV have close charge to mass ratios. By simple vector considerations V.B. Taranov 2 so called extended symmetry transformations are obtained which allow us to reduce the number
of equations. so called extended symmetry transformations are obtained which allow us to reduce the number
of equations. Conclusions are made in Section 6 Conclusions are made in Section 6. 2
Electron magnetohydrodynamics Let us consider the equations of the electron magnetohydrodynamics (EMHD) [4 ∂Ψ
∂t = ∇× (v × Ψ),
Ψ = B −∆B,
v = −∇× B,
∇· B = 0, where Ψ is the generalized vorticity, B is magnetic field strength and v is the hydrodynamic
velocity of the electron plasma component. Nine functions, Ψ, v, B, of four independent variab-
les t, x, y, z are present in this system of partial differential equations. Nevertheless, its
symmetry has been obtained by the standard Maple 11 package program. y
y
p
p
g p
g
The infinitesimal operators of Lie point symmetries of the model are as follows: X1 = ∂
∂t,
X2 = ∂
∂x,
X3 = ∂
∂y,
X4 = ∂
∂z ,
X5 = t ∂
∂t −v ∂
∂v −B ∂
∂B −Ψ ∂
∂Ψ,
X6 = r × ∂
∂r + v × ∂
∂v + B × ∂
∂B + Ψ × ∂
∂Ψ. X1 = ∂
∂t,
X2 = ∂
∂x,
X3 = ∂
∂y,
X4 = ∂
∂z ,
X5 = t ∂
∂t −v ∂
∂v −B ∂
∂B −Ψ ∂
∂Ψ,
X6 = r × ∂
∂r + v × ∂
∂v + B × ∂
∂B + Ψ × ∂
∂Ψ. The Maple 11 program was tested previously on Korteweg–de Vries, nonlinear Schr¨odinger,
Hasegawa–Mima and other nonlinear plasma theory models and reproduced well known sym-
metries. So we can expect that operators X1–X6 form a full basis of EMHD Lie point symmetry
algebra. A little doubt remains, however: maybe some additional symmetries could be obtained by
the direct ‘manual’ use of the standard Lie algorithm. Due to time and space homogeneity (symmetries X1 to X4) the simplest exact solution exists
corresponding to the constant external magnetic field which can be directed along the Oz axis
without the loss of generality (because of the rotation symmetry X6). The field amplitude can
be set as B0 = 1 due to the similarity transform generated by X5: (1) B = ez,
v = 0. B = ez,
v = 0. This solution can be chosen as a background for the perturbation theory. 3
Collision less electron plasma Let us consider electron collisionless plasma. In this case the Vlasov–Maxwell integro-differential
system of equations holds: ∂f
∂t + v∂f
∂x −E ∂f
∂v = 0,
(4)
∂E
∂x = 1 −
Z ∞
−∞
fdv,
∂E
∂t =
Z ∞
−∞
vfdv,
(5) (4) (5) where f(t, x, v) is the distribution function of the electron component of the plasma, E(t, x) is
the electric field strength. where f(t, x, v) is the distribution function of the electron component of the plasma, E(t, x) is
the electric field strength. We cannot find symmetries of this integro-differential model by the traditional Lie method. This can be done, however, by the indirect algorithm [3]. [ ]
First, let us introduce the moments: First, let us introduce the moments: First, let us introduce the moments: Mk(t, x) =
Z ∞
−∞
vkfdv,
k = 0, 1, 2, . . . (6) (6) and obtain an infinite set of partial differential equations for them, which is as follows: ∂Mk
∂t
+ ∂Mk+1
∂x
+ EkMk−1 = 0,
∂E
∂x = 1 −M0,
∂E
∂t = M1. Then, we consider the reduced system involving only first N + 1 equations of this set, and
we can obtain by the standard Lie procedure the symmetries of this system: Then, we consider the reduced system involving only first N + 1 equations of this set, and
we can obtain by the standard Lie procedure the symmetries of this system: X1 = ∂
∂t,
X2 = ∂
∂x,
X3 = x ∂
∂x + E ∂
∂E +
N+1
X
k=1
kMk
∂
∂Mk
,
X4 = cos (t)
∂
∂x + ∂
∂E
−sin (t)
N+1
X
k=1
kMk−1
∂
∂Mk
,
(7)
X5 = sin (t)
∂
∂x + ∂
∂E
+ cos (t)
N+1
X
k=1
kMk−1
∂
∂Mk
, (7) (7) and an additional symmetry and an additional symmetry and an additional symmetry XFGN = F(t)
∂
∂MN
+
G(t) −x∂F(t)
∂t
∂
∂MN+1
, F(t) and G(t) being arbitrary functions of t. The standard Maple 11 program can help us to check these results for not very large N
(depending on computer capacity). Only two highest moments MN and MN+1 are present in XFGN. So only moments with
numbers greater than any previously chosen are involved in the transform generated by XFGN. 2
Electron magnetohydrodynamics After the substi-
tution B →ez + B, Ψ →ez + Ψ the first EMHD equation becomes ∂Ψ
∂t = ∂v
∂z + ∇× (v × Ψ)
(2) ∂Ψ
∂t = ∂v
∂z + ∇× (v × Ψ) (2) but other equations remain unchanged. In this way the system of equations describing helicon
waves in a plasma is usually obtained. but other equations remain unchanged. In this way the system of equations describing helicon
waves in a plasma is usually obtained. We can expect that EMHD model is much more symmetric. For example, nonlinear term
in (2) vanishes under an additional condition: v × Ψ = ∇F,
(3) (3) v × Ψ = ∇F, where F is arbitrary scalar function. In this way we obtain the set of linear partial differential
equations and the nonlinear algebraic restriction (3). This property can be a source of rich
additional symmetries. where F is arbitrary scalar function. In this way we obtain the set of linear partial differential
equations and the nonlinear algebraic restriction (3). This property can be a source of rich
additional symmetries. Symmetry Extensions in the Kinetic and Hydrodynamic Plasma Models 3 3
Collision less electron plasma In the limit N →∞the symmetry group is generated by X1–X5. In general, restoring kinetic symmetries from the symmetries of the corresponding set of the
moments equations is a hard problem in pure mathematics. Nevertheless, for the relatively
simple operators X1–X5 it was shown in [3] that in the case considered they correspond to the
following infinitesimal operators of the kinetic theory: X1 = ∂
∂t,
X2 = ∂
∂x,
X3 = x ∂
∂x + v ∂
∂v + E ∂
∂E −f ∂
∂f ,
(8) (8) V.B. Taranov 4 X4 = cos (t)
∂
∂x + ∂
∂E
−sin (t) ∂
∂v,
X5 = sin (t)
∂
∂x + ∂
∂E
+ cos (t) ∂
∂v. (9) (9) Strictly speaking, from the definition (6) it can be readily seen that the transformations gen-
erated by the operators (8), (9) in the space of variables t, x, v and f lead to the transformations
generated by the infinitesimals (7) in the space of variables t, x and Mk, k = 0, 1, 2, . . . . Strictly speaking, from the definition (6) it can be readily seen that the transformations gen-
erated by the operators (8), (9) in the space of variables t, x, v and f lead to the transformations
generated by the infinitesimals (7) in the space of variables t, x and Mk, k = 0, 1, 2, . . . . The symmetries of the same electron collisionless plasma kinetic model (4), (5) were found
in [5] by the direct algorithm, i.e. without using of an infinite set of moments equations. The
same generators (8), (9) were obtained. 4
Hasegawa–Mima model Let us consider an inhomogeneous plasma slab in the external homogeneous magnetic field. Electrons, unlike ions, are magnetized, smoothing an electrostatic potential Φ along the magnetic
field lines. In this case, Hasegawa–Mima model equations hold [6]: ∂Ψ
∂t + J(Φ, Ψ) = ∂Φ
∂y ,
Ψ = Φ −∆⊥Φ, ∂Ψ
∂t + J(Φ, Ψ) = ∂Φ
∂y ,
Ψ = Φ −∆⊥Φ,
(10) Ψ = Φ −∆⊥Φ,
(10) (10) where Ψ is the generalized vorticity, where Ψ is the generalized vorticity, J(F, G) ≡∂F
∂x
∂G
∂y −∂G
∂x
∂F
∂y ,
∆⊥≡∂2
∂x2 + ∂2
∂y2 . J(F, G) ≡∂F
∂x
∂G
∂y −∂G
∂x
∂F
∂y ,
∆⊥≡∂2
∂x2 + ∂2
∂y2 . The simultaneous presence of the Φ and ∆⊥Φ in the second equation of (10) leads to the
symmetry reduction and, as a consequence, to the absence of self-similar solutions [7]. Nevertheless, self-similar solutions can exist as a consequence of conditional symmetries [8]. The notion of conditional symmetry is discussed and large bibliography is presented in a recent
paper [9]. For example, if an additional condition ∂2Φ
∂x2 = 0
(11) ∂2Φ
∂x2 = 0
(11) ∂2Φ
∂x2 = 0 (11) is fulfilled, the term ∆⊥Φ is reduced to ∂2Φ
∂y2 , which leads to the symmetry extension. In this
way we obtain more symmetric, but still nonlinear, set of equations, which admits the following
similarity transform: is fulfilled, the term ∆⊥Φ is reduced to ∂2Φ
∂y2 , which leads to the symmetry extension. In this
way we obtain more symmetric, but still nonlinear, set of equations, which admits the following
similarity transform: X1 = x ∂
∂x + Φ ∂
∂Φ + Ψ ∂
∂Ψ, which allows us to try solutions in the form: which allows us to try solutions in the form: Φ = xF(t, y),
Ψ = xG(t, y). G = −1 is a trivial solution which is not interesting for physics. So only the simple nonlinear
exact solution with G ̸= −1 important for physics as a nonlinear wave asymptotic, will be
considered below. More complicated solutions can be presented and discussed in future papers. For the functions F(t, y) and G(t, y) we obtain the equations For the functions F(t, y) and G(t, y) we obtain the equations ∂(1/(G + 1))
∂t
+ ∂(F/(G + 1))
∂y
= 0,
G = F −∂2F
∂y2 . 4
Hasegawa–Mima model (12) ∂(1/(G + 1))
∂t
+ ∂(F/(G + 1))
∂y
= 0,
G = F −∂2F
∂y2 . (12) Equations (12) have a particular solution [8]: (13) F = α(1 + β cos((ω + δω)t + qy)), Symmetry Extensions in the Kinetic and Hydrodynamic Plasma Models 5 so that so that Φ = αx(1 + β cos((ω + δω)t + qy)),
(14) αx(1 + β cos((ω + δω)t + qy)),
(14) (14) Φ = αx(1 + β cos((ω + δω)t + qy)), where α is arbitrary constant amplitude, constant factor β determines the relative weight of the
zonal flow and the monochromatic wave, the frequency is ω = q/(1+q2) and the frequency shift
is δω = −αq3/(1 + q2). where α is arbitrary constant amplitude, constant factor β determines the relative weight of the
zonal flow and the monochromatic wave, the frequency is ω = q/(1+q2) and the frequency shift
is δω = −αq3/(1 + q2). Another class of conditionally symmetric invariant solutions became possible under the fol-
lowing condition: ∂2Φ
∂y2 = 0. ∂2Φ
∂y2 = 0. In this case the following similarity transform appears as a conditional symmetry: In this case the following similarity transform appears as a conditional symmetry: X2 = t ∂
∂t + y ∂
∂y X2 = t ∂
∂t + y ∂
∂y X2 = t ∂
∂t + y ∂
∂y and another class of invariant solutions is possible Φ = F(x)y
t + G(x),
Ψ = (F(x) −F ′′(x))y
t + G(x) −G′′(x). Pure self similar solutions G(x) = 0 are absent in this case, but there exist solutions corre-
sponding to the arbitrary shear flow F(x) = 0, G(x) being an arbitrary function of x. 5
Multi-component collisionless plasma So the transformation [8] f′
1 = f1 −e2F(f1, f2),
f′
2 = f2 + e1F(f1, f2), (19) where F(f1, f2) is an arbitrary function of its arguments, leaves the Vlasov–Maxwell equations
invariant, at least if we perform in (17) the summation first and the integration later. The transformation (19) does not preserve, in general, the positiveness of the distribution
functions. Moreover, they can lead to the divergence of the moments, which can be shown, for
example, by choosing F = const. y
g
eless, we can choose F(f1, f2) = f1/e2 to obtain the simplifying transformation vertheless, we can choose F(f1, f2) = f1/e2 to obtain the simplifying transformation Nevertheless, we can choose F(f1, f2) = f1/e2 to obtain the simplifying transf e2f′
2 = e1f1 + e2f2. (20) f′
1 = 0,
e2f′
2 = e1f1 + e2f2. (20) The simplifying transform (20) can also be obtained from the symmetry deduced from the
symmetry of moments’ equations, f′
1 = f1 exp(a),
f′
2 = f2 + e1
e2
(1 −exp(a))f1, a is arbitrary constant, in the limit a →−∞. a is arbitrary constant, in the limit a →−∞. According to (20), we can choose the invariant (18) as a new distribution function and solve
the system (15), (16) for e1f1 + e2f2,
f3,
. . . ,
fN e1f1 + e2f2,
f3,
. . . ,
fN and the fields E and B, so the number of equations is reduced by one. and the fields E and B, so the number of equations is reduced by one. 5
Multi-component collisionless plasma Let us consider N-component collisionless plasma. In this case the Vlasov–Maxwell integro-
differential system of equations holds: ∂fα
∂t + v∂fα
∂r + eα
mα
E + 1
c[v × B]
∂fα
∂v = 0,
(15)
∇× E + 1
c
∂B
∂t = 0,
∇· E = 4πρ,
∇× B = 1
c
∂E
∂t + 4π
c j,
∇· B = 0,
(16) (15) 1
c
∂B
∂t = 0,
∇· E = 4πρ,
∇× B = 1
c
∂E
∂t + 4π
c j,
∇· B = 0,
(16) (16) where fα(t, r, v) is the distribution function of the α−th component of the plasma, α = 1, . . . , N. Charge and mass of particles of the αth component are denoted by eα and mα, respectively. Charge and current densities have the form where fα(t, r, v) is the distribution function of the α−th component of the plasma, α = 1, . . . , N. Charge and mass of particles of the αth component are denoted by eα and mα, respectively. Charge and current densities have the form ρ =
N
X
α=1
eα
Z ∞
−∞
fαdv,
j =
N
X
α=1
eα
Z ∞
−∞
vfαdv. (17) (17) Let α = 1 and α = 2 correspond to plasma components with equal charge to mass ratio of
particles, for example, alpha particles and deuterium ion components, so we assume (and it is
a good approximation: e1
m1
= e2
m2
. e1
m1
= e2
m2
. According to (17), these components enter the Maxwell equations only as the sum According to (17), these components enter the Maxwell equations only as the sum e1f1 + e2f2,
(18) e1f1 + e2f2,
(18) (18) e1f1 + e2f2, V.B. Taranov 6 V.B. Taranov V.B. Taranov which is in fact the distribution function of the charge density of the components α = 1 and
α = 2. In addition, according to (15), the functions f1, f2 and e1f1 + e2f2 satisfy the same
Vlasov equation. So the transformation [8] which is in fact the distribution function of the charge density of the components α = 1 and
α = 2. In addition, according to (15), the functions f1, f2 and e1f1 + e2f2 satisfy the same
Vlasov equation. 6
Conclusions Lie point symmetries of electron magnetohydrodynamics are obtained (Section 2), which include
time and space homogeneity, self similarity and rotations. As a consequence of the symmetries,
simple invariant solution (1) exists which is in fact the background solution which allows to
explore nonlinear waves evolution by a perturbation formalism. Rich conditional symmetries
are possible in this model due to additional conditions like (3). Lie point symmetries of the integro differential Vlasov–Maxwell model for collisionless elec-
tron plasma obtained in [3] by the moments’ method, i.e. from the symmetry of an infinite set
of partial differential equations for the moments of the distribution functions, are presented in
Section 3. It is shown in Section 4 that hydrodynamic Hasegawa–Mima model for drift waves in magne-
tized plasma, which, in general, is not self similar, has exact asymptotic self similar solutions (14). It is possible due to the conditional symmetry of the model as a consequence of an additional
condition (11). The physical reason for this symmetry extension is that nonlinear waves have
a simpler shape near their critical points – nodes, crests etc. In Section 5, by simple vector considerations, it is shown that multi-component collisionless
plasma containing components with equal charge to mass ratio of particles is invariant under
the transformation (19), if we perform the summation first and the integration later in the
expressions for charge and current densities (17). This transformation, called the extended
symmetry, does not preserve, in general, the global conditions of non negativity of distribution
functions and existence of their moments. So, in contrast with the conditional symmetries, this
symmetry extension is a consequence not of imposing additional conditions, but of neglecting Symmetry Extensions in the Kinetic and Hydrodynamic Plasma Models 7 some global constraints. Nevertheless, among the extended symmetries (19) we can find the
simplifying transform (20), which allows us to reduce the number of equations by 1. The physical reason for the extended symmetry (19) is that particles with equal charge to
mass ratios under the same initial conditions (coordinate and velocity) in a given electric and
magnetic fields move along the same trajectories. The second physically important reason is
that we consider very fast evolution of a plasma, faster than any collisions. References [1] Ibragimov N.H., Kovalev V.F., Pustovalov V.V., Symmetries of integro-differential equations: a survey of
methods illustrated by the Benney equation, Nonlinear Dynam. 28 (2002), 135–165, math-ph/0109012. [2] Cicogna G., Ceccherini F., Pegoraro F., Applications of symmetry methods to the theory of plasma physics,
SIGMA 2 (2006), 017, 17 pages, math-ph/0602008. [3] Taranov V.B., On the symmetry of one-dimensional high frequency motions of a collisionless plasma, Sov. J. Tech. Phys. 21 (1976), 720–726. [4] Gordeev A.V., Kingsep A.S., Rudakov L.I., Electron magnetohydrodynamics, Phys. Rep. 243 (1994), 215–
465. [5] Meleshko S.V., Application of group analysis in gas kinetics, in Proc. Joint ISAMM/FRD Inter-Disciplinary
Workshop “Symmetry Analysis and Mathematical Modelling”, 1998, 45–60. [6] Horton W., Drift waves and transport, Rev. Modern Phys. 71 (1999), 735–778. [7] Taranov V.B., Drift and ion-acoustic waves in magnetized plasmas, symmetries and invari
Ukrainian J. Phys. 49 (2004), 870–874. ] Taranov V.B., Symmetry extensions in kinetic and hydrodynamic plasma models, in Proceedin
International Congress on Plasma Physics (2006, Kyiv), 2006, A041p, 4 pages. [9] Cicogna G., Laino M., On the notion of conditional symmetry of differential equations, Rev. Math. Phys. 18 (2006), 1–18, math-ph/0603021.
|
https://openalex.org/W2230507077
|
https://seer.ufs.br/index.php/revtee/article/download/7512/6018, https://dialnet.unirioja.es/descarga/articulo/8640964.pdf, https://seer.ufs.br/index.php/revtee/article/download/16043/11935, https://seer.ufs.br/index.php/revtee/article/download/10320/7903
|
es
|
Lista de Pareceristas Avaliadores - 2017
|
Revista Tempos e Espaços em Educação
| 2,017
|
cc-by
| 340
|
REVISTA TEMPOS E ESPAÇOS EM EDUCAÇÃO
AVALIADORES AD HOC 2017
Alice Copetti Dalmaso (UFSM)
Amália Cristina Dias da Rocha Bezerra (UERJ)
Ana Lucia Pereira (UEPG)
André Luis Castro de Freitas (UFRN)
Andreia Aparecida Guimarães Strohschoen (UNIVATES)
Anne Alilma Silva Souza Ferrete (UFS)
Antônio Amorim (UNEB)
Antônio Pereira (UNEB)
Antônio Roberto Xavier (UNILAB)
Bréscia França Nonato (UFMG)
Carlos Nogueira Fino (Universidade da Madeira – Portugal)
Cassia Hack (UNIFAP)
Cláudia Moraes (UFSC)
Cláudio Pinto Nunes (UESB)
Claudia Tavares Amaral (UFG)
Célia Souza da Costa (UNIFAP)
Cristiano Feronatto (UNIT)
Cristiano Mezzaroba (UFS)
Eduardo Galak (Universid Nacional de La Plata – Argentina)
Eimard Gomes Antunes do Nascimento (Universidade de Aveiro – Portugal)
Ezer Wellington Gomes Lima (UNIVASF)
Fabiana Sena (UFPB)
Gabriella Eldereti Machado (UFSM)
Galdino Sousa (UFES)
Gilberto Ferreira da Silva (UNILASALLE)
Giovani Ferreira Bezerra (UFMS)
Gislene de Araújo Alves (IFRN)
Gustavo Roese Sanfelice (FEEVALE)
Horácio Miguel Zapata (UNCAUS/UNNE/ISFDPAG)
Jadson Justi (UFAM)
Jean Carlos Miranda (UFF)
Jenerton Arlan Schütz (UNIJUI)
Joanice Conceição (UFF)
Joaquim Tavares da Conceição (UFS)
João Manuel de Oliveira (Instituto Universitário de Lisboa – Portugal)
Jonathas Chaguri (Governo do Estado do Paraná)
Lyana Virgínia Thédiga de Miranda (UFSC)
Lucas Romário (UFPB)
Lúcia Gracia Ferreira (UFRB)
Lucken Bueno Lucas (UENP)
Marcio Romeu Ribas de Oliveira (UFRN)
Maria do Mar Ferreira (University of Warwick – Reino Unido)
Maria Helena Santana Cruz (UFS)
Maria Gouvêa Campesato (Prefeitura Municipal de Porto Alegre / RS)
Mariana Mendonça Lisboa (UFSC)
Marta Maria de Araújo (UFRN)
Michelle Carreirão Gonçalves (UFRJ)
Nilma Margarida de Castro Crusoé (UESB)
Núbia Regina Moreira (UESB)
Osmar Hélio Alves Araújo (UFC)
Paulo Gerson Rodrigues Stefanello (UFSCar)
Paulo Sergio Marchelli (UFS)
Raphael Feitosa (UFC)
Reginaldo Santos Pereira (UESB)
Rita de Cássia Santos Souza (UFS)
Rodrigo Duarte Ferrari (UFSC)
Ronaldo Santos Santana (Secretaria Estadual de Educação de São Paulo/ SP)
Rosana Carla Nascimento Givigi (UFS)
Rosaria Nakashima (UFT)
Sérgio Dorenski Dantas Ribeiro (UFS)
Sílvia Ester Orrú (UNB)
Simone Ferreira de Lucena (UFS)
Tarcia Regina Silva (UFPE)
Verônica Werle (UFSC)
William Timóteo (UNIT)
Wilma Amâncio da Silva Amâncio da Silva (Secretaria Municipal de Educação de
Delmiro Gouveia/AL)
|
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https://openalex.org/W2890832593
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https://openresearch.surrey.ac.uk/view/delivery/44SUR_INST/12139765920002346/13140461640002346
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English
| null |
Metal Ion Interactions with Crude Oil Components: Specificity of Ca2+ Binding to Naphthenic Acid at an Oil/Water Interface
|
Colloids and interfaces
| 2,018
|
cc-by
| 16,184
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Received: 17 August 2018; Accepted: 14 September 2018; Published: 18 September 2018 Abstract: On the basis of dynamic interfacial tension measurements, Ca2+ has been shown specifically
to interact with naphthenic acid (NA) at the n-heptane/water interface, consistent with NA adsorption
followed by interfacial complexation and formation of a more ordered interfacial film. Optimum
concentrations of Ca2+ and NA have been found to yield lower, time-dependent interfacial tensions,
not evident for Mg2+ and Sr2+ or for several alkali metal ions studied. The results reflect the specific
hydration and coordination chemistry of Ca2+ seen in biology. Owing to the ubiquitous presence of
Ca2+ in oilfield waters, this finding has potential relevance to the surface chemistry underlying crude
oil recovery. For example, “locking” acidic components at water/oil interfaces may be important for
crude oil emulsion stability, or in bonding bulk oil to mineral surfaces through an aqueous phase,
potentially relevant for carbonate reservoirs. The relevance of the present results to low salinity
waterflooding as an enhanced crude oil recovery technique is also discussed. Keywords: dynamic interfacial tension; interfacial complexation; low salinity waterflooding;
metal ion interactions; naphthenic acid; oil recovery Article
Metal Ion Interactions with Crude Oil Components:
Specificity of Ca2+ Binding to Naphthenic Acid at
an Oil/Water Interface Spencer E. Taylor *
and Hiu Tung Chu
Centre for Petroleum and Surface Chemistry, Department of Chemistry, University of Surrey, Guildford,
Surrey GU2 5XH, UK; tungchi0906@gmail.com
* Correspondence: s.taylor@surrey.ac.uk; Tel.: +44-1483-681999 Spencer E. Taylor *
and Hiu Tung Chu
Centre for Petroleum and Surface Chemistry, Department of Chemistry, University of Surrey, Guildford,
Surrey GU2 5XH, UK; tungchi0906@gmail.com
* Correspondence: s.taylor@surrey.ac.uk; Tel.: +44-1483-681999 Spencer E. Taylor *
and Hiu Tung Chu
Centre for Petroleum and Surface Chemistry, Department of Chemistry, University of Surrey, Guildford,
Surrey GU2 5XH, UK; tungchi0906@gmail.com
* Correspondence: s.taylor@surrey.ac.uk; Tel.: +44-1483-681999 Spencer E. Taylor *
and Hiu Tung Chu
Received: 17 August 2018; Accepted: 14 September 2018; Published: 18 September 2018 colloids
and interfaces colloids
and interfaces colloids
and interfaces colloids
and interfaces www.mdpi.com/journal/colloids 1. Introduction Carboxylic acids are fundamental to the chemistry of all crude oils [1]. These are oil-soluble
compounds with the general formula CnH2n+zO2 containing acyclic (z = 0) and alicyclic (z > 0)
hydrocarbyl groups; in crude oils, they are collectively referred to as ‘naphthenic acid’ (NA), since the
earliest examples identified contained saturated cyclopentyl and cyclohexyl groups [2,3]. Carboxylic acids are fundamental to the chemistry of all crude oils [1]. These are oil-soluble
compounds with the general formula CnH2n+zO2 containing acyclic (z = 0) and alicyclic (z > 0)
hydrocarbyl groups; in crude oils, they are collectively referred to as ‘naphthenic acid’ (NA), since the
earliest examples identified contained saturated cyclopentyl and cyclohexyl groups [2,3]. As is the case for other polar species present in petroleum, organic acids are often implicated
in various upstream and downstream operational problems. Thus, during crude oil production,
the surface activity of NA can lead to alterations in reservoir wettability [4,5] and increased stability of
water-in-oil (w/o) emulsions [6]. Additionally, under high pH conditions, ionized acid groups stabilize
oil-in-water (o/w) emulsions [7] when dissociation of oil-soluble acids produces anionic surface-active
species which strongly affect properties, such as interfacial tension and interfacial rheology [8]. During crude oil refining, the acidity of NA (pKa ~5 [9]) is responsible for corrosion in distillation
units under the high temperature process conditions. The total acidity of the crude oil depends on its
source, and oil-specific measures are often required to reduce potential damage when processing the
highest acidity crudes (typically total acid number [TAN] values in excess of approximately 0.8 mg
KOH/g) [10]. Naphthenic acids also exhibit specific interactions with calcium in certain oilfields,
where Ca2+ coordination leads to precipitation and deposition of naphthenate-rich scale [11]. In this
case, the naphthenates are tetrameric species, the original C80-tetra-acid member being identified
as C80H142O8, molecular weight 1230 g/mol [12]. Following this discovery, other similar members Colloids Interfaces 2018, 2, 40; doi:10.3390/colloids2030040 www.mdpi.com/journal/colloids 2 of 21 2 of 21 Colloids Interfaces 2018, 2, 40 were identified as being responsible for naphthenate deposits in diverse locations [13,14]. Regarding
the formation of these species, two points are noteworthy. First, if the tetra-acids are uncharged, it is
likely that interactions must occur at an aqueous/oil interface. Second, from the composition of the
deposits, there should be a specific affinity between the organic acid and aqueous Ca2+ ions, given the
mixed cation composition of oilfield waters. 1. Introduction This is also confirmed using a range of techniques and a
synthetic tetra-acid, for which the strongest affinity was found for Ca2+ [15,16]. There is evidence in the literature to show that acidic species contribute to the structure and
properties of crude oil–water interfaces [17]. Dissociation of carboxylic acids, for example, explains
the pH-dependent behavior of crude oil/water interfaces [18,19] leading to increased o/w emulsion
stability at high pH. It is also evident that crude oil/water interfacial properties can be significantly
influenced by the presence of divalent metal ions. Thus, Alvarado et al. [20] reported the increased
stabilization of water-in-crude oil emulsions in the presence of Ca2+ ions compared with Na+ ions of
the same ionic strength, suggesting that stronger interactions occur with Ca2+, although the authors
suggested that further investigation is necessary [21]. Carboxylic acid groups at crude oil interfaces have also been implicated in improved recovery
mechanisms during low salinity waterflooding (involving the so-called “low salinity effect”, LSE). One popular suggested LSE mechanism starts with the premise that oil is naturally bound to
specific reservoir mineral surfaces via divalent metal ion bridging involving oil-based carboxylate
groups [22–30]. It is considered that reducing the water salinity disrupts the bridges through ion
exchange, resulting in reduced oil adhesion, thereby improving oil displacement and recovery. Other studies into the LSE have suggested that interfacial properties should be sensitive to
complexation across oil/water interfaces [31,32]. Indeed, the specificity of interfacial complexation
between metal ions and oil-soluble components is important elsewhere in liquid–liquid extraction
processes [33]. Early work [34–36] identified that interfacial reactions involving metal ions and spread
carboxylic acid monolayers at the air-water interface are relatively slow, with half-lives of several
minutes. More recently, de Ruiter et al. studied the evolution of the interfacial tension upon placing
aqueous solutions of metal ions in contact with a decane solution of stearic acid (HSt) [31]. When the
solution pH > pKa, the acidity constant of stearic acid, distinct stages in the formation of metal stearate
layers were noted, and interpreted as [31]: (i) deprotonation of adsorbed stearic acid (HSt), forming
MSt2 complexes at the interface; (ii) nucleation of the MSt2 complexes, allowing further acid adsorption
and complexation; (iii) partitioning of the three-dimensional MSt2 structures into the oil phase. 1. Introduction The present study is concerned with the influence of alkali and alkaline earth metal ions (as the
respective chloride salts) on oil/water interfaces containing a commercial (technical-grade) sample of NA. Previous work by Brandal et al. investigated the effects of divalent metal ions (Mg2+, Ca2+, Sr2+ and Ba2+)
on the interfacial tension behavior of different carboxylic acids under pH conditions where the acids were
completely ionized [32]. However, we were keen to explore interactions between aqueous metal ions and
the oil-soluble NA, initially in its unionized form as would be present in a native crude oil. 2.2.1. Synthesis of Calcium Naphthenate Calcium naphthenate was synthesized based on a method used by Pereira et al. to prepare
calcium salts of long-chain carboxylates [38]. Thus, KOH (0.752 g, 13.0 mmol) was dissolved in ethanol
(20 mL) and heated gently on a hot-plate until boiling. NA (3.270 g, ~3.3 mmol based on the above TAN
value) was then added and boiling maintained. Separately, Ca(OAc)2·xH2O (1.062 g, ~6.5 mmol) was
dissolved in the minimum quantity of water with sonication, after which this solution was carefully
added dropwise to the boiling ethanolic KOH/NA. The resultant mixture was then allowed to cool and
evaporate, whereupon an amber-colored gel separated, together with residual water (~2 mL). The water
was decanted, and the gel product was washed with an approximately equal volume of deionized
water, heated to 80 ◦C, and then allowed to cool. Again, the residual water was decanted. This washing
process was repeated one further time. The drained product was then stirred with petroleum ether
(40–60 ◦C boiling range) until dissolved (~40 mL required). This solution was filtered (Whatman
#1 filter paper), which also served to remove residual water from the resultant clear amber filtrate. Solvent evaporation yielded viscous product (3.079 g, 87%). The stoichiometry of the product was
determined by thermogravimetric analysis (TA Instruments TGA Q500 thermogravimetric analyzer,
Elstree, Hertfordshire, UK). The sample was heated from ambient temperature to 900 ◦C at 10 ◦C/min
in air to yield a residue (calcium oxide) corresponding to 7.14% Ca. Using the average NA molecular
mass of 243 calculated above, the product has a composition consistent with the expected 1:2 Ca:NA
stoichiometry. Figure 1 shows a comparison of the infrared spectra (Bruker Alpha spectrometer with
Diamond ATR (Bruker UK Ltd., Coventry, UK), each sample applied in neat form) for the original
naphthenic acid and its calcium salt, from which the shift of the carbonyl band from 1700 to 1543 cm−1
is characteristic of neutralized carboxylic acids [39,40]. The small band at 1709 cm−1 in the calcium
naphthenate spectrum indicates a trace amount of unreacted naphthenic acid remains in the product. Loss of bands at 1265 and 943 cm−1 are also consistent with calcium salt formation [40]. Colloids Interfaces 2018, 2, x
4 of 21 Figure 1. Attenuated Total Reflection (ATR)-infrared spectra of naphthenic acid (red, dashed) and its
calcium salt (blue, continuous). Figure 1. Attenuated Total Reflection (ATR)-infrared spectra of naphthenic acid (red, dashed) and its
calcium salt (blue, continuous). 2.1. Materials Organic solvents (ethanol and n-heptane, both >98% purity) and the reagents naphthenic
acid (technical grade), calcium chloride dihydrate (CaCl2·2H2O), strontium chloride hexahydrate
(SrCl2·6H2O), magnesium chloride hexahydrate (MgCl2·6H2O), sodium chloride (NaCl), potassium
chloride (KCl), lithium chloride (LiCl), cesium chloride (CsCl), potassium hydroxide (KOH), calcium
acetate (Ca(OAc)2·xH2O) were purchased from Sigma-Aldrich Co. Ltd. (Gillingham, Dorset, UK)
and apart from n-heptane were used without further purification. The latter was passed through an
alumina column to remove any surface-active impurities before use. According to the supplier, the commercial naphthenic acid is a technical mixture of alkylated
cyclopentane carboxylic acids, with a total acid number (TAN) value of 230.5 mg KOH/g, 3 of 21 3 of 21 Colloids Interfaces 2018, 2, 40 which equates to an average molecular mass of 56/0.2305 = 243 g/mol with the reasonable assumption
that the mixture only contains monoprotic acids. However, consistent with our own analysis,
Hindle et al. reported that this product contains a high proportion of saturated fatty acids, with
relatively low levels of true naphthenic components [37]. Notwithstanding the relatively low content
of authentic naphthenic structures, its composition does provide a representative range of carbon chain
lengths to investigate interactions which mainly involve the carboxylic acid function of crude oils. g
g
y
y
Deionized water (resistivity = 18.2 MΩ·cm) was from a Millipore Direct-Q system, with a pH of
~5.6 at room temperature (21 ◦C). 2.2.1. Synthesis of Calcium Naphthenate Figure 1. Attenuated Total Reflection (ATR)-infrared spectra of naphthenic acid (red, dashed) and its
calcium salt (blue, continuous). Figure 1. Attenuated Total Reflection (ATR)-infrared spectra of naphthenic acid (red, dashed) and its
calcium salt (blue, continuous). 4 of 21 Colloids Interfaces 2018, 2, 40 2.2.2. Dynamic Interfacial Tension Measurements 2.2.2. Dynamic Interfacial Tension Measurements The dynamic tension of water/n-heptane interfaces was determined using an in-house dynamic
drop volume (DDV) method. The general basis of this approach has been detailed by van Hunsel and
Joos [41] and is similar to the method used by Deshiikan et al. [42]. Briefly, aqueous drops are formed
at the tip of a capillary using different (known) flow rates, f, and the corresponding drop formation
times, td, measured. By knowing f and td allows corresponding drop volumes to be determined
under dynamic conditions. From a comparison with the Ward-Tordai equation, the adsorption time
is ~3/7td [43]. Strictly, drop volume (or weight) methods are used to determine equilibrium surface
or interfacial tension for drops formed infinitely slowly, because at finite rates of drop formation
hydrodynamic effects cause excess liquid to be incorporated into the detaching drop [41]. However,
under dynamic conditions, the drop volume, V(td), is related to the drop formation time by the
empirical relationship attributed to McGee (see ref. [41]): V(td) = V0 + St−0.75
d
(1) (1) in which V0 is the drop volume at an infinitely slow formation time and S is the so-called McGee
slope [41]. The latter is determined by measuring drop volumes at different flow rates for a series of
pure liquids, under conditions of constant interfacial tension. Surface or interfacial tensions are then
obtained using the established static pendant drop volume relationship: in which V0 is the drop volume at an infinitely slow formation time and S is the so-called McGee
slope [41]. The latter is determined by measuring drop volumes at different flow rates for a series of
pure liquids, under conditions of constant interfacial tension. Surface or interfacial tensions are then
obtained using the established static pendant drop volume relationship: γ = V0∆ρg
2πrF
(2) (2) in which r is the external tip radius, ∆ρ the density difference between the two phases, and F is a
correction factor accounting for drop shape which was determined and tabulated as a function of
r/V01/3 by Harkins and Brown [44] and extended by Wilkinson [45]. The more recent polynomial
developed by Lee et al. [46] shown in Equation (3), which correlates F with r/V01/3 within the range
0 ≥r/V01/3 ≥1.2, was used in the present analysis. 2.2.2. Dynamic Interfacial Tension Measurements in which r is the external tip radius, ∆ρ the density difference between the two phases, and F is a
correction factor accounting for drop shape which was determined and tabulated as a function of
r/V01/3 by Harkins and Brown [44] and extended by Wilkinson [45]. The more recent polynomial
developed by Lee et al. [46] shown in Equation (3), which correlates F with r/V01/3 within the range
0 ≥r/V01/3 ≥1.2, was used in the present analysis. F = 1 −0.9121(r/V01/3) −2.109(r/V01/3)2 + 13.38(r/V01/3)3 −27.29(r/V01/3)4 +
27.53(r/V01/3)5 −13.58(r/V01/3)6 + 2.593(r/V01/3)7
(3) (3) By using pure low viscosity liquids of constant surface tension over the range of drop formation
times, hydrodynamic contributions to drop formation are effectively eliminated, which allows the
relationship between the McGee slope and V0 to be defined. For our system, we found that the McGee
slope is described well by S = 0.22V0
(4) (4) such that for a given drop time, Equation (2) becomes: such that for a given drop time, Equation (2) becomes: uch that for a given drop time, Equation (2) becomes: γ(td) =
V(td)∆ρg
2πrF
1 + 0.22t−0.75
d
(5) (5) In the present method, water or aqueous salt solution drops were formed below the surface of
the n-heptane phase from the tip of a stainless steel syringe needle, the end of which was cleaned
and roughened with a fine grade sand paper before each measurement to ensure effective contact
of the aqueous phase is made with the outer diameter (3.413 mm). Drop volumes were in the range
~30–120 mm3. The pH of the unadjusted aqueous phases was in the range 5.4–5.7 consistent with
Robertson et al. [47]. Drops are then produced at up to ten known rates using a pre-calibrated syringe
pump (B. Braun, Melsungen, Germany) at 21 ± 1 ◦C, and the drop times calculated by averaging
the time taken for several drops to be produced. Drop counting was performed visually with an 5 of 21 Colloids Interfaces 2018, 2, 40 estimated mechanical (stopwatch) response time of ±0.02 s for a single drop; this limits the realistic
rate of formation typically to ~1 drop per second. The procedure also requires the density difference between the liquid phases to be known under
the same temperature conditions, and these were determined by pycnometry from the mass of a known
volume of the respective liquids. 3. Results and Discussion In advance of discussing the results of this study, it is pertinent to consider the ionization behavior
of carboxylic acids at oil/water interfaces. Thus, on electrostatic grounds it would be expected that
ionization equilibria will be affected by the presence of an interface, which would then be reflected in
the relevant dissociation constants. To illustrate this, if the acid dissociation constant of a carboxylic
acid in bulk (aqueous) solution is pKa(∞), then the electrostatic image potential Ψ experienced by the
acid group at a distance x from the interface is given by [49]: Ψ(x) = −
Ce
2Dwx
(6) (6) where C = 1/4πε0, e is the electronic charge (1.602 × 10−19 C), ε0 is the permittivity of a vacuum
(8.854 × 10−12 F/m), and Dw is the dielectric constant of water (~80). By further considering the acid-base equilibrium as a function of distance from the interface, Dill
and Bromberg [49] showed that the apparent pKa at a distance x from the interface is related to the
bulk value by pKa(x) = pKa(∞) −0.4343eΨ(x)
kBT
= pKa(∞) + 0.4343Ce2
2xDwkBT
(7) (7) in which kB is Boltzmann’s constant (1.381 × 10−23 J/K) and T is the absolute temperature. This simple
model suggests that adsorbed acid groups within, say, 0.2–0.5 nm of an oil/water interface, would
experience an increase in pKa of 0.3–0.8 units, purely on electrostatic grounds, without introducing
specific ionic refinements. However, by using more sophisticated computational approaches, Andersson et al. found that the
pKa of isolated acid molecules at an oil/water interface increased by an average of ~1 unit compared
with the aqueous bulk solution value [50]. This reflects the greater affinity of the neutral molecule
for the interface. The consequences of molecular confinement, at an interface, for example, were also
probed using the same computational methods. In this situation, the possibility of deprotonated
carboxylate groups stabilizing neighboring undissociated (neutral) carboxylic acid molecules arises
through hydrogen bonding. Thus, the same workers also showed how the pKa shift produced in the
first molecule, as described above, is almost exactly compensated by the stabilizing effect induced on
the second molecule [50]. 2.2.2. Dynamic Interfacial Tension Measurements Thus, by measuring td at different flow rates allows the determination
of γ(td) as a function of drop time using Equation (5). The combination of interfacial tension, drop
volume, needle diameter and density difference translates for most of the measurements into values of
the newly proposed Worthington number (Wo = ∆ρgVd/2πrγ) in the range ~0.7–0.9, for which a value
of unity would be the ideal situation [48]. Additionally, replicate measurements using pure liquids
with time-invariant interfacial tension, indicate a standard error of <3% of the quoted values, which is
often better than ± 1 mN/m. 3.1. NA Adsorption at the n-Heptane/Water Interface
Dynamic drop volume measurements betwee
d
f
l
d
f 3.1. NA Adsorption at the n-Heptane/Water Interface
Dynamic drop volume measurements betwee
d
f
l
d
f Dynamic drop volume measurements between n-heptane containing NA and deionized water
produce constant interfacial tensions over drop formation times in the approximate range ~0.3–500 s
as shown in Figure 2a. The time-invariant values seen for different bulk NA concentrations (CNA) are
due to the changes in the composition of the n-heptane phase, reducing slightly from the value for
the n-heptane/water interface. The results also indicate that adsorption is rapid compared with the
experimental timescale, being complete in <0.3 s. Using a microfluidic technique, adsorption at the
hydrocarbon/water interface has recently been shown [55] to occur on the millisecond timescale, more
rapidly than previously thought. roduce constant interfacial tensions over drop formation times in the approximate range ~0.3–500
s shown in Figure 2a. The time-invariant values seen for different bulk NA concentrations (CNA) ar
ue to the changes in the composition of the n-heptane phase, reducing slightly from the value fo
he n-heptane/water interface. The results also indicate that adsorption is rapid compared with th
xperimental timescale, being complete in <0.3 s. Using a microfluidic technique, adsorption at th
ydrocarbon/water interface has recently been shown [55] to occur on the millisecond timescale
more rapidly than previously thought. Figure 2. (a) Examples of the dynamic interfacial tension of naphthenic acid solutions in n-heptane
against deionized water. (b) Corresponding results against water containing 0.3 mol/L Ca2+. (c)
Interfacial tension-log (naphthenic acid concentration) profiles for deionized water and the
individual γi and γ∞ contributions for 0.3 mol/L Ca2+ (see Equation (8)). The inset shows the
corresponding plot for n-heptane solutions of Ca(NA)2 against deionized water. All curves are fits to
the Szyszkowski equation (Equation (9)). (d) The effect of Ca2+ concentration on the interfacial
tension of the n-heptane/water interface. The line is based on the empirical equation γ = γ0 − Aγ0.5 +
Figure 2. (a) Examples of the dynamic interfacial tension of naphthenic acid solutions in n-heptane
against deionized water. (b) Corresponding results against water containing 0.3 mol/L Ca2+. (c) Interfacial tension-log (naphthenic acid concentration) profiles for deionized water and the
individual γi and γ∞contributions for 0.3 mol/L Ca2+ (see Equation (8)). The inset shows the
corresponding plot for n-heptane solutions of Ca(NA)2 against deionized water. All curves are fits to
the Szyszkowski equation (Equation (9)). 3. Results and Discussion Several experimental studies have identified increased pKa values for acid groups present at
surfaces and interfaces, including self-assembled monolayers [51], fatty acid micelles [52,53] and, more
recently, at the air/water interface for fatty acid monolayers [54], the latter study also identifying the
effect of chain length on the magnitude of the pKa change. In the context of the present paper, this will
mean that adsorbed NA molecules will remain largely undissociated as the effective (interfacial) pKa is
higher than the pH of the aqueous solutions. Colloids Interfaces 2018, 2, 40
(
) p
1 NA Ad
ti
t th 6 of 21 6 of 21 3.1. NA Adsorption at the n-Heptane/Water Interface
Dynamic drop volume measurements betwee
d
f
l
d
f (d) The effect of Ca2+ concentration on the interfacial tension
of the n-heptane/water interface. The line is based on the empirical equation γ = γ0 −Aγ0.5 + Bγ as
described in the text. Figure 2. (a) Examples of the dynamic interfacial tension of naphthenic acid solutions in n-heptane
gainst deionized water. (b) Corresponding results against water containing 0.3 mol/L Ca2+. (c)
nterfacial tension-log (naphthenic acid concentration) profiles for deionized water and the
ndividual γi and γ∞ contributions for 0.3 mol/L Ca2+ (see Equation (8)). The inset shows the
orresponding plot for n-heptane solutions of Ca(NA)2 against deionized water. All curves are fits to
he Szyszkowski equation (Equation (9)). (d) The effect of Ca2+ concentration on the interfacial
ension of the n-heptane/water interface. The line is based on the empirical equation γ = γ0 − Aγ0.5 +
Figure 2. (a) Examples of the dynamic interfacial tension of naphthenic acid solutions in n-heptane
against deionized water. (b) Corresponding results against water containing 0.3 mol/L Ca2+. (c) Interfacial tension-log (naphthenic acid concentration) profiles for deionized water and the
individual γi and γ∞contributions for 0.3 mol/L Ca2+ (see Equation (8)). The inset shows the
corresponding plot for n-heptane solutions of Ca(NA)2 against deionized water. All curves are fits to
the Szyszkowski equation (Equation (9)). (d) The effect of Ca2+ concentration on the interfacial tension
of the n-heptane/water interface. The line is based on the empirical equation γ = γ0 −Aγ0.5 + Bγ as
described in the text. Bγ as described in the text. Compared with the time-invariant interfacial tension behavior seen at the deionized water
interface, the presence of 0.3 mol/L Ca2+ is accompanied by the appearance of time-dependent
behavior, as shown in Figure 2b for equivalent naphthenic acid concentrations. Bγ as described in the text. Compared with the time-invariant interfacial tension behavior seen at the deionized water
interface, the presence of 0.3 mol/L Ca2+ is accompanied by the appearance of time-dependent
behavior, as shown in Figure 2b for equivalent naphthenic acid concentrations. These latter profiles suggest that the approach to equilibrium of a growing drop in the presence
of Ca2+ occurs in (at least) two stages. 3.1. NA Adsorption at the n-Heptane/Water Interface
Dynamic drop volume measurements betwee
d
f
l
d
f The first stage involves a rapid γ decrease, as described above,
as the interface re-equilibrates from γ0 ~48 mN/m to an “instantaneous” value (within the timescale Colloids Interfaces 2018, 2, 40 7 of 21 of the present technique), γi, which, for 5 mg/L ≤CNA ≤200 mg/L and 0.3 mol/L Ca2+, is lower than
the corresponding values obtained for the deionized water interface. of the present technique), γi, which, for 5 mg/L ≤CNA ≤200 mg/L and 0.3 mol/L Ca2+, is lower than
the corresponding values obtained for the deionized water interface. However, it is also evident in Figure 2b that within approximately the same CNA range given
above, the interfacial tension behavior exhibits a slower second process which finally results in an
equilibrium value, γ∞, i.e., γ0 →γi, followed by γi →γ∞. The interfacial tension curves for the slower
process, as shown in Figure 2b, are satisfactorily represented by first-order kinetic curves constructed
according to Equation (8), each of which being characterized by a first-order rate constant, k. γ = γ∞+ (γi −γ∞) exp(−kt)
(8) (8) Since γ0 for each Ca2+ solution in the presence of NA is inaccessible using the present technique,
we felt that it is reasonable to assume that it will be the same as the time-invariant value for the
corresponding n-heptane/0.3 mol/L Ca2+ interface in the absence of NA. Thus, in Figure 2c are shown plots of γi and γ∞for a Ca2+ concentration, CCa of 0.3 mol/L as
a function of CNA, together with the corresponding γ∞behavior at the deionized water interface,
the latter being in good agreement with data published by Havre et al. [9] for a similar commercial NA
sample. It is immediately apparent that the interfacial tension decreases in the presence of 0.3 mol/L
Ca2+ in the aqueous phase for ~5 mg/L ≤CNA ≤200 mg/L, and increases once again above this
concentration, towards the deionized water isotherm. In Figure 2c (inset), we also show interfacial tension data for the synthesized calcium salt, Ca(NA)2. In this case, only very small time-dependent effects were observed, the error bars reflecting the spread
of results over the timescale of the measurements. However, it is evident that the calcium salt follows
the same general behavior as NA, inasmuch as the interfacial tension is largely unaffected up to
~100 mg/L, but thereafter the data diverge. 3.1. NA Adsorption at the n-Heptane/Water Interface
Dynamic drop volume measurements betwee
d
f
l
d
f Within the NA concentration range considered, we find that the interfacial tension-log profiles in
ure 2c are satisfactorily represented by the Szyszkowski equation [56]: γ = γ0 −RTΓm ln
CNA
a
+ 1
(9) (9) where CNA is the bulk NA concentration in the organic phase, Γm is the interfacial concentration of
NA at monolayer adsorption (mol/m2) and a is a molecule-dependent constant (expressed as mol/L)
related to the Gibbs energy of adsorption (∆G0) at infinite dilution by [57]: a = 55.3 exp
∆G0
RT
(10) (10) From Table 1 it is apparent that the “instantaneous” interfacial concentration of NA in the presence
of Ca2+ is some ten times greater than in its equilibrium concentration. This translates into a much
smaller apparent molecular area. The latter dimension is unreasonable, however, and most probably
relates to a disordered interfacial accumulation of NA molecules in the initial stages, which would be
consistent with the initial rapid response of the interfacial tension. The situation then transforms into
a more structured interfacial film characterized by a more realistic molecular area, which is almost
identical to the value at the deionized water interface. Moreover, the Gibbs energy of adsorption of NA
is ~20 kJ/mol more favorable (i.e., lower) in the presence of Ca2+ ions, which presents an intriguing
insight into the role played by Ca2+ ions in facilitating NA adsorption. Interestingly, this value is
very close to the Gibbs energy for Ca2+ binding to hydrated carboxylate side chains of the protein
β-lactoglobulin (−18 ± 1 kJ/mol) [58]. Whilst the interfacial packing of NA molecules appears to
remain the same, the adsorption process itself is significantly more favorable, which suggests some
degree of interaction occurs between NA and Ca2+ ions across the interface. 8 of 21 Colloids Interfaces 2018, 2, 40 Table 1. Summary of the adsorption parameters for NA in the presence and absence of Ca2+ ions based
on the Szyszkowski equation. Table 1. Summary of the adsorption parameters for NA in the presence and absence of Ca2+ ions based
on the Szyszkowski equation. 3.1. NA Adsorption at the n-Heptane/Water Interface
Dynamic drop volume measurements betwee
d
f
l
d
f Interface Species
Γm (mol/m2)
Molecular Area (nm2)
∆G0 (kJ/mol)
NA
1.58 × 10−6
1.05
−27.9
NA + Ca2+ (i)
1.66 × 10−5
0.10
−27.5
NA + Ca2+ (∞)
1.60 × 10−6
1.04
−48.7
Ca(NA)2
3.34 × 10−6
0.50
−29.1 Finally, as a relevant aside, in Figure 2d are shown the effects of CCa on the n-heptane/water
interfacial tension in the absence of NA. It is well-known that high electrolyte concentrations generally
show an increase in surface and interfacial tension as a consequence of “negative adsorption”. However, relatively low electrolyte concentrations cause a small decrease in tension, a phenomenon
known as the Jones-Ray effect, after the first researchers to report it, in 1937 [59]. While the original
work focused on electrolyte effects on surface tension, some recent studies have considered the behavior
at hydrocarbon/water interfaces [60], including the crude oil/water interface [61], because of their
relevance to some of the applications mentioned earlier in the present paper. In each of these earlier studies, interfacial tension minima have been seen at low CCa, which is
also evident in the present results. Thus, Figure 2d shows that the interfacial tension reduction for
<0.1 mol/L Ca2+ is ~1.4 mN/m, similar to values obtained elsewhere [60]. Suggested reasons for this
effect have been debated, and have included cation [60], anion [62–64] or impurity [65] adsorption,
or electrostatic effects influencing water structure at the interface [66,67]. The curve fitted through the
experimental data in Figure 2d is of the form γ = γ0 −Aγ0.5 + Bγ, where A and B are empirical constants
(0.158 (mN/m)0.5 and 0.217, respectively) and γ0 is the interfacial tension of the n-heptane/water
interface (48.4 mN/m). derived from Equation (13).
3 3 D
d
f I t f
i l T
i
C
d C
3.3. Dependence of Interfacial Tension on CNA and CCa q
(
)
3 3 D
d
f I
f
i l T
i
C
d C
3.3. Dependence of Interfacial Tension on CNA and CCa 3.3. Dependence of Interfacial Tension on CNA and CCa
The theoretical curves drawn through the data in Figure 3b were derived on the assumption
that the interface can be described as a combination of two limiting situations, one based on pure
water and the other containing a certain interfacial excess of Ca2+ (possibly representing a
“saturated” interfacial film). The former condition is simply represented by deionized water data in
Figure 2c, whilst the latter has to be obtained from the limiting interfacial tension data determined in
the presence of different Ca2+ concentrations shown in Figure 3b (red line). As discussed above, both
limiting conditions have been shown to be adequately described by the Szyszkowski equation (see
Figure 2c). In the subsequent analysis, therefore, we have assumed that the overall adsorption
conditions will be a combination of these two limiting cases, the extent to which each contributes
being dependent on the (bulk) concentration ratio between NA and Ca2+. Thus, the interfacial
tensions produced by NA adsorption at a water interface and at a saturated Ca2+-containing interface
are given by Equations (9) and (11), respectively; it is unnecessary to define the bulk Ca2+
concentration responsible for the latter condition at this stage of the analysis. In Equation (11), Гm’
d ’
h
i
f
i
d
l
l
f NA i
h
f h
The theoretical curves drawn through the data in Figure 3b were derived on the assumption
that the interface can be described as a combination of two limiting situations, one based on pure
water and the other containing a certain interfacial excess of Ca2+ (possibly representing a “saturated”
interfacial film). The former condition is simply represented by deionized water data in Figure 2c,
whilst the latter has to be obtained from the limiting interfacial tension data determined in the presence
of different Ca2+ concentrations shown in Figure 3b (red line). As discussed above, both limiting
conditions have been shown to be adequately described by the Szyszkowski equation (see Figure 2c). In the subsequent analysis, therefore, we have assumed that the overall adsorption conditions will be
a combination of these two limiting cases, the extent to which each contributes being dependent on
the (bulk) concentration ratio between NA and Ca2+. 3.2. NA Adsorption at the n-Heptane/Water Interface in the Presence of Metal Ions With the exception of Ca2+, none of the other metal ions studied has been found to exhibit
time-dependent interfacial tensions using the present experimental method. Thus, in Figure 3a,
NA adsorption at the n-heptane/metal chloride solution (1.0 mol/L) interface is indistinguishable
from the deionized water curve shown in Figure 2c. These data include the two other divalent metal ions
studied, Mg2+ and Sr2+, which do not show the same interfacial tension behavior as a function of CNA
as seen for Ca2+, i.e., interfacial tension minima that are dependent on CCa as highlighted in Figure 3b. As observed above, for NA concentrations beyond the minimum interfacial tension, the behavior
returns to the curve defined by the other metal ion solutions and deionized water. This suggests a
reduced influence of Ca2+ ions in lowering the Gibbs energy of adsorption of NA molecules (Table 1)
as the interface becomes more saturated with NA. Crucially, however, it also suggests that the presence
of specific calcium naphthenate species with higher surface activity may not be directly responsible
for the increased interfacial activity of NA, but rather the Ca2+ ions promote NA adsorption, through
specific interactions as will be discussed subsequently. 9 of 21
9 of 21 Colloids Interfaces 2018, 2, 40
Colloids Interfaces 2018, 2, x Figure 3. (a) Interfacial tension-log (naphthenic acid concentration) profiles at the n-heptane/water
interface in the presence of the indicated cations (all chloride salts at 1 mol/L). (b) Corresponding
profiles for different aqueous Ca2+ concentrations. The blue and red curves represent the respective
limiting conditions for deionized water and a “Ca2+-saturated” interface, and the gray curves are
derived from Equation (13). Figure 3. (a) Interfacial tension-log (naphthenic acid concentration) profiles at the n-heptane/water
interface in the presence of the indicated cations (all chloride salts at 1 mol/L). (b) Corresponding
profiles for different aqueous Ca2+ concentrations. The blue and red curves represent the respective
limiting conditions for deionized water and a “Ca2+-saturated” interface, and the gray curves are
derived from Equation (13). Figure 3. (a) Interfacial tension-log (naphthenic acid concentration) profiles at the n-heptane/water
interface in the presence of the indicated cations (all chloride salts at 1 mol/L). (b) Corresponding
profiles for different aqueous Ca2+ concentrations. The blue and red curves represent the respective
limiting conditions for deionized water and a “Ca2+-saturated” interface, and the gray curves are
derived from Equation (13)
Figure 3. 3.2. NA Adsorption at the n-Heptane/Water Interface in the Presence of Metal Ions (a) Interfacial tension-log (naphthenic acid concentration) profiles at the n-heptane/water
interface in the presence of the indicated cations (all chloride salts at 1 mol/L). (b) Corresponding
profiles for different aqueous Ca2+ concentrations. The blue and red curves represent the respective
limiting conditions for deionized water and a “Ca2+-saturated” interface, and the gray curves are
derived from Equation (13). derived from Equation (13).
3 3 D
d
f I t f
i l T
i
C
d C
3.3. Dependence of Interfacial Tension on CNA and CCa The appearance of a plateau
region in Figure 4 at CNA ~ 0.4 mol/L suggests that this bulk concentration corresponds to the
interfacial “saturation” condition for Ca2+ ions, as alluded to earlier, and for which the maximum
influence on NA adsorption is seen. 3 4 NA Ca2+ Interfacial Tension Kinetics derived from Equation (13).
3 3 D
d
f I t f
i l T
i
C
d C
3.3. Dependence of Interfacial Tension on CNA and CCa Thus, the general interfacial tension expression for
different Ca2+ and NA concentrations is given by combining Equations (9) and (11), to give
𝛾= 𝛾0 −𝑅𝑇(𝑚(1 −exp (𝐶Ca
𝐶NA
)) ln (𝐶NA
𝑎+ 1) + 𝑚′exp (𝐶Ca
𝐶NA
) ln (𝐶NA
𝑏+ 1))
(12) γ = γ0 −RT
Γm
1 −exp
φCCa
CNA
ln
CNA
a
+ 1
+ Γm′ exp
φCCa
CNA
ln
CNA
b
+ 1
(12
Initial attempts at fitting the data shown in Figure 3b to Equation (12) indicated that, perha
nsurprisingly, Гm ~ Гm’, allowing the simplified Equation (13) to be used to generate the fitt (12)
erhaps Initial attempts at fitting the data shown in Figure 3b to Equation (12) indicated that, perhaps
unsurprisingly, Γm ~Γm’, allowing the simplified Equation (13) to be used to generate the fitted curves
shown in Figure 3b. For the purpose of this demonstration, the following values were used for all Ca2+
concentrations: γ0 (47.0 mN/m); Γm (1.60 × 10−6 mol/m2), a (0.03 mol/L) and b (145 mol/L), leaving
the interaction term ϕ (mg/mol) as the only adjustable parameter. p
g y
g
p
q
g
curves shown in Figure 3b. For the purpose of this demonstration, the following values were used
for all Ca2+ concentrations: γ0 (47.0 mN/m); Гm (1.60 × 10−6 mol/m2), a (0.03 mol/L) and b (145 mol/L)
leaving the interaction term φ (mg/mol) as the only adjustable parameter. 𝛾= 𝛾0 −𝑅𝑇𝑚(1 −exp (𝐶Ca
𝐶
)) ln (𝐶NA
𝑎+ 1) + exp (𝐶Ca
𝐶
) ln (𝐶NA
𝑏+ 1)
(13) Initial attempts at fitting the data shown in Figure 3b to Equation (12) indicated that, perhaps
unsurprisingly, Γm ~Γm’, allowing the simplified Equation (13) to be used to generate the fitted curves
shown in Figure 3b. For the purpose of this demonstration, the following values were used for all Ca2+
concentrations: γ0 (47.0 mN/m); Γm (1.60 × 10−6 mol/m2), a (0.03 mol/L) and b (145 mol/L), leaving
the interaction term ϕ (mg/mol) as the only adjustable parameter. p
g y
g
p
q
g
curves shown in Figure 3b. For the purpose of this demonstration, the following values were used
for all Ca2+ concentrations: γ0 (47.0 mN/m); Гm (1.60 × 10−6 mol/m2), a (0.03 mol/L) and b (145 mol/L)
leaving the interaction term φ (mg/mol) as the only adjustable parameter. derived from Equation (13).
3 3 D
d
f I t f
i l T
i
C
d C
3.3. Dependence of Interfacial Tension on CNA and CCa Thus, the interfacial tensions produced by NA
adsorption at a water interface and at a saturated Ca2+-containing interface are given by Equations
(9) and (11), respectively; it is unnecessary to define the bulk Ca2+ concentration responsible for the
latter condition at this stage of the analysis. In Equation (11), Γm’ and a’ are the respective surface
concentration and molecular constant of NA in the presence of the saturation concentration of Ca2+. Colloids Interfaces 2018, 2, 40
Colloids Interfaces 2018, 2, x 10 of 21
10 of 21 γ = γ0 −RTΓm′ ln
CNA
a′
+ 1
(11)
ases containing Ca2+ ions, and n-heptane phases containing NA, we
nteraction at the water/n-heptane interface is a function of the (11)
NA, we
of the In the case of aqueous phases containing Ca2+ ions, and n-heptane phases containing NA,
we assume that the extent of interaction at the water/n-heptane interface is a function of the
concentration ratio CCa/CNA, such that the contributions arising from pure water and aqueous Ca2+
solution interfaces are [1 −exp(φCCa/CNA] and exp(φCCa/CNA), respectively, where the factor ϕ
takes into account differences between concentration units as well as the relative effectiveness of Ca2+
and NA interactions at the interface. Thus, the general interfacial tension expression for different Ca2+
and NA concentrations is given by combining Equations (9) and (11), to give
concentration ratio CCa/CNA, such that the contributions arising from pure water and aqueous Ca2
solution interfaces are [1 −exp(𝐶Ca/𝐶NA] and exp(𝐶Ca/𝐶NA), respectively, where the factor φ
takes into account differences between concentration units as well as the relative effectiveness o
Ca2+ and NA interactions at the interface. derived from Equation (13).
3 3 D
d
f I t f
i l T
i
C
d C
3.3. Dependence of Interfacial Tension on CNA and CCa 𝛾= 𝛾0 −𝑅𝑇𝑚(1 −exp (𝐶Ca
𝐶
)) ln (𝐶NA
𝑎+ 1) + exp (𝐶Ca
𝐶
) ln (𝐶NA
𝑏+ 1)
(13) γ = γ0 −RTΓm
1 −exp
φCCa
CNA
ln
CNA
a
+ 1
+ exp
φCCa
CNA
ln
CNA
b
+ 1
(13)
NA
NA
t is evident from Figure 3b that this simplistic analysis produces reasonable fits to the
imental data and produces a sigmoidal variation of φ with the bulk Ca2+ concentration as (13)
to the It is evident from Figure 3b that this simplistic analysis produces reasonable fits to the experimental
data and produces a sigmoidal variation of ϕ with the bulk Ca2+ concentration as shown in Figure 4. The behavior seen in Figure 3b reflects the increasing concentration of Ca2+-NA interactions in the
interface as the bulk Ca2+ concentration is raised. The appearance of a plateau region in Figure 4 at CNA
~0.4 mol/L suggests that this bulk concentration corresponds to the interfacial “saturation” condition for
Ca2+ ions, as alluded to earlier, and for which the maximum influence on NA adsorption is seen. shown in Figure 4. The behavior seen in Figure 3b reflects the increasing concentration of Ca2+-NA
interactions in the interface as the bulk Ca2+ concentration is raised. The appearance of a plateau
region in Figure 4 at CNA ~ 0.4 mol/L suggests that this bulk concentration corresponds to the
interfacial “saturation” condition for Ca2+ ions, as alluded to earlier, and for which the maximum
influence on NA adsorption is seen. 3 4 NA Ca2+ Interfacial Tension Kinetics It is evident from Figure 3b that this simplistic analysis produces reasonable fits to the experimental
data and produces a sigmoidal variation of ϕ with the bulk Ca2+ concentration as shown in Figure 4. The behavior seen in Figure 3b reflects the increasing concentration of Ca2+-NA interactions in the
interface as the bulk Ca2+ concentration is raised. The appearance of a plateau region in Figure 4 at CNA
~0.4 mol/L suggests that this bulk concentration corresponds to the interfacial “saturation” condition for
Ca2+ ions, as alluded to earlier, and for which the maximum influence on NA adsorption is seen. shown in Figure 4. The behavior seen in Figure 3b reflects the increasing concentration of Ca2+-NA
interactions in the interface as the bulk Ca2+ concentration is raised. 3.4. NA-Ca2+ Interfacial Tension Kinetics
f
The additional decrease in inter A comparison with the data in
Figure 3b shows that the maximum k corresponds to the minimum interfacial tension of ~15 mN/m
Also for the low CNA region, the behavior of the second-order rate constant is shown as Also for the low CNA region, the behavior of the second-order rate constant is shown as a function
of CCa in Figure 5b. Here, it is seen that a minimum rate region is apparent at CCa ~0.5–1.0 mol/L,
consistent with the onset of the plateau region in Figure 4. g
function of CCa in Figure 5b. Here, it is seen that a minimum rate region is apparent at CCa ~ 0.5–1.0
mol/L, consistent with the onset of the plateau region in Figure 4. We speculate that the cause of the slower reduction of interfacial tension shown in Figure 2b i We speculate that the cause of the slower reduction of interfacial tension shown in Figure 2b
is due to complex formation involving Ca2+ and NA, and reorientation at the interface. This is
evident only in the case of Ca2+, but not for any of the other studied cations. With reference to the
adsorption parameters in Table 1, it appears probable that the hydrophobic side of the interfacial
region rapidly becomes saturated with randomly-distributed NA molecules upon contacting water
containing Ca2+ ions. The increased local NA concentration then reconfigures into a more ordered
adsorbed layer with the same adsorbed area/molecule as for NA at the n-heptane/deionized water
interface. With respect to the adsorption data for pre-formed Ca(NA)2 in Table 1, this finding suggests
that complete complexation between NA and Ca2+ is unlikely to have taken place. The question then
remains as to the role of Ca2+ ions in the adsorption process. To try to answer this, consideration needs
to be given to the behavior of metal ions in aqueous solution, and especially with regard to interactions
with carboxylates. p
g
due to complex formation involving Ca2+ and NA, and reorientation at the interface. This is eviden
only in the case of Ca2+, but not for any of the other studied cations. With reference to the adsorption
parameters in Table 1, it appears probable that the hydrophobic side of the interfacial region rapidly
becomes saturated with randomly-distributed NA molecules upon contacting water containing Ca2
ions. 3.4. NA-Ca2+ Interfacial Tension Kinetics
f
The additional decrease in inter The additional decrease in interfacial tension of NA at the n-heptane/water interface in the
presence of Ca2+ ions shown in Figure 2b occurs at rates that are measurable using the present technique. Kinetic data associated with these changes are given in Figure 5 in the form of two plots. The first
shows the effects of CNA on the first-order rate constant k at constant CCa (0.3 mol/L; Figure 5a), and the
second shows the variation in the second-order rate constant (=k/CNA; for CNA = 5 and 10 mg/L;
Figure 5b) as a function of CCa. presence of Ca2+ ions shown in Figure 2b occurs at rates that are measurable using the present
technique. Kinetic data associated with these changes are given in Figure 5 in the form of two plots
The first shows the effects of CNA on the first-order rate constant k at constant CCa (0.3 mol/L; Figure
5a), and the second shows the variation in the second-order rate constant (=k/CNA; for CNA = 5 and 10
mg/L; Figure 5b) as a function of CCa. Figure 4. The effect of Ca2+ concentration on the interaction factor φ used in the construction of the
fitted curves in Figure 3b. Figure 4. The effect of Ca2+ concentration on the interaction factor ϕ used in the construction of the
fitted curves in Figure 3b. Figure 4. The effect of Ca2+ concentration on the interaction factor φ used in the construction of the
fitted curves in Figure 3b. Figure 4. The effect of Ca2+ concentration on the interaction factor ϕ used in the construction of the
fitted curves in Figure 3b. 11 of 21
11 of 2 Colloids Interfaces 2018, 2, 40
Colloids Interfaces 2018, 2, x 11 of 21
11 of 2 Figure 5a illustrates the NA concentration range over which dynamic interfacial tension effects are
observed for a Ca2+ concentration of 0.3 mol/L. This indicates that there is a CNA “window”, spanning
almost three orders of magnitude, in which time-dependent interfacial processes are occurring. Moreover, for CNA < 50 mg/L at this particular Ca2+ concentration, k is proportional to the NA
concentration, the slope of the plot shown in the inset to Figure 5a representing the second-order rate
constant for the process. 3.4. NA-Ca2+ Interfacial Tension Kinetics
f
The additional decrease in inter As CNA increases, however, not only is a maximum rate constant apparent at
~100 mg/L, but the uncertainties in the data are also seen to increase, as a result of the changes in the
i t
f
i l t
i
b
i
ll
i
thi
i
A
i
ith th d t i
Fi
3b h
g
g
y
are observed for a Ca2+ concentration of 0.3 mol/L. This indicates that there is a CNA “window”
spanning almost three orders of magnitude, in which time-dependent interfacial processes ar
occurring. Moreover, for CNA < 50 mg/L at this particular Ca2+ concentration, k is proportional to th
NA concentration, the slope of the plot shown in the inset to Figure 5a representing the second-orde
rate constant for the process. As CNA increases, however, not only is a maximum rate constan
apparent at ~100 mg/L, but the uncertainties in the data are also seen to increase, as a result of th
changes in the interfacial tension becoming smaller in this region. A comparison with the data in
Fi
3b h
h
h
i
k
d
h
i i
i
f
i l
i
f 15
N/ p
NA
,
,
y
pp
~100 mg/L, but the uncertainties in the data are also seen to increase, as a result of the changes in the
interfacial tension becoming smaller in this region. A comparison with the data in Figure 3b shows
that the maximum k corresponds to the minimum interfacial tension of ~15 mN/m. apparent at 100 mg/L, but the uncertainties in the data are also seen to increase, as a result of th
changes in the interfacial tension becoming smaller in this region. A comparison with the data in
Figure 3b shows that the maximum k corresponds to the minimum interfacial tension of ~15 mN/m
Also for the low CNA region, the behavior of the second-order rate constant is shown as ~100 mg/L, but the uncertainties in the data are also seen to increase, as a result of the changes in the
interfacial tension becoming smaller in this region. A comparison with the data in Figure 3b shows
that the maximum k corresponds to the minimum interfacial tension of ~15 mN/m. pp
g
changes in the interfacial tension becoming smaller in this region. 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding In order to provide a plausible explanation for the observed behavior described above, three
particular factors need to be considered: (i) the behavior of hydrated cations at interfaces; (ii) exchange
of ligated water in the inner coordination sphere; and (iii) specific interactions between cations and
carboxylate groups. It has long been known that aqueous ions influence interfacial behavior [68]. Typically, aqueous
salts are considered to “negatively adsorb” at air/water or oil/water interfaces, producing an
increase in the respective surface or interfacial tensions in accord with the Gibbs adsorption theory,
and to an extent that is specific to each ion [69]. Ion theories have been developed over the years,
with increasingly sophisticated models proposed to predict their behavior at interfaces [70,71]. Herein, we have considered several different monovalent and divalent cations and found that,
of these, only Ca2+ affects the n-heptane/water interfacial tension in the presence of NA. All cations
have a tendency to be hydrated, owing to their ionic charge [72], and as a consequence will be excluded
from the interfacial region [73,74]. Therefore, we have to examine why Ca2+ behaves differently from
the rest of the studied cations. Ikeda et al. compared Ca2+ and Mg2+ ions in solution using constrained molecular dynamics,
and found that the hydrated structure of Ca2+ is highly variable compared with that of Mg2+ [75]. Using density functional theory, Pavlov et al. identified that Ca2+ has a lower water binding energy
compared with Mg2+ (as well as Be2+ and Zn2+), notwithstanding the different water structures around
the respective ions [76]. This implies that Ca2+ will show potentially greater lability of its inner-shell
coordinated water molecules, which is of importance for adsorption as well as coordination to ligands. Specific ion effects were originally classified based on the different abilities of salts to precipitate
proteins from aqueous solution, a phenomenon which is accredited to Hofmeister in 1888 [77,78]. The “Hofmeister effect”, as it came to be known, now refers more generally to the relative effects
of either cations or anions on colloidal, surface or biological processes. The commonly-held view
that has been developed, discussed and reviewed by many authors over the years is that ion
hydration is fundamental to the Hofmeister effect [73,74,79]; this includes the behavior of ions at
liquid interfaces [70]. 3.4. NA-Ca2+ Interfacial Tension Kinetics
f
The additional decrease in inter The increased local NA concentration then reconfigures into a more ordered adsorbed laye
with the same adsorbed area/molecule as for NA at the n-heptane/deionized water interface. With
respect to the adsorption data for pre-formed Ca(NA)2 in Table 1, this finding suggests that complete
complexation between NA and Ca2+ is unlikely to have taken place. The question then remains as to
the role of Ca2+ ions in the adsorption process. To try to answer this, consideration needs to be given
to the behavior of metal ions in aqueous solution, and especially with regard to interactions with
carboxylates. Figure 5. (a) The effect of NA concentration on the first-order kinetic rate constant for equilibration of
the n-heptane/water interfacial tension in the presence of 0.3 mol/L Ca2+ ions. The inset shows the
second-order plot for low NA concentrations. (b) The effect of Ca2+ concentration on the
second-order rate constant in the low NA concentration range (red and blue symbols are for CNA = 5
and 10 mg/L, respectively). Figure 5. (a) The effect of NA concentration on the first-order kinetic rate constant for equilibration
of the n-heptane/water interfacial tension in the presence of 0.3 mol/L Ca2+ ions. The inset
shows the second-order plot for low NA concentrations. (b) The effect of Ca2+ concentration on
the second-order rate constant in the low NA concentration range (red and blue symbols are for
CNA = 5 and 10 mg/L, respectively). Figure 5. (a) The effect of NA concentration on the first-order kinetic rate constant for equilibration of
the n-heptane/water interfacial tension in the presence of 0.3 mol/L Ca2+ ions. The inset shows the
second-order plot for low NA concentrations. (b) The effect of Ca2+ concentration on the
second-order rate constant in the low NA concentration range (red and blue symbols are for CNA = 5
and 10 mg/L, respectively). Figure 5. (a) The effect of NA concentration on the first-order kinetic rate constant for equilibration
of the n-heptane/water interfacial tension in the presence of 0.3 mol/L Ca2+ ions. The inset
shows the second-order plot for low NA concentrations. (b) The effect of Ca2+ concentration on
the second-order rate constant in the low NA concentration range (red and blue symbols are for
CNA = 5 and 10 mg/L, respectively). 12 of 21 Colloids Interfaces 2018, 2, 40 12 of 21 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding The size of the ion and its nuclear charge control its hydration properties
in aqueous solution and serve to define kosmotropic (strongly hydrated) and chaotropic (weakly
hydrated) ions. Thus, kosmotropes are likely to possess high charge densities and their consequent
strong hydration contributes to repulsion from an interface [73,74]. Adsorption of these ions would be
accompanied by liberation of the strongly-bound hydration water, requiring large positive enthalpic
contributions to the Gibbs energy which would not necessarily be outweighed by the corresponding
entropy changes, i.e., ∆Hads −T∆Sads > 0. On the other hand, water molecules bound to chaotropic
ions are more easily lost on approaching an interface [73,74], making adsorption more favorable,
i.e., ∆Hads −T∆Sads < 0. Considering the second factor mentioned above, that of the lability of the coordinated inner-shell
water molecules, consistent with the theoretical investigations [75,76], it was found that water exchange
rates from the innermost hydration shells are almost four orders of magnitude faster for Ca2+ than
for Mg2+ ions [80]. This trend reflects the relative sizes of the anhydrous cations, and the greater
charge density of Mg2+. Using the same arguments, this would suggest that Sr2+ should behave in
a similar fashion to Ca2+, but we have not found this to be the case. We therefore turn to the third
factor mentioned above, that of specific effects involving the coordination chemistry of Ca2+ and the
carboxylate group. In a recent example, Kherb et al. studied the strength of association of monovalent and
divalent cations with carboxylate sites on a polypeptide biopolymer, and identified the divalent ion
sequence Zn2+ > Ca2+ > Ba2+ > Sr2+ > Mg2+, whilst monovalent cations showed weaker binding [81]. The inference made from this was that the most hydrated cations bind more strongly to carboxylate [81]. The structures of metal carboxylate complexes have been extensively studied over the years and
have allowed the detailed examination of X-ray crystallographic data by Einspahr and Bugg [82] 13 of 21
13 of 21 Colloids Interfaces 2018, 2, 40
Colloids I terfaces 2018 2 and Carrell et al. [83], which, together with other investigations [47,84–86] has culminated in the
principal binding modes shown in Figure 6 being identified. Thus, specifically for Ca2+, monodentate
coordination with one carboxylate O atom and hydrogen bonded water, bidentate coordination,
with both carboxylate O atoms, or bridging (see Figure 6a–c) are notable [82]. 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding On the other hand,
a major difference in bonding mode with carboxylic acids in the case of magnesium is the formation of
uncoordinated ionic species of the form shown in Figure 6d [87]. The crystal structure of the magnesium
complex of p-anisic acid (4-methoxybenzoic acid) contains a layer structure of alternating [Mg(H2O)6]2+
cations and p-anisate anions with the lattice water functioning as a link between the layers. On the
other hand, the monohydrated Ca(II) salt is a two-dimensional coordination polymer [87]. principal binding modes shown in Figure 6 being identified. Thus, specifically for Ca2+, monodentate
coordination with one carboxylate O atom and hydrogen bonded water, bidentate coordination,
with both carboxylate O atoms, or bridging (see Figure 6a–c) are notable [82]. On the other hand, a
major difference in bonding mode with carboxylic acids in the case of magnesium is the formation of
uncoordinated ionic species of the form shown in Figure 6d [87]. The crystal structure of the
magnesium complex of p-anisic acid (4-methoxybenzoic acid) contains a layer structure of
alternating [Mg(H2O)6]2+ cations and p-anisate anions with the lattice water functioning as a link
between the layers. On the other hand, the monohydrated Ca(II) salt is a two-dimensional
coordination polymer [87]. Figure 6. Bonding modes identified for divalent ion interactions with carboxylates [47,84–86]. 1:1
coordination is shown in binding modes (a) and (b), with bridging exhibited in (c). (d) shows ionic
bonding. Figure 6. Bonding modes identified for divalent ion interactions with carboxylates [47,84–86]. 1:1 coordination is shown in binding modes (a) and (b), with bridging exhibited in (c). (d) shows
ionic bonding. Figure 6. Bonding modes identified for divalent ion interactions with carboxylates [47,84–86]. 1:1
coordination is shown in binding modes (a) and (b), with bridging exhibited in (c). (d) shows ionic
bonding. Figure 6. Bonding modes identified for divalent ion interactions with carboxylates [47,84–86]. 1:1 coordination is shown in binding modes (a) and (b), with bridging exhibited in (c). (d) shows
ionic bonding. The comparative behavior between Ca2+ and Mg2+ interactions at carboxylate monolayers has
also been studied by Tang et al. using vibrational sum frequency generation spectroscopy [88,89]. 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding Thus, binding of these ions to palmitic acid monolayers indicated different concentration-dependent
metal ion binding modes [88], concluding that as the Ca2+ concentration increases (to 0.3 mol/L),
complexation favors the 2:1 bridging configuration shown in Figure 6c, whereas Mg2+ adopts the
ionic complex in Figure 6d consistent with solution and crystallographic results. Moreover, the
stronger Ca2+ binding was shown to have a much greater impact on the underlying hydrogen
bonding below the carboxylic acid monolayer [89], which is arguably consistent with the differences
seen between Ca2+ and Mg2+ ions in the present study at the oil/water interface
The comparative behavior between Ca2+ and Mg2+ interactions at carboxylate monolayers has
also been studied by Tang et al. using vibrational sum frequency generation spectroscopy [88,89]. Thus, binding of these ions to palmitic acid monolayers indicated different concentration-dependent
metal ion binding modes [88], concluding that as the Ca2+ concentration increases (to 0.3 mol/L),
complexation favors the 2:1 bridging configuration shown in Figure 6c, whereas Mg2+ adopts the ionic
complex in Figure 6d consistent with solution and crystallographic results. Moreover, the stronger
Ca2+ binding was shown to have a much greater impact on the underlying hydrogen bonding below
the carboxylic acid monolayer [89], which is arguably consistent with the differences seen between
Ca2+ and Mg2+ ions in the present study at the oil/water interface. seen between Ca and Mg ions in the present study at the oil/water interface. We have also looked to biological systems to provide an understanding of the differences
between divalent ion coordination modes, especially with respect to carboxylate ligands found in a
wide range of protein environments [90–94]. Williams provides a clear indication that the most
significant factors responsible for ion binding in biological systems are, inter alia, ion charge, ion size,
ligand donor atom, and preferred coordination geometry [95]. Thus, in connection with biological
systems, binding of ions such as those considered in the present study is restricted to oxygen donor
ligands, and specificity of Ca2+ binding (e.g., relative to Mg2+) is due to its size and less-restricted
steric preferences compared with other ions. Ca2+ shows a greater tendency for bidentate binding to
carboxylate, compared with Mg2+ [91], and a lower affinity for water [92,94]
We have also looked to biological systems to provide an understanding of the differences between
divalent ion coordination modes, especially with respect to carboxylate ligands found in a wide range
of protein environments [90–94]. 3.6. Implications for Crude Oil Surface Chemistry
3.6. Implications for Crude Oil Surface Chemistry Over the past two decades in particular, the importance of injection water composition on crude
oil reservoir chemistry has been established through various laboratory coreflood investigations and
field evaluations. Studies have shown that low salinity conditions contribute to improvements in
oil recovery, but although various mechanisms have been proposed, a full understanding of the low
salinity effect is still lacking. However, a common factor throughout some proposed mechanisms
is the role of the ionic composition of the injection and formation waters [98] and especially their
effects on reservoir wettability and the corresponding affinity of the oil for the rock surface [99]. Based on our present findings the specific role of Ca2+ is considered to be significant. From the data
given in Figure 3b, for example, it is evident that the minimum interfacial tension of the present NA
system is dependent on the Ca2+ concentration, as shown in Figure 7a. This representation of the data
assumes that there is an excess of NA in the oil phase and that the bulk Ca2+ concentration governs
the interfacial tension. Over the past two decades in particular, the importance of injection water composition on crude
oil reservoir chemistry has been established through various laboratory coreflood investigations and
field evaluations. Studies have shown that low salinity conditions contribute to improvements in oil
recovery, but although various mechanisms have been proposed, a full understanding of the low
salinity effect is still lacking. However, a common factor throughout some proposed mechanisms is
the role of the ionic composition of the injection and formation waters [98] and especially their
effects on reservoir wettability and the corresponding affinity of the oil for the rock surface [99]. Based on our present findings the specific role of Ca2+ is considered to be significant. From the data
given in Figure 3b, for example, it is evident that the minimum interfacial tension of the present NA
system is dependent on the Ca2+ concentration, as shown in Figure 7a. This representation of the data
assumes that there is an excess of NA in the oil phase and that the bulk Ca2+ concentration governs
the interfacial tension. Figure 7. (a) The effect of Ca2+ ion concentration (expressed here in mg/L) on the minimum interfacial
tension of the n-heptane/water interface when NA is in excess. 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding Williams provides a clear indication that the most significant factors
responsible for ion binding in biological systems are, inter alia, ion charge, ion size, ligand donor atom,
and preferred coordination geometry [95]. Thus, in connection with biological systems, binding of
ions such as those considered in the present study is restricted to oxygen donor ligands, and specificity
of Ca2+ binding (e.g., relative to Mg2+) is due to its size and less-restricted steric preferences compared
with other ions. Ca2+ shows a greater tendency for bidentate binding to carboxylate, compared with
Mg2+ [91], and a lower affinity for water [92,94]. carboxylate, compared with Mg [91], and a lower affinity for water [92,94]. Ca2+ binding to carboxylate is also considered to be advantageous based on Collins’ “matching
water affinity” approach, according to which specific pairing interactions between ions, resulting in
the formation of ion pairs, is most favorable for ions of comparable hydration energy [96]. The latter
is quantified [97] by the apparent dynamic hydration number (ADHN) which, for Ca2+ and
carboxylate are 2.1 and 2.0, respectively, which therefore match up surprisingly well (cf. 5.7 for
Mg2+). Kiriukhin and Collins consider that interaction between these two species will result in
dissipation of the hydration shell, thereby benefitting coordination [97], which we suggest may also
feature in specific Ca2+-naphthenic acid interactions across an interface. Such interactions would be
likely to require water to be displaced from the first coordination shell
Ca2+ binding to carboxylate is also considered to be advantageous based on Collins’ “matching
water affinity” approach, according to which specific pairing interactions between ions, resulting in
the formation of ion pairs, is most favorable for ions of comparable hydration energy [96]. The latter is
quantified [97] by the apparent dynamic hydration number (ADHN) which, for Ca2+ and carboxylate
are 2.1 and 2.0, respectively, which therefore match up surprisingly well (cf. 5.7 for Mg2+). Kiriukhin
and Collins consider that interaction between these two species will result in dissipation of the
hydration shell, thereby benefitting coordination [97], which we suggest may also feature in specific
Ca2+-naphthenic acid interactions across an interface. Such interactions would be likely to require
water to be displaced from the first coordination shell. 14 of 21 14 of 21 Colloids Interfaces 2018, 2, 40 In summary, there is compelling evidence for specific Ca2+ interactions with carboxylic acids from
biology. 3.5. Interfacial Behavior of Ions and Metal Ion–Carboxylate Bonding Therefore, a combination of evidence from biological and coordination chemistry appears to
offer an interpretation of the present surface chemical findings, which may be of relevance in other
practical and technological areas, such as our current interest in crude oil surface chemistry. Colloids Interfaces 2018, 2, x
14 of 21
appears to offer an interpretation of the present surface chemical findings, which may be of
relevance in other practical and technological areas, such as our current interest in crude oil surface
chemistry. 3.6. Implications for Crude Oil Surface Chemistry
3.6. Implications for Crude Oil Surface Chemistry Young’s equation is derived from the balance of forces acting on the contact line of a drop on a
ace (Figure 7b), i.e., γWS=γOS + γWO cos θ
(14) (14) where the γ terms are the respective Gibbs energies of the water/solid (WS), oil/solid (OS) and
oil/water (OW) interfaces and θ is the contact angle made by oil on the solid surface. We can therefore
consider two hypothetical situations by way of illustrating the potential role of Ca2+ ions in oil recovery,
based on an initial oil/rock system in equilibrium with formation water, and a final waterflooded
system with reduced salinity, and a concomitant lower Ca2+ concentration. We assume that Equation (14) can be applied to each situation, such that the change in Ca2+
concentration causes a resultant change in the water/oil interfacial tension, γWO. For this elementary
analysis, we also assume that γWS and γOS remain relatively unaffected by the change in Ca2+
concentration. However, although we recognize that, in practice it is likely that wetting films
comprising, for example, hydrophobic species [101] or hydration layers [102], can modify water/solid
and oil/solid interfaces and their respective interfacial energies, predicting the effects on the contact
angle, for example, is not easy. Therefore, we consider, as an example, the simple analysis involving
the initial and final conditions as introduced above, for which it is apparent that (15) γi
OW cos θi = γ f
OW cos θ f
(15) where i and f refer to initial and final states, respectively. As shown in Table 2, formation waters are
generally more saline than sea water, the ionic composition being source-dependent but also dependent
on the geologic period [103]. Thus, it is possible that during geologic time the oil/rock system will
have been “conditioned” with a relatively highly saline brine containing a high Ca2+ concentration,
say ~20,000 mg/L as an illustrative value (Table 2). Therefore, with reference to Figure 7a, dilution of
the formation brine to ~5000 mg/L Ca2+ would approximately double the interfacial tension, such that (16) cos θ f = (γi
OW/γ f
OW) cos θi ≈0.5 cos θi
(16) cos θ f = (γi
OW/γ f
OW) cos θi ≈0.5 cos θi
(16) Table 2. Na+ and Ca2+ compositions of formation waters from different geologic periods. For comparison,
sea water contains ~10,500 mg/L Na+ and ~400 mg/L Ca2+. Data compiled from Collins [103]. Table 2. 3.6. Implications for Crude Oil Surface Chemistry
3.6. Implications for Crude Oil Surface Chemistry (b) Schematic of an oil drop adhering
to a rock surface in water showing the three-phase contact point defining the contact angle. (c)
Demonstrating the effect of a decrease in Ca2+ concentration from 20,000 mg/L to 5000 mg/L on the
final contact angle for different initial contact angles, and a simulated visual result on a sand surface
to illustrate. Figure 7. (a) The effect of Ca2+ ion concentration (expressed here in mg/L) on the minimum interfacial
tension of the n-heptane/water interface when NA is in excess. (b) Schematic of an oil drop adhering to a
rock surface in water showing the three-phase contact point defining the contact angle. (c) Demonstrating
the effect of a decrease in Ca2+ concentration from 20,000 mg/L to 5000 mg/L on the final contact angle
for different initial contact angles, and a simulated visual result on a sand surface to illustrate. Figure 7. (a) The effect of Ca2+ ion concentration (expressed here in mg/L) on the minimum interfacial
tension of the n-heptane/water interface when NA is in excess. (b) Schematic of an oil drop adhering
to a rock surface in water showing the three-phase contact point defining the contact angle. (c)
Demonstrating the effect of a decrease in Ca2+ concentration from 20,000 mg/L to 5000 mg/L on the
final contact angle for different initial contact angles, and a simulated visual result on a sand surface
ill
Figure 7. (a) The effect of Ca2+ ion concentration (expressed here in mg/L) on the minimum interfacial
tension of the n-heptane/water interface when NA is in excess. (b) Schematic of an oil drop adhering to a
rock surface in water showing the three-phase contact point defining the contact angle. (c) Demonstrating
the effect of a decrease in Ca2+ concentration from 20,000 mg/L to 5000 mg/L on the final contact angle
for different initial contact angles, and a simulated visual result on a sand surface to illustrate. Colloids Interfaces 2018, 2, 40 15 of 21 With reference to Figure 7b, showing a schematic of oil coating a solid (rock) surface,
two equilibrium conditions can be envisaged during oil recovery. The first is the original condition
within the reservoir, in which the crude oil and rock are in equilibrium with the formation brine,
and the second is during waterflooding conditions where sea water, softened water or low salinity
water will change the aqueous phase composition [100]. 3.6. Implications for Crude Oil Surface Chemistry
3.6. Implications for Crude Oil Surface Chemistry Na+ and Ca2+ compositions of formation waters from different geologic periods. For comparison,
sea water contains ~10,500 mg/L Na+ and ~400 mg/L Ca2+. Data compiled from Collins [103]. Geologic Period
Average (Maximum) Metal Ion Concentration (mg/L)
Na+
Ca2+
Oligocene/Paleogene
39,000 (103,000)
2530 (38,800)
Cretaceous
31,000 (88,600)
7000 (37,400)
Jurassic
57,300 (120,000)
25,800 (56,300)
Permian
47,000 (109,000)
8600 (22,800)
Carboniferous (Pennsylvanian)
43,000 (101,000)
9100 (205,000)
Carboniferous (Mississippian)
41,500 (115,800)
8900 (37,800)
Devonian
48,000 (101,000)
18,000 (129,000) Thus, Equation (16) enables the plot shown in Figure 7c to be constructed, illustrating how the
contact angle between the oil and rock surface is affected by the Ca2+ concentration change in the
example given, from 20,000 mg/L to 5000 mg/L during waterflooding. Here, it is evident that the
lowest initial contact angles, indicating high oil affinity with the surface, are influenced most by the 16 of 21 Colloids Interfaces 2018, 2, 40 16 of 21 change in interfacial tension. In this region, contact angles increase by as much as 50◦, potentially
improving oil recovery. Such a scenario is depicted in the inset images in Figure 7c, added for
illustrative purposes only, but is consistent with the results of the study by Shehata and Nasr-El-Din
regarding the dominant role of formation water salinity and composition in improving oil recovery
during low salinity waterflooding [98]. Thus, the findings presented herein may provide further insight into potential mechanisms for the
low salinity effect. In addition, it is apparent that the ability to predict the effect of water chemistry on
oil recovery reservoir requires not only knowledge of the interfacial chemistry of inorganic ions, but also
the range of surface-active species present in the crude oil; here, we have only examined the effects of
naphthenic acids. For real crude oils, however, the increased complexity and competition resulting
from the presence of other interfacially active species, such as asphaltenes and resin components,
would be expected to produce a different interfacial behavior [104]. This has been borne out in studies
on crude oil fractions to be reported elsewhere. 4. Conclusions It seems to be a recurring theme that Ca2+ plays significant roles in the aqueous colloid and
interfacial chemistry of crude oil and, in this respect, is more important than other metal ions. As discussed in the present paper, the characteristics of this ion make it favorable to undergo specific
interactions with carboxylic components found in crude oil, to some extent mimicking its unique
position in biology [95]. Of the metal ions studied herein, both monovalent and divalent, only Ca2+ has
been shown to exhibit interfacial coordination with carboxylate groups of a commercial naphthenic
acid sample. The present findings have indicated that, on their own, naphthenic acids (NAs) are only weakly
active at the n-heptane/water interface at concentrations below ~200 mg/L as determined using the
dynamic drop volume technique. However, this behavior is modified in the specific presence of Ca2+,
whereas Mg2+ and Sr2+, as well as several alkali metal cations, exhibit no comparable effect. We have
found that, for a given aqueous Ca2+ concentration, there exists an “optimum” NA concentration
in the n-heptane phase at which a minimum interfacial tension is attained. As shown in Figure 7,
the lowest interfacial tension is ~10–15 mN/m, to be compared with a value of ~45 mN/m for the other
cations and deionized water. The interaction is not stoichiometric with respect to the respective bulk
concentrations, in that the minimum interfacial tension occurs at a bulk Ca2+:NA molar concentration
ratio of ~1000:1. The interfacial stoichiometry is unclear at present, but probably relates to the
bidentate preference of Ca2+, but further interfacial studies will be necessary in order to provide more
definitive evidence. In fact, there may be no single factor responsible for the results presented in this study. Rather,
for Ca2+, the combination of its ionic hydration structure and its thermodynamic and kinetic properties
as well as its coordination preferences, particularly relating to carboxylate, may contribute to its
behavior at the oil/water interface. Conclusions from studies undertaken some 80 years ago by
Alexander et al. also highlighted the specific relevance of Ca2+ ions (compared with Na+, K+ and Mg2+)
to the chemistry of biological films, such as cephalin (kephalin) at the benzene/water interface [105]. Author Contributions: S.E.T. conceptualized and supervised the research and conducted some of the experiments. H.T.C. conducted the majority of the experiments and prepared an initial interpretation of the results. S.E.T. wrote
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on Low-salinity waterflooding in sandstone reservoirs. SPE Reserv. Eval. Eng. 2017, 20, 59–76. [CrossRef] 99. Lashkarbolooki, M.; Ayatollahi, S.; Riazi, M. References The impacts of aqueous ions on interfacial tension and
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104. Sauerer, B.; Stukan, M.; Buiting, J.; Abdallah, W.; Andersen, S. Dynamic asphaltene-stearic acid competition
at the oil–water interface. Langmuir 2018, 34, 5558–5573. [CrossRef] [PubMed] 105. Alexander, A.E.; Teorell, T.; Åborg, G.C. A study of films at the liquid/liquid interface. Part III. A specific
effect of calcium ions on kephalin monolayers. Trans. Faraday Soc. 1939, 35, 1200–1205. [CrossRef] © 2018 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access
article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/).
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What Explains Vietnam’s Exceptional Performance in Education Relative to Other Countries? Analysis of the 2012 and 2015 PISA Data
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Abstract On the 2012 and 2015 PISA assessments, Vietnam’s performance ranks much higher than that of all other
developing countries, and even ahead of richer countries such as the U.S. and the U.K. This is especially
remarkable since Vietnam was the poorest or second-poorest PISA participant. We provide the first rigorous
investigation of Vietnam’s strong performance. Despite various corrections for potentially non-representative
PISA samples, including bias due to Vietnam’s large out-of-school population, Vietnam remains a large positive
outlier conditional on its income. Possible higher motivation of Vietnamese students at most only partly explains
Vietnam’s performance. We also find that household- and school-level variables explain little of Vietnam’s
exceptional performance. Finally, Blinder-Oaxaca decompositions indicate that the gap in average test scores
between Vietnam and the other participating countries is due not to differences in students’ and schools’
observable characteristics, but is mostly caused by the greater “productivity” of those characteristics in Vietnam. JEL Classifications: H0, I2, O1, P3 JEL Classifications: H0, I2, O1, P3 JEL Classifications: H0, I2, O1, P3
Keywords: education, student learning, test scores, enrolment, PISA, Vietnam Keywords: education, student learning, test scores, enrolment, PISA, Vietnam What Explains Vietnam’s Exceptional Performance in Education Relative to Other Countries?
Analysis of the 2012 and 2015 PISA Data Hai-Anh Dang
World Bank Acknowledgements: Acknowledgements: We thank Francesco Avvisati, Luis Felipe Saenz, and Nic Spaull, as well as seminar participants at
Columbia University (Teachers College), the City University of New York, the University of Copenhagen,
the 2017 LACEA conference in Buenos Aires, the 2017 RISE conference, the University of Hawaii, and
the OECD, for helpful comments. This is one of a series of working papers from “RISE”—the large-scale education systems research
programme supported by funding from the United Kingdom’s Department for International Development
(DFID), the Australian Government’s Department of Foreign Affairs and Trade (DFAT), and the Bill and
Melinda Gates Foundation. The Programme is managed and implemented through a partnership between
Oxford Policy Management and the Blavatnik School of Government at the University of Oxford. Please cite this paper as: Dang, H., Glewwe, P., Lee, J., and Vu, K. (2020). What Explains Vietnam’s Exceptional Performance in
Education Relative to Other Countries? Analysis of the 2012 and 2015 PISA Data. RISE Working Paper
Series. 20/036. https://doi.org/10.35489/BSG-RISE-WP_2020/036 Dang, H., Glewwe, P., Lee, J., and Vu, K. (2020). What Explains Vietnam’s Exceptional Performance in
Education Relative to Other Countries? Analysis of the 2012 and 2015 PISA Data. RISE Working Paper
Series. 20/036. https://doi.org/10.35489/BSG-RISE-WP_2020/036 Use and dissemination of this working paper is encouraged; however, reproduced copies may not be
used for commercial purposes. Further usage is permitted under the terms of the Creative Commons
License. The findings, interpretations, and conclusions expressed in RISE Working Papers are entirely those of the
author(s) and do not necessarily represent those of the RISE Programme, our funders, or the authors’
respective organisations. Copyright for RISE Working Papers remains with the author(s). Research on Improving Systems of Education (RISE)
www.riseprogramme.org
information@riseprogramme.org Research on Improving Systems of Education (RISE)
www.riseprogramme.org
information@riseprogramme.org The lower secondary rate is from Dang and Glewwe (2018), while the primary completio
he authors using the 2014 VHLSS data; the VHLSS data are described further below. y
, w
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, w
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aiwan as countries, although the first two are Chinese territories, and Taiwan’s status is disputed. er considers only countries, and thus excludes Shanghai, which is obviously not representative of C
nd the territory of Perm, which not representative of Russia. For convenience, we refer to Hong Ko ower secondary rate is from Dang and Glewwe (2018), while the primary complet
hors using the 2014 VHLSS data; the VHLSS data are described further below. g
2 This paper considers only countries, and thus excludes Shanghai, which is obviously not representative of China as
a whole, and the territory of Perm, which not representative of Russia. For convenience, we refer to Hong Kong, 3 The GDP per capita for Indonesia was $US 3,347 in 2015, which is about 50 percent higher than that of Vietnam
($US 2,110) in the same year (World Bank, 2017). g
2 This paper considers only countries, and thus excludes Shanghai, which is obviously not representative of China as
a whole, and the territory of Perm, which not representative of Russia. For convenience, we refer to Hong Kong,
Macao and Taiwan as countries, although the first two are Chinese territories, and Taiwan’s status is disputed. 1 The lower secondary rate is from Dang and Glewwe (2018), while the primary completion rate was calculated by g
;
er considers only countries, and thus excludes Shanghai, which is obviously not representative of C The lower secondary rate is from Dang and Glewwe (2018), while the primary completion rate wa
he authors using the 2014 VHLSS data; the VHLSS data are described further below the authors using the 2014 VHLSS data; the VHLSS data are described further below.
2 This paper considers only countries, and thus excludes Shanghai, which is obviously not representative of China as
a whole, and the territory of Perm, which not representative of Russia. For convenience, we refer to Hong Kong,
Macao and Taiwan as countries, although the first two are Chinese territories, and Taiwan’s status is disputed.
3 The GDP per capita for Indonesia was $US 3,347 in 2015, which is about 50 percent higher than that of Vietnam
($US 2 110) in the same year (World Bank 2017) 1 The lower secondary rate is from Dang and Glewwe (2018), while the primary completion rate was calculated by
the authors using the 2014 VHLSS data; the VHLSS data are described further below.
2 This paper considers only countries, and thus excludes Shanghai, which is obviously not representative of China as
a whole, and the territory of Perm, which not representative of Russia. For convenience, we refer to Hong Kong,
Macao and Taiwan as countries, although the first two are Chinese territories, and Taiwan’s status is disputed.
3 The GDP per capita for Indonesia was $US 3,347 in 2015, which is about 50 percent higher than that of Vietnam
($US 2,110) in the same year (World Bank, 2017). 1 The lower secondary rate is from Dang and Glewwe (2018), while the primary completion rate was calculated by
the authors using the 2014 VHLSS data; the VHLSS data are described further below. a whole, and the territory of Perm, which not representative of Russia. For convenience, we refer to Hong Kong,
Macao and Taiwan as countries, although the first two are Chinese territories, and Taiwan’s status is disputed.
3 The GDP per capita for Indonesia was $US 3,347 in 2015, which is about 50 percent higher than that of Vietnam
($US 2,110) in the same year (World Bank, 2017). I. Introduction Vietnam’s rapid economic growth in the last 30 years has transformed it from one of the
world’s poorest countries to a middle-income country (World Bank, 2013). While Vietnam’s
economic achievements have attracted much attention, in more recent years its accomplishments
in education have also generated a great deal of international interest. Vietnam’s high performance in the “quantity” of education is exemplified by its high
primary school completion rate of 97%, and its high lower secondary enrollment rate of 95%.1
More striking still are the results of the 2012 PISA assessment: Vietnam’s performance ranked
16th in math and 18th in reading out of 63 countries and territories,2 ahead of both the US and the
UK and much higher than that of any other developing country (OECD, 2014a). Its 2012 PISA
mathematics and readings scores (at 511 and 508), for example, were more than one standard
deviation higher than those of Indonesia (375 and 396), a nearby country whose GDP per capita
is most similar to that of Vietnam among all countries that participated in the 2012 PISA.3 A visual depiction of Vietnam’s performance on the PISA in 2012, given its income, is
shown in Figures 1 and 2, which plot PISA scores in math and reading by the log of per capita
GDP for all 63 countries. Vietnam is in the upper left in both figures, higher than any other
country above the line that shows the expected test score given per capita GDP. Vietnam is also
the largest positive outlier when PPP (purchasing power parity) GDP is used (see Figures B1 and
B2 in Appendix B). More recently, Vietnam was again the largest positive outlier in the 2015
PISA assessment, as seen in Figures 3 and 4. Finally, if the sample is limited to the nine East 1 1 Asian and Southeast Asian participants in the 2012 PISA, Vietnam is still the largest positive
outlier (see Figures B3 and B4 in Appendix B). Asian and Southeast Asian participants in the 2012 PISA, Vietnam is still the largest positive
outlier (see Figures B3 and B4 in Appendix B). This paper uses the 2012 and 2015 PISA data to try to understand Vietnam’s unusually
high performance on these assessments of student learning. More specifically, it has three
objectives. I. Introduction First, it examines whether Vietnam’s impressive performance on these PISA
assessments may be exaggerated because: i) the 15-year-old Vietnamese students who
participated in these assessments are not representative of 15-year-old students in Vietnam; ii)
the enrollment rate of 15-year-olds in Vietnam is much lower than those rates in other PISA
countries; or iii) Vietnamese students put more effort into those assessments than other students. Second, it uses regression methods to investigate whether family, teacher or school
characteristics in the PISA data can explain Vietnamese students’ high performance. Third, it
applies the Oaxaca-Blinder decomposition to disaggregate the difference in average test scores
between Vietnamese students and students in the other countries that participated in the 2012 and
2015 PISA assessments. This paper’s first finding is that Vietnam’s striking performance on the 2012 and 2015
PISA assessments is at most only partially reduced by adjustments to reduce the possible sources
of upward bias discussed above. In particular, adjusting Vietnam’s PISA scores to control for
possible oversampling of wealthier, more urban students reduces its scores by only 20-24 (14-15)
points in 2012 (2015). Moreover, three different methods to adjust for Vietnam’s low coverage
(enrollment) rates (after correcting the OECD coverage rates for Vietnam) have little effect on its
outlier status. Finally, back-of-the-envelope calculations to account for possible higher
motivation of Vietnamese students on the PISA account for less than half of Vietnam’s outlier
status. 2 The second finding it that accounting for household- and school-level variables in the
PISA data explains little of Vietnam’s high performance on the 2012 and 2015 PISA assessments
relative to its income level. A third finding is that Oaxaca-Blinder decompositions show that the
gap in average test scores between Vietnam and the other countries in the 2012 and 2015 PISA is
not due to differences in observable child, household and school characteristics; it is mostly due
to Vietnam’s higher “productivity” of those characteristics relative to the other PISA countries. This paper focuses on Vietnam, and is most relevant to education policies in that country. I. Introduction But to the best of our knowledge, it also the first study to offer a rigorous and detailed analysis of
the performance of one PISA country participant in comparison with the others.4 As such, the
framework of analysis used here may be useful for studies of other countries’ performance in a
similar context, not only for the PISA but also other skill assessments of multiple countries. This paper consists of five sections. It examines the possible mechanisms that could
exaggerate Vietnam’s performance in the next section, before employing regression methods to
investigate the role of family, teacher and school characteristics in explaining this performance in
Section III. It then provides in Section IV Oaxaca-Blinder decompositions of the test score gap
between Vietnam and the other PISA participants, focusing on child, household and school
characteristics. Conclusions are offered in Section V. ent investigation of the qualitative aspects of the PISA, see Waldow and Steiner-Khamsi (2019). See, for example, the comments by Deputy Minister of Education Nguyen Vinh Hien in Thanh Ni 4 For a recent investigation of the qualitative aspects of the PISA, see Waldow and Steiner-Kh g
q
p
,
(
)
e, the comments by Deputy Minister of Education Nguyen Vinh Hien in Thanh Nien News (2013). 6 Most PISA countries, including Vietnam, conducted testing on April 12-14 of 2012. Thus children born in 1996
would be from 15 years and 3 (completed) months of age (born in December of 1996) and 16 years to 2 (completed)
months (born in January of 1996). The target population was defined as “all students aged from 15 years and 3 com-
pleted months to 16 years and 2 completed months at the beginning of the assessment period” (OECD, 2014b, p.66).
7 PISA’s urban classification refers to schools, not students. Some students may live in rural areas and attend urban
schools; they would be classified as urban in the PISA but as rural in the VHLSS and so may explain some of the
urban/rural difference in the two samples. Yet this cannot explain the other differences in these samples in Table 1. 6 Most PISA countries, including Vietnam, conducted testing on April 12-14 of 2012. Thus children born in 1996
would be from 15 years and 3 (completed) months of age (born in December of 1996) and 16 years to 2 (completed)
months (born in January of 1996). The target population was defined as “all students aged from 15 years and 3 com-
pleted months to 16 years and 2 completed months at the beginning of the assessment period” (OECD, 2014b, p.66).
7 PISA’s urban classification refers to schools not students Some students may live in rural areas and attend urban p
y
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p
(
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, p
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7 PISA’s urban classification refers to schools, not students. Some students may live in rural areas and attend urban
schools; they would be classified as urban in the PISA but as rural in the VHLSS and so may explain some of the
urban/rural difference in the two samples. Yet this cannot explain the other differences in these samples in Table 1. II. Is Vietnam’s Performance on the 2012 and 2015 PISA Assessments Exaggerated? Some observers, both Vietnamese and international, of Vietnam’s high performance on
the 2012 and 2015 PISA assessments have expressed surprise that Vietnam could perform so
well.5 This section investigates three possible mechanisms that could exaggerate Vietnam’s 3 performance: 1. The 15-year-old Vietnamese PISA participants in 2012 and 2015 were not
representative of 15-year-old Vietnamese students in those years; 2. Vietnam’s relatively low
enrollment rate for 15-year-olds selects higher-performing PISA participants; and 3. Vietnamese
students exert more effort when participating in the PISA assessments. A. Were Vietnam’s PISA Participants “Better than Average” 15-year-old Students? Were the Vietnam’s PISA participants in 2012 and 2015 representative of the students that those
assessments were intended to sample? Consider the 2012 PISA. In each participating country,
the 2012 PISA participants were to be a random sample of all children born in 1996 (and thus
were 15 years old in January of 2012) who were enrolled in school in 2012 (OECD, 2014b).6
Whether the students who participated in the 2012 PISA in Vietnam are a representative sample
of individuals born in 1996 who were students in 2012 can be considered using data from the
2012 Vietnam Household Living Standards Survey (VHLSS), which Vietnam’s General
Statistical Office conducts every two years on a sample of about 9,000 households in Vietnam. The 2012 VHLSS can be used to compare Vietnamese children in that survey who were born in
1996 and were students in 2012 with the Vietnamese students who participated in the 2012
PISA. Table 1 uses data from the 2012 PISA assessment and the 2012 VHLSS to assess the
representativeness of Vietnam’s PISA participants in 2012. These two data sources have several
discrepancies. Compared to the VHLSS data, the students who participated in the 2012 PISA are
more likely to be from urban areas (50% vs. 26%),7 are slightly more likely to be in grade 10 4 (86% vs. 84%) and less likely to be in grade 9 (10% vs. 14%), have more educated fathers (9.0
vs. 7.2 years of schooling) and mothers (8.3 vs. 6.8 years), and are more likely to live in homes
with air conditioners, cars, computers and televisions, and so are from wealthier families. 5-year-old students interviewed in the first two rounds of the 2012 VHLSS, about half were inter-
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d
i 8 Of the 236 15-year-old students interviewed in the first two rounds of the 2012 VHLSS, about half were inter-
viewed in March or April, and about half were interviewed in June. Only 5 were interviewed in May, and 4 in July. or April, and about half were interviewed in June. Only 5 were interviewed in May, and 4 in July. II. Is Vietnam’s Performance on the 2012 and 2015 PISA Assessments Exaggerated? The discrepancy regarding the likelihood of being in grades 9 and 10 is larger if one notes
that the 2012 PISA assessment was administered in Vietnam in April of 2012, which is a time
when 22% of the children born in 1996 are still in grade 9, as seen in the third column of Table
1. More specifically, of the children born in 1996 who were still in school and were interviewed
between March and July in the 2012 VHLSS (and thus had not yet reached the next grade of
schooling in September of 2012),8 76% were in grade 10, while 22% were in grade 9; in contrast,
of PISA participants in April of 2012, 86% were in grade 10 and only 10% were in grade 9. The
distinction between grades 9 and 10 is important in Vietnam, because almost all children
complete grade 9, but in many provinces, students must pass provincial entrance exams to enroll
in grade 10. Thus 86% of the students in the PISA sample are students who have passed an
exam that selects better performing students for upper secondary school, but the VHLSS data
indicate that only about 76% of children in Vietnam who were eligible to participate in the PISA
exam when it was administered (in April of 2012) were in grade 10 and thus had passed that
exam. Similar patterns are seen in the last four columns of Table 1, which compare the students
in the 2015 PISA (who were 15 years old when they took that test, and so should be a random
sample of students born in 1999) to an average of students who were 15 years old in 2014 (more
precisely, born in 1998) in the 2014 VHLSS data and students who were 15 years old in 2016
(more precisely, born in 2000) in the 2016 VHLSS data (this average is used because there is no 5 VHLSS for 2015). Relative to the averaged VHLSS data (focusing on those interviewed between
March and July), the 2015 PISA participants are more likely to be in urban areas (50% vs. 29%),
have more educated mothers (8.4 vs. 6.9 years of schooling) and fathers (7.9 vs. 6.4 years), and
are more likely to live in homes with air conditioners, cars, computers and televisions. II. Is Vietnam’s Performance on the 2012 and 2015 PISA Assessments Exaggerated? In contrast
to the findings for 2012, the students in the 2015 PISA assessment are only slightly more likely
to be in grade 10 (85.5%) relative to their counterparts in the averaged VHLSS data (84.3%). The differences in Table 1 between the PISA and the VHLSS data raise a question: How
would Vietnam’s students have scored on the PISA if the PISA sample had had the same student
characteristics as the VHLSS sample? This can be assessed by using the PISA data for Vietnam
to predict Vietnamese students’ performance on the PISA, assuming that this predictive power of
the student-level characteristics is valid for the same characteristics as measured by the VHLSS. More specifically, consider an ordinary least squares (OLS) regression that uses the PISA
data for Vietnam to predict students’ scores on that assessment based on the variables in Table 1:
PISAscorei = βʹXi + ui
(1) (1) where Xi is a vector, for student i, of the student characteristics in Table 1. The regressions for
the 2012 and 2015 PISA data, using Equation (1), are shown in Tables B1 and B2, Appendix B.9 A convenient property of OLS regressions is that the mean values of the explanatory
variables perfectly predict the mean value of the dependent variable. That is:
PISAscore
̅̅̅̅̅̅̅̅̅̅̅̅̅ = 𝛃̂OLSʹ𝐗̅PISA
(2) A convenient property of OLS regressions is that the mean values of the
variables perfectly predict the mean value of the dependent variable. That is: PISAscore
̅̅̅̅̅̅̅̅̅̅̅̅̅ = 𝛃̂OLSʹ𝐗̅PISA
(2) (2) PISAscore
̅̅̅̅̅̅̅̅̅̅̅̅̅ = 𝛃̂OLSʹ𝐗̅PISA where the horizontal bars indicate mean values and 𝛃̂OLS is the OLS estimate of β. This is shown
in the top halves of Tables 2 (math) and 3 (reading); the first columns depict 𝐗̅ from the 2012
PISA data in Table 1, the fourth column shows the β coefficients (from Table B1, Appendix B), 9These regressions have high predictive power; for example, the R2 is 0.341 for reading and 0.310 for math for the
2012 PISA data. Most variables are highly significant, and almost all of the signs are in the expected direction. 6 and the fifth column shows the product of each variable with its respective coefficient. Summing
the fifth column produces the actual PISA scores, 512.7 for math and 509.8 for reading. and the fifth column shows the product of each variable with its respective coefficient. II. Is Vietnam’s Performance on the 2012 and 2015 PISA Assessments Exaggerated? Summing
the fifth column produces the actual PISA scores, 512.7 for math and 509.8 for reading. These regression coefficients can also be used to predict what the 2012 PISA score would
have been if 𝐗̅ had been the means in the 2012 VHLSS data. The 2012 VHLSS means for the
interviews conducted from March to July of 2012 (since the PISA was administered in April of
2012), from the third column of Table 1, are shown in the second column of Tables 2 and 3, the
products of these variables and their coefficients are in the sixth column, and the predicted 2012
PISA scores are at the bottom of that column. When the 2012 VHLSS means are used to predict
the PISA scores, the math score by about 24 points, to 489.0, and the reading score declines by
about 20 points, to 489.5. Almost half of the difference between the 2012 PISA score and the
predicted score that adjusts for the potential non-representative sample is due to the larger
percentage of grade 10 students in the PISA sample, as seen in the last columns of Tables 2 and
3 10 Albania’s coverage rate, which is much higher for both the 2009 and 2015 PISA assessments, could be an error.
We thank Francesco Avvisati for pointing this out. 3. A similar analysis based on equation (2) for the 2015 PISA data and the average of the
2014 and 2016 VHLSS data is shown in the bottom of Tables 2 and 3. Using the means 𝐗̅ from
the averaged VHLSS data (the households interviewed from March to July), instead of the 2015
PISA data, Table 2 shows that the math score decreases by 13.6 points, and Table 3 shows that
the reading score decreases by 14.7 points. These are smaller than the drops for the 2012 data. The overall message from this exercise is that the differences in child, parent and house-
hold characteristics seen in Table 1 between the 2012 PISA sample and the 2012 VHLSS sample
imply a drop of only about 20-24 points (or 0.20-0.24 standard deviations) of Vietnam’s
performance on the 2012 PISA. Yet a quick glance at Figures 1 and 2 shows that Vietnam is still
an outlier even after doing this adjustment. A similar adjustment comparing the 2015 PISA with 7 7 the 2014 and 2016 VHLSS has an even smaller effect on Vietnam’s outlier status in Figures 3
and 4. Correcting, and Three Methods to Adjust for, Vietnam’s Low Enrollment Rate B. Correcting, and Three Methods to Adjust for, Vietnam’s Low Enrollment Rate. Another possible explanation for Vietnam’s strong PISA performance is that many Vietnamese
15-year-olds are not enrolled in school, and those who are not in school are likely to have lower
academic skills than those who are. Thus, one possible explanation for Vietnam’s strong
performance on the PISA assessments is that, relative to other PISA countries, a larger
proportion of Vietnam’s less academically inclined 15-year-olds did not participate in the PISA
assessment, which includes only 15-year-olds enrolled in school. Indeed, Vietnam’s “coverage
index” indicates that only 55.7% of its 15-year-olds participated in the 2012 PISA, primarily
because of this age group’s low enrollment rate (OECD, 2014a, Table A2.1). This is the third
lowest coverage rate of the 63 countries that participated in the 2012 PISA assessment; only
Albania (55.2%) and Costa Rica (49.6%) had lower rates.10 Vietnam’s lower coverage rate is
even more extreme in the 2015 PISA assessment; of the 66 participating countries, Vietnam’s
coverage rate was the lowest, at only 49% (OECD, 2016, Table I.6.1). The next lowest country,
Mexico, had a much higher rate of 62%. This subsection first corrects the Vietnam coverage
rates reported by the OECD, and then presents three different methods to adjust for Vietnam’s
(corrected) lower coverage rate. Even after this correction and these adjustments, Vietnam still
remains a positive outlier, given its low income, in its performance on the 2012 and 2015 PISA
assessments. 1. Correcting PISA Coverage Rates. Analysis of the PISA data demonstrates that
Vietnam’s coverage rates were incorrectly calculated, and correcting these errors leads to 8 sizeable increases in those rates. As explained in Appendix A, census data from Vietnam were
incorrectly used to calculate the number of 15-year-olds in Vietnam in 2012 and 2015. Correctly
applying the census data to the school enrollment data in the OECD reports shows that the
correct PISA coverage rates for 2012 and 2015 are, respectively, 65.9% and 65.6%. Yet even
after these corrections, 34.1% of 15-year-olds in Vietnam did not participate in the 2012 PISA
assessment, and 34.4% did not participate in the 2015 PISA assessment. These individuals were
likely weak students before leaving school, since most of the PISA participants were in grade 10
and, unlike grade 9, grade 10 students are a selected group, as explained above. 11 These differ slightly from the numbers in OECD (2014a) because sample weights were not used, for comparabil-
ity with columns 3 and 4, which cannot use sampling weights to exclude 15-year-olds who did not participate. Correcting, and Three Methods to Adjust for, Vietnam’s Low Enrollment Rate The rest of this
subsection applies three different methods to adjust PISA scores for differences in countries’
coverage rates. Method 1: Focus on Top 50%. One way to adjust each country’s performance to
account for differential participation in the PISA is to focus on the “top 50%” of 15-year-olds. This can be done by assuming that non-participating 15-year-olds would have scored in the
lowest 50% of the distribution of test scores among all 15-year-olds in their respective countries
if they had participated in the assessment, and then excluding the bottom 50% of 15-year-olds for
all countries. In fact, for countries with a lower coverage rate, such as Vietnam, this assumption
leads to underestimation of the performance of the top 50% of students since, for these countries,
it is more likely that some not in school would be in the top 50% if they were in school, which
means that some 15-year-olds classified as in the top 50% for these countries were in fact in the
bottom 50%. The results of doing so for the 2012 PISA assessment are shown in Table 4A. The first two columns of Table 4A show the widely reported scores in the PISA reports,
which include all test participants (and, of course, exclude nonparticipants).11 Vietnam ranks 16 11 These differ slightly from the numbers in OECD (2014a) because sample weights were not used, for comparabil-
ity with columns 3 and 4, which cannot use sampling weights to exclude 15-year-olds who did not participate. 9 out of 63 in math and 18 out of 63 in reading. However, when 15-year-olds in the bottom 50%
of the population of all 15-year-olds are excluded, using the method described above, 12 the
performance of Vietnam’s “top 50%” of 15-year-olds is less impressive, ranking only 34 out of
63 in math and 39 out of 63 in reading (see the third and fourth columns of Table 4A). The same exercise using the 2015 PISA results yields a similar conclusion. As seen in
Table 4B, Vietnam ranks 24 out of 66 countries in math and 28 out of 66 in reading. While these
rankings are not as high as the 2012 PISA rankings, this is still a very strong performance given
that Vietnam is the second poorest of all participating countries (see Figures 3 and 4). 12 Table 4A shows the mean scores of the top 50% of 15-year-olds under the assumption that those who did not par-
ticipate would not have scored in the top 50% had they participated. Mathematically, denote the coverage rate by c,
which is ≥ 50 for all countries except Costa Rica (its 49.6% rate is set to 50%). The goal is to drop the d% of the test
participants who were not in the top 50%, thus d = c - 50. Thus one must drop (d/c)×100%, i.e. ((c-50)/c)×100%, of
test participants. For each country the bottom ((c-50)/c)×100% of test takers were dropped, separately for each test. Correcting, and Three Methods to Adjust for, Vietnam’s Low Enrollment Rate However,
when 15-year-olds in the bottom 50% of Vietnam’s 15-year-olds are excluded, as shown in the
third and fourth columns of Table 4B, the performance of Vietnam’s “top 50%” of 15-year-olds
is less impressive, ranking only 37 out of 66 in math and 41 out of 66 in reading. Yet these lower rankings are still impressive when one recalls Vietnam’s relatively low
income. First, it still outperforms almost all other developing countries in the PISA, the sole
exception being that Chile’s top 50% of 15-year-olds outperformed Vietnam’s top 50% on the
2015 reading assessment (and note that Chile is much wealthier than Vietnam). Second, as
Figures 5 and 6 show, given that Vietnam was the poorest of the 63 countries in the 2012 PISA it
is still by far the largest positive outlier when the scores of the “top 50% of all 15-year-olds” are
plotted against log of per capita GDP. Although Vietnam’s “top 50%” scores in mathematics
and reading are not much higher than their “unadjusted” scores, and the increase in test scores of
the “top 50% of 15-year-olds” was much higher for other countries, the increases were highest in 10 the wealthier countries, which usually have high PISA participation rates. This increases the
slope of the lines in Figures 5 and 6, relative to Figures 1 and 2, and since Vietnam is at the far
left in these figures the higher slope makes it more of an outlier. This is also the case for the
2015 PISA assessment, as shown in Figures 7 and 8. Method 2: Bounds Analysis – Inferring Full Distribution Mean from Truncated
Distribution Mean. A second method to correct for Vietnam’s relatively low enrollment rate is
to build on the intuition of Method 1 that individuals not in school have lower academic skills in
order to estimate bounds on the average test score of all 15-year-olds for the countries that
participated in the PISA. To begin, assume that the PISA test scores follow a normal distribution
when the entire population of 15-year-olds is included. Figure B5 in Appendix B shows that this
assumption is reasonable for four 2015 PISA countries from different regions of the world that
had enrollment (coverage) rates above 90%: Australia, Germany, South Korea and Tunisia. Correcting, and Three Methods to Adjust for, Vietnam’s Low Enrollment Rate One
can derive a lower bound for the mean of the distribution of the test scores of all 15-year-olds
(students and non-students) by making a second assumption: that the test scores of all children
not in school, if they had participated in the PISA, would be lower than those of all children in
school. This assumption is illustrated in Figure B6. Under these two assumptions, PISA
participants constitute a normal distribution that is truncated from below (as in Figure B6). However, assuming that the test score of each 15-year-old PISA participant is higher than
the test scores of all 15-year-old non-participants is quite extreme. It is almost certain that some
PISA non-participants would have scored higher on that assessment than some participants. If
so, the adjustment described below would underestimate (and so provide a lower bound for) the
true mean (for all 15-year-olds). To see the intuition, consider Figure B6. Suppose that some
15-year-olds to the right of the truncation point did not participate in the PISA, and that some 15- 11 11 year-olds to the left of that point did participate. This would reduce the extent to which the mean
of the distribution of the PISA participants overestimates the mean of the distribution for all 15-
year-olds; applying the truncation formula that is proved below in Proposition 1 overcorrects and
so provides a lower bound of that mean. While standard formulas for the mean of a truncated normally distributed variable obtain
that mean using the mean of the overall (untruncated) distribution, our goal is to go in the other
direction; we want to obtain the mean of the overall distribution using the mean of the truncated
distribution. Proposition 1 below provides the formulas for doing this. Proposition 1: Estimating lower bounds and upper bounds of test scores Assume that the test scores of the entire population of 15-year-olds follow a normal distribution
with mean µ and standard deviation 𝜎. The truncated mean of this distribution is given by the
sample mean test scores 𝑇̅k, where k indexes truncation from above (a) or below (b), with 𝜏 being
the truncation point. Define 𝛼 as
𝜏−µ
𝜎, and let r represent the given school enrollment rate. 1.1. If the PISA’s tested samples capture only academically better-performing children (as in
Figure B6) the true mean test scores, denoted by µ lt (lt denotes lower truncation), is given by:
̅ 𝜇𝑙𝑡 = 𝑇̅𝑏− 𝜆𝑏(𝛼)
𝑇̅𝑏−𝑇𝑚𝑖𝑛
𝜆𝑏(𝛼)−𝛼
(1)
where 𝑇̅𝑏= 𝐸(𝑇|𝑇> 𝜏) , 𝛼= Φ−1(1 −𝑟), 𝜆𝑏(𝛼) =
𝜙(𝛼)
1−Φ(𝛼) , and the truncation point 𝜏 is
given by Tmin, the lowest observed test score in the data. (1) 1.2. If the PISA’s tested samples capture only the academically worse-performing children,
the true mean test scores, denoted by µ ut (ut denotes upper truncation) is given by
̅ 1.2. If the PISA’s tested samples capture only the academically worse-performing children,
the true mean test scores, denoted by µ ut (ut denotes upper truncation) is given by 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝑇𝑚𝑎𝑥−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼
(2) 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝑇𝑚𝑎𝑥−𝑇𝑎
𝜆𝑎(𝛼)+𝛼
(2)
where 𝑇̅ = 𝐸(𝑇|𝑇< 𝜏), 𝛼= Φ−1(𝑟), 𝜆𝑎(𝛼) =
𝜙(𝛼)
Φ(α) , and the truncation point 𝜏 is given
by Tmax, the highest observed test score in the data. (2) 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝑇𝑚𝑎𝑥−𝑇𝑎
𝜆𝑎(𝛼)+𝛼
(2)
where 𝑇̅ = 𝐸(𝑇|𝑇< 𝜏), 𝛼= Φ−1(𝑟), 𝜆𝑎(𝛼) =
𝜙(𝛼)
Φ(α) , and the truncation point 𝜏 is given
by Tmax, the highest observed test score in the data. The proof is given in Appendix A. Yet, since the assumptions that the PISA’s tested samples
capture only either the academically better-performing or the academically worse-performing
children in Proposition 1 are rather extreme, Equations (1) and (2) would provide, respectively, 12 lower bound and upper bound estimates of the true mean test scores. Equation (1) is the
derivation of interest given that the mean test score of the PISA participants is almost certainly
higher than the mean of the PISA non-participants (this is also shown below for the Young Lives
data from Vietnam). We show Equation (2) only for mathematical completeness. 13 For more information on the Young Lives data, consult its website: www.younglives.org.uk g
,
y
g
g
14 The only other PISA country for which there are Young Lives data is Peru. 15 Another possibility is that Vietnamese teachers provided answers to students, along the lines that Jacob and Levitt
(2003) found in Chicago public schools. Unfortunately, we cannot apply most of the methods that that paper used to
check for such cheating because we do not have panel data. Also, students taking the PISA exam are given many
different versions of the test so that any given student does not have the identical questions as the students sitting
nearby. Multiple versions of the test also make it much harder for teachers to provide students the correct answers. Proposition 1: Estimating lower bounds and upper bounds of test scores Applying Equation (1) of Proposition 1 to the PISA math test, Figure 9 shows these
estimates, as well as the sample means of the scores of the observed (truncated) distributions of
2012 PISA participants, which one can view as upper bounds, as explained above. Figure 10
shows the same for the 2015 PISA assessments. The gap between these two bounds is a
decreasing function of the enrollment (coverage) rate, and will equal zero when the enrollment
rate is 100%. The bounds in Figures 9 and 10 for Vietnam, and for some other countries, are
rather wide. A “natural” approximation of the test score means that would be observed if all 15-
year-olds in each country were tested is the mid-point between the lower bound derived in
Proposition 1 and the upper bound given by the observed mean for the truncated sample, that is
the midpoints of the gaps in Figure 9. These mid-point values are shown in Figures 11-14,
where the countries are again plotted against their log of per capita GDP. Vietnam still stands
out as an outlier, especially for math. Method 3: Adjustment with Auxiliary Data. A third way to adjust the mean of the test
scores for Vietnamese students to include the scores of the PISA non-participants is to use the
Young Lives data.13 This is done only for Vietnam, because only one other PISA country has
such data;14 yet doing this adjustment only for Vietnam will be biased against Vietnam being an
outlier. The younger cohort in the Young Lives Study were 15 years old in Round 5 of that
study. This round, which took place in 2016, included administration of math and reading 13 comprehension tests to all 15-year-olds in that sample, about 1,940 15-year-olds, both those in,
and those not in, school; as expected, those not in school had lower average math (9.4 out of 21)
and reading (10.9 out of 25) scores than those in school (15.5 and 14.8). Assuming that the
Young Lives reading and math scores rank 15-year-olds in a way that is similar to the PISA test
rankings, one can adjust the observed PISA test scores to include 15-year-olds who are not in
school; see Appendix A for the details of this method. Proposition 1: Estimating lower bounds and upper bounds of test scores Making this adjustment to the 2012 PISA
scores reduces the mean math score for Vietnam by about 13 points, from 511.2 to 498.4, and the
mean reading score by about 11 points, from 508.2 to 496.9. The same adjustment for the 2015
PISA has similar reductions, about 12 points for math and 11 for reading. These relatively small
changes do not change the overall finding that Vietnam’s PISA performance was exceptional. C. Were Vietnamese Students More Motivated and Better Prepared for the PISA? A final possible explanation for Vietnam’s performance in the PISA assessments is that
Vietnamese students really did outperform those of most other countries, but not due to higher
skills; rather, they were highly motivated when they took the PISA tests, and they received
extensive preparation for those tests.15 No studies have examined the motivation of Vietnamese
students when taking international tests, but there are many anecdotes that Vietnamese students
(and their teachers) are very competitive test takers. In contrast, there is evidence that students in
developed countries exert little effort on tests for which there are no consequences. Gneezy et al. (2019) administered tests based on questions from previous PISA math
t
t t Chi
t d
t
d U S
t d
t
Th Chi
t d
t
d
h hi h
th
U S Gneezy et al. (2019) administered tests based on questions from previous PISA math
tests to Chinese students and U.S. students. The Chinese students scored much higher than U.S. Gneezy et al. (2019) administered tests based on questions from previous PISA math tests to Chinese students and U.S. students. The Chinese students scored much higher than U.S. 14 students under standard conditions. However, randomly selected U.S. students who were offered
financial incentives for high scores on the exam performed much better (22-24 points higher),
while Chinese students performed no differently. The lack of an effect for Chinese students
suggests that they are highly motivated to take tests despite no direct benefits. Vietnamese
culture has many similarities to Chinese culture, and thus it is possible that Vietnamese students’
intrinsic motivation to do well on tests increased their PISA scores by 22-24 points relative to
U.S., and perhaps other, students. There is also evidence that teachers and schools prepared Vietnamese students to take the
2012 and 2015 PISA tests.16 Studies in the U.S. and elsewhere have shown that preparation
sessions for academic tests can greatly increase students’ scores. For example, Bangert-Drowns
et al. (1983), summarizing a large number of studies, found that programs involving coaching
sessions of more than nine hours total duration increased average test scores by 0.39 standard
deviations (of the distribution of test scores, which for the PISA is equivalent to 39 points). A rough estimate of the combined impact of intrinsic motivation and preparation for the
PISA exam would be 62 points (23 from being more motivated, and 39 from exam preparation). 16 When Vietnamese students took a draft version of the PISA exam in 2011 in preparation for the 2012 PISA, their
performance was lower than expected, and Vietnam’s Ministry of Education and Training took several steps to
increase their performance. This does not violate rules of the PISA assessment; schools can have students practice,
using old exams, to become “accustomed” to PISA exams. In each country, the schools that participate in the PISA
exam are selected several months before the exam, and the students who participate are selected 3-4 weeks before the
exams. The selected Vietnamese students were told that a strong performance would bring honor to Vietnam, and
were given special t-shirts indicating that they were PISA participants. The information on the implementation of the
PISA in Vietnam is based on emails and discussions with Francesco Avvisati, who works on the PISA for the OECD. 17 For example, when presenting an earlier version of this paper in Colombia, we were told that Colombia has made
similar efforts to increase its students’ performance on the PISA, but their efforts were not particularly effective. C. Were Vietnamese Students More Motivated and Better Prepared for the PISA? This would explain about half of Vietnam’s exceptional performance in terms of the positive
residuals discussed below (Table 5, columns 1 and 2), assuming that all other countries that
participated in the PISA took no steps to increase their students’ test scores and had students as
unmotivated as U.S. students. Yet this assumption is rather extreme, and anecdotal evidence 15 suggests that other countries also try to increase their students’ performance on the PISA.17 Thus
these two factors together likely explain less than half of Vietnam’s exceptional performance. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? The evidence in Section II shows that 15-year-old students in Vietnam scored unusually
high on the 2012 and 2015 PISA assessments given Vietnam’s low GDP per capita, even after
adjusting for the possibility that the PISA sample was not representative of the 15-year-olds
enrolled in school and for the low enrollment rate of 15-year-olds in Vietnam. Presumably there
is some reason why Vietnamese students outperform those in other countries conditional on per
capita GDP. This section uses the PISA data to investigate Vietnam’s performance on the PISA. A. From Country Level to Student Level Regressions. Figures 1-4 in Section II are
based on the following simple linear regression equation: Test Score = β0 + βgdp×Log(GDP/capita) + u Test Score = β0 + βgdp×Log(GDP/capita) + u
(3)
In these figures, the gap between any country’s actual performance on the test and its predicted
performance given its (log) GDP per capita is given by u in equation (3). These figures show that
Vietnam has a very high value of u. The regressions that generated these figures have one
observation per country, yet analogous regressions with one observation per student for each
country participating in the PISA assessments yield the same finding. Such regressions, which
regress the student-level PISA test score data on a constant term and the log of per capita GDP,
are shown for the 2012 PISA data in the first two columns of the top half of Table 5. As
expected, the coefficient on GDP per capita is positive: countries with a higher GDP tend to have
higher scores. However, Vietnam’s test scores in the 2012 PISA are much higher than those (3) 16 predicted by this regression. In particular, for the math regression Vietnam’s average value of u
(the residual, shown in bold in the fifth row of Table 5) is 135.8, and for the reading regression it
is 119.0. These are the highest values among all the countries included in the regression, as
indicated by the “Residual Rank” row in Table 5, just as Vietnam is the largest positive outlier in
the country-level regressions that generated Figures 1-4. The question is: Why is Vietnam’s residual so high? In particular, would adding more
variables to the regression result in a “better fit” in which the (average) residual for Vietnam
would not be so high? oner could not be used since it was collected for some countries (including Vietnam) but not others. 18 Air conditioner could not be used since it was collected for some countries (including Vietnam) but not others. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? This question is addressed in the rest of this section, first by adding
household and student level characteristics, and then adding by school characteristics, using
information from the 2012 and 2015 PISA data, which not only administered reading and
mathematics tests but also collected data from students, parents and schools. The remaining columns in Table 5 explore the simple relationship between student test
scores and national and household level income and wealth. One disadvantage of the regressions
in the first two columns of that table is that the variable for the log of GDP per capita does not
vary over students in the same country; ideally, it would be useful to have a wealth or income
variable that varies within countries, which should provide more explanatory power in student-
level regressions. A wealth variable can be generated from the PISA data by using information
on students’ households from the student questionnaire. This was done by applying principle
components analysis to the following household level variables in the PISA: internet connection,
dishwasher, DVD, number of cell phones, number of televisions, number of computers, and
number of cars.18 The first estimated principle component is used as a wealth variable in the
analysis of this section. The third and fourth columns of Table 5 show that, for the 2012 PISA, 17 when this variable is used instead of the log of GDP per capita, Vietnam is still the largest outlier
in the math regression, though it is only the second largest outlier in the reading regression, after
Hong Kong. The rest of the analysis of this paper will use this wealth variable instead of log of
GDP per capita because the former varies across students within each country in the PISA data. Before adding other variables to equation (3), which is the focus of this section, the last
four columns of Table 5 explore two aspects of the wealth variable that was generated by
principal components analysis. First, the third and fourth columns in Table 5 use country
averages of the wealth variable, for comparability with the first two columns in that table, which
are based on the log of GDP per capita. In contrast, the fifth and sixth columns allow each
student to have his or her own household-specific value of wealth, instead of the national
average. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? This allows the wealth variable to explain not only the differences in test scores across
countries but also within countries. This reduces the coefficients on the wealth variable
somewhat, but it is still highly significant. More interesting is that Vietnam falls slightly in
terms of its outlier status. For math it is now the fourth highest outlier, while for reading it is the
second highest. The main reason for this is that the predictive power of the wealth variable falls
by about one fifth when it varies within countries, which indicates that it has a stronger role to
play when explaining differences between countries than within them. This drop in the
coefficient in effect leads to a less steep slope in the fitted lines in Figures 1-4, reducing the size
of Vietnam’s residual and increasing the size of the residuals for the wealthiest top performers,
such as Hong Kong, Singapore and South Korea. Yet Vietnam is still a large outlier, and much
poorer than these other outlier countries. Second, the last two columns of Table 5 add country fixed effects, which again reduces
the impact of wealth somewhat. The reported residuals in those two columns are simply the 18 estimated country fixed effects. Again, Vietnam is still an outlier, although slightly less of an
outlier in that it has the fifth highest fixed effect for math and the third highest for reading. A similar analysis is done for the 2015 PISA data in the bottom half of Table 5. The
overall pattern is the same. The average residual for Vietnam slightly decreases when the
average wealth variable is used instead of GDP per capita, and decreases slightly more when the
wealth variable is allowed to vary at the student level, but Vietnam is still one of the largest, if
not the largest, outlier. This is also the case when country fixed effects are used. Again, the
countries that occasionally are larger outliers than Vietnam are much wealthier than Vietnam. B. Adding Other Variables to Explain Vietnam’s Performance. The student-level
regressions with country fixed effects in the last two columns of Table 5 are a useful starting
point for a more systematic analysis to find characteristics of Vietnamese students, households,
teachers and schools that explain Vietnam’s outlier status in the 2012 and 2015 PISA
assessments. To begin, assume that the underlying skill (e.g. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? math) measured by the PISA test
score of student i in country c, denoted by Sic, is a linear function of the characteristics of that
student, of his or her household, the teachers he or she has had, and the school(s) he or she has
attended: assessments. To begin, assume that the underlying skill (e.g. math) measured by the PISA test
score of student i in country c, denoted by Sic, is a linear function of the characteristics of that
student, of his or her household, the teachers he or she has had, and the school(s) he or she has
attended: Sic = βʹxic + εic
(4) Sic = βʹxic + εic
(4) (4) (4) where the xic variables are all the student, household, teacher and school characteristics that
affect students’ underlying skills, β measures the causal impacts of those characteristics on that
skill, and εic is measurement error in the PISA test. The linearity assumption is not very
restrictive since xic could include higher order and interaction terms. An important distinction to make is between the observed and unobserved xic variables:
5 An important distinction to make is between the observed and unobserved xic variables: An important distinction to make is between the observed and unobserved xic variables:
Sic = βoʹxico + βuʹxicu + εic
(5) important distinction to make is between the observed and unobserved xic variables Sic = βoʹxico + βuʹxicu + εic
(5) (5) Sic = βoʹxico + βuʹxicu + εic
( Sic = βoʹxico + βuʹxicu + εic 19 = βoʹxico + βuʹ𝐱̅cu + βuʹxicu,d + εic = βoʹxico + βuʹ𝐱̅cu + βuʹxicu,d + εic = βoʹxico + βuʹ𝐱̅cu + βuʹxicu,d + εic where the superscripts o and u indicate observed and unobserved, respectively. The second line
of equation (5) disaggregates xicu into its country specific mean, 𝐱̅cu, and the within-country
deviation from that mean for student i, xicu,d, where the superscript d indicates that deviation. This disaggregation implies that the within-country mean of xicu,d equals zero for all countries. where the superscripts o and u indicate observed and unobserved, respectively. The second line
of equation (5) disaggregates xicu into its country specific mean, 𝐱̅cu, and the within-country
deviation from that mean for student i, xicu,d, where the superscript d indicates that deviation. This disaggregation implies that the within-country mean of xicu,d equals zero for all countries. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? In a regression with country fixed effects, the fixed effect for country c would be βuʹ𝐱̅cu,
and the error term would be βuʹxicu,d + εic. The last two regressions in Table 5 have only one
observed variable, the wealth indicator. The goal of the rest of this section is to add additional
variables to equation (5), which in effect moves those variables out of xicu and into xico in that
equation, to see whether Vietnam’s outlier status can be explained by observed variables in the
PISA data. This approach was used by Fryer and Levitt (2004) to investigate the gap in test
scores between black and white students in the U.S., and by Singh (2019) to explain differences
in test scores of primary and secondary school age children across Ethiopia, India (Andhra
Pradesh), Peru and Vietnam. If the PISA data contain the key factors that explain Vietnamese
students’ success, then adding them as regressors will yield small and statistically insignificant
country fixed effect for Vietnam by removing the variables that contribute to the βuʹ𝐱̅cu term in
the second line of Equation (5). If all variables are included that explain the performance of all
the countries in the PISA data set, then all country fixed effects will become insignificant and the
error term will become the (within-country) variation in the measurement error, εic. Even if the PISA data lack some of the key variables that explain Vietnam’s success, and
more generally explain student learning in all the countries that participated in the 2012 and 2015
PISA assessments, it may be that the country fixed effects, while statistically significant, are
greatly reduced and thus at least part of the reasons for Vietnam’s success would be explained by 20 20 the PISA data. Even if Vietnam is still one of the largest outliers, it may be a much smaller
outlier – relative to the overall variation in the PISA test score data – after adding the variables
available in the PISA data. In contrast, if the student, household, teacher and school variables
that explain Vietnam’s success are for the most part not in the PISA data, then Vietnam will
continue to be a large, positive outlier and the reason(s) for its outlier status will be due to factors
that are not measured, or at least not well measured, in the PISA data. the PISA data. 19 Missing values were particularly common for the sibling index. To avoid losing observations due to that variable
being missing, missing values were assigned its average value and a variable was added indicating that it is missing. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? Even if Vietnam is still one of the largest outliers, it may be a much smaller
outlier – relative to the overall variation in the PISA test score data – after adding the variables
available in the PISA data. In contrast, if the student, household, teacher and school variables
that explain Vietnam’s success are for the most part not in the PISA data, then Vietnam will
continue to be a large, positive outlier and the reason(s) for its outlier status will be due to factors
that are not measured, or at least not well measured, in the PISA data. To begin, student and household level variables from the 2012 PISA assessment are
added to the regression equation in Table 6A. The first two columns of that table show
regressions identical to those in the last two columns of Table 5, except that the sample size is
reduced so that the sample is identical to that in the third and fourth columns of Table 6A, which
add four additional household variables. The estimates in the first two columns of Table 6A are
very similar to those in the last two columns of Table 5; the rank of Vietnam’s estimated country
fixed effects is the same, and the countries with larger fixed effects are also the same. The third and fourth columns of Table 6A add four additional household characteristics
that are “pre-determined” and may also explain students’ test performance: a dummy variable for
girl students, an index of the number of siblings in the home (0 = none, 1 = brothers but no
sisters, or sisters but no brothers, and 2 = sisters and brothers); and mother’s and father’s years of
schooling. Each of these household variables has some missing values, which reduces the
sample size to 401,489, compared to 455,971 in the last two columns of Table 6.19 The key question for Table 6A is whether adding these additional household-level
variables “explains” much of the very large country fixed effect found for Vietnam when house-
hold wealth is the only regressor. The third and fourth columns in Table 6A shows that adding 21 these four variables to the regression reduces the explanatory power of the wealth index variable
by about one third (although it is still highly significant) but it has very little impact on the
Vietnam country fixed. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? Indeed, these fixed effects increase slightly, from 78.2 to 80.6 for the
math test and from 68.3 to 70.7 for the reading test. Vietnam’s outlier status is also largely
unchanged; its fixed effect in the math regression drops from fifth place to sixth place, but
increases from third place to second place in the reading regression. Thus, these four household-level variables in the PISA data do not explain Vietnam’s
strong performance on the 2012 PISA assessment. This is not surprising when the means of
these variables are compared for Vietnam and these other countries. In particular, Table 8 shows
that the average of Vietnam’s sibling index is almost identical to that of the other PISA countries
combined (1.048 vs. 1.086, respectively), and that Vietnamese parents have, on average, fewer
years of schooling (8.3 for mothers and 8.9 for fathers) than do parents in the other PISA
countries (11.0 for mothers and 11.1 for fathers), so these variables cannot explain why Vietnam
outperforms other countries; indeed, its lower parental education levels make its performance all
the more remarkable. The 2012 PISA data contain several variables that are directly related to students’
education, such as the grade they are in, years of preschool, several educational inputs, days of
school attendance (in the past two weeks), books in the home not related to the child’s schooling,
and hours per week in tutoring classes. These variables are likely to be endogenous (parents may
provide more educational inputs to children not doing well at school, or perhaps to their most
promising children), so adding them to the regression analysis likely produces biased estimates
of the causal impacts of these variables. Despite this possible bias, these variables may provide
informative explanatory power that may shed light on why Vietnamese students perform so well 22 on the PISA. For example, Table 8 shows that, on average, Vietnamese students spend more
hours per week in tutoring classes (1.3 for reading, 2.7 for math) than do students in other PISA on the PISA. 0 Unfortunately, data on tutoring classes and on hours studying at home were not collected for Viet y,
g
y g
PISA, so those variables can be used only for the 2012 analysis. y
y
21 The education input index is the first principal component of the following variables: quiet place to study, desk,
educational software, classical literature books, poetry books, educational books, technical books, and a dictionary. 20 Unfortunately, data on tutoring classes and on hours studying at home were not collected for Vietnam for the 2015
PISA so those variables can be used only for the 2012 analysis data o tuto
g c asses a d o
ou s study g at o e we e ot co ected o V et a
o t e 0 5
ariables can be used only for the 2012 analysis. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? For example, Table 8 shows that, on average, Vietnamese students spend more
hours per week in tutoring classes (1.3 for reading, 2.7 for math) than do students in other PISA
countries (0.9 for reading, 1.3 for math), so even if one cannot estimate the causal impact of
these classes on student test scores, even biased estimates may reduce Vietnam’s outlier status.20 The last two columns of Table 6A add these more education-focused child and household
variables to the regression that had only gender, parental education and the sibling index variable
(this reduces the sample size, so the fifth and sixth columns show the results with only household
wealth but the same samples as in the last two columns). Adding these variables further reduces
the coefficient on household wealth, and reduces Vietnam’s estimated country fixed effect (from
79.1 to 65.0 for math, and from 68.9 to 55.1 for reading), but it does not reduce Vietnam’s outlier
status: it remains the fifth highest outlier for math and the third largest for reading. Again, the
reason for this is that, for some education variables added to the regression, Vietnamese students
have lower average values than do the students in the other PISA countries. For example,
Vietnamese students have fewer educational inputs21 and fewer books at home (see Table 8). Vietnamese students have fewer educational inputs21 and fewer books at home (see Table 8). A similar analysis for the 2015 PISA data is presented in Table 6B. The overall results
are similar. The country fixed effect for Vietnam changes very little when additional household-
level variables are added to the regression, and Vietnam is always one of the top five positive
outliers. Also, all of the other top five outliers are much wealthier than Vietnam. Since child and household variables in the PISA data do little to explain Vietnam’s
exceptional performance (outlier status) in education, perhaps that performance is due to better
schools and teachers. This is examined in Tables 7A and 7B, which add school and teacher 23 characteristics to the regressions. As before, the first two columns show, for comparison
purposes, regressions that include only the wealth variable, but have the same samples as the
regressions that include the school and teacher variables. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? The third and fourth columns of Tables 7A and 7B show regression results that add not
only child and household variables (which are not shown to reduce clutter) but also school and
teacher variables. The school and teacher variables are: class size; the proportion of teachers
who have the required qualifications; computers per student; a variable indicating whether
student performance is used to assess teachers’ performance (a higher value indicates a “no”
response); an indicator of teacher absenteeism; an index of whether parents put pressure on
teachers (2012 only); two variables indicating whether school principals and outside inspectors,
respectively, observe teachers in the classroom; an indicator of the extent to which student
performance determines teacher pay (2012 only); and an index that measures teacher mentoring. Most of these school characteristics have the expected signs, but the key question is whether they
can “explain” at least part of Vietnam’s outlier status as measured by its country fixed effect. The results for the 2012 PISA in Table 7A show that adding school and teacher variables
reduces Vietnam’s outlier status in the sense that Vietnam’s estimated fixed effects are reduced
by nearly one fourth (from 76.7 to 58.1) for math and almost one third (from 66.1 to 44.7) for
reading. Yet little has changed in the sense that adding these variables to the math test yields
only a small reduction in the rank of Vietnam’s estimated fixed effect (from five to eight), and its
rank for the reading test (four) is unchanged. The same analysis using the 2015 PISA data is shown in Table 7B, and the results are
similar. In particular, adding school and teacher variables reduces Vietnam’s country fixed 24 effects by about one fifth for math (from 71.4 to 59.2) and for reading (from 59.2 to 46.5), but its
relative rank declines by only one for math and five for reading. effects by about one fifth for math (from 71.4 to 59.2) and for reading (from 59.2 to 46.5), but its
relative rank declines by only one for math and five for reading. To summarize, this section shows that the observed child, household, school and teacher
variables in the PISA data explain very little of Vietnam’s impressive performance on the 2012
and 2015 PISA assessments relative to its income level. III. What Observed Variables in PISA Explain the Gaps Conditional on Income? At most, adding these variables explains
one fourth of Vietnam’s exceptional performance in math and one third of its exceptional
performance in reading. Thus, most of the explanation for that performance must be found
elsewhere. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? The analysis in the previous section assumed that the impact of each variable on test
scores is the same for all 63 countries in the analysis. But perhaps Vietnam’s exceptional
performance is partly due to it being “more effective” in using various “inputs”. For example, it
may be that each year of Vietnamese parents’ years of schooling represents a higher level of
cognitive skills than does the average year of parental schooling in the other PISA countries. To explore this possibility, a Oaxaca-Blinder decomposition (Blinder, 1973; Oaxaca,
1973) is applied to differences in test scores between Vietnam and all other countries. Test
scores (S) are assumed to be linear functions of the variables used in the last two columns of
Tables 7A and 7B, again denoted by x. The impacts of these variables on test scores, denoted by
β, are allowed to differ between Vietnam and the other countries in the PISA assessment. This
yields the following regression equations (omitting the i subscript to reduce clutter): Svn = βvnʹxvn + uvn (Vietnam)
(6)
So = βoʹxo + uo (Other countries)
(7) Svn = βvnʹxvn + uvn (Vietnam)
(6)
So = βoʹxo + uo (Other countries)
(7) 25 The constant term in both of these equations can be normalized so that the means of the residuals
equal 0. Taking the mean of both sides of each regression equation gives the following The constant term in both of these equations can be normalized so that the means of the residuals
equal 0. 22 Note that this decomposition holds algebraically even when 𝛃̅ is replaced by any vector β of the same dimension;
the arithmetic average is used here for its intuitive appeal. Other β’s have been suggested in the literature. See
Fortin et al. (2011) and Jann (2008) for further discussion of this decomposition. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? Taking the mean of both sides of each regression equation gives the following
i
f
th
t
t
i Vi t
S̅
d i th
th
62 PISA
t i
S̅ expressions for the average test scores in Vietnam, S̅vn, and in the other 62 PISA countries, S̅o: expressions for the average test scores in Vietnam, S̅vn, and in the other 62 PISA countries, S̅o: S̅vn = βvnʹ𝐱̅vn (8)
S̅o = βoʹ𝐱̅o
(9) S̅o = βoʹ𝐱̅o
(9) (9) The standard Oaxaca-Blinder decomposition uses equations (8) and (9) to express the
difference in the mean test scores between Vietnam and the other PISA countries as follows: S̅vn – S̅o = βvnʹ𝐱̅vn – βoʹ𝐱̅o
(10)
= βvnʹ𝐱̅vn – βoʹ𝐱̅o + βoʹ𝐱̅vn – βoʹ𝐱̅vn
= βoʹ(𝐱̅vn –𝐱̅o) + (βvn – βo)ʹ𝐱̅vn (10) = βoʹ(𝐱̅vn –𝐱̅o) + (βvn – βo)ʹ𝐱̅vn Thus, the difference in the average test scores in Vietnam and the average scores in the other
countries has two components. The first is the difference in the means of the x variables between
Vietnam and the other countries, multiplied by the β for the other countries (denoted by βo). The
second is the difference between Vietnam and the other countries in the “effectiveness” of the x
variables, βvn – βo, multiplied by the means of Vietnam’s x variables (denoted by 𝐱̅vn). Thus, the difference in the average test scores in Vietnam and the average scores in the other One criticism of equation (10) is that the differences in the means of the explanatory
variables (𝐱̅vn –𝐱̅o) are “weighted” by the coefficient for the other 62 countries (βo) while the
differences in the coefficients (βvn + βo) are weighted by the means of the explanatory variables
for Vietnam (𝐱̅vn). Intuitively, it seems unbalanced that these weights are all from one group or
the other group; it may be better for the weights to be weighted averages of the β’s and the 𝐱̅’s of
both groups. 23 These means are for the sample for which the Oaxaca-Blinder composition is implemented. Observations with
missing values for the x variables are dropped, and so the means are slightly different from those in previous tables. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? One way to do this is to take the average of the two vectors of β coefficients and
use that β as the weight for the differences in the means; this yields the following decomposition: (11) S̅vn – S̅o = βvnʹ𝐱̅vn – βoʹ𝐱̅o 26 = 𝛃̅ʹ(𝐱̅vn –𝐱̅o) + [(βvn – 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo)ʹ𝐱̅o] where 𝛃̅ = (βvn + βo)/2.22 The first term, 𝛃̅ʹ(𝐱̅vn –𝐱̅o), weights the influence of the differences in
the x variables by the simple average of the two β coefficients. The second term, (βvn – 𝛃̅)ʹ𝐱̅vn +
(𝛃̅ – βo)ʹ𝐱̅o, accounts for the influence of the differences in the βvn and βo coefficients by splitting
that difference into two parts, the difference between βvn and 𝛃̅, weighted by 𝐱̅vn, and the differ-
ence between 𝛃̅ and βo, weighted by 𝐱̅o. As in the original Oaxaca-Blinder decomposition, the
first term “explains” how much of the difference in the mean test scores between Vietnam and
the other countries is due to Vietnamese students having different characteristics than the other
students, and the second term “explains” how much of the difference stems from the differences
in the impacts of the various x variables, as measured by the difference between βvn and βo. In addition to decomposing the differences in the mean test scores, S̅vn – S̅o, into the
above two components (the “explained” part due to differences in the x’s and the “unexplained”
part due to differences in the β’s), both components can be further decomposed into the
contributions of the individual variables, which sum up to equal the overall component. For
example, one variable used below measures hours per week that children receive math tutoring,
which for the 2012 PISA is much higher in Vietnam (2.7) than the average for the other PISA
countries (1.3), as seen in Table 8. This tutoring could explain Vietnam’s strong performance by
contributing to the first component, 𝛃̅ʹ(𝐱̅vn –𝐱̅o). That is, part of this component is β̅t(x̅vn,t – x̅o,t),
where the t subscript indicates that this is the tutoring variable. Similarly, the impact of tutoring
could also contribute to the second component via the difference in the βt coefficients; its
contribution to the second component is [(βvn,t – β̅t)ʹx̅vn,t + (β̅t – βo,t)ʹx̅o,t]. 27 However, there are some potential problems with determining the roles played by
specific variables in these decompositions. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? First, sets of categorical variables, such as region or
ethnic group codes, require an omitted (base) category, and different base categories can produce
different results when assessing the impacts of the differences in the mean values of the x However, there are some potential problems with determining the roles played by
specific variables in these decompositions. First, sets of categorical variables, such as region or
ethnic group codes, require an omitted (base) category, and different base categories can produce
different results when assessing the impacts of the differences in the mean values of the x
variables across the two groups. Fortunately, this is not a problem here because this paper does
not use any sets of categorical variables. Second, variables that do not have “natural” zero points
can yield different results when assessing the impacts of the differences in the values of the β
terms across the two groups; adding an arbitrary constant to such variables will change the
contribution of the difference in the β terms because that difference is multiplied by the mean of
that variable, and the mean has changed. While almost all variables in the regressions have
natural “zeros”, as explained above the wealth index and the education inputs index were
constructed by using principal components analysis, and the first principal components for both
of these variables take both positive and negative numbers. Both of these variables are “re-
centered” by adding a constant that ensures that their minimum values are close to zero. Table 8 shows the means of the x variables separately for Vietnam and for the other PISA
countries for both PISA assessments. The 2012 means are also shown in the second and fifth
columns of Table 9A (and Table 10A). The bottom of the Table 9A shows the mean math test
score for Vietnam, 516.5 (in the third column), which is S̅vn, and the mean math test score for the
other 62 countries, 462.8 (the sixth column), which is S̅o.23 The gap between these two means is
53.7. Similarly, Table 10A shows that the gap between the two mean reading scores is 40.3. 24 The impact of this variable is similar in both Vietnam (6.77) and in the other 62 countries (7.72), although it is
statistically significant only for the 62 countries, which reflects the much larger sample size for that estimate. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? These gaps are smaller than the average residuals for Vietnam in Table 5 because those residuals
effectively compared Vietnam to a hypothetical “typical” other country that had the same level These gaps are smaller than the average residuals for Vietnam in Table 5 because those residuals
effectively compared Vietnam to a hypothetical “typical” other country that had the same level 28 of wealth as Vietnam, while the gaps in Tables 9A and 10A compare Vietnam, which has a
relatively low wealth of 4.14, with the other 62 countries, that have a higher wealth of 6.10. Returning to Table 9A, the x variables that have higher means in Vietnam than in the
other 62 countries, and for which the corresponding 𝛃̅ coefficients are positive, can potentially
explain part of the gap between the mean test scores in Vietnam and the other 62 countries in the
2012 PISA assessment. That is, the contribution of such variables to the 𝛃̅ʹ(𝐱̅vn – 𝐱̅o) component
in equation (10) is positive. The contribution is also positive when the mean for Vietnam is
lower than for the other 62 countries and the corresponding 𝛃̅ coefficient is negative. An
example of the former is the teacher mentoring variable. This is higher in Vietnam than in other
countries, and as expected mentoring of teachers is estimated to increase students’ test scores.24 In contrast, if Vietnam’s mean is higher but the corresponding 𝛃̅ coefficient is negative,
or Vietnam’s mean is lower and the corresponding 𝛃̅ coefficient is positive, this widens the gap
and thus makes the gap even harder to explain. For example, the mean years of schooling of
mothers and of fathers is lower in Vietnam than in the other 62 countries, and as expected the
corresponding β coefficients are positive, so the parent education variables cannot explain why
Vietnamese students’ scores are higher than those of other countries’ students, and so these
variables “increase the burden” on other variables to explain that gap. Table 9A provides the results of the Oaxaca-Blinder decomposition for the 2012 PISA
mathematics test. Recall that the overall gap to explain is 53.7 points. The differences in the x
variables, which are expressed as the βoʹ(𝐱̅vn –𝐱̅o) component of the decomposition, cannot
explain the gap. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? Indeed, summing over all of the x variables shows that the values of the x 29 variables lead one to expect a slightly larger gap, with an overall contribution of -1.6 (see the
bottom of the second to last column in Table 9A). Instead, the decomposition indicates that the
entire gap is due to the difference in the β coefficients; on average, Vietnam is “more efficient”
in “converting” x variables into higher test scores; this is seen in the last column in Table 9A. Given that it is the differences in the β coefficients that “explain” Vietnam’s remarkable
performance on the 2012 PISA assessment, it is of interest to see which specific variables seem
to contribute the most to this component of the decomposition. The variable in the last column
of Table 9A that plays the most important role is the percentage of students in grade 10, which
accounts for about seven eighths (88%) of this differential efficiency of the β coefficients. Quite
simply, on average for other 2012 PISA countries, moving a student from grade 9 to grade 10
without changing any other characteristic increases his or her test score by 18.9 points, but in
Vietnam this increases a students’ score by 85.9 points. This almost certainly reflects the fact
that movement from grade 9 to grade 10 in Vietnam is a selection process based on province-
level exams, which removes the grade 9 students with the weakest academic performance. But it
is also likely that, apart from this selection effect, an additional year of school in Vietnam leads
to more learning than an addition year in other countries; recall from Section II that correcting
for the very low “coverage rate” in Vietnam explained only a small part of its exceptional
performance. Moreover, as seen in the next paragraph, additional days of attendance have a
larger impact in Vietnam than in the other 2012 PISA countries (on average). The other variables for which the higher β coefficient for Vietnam explains a substantial
portion of the gap (a contribution greater than 20 points) are: 1. Using student performance as
part of the formal process to evaluate teachers’ effectiveness; 2. Number of days attended in the
past two weeks; and 3. Proportion of teachers who are “qualified”. 25 In fact, Vietnam is more likely to use student performance to evaluate teachers, but the effect of this variable’s
difference in means (second to last column of Table 9A) is much smaller (3.0) than its differential effect (25.0). IV. What Can Be Learned from Oaxaca-Blinder Decompositions? It is important to keep in 30 mind that these effects do not reflect that these variables have higher mean values in Vietnam;
the 25 point impact using student performance to assess teacher effectiveness in the last column
of Table 9A does not reflect that Vietnam is more likely to use student performance when
assessing teachers’ performance,25 but rather the results suggest that this policy is more effective
in Vietnam than in other countries. Similarly, the role played by student attendance is not so
much due to higher student attendance in Vietnam, but due to students learning more per day of
school attended. Finally, note that the third largest contributor is in fact negative: While, as one
would expect, having more “qualified” teachers increases student learning, the estimated impact
of such teachers in Vietnam (18.2) is much lower than the estimated impact for the other
countries that participated in the 2012 PISA assessment (46.1). Table 10A yields similar results for the reading decomposition using the 2012 PISA data,
although they are somewhat more difficult to interpret. The overall gap to be explained is 40.3
points. As with the math score, the differences in the x variables explain little, and in fact they
slightly widen the gap to be explained by 1 point. In contrast, the “greater efficiency” of the x
variables explains virtually all of the gap by accounting for 41.3 points in that gap. Two x
variables stand out as making the biggest contribution to explaining this gap. First, as with the
mathematical results, the differential efficiency of being in grade 10 can account for virtually all
of the gap (42.7 points). Again, this may reflect not only differential efficiency but also a
screening process between grades 9 and 10 that removes the weakest students from the overall
population of 15-year-old students. Second, the differential impact of school attendance makes
an even larger contribution, unlike the results for the math exam. The contribution of this
differential school attendance, 80.5 points, is so large that it requires some variable to have a 31 large negative effect, and that variable is the constant term, which is much larger in the other
countries than in Vietnam, a result also not seen with the math test. 26 The application of the Oaxaca-Blinder decomposition in this paper could be misleading in that the estimated β
terms for Vietnam are based on only within-country variation in the variables, while the estimates of β for all other
countries are based on both within-country and between-country variation. It is possible that the decomposition
results could change if the latter estimates were also based on only within-country variation, which can be done by
estimating βo using a country-fixed-effects specification. This was done, and the main results still hold, as seen in
Appendix Tables B.4 and B.5. In particular, the contribution of the difference in the x variables, as measured by
𝛃̅ʹ(𝐱̅vn-𝐱̅o), remains very small. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? This somewhat puzzling
result for the reading decomposition, and unfortunately it is difficult to interpret this large
difference in the constant terms between Vietnam and the other 62 countries in the 2012 PISA
assessment.26 These Oaxaca-Blinder decompositions were also done for the 2015 PISA data, and the
results are similar. The relevant tables for these results are Tables 9B and 10B. Note first that,
as seen in Table 8, there are seven fewer variables in the 2015 PISA with which to do this
decomposition, but that still leaves 15 variables. The decomposition of the difference in the mathematics scores for the 2015 PISA
assessment is shown in Table 9B. As in the 2012 PISA, the differences in the x variables do not
explain the 44.6 point gap in the mean test score between Vietnam and the 65 other countries that
participated in the 2015 PISA. Indeed, those differences add 16.7 points to the gap. Thus, all of
the differences are due to the differences in the β terms for Vietnam and for the other countries. The decomposition of the difference in the mathematics scores for the 2015 PISA
assessment is shown in Table 9B. As in the 2012 PISA, the differences in the x variables do not
explain the 44.6 point gap in the mean test score between Vietnam and the 65 other countries that
participated in the 2015 PISA. Indeed, those differences add 16.7 points to the gap. Thus, all of
the differences are due to the differences in the β terms for Vietnam and for the other countries. The decomposition of the difference in the reading scores is shown in Table 10B, and the results
are very similar to those in Table 9B; the differences in the x variables do not explain the 27.0
point gap in the mean test score between Vietnam and the 65 other countries, and instead they in
effect widen the gap. Again, the entire explanation for the gap must come from the differences
in the β terms between Vietnam and the other countries. The decomposition of the difference in the reading scores is shown in Table 10B, and the results
are very similar to those in Table 9B; the differences in the x variables do not explain the 27.0
point gap in the mean test score between Vietnam and the 65 other countries, and instead they in
effect widen the gap. IV. What Can Be Learned from Oaxaca-Blinder Decompositions? Again, the entire explanation for the gap must come from the differences
in the β terms between Vietnam and the other countries. 32 To summarize this section, the Oaxaca-Blinder decompositions for both 2012 and 2015
indicate that Vietnam’s exceptional performance on the PISA assessment in both years is not due
to Vietnam having “better” observable child, household or school characteristics. Instead
Vietnam seems to be more effective in transforming those factors into test scores. In other
words, these decompositions indicate that virtually all of Vietnam’s strong performance in these
two PISA assessments comes from the “unexplained” portion of this decomposition method, and
thus this decomposition sheds very little light on the underlying reasons for that performance. V. Conclusion Vietnam’s very high performance on the 2012 and 2015 PISA assessments has raised the
question of why Vietnam does so well, and whether other countries can improve their student
learning outcomes by applying what works well in Vietnam. This paper has used the 2012 and
2015 PISA data to do three types of analysis to explore the reasons behind Vietnam’s apparent
success. The analysis done thus far has led to three sets of general results. First, one important, albeit partial, explanation of Vietnam’s very strong performance on
the 2012 and 2015 PISA assessments is that the weakest students are excluded from grade 10. In
particular, only about 66% of Vietnamese 15-year-olds participated in the 2012 and 2015 PISA
assessments, presumably because most or all of the other 34% were no longer in school. (Note
that the OECD reports even lower participation rates, but this paper corrects those erroneous
rates.) Yet applying three different methods to adjust for Vietnam’s low coverage (enrollment)
rates has little effect on its outlier status. Moreover, back-of-the-envelope calculations to
account for possible higher motivation of Vietnamese students on the PISA account for at most
less than half of Vietnam’s outlier status. A related point is that the 15-year-old students who 33 participated in the PISA assessments appear to be better off when compared to the 15-year-olds
enrolled in school in the 2012, 2014 and 2016 Vietnam Household Living Standards Surveys. Yet even after adjusting Vietnam’s test scores for these differences between the two datasets,
those scores are still well above what one would predict based on Vietnam’s income alone. A
similar result also holds for the 2015 PISA scores. Thus, even after accounting for these
apparent differences between the PISA and VHLSS data, Vietnam still performs unusually well
on the PISA assessments relative to what one would expect given its income level. Second, taking the PISA data at face value, this paper has used regression methods to
investigate which family, teacher or school characteristics in the PISA data can “explain” the
high performance of Vietnamese students. The general finding of this analysis is that accounting
for household level and school level variables in the PISA data explains at most only one fourth
or one third of Vietnam’s high performance on the 2012 PISA relative to its income level. V. Conclusion Moreover, adding these variables to an initial regression of PISA test scores on household wealth
does very little to Vietnam’s “outlier” status. Similar results are found for the 2015 PISA. Third, this paper has applied the Oaxaca-Blinder decomposition method to better
understand the difference in average test scores between Vietnamese students and students in the
other countries that participated in the 2012 and 2015 PISA assessments. Unlike the analysis
discussed in the previous paragraph, this approach is more flexible in that it allows the impacts
(the β’s) of the household and school variables (the x variables) to differ between Vietnam and
all other countries that participated in the PISA assessments. The decompositions indicate that
all of the gap in average test scores between Vietnam and the other countries (62 in 2012 and 65
in 2015) is due to the greater “productivity” of various household and school variables in
Vietnam, relative to the “productivity” of those variables in other countries, especially the 34 greater “productivity” that occurs when grade 9 students in Vietnam are enrolled in grade 10. This could reflect the fact that only Vietnamese students who pass an entrance exam are allowed
to proceed from grade 9 to grade 10, which is the first grade of the upper secondary cycle in
Vietnam. This is most clear for the mathematics results; the reading results show a similar
pattern but also have other factors at work (in particular, more productivity from each day of
school attendance). In contrast, differences in household and school characteristics explain none
of the gap between Vietnam and the other countries that participated in the PISA assessment;
indeed, they add to the gap to be explained. While the analysis in this paper has shed some light on the reasons for Vietnam’s
exceptional performance on the 2012 and 2015 PISA assessments, in the end the main
contribution of this paper is that it has shown what does not explain that exceptional
performance. In particular, it does not appear to be due to the low participation of Vietnam’s 15-
year-olds, nor to any possible selection of “better” students (as measured by urban location,
parental education and household wealth) into the PISA assessments. V. Conclusion Observable child,
household, and school characteristics explain little or none of the differential performance; in
particular the Oaxaca-Blinder decompositions attribute none of the gap in test scores to such
factors, and instead attribute all to the “unexplained” differences between Vietnam and the PISA
participants in the coefficients that determine the impacts of these observed factors. Future
research on Vietnam’s exceptional performance will need to use different data, and perhaps
different methodologies, to understand that country’s impressive performance in education. 35 References Blinder, Alan. 1973. “Wage Discrimination: Reduced Form and Structural Estimates”. Journal
of Human Resources 8(4):436-455. Dang, Hai-Anh, and Paul Glewwe. 2018. “Well Begun, But Aiming Higher: A Review of
Vietnam’s Education Trends in the Past 20 Years and Emerging Challenges”. Journal of
Development Studies 54(7):1171-1195. Fortin, Nicole, Thomas Lemieux and Sergio Firpo. 2011. “Decomposition Methods in
Economics”, in O. Ashenfelter and D. Card, eds., Handbook of Labor Economics,
Volume 4A. North-Holland: Amsterdam. Fryer, Roland, and Steven Levitt. 2004. “Understanding the Black-White Test Score Gap in the
First Two Years of School”. Review of Economics and Statistics 86(2):447-464. General Statistics Office. 2010. The 2009 Vietnam Population and Housing Census: Completed
Results. Hanoi: Vietnam. Available online at:
https://www.gso.gov.vn/default_en.aspx?tabid=515&idmid=5&ItemID=10799 Gneezy, Uri, John A. List, Jeffrey A. Livingston, Xiangdong Qin, Sally Sadoff, and Yang
Xu. 2019. “Measuring Success in Education: The Role of Effort on the Test
Itself.” American Economic Review: Insights 1(3): 291-308. Jacob, Brian, and Steven Levitt. 2003. “Rotten Apples: An Investigation of the Prevalence and
Predictors of Teacher Cheating”. Quarterly Journal of Economics 118(3):843-877. Jann, Ben. 2008. “The Blinder-Oaxaca Decomposition for Linear Regression Models”. The
Stata Journal 8(4):453-479. Oaxaca, Ronald. 1973. “Male-Female Differentials in Urban Labor Markets”. International
Economic Review 3(4):693-709. OECD. 2014a. PISA 2012 Results: What Students Know and Can Do – Student Performance in
Mathematics, Reading and Science, Volume I (Revised Edition). Paris: Organization for
Economic Cooperation and Development. OECD. 2014b. PISA 2012 Technical Report. Paris: Organization for Economic Cooperation
and Development. OECD. 2016. PISA 2015 Results: Excellence and Equity in Education, Volume I. Paris:
Organization for Economic Cooperation and Development. Singh, Abhijeet. 2019. “Learning More with Every Year: School Year Productivity and
International Learning Divergence”. Journal of European Economic Association. https://doi.org/10.1093/jeea/jvz033. 36 Thanh Nien News. 2013. “Vietnam Deputy Education Minister not Convinced by Global Test”. December 7, 2013. http://www.thanhniennews.com/education-youth/vietnam-deputy-
education-minister-not-convinced-by-global-test-18276.html. Waldow, Florian and Gita Steiner-Khamsi. 2019. Understanding PISA’s Attractiveness: Critical
Analyses in Comparative Policy Studies. V. Conclusion Bloomsbury Academic, London. World Bank. 2013. “Skilling up Vietnam: Preparing the workforce for a modern market
economy”. Vietnam Development Report 2014. World Bank, Washington, DC. . World Development Indicators Online database. World Bank, Washington, DC. 37 37 Figure 1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita
Figure 2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita Figure 1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita gure 1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita Figure 2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita Figure 2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita Figure 2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP 38 Figure 3. Mean Age 15 Math Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 3. Mean Age 15 Math Scores in 2015 PISA, by 2015 Log Real GDP/capita
Figure 4. Mean Age 15 Language Scores in 2015 PISA, by 2015 Log Real GDP/capita Mean Age 15 Math Scores in 2015 PISA, by 2015 Log Real GDP/capita 015 PISA, by 2015 Log Real GDP/capita Figure 4. Mean Age 15 Language Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 4. Mean Age 15 Language Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 4. Mean Age 15 Language Scores in 2015 PISA, by 2015 Log Real GDP 39 Figure 5. Mean Age 15 Top 50% Math Scores in 2012 PISA, by 2010 Log Real GDP/capita
Figure 6. Mean Age 15 Top 50% Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita Figure 5. Mean Age 15 Top 50% Math Scores in 2012 PISA, by 2010 Log Real GDP/capita an Age 15 Top 50% Math Scores in 2012 PISA, by 2010 Log Real GDP/capita 40
Figure 6. Mean Age 15 Top 50% Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita Figure 6. Mean Age 15 Top 50% Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita an Age 15 Top 50% Reading Scores in 2012 PISA, by 2010 Log Real GDP/capi Figure 6. Mean Age 15 Top 50% Reading Scores in 2012 PISA, by 2010 Log R 40 41
Figure 7. V. Conclusion Mean Age 15 Top 50% Math Scores in 2015 PISA, by 2015 Log Real GDP/capita
Figure 8. Mean Age 15 Top 50% Reading Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 7. Mean Age 15 Top 50% Math Scores in 2015 PISA, by 2015 Log Real GDP/capita an Age 15 Top 50% Math Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 8. Mean Age 15 Top 50% Reading Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 8. Mean Age 15 Top 50% Reading Scores in 2015 PISA, by 2015 Log R igure 8. Mean Age 15 Top 50% Reading Scores in 2015 PISA, by 2015 Log Real GDP/capita Figure 8. Mean Age 15 Top 50% Reading Scores in 2015 PISA, by 2015 Log Real GDP/capita 41 42
Figure 9: Lower and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2012
Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. Figure 10: Lower and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2015
Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. Figure 9: Lower and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2012
Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. Figure 9: Lower and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2012 and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2012 Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. es are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. V. Conclusion Figure 10: Lower and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2015
Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. Figure 10: Lower and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 2015
Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. wer and Upper (observed) Bounds of Math Scores for All 15-year-olds, PISA 20 Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. Note: All countries are sorted in an increasing order of the upper bound (observed mean from PISA participants) of
test scores. The dots represent the observed mean test scores. Vietnam is indicated by the larger diamond. 42 Figure 11: Midpoint of Upper and Lower Bounds of 2012 PISA Math Scores, by Log of
GDP/capita
Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. Midpoint of Upper and Lower Bounds of 2012 PISA Math Scores, by Log of Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. Figure 12: Midpoint of Upper and Lower Bounds of 2012 PISA Reading Scores, by Log of
GDP/capita
Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. Figure 12: Midpoint of Upper and Lower Bounds of 2012 PISA Reading Scores, by Log of
GDP/
it Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. 43 Figure 13: Midpoint of Upper and Lower Bounds of 2015 PISA Math Scores, by Log of
GDP/capita
Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. V. Conclusion Figure 13: Midpoint of Upper and Lower Bounds of 2015 PISA Math Scores, by Log of Midpoint of Upper and Lower Bounds of 2015 PISA Math Scores, by Log of Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. Figure 14: Midpoint of Upper and Lower Bounds of 2015 PISA Reading Scores, by Log of
GDP/capita
N t
Adj
t d t
t
f
h
t
th
id
i t
l
f th
b
d t
t
d th th
ti
l Figure 14: Midpoint of Upper and Lower Bounds of 2015 PISA Reading Scores, by Log of
GDP/capita
Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. point of Upper and Lower Bounds of 2015 PISA Reading Scores, by Log of Figure 14: Midpoint of Upper and Lower Bounds of 2015 PISA Reading Figure 14: Midpoint of Upper and Lower Bounds of 2015 PISA Reading Scores, by Log of
GDP/capita Note: Adjusted test scores for each country are the mid-point value of the observed test scores and the theoretical
lower bounds based on Proposition 1. 44 Table 1: Student Characteristics in 2012 (born in 1996) and 2015 (born in 1999): PISA vs. V. Conclusion VHLSS
2012 PISA and 2012 VHLSS
2015 PISA and 2014 & 2016 VHLSS
PISA
VHLSS (PISA-eligible only)
PISA VHLSS (PISA-eligible only)
Variable
(1)
All
(2)
Mar.-July
(3)
Difference
(3) – (1)
(4)
All
(5)
Mar.-July
(6)
Difference
(6) – (4)
Urban
50.3%
26.0%
25.3%
-24.9***
49.6%
30.5%
28.6%
-21.0***
(4.2)
(2.3)
(3.2)
(5.2)
(4.0)
(1.9)
(2.7)
(4.9)
Female
53.8%
51.7%
51.7%
-2.1
51.4%
51.4%
47.1%
-4.3
(0.8)
(2.6)
(3.5)
(3.6)
(1.0)
(1.9)
(2.6)
(2.8)
Current grade: 10 or higher
86.1%
84.3%
75.7%
-10.4***
85.5%
90.5%
84.3%
-1.2
(2.6)
(1.8)
(3.0)
(3.9)
(3.0)
(1.0)
(1.8)
(3.5)
Current grade: 9 or lower
10.3%
14.0%
22.2%
11.9***
9.0%
8.5%
15.1%
6.2**
(2.2)
(1.7)
(2.8)
(3.6)
(2.2)
(1.0)
(1.8)
(2.8)
Current grade: unknown/othera/ 3.6%
1.7%
2.1%
-1.5
5.5%
1.0%
0.6%
-4.9**
(1.5)
(0.7)
(1.3)
(2.0)
(2.3)
(0.4)
(0.4)
(2.3)
Father’s years of schooling
8.95
7.18
7.19
-1.76***
8.4
7.1
6.9
-1.47***
(0.17)
(0.22)
(0.32)
(0.37)
(0.17)
(0.17)
(0.23)
(0.29)
Mother’s years of schooling
8.34
6.80
6.93
-1.41***
7.9
6.6
6.4
-1.41***
(0.19)
(0.19)
(0.26)
(0.32)
(0.20)
(0.15)
(0.22)
(0.29)
Owns an air-conditioner
16.0%
7.1%
7.1%
-8.8***
20.7%
19.2%
15.2%
-5.5**
(2.1)
(1.4)
(2.1)
(3.0)
(1.6)
(1.7)
(2.2)
(2.8)
Owns a motorbike
93.1%
91.0%
90.7%
-2.4
93.9%
94.0%
93.8%
-0.2
(0.5)
(1.4)
(2.0)
(2.1)
(0.5)
(0.8)
(1.3)
(1.4)
Owns a car
7.3%
0.7%
1.0%
-6.3***
7.9%
2.0%
2.6%
-5.3***
(0.8)
(0.3)
(0.7)
(1.1)
(0.7)
(0.5)
(0.9)
(1.1)
Owns a computer
39.1%
24.5%
25.1%
-14.1***
44.1%
29.5%
28.5%
-15.6***
(2.2)
(2.3)
(3.2)
(3.9)
(1.9)
(1.8)
(2.4)
(3.1)
Number of televisions owned
1.39
1.00
1.00
-0.38***
1.42
1.09
1.05
-0.36***
(0.03)
(0.02)
(0.03)
(0.04)
(0.03)
(0.02)
(0.03)
(0.04)
Sample size
4,771
455
236
5687
849
415
PISA coverage/eligibility rate
56%
75%
78%
49%
76.4%
77.8%
Robust standard errors, clustered at school level in the PISA sample and at commune level in the racteristics in 2012 (born in 1996) and 2015 (born in 1999): PISA vs. VHLSS Robust standard errors, clustered at school level in the PISA sample and at commune level in the
VHLSS sample, are shown in parentheses. The difference column reports mean differences between the PISA sample and the VHLSS
subsample interviewed from March to July, as well as their standard errors; t-tests are conducted
to test whether the mean difference of each variable is significantly different from zero, for
which: *** p<0.01, ** p<0.05, * p<0.1. V. Conclusion The difference column reports mean differences between the PISA sample and the VHLSS
subsample interviewed from March to July, as well as their standard errors; t-tests are conducted
to test whether the mean difference of each variable is significantly different from zero, for
which: *** p<0.01, ** p<0.05, * p<0.1. a/ In the PISA sample, this category consists of observations originally categorized as
“Ungraded”, with no further information; in the VHLSS sample, this category consists of
observations originally categorized as “Attending vocational schools”. a/ In the PISA sample, this category consists of observations originally categorized as
“Ungraded”, with no further information; in the VHLSS sample, this category consists of
observations originally categorized as “Attending vocational schools”. 45 Table 2: Predicted PISA Math Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means of VHLSS data)
A. 2012 PISA Data and 2012 VHLSS Data
Variable
Variable Means
Difference
in Means
Math
Coeff. Math Coefficient Multiplied by:
PISA
VHLSS
PISA
Mean
VHLSS
Mean
Difference
in Means
Rural
0.497
0.747
-0.250
-18.04
-9.0
-13.5
4.5
Female
0.538
0.517
0.021
-16.58
-8.9
-8.6
-0.4
Grade 10
0.861
0.757
0.104
105.8
91.0
80.1
11.0
Dad Yrs. Sch. 8.81
7.19
1.62
2.231
19.7
16.0
3.6
Mom yrs. sch. 8.23
6.93
1.306
1.879
15.5
13.0
2.4
Air condit. 0.160
0.071
0.089
5.456
0.9
0.4
0.5
Car
0.094
0.010
0.084
-6.723
-0.6
-0.1
-0.6
Computer
0.391
0.251
0.140
17.35
6.8
4.4
2.4
TVs
1.39
1.00
0.39
0.526
0.7
0.5
0.2
Constant
1.000
1.000
0.000
396.7
396.7
396.7
0.0
Column sum
--
--
--
--
512.7
489.0
23.7 Table 2: Predicted PISA Math Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means of VHLSS data) edicted PISA Math Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means of VHLSS data) B. 2015 PISA Data and 2014 and 2016 VHLSS Data B. 2015 PISA Data and 2014 and 2016 VHLSS Data
Variable
Variable Means
Math Coefficient Multiplied by:
Difference
Math
PISA
VHLSS
Difference
PISA
VHLSS
in Means
Coeff. Mean
Mean
in Means
Rural
0.504
0.714
-0.210
-9.822
-5.0
-7.0
2.1
Female
0.514
0.471
0.043
-8.461
-4.3
-4.0
-0.4
Grade 10
0.855
0.843
0.012
74.61
63.8
62.9
0.9
Dad yrs. sch. 8.40
6.446
1.410
2.041
17.1
9.4
2.1
Mom yrs. sch. 7.86
6.932
1.467
1.460
11.5
14.2
3.0
Air condit. V. Conclusion 0.207
0.152
0.055
-2.685
-0.6
-0.4
-0.15
Motorbike
0.939
0.938
0.002
6.451
6.1
6.0
0.01
Car
0.079
0.026
0.053
-1.249
-0.1
0.0
-0.1
Computer
0.441
0.285
0.156
23.40
10.3
6.7
3.7
TVs
1.416
1.054
0.363
6.734
9.5
7.1
2.4
Constant
1.000
1.000
0.000
386.4
386.4
386.4
0.0
Column sum
494.7
481.2
13.6 46 Table 3 Predicted Reading Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means for the VHLSS data)
A. 2012 PISA and 2012 VHLSS Data
Variable
Variable Means
Difference
in Means
Reading
Coeff. Reading Coefficient Multiplied by:
PISA
VHLSS
PISA
Mean
VHLSS
Mean
Difference
in Means
Rural
0.497
0.747
-0.250
-11.56
-5.7
-8.6
2.9
Female
0.538
0.517
0.021
24.61
13.2
12.7
0.5
Grade 10
0.861
0.757
0.104
95.14
81.9
72.0
9.9
Dad Yrs. Sch. 8.81
7.19
1.62
1.536
13.5
11.0
2.5
Mom yrs. sch. 8.23
6.93
1.30
1.661
13.7
11.5
2.2
Air condit. 0.160
0.071
0.089
-0.626
-0.1
-0.0
-0.1
Car
0.094
0.010
0.084
-3.442
-0.3
-0.0
-0.3
Computer
0.391
0.251
0.140
10.86
4.2
2.7
1.5
TVs
1.39
1.00
0.39
2.977
4.1
3.0
1.1
Constant
1.000
1.000
0.000
385.2
385.2
385.2
0.0
Column sum
--
--
--
--
509.8
489.5
20.3
B. 2015 PISA Data and 2014 and 2016 VHLSS Data
Variable
Variable Means
Reading Coefficient Multiplied by:
Difference Reading
PISA
VHLSS
Difference
PISA
VHLSS
in Means
Coeff. Mean
Mean
in Means
Rural
0.504
0.714
-0.210
-18.86
-9.5
-13.5
4.0
Female
0.514
0.471
0.043
15.97
8.2
7.5
0.7
Grade 10
0.855
0.843
0.012
69.85
59.7
58.9
0.9
Dad yrs. sch. 8.40
6.446
1.410
1.646
13.8
5.8
1.3
Mom yrs. sch. 7.86
6.932
1.467
0.893
7.0
11.4
2.4
Air condit. 0.207
0.152
0.055
-0.712
-0.1
-0.1
-0.04
Motorbike
0.939
0.938
0.002
15.83
14.9
14.8
0.03
Car
0.079
0.026
0.053
5.202
0.4
0.1
0.3
Computer
0.441
0.285
0.156
16.61
7.3
4.7
2.6
TVs
1.416
1.054
0.363
7.284
10.3
7.7
2.6
Constant
1.000
1.000
0.000
376.9
376.9
376.9
0.0
Column sum
489.0
474.3
14.7 Table 3 Predicted Reading Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means for the VHLSS data)
A. 2012 PISA and 2012 VHLSS Data
Variable
Variable Means
Difference
in Means
Reading
Coeff. V. Conclusion Reading Coefficient Multiplied by:
PISA
VHLSS
PISA
Mean
VHLSS
Mean
Difference
in Means
Rural
0.497
0.747
-0.250
-11.56
-5.7
-8.6
2.9
Female
0.538
0.517
0.021
24.61
13.2
12.7
0.5
Grade 10
0.861
0.757
0.104
95.14
81.9
72.0
9.9
Dad Yrs. Sch. 8.81
7.19
1.62
1.536
13.5
11.0
2.5
Mom yrs. sch. 8.23
6.93
1.30
1.661
13.7
11.5
2.2
Air condit. 0.160
0.071
0.089
-0.626
-0.1
-0.0
-0.1
Car
0.094
0.010
0.084
-3.442
-0.3
-0.0
-0.3
Computer
0.391
0.251
0.140
10.86
4.2
2.7
1.5
TVs
1.39
1.00
0.39
2.977
4.1
3.0
1.1
Constant
1.000
1.000
0.000
385.2
385.2
385.2
0.0
Column sum
--
--
--
--
509.8
489.5
20.3 Table 3 Predicted Reading Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means for the VHLSS data) Table 3 Predicted Reading Scores Based on VHLSS Data, Decomposed by Variable
(Using March – July Means for the VHLSS data) B. 2015 PISA Data and 2014 and 2016 VHLSS Data
Variable
Variable Means
Reading Coefficient Multiplied by:
Difference Reading
PISA
VHLSS
Difference
PISA
VHLSS
in Means
Coeff. Mean
Mean
in Means
Rural
0.504
0.714
-0.210
-18.86
-9.5
-13.5
4.0
Female
0.514
0.471
0.043
15.97
8.2
7.5
0.7
Grade 10
0.855
0.843
0.012
69.85
59.7
58.9
0.9
Dad yrs. sch. 8.40
6.446
1.410
1.646
13.8
5.8
1.3
Mom yrs. sch. 7.86
6.932
1.467
0.893
7.0
11.4
2.4
Air condit. 0.207
0.152
0.055
-0.712
-0.1
-0.1
-0.04
Motorbike
0.939
0.938
0.002
15.83
14.9
14.8
0.03
Car
0.079
0.026
0.053
5.202
0.4
0.1
0.3
Computer
0.441
0.285
0.156
16.61
7.3
4.7
2.6
TVs
1.416
1.054
0.363
7.284
10.3
7.7
2.6
Constant
1.000
1.000
0.000
376.9
376.9
376.9
0.0
Column sum
489.0
474.3
14.7 B. 2015 PISA Data and 2014 and 2016 VHLSS Data 47 PISA Assessment Country 2012 Rankings, Overall and Top 50% of Overall 48
Table 4A: PISA Assessment Country 2012 Rankings, Overall and Top 50% of Overall
Population
Math (all students)
Reading (all students)
Math (top 50% of pop.)
Reading (top 50% of pop.)
Rank
Country
Avg. score
Country
Avg. score
Country
Avg. score
Country
Avg. V. Conclusion score
1
Singapore
573
Hong Kong
545
Singapore
648
Singapore
612
2
Hong Kong
561
Singapore
542
Taiwan
639
Japan
607
3
Taiwan
559
Japan
538
Hong Kong
623
Hong Kong
599
4
South Korea
554
South Korea
536
South Korea
622
South Korea
594
5
Macao
538
Finland
524
Japan
602
Belgium
587
6
Japan
536
Canada
523
Belgium
596
Finland
585
7
Liechtenstein
535
Taiwan
523
Macao
595
Taiwan
585
8
Switzerland
531
Ireland
523
Netherlands
592
New Zealand
585
9
Netherlands
523
Poland
518
Liechtenstein
589
France
584
10
Estonia
521
Liechtenstein
516
Switzerland
586
Ireland
583
11
Finland
519
Estonia
516
Germany
586
Netherlands
580
12
Poland
518
New Zealand
512
Poland
583
Poland
580
13
Canada
518
Australia
512
Czech Republic
583
Germany
575
14
Belgium
515
Netherlands
511
Estonia
578
Estonia
573
15
Germany
514
Macao
509
Finland
573
Norway
572
16
Vietnam
511
Belgium
509
Austria
569
Czech Republic
571
17
Austria
506
Switzerland
509
New Zealand
568
Canada
569
18
Australia
504
Vietnam
508
France
566
Israel
567
19
Ireland
501
Germany
508
Canada
563
Australia
567
20
Slovenia
501
France
505
Ireland
560
United Kingdom
565
21
Denmark
500
Norway
504
Iceland
559
Liechtenstein
565
22
New Zealand
500
United Kingdom
499
Slovakia
558
Sweden
561
23
Czech Republic
499
United States
498
Australia
557
Switzerland
560
24
France
495
Denmark
496
United Kingdom
556
United States
560
25
United Kingdom
494
Czech Republic
493
Luxembourg
556
Luxembourg
560
26
Iceland
493
Austria
490
Spain
555
Macao
558
27
Latvia
491
Italy
490
Norway
553
Italy
557
28
Luxembourg
490
Latvia
489
Italy
551
Spain
555
29
Norway
489
Spain
488
Slovenia
549
Austria
553
30
Portugal
487
Luxembourg
488
Portugal
548
Iceland
553
31
Italy
485
Portugal
488
Denmark
547
Hungary
551
32
Spain
484
Hungary
488
Latvia
547
Latvia
549
33
Russian Federation
482
Israel
486
Sweden
544
Portugal
548
34
Slovakia
482
Croatia
485
Vietnam
543
Denmark
546
35
United States
481
Iceland
483
Russian Federation
543
Croatia
546
36
Lithuania
479
Sweden
483
United States
542
Slovakia
542
37
Sweden
478
Slovenia
481
Israel
541
Greece
541
38
Hungary
477
Greece
477
Hungary
540
Russian Federation
538
39
Croatia
471
Lithuania
477
Lithuania
536
Vietnam
537
40
Israel
466
Turkey
475
Croatia
533
Lithuania
534
41
Greece
453
Russian Federation
475
Greece
510
Slovenia
530
42
Serbia
449
Slovakia
463
Romania
504
Turkey
512
43
Turkey
448
Serbia
446
Serbia
503
Chile
511
44
Romania
445
United Arab Emirates
442
Bulgaria
492
Bulgaria
509
45
Bulgaria
439
Chile
441
Chile
499
Romania
505
46
United Arab Emirates
434
Costa Rica
441
United Arab Emirates 486
Serbia
503
47
Kazakhstan
432
Thailand
441
Turkey
486
United Arab Emirates 499
48
Thailand
427
Romania
438
Thailand
482
Thailand
492
49
Chile
423
Bulgaria
436
Kazakhstan
471
Montenegro
482
50
Malaysia
421
Mexico
424
Malaysia
468
Qatar
470
51
Mexico
413
Montenegro
422
Montenegro
460
Tunisia
464
52
Montenegro
410
Uruguay
411
Uruguay
453
Argentina
462
53
Uruguay
409
Brazil
410
Qatar
447
Uruguay
461
54
Costa Rica
407
Tunisia
404
Mexico
443
Mexico
456
55
Albania
394
Colombia
403
Argentina
440
Jordan
455
56
Brazil
391
Jordan
399
Tunisia
438
Malaysia
449
57
Argentina
388
Malaysia
398
Jordan
430
Colombia
443
58
Tunisia
388
Indonesia
396
Brazil
414
Costa Rica
441
59
Jordan
386
Argentina
396
Albania
412
Brazil
437
60
Qatar
376
Albania
394
Colombia
410
Kazakhstan
437
61
Colombia
376
Kazakhstan
393
Peru
406
Peru
429
62
Indonesia
375
Qatar
388
Costa Rica
406
Indonesia
423
63
Peru
368
Peru
384
Indonesia
399
Albania
419 Table 4A: PISA Assessment Country 2012 Rankings, Overall and Top 50
P
l ti 48 49
Table 4B: PISA Assessment 2015 Country Rankings, Overall and Top 50% of Overall Population
Math (all students)
Reading (all students)
Math (Top 50% of pop.)
Reading (Top 50% of pop.)
Rank
Country
Avg score Country
Avg score
Country
Avg score Country
Avg score
1 Singapore
564 Singapore
533 Singapore
631 Singapore
603
2 Hong Kong
547 Canada
527 Hong Kong
609 Finland
595
3 Macao
543 Hong Kong
527 Japan
596 Germany
585
4 Japan
533 Finland
527 Macao
595 Hong Kong
584
5 South Korea
524 Ireland
520 South Korea
593 Ireland
583
6 Switzerland
520 Estonia
519 Switzerland
588 South Korea
583
7 Estonia
519 Japan
515 Netherlands
583 Norway
582
8 Canada
516 South Korea
515 Belgium
580 Japan
582
9 Netherlands
513 Norway
514 Estonia
576 New Zealand
582
10 Denmark
512 Germany
509 Germany
572 Estonia
582
11 Finland
511 New Zealand
509 Finland
572 France
580
12 Slovenia
510 Macao
508 Poland
565 Netherlands
580
13 Belgium
507 Poland
506 France
565 Canada
577
14 Germany
505 Slovenia
505 Czech Rep. V. Conclusion 564 Belgium
575
15 Poland
505 Netherlands
503 Malta
564 Sweden
573
16 Ireland
503 Australia
503 Canada
562 Poland
569
17 Norway
500 Denmark
500 Slovenia
562 Czech Rep. 567
18 Austria
496 Sweden
500 Ireland
562 Australia
567
19 New Zealand
494 Belgium
499 Norway
557 Israel
565
20 Sweden
494 France
498 Sweden
557 Switzerland
563
21 Australia
494 Portugal
498 New Zealand
557 Macao
562
22 Russia
494 United Kingdom
497 Denmark
557 United States
560
23 France
494 United States
496 Austria
557 Slovenia
560
24 Vietnam
493 Russia
495 Russia
553 Spain
558
25 Czech Rep. 492 Spain
495 Iceland
553 Russia
557
26 Portugal
492 Switzerland
492 Australia
551 United Kingdom
556
27 United Kingdom
492 Latvia
489 Portugal
550 Luxembourg
553
28 Italy
489 Vietnam
488 Luxembourg
550 Portugal
553
29 Iceland
487 Czech Rep. 487 United Kingdom
549 Iceland
552
30 Luxembourg
487 Croatia
487 Italy
548 Denmark
551
31 Spain
486 Italy
487 Spain
547 Croatia
551
32 Latvia
482 Austria
486 Hungary
546 Austria
550
33 Lithuania
479 Iceland
481 Israel
545 Italy
547
34 Malta
477 Luxembourg
480 Slovakia
542 Latvia
546
35 Hungary
477 Israel
479 Lithuania
535 Hungary
543
36 Slovakia
475 Lithuania
472 Latvia
534 Greece
542
37 United States
470 Hungary
469 Vietnam
528 Malta
541
38 Israel
468 Greece
466 United States
526 Lithuania
535
39 Croatia
463 Chile
458 Croatia
525 Slovakia
524
40 Greece
455 Slovakia
453 Greece
524 Chile
521
41 Romania
443 Malta
448 Romania
504 Vietnam
518
42 Bulgaria
442 Uruguay
438 Bulgaria
501 United Arab Em. 507
43 United Arab En. 427 Romina
433 United Arab Em. 494 Bulgaria
504
44 Chile
423 United Arab Em. 432 Chile
483 Romania
501
45 Turkey
421 Bulgaria
431 Moldova
482 Montenegro
489
46 Uruguay
420 Turkey
429 Qatar
473 Qatar
486
47 Moldova
419 Trinidad & Tob. 428 Montenegro
472 Moldova
485
48 Trinidad & Tob. 419 Costa Rica
427 Trinidad & Tob. 469 Trinidad & Tob. V. Conclusion 485
49 Montenegro
416 Montenegro
426 Albania
464 Uruguay
485
50 Thailand
415 Colombia
425 Uruguay
460 Colombia
478
51 Albania
412 Mexico
423 Georgia
458 Jordan
470
52 Mexico
408 Moldava
417 Turkey
456 Turkey
465
53 Georgia
405 Thailand
408 Thailand
455 Albania
464
54 Qatar
402 Brazil
408 Lebanon
439 Georgia
462
55 Costa Rica
400 Jordan
408 Macedonia
436 Costa Rica
454
56 Lebanon
398 Albania
405 Jordan
436 Mexico
451
57 Colombia
390 Qatar
403 Mexico
434 Thailand
447
58 Indonesia
387 Georgia
402 Colombia
434 Peru
444
59 Peru
386 Indonesia
398 Peru
427 Brazil
443
60 Jordan
381 Peru
397 Tunisia
423 Indonesia
433
61 Brazil
377 Tunisia
359 Costa Rica
422 Macedonia
420
62 Macedonia
372 Dominican Rep. 358 Indonesia
421 Tunisia
416
63 Tunisia
365 Macedonia
352 Brazil
407 Dominican Rep. 397
64 Kosovo
362 Algeria
348 Algeria
395 Lebanon
394
65 Algeria
360 Kosovo
347 Kosovo
393 Algeria
385
66 Dominican Rep. 329 Lebanon
347 Dominican Rep. 360 Kosovo
381 49 Table 5. Regressions of PISA Test Scores on Log(GDP)/capita or Wealth/capita: Student-Level Data
(1)
(2)
(3)
(4)
(5)
(6)
(7)
(8)
Variables
Math
Reading
Math
Reading
Math
Reading
Math
Reading
A. 2012 PISA Assessment
Log of per capita GDP
34.14*** 31.53***
(1.57)
(1.44)
Wealth (national average)
28.84*** 26.63***
(1.10)
(1.04)
Wealth (student specific)
22.35*** 20.82*** 16.26*** 15.16***
(0.51)
(0.50)
(0.52)
(0.46)
Constant
126.08*** 159.47*** 454.86*** 463.16*** 458.27*** 466.98***
--
--
(15.47)
(14.13)
(1.18)
(1.11)
(1.06)
(0.99)
Vietnam residual (average)
135.8
119.0
111.6
96.7
98.2
83.6
82.8
73.4
Residual rank
1
1
2
1
4
2
5
3
More highly ranked
none
none
HK
none
HK
S. Korea
Singap. HK
HK
S. Korea
Singap. Taiwan
HK
S. Korea
Observations
473,236
473,236
473,236
473,236
455,971
455,971
455,971
455,971
R-squared
0.117
0.103
0.126
0.111
0.155
0.140
0.350
0.280
B. 2015 PISA Assessment
Log of per capita GDP
34.41*** 34.95***
(1.242)
(1.170)
Wealth (national average)
30.53*** 30.58***
(1.004)
(0.951)
Wealth (student specific)
22.89*** 23.03*** 16.24*** 16.22***
(0.445)
(0.440)
(0.476)
(0.443)
Constant
118.1*** 123.4*** 445.9*** 456.3*** 450.5*** 461.1***
(12.06)
(11.29)
(1.096)
(1.042)
(0.957)
(0.901)
Vietnam residual
(average)
114.6
99
106.2
89.7
89.7
72.8
73.2
63.4
Residual rank
1
1
1
1
3
1
4
3
More highly ranked
None
none
none
none
HK
Singap. none
Singap. HK
Macao
HK
Singap. V. Conclusion Observations
464,518
464,518
460,701
460,701
428,716
428,716
428,716
428,716
R-squared
0.128
0.127
0.140
0.136
0.170
0.167
0.331
0.262
Country fixed effects
No
No
No
No
No
No
Yes
Yes
Robust standard errors, clustered at the school level, in parentheses *** p<0.01, ** p<0.05, * p<0.1
For fixed effects regressions, residual = fixed effect – constant in regression without fixed effects. Table 6A: Regressions of 2012 Test Scores on Wealth/capita and Student and Household Variables
Variables
Math
Reading
Math
Reading
Math
Reading
Math
Reading
Wealth index
15.92*** 14.66*** 9.998*** 9.548*** 15.77*** 14.49*** 5.694***
5.080***
(0.52)
(0.48)
(0.433)
(0.406)
(0.53)
(0.48)
(0.384)
(0.355)
Girl
-8.705***
33.31***
-15.39*** 26.55***
(0.767)
(0.757)
(0.697)
(0.677)
Sibling index
-1.905*** -2.457***
-1.930*** -2.392***
(0.524)
(0.542)
(0.506)
(0.514)
Sibling index missing
-19.59*** -15.66***
-17.54*** -13.51***
(0.798)
(0.843)
(0.763)
(0.799)
Mom years school
2.978*** 2.872***
1.800*** 1.702***
(0.142)
(0.143)
(0.131)
(0.130)
Dad years school
3.310*** 3.065***
2.046*** 1.841***
(0.131)
(0.133)
(0.120)
(0.121)
Grade10
22.87*** 23.87***
(1.36)
(1.379
Years of preschool
10.74*** 10.05***
(0.680)
(0.693)
Educational input index
7.432*** 7.985***
(0.286)
(0.306)
Attendance (past 2 weeks)
7.710*** 7.638***
(0.365)
(0.364)
Books at home
0.069*** 0.0595***
(0.003)
(0.003)
Hours of study
3.170*** 3.017***
(0.094)
(0.089)
Extra math classes (tutored)
-0.558*
(0.212)
Extra math variable missing
-2.929***
(0.544)
Extra read. classes (tutored)
-4.440***
(0.2331)
Extra read. variable missing
-3.052***
(0.577)
Vietnam fixed effect
78.2
68.3
80.6
70.7
79.1
68.9
65.0
55.1
Fixed effect rank
5
3
6
2
5
3
5
3
More highly ranked:
HK
S. Korea
Singap. Taiwan
HK
S. Korea
HK
Macao
Singap. S. Korea
Taiwan
HK
HK
S. Korea
Singap. Taiwan
HK
S. Korea
HK
Macao
Singap. Taiwan
Finland
HK
Observations
401,489
401,489
401,489
401,489
393,730
393,730
393,730
393,730
R-squared
0.366
0.295
0.399
0.350
0.360
0.291
0.464
0.421
Robust standard errors, clustered at the school level, in parentheses. All regressions use country fixed effects. *** p<0.01, ** p<0.05, * p<0.1 Table 6A: Regressions of 2012 Test Scores on Wealth/capita and Student and of 2012 Test Scores on Wealth/capita and Student and Household Variables Variables
Wealth index
1
Girl
Sibling index
Sibling index missing
Mom years school
Dad years school
Grade10
Years of preschool
Educational input index
Attendance (past 2 weeks)
Books at home
Hours of study
Extra math classes (tutored)
Extra math variable missing
Extra read. classes (tutored)
Extra read. V. Conclusion variable missing
Vietnam fixed effect
Fixed effect rank
More highly ranked:
S
S
T
Observations
4
R-squared
Robust standard errors, cl 51 Table 6B: Regressions of 2015 Test Scores on Wealth/capita and Student and Household Variables Table 6B: Regressions of 2015 Test Scores on Wealth/capita and Student and Household Variables
Variables
Math
Reading
Math
Reading
Math
Reading
Math
Reading
Wealth Index
16.13*** 16.03*** 10.26***
10.76*** 16.02*** 15.89*** 7.042***
7.222***
(0.482)
(0.452)
(0.397)
(0.391)
(0.480)
(0.453)
(0.373)
(0.378)
Girl
-6.680*** 23.62***
-9.667*** 20.45***
(0.671)
(0.691)
(0.636)
(0.669)
Mom years school
2.863***
2.658***
2.070***
1.841***
(0.122)
(0.131)
(0.116)
(0.126)
Dad years school
3.151***
2.990***
2.327***
2.129***
(0.121)
(0.127)
(0.114)
(0.122)
Grade 10
24.58***
25.17***
(1.429)
(1.447)
Educational input
index
7.580***
(0.283)
8.756***
(0.302)
Books at home
0.063***
0.059***
(0.002)
(0.003)
Vietnam fixed effect
73.0
62.4
82.6
71.8
72.2
61.4
68.7
57.0
Fixed effect rank
5
3
4
2
5
3
4
3
More highly ranked: Singap. HK
Macao
Japan
HK
Singap. Singap. HK
Macao
HK
Singap. HK
Macao
Japan
HK
Singap. Macao
HK
Singap. HK
Finland
Observations
389,472
389,472
389,472
389,472
387,092
387,092
387,092
387,092
R-squared
0.341
0.276
0.372
0.314
0.338
0.273
0.404
0.348
Robust standard errors, clustered at the school level, in parentheses. All regressions use country fixed effects. *** p<0.01, ** p<0.05, * p<0.1 ons of 2015 Test Scores on Wealth/capita and Student and Household Variable Table 6B: Regressions of 2015 Test Scores on Wealth/capita and Student a Variables
Math
Reading
Math
Reading
Math
Reading
Math
Reading
Wealth Index
16.13*** 16.03*** 10.26***
10.76*** 16.02*** 15.89*** 7.042***
7.222***
(0.482)
(0.452)
(0.397)
(0.391)
(0.480)
(0.453)
(0.373)
(0.378)
Girl
-6.680*** 23.62***
-9.667*** 20.45***
(0.671)
(0.691)
(0.636)
(0.669)
Mom years school
2.863***
2.658***
2.070***
1.841***
(0.122)
(0.131)
(0.116)
(0.126)
Dad years school
3.151***
2.990***
2.327***
2.129***
(0.121)
(0.127)
(0.114)
(0.122)
Grade 10
24.58***
25.17***
(1.429)
(1.447)
Educational input
index
7.580***
(0.283)
8.756***
(0.302)
Books at home
0.063***
0.059***
(0.002)
(0.003)
Vietnam fixed effect
73.0
62.4
82.6
71.8
72.2
61.4
68.7
57.0
Fixed effect rank
5
3
4
2
5
3
4
3
More highly ranked: Singap. HK
Macao
Japan
HK
Singap. Singap. HK
Macao
HK
Singap. HK
Macao
Japan
HK
Singap. Macao
HK
Singap. HK
Finland
Observations
389,472
389,472
389,472
389,472
387,092
387,092
387,092
387,092
R-squared
0.341
0.276
0.372
0.314
0.338
0.273
0.404
0.348
Robust standard errors, clustered at the school level, in parentheses. All regressions use country fixed effects. p
p
p
and household variables not shown. All regressions use country fixed effects. V. Conclusion *** p<0.01, ** p<0.05, * p<0.1 52 52 Table 7A: Regressions 2012 Test Scores on Wealth/capita and Student, Household and
School Variables
Variables
Math
Reading
Math
Reading
Wealth
15.32***
13.75***
5.436***
3.869***
(0.57)
(0.51)
(0.414)
(0.387)
Class size (student/teacher ratio)
0.094
0.271***
(0.087)
(0.082)
Ratio qualified teachers
13.28***
10.44***
(3.33)
(3.30)
Qual. tchr. ratio missing
-1.370
-2.833
(3.320)
(3.038)
Square root of computers/pupil
-2.087
-0.710
(3.211)
(2.989)
Stud. perf. used to assess tchrs
1.728
2.049
(1.841)
(1.803)
Teacher absenteeism
-3.302***
-2.961***
(0.959)
(0.969)
Parents pressure teachers
11.59***
11.33***
(1.25)
(1.22)
Principal observes teachers
-2.741
0.117
(1.988)
(1.889)
Inspector observes teachers
-4.735***
-6.698***
(1.746)
(1.790)
Tchr pay linked to stud perf
-2.232**
-2.501***
(0.947)
(0.911)
Teacher mentoring index
5.244***
5.906***
(1.745)
(1.784)
Vietnam fixed effect
76.7
66.1
58.1
44.7
Fixed effect rank
5
4
8
4
More highly ranked:
HK
S. Korea
Singap. Taiwan
HK
S. Korea
Singap. HK
Liecht. Macao
S. Korea
Singap. Switz. Taiwan
Finland
HK
Liecht. Observations
341,409
341,409
341,409
341,409
R-squared
0.354
0.286
0.460
0.405
Robust standard errors, clustered at the school level, in parentheses. *** p<0.01, ** p<0.05, * p<0.1
Note: Student and household variables not shown. All regressions use country fixed effects. gressions 2012 Test Scores on Wealth/capita and Student, Household and
bl Note: Student and household variables not shown. All regressions use country fixed effects. 53 Table 7B: Regressions of 2015 Test Scores on Wealth/capita and Student, Household and
School Variables
Variables
Math
Reading
Math
Reading
Wealth Index
15.37***
15.12***
6.174***
6.302***
(0.532)
(0.488)
(0.401)
(0.402)
Class size (student/teacher ratio)
0.574***
0.704***
(0.104)
(0.108)
Ratio qualified teachers
9.323***
7.501***
(3.031)
(2.905)
Qual. tchr. ratio missing
-4.036
-4.125
(3.894)
(4.371)
Square root of computers/pupil
-0.435
0.759
(2.884)
(2.856)
Stud. perf. used to assess tchrs
3.996
1.316
(2.538)
(2.479)
Teacher absenteeism
-4.579***
-3.815***
(1.122)
(1.093)
Principal observes teachers
-1.984
0.675
(2.471)
(2.615)
Inspector observes teachers
-0.966
-2.204
(1.764)
(1.885)
Teacher mentoring index
0.449
0.276
(1.161)
(1.162)
Vietnam fixed effect
71.4
60.6
59.2
46.5
Fixed effect rank
4
3
4
8
More highly ranked:
Singap. HK
Macao
HK
Singap. Macao
HK
Singap. Switz. Finland
HK
Germany
Ireland
N. Zealand
Estonia
Singap. Observations
317,006
317,006
317,006
317,006
R-squared
0.320
0.262
0.391
0.341
Robust standard errors, clustered at the school level, in parentheses. *** p<0.01, ** p<0.05, * p<0.1
Note: Student and household variables not shown. All regressions use country fixed effects. V. Conclusion gressions of 2015 Test Scores on Wealth/capita and Student, Household and
bl Regressions of 2015 Test Scores on Wealth/capita and Student, Household and
riables Note: Student and household variables not shown. All regressions use country fixed effects. 54 gression Variables, for Vietnam and for Other Countries, 2012 and 2015 Table 8: Means of Regression Variables, for Vietnam and for Other Countries, 2012 and 2015
2012 PISA Assessment
2015 PISA Assessment
Variable (x)
Vietnam
Other PISA
Countries
Vietnam
Other PISA
Countries
Math test score
516.5
462.8
501.2
456.6
Reading test score
512.8
472.5
495.1
468.1
Wealth
4.143
6.101
3.21
5.63
Grade 10
0.874
0.584
0.89
0.59
Sibling index
1.048
1.086
--
--
Sibling index missing
0.149
0.238
--
--
Mom years schooling
8.313
10.98
8.03
11.45
Dad years schooling
8.883
11.09
8.53
11.54
Years preschool enrollment
1.600
1.487
--
--
Education inputs index (desk, books)
4.680
5.154
4.87
5.48
Books in home
57.59
114.1
69.17
113.15
Days attended in past 2 weeks
9.849
9.622
--
--
Hours of study per week
5.756
5.362
--
--
Extra reading classes (tutoring), hours/week
1.290
0.944
--
--
Extra reading classes variable missing
0.337
0.358
--
--
Extra math classes (tutoring), hours/week
2.741
1.325
--
--
Extra math classes variable missing
0.336
0.358
--
--
Class size
44.81
32.61
40.61
31.08
Proportion of teachers who are qualified
0.800
0.834
0.85
0.80
Proportion qualified teacher missing
0.069
0.188
0.07
0.08
Square root of computers/pupil
0.417
0.623
0.44
0.65
Student performance used to assess teachers
0.992
0.708
0.99
0.88
Teacher absenteeism
0.692
0.778
1.60
1.83
Parents pressure teachers
1.311
0.957
--
--
Principal observes teachers
0.965
0.802
0.99
0.87
Outside Inspector observes teachers
0.847
0.406
0.77
0.57
Teacher pay linked to student performance
1.487
0.703
--
--
Teachers are mentored
0.845
0.684
1.81
1.46
Sample size
4,421
336,988
4,895
312,111
Notes: 1. Averages over countries are weighted by country populations. 2. The following variables were not collected for all countries, or not for Vietnam, in
2015, and so are excluded from the analysis for that year: siblings, years in pre-school,
days attended, hours of study per week, extra classes, parents pressure teachers, and
teacher pay is linked to student performance. 2. The following variables were not collected for all countries, or not for Vietnam, in
2015, and so are excluded from the analysis for that year: siblings, years in pre-school,
days attended, hours of study per week, extra classes, parents pressure teachers, and
teacher pay is linked to student performance. Sample size 2. The following variables were not collected for all countries, or not for Vietnam, in
2015, and so are excluded from the analysis for that year: siblings, years in pre-school,
days attended, hours of study per week, extra classes, parents pressure teachers, and
teacher pay is linked to student performance. 55 Table 9A: Math Decomposition, 2012 (diff = 516.54– 462.80 = 53.74) Table 9A: Math Decomposition, 2012 (diff = 516.54– 462.80 = 53.74)
Variable
βvn
𝐱̅vn
βvnʹ𝐱̅vn
βo
𝐱̅o
βoʹ𝐱̅o 𝛃̅ (= (βvn+βo)/2 𝛃̅ʹ(𝐱̅vn-𝐱̅o) (βvn - 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo)
Wealth
6.764***
4.143
28.02
9.633***
6.101
58.77
8.198
-16.05
-14.69
Grade 10
85.85***
0.874
75.01
18.93***
0.584
11.05
52.39
15.19
48.76
Sibling index
3.152*
1.048
3.30
-1.697***
1.086
-1.84
0.728
-0.03
5.17
Sibling index missing
-0.576
0.149
-0.09
-17.87***
0.238
-4.25
-9.225
0.82
3.35
Mom years schooling
0.962**
8.313
8.00
1.786***
10.975
19.60
1.374
-3.66
-7.95
Dad years schooling
1.511***
8.883
13.42
2.390***
11.086
26.50
1.950
-4.30
-8.78
Years in preschool
6.533***
1.600
10.45
13.07***
1.487
19.43
9.799
1.10
-10.08
Education inputs index
4.397***
4.680
20.58
7.337***
5.154
37.81
5.867
-2.78
-14.46
Books in home
0.0089
57.59
0.51
0.0882***
114.07
10.07
0.049
-2.74
-6.81
Days attend past 2 wks
10.43***
9.849
102.72
8.094***
9.622
77.88
9.261
2.10
23.74
Hours study per week
2.920***
5.756
16.81
2.425***
5.362
13.00
2.672
1.05
2.75
Extra math class, hrs/wk
3.904***
2.741
10.70
-0.633**
1.325
-0.84
1.636
2.32
9.22
Extra math class missing
8.890***
0.336
2.98
-3.188***
0.358
-1.14
2.851
-0.06
4.19
Class size
0.0643
44.81
2.88
0.148*
32.61
4.82
0.106
1.29
-3.24
Proport. qualified tchrs
18.18**
0.800
14.55
46.08***
0.834
38.42
32.13
-1.09
-22.79
Prop. qual. tchr. missing
-17.15
0.069
-1.18
-23.14***
0.188
-4.35
-20.14
2.40
0.77
Square root comp/pupil
-0.0392
0.417
-0.02
4.925
0.623
3.07
2.443
-0.50
-2.58
Stud perf. to assess tchrs
25.08***
0.992
24.89
-4.267**
0.708
-3.02
10.40
2.96
24.95
Teacher absenteeism
-0.759
0.692
-0.53
-6.600***
0.778
-5.13
-3.679
0.32
4.29
Parents pressure tchrs
15.71***
1.311
20.60
6.686***
0.957
6.40
11.20
3.97
10.24
Principal observes tchrs
14.12
0.965
13.63
-3.816**
0.802
-3.06
5.154
0.84
15.85
Inspector observes tchrs
-16.73
0.847
-14.17
-10.15***
0.406
-4.12
-13.44
-5.93
-4.13
Tchr pay link stud. perf. Sample size 2.209
1.487
3.28
-2.279**
0.703
-1.60
-0.035
-0.03
-4.92
Teachers are mentored
6.766
0.845
5.72
7.722***
0.684
5.28
7.244
1.17
-0.73
Constant
154.46***
1.000
154.46
160.07***
1.000
160.07
157.26
0.00
-5.61
Column sum:
--
--
516.54
--
--
462.80
--
-1.62
55.36 Table 9B: Math Decomposition, 2015 (diff = 501.23– 456.61 = 44.62)
Variable
βvn
𝐱̅vn
βvnʹ𝐱̅vn
βo
𝐱̅o
βoʹ𝐱̅o 𝛃̅ (= (βvn+βo)/2
𝛃̅ʹ(𝐱̅vn-𝐱̅o)
(βvn - 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo)ʹ𝐱̅o
Wealth
6.573***
2.275
14.95
10.590***
4.198
44.47
8.581
-16.51
-13.00
Grade 10
79.641***
0.894
71.20
22.482***
0.592
13.30
51.061
6.80
51.10
Mom years schooling
1.214**
8.030
9.75
2.493***
11.449
28.55
1.853
-8.53
-10.27
Dad years schooling
2.059***
8.532
17.57
3.005***
11.537
34.67
2.532
-9.03
-8.07
Education input index
8.585***
4.031
34.61
7.360***
4.415
32.49
7.972
-3.05
5.17
Books in home
-0.021**
69.165
-1.42
0.089***
113.154
10.10
0.034
-3.93
-7.59
Class size
0.045
40.612
1.81
-0.033*
31.084
-1.02
0.006
-0.31
3.14
Proport. qualified tchrs
-5.666*
0.850
-4.82
26.023***
0.805
20.94
10.178
1.19
-26.95
Prop. Qual. tchr. missing
-5.084
0.070
-0.36
-11.814***
0.085
-1.00
-8.449
0.17
0.47
Square root comp/pupil
11.477**
0.442
5.07
10.850***
0.649
7.05
11.163
-2.25
0.28
Stud perf. Sample size to assess tchrs
-19.137*
0.990
-18.94
-5.876***
0.879
-5.16
-12.507
-0.65
-13.13
Teacher absenteeism
-2.328
1.604
-3.74
-7.533***
1.833
-13.81
-4.931
1.72
8.35
Principal observes tchrs
42.505**
0.994
42.26
1.178**
0.875
1.03
21.842
0.14
41.09
nspector observes tchrs
4.869*
0.766
3.73
-1.833***
0.568
-1.04
1.518
-0.36
5.14
Teacher are mentored
12.247***
1.808
22.15
-1.167***
1.462
-1.71
5.540
-0.40
24.26
Constant
307.41***
1.000
307.41
287.75***
1.000
287.75
297.58
0.00
19.66
Column sum:
--
--
501.23
--
--
456.61
--
-16.69
61.31 Table 9B: Math Decomposition, 2015 (diff = 501.23– 456.61 = 44.62) 57 Table 10A: Reading Decomposition, 2012 (diff = 512.82– 472.52 = 40.30)
Variable
βvn
𝐱̅vn
βvnʹ𝐱̅vn
βo
𝐱̅o
βoʹ𝐱̅o
𝛃̅ (= (βvn+βo)/2 𝛃̅ʹ(𝐱̅vn-𝐱̅o) (βvn - 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo
Wealth
4.748***
4.143
19.67
9.305***
6.101
56.77
7.026
-13.75
-23.34
Grade 10
79.18***
0.874
69.18
20.58***
0.584
12.01
49.88
14.46
42.70
Sibling index
4.045**
1.048
4.24
-1.736***
1.086
-1.89
1.154
-0.04
6.17
Sibling index missing
-0.428
0.149
-0.06
-12.01***
0.238
-2.86
-6.217
0.55
2.24
Mom years schooling
0.721**
8.313
5.99
1.083***
10.975
11.88
0.902
-2.40
-3.49
Dad years schooling
0.694**
8.883
6.17
1.877***
11.086
20.81
1.286
-2.83
-11.81
Years in preschool
4.884**
1.600
7.81
10.98***
1.487
16.34
7.933
0.89
-9.41
Education inputs index
5.657***
4.680
26.47
8.061***
5.154
41.55
6.859
-3.25
-11.82
Books in home
0.00231
57.59
0.13
0.0741***
114.07
8.45
0.038
-2.16
-6.16
Days attend past 2 wks
16.08***
9.849
158.34
7.806***
9.622
75.11
11.94
2.71
80.52
Hours study per week
2.335***
5.756
13.44
2.786***
5.362
14.94
2.651
1.01
-2.51
Extra reading class hr/wk -1.547***
2.741
-1.99
-4.887***
1.325
-4.61
-3.217
-1.11
3.73
Extra reading class miss. 0.712
0.336
0.24
-3.434***
0.358
-1.23
-1.361
0.03
1.44
Class size
0.258
44.81
11.58
0.358***
32.61
11.67
0.308
3.76
3.85
Proport. qualified tchrs
16.22***
0.800
12.98
35.92***
0.834
29.95
26.07
-0.88
-16.09
Prop. qual. tchr. missing
-17.21***
0.069
-1.19
-16.85***
0.188
-3.17
-17.03
2.03
-0.05
Square root comp/pupil
-4.467
0.417
-1.86
7.049***
0.623
4.40
1.291
-0.27
-5.99
Stud perf. to assess tchrs
1.901
0.992
1.89
-4.253**
0.708
-3.01
-1.176
-0.33
5.23
Teacher absenteeism
-1.489
0.692
-1.03
-5.874***
0.778
-4.57
-3.681
0.32
3.22
Parents pressure tchrs
9.980**
1.311
13.08
8.313***
0.957
7.96
9.146
3.24
1.89
Principal observes tchrs
34.74***
0.965
33.53
-1.893
0.802
-1.52
16.42
2.68
32.37
Inspector observes tchrs
-18.02**
0.847
-15.26
-11.80***
0.406
-4.79
-14.91
-6.57
3.90
Tchr pay link stud. perf. Sample size Yet these
1,717,966 individuals would be 18 years old in 2012, not 15 years old. The same census report
shows 1,450,815 12-year-olds in 2009, and these individuals would then be 15 in 2012. Thus the
correct PISA coverage rate for 2012 should be 65.9% (956,517/1,450,815). Appendix A: Further Derivations and Proof Appendix A: Further Derivations and Proof
A1. Re-calculations of the PISA Coverage Rates
The 55.7% coverage rate in the 2012 PISA report was obtained by taking Ministry of Education
and Training (MoET) records, which showed a “weighted number of participating students” of
956,517 students enrolled in school who were 15 years old, divided by 1,717,996 15-year-olds in
Vietnam (see Table 11.1 in OECD, 2014b). The 1,717,996 figure was obtained from the 2009
Census (General Statistics Office, 2010, Table 3); it is the number of 15-year-olds in Vietnam in
2009, and an implicit assumption was made that this number would be the same in 2012. Yet these
1,717,966 individuals would be 18 years old in 2012, not 15 years old. The same census report
shows 1,450,815 12-year-olds in 2009, and these individuals would then be 15 in 2012. Thus the
correct PISA coverage rate for 2012 should be 65.9% (956,517/1,450,815). The discrepancy for the 2015 PISA is even larger. The OECD report states that there were
1,803,552 15-year-olds in Vietnam in 2015 (OECD, 2016, Table A2.1), which seems to be based
on an assumption of slow population growth based on the 1,717,966 figure used for 2012. Yet the
2009 census shows only 1,332,822 9-year-old children in 2009, and these are the individuals who
would have been 15 years old in 2015. The 2015 PISA report (OECD, 2016, Table A2.1) shows
a “weighted number of participating students” of 874,859 students enrolled in school who were 15
years old in 2015, and dividing this figure by 1,803,552 gives a coverage rate of only 48.5%
(874,859/1,803,552). Yet the correct coverage rate should be 65.6% (874,859/1,332,822). A2. Proof of Proposition 1 p
Assume that the true test scores follow a normal distribution, with mean 𝜇 and standard deviation
𝜎. The truncated mean from below is given by 𝑇̅𝑏= 𝐸(𝑇|𝑇> 𝜏). The given school enrollment
rate is r. Sample size 3.676
1.487
5.47
-4.785***
0.703
-3.36
-0.555
-0.43
9.27
Teachers are mentored
9.211
0.845
7.78
7.342***
0.684
5.02
8.276
1.34
1.43
Constant
136.21***
1.000
136.21
186.68***
1.000
186.61
161.45
0.00
-50.47
Column sum:
--
--
512.82
--
--
472.52
--
-1.02
41.32 Table 10A: Reading Decomposition, 2012 (diff = 512.82– 472.52 = 40.30) 58 Table 10B: Reading Decomposition, 2015 (diff = 495.09– 468.06 = 27.03) Table 10B: Reading Decomposition, 2015 (diff = 495.09– 468.06 = 27.03)
Variable
βvn
𝐱̅vn
βvnʹ𝐱̅vn
βo
𝐱̅o
βoʹ𝐱̅o 𝛃̅(=(βvn+βo)/2 𝛃̅ʹ(𝐱̅vn-𝐱̅o) (βvn - 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo)ʹ𝐱̅o
Wealth
7.217***
2.275
16.42
11.487***
4.199
48.23
9.352
-17.99
-13.82
Grade 10
90.124***
0.894
80.57
21.122***
0.592
12.50
55.623
6.38
61.69
Mom years schooling
0.522*
8.030
4.19
2.399***
11.449
27.47
1.461
-8.20
-15.07
Dad years schooling
1.716***
8.532
14.64
2.252***
11.537
25.98
1.984
-6.77
-4.57
Education input index
6.883***
4.032
27.75
8.435***
4.415
37.24
7.659
-2.93
-6.55
Books in home
-0.019**
69.165
-1.30
0.077***
113.154
8.71
0.029
-3.39
-6.63
Class size
0.066
40.612
2.69
0.134***
31.084
4.17
0.100
1.28
-2.75
Proport. qualified tchrs
0.242
0.850
0.21
15.364***
0.805
12.36
7.803
0.70
-12.86
Prop. Qual. tchr. missing -12.390***
0.070
-0.87
-5.350***
0.085
-0.45
-8.870
0.08
-0.49
Square root comp/pupil
23.429***
0.442
10.36
18.271***
0.649
11.87
20.850
-3.79
2.28
Stud perf. to assess tchrs
-28.768**
0.990
-28.48
-2.588***
0.879
-2.27
-15.678
-0.29
-25.91
Teacher absenteeism
1.231
1.604
1.98
-3.973***
1.833
-7.28
-1.371
0.91
8.35
Principal observes tchrs
37.289**
0.994
37.08
-0.191
0.875
-0.17
18.549
-0.02
37.27
Inspector observes tchrs
0.567
0.766
0.43
-4.498***
0.568
-2.55
-1.966
-0.89
3.88
Teacher are mentored
11.366***
1.808
20.56
-0.563**
1.462
-0.82
5.402
-0.20
21.57
Constant
308.871***
1.000
308.87 293.091***
1.000
293.09
300.981
0.000
15.78
Column sum:
--
--
495.09
--
--
468.06
--
-16.96
44.00 59 Appendix A: Further Derivations and Proof
A1. Re-calculations of the PISA Coverage Rates
The 55.7% coverage rate in the 2012 PISA report was obtained by taking Ministry of Education
and Training (MoET) records, which showed a “weighted number of participating students” of
956,517 students enrolled in school who were 15 years old, divided by 1,717,996 15-year-olds in
Vietnam (see Table 11.1 in OECD, 2014b). The 1,717,996 figure was obtained from the 2009
Census (General Statistics Office, 2010, Table 3); it is the number of 15-year-olds in Vietnam in
2009, and an implicit assumption was made that this number would be the same in 2012. and 𝜇𝑢𝑡 =𝑇̅𝑎+ 𝜎 𝜆𝑎(𝛼) 𝜇𝑢𝑡 =𝑇̅𝑎+ 𝜎 𝜆𝑎(𝛼) Combining Equations (1.1) and (1.11), we can solve for 𝜎 in this case as
̅ Combining Equations (1.1) and (1.11), we can solve for 𝜎 in this case as
𝜏−𝑇̅𝑎
(1 12) 𝜎=
𝜏−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼
(1.1 𝜎=
𝜏−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼 𝜎=
𝜏−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼 𝑎( )
Plugging this result into Equation (1.1), we have the stated result
̅ 𝑎( )
Plugging this result into Equation (1.1), we have the stated result
𝑇̅ 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝜏−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼
(1.13) 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝜏−𝑇𝑎
𝜆𝑎(𝛼)+𝛼 (1.13) Note that the truncation point 𝜏 can be empirically estimated with the maximal observed test score
in the data Tmax, which results in the following estimating equation for Equation (1.13) Note that the truncation point 𝜏 can be empirically estimated with the maximal observed test score
in the data Tmax, which results in the following estimating equation for Equation (1.13) 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝑇𝑚𝑎𝑥−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼
(1.14) 𝜇𝑢𝑡 = 𝑇̅𝑎+ 𝜆𝑎(𝛼)
𝑇𝑚𝑎𝑥−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼
(1.14) (1.14) 𝜎=
𝜏−𝑇̅𝑎
𝜆𝑎(𝛼)+𝛼 A2. Proof of Proposition 1 However, since we
now assume that the PISA students are all worse-performing children, we need to rewrite Equation
(1.2) to reflect this assumption as follows For equation (2) in Proposition 1, define 𝜆𝑎(𝛼) instead as
𝜙(𝛼)
Φ(α), also using Theorem 19.2 in Greene
(2018), Equation (1.1) still holds for the case of the truncation from above. However, since we
now assume that the PISA students are all worse-performing children, we need to rewrite Equation
(1.2) to reflect this assumption as follows
𝑃(𝑇≤)
(1 9) 𝑃(𝑇≤𝜏) = 𝑟
(1.9) 𝑃(𝑇≤𝜏) = 𝑟 (1.9) Again, making use of the assumption that T follows a normal distribution, after some similar
straightforward manipulations for Equation (1.9) as with the proof for Proposition 1.1, we have
𝛼= Φ−1(𝑟)
(1.10) Again, making use of the assumption that T follows a normal distribution, after some similar
straightforward manipulations for Equation (1.9) as with the proof for Proposition 1.1, we have
𝛼= Φ−1(𝑟)
(1.10) A2. Proof of Proposition 1 Define 𝛼 as
𝜏−µ
𝜎 and 𝜆𝑏(𝛼) as
𝜙(𝛼)
1−Φ(𝛼) , using Theorem 19.2 in Greene (2018), we have 𝜎
𝑏( )
1−Φ(𝛼) ,
g
(
),
𝜇𝑙𝑡 = 𝑇̅𝑏−𝜎 𝜆𝑏(𝛼)
(1.1)
Let r represent the given school enrollment rate, we also have
𝑃(𝑇> 𝜏) = 𝑟
(1.2)
Since T follows a normal distribution, subtracting the two sides of Equation (1.2) from unity yields
Φ (
𝜏−µ
𝜎) ≡Φ(𝛼) = 1 −𝑟
(1.3)
This leads to the following results
𝛼= Φ−1(1 −𝑟)
(1.4)
and
𝜇𝑙𝑡= 𝜏−𝜎𝛼
(1.5)
Combining Equations (1.1) and (1.5), we can solve for 𝜎 as
𝜎=
𝑇̅𝑏−𝜏
𝜆𝑏(𝛼)−𝛼
(1.6)
Plugging this result into Equation (1.1), we have the stated result
𝜇𝑙𝑡 = 𝑇̅𝑏− 𝜆𝑏(𝛼)
𝑇̅𝑏−𝜏
𝜆𝑏(𝛼)−𝛼
(1.7) 1 Φ(𝛼)
𝜇𝑙𝑡 = 𝑇̅𝑏−𝜎 𝜆𝑏(𝛼)
(1.1) ( )
𝜇𝑙𝑡 = 𝑇̅𝑏−𝜎 𝜆𝑏(𝛼) t the given school enrollment rate, we also have
𝑃(𝑇> )
(1 2) Let r represent the given school enrollment rate, we also have
𝑃(𝑇> 𝜏) = 𝑟
(1.2) Let r represent the given school enrollment rate, we also have
𝑃(𝑇> 𝜏) = 𝑟
(1.2) ,
𝑃(𝑇> 𝜏) = 𝑟
b
i
h 𝑃(𝑇> 𝜏) = 𝑟
(1.2) Since T follows a normal distribution, subtracting the two sides of Equation (1.2) s a normal distribution, subtracting the two sides of Equation (1.2) from unity yields g
Φ (
𝜏−µ
𝜎) ≡Φ(𝛼) = 1 −𝑟 Φ (
𝜏−µ
𝜎) ≡Φ(𝛼) = 1 −𝑟 Φ (
µ
𝜎) ≡Φ(𝛼) = 1 −𝑟
(1.3)
This leads to the following results
𝛼= Φ−1(1 −𝑟)
(1.4)
and 𝜇𝑙𝑡= 𝜏−𝜎𝛼
Combining Equations (1.1) and (1.5), we can solve for 𝜎 as
̅ 𝜇𝑙𝑡= 𝜏−𝜎𝛼
(1.5)
Combining Equations (1.1) and (1.5), we can solve for 𝜎 as
𝜎=
𝑇̅𝑏−𝜏
(1 6) 𝜎=
𝑇̅𝑏−𝜏
𝜆𝑏(𝛼)−𝛼 𝜎=
𝑇̅𝑏−𝜏
𝜆𝑏(𝛼)−𝛼 𝑏( )
Plugging this result into Equation (1.1), we have the stated result
̅ 𝑏( )
Plugging this result into Equation (1.1), we have the stated result
̅ )
𝜇𝑙𝑡 = 𝑇̅𝑏− 𝜆𝑏(𝛼)
𝑇̅𝑏−𝜏
𝜆𝑏(𝛼)−𝛼
(1.7) (1.7) 𝑏( )
Note that the truncation point 𝜏 can be empirically estimated with the minimal observed test score
in the data Tmin, which results in the following estimating equation for Equation (1.7)
̅ 𝑏( )
Note that the truncation point 𝜏 can be empirically estimated with the minimal observed test score
in the data Tmin, which results in the following estimating equation for Equation (1.7)
̅ 𝜇𝑙𝑡 = 𝑇̅𝑏− 𝜆𝑏(𝛼)
𝑇̅𝑏−𝑇𝑚𝑖𝑛
𝜆𝑏(𝛼)−𝛼
(1.8) 𝜇𝑙𝑡 = 𝑇̅𝑏− 𝜆𝑏(𝛼)
𝑇̅𝑏−𝑇𝑚𝑖𝑛
𝜆𝑏(𝛼)−𝛼
(1.8) (1.8) For equation (2) in Proposition 1, define 𝜆𝑎(𝛼) instead as
𝜙(𝛼)
Φ(α), also using Theorem 19.2 in Greene
(2018), Equation (1.1) still holds for the case of the truncation from above. A3. Procedure to Adjust PISA Test Scores with Young Lives Data The adjustments to the PISA scores using the Young Lives data were done as follows. First, the
Young Lives sample was sorted into 10 deciles based on the average test scores over the math and
reading comprehension tests, where Decile 1 has the 10% of the Young Lives sample with the
lowest scores, Decile 2 has the 10% with the next lowest test scores, and so forth, up to Decile 10,
which has the 10% of the sample with the highest scores. For all 10 deciles, the proportion of
Young Lives students who were still in school was calculated, which ranged from 0.582 for Decile
1 to 1.000 for Decile 10. These proportions are shown in column (1) of Table B3 in Appendix B. These proportions serve two purposes. First, they are used to assign students in the Vietnam PISA
sample to deciles (ranked by student performance) of the distribution of all 15-year-olds, including
those not in school. An initial step is to adjust the proportions in school (for the Young Lives
sample) in column (1) so that they have a mean of 1.000; this shows how 15-year-olds in that
sample who are in school are distributed across the deciles of the distribution of academic
performance for all 15-year-olds. Thus column (2) in Table B3 shows that, of all 15-year-olds in
school, 7.01% are in the bottom decile of the distribution of the academic performance of all 15-
year-olds, 7.76% are in the second decile of that distribution, and so forth, and finally 12.03% are
in the top decile; note that these percentages sum to 100%. Then the bottom 7.01% (in terms of
academic performance on the PISA) of the 15-year-old PISA participants are assigned to the
bottom decile of this “all 15-year-olds” distribution, the next 7.76% are assigned to the second
decile, and so forth. The second purpose of the proportions in column (1) is to generate “inflation factors” for the PISA
students assigned to these deciles. These inflation factors, which are shown in column (3) and are 61 the inverses of the proportions in column (2), are applied to the PISA participants assigned to these
deciles to approximate the distribution of test scores that would have been generated if the entire
population of 15-year-olds in Vietnam had participated in the PISA. A3. Procedure to Adjust PISA Test Scores with Young Lives Data For example, the 7.01% of
the PISA participants assigned to the first decile are given a weight of 1.427 (= 1/7.01) so that they
represent the bottom 10% of the entire population of 15-year-olds. This is last step in the
adjustment, which is to assume that the mean scores of 15-year-olds not in school in each decile
of the population are equal to the mean scores of the 15-year-olds in the respective deciles who
are in school, and thus participated in the PISA. The means for the latter (which by this assumption
are also assigned to the former) for the 2012 PISA are shown in column (4) of Table B3, separately
for mathematics and reading. They can be compared to the actually means, by decile, in the 2012
PISA, which are shown in column (5). The overall results show that this adjustment decreases the
2012 PISA scores for math by only 12.8 points and the 2012 PISA scores for reading by only 11.3
points. The same adjustments for the 2015 PISA are shown in columns (6) and (7); they show that
this this adjustment decreases the 2015 PISA scores for math by only 12.4 points and the 2015
PISA scores for reading by only 10.9 points. 62 62 Appendix B: Additional Tables and Figures
Figure B1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita 63
Appendix B: Additional Tables and Figures
Figure B1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita
Figure B2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita Appendix B: Additional Tables and Figures
Figure B1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita Appendix B: Additional Tables and Figures
Figure B1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita pp
g
Figure B1. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real PPP G 63
re B2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita igure B2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita ean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real PPP GDP/capita e B2. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real PPP GDP/ 63 Figure B3. A3. Procedure to Adjust PISA Test Scores with Young Lives Data Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita, East
Asia Only
i
4
A
1
i
S
i
2012
SA
2010
G
/
i
MYS
IDN
THA
MAC
JPN
HKG
KOR
SGP
VNM
350
400
450
500
550
600
7
8
9
10
11
lgdppc2010real
PISA 2012 Avg. Math Score
Fitted values Figure B3. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita, East ean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita, East Figure B3. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita, East
Asia Only
Figure B4. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita, East
Asia Only
MYS
IDN
THA
MAC
JPN
HKG
KOR
SGP
VNM
350
400
450
500
550
600
7
8
9
10
11
lgdppc2010real
PISA 2012 Avg. Math Score
Fitted values
MYS
IDN
THA
MAC
JPN
SGP
HKG
KOR
VNM
400
450
500
550 Figure B3. Mean Age 15 Math Scores in 2012 PISA, by 2010 Log Real GDP/capita, East
Asia Only
Figure B4. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita, East
Asia Only
MYS
IDN
THA
MAC
JPN
HKG
KOR
SGP
VNM
350
400
450
500
550
600
7
8
9
10
11
lgdppc2010real
PISA 2012 Avg. Math Score
Fitted values 9
lgdppc2010real Fitted values Figure B4. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita, Ea
Asia Only
MYS
IDN
THA
MAC
JPN
SGP
HKG
KOR
VNM
400
450
500
550
7
8
9
10
11
lgdppc2010real
PISA 2012 Avg. Reading Score
Fitted values Figure B4. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita, East
Asia Only
MYS
IDN
THA
MAC
JPN
SGP
HKG
KOR
VNM
400
450
500
550
7
8
9
10
11
lgdppc2010real
PISA 2012 Avg. Reading Score
Fitted values ean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita, East Figure B4. Mean Age 15 Reading Scores in 2012 PISA, by 2010 Log Real GDP/capita, East 9
lgdppc2010real 64 64 Figure B5. Distributions of 2015 PISA Scores for Four Countries with High Enrollment Rates igure B5. Distributions of 2015 PISA Scores for Four Countries with High Enr Figure B5. A3. Procedure to Adjust PISA Test Scores with Young Lives Data Distributions of 2015 PISA Scores for Four Countries with High Enrollment Rates ns of 2015 PISA Scores for Four Countries with High Enrollment Rates or Four Countries with High Enrollment Rates Figure B5. Distributions of 2015 PISA Scores f 65 Figure B6. Distribution of Test Scores, Truncated from Below Figure B6. Distribution of Test Scores, Truncated from Below Students in
the sample (enrolled in school) Students not
in the sample
(not enrolled in school)
Test Score Truncation
Point 66 Table B1: Predictors of 2012 PISA Scores in Vietnam
Variables
Math
Reading
Rural
-18.04***
-11.56**
(6.775)
(5.699)
Female
-16.58***
24.61***
(2.317)
(2.009)
Grade 10
105.8***
95.14***
(6.809)
(6.077)
Father years of schooling
2.231***
1.536***
(0.495)
(0.395)
Mother years of schooling
1.879***
1.661***
(0.489)
(0.422)
Owns an air conditioner
5.456
-0.626
(6.279)
(4.450)
Owns a car
-6.723
-3.442
(4.645)
(3.892)
Owns a computer
17.35***
10.86***
(3.511)
(2.810)
Number of televisions owned
0.526
2.977
(2.425)
(2.187)
Constant
396.7***
385.2***
(8.881)
(8.545)
Observations
4771
4771
R-squared
0.310
0.341
Robust standard errors, clustered at the school level, in parentheses
*** p<0.01, ** p<0.05, * p<0.1 Table B1: Predictors of 2012 PISA Scores in Vietnam 67 Table B2: Predictors of 2015 PISA Scores in Vietnam Table B2: Predictors of 2015 PISA Scores in Vietnam Table B2: Predictors of 2015 PISA Scores in Vietnam
Variable
Math
Reading
Rural
-18.86***
-9.822
(4.98)
(5.908)
Female
15.97***
-8.461***
(2.05)
(2.272)
Grade 10
69.85***
74.61***
(7.19)
(6.07)
Mother years of schooling
0.893**
1.460***
(0.408)
(0.541)
Father years of schooling
1.646***
2.041***
(0.328)
(0.373)
Owns an air conditioner
-0.712
-2.685
(4.126)
(4.971)
Owns a motorbike
15.83***
6.451
(5.01)
(5.974)
Owns a car
5.202
-1.249
(4.758)
(5.950)
Own a computer
16.61***
23.39***
(2.611)
(3.34)
Number of televisions owned
7.284***
6.734**
(2.141)
(2.601)
Constant
376.9***
386.4***
(9.31)
(10.41)
Observations
5687
5687
R2
0.274
0.207
Robust standard errors, clustered at the school level, in parentheses
* p < 0.10, ** p < 0.05, *** p < 0.01 68 Table B3. A3. Procedure to Adjust PISA Test Scores with Young Lives Data Adjusted PISA 2012 and 2015 Test Scores Using Young Lives Attrition Data
Test
Score
Decile
(1)
Proportion
in School in
Young Lives
Data
(2)
Proportions
Divided by
0.831
((1) ÷ 0.831)
(3)
Inflation
Factor for
PISA Sample
(1/(2))
(4)
Adjusted 2012
PISA Scores, by
Decile (all 15-
year-olds)
(5)
Original 2012
PISA Scores, by
Decile (in school
only)
(6)
Adjusted 2015
PISA Scores, by
Decile (all 15-
year-olds)
(7)
Original 2015
PISA Scores, by
Decile (in school
only)
Math
Reading
Math
Reading
Math
Reading
Math Reading
1
0.582
0.701
1.427
358.0
363.4
364.3
370.3
340.8
349.0
352.2
359.6
2
0.646
0.776
1.289
409.4
419.2
421.5
432.1
391.2
396.1
405.2
409.9
3
0.746
0.897
1.115
442.1
449.6
454.3
461.7
419.9
424.8
434.6
438.7
4
0.761
0.915
1.093
463.2
472.1
477.2
484.8
444.1
447.2
458.9
460.1
5
0.849
1.022
0.978
483.2
492.6
498.7
502.7
466.3
467.0
479.8
478.8
6
0.885
1.065
0.939
507.2
509.4
521.2
520.1
488.0
485.7
502.5
497.4
7
0.920
1.106
0.904
530.1
528.2
543.8
539.5
512.7
506.1
525.7
516.3
8
0.951
1.144
0.874
555.4
548.1
568.5
558.8
539.6
527.9
550.9
537.4
9
0.973
1.171
0.854
586.6
570.8
600.6
583.2
571.5
556.1
580.7
564.5
10
1.000
1.203
0.831
648.7
615.9
662.6
630.1
635.6
608.7
643.6
614.8
Average
0.831
1.000
498.4
496.9
511.2
508.2
481.0
476.9
493.4
487.8 Adjusted PISA 2012 and 2015 Test Scores Using Young Lives Attrition Data Table B.4: Math Decomposition Using Fixed-Effects Estimates of βo (diff = 516.54– 462.80 = 53.74)
Variable
βvn
𝐱̅vn
βvnʹ𝐱̅vn
βo
𝐱̅o
βoʹ𝐱̅o 𝛃̅ (= (βvn+βo)/2 𝛃̅ʹ(𝐱̅vn-𝐱̅o) (βvn - 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo)ʹ𝐱̅o
Wealth
6.764***
4.143
28.02
5.316***
6.101
32.43
6.040
-11.83
7.42
Grade 10
85.85***
0.874
75.01
19.34***
0.584
11.29
52.595
15.25
48.49
Sibling index
3.152*
1.048
3.30
-2.343***
1.086
-2.54
0.405
-0.02
5.86
Sibling index missing
-0.576
0.149
-0.09
-18.19***
0.238
-4.33
-9.383
0.84
3.41
Mom years schooling
0.962***
8.313
8.00
1.657***
10.975
18.19
1.310
-3.49
-6.70
Dad years schooling
1.511***
8.883
13.42
1.991***
11.086
22.07
1.751
-3.86
-4.79
Years in preschool
6.533***
1.600
10.45
9.972***
1.487
14.83
8.253
0.93
-5.31
Education inputs index
4.397***
4.680
20.58
6.858***
5.154
35.35
5.628
-2.67
-12.10
Books in home
0.00887
57.59
0.51
0.0677***
114.07
7.72
0.038
-2.16
-5.05
Days attend past 2 wks
10.43***
9.849
102.72
7.040***
9.622
67.74
8.735
1.98
33.00
Hours study per week
2.920***
5.756
16.81
2.882***
5.362
15.45
2.901
1.14
0.21
Extra math class, hrs/wk
3.904***
2.741
10.70
-0.858***
1.325
-1.14
1.523
2.16
9.68
Extra math class missing 8.890***
0.336
2.98
-2.590***
0.358
-0.93
3.150
-0.07
3.98
Class size
0.0643
44.81
2.88
0.0657***
32.61
2.14
0.065
0.79
-0.05
Proport. A3. Procedure to Adjust PISA Test Scores with Young Lives Data qualified tchrs
18.18***
0.800
14.55
12.62***
0.834
10.53
15.400
-0.52
4.54
Prop. qual. tchr. missing -17.15***
0.069
-1.18
-0.486
0.188
-0.09
-8.818
1.05
-2.14
Square root comp/pupil
-0.0392
0.417
-0.02
-0.782
0.623
-0.49
-0.411
0.08
0.39
Stud perf. to assess tchrs
25.08**
0.992
24.89
1.708***
0.708
1.21
13.394
3.80
19.87
Teacher absenteeism
-0.759
0.692
-0.53
-3.475***
0.778
-2.70
-2.117
0.18
2.00
Parents pressure tchrs
15.71***
1.311
20.60
11.23***
0.957
10.75
13.470
4.77
5.08
Principal observes tchrs
14.12**
0.965
13.63
-2.586***
0.802
-2.07
5.767
0.94
14.76
Inspector observes tchrs -16.73***
0.847
-14.17
-4.317***
0.406
-1.75
-10.524
-4.64
-7.78
Tchr pay link stud. perf. 2.209
1.487
3.28
-2.397***
0.703
-1.69
-0.094
-0.07
5.04
Teachers are mentored
6.766**
0.845
5.72
5.030***
0.684
3.44
5.898
0.95
1.33
Constant
154.46***
1.000
154.46
227.4***
1.000
227.39
190.93
0.00
-72.93
516.54
462.80
5.55
48.21 mposition Using Fixed-Effects Estimates of βo (diff = 516.54– 462.80 = 53.74) Table B.5: Reading Decomposition Using Fixed-Effects Estimates of βo (diff = 512.82– 472.52 = 40.30)
Variable
βvn
𝐱̅vn
βvnʹ𝐱̅vn
βo
𝐱̅o
βoʹ𝐱̅o
𝛃̅ (= (βvn+βo)/2 𝛃̅ʹ(𝐱̅vn-𝐱̅o) (βvn - 𝛃̅)ʹ𝐱̅vn + (𝛃̅ – βo)ʹ𝐱̅
Wealth
4.748***
4.143
19.67
3.859***
6.101
23.54
4.3035
-8.43
4.55
Grade 10
79.18***
0.874
69.18
21.15***
0.584
12.35
50.165
14.55
42.30
Sibling index
4.045***
1.048
4.24
-2.214***
1.086
-2.40
0.9155
-0.03
6.68
Sibling index missing
-0.428
0.149
-0.06
-12.56***
0.238
-2.99
-6.494
0.58
2.35
Mom years schooling
0.721**
8.313
5.99
1.297***
10.975
14.23
1.009
-2.69
-5.55
Dad years schooling
0.694**
8.883
6.17
1.702***
11.086
18.87
1.198
-2.64
-10.06
Years in preschool
4.884***
1.600
7.81
10.28***
1.487
15.29
7.582
0.86
-8.33
Education inputs index
5.657***
4.680
26.47
8.447***
5.154
43.54
7.052
-3.34
-13.72
Books in home
0.00231
57.59
0.13
0.0572***
114.07
6.52
0.029755
-1.68
-4.71
Days attend past 2 wks
16.08***
9.849
158.34
7.325***
9.622
70.48
11.7025
2.66
85.23
Hours study per week
2.335***
5.756
13.44
3.225***
5.362
17.29
2.78
1.10
-4.95
Extra reading class hr/wk -1.547***
2.741
-1.99
-4.460***
1.325
-5.91
-3.0035
-4.25
5.92
Extra reading class miss. 0.712
0.336
0.24
-3.047***
0.358
-1.09
-1.1675
0.03
1.30
Class size
0.258***
44.81
11.58
0.261***
32.61
8.51
0.2595
3.17
-0.12
Proport. qualified tchrs
16.22***
0.800
12.98
9.841***
0.834
8.21
13.0305
-0.44
5.21
Prop. qual. tchr. missing -17.21***
0.069
-1.19
-2.079***
0.188
-0.39
-9.6445
1.15
-1.94
Square root comp/pupil
-4.467
0.417
-1.86
0.639
0.623
0.40
-1.914
0.39
-2.66
Stud perf. A3. Procedure to Adjust PISA Test Scores with Young Lives Data to assess tchrs
1.901
0.992
1.89
2.067***
0.708
1.46
1.984
0.56
-0.14
Teacher absenteeism
-1.489
0.692
-1.03
-3.003***
0.778
-2.34
-2.246
0.19
1.11
Parents pressure tchrs
9.980***
1.311
13.08
11.22***
0.957
10.74
10.6
3.75
-1.41
Principal observes tchrs
34.74***
0.965
33.53
-0.136
0.802
-0.11
17.302
2.82
30.81
Inspector observes tchrs -18.02***
0.847
-15.26
-6.500***
0.406
-2.64
-12.26
-5.41
-7.22
Tchr pay link stud. perf. 3.676***
1.487
5.47
-2.740***
0.703
-1.93
0.468
0.37
7.03
Teachers are mentored
9.211***
0.845
7.78
5.721***
0.684
3.91
7.466
1.20
2.67
Constant
136.21***
1.000
136.21
237.0***
1.000 236.97***
186.59
0.00
-100.76
512.82
472.52
4.45
38.04 71
|
https://openalex.org/W2801091560
|
https://europepmc.org/articles/pmc5943252?pdf=render
|
English
| null |
Contrasting roles for actin in the cellular uptake of cell penetrating peptide conjugates
|
Scientific reports
| 2,018
|
cc-by
| 10,794
|
Contrasting roles for actin in the
cellular uptake of cell penetrating
peptide conjugates Received: 15 December 2017
Accepted: 25 April 2018
Published: xx xx xxxx L. He1, E. J. Sayers 1, P. Watson
2 & A. T. Jones 1 The increased need for macromolecular therapeutics, such as peptides, proteins and nucleotides,
to reach intracellular targets necessitates more effective delivery vectors and a higher level of
understanding of their mechanism of action. Cell penetrating peptides (CPPs) can transport a range
of macromolecules into cells, either through direct plasma membrane translocation or endocytosis. All known endocytic pathways involve cell-cortex remodelling, a process shown to be regulated
by reorganisation of the actin cytoskeleton. Here using flow cytometry, confocal microscopy and a
variety of actin inhibitors we identify how actin disorganisation in different cell types differentially
influences the cellular entry of three probes: the CPP octaarginine – Alexa488 conjugate (R8-Alexa488),
octaarginine conjugated Enhanced Green Fluorescent Protein (EGFP-R8), and the fluid phase probe
dextran. Disrupting actin organisation in A431 skin epithelial cells dramatically increases the uptake
of EGFP-R8 and dextran, and contrasts strongly to inhibitory effects observed with transferrin and
R8 attached to the fluorophore Alexa488. This demonstrates that uptake of the same CPP can occur
via different endocytic processes depending on the conjugated fluorescent entity. Overall this study
highlights how cargo influences cell uptake of this peptide and that the actin cytoskeleton may act as a
gateway or barrier to endocytosis of drug delivery vectors. Cell penetrating peptides (CPPs) are a group of short sequences typically containing 5–30 amino acids that have
been extensively investigated as carriers for intracellular delivery of various cargos including genetic material,
peptides, proteins and nanoparticles1–4 Numerous efforts have been made to unveil the mechanisms of CPP trans-
location to the cytoplasm and cytosol of cells, and it is now well accepted that two modes of cell entry exist: direct
membrane translocation, which may be energy and temperature independent, and uptake via one or more energy
dependent endocytic pathways5,6. The propensity for uptake via these mechanisms is dependent on the peptide
sequence, choice of cargo, in vitro model and can be influenced by experimental factors, including incubation
temperature and the presence or absence of serum in media7. In a number of CPP studies an intact actin cytoskel-
eton has been proposed to be required for cell internalisation and CPPs inside and outside of cells can modify the
actin cytoskeleton to influence cellular processes including CPP entry8–11. www.nature.com/scientificreports www.nature.com/scientificreports www.nature.com/scientificreports Materials and Methods The cells were then incubated for 1 hr (or
30 min for TF-647) at 37 °C/5% CO2 before washing 3x with PBS (TF-647 or Dex-647) or PBS 0.5 mg/mL heparin
(R8-conjugates). They were then washed once with PBS, prior to the addition of 2 mL of imaging medium (phenol
red free RPMI, 20 mM HEPES pH 7.4) and analysis by live cell confocal microscopy. Actin inhibition and confocal microscopy of endocytic probes and R8 conjugates. Cells were
seeded on MatTek dishes as previously described and were washed with PBS before preincubation at 37 °C with
diluent control (DMSO) or actin inhibitors: 10 µM Cyt D for 15 min; 0.1–1 µM Lat B for 30 min; 0.2–4 µM JAS for
45 min; 50–100 µM Y27632 for 4 hr in serum-free D-MEM at 37 °C/5% CO2. Endocytic probes (TF-647, Dex-647)
or R8-conjugates were then incubated with the cells in the absence (control) or presence of the inhibitors and
further incubated at 37 °C/5% CO2 for times specified in figure legends prior to analysis by confocal microscopy. For pulse chase experiments, A431 cells were incubated with 0.1 mg/ml Dex-647 for 2 hr (pulse) followed by
washing and a further 4 hr (chase) in fresh complete growth medium. EGFP-R8 (2 µM) was then added for 1 hr in
the absence or presence of 10 µM Cyt D. EGFP-R8 and Dex-647 subcellular distribution was then analysed in live
cells using confocal microscopy. For low temperature experiments, cells were preincubated with Cyt D as above
and then placed on ice for 2 min prior to addition of ice cold EGFP-R8. The cells were then maintained under
these conditions for 1 hr prior to analysis by confocal microscopy. Confocal microscopy. Analysis was performed on a Leica SP5 confocal laser scanning microscope,
equipped with laser lines: 405 Blue Diode (Excitation wavelength 405 nm for Hoechst 33342) Argon (Excitation
wavelength 488 nm for Alexa 488 or EGFP), Helium Neon Laser 1 (Excitation wavelength 543 nm for
Tetramethyl-Rhodamine Phalloidin (Rh-P)) and Laser 2 (Excitation wavelength 633 nm for Alexa 647 con-
jugates). All images presented in this study were obtained using a HCX PL APO 63x 1.4 NA oil immersion
objective with Leica Type F immersion oil. Parameters “Gain” and “Off-set” of the individual photomultiplier
tubes were adjusted between experiments in order to obtain optimal image acquisition avoiding saturation. Materials and Methods Cell culture. HeLa (cervical carcinoma, epithelial) cells (ATCC Code CCL-2) or A431 (skin epidermal) cells
(ATCC Code CRL-1555) were maintained in growth Dulbecco’s Modified Eagle’s medium (D-MEM) supple-
mented with foetal bovine serum (10%, final volume) and penicillin/streptomycin (100 units/ml and 100 μg/ml
respectively), herein referred to as complete medium in a humidified 5% CO2/37 °C incubator as a subconfluent
monolayer. Cells were obtained from the ATCC and were routinely tested for mycoplasma. Peptide Labelling. Peptide octaarginine (R8), extended with GC at its C-terminal (RRRRRRRR-GC), was
synthesized by American Peptide Company (California, USA) and labelled with Alexa Fluor® 488 C5 maleimide
salt (Invitrogen, Paisley, UK) as previously described30. Labelling and conjugate mass (2126.5 Da) was confirmed
by matrix-assisted laser-desorption ionization–time-of-flight spectrometry and quantified by UV spectrometry. All labelled peptides were resuspended to a stock concentration of 1 mM in autoclaved distilled water, aliquoted
and frozen at −80 °C until needed. Expression and purification of recombinant proteins. Plasmids encoding histidine tagged -
EGFP-octaarginine, or - EGFP referred to here as EGFP-R8 or EGFP, were a kind gift from Professor Shiroh
Futaki, University of Kyoto, Japan. To produce the plasmid the DNA fragment encoding EGFP was amplified
from pEGFP-N1 (Clontech) using primers possessing the sequences for NdeI followed by His6, and R8 and stop
codon followed by EcoRI. The fragment was cleaved by NdeI and EcoRI and cloned into a pET3b (Novagen)
derivative without N-terminal T7.Tag sequence to construct E. Coli expression vector pEV3b31 that has NdeI
and EcoRI sites within the multi cloning site. Proteins were produced in E. coli BL21 (DE3) cells and assessed for
purity and molecular weight as described in Supplementary Information and shown in Supplementary Fig. 1. Live cell imaging of endocytic probes and R8 conjugates. HeLa or A431 cells were seeded on 35 mm
MatTek dishes (Ashland, US) at respective densities of 5 × 105 cells/dish and 1.0 × 106 cells/dish and maintained
in 2.5 mL D-MEM containing 10%, 100 IU/mL penicillin and 100 µg/mL streptomycin. Cells were then allowed
to adhere for 24 hr at 37 °C/5% CO2 to reach a confluence of ~80%. On the day of experimentation, they were
washed with PBS and then incubated with 100 µl serum-free D-MEM (thus reducing well described serum
binding effects32) containing either 2 µM CPP conjugate (R8-Alexa488 or EGFP-R8), 5 µg/ml transferrin-Alexa
647 (TF-647) or 0.1 mg/ml dextran (10 kDa)-Alexa 647 (Dex-647). Contrasting roles for actin in the
cellular uptake of cell penetrating
peptide conjugates yl
g
y
One endocytic pathway that is absolutely reliant on actin is macropinocytosis. When activated this process
has the capacity to form large plasma membrane derived intracellular vesicles termed macropinosomes12–15. Classically macropincytosis is induced in response to growth factor activation such as epidermal growth factor
(EGF) binding to the EGF receptor, initially leading to extensive actin-dependent ruffling on the plasma mem-
brane. This induces a “gulping effect” manifest as an increased uptake of extracellular fluid13,14,16. Much of the
information known regarding growth factor induced and actin dependent macropinocytosis comes from studies
on high EGFR expressing A431 skin epithelia cells and their response to EGF13,17,18. Of interest are observations
that some CPPs under defined experimental conditions may induce plasma membrane effects similar to that seen
upon growth factor activation19–21 and in line with this that they promote the concomitant uptake of dextran, a
well characterised marker of fluid phase endocytosis22–24. Dextran itself, in addition to being widely used as a fluid
phase endocytic probe has been extensively investigated as a drug delivery vector25. Tools used routinely to examine the roles of the actin cytoskeleton in various cellular processes, including
endocytosis and CPP entry are pharmacological/chemical inhibitors. The most notable such agent is the fungal 1Cardiff School of Pharmacy and Pharmaceutical Sciences, Redwood Building, Cardiff University, Cardiff, Wales, CF10
3NB, UK. 2Cardiff School of Biosciences, The Sir Martin Evans Building, Cardiff University, Cardiff, Wales, CF10 3AX,
UK. L. He and E. J. Sayers contributed equally to this work. Correspondence and requests for materials should be
addressed to P.W. (email: watsonpd@cardiff.ac.uk) or A.T.J. (email: jonesat@cardiff.ac.uk) Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 1 www.nature.com/scientificreports/ metabolite cytochalasin D (Cyt D) which disrupts actin polymerisation and is a well characterised inhibitor of var-
ious endocytic mechanisms26–28. Other natural compounds and synthetic products such as Latrunculin B (Lat B)
and Jasplakinolide (JAS) have been identified or developed to target the actin directly or indirectly and to disrupt
its organisation and function29. Contrasting roles for actin in the
cellular uptake of cell penetrating
peptide conjugates Very few studies have investigated the effects of these other actin disrupters on
CPP uptake though it is generally recognised that actin disruption universally inhibits CPP entry.f p
g
g
y
g
p
y
y
Here we show that the effects of actin disruption on uptake of CPPs and dextran is cell type dependant and
in A431 skin epithelia, in complete contrast to HeLa cells, leads to a dramatic increase in uptake of EGFP-R8
and dextran but inhibits the uptake of R8-Alexa488. Together the data indicate that actin organisation has very
different influences on uptake of these octaarginine and fluid phase conjugates and that actin could be targeted to
enhance cellular uptake of drug delivery vectors. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 Results
ff Cyt D effects on actin and endocytosis in HeLa and A431 cells. Cyt D binds to the fast-growing ends
of actin nuclei and filaments, preventing addition of monomeric actin to these sites. In our previous studies using
flow cytometry we showed that Cyt D inhibited the uptake of Alexa488-CPPs R8 and Tat in two different cell mod-
els, HeLa and A43133. But interestingly, the effect of actin disruption with this agent on the uptake of dextran was
highly dependent on the choice of cell line as there was no effect in HeLa cells and an increased uptake in A431
cells. Here we further explored these contradictory effects by also performing uptake studies with recombinant
EGFP-R8 following purification from E. Coli as described in Supplementary methods and Supplementary Fig. 1. g pi
pp
y
pp
y
g
At 2 µM extracellular concentration, both R8 conjugates, in the two cell lines, labelled vesicular structures
scattered throughout the cytoplasm as expected (Supplementary Figs 2 and 3), no cell associated fluorescence
was observed for EGFP alone. We then performed the same experiments in cells incubated with Cyt D, focusing
on effects on endocytosis and also cell morphology. 10 μM of the inhibitor was used as this concentration has
previously been shown by different groups to inhibit the uptake of cationic CPPs such as HIV-Tat and R8 linked
to different cargoes33–35. DIC microscopy highlighted the dramatic morphological effects on both cell lines incu-
bated with this agent (Fig. 1 and Supplementary Fig. 4). Cyt D inhibited the internalisation of R8-Alexa 488 in
HeLa cells (Fig. 1A), confirming our published data using flow cytometry33. There was no clear evidence of Cyt D
mediated inhibition of EGFP-R8 but the drug caused a scattering of the fluorescent structures compared to
untreated cells that usually displayed strong juxtanuclear labelling (Fig. 1B). In contrast to HeLa cells there was
less of a noticeable reduction in uptake of R8-Alexa488 in A431 cells but a clear change in subcellular distribution
from evenly scattered throughout the cytoplasm in control cells to a largely clustered phenotype in treated cells
(Fig. 1C). In contrast to the results obtained with R8-Alexa 488, Cyt D dramatically enhanced the internalisation
of EGFP-R8 into small and large cytoplasmic structures (Fig. 1D). This data was confirmed in A431 cells using
flow cytometry showing a highly significant 4.5 fold increase in intracellular fluorescence (Fig. Materials and Methods For
multi-channel image acquisition, the channels were scanned in a sequential recording mode to avoid spectral
cross-talk caused by overlapping excitation and/or emission spectra of fluorophores. Acquisition mode “XYZ”
and image resolutions (pixels/image) of 512 × 512 or 1024 × 1024 were selected and pinhole size was set to 1
Airy Unit. Depending on acquisition requirements the scan speed was set as either 400 or 700 Hz and for single
section images, the same optical section was scanned 3x to generate a line average image. Captured images were Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 2 www.nature.com/scientificreports/ then analysed using ImageJ. In some cases (live cell imaging) the same cells were also imaged using Differential
Imaging Contrast microscopy (DIC). Actin cytoskeleton visualisation. A recently described method was utilised giving information on actin
architecture at different regions of the cells termed basal and cell body and apex (CBA). For actin staining, HeLa
(0.5 × 106 cells/well) or A431 (1.0 × 106 cells/well) cells in 2.5 mL D-MEM containing 10% FBS were seeded on
round borosilicate No.1.5 cover slips in 6 –well plates and allowed to adhere for 24 hr at 37 °C/5% CO2, to reach a
confluence of ~80%. The cells were then treated with actin disrupters as described above, washed 3x with PBS and
fixed for 15 min at room temperature with 3% PFA in PBS. Cells were then washed 3x with PBS and permeabilised
with 0.2% Triton X-100 in PBS at room temperature for 5 min, washed 3x with PBS and stained for cell nuclei
with 1.0 µg/ml Hoechst 33342 in PBS and for polymerised actin with 1.0 µg/ml Rh-P in PBS for 15 min at room
temperature. The cells were finally washed 3x with PBS and once with distilled water, coverslips were mounted in
DAKO mounting medium on glass microscope slides and imaged by confocal microscopy19. Flow Cytometry. A431 cells were seeded onto 12-well plates and cultured for 24 hr to be ~80% confluent on
the day of experiment. Cells were washed twice with PBS and then incubated with diluent control (DMSO/dH2O)
or actin inhibitors as previously described. They were then loaded with serum-free DMEM containing either
2 µM EGFP-R8, 5 µg/ml TF-647 or 0.1 mg/ml Dex-647 in the presence or absence of actin inhibitors for times
stipulated in the figure legends. Materials and Methods The plates were then placed on ice to prevent further uptake and washed 3x with
either ice-cold PBS or 0.5 mg/ml heparin in PBS. To remove the plasma membrane bound TF, cells were washed
3x with ice-cold PBS followed by an incubation for 1 min in ice-cold acid wash (0.2 M acetic acid, 0.2 M NaCl,
pH 2.0)33. All cells were then washed twice with PBS at room temperature before trypsinisation at 37 °C for 5 min
before placing the detached cells in centrifuge tubes and centrifuging at 4 °C for 3 min at 800 × g. The obtained cell
suspension was washed 3x with ice-cold PBS and finally resuspended in 300 µl ice-cold PBS for measurement of
cell-associated fluorescence (cell gate 50,000 cells) by flow cytometry on a BD FACSVerse utilising a 488 laser and
527/32 filter for EGFP analysis and a 633 nm laser and 660/10 filter for Alexa 647. Cells were gated using FSC-A/
SSC-A to isolate single cell populations and analysed using FlowJo. Statistics. Flow cytometry measurements represent the mean fluorescence from three independent experi-
ments performed in duplicate, where each individual data point was the geometric mean. A two-tailed unpaired
Students t-test was used to test for differences within experiments. Significant differences were classed as having
a P value of < 0.05. Data Availability. The datasets generated during the current study are available from the corresponding
author on reasonable request. Results
ff 2), P = 0.0036.f lil
In these two cell lines, we also investigated the effects of Cyt D on the uptake of dextran, that enters cells via
fluid phase uptake, and transferrin that enters via clathrin mediated endocytosis. In agreement with our previous
flow cytometry and microscopy studies with 40 KDa FITC and Alexa488 dextran33, uptake of dextran-Alexa647
was dramatically increased in A431 cells (Supplementary Fig. 5A, C).lf Cell associated transferrin fluorescence was not markedly different between control and Cyt D treated HeLa
cells (Supplementary Fig. 5B) confirming previous reports performed at the same concentration in this cell line36. Transferrin fluorescence in Cyt D treated HeLa cells was, however, more prominent on the plasma membrane com-
pared to a strong perinuclear enrichment of fluorescence in control cells. Flow cytometry utilising acid washing of
surface label confirmed that there was no difference in fluorescence uptake of transferrin in A431 cells treated with
this drug (Supplementary Fig. 6), confirming previous studies in this cell line27. We focused our further investigations
on the A431 cell line and EGFP-R8, as this was the conjugate that gave unexpected Cyt D mediated uptake effects. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 3 www.nature.com/scientificreports/ Figure 1. Cytochalasin D effects on the cellular uptake of R8 conjugates in HeLa or A431 cells. HeLa (A,B) or
A431 (C,D) were preincubated with 10 µM Cyt D or diluent control for 15 min prior to incubation with 2 µM
R8 conjugates (R8-Alexa 488, A,C, or EGFP-R8, B,D) for 1 hr in the presence or absence of 10 µM Cyt D before
washing in heparin and analysis by confocal microscopy. Shown are single projection images of fluorescence
only (R8-Alexa 488/EGFP-R8), DIC and merges of fluorescence and DIC of the same cells. Scale bars 10 µm. Figure 1. Cytochalasin D effects on the cellular uptake of R8 conjugates in HeLa or A431 cells. HeLa (A,B) or
A431 (C,D) were preincubated with 10 µM Cyt D or diluent control for 15 min prior to incubation with 2 µM
R8 conjugates (R8-Alexa 488, A,C, or EGFP-R8, B,D) for 1 hr in the presence or absence of 10 µM Cyt D before
washing in heparin and analysis by confocal microscopy. Shown are single projection images of fluorescence
only (R8-Alexa 488/EGFP-R8), DIC and merges of fluorescence and DIC of the same cells. Scale bars 10 µm. Figure 2. Results
ff Cells on coverslips were treated either with
diluent control (0.1% DMSO) or 0.1–1.0 μM Lat B for 30 min before fixing and staining with Rh-P and Hoechst. Single sections (A) show the overall distribution of the actin relative to the nucleus. (B) Actin arrangement from
the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. To try and identify the subcellular compartments occupied by EGFP-R8 we performed pulse/chase experi-
ments with Dex-647: a 2 hr pulse labelled endolysosomal organelles was followed by a 4 hr chase to specifically
label late endosomes and lysosomes37. EGFP-R8 was then incubated with the cells for 1 hr in the presence or
absence of Cyt D. Control cells (Supplementary Fig. 8A, top row, indicated by arrows) show colocalisation of the
two fluorophores demonstrating trafficking of some EGFP-R8 to dextran labelled late endosomes and lysosomes. However, EGFP only structures were also clearly visible suggesting that this fraction had not yet entered these
structures or were trafficking to a different location. To verify that the EGFP fluorescence was being emitted from
inside the cells we captured images of the Cyt D treated cells through the Z axis starting 1.44 µm from the glass
surface to a height of 14.5 µm. This allowed us to visualise the dextran and EGFP-R8 structures at different focal
planes and the data confirmed that the EGFP fluorescence was of intracellular origin (Supplementary Fig. 8B). Effects of other actin reagents on actin and cellular uptake of EGFP-R8. Lat B, binds to and com-
plexes with actin monomers at a 1:1 molar ratio and prevents polymerisation38. We initially investigated the
concentration dependent effects of this drug on the actin cytoskeleton. At the lowest concentration (0.1 μM) the
drug caused the formation of small actin aggregates at both the basal and cell body and apex (CBA) sections of
the cells (Fig. 3). At 0.5 μM, these aggregates were extremely prominent especially at the basal section. At 1 μM the
drug caused major disruption of the actin organisation manifest as amorphous structures scattered throughout
the cytoplasm. The effects of this drug on cell morphology is further highlighted in Supplementary Fig. 4. yhf
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gy
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g
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g
We subsequently performed CPP uptake experiments in A431 cells pre-incubated with 0.5 µM Lat B prior to
addition of EGFP-R8. The effect of Lat B (Fig. Results
ff Cytochalasin D treatment increases the cellular uptake of EGFP-R8 in A431 cells. Cells were
preincubated with 10 µM Cyt D or diluent control for 15 min prior to incubation with 2 µM EGFP-R8 in
the absence or presence of 10 µM Cyt D for 1 hr. Cells were then washed, trypsinised and analysed by flow
cytometry. Data represent the geometric means ± S.D. from three independent experiments. Figure 2. Cytochalasin D treatment increases the cellular uptake of EGFP-R8 in A431 cells. Cells were
preincubated with 10 µM Cyt D or diluent control for 15 min prior to incubation with 2 µM EGFP-R8 in
the absence or presence of 10 µM Cyt D for 1 hr. Cells were then washed, trypsinised and analysed by flow
cytometry. Data represent the geometric means ± S.D. from three independent experiments. To identify whether the enhanced cellular uptake of EGFP-R8 in A431 cells by Cyt D treatment was an energy
dependant process, the same experiments were performed with cells pre-incubated at 37 °C with the drug, then
placed on ice and maintained at 4 °C throughout the subsequent incubation with EGFP-R8. There was very little
evidence above background fluorescence that EGFP-R8 entered cells at 4 °C in the presence or absence of Cyt D
(Supplementary Fig. 7) confirming that the enhanced EGFP-R8 internalisation induced by the inhibitor was
energy dependent. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 4 www.nature.com/scientificreports/ Figure 3. Effects of Lat B on the actin architecture in A431 cells. Cells on coverslips were treated either with
diluent control (0.1% DMSO) or 0.1–1.0 μM Lat B for 30 min before fixing and staining with Rh-P and Hoechst. Single sections (A) show the overall distribution of the actin relative to the nucleus. (B) Actin arrangement from
the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. Figure 3. Effects of Lat B on the actin architecture in A431 cells. Cells on coverslips were treated either with
diluent control (0.1% DMSO) or 0.1–1.0 μM Lat B for 30 min before fixing and staining with Rh-P and Hoec
Single sections (A) show the overall distribution of the actin relative to the nucleus (B) Actin arrangement fr Figure 3. Effects of Lat B on the actin architecture in A431 cells. Results
ff At 2 µM JAS the actin staining appeared in large clumps,
previously identified as aggresomes43, or was absent, and was completely lost from all cells treated at 4 µM (Fig. 5A). This loss of fluorescence may be due to the fact that this drug has been shown to be a competitive inhibitor of
phalloidin-actin binding39. Supplementary Fig. 4 shows the significant morphological effects of this drug on the cells. f
l
d
k
d
ll (
)
d h d
d cells, no visible changes in actin organisation were observed in both basal and CBA sections of cells until the con-
centration of JAS reached 0.2 µM (data not shown and Fig. 5). At this concentration, actin staining was more prom-
inent on the cell periphery at both basal and CBA regions. At 2 µM JAS the actin staining appeared in large clumps,
previously identified as aggresomes43, or was absent, and was completely lost from all cells treated at 4 µM (Fig. 5A). This loss of fluorescence may be due to the fact that this drug has been shown to be a competitive inhibitor of
phalloidin-actin binding39. Supplementary Fig. 4 shows the significant morphological effects of this drug on the cells. Confocal microscopy dramatic increase in EGFP-R8 uptake in JAS treated cells (Fig. 6A) and the drug caused
a pronounced clustering of fluorescence in the cytoplasm. Equivalent experiments analysed by flow cytom-
etry (Fig. 6B) indicated that JAS induced a four-fold increase in fluorescence that was statistically significant
(P = 0.0025). The diluent control experiments with JAS required adding DMSO to the cells at a final concentration
of 0.4% v/v; the highest used in this study. This concentration did not affect uptake of EGFP-R8 or R8-Alexa488
(Supplementary Fig. 9). p
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pp
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gi
p
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Confocal microscopy dramatic increase in EGFP-R8 uptake in JAS treated cells (Fig. 6A) and the drug caused
a pronounced clustering of fluorescence in the cytoplasm. Equivalent experiments analysed by flow cytom-
etry (Fig. 6B) indicated that JAS induced a four-fold increase in fluorescence that was statistically significant
(P = 0.0025). The diluent control experiments with JAS required adding DMSO to the cells at a final concentration
of 0.4% v/v; the highest used in this study. This concentration did not affect uptake of EGFP-R8 or R8-Alexa488
(Supplementary Fig. 9). Results
ff 4) was comparable to that of Cyt D (Fig. 2): the inhibitor treated
cells showing much higher fluorescence compared to controls (Fig. 4). Flow cytometry analysis confirmed these
findings and the 6.23 fold increase in uptake was highly statistically significant (P = 0.0045).i i
g
p
g y
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gi
(
)
An alternative actin inhibitor, Jasplakinolide (JAS) binds directly to filamentous actin and unlike Cyt D and Lat
B induces stabilisation of the existing actin filaments thus inhibiting actin disassembly39–42. Compared with control Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 5 www.nature.com/scientificreports/ Figure 4. Effects of Lat B on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with either
with diluent control or 0.5 μM Lat B for 30 min prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence
(control) or presence of 0.5 µM Lat B and washed in heparin. (A) Cell associated fluorescence was analysed
using confocal microscopy and shown are single projection images of fluorescence only (top rows) and merges
of fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells treated as in A were washed, trypsinised
and analysed by flow cytometry. Data represent the mean of geometric means ± S.D. from three independent
experiments. Figure 4. Effects of Lat B on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with either
with diluent control or 0.5 μM Lat B for 30 min prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence
(control) or presence of 0.5 µM Lat B and washed in heparin. (A) Cell associated fluorescence was analysed
using confocal microscopy and shown are single projection images of fluorescence only (top rows) and merges
of fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells treated as in A were washed, trypsinised
and analysed by flow cytometry. Data represent the mean of geometric means ± S.D. from three independent
experiments. cells, no visible changes in actin organisation were observed in both basal and CBA sections of cells until the con-
centration of JAS reached 0.2 µM (data not shown and Fig. 5). At this concentration, actin staining was more prom-
inent on the cell periphery at both basal and CBA regions. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 Discussion Since the discovery of CPPs, considerable efforts have been made to elucidate their internalisation mechanisms. Focusing mainly on arginine rich variants, macropinocytosis has been proposed to be a major contributing path-
way for delivery of low molecular weight fluorophores or larger cargo by these peptides20,24,49. This is based on
studies utilising chemical macropinocytosis inhibitors including, and almost exclusively Cyt D for actin inhi-
bition. It should however be noted that the involvement of actin activity has been shown for other endocytic
pathways such as clathrin-mediated endocytosis (CME)27,50. As shown here and in several other studies, Cyt D
at concentrations >5 µM, completely destroys actin organisation and, unsurprisingly, cell morphology is also
compromised. A number of other natural compounds and synthetic products have been identified or developed
to target the actin cytoskeleton directly or indirectly and to disrupt its arrangement/function. These include Lat B,
JAS and Y27632 and these inhibitors have not been used to support Cyt D data for drug delivery endocytosis
studies, including CPPs. To our knowledge this is the first study that has analysed CPP uptake with actin inhibi-
tors employing different mechanisms of action.l p y
gf
By employing high content analysis of the actin architecture using confocal microscopy and then flow cytom-
etry we were able to correlate actin disorganisation with cellular uptake of EGFP-R8 as a model protein-CPP con-
jugate. EGFP has been shown to be delivered into cells by R8 and R935,51 but correlative information regarding the
uptake mechanism of the proteins relative to that of the peptides attached to fluorophores is lacking. EGFP-Tat
uptake has been proposed to be mediated via macropinocytosis and caveolae24,52. Both Tat peptide, octaarginine,
and shorter number of arginine residues (R7, R9) have also been shown to deliver proteins into cells to mediate
biological effects53–55. gf
In this study, HeLa cells reacted in a somewhat predictable manner in response to actin breakdown with Cyt D
showing a reduction in uptake of R8-Alexa488 and dextran. In this cell line EGFP-R8 uptake was also inhibited by
Cyt D treatment suggesting a similar mechanism of uptake. We have previously shown that cortical actin in A431
cells is particularly prominent and suggested that this may have a role in the uptake of dextran that, under normal
conditions, enters cells via constitutive pinocytosis; also termed fluid phase endocytosis. This can be differentiated
from stimulus induced macropinocytosis56,57. Results
ff Actin dynamics and regulation are achieved through the concerted action of actin regulatory proteins whose
activities are finely modulated by protein kinases. These include Rho-associated coiled-coil forming serine/thre-
onine kinases ROCK I and II that have been identified as important downstream effectors of RhoA through
their interaction with its GTP-bound form44,45. A number of chemical inhibitors of the Rho-ROCK signalling
cascade have been developed, including compound Y27632 [(+)-(R)-trans-4-(1-aminoethyl)-N-(4-pyridyl)
cyclohexanecarboxamide dihydrochloride]. This has been shown to inhibit the kinase activity of a number of
ROCKs by binding to their catalytic sites46,47. Initially, 1 and 10 μM concentrations of Y27632 were investigated
on actin localisation based on published observations showing actin deformities at these concentrations46,48. Supplementary Fig. 10 demonstrates that no visible changes in the organisation of the actin cytoskeleton were
detected up to 10 μM drug. Higher concentrations caused extensive actin deformities including the formation of
a large number of actin needles, the majority of which ran perpendicular to the plasma membrane (Fig. 7). Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 6 www.nature.com/scientificreports/ Figure 5. Effects of JAS on the actin architecture in A431 cells. Cells on coverslips were treated either with
diluent control (0.1% DMSO) or 0.2, 2.0 or 4.0 μM JAS for 45 min before fixing and staining with Rh-P and
Hoechst. (A) Single sections show the overall distribution of the actin relative to the nucleus. (B) Actin
arrangement from the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. Figure 5. Effects of JAS on the actin architecture in A431 cells. Cells on coverslips were treated either with
diluent control (0.1% DMSO) or 0.2, 2.0 or 4.0 μM JAS for 45 min before fixing and staining with Rh-P and
Hoechst. (A) Single sections show the overall distribution of the actin relative to the nucleus. (B) Actin
arrangement from the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. In CPP uptake experiments, Y27632 treated cells displayed scattered EGFP-R8 fluorescence in small vesicles
in contrast to the vesicle aggregates seen in cells incubated with the other actin disrupting agents (Figs 1,4 and 8A)
while DIC microscopy did not show the gross morphological effects caused by the other actin inhibitors. Flow
cytometry analysis indicated that cell uptake of EGFP-R8 uptake was significantly inhibited by Y27632 but only
by ~20% (Fig. 8B P = 0.02). Results
ff To exclude the possibility that any of the actin inhibitors was affecting plasma mem-
brane permeability, we incubated A431 cells with the inhibitors at the concentrations and incubations used above
in the presence of live cell impermeable dye - DRAQ7. Using Tritin X-100 as positive control, none of the inhibi-
tors showed any evidence of inducing plasma membrane permeabilisation (Supplementary Fig. 4). Discussion This potential for the involvement of cortical actin was identified by
discovering a rather unexpected and highly significant increase in the uptake of 10 and 40 KDa dextran in Cyt D
treated A431 cells that was not observed with Alexa conjugates of transferrin, Tat and R833. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 7 www.nature.com/scientificreports/ Figure 6. Effects of JAS on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with diluent
control or 2–4 µM JAS for 45 min prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence (control) or
presence of 2–4 µM JAS and washed thoroughly using heparin. (A) Cell associated fluorescence was analysed
using confocal microscopy and shown are single projection images of fluorescence only (top row) and merges
of fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells treated as in A with 4 µM JAS were then
washed, collected and analysed by flow cytometry. Data represent the mean of geometric means ± S.D. from
three independent experiments. Figure 6. Effects of JAS on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with diluent
control or 2–4 µM JAS for 45 min prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence (control) or
presence of 2–4 µM JAS and washed thoroughly using heparin. (A) Cell associated fluorescence was analysed
using confocal microscopy and shown are single projection images of fluorescence only (top row) and merges
of fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells treated as in A with 4 µM JAS were then
washed, collected and analysed by flow cytometry. Data represent the mean of geometric means ± S.D. from
three independent experiments. Figure 6. Effects of JAS on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with dilue In this study in we show that dextran and also EGFP-R8 uptake is significantly increased in A431 treated with
Cyt D and contrasts to that observed for R8-Alexa488 (inhibition) and transferrin (no change). This strongly
suggests that the process by which these two R8 conjugates interact with and/or enter cells is different. It should
be noted that the structure of both conjugates differ in their cargo-CPP orientation and also the linkers utilised
between them. Discussion We consider that it is highly unlikely that these factors mediate the differences in endocytic pro-
files of these conjugates. Due to the similarities we see in entry for dextran and EGFP-R8, we hypothesise that
the differences are driven by the cargo and not the orientation of the CPP. But this warrants further study, noting
that EGFP alone does not enter these cells at the concentrations used here, demonstrating a clear role for R8 that
may be interacting differently with cells when attached to a fluorophore or protein. In none of the experiments
performed did we observe clearly defined cytosolic EGFP fluorescence in Cyt D treated cells demonstrating that
the effects of this drug maintained the localisation of the protein within a membrane bound compartment.fi f
g
p
p
We have previously shown that R8 and Tat peptides as Alexa conjugates traffic rapidly by endocytosis to
reach lysosomes30 and by performing pulse chase experiments we now show that the same is true for EGFP-R8,
demonstrating that it is trafficked to the same degradative compartments. From these combined studies we also
highlight that traffic to lysosomes of R8 attached to different cargo is also entirely dependent on functional actin.i reach lysosomes30 and by performing pulse chase experiments we now show that the same is true for EGFP-R8,
demonstrating that it is trafficked to the same degradative compartments. From these combined studies we also
highlight that traffic to lysosomes of R8 attached to different cargo is also entirely dependent on functional actin. Our findings prompted us to further study EGFP-R8 uptake in cells treated with inhibitors of actin that oper-
ate via different mechanisms and affect actin dynamics in different ways. Like Cyt D, at effective actin disrupting
concentrations, JAS and Lat B also affected cell morphology and increased EGFP-R8 uptake that was again highly
enriched in large structures in particular regions of the cells. Previous studies in non-polarised epithelial MDCK
cells demonstrated that JAS treated cells had much higher levels of internalised and retained dextran compared fif
Our findings prompted us to further study EGFP-R8 uptake in cells treated with inhibitors of actin that oper-
ate via different mechanisms and affect actin dynamics in different ways. Like Cyt D, at effective actin disrupting
concentrations, JAS and Lat B also affected cell morphology and increased EGFP-R8 uptake that was again highly
enriched in large structures in particular regions of the cells. Discussion Previous studies in non-polarised epithelial MDCK
cells demonstrated that JAS treated cells had much higher levels of internalised and retained dextran compared Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 8 www.nature.com/scientificreports/ Figure 7. Effects of Y27632 on the actin architecture of A431 cells. Cells on coverslips were treated either with
diluent control (D-MEM) or 50, 100 μM of Y27632 for 4 hr before fixing and staining with Rh-P and Hoechst. (A) Single sections show the overall distribution of the actin relative to the nucleus. (B) Actin arrangement from
the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. Figure 7. Effects of Y27632 on the actin architecture of A431 cells. Cells on coverslips were treated either with
diluent control (D-MEM) or 50, 100 μM of Y27632 for 4 hr before fixing and staining with Rh-P and Hoechst. (A) Single sections show the overall distribution of the actin relative to the nucleus. (B) Actin arrangement from
the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. Figure 7. Effects of Y27632 on the actin architecture of A431 cells. Cells on coverslips were treated either with
diluent control (D-MEM) or 50, 100 μM of Y27632 for 4 hr before fixing and staining with Rh-P and Hoechst. (A) Single sections show the overall distribution of the actin relative to the nucleus. (B) Actin arrangement from
the basal, CBA regions and composite (Merge) of basal and CBA. Scale bars 10 μm. with control cells57. In polarised cells, however, this effect was limited to dextran entering via the basolateral
surface and the retained dextran was observed in large structures as we observe for EGFP-R8. It remains to be
determined whether EGFP-R8 would also behave in a similar manner when introduced to different surfaces of
polarised cells and whether this is influenced by any differences in cortical tension between these surfaces. The
nature of these large structures is unknown but may be formed by tubulation of the plasma membrane that has
been shown to be caused by Cyt D58. The plasma membrane tension of mammalian cells is coordinated and main-
tained through the actin network, and this profoundly influences endocytosis59,60. The reduced cortical tension
resulting from actin disruption in MDCK basolateral surfaces57 and here in A431, cells may favour the formation
of invaginations and tubulation to allow entry of macromolecules. Discussion Cells were preincubated with
diluent control or 100 µM Y27632 for 4 hr prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence
(control) or presence of 100 µM Y27632 and washed with heparin. (A) Cell associated fluorescence was
analysed using confocal microscopy and shown are single projection images of fluorescence only and merge of
fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells were washed, trypsinised and analysed by
flow cytometry. Data represent the mean of geometric means ± S.D. from three independent experiments. Figure 8. Effects of Y27632 on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with
diluent control or 100 µM Y27632 for 4 hr prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence
(control) or presence of 100 µM Y27632 and washed with heparin. (A) Cell associated fluorescence was
analysed using confocal microscopy and shown are single projection images of fluorescence only and merge of
fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells were washed, trypsinised and analysed by
flow cytometry. Data represent the mean of geometric means ± S.D. from three independent experiments. Conclusions Interpretation of endocytosis data in in vitro models obtained using actin modifying agents is difficult as the net-
work that this protein nucleates is a regulator of so many processes; global cell morphology is often affected by its
perturbation. Here we highlight that disruption of actin in some cell types promotes cell uptake of specific mol-
ecules whilst having no effects, or even inhibitory effects on others. From careful scrutiny of the literature there
is precedent for this in other cell types and distinct plasma membrane regions in polarised cells. The possibility
exists that these cells have a specific but common cortical organisation of actin that could be exploited to drive
internalisation of drug delivery vectors carrying macromolecular therapeutics. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 Discussion This strongly supports studies showing an
increased uptake of 70 KDa dextran and the folic acid receptor in Cyt D treated cells61,62. Both folic acid and dex-
tran have been extensively utilised as, respectively, targeting and passive vectors for drug delivery25,63.h Through interaction with the Rho GTPase, ROCKs play a central role in actin network assembly and mem-
brane association64. In A431 cells the effects of Y27632 on the actin cytoskeleton was more subtle than that
observed with the agents that directly targeted actin. No change in cell morphology was observed in treated cells
but actin distribution was very different at high 100 µM concentrations. Effects of this agent in human trabecular
meshwork eye cell actin has been shown to be extremely prominent at much lower concentrations, showing a loss
of filamentous actin65. In contrast to results observed with direct actin inhibitors, we observed a slight but signifi-
cant decrease in the uptake of EGFP-R8 in cells treated with Y27632. To our knowledge no CPP studies have been
performed with this drug though it has previously been shown to affect the internalisation of plasma membrane
receptors and also to inhibit (~50%) the uptake of 70 KDa dextran in macrophages66–68. This again supports our
suggestion that dextran and EGFP-R8 uptake is responsive to the same inhibition in A431 cells but that they may
not necessarily enter via the same mechanism. This supports recent observations showing that cell uptake profiles
of HIV-TAT peptide and dextran can be separated69. Scientific REPorTS | (2018) 8:7318 | DOI:10.1038/s41598-018-25600-8 9 www.nature.com/scientificreports/ Figure 8. Effects of Y27632 on the cellular uptake of EGFP-R8 in A431 cells. Cells were preincubated with
diluent control or 100 µM Y27632 for 4 hr prior to incubation with 2 µM EGFP-R8 for 1 hr in the absence
(control) or presence of 100 µM Y27632 and washed with heparin. (A) Cell associated fluorescence was
analysed using confocal microscopy and shown are single projection images of fluorescence only and merge of
fluorescence and DIC of the same cells. Scale bars 10 µm. (B) Cells were washed, trypsinised and analysed by
flow cytometry. Data represent the mean of geometric means ± S.D. from three independent experiments. Figure 8. Effects of Y27632 on the cellular uptake of EGFP-R8 in A431 cells. References
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The study presented here is funded by Cardiff University (LH, PW, ATJ) and has received support from the
Innovative Medicines Initiative Joint Undertaking under grant agreement no. [115363], resources of which are
composed of financial contribution from the European Union’s Seventh Framework Programme (FP7/2007–
2013) and EFPIA companies’ in-kind contribution (EJS, PW, ATJ). We would like to acknowledge Thomas
Williams from School of Chemistry Cardiff University for his help with the mass spectrometry funded in part
by EPSRC (EP/L027240/1). We would also like to thank Prof. Shiro Futaki from the Division of Biochemistry,
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The effect of corticosteroid versus platelet-rich plasma injection therapies for the management of lateral epicondylitis: A systematic review
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1 The University of Salford, Salford, UK
2 y
,
,
ma and Orthopaedics department, Central Manchester University Hospitals NHS Foundation Trust, UK 2 Trauma and Orthopaedics department, Central Manchester University Hospitals NHS Fo Received 27 April 2017, Accepted 29 December 2017, Published online 21 March 2018 Abstract -- Introduction: Lateral epicondylitis is a common musculoskeletal disorder of the upper limb. Corticosteroid injection has been widely used as a major mode of treatment. However, better understanding of
the pathophysiology of the disease led to a major change in treating the disease, with new options including
platelet-rich plasma (PRP) are currently used. (
)
y
bjectives/research aim: To systematically evaluate the effect of corticosteroid versus PRP injections for the
eatment of LE. Hypothesis: PRP injections provide longer-term therapeutic effect and less rate of complications compared to
corticosteroid injection. j
Level of evidence: Level 2 evidence (4 included studies are of level 1 evidence, 1 study of level 2 evidence). Design: Systematic Review (according to PRISMA guidelines). Methods: Eleven databases used to search for relevant primary studies comparing the effects of corticosteroid
and PRP injections for the treatment of LE. Quality appraisal of studies performed using Cochrane Handbook
for Systematic Reviews of Interventions Version 5.1.0, CASP Randomised Controlled Trial Checklist, and
SIGN Methodology Checklist 2. Results: 732 papers were identified. Five randomised controlled trials (250 Patients) met the inclusion criteria. Clinical findings: Corticosteroid injections provided rapid symptomatic improvement with maximum effect at
6/8/8 weeks before symptoms recurrence, whereas PRP showed slower ongoing improvements up to 24/52/104
weeks (3 studies). Corticosteroid showed more rapid symptomatic improvement of symptoms compared to PRP
up to the study end-point of 3 months (1 study). Comparable therapeutic effects of corticosteroid and PRP were
observed at 6 weeks (1 study). Ultrasonographic Findings: (1) Doppler activity decreased more significantly in
patients who received corticosteroid compared to PRP. (2) Reduced tendon thickness and more patients with
cortical erosion noted in corticosteroid group whereas increased tendon thickness and less number of patients
with common extensor tendon tears noted in PRP group. (3) Fewer patients reported Probe-induced tenderness
and oedema in the common extensor tendon in both corticosteroid and PRP groups (2 studies). onclusion: Corticosteroid injections provide rapid therapeutic effect in the short-term with recurrence of
ymptoms afterwards, compared to the relatively slower but longer-term effect of platelet-rich plasma. Key words: Tennis elbow, Lateral epicondylitis, Epicondylopathy, Epicondylalgia Corticosteroid, Stero
injections, Platelet-rich plasma, PRP injections This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. *Corresponding author: W.O.Ben-Nafa@edu.salford.ac.uk SICOT-J 2018, 4, 11
© The Authors, published by EDP Sciences, 2018
https://doi.org/10.1051/sicotj/2017062 SICOT-J 2018, 4, 11
© The Authors, published by EDP Sciences, 2018
https://doi.org/10.1051/sicotj/2017062 Available online at:
www.sicot-j.org Available online at:
www.sicot-j.org REVIEW ARTICLE REVIEW ARTICLE The effect of corticosteroid versus platelet-rich plasma injection
therapies for the management of lateral epicondylitis: A systematic
review Walid Ben-Nafa1,2,* and Wendy Munro1 1 The University of Salford, Salford, UK
2 Methodology Study selection criteria Although the main focus
was on RCTs, all interventional studies were eligible for
inclusion in this study. The search strategy of this review
was structured using Population, Intervention, Compari-
son, Outcome framework (PICO). Free-text searching and
Medical Subject Headings were also considered. j
g
Search strategy The search strategy was conducted
and reported in accordance with PRISMA guidelines [37]
and Cochrane Collaboration [38] whenever possible. It
also adhered to the published guidance used when
undertaking reviews in healthcare [39]. Eleven databases
were systematically searched, namely, Web of science,
Scopus, PubMed (MedLine), ScienceDirect, CINAHL,
EMBASE, Ovid, NICE, Physiotherapy Evidence Data-
base, Cochrane Library and ClinicalTrials database
(Figure 1). As we aimed to search for only the recent
and relevant evidence, only papers published from 2005
onwards were accessed, and hand-searched for their
bibliographic references to look for any relevant refer-
ences. The search was also enhanced using forward
citation searching strategy, with searching by author and
study title was also considered using Web of Science
Service. Cochrane Database of Systematic Reviews,
Cochrane Central Register of Controlled Trials, and
database of abstracts of reviews of effectiveness were also
consulted. Potentially
relevant
studies,
which
are
labelled as ongoing studies, were searched for using
International Clinical Trials Registry Platform, UK
Clinical Research Network Study Portfolio, US National
Library of Medicine Database of Clinical Trials and
metaRegister of Controlled Trials. Database bias was
eliminated as possible, by reviewing journals which were
not indexed in major bibliographic databases. Finally,
potentially relevant studies in the grey literature was also
considered to identify any unpublished research or
research produced outside the traditional distribution
channels. With
recent
histological
findings
showing
that
tendinosis is not an acute inflammatory condition but
rather
a
failure
of
normal
tendon
repair
[18,19]
researchers started to focus on the function of biologics
in treating LE [20], including Platelet-rich Plasma (PRP)
injection [21]. Firstly introduced by Whitman, Berry, &
Green [22], PRP has a 3–5 fold increase of platelet
concentration compared to whole blood levels [23], and
includes many growth factors essential for bone-to-tendon
healing, as well as other vascular, epidermal and connective
tissue growth factors [24]. Different laboratory studies
showedenhanced tenocyteproliferationafterinjection with
platelet-released growth factors [25,26]. Introduction including pain and tenderness over the origin of extensor
muscles of the wrist and fingers [3,4]. With a prevalence
rate of more than 1% among the general population, and
with a slight predominance among females [5,6] the disease
mostly affects people aged between 35–50 years, who have
a history of repetitive activities involving the upper limb
[7,8]. While the exact pathophysiology behind the condi-
tion is not yet clear, and despite the presence of
inflammatory cells locally, there is a strong argument Tennis elbow or lateral epicondylitis (LE), is one of the
most common and painful musculo-skeletal conditions,
which has a significant impact on the healthcare industry
and society in general [1,2]. Lateral epicondylitis is a
common term used to describe a group of symptoms W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 2 that LE can be regarded as a degenerative process caused
by muscle overuse, with subsequent tendinosis, micro-
trauma and tear of the extensor carpi radialis brevis
tendon [9,10]. injections, leaving a gap in the literature, with inadequate
knowledge available to inform the relevant clinical
practices. In this systematic review, we aim to systemati-
cally accumulate the evidence available from the relevant
primary studies about the effect of these two treatments. We hypothesise that PRP injection provides longer-term
therapeutic effect and less rate of complications compared
to corticosteroid injection. [ ,
]
The relatively large number of options currently
available for
the treatment of lateral epicondylitis
[11,12], could be attributed to the scarce evidence
available about the disease aetiology and the lack of
agreement about a definitive treatment for the condition. Corticosteroid injection has been considered a major
treatment option for LE [9]. As corticosteroid mainly aims
to reduce inflammation, it is apparently questionable
whether it has any possible long-term healing potential
with the disease’s degenerative changes [10]. Besides the
known local and systemic side effects associated with its
use [11,13–16], some researchers believe that corticoster-
oid injections may even delay the natural recovery
expected with watchful waiting or other management
options [17], in addition to the high rate of symptoms
recurrence noted with its use. Methodology However, and with
the insufficient evidence available to support its clinical use
[27], PRP injections continues to be debated in the
literature, especially with the lack of its comparable rapid
therapeutic effect compared to other pharmacological
treatments [28,29]. Although a number of randomised controlled trials
compared the effect of corticosteroid and PRP injection
for treating LE, an up-to-date systematic review of these
primary studies is lacking within the literature. While
some reviews [17,30] assessed the effect of various injection
therapies for LE, none of these studies investigated the
effect of PRP injection as a comparative therapy for the
treatment of LE. On the other hand, and while a number of
systematic reviews [31–33] focused on the effect of PRP
injections, none of these compared the outcome of this
injection therapy with other injection modalities, includ-
ing corticosteroid injections. Some reviews and meta-
analyses recently included more primary studies about the
effect of PRP and steroid injections [34–36], but again,
none of these reviews focused or specifically compared the
effect of steroid and PRP injection therapies. This
attributed to the fact that these reviews were aiming at
collectively comparing the effects of different injection
therapies to obtain a general conclusion about their effect
in LE. As a result, this has hindered for a conclusion to be
drawn about the effects peculiar to steroid and PRP Only Papers published in English language were
included in this review, although it was aimed initially
to consider papers written in non-English languages too,
including two papers in Turkish language. Nevertheless,
and as the accuracy of quality assessment and data
extraction could be affected when translating non-English
papers, it was finally decided to restrict the work on papers
written in English language. This was also supported by
the evidence that language restriction does not appear to
change the results of systematic reviews conducted on
RCTs of conventional interventions [40]. 3 W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 7 study papers
considered
732 potentially relevant study papers identified by search strategy:
199 (Web of science)
84 (PubMed: Including MEDLINE)
109 (Scopus)
340 (Other databases - refer to the methodology section)
689 articles were excluded because
of the following reasons:
- Duplicate papers. - Review articles. - Conference papers,
periodicals, editorials, etc. Methodology 43 papers selected
36 study papers were excluded
(after reading the papers abstracts)
because of the following reasons:
- Protocol-only study papers
- No comparative assessment of
interventions
- Comparison of irrelevant
treatment interventions
2 study papers written
in non-English
language (Turkish)
5 primary studies included
in this systematic review
Figure 1. PRISMA Flow Chart showing the studies’ selection process which led to the inclusion of the five study. 732 potentially relevant study papers identified by search strategy:
199 (Web of science)
84 (PubMed: Including MEDLINE)
109 (Scopus)
340 (Other databases - refer to the methodology section) 7 study papers
considered 2 study papers written
in non-English
language (Turkish) 5 primary studies included
in this systematic review Figure 1. PRISMA Flow Chart showing the studies’ selection process which led to the inclusion of the five study. reviewers before embarking on the data extraction
process, to test the comprehensiveness and objectivity
of the extraction form. Some of the experts in the field pertinent to this review,
including some of the included studies’ authors, were
contacted to inquire about any ongoing or unpublished
relevant studies. No relevant studies were declared by the
authors. To objectively illustrate all the potential biases of the
studies in a comparative style, a table has been compiled
to
evaluate
the
included
studies
against
different
checklist elements used to assess potential biases in
RCTs [41–44]. The reviewers independently appraise
and eventually agreed on the quality of the included
studies (Table 1). The process of inclusion of relevant studies was
independently conducted by the two authors, and consen-
sus has been reached between the two reviewers with no
need for a third opinion. Exclusion criteria included old
studies conducted before 2005, study population with
elbow pain/pathologies other than LE, and injection
materials other than corticosteroid or PRP, including
autologus blood and autologous conditioned plasma
injections, which were excluded due to their irrelevancy. Results Eleven electronic databases were searched up to
February 2017 without language, publication, or study-
designs restrictions (Figure 1). Out of 732 papers identified
in the body of literature, 43 study papers were considered
after excluding duplicate papers, review articles, confer-
ence papers and other irrelevant articles. Five studies
[29,45–48], including 250 patients, were finally considered The data extraction process was performed before
assessing the quality of the studies, to enhance the
objectivity in collecting data regardless of the quality. The process of data extraction was carried out indepen-
dently by the two reviewers as well. A pilot data
extraction form [41] was created and tested by the two W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 4 Y: Yes; N: No; C: Consecutive sampling; U: Unclear; RCT: Randomised controlled trials; NS: Not stated; DD: Demographic details; DC: Disease characteristics; P: Population; I:
Intervention; O: Outcomes; OM: Outcome measures; DSL: Different study locations; HD: Hand dominance; SH: Smoking history; SD: Symptoms durations; PI: Previous
interventions; IPB: Inter-personal bias. Table 1. The result of quality assessment of the included studies. Study
authors
Internal validity Items
External
validity items
Summary of biases
Study design
Were
groups
similar at
the start
of the
trial? SelecƟon bias
Performance
bias
Blinding of outcome
assessors? Co-intervenƟon bias
Incomplete outcome data
Likelihood of aƩriƟon bias? + (Drop-out raƟo)
SelecƟve reporƟng
(ReporƟng bias)
ReporƟng details are clear
Groups analysed according to
their original allocaƟon
(intenƟon-to-treat analysis)? Other sources of bias
Confounding factors
Sampling method
Are ParƟcipants
representaƟve of the wider
populaƟon? DD
DC
Random
sequence
generaƟon
AllocaƟon
concealmen
Blinding of
parƟcipants
Blinding of
treatment
providers
Peerbooms
et al. (2010)
RCT
Y
N
Y
Y
Y
U
U
Y
N
N
(6:100)
N
P
Y
Y/N
- PI
- IPB
DSL
C
Y
- Different demographic
variables. - Possible performance,
inter-personal and co-
intervenƟon biases. I
Y
O
Y
Gosens et
al. (2011)
RCT
Y
N
Y
Y
Y
U
U
Y
N
N
(6:100)
N
P
Y
Y/N
- PI
- IPB
DSL
C
Y
As above
I
Y
I
Y
Krogh et al. (2013)
RCT
N
N
Y
Y
Y
N
Y
Y
Y
Y
(44:60)
at 1
year
Y
P
Y
Y/N
- PI
- HD
- SH
- SD
C
/U
Y
- Different demographic
variables. - Possible performance,
aƩriƟon, co-intervenƟon
and reporƟng biases. - confounding factors. I
Y
O
Y
Gautam et
al. Results (2015)
NS/
RCT
U
U
U
U
U
U
U +
Y
Y
N
U
N
P
N
U
- PI
U
U
Possible selecƟon,
performance, co-
intervenƟon and
sampling biases. I
Y
O
Y
Omar et al. (2012)
RCT
U
Y/U
U
U
U
U
U
Y
N
U
N
P
Y / U
U
- PI
U
Y/U
As above
I
Y
O
Y
Y: Yes; N: No; C: Consecutive sampling; U: Unclear; RCT: Randomised controlled trials; NS: Not stated; DD: Demographic details; DC: Disease characteristics; P: Population; I
Intervention; O: Outcomes; OM: Outcome measures; DSL: Different study locations; HD: Hand dominance; SH: Smoking history; SD: Symptoms durations; PI: Previou
interventions; IPB: Inter-personal bias. Internal validity Items W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 5 Study
Peerbooms et al. (2010)
Gosen et al. (2011)
Krogh et al. (2013)
Gautam et al. (2015)
Omar et al. (2012)
Study Ɵtle
PosiƟve Effect of an Autologous Platelet
Concentrate in Lateral EpicondyliƟs in a
Double-Blind Randomized Controlled Trial:
Platelet-Rich Plasma Versus CorƟcosteroid
InjecƟon With a 1-Year Follow-up. Ongoing PosiƟve Effect of Platelet-Rich
Plasma Versus CorƟcosteroid InjecƟon in
Lateral EpicondyliƟs: A Double-Blind
Randomized Controlled Trial With 2-year
Follow-up. Treatment of Lateral EpicondyliƟs With Platelet-Rich Plasma,
GlucocorƟcoid, or Saline: A Randomized, Double-
Blind, Placebo-Controlled Trial. Platelet-rich plasma versus
corƟcosteroid injecƟon for
recalcitrant lateral epicondyliƟs:
clinical and ultrasonographic
evaluaƟon. Local injecƟon of autologous
platelet rich plasma and
corƟcosteroid in treatment of
lateral epicondyliƟs and
plantar fasciiƟs: Randomized
clinical trial. When trial
conducted? May 2006 To January 2008
May 2006 - January 2008
January 2009 To July 2010
May 2011 To October 2012
October 2009 – May 2010
Study design
RCT
RCT
RCT
Not menƟoned. (Most probably
an RCT, as per study details)
RCT
Conflict of
interest
No
PotenƟal conflict of interest :
One or more of the authors declared that
Biomet has sponsored the study and
supplied the Recover system used at a
discounted rate. However, Biomet did not
have any influence on the collecƟon and
analysis of the data from this study. PotenƟal conflict of interest :
One or more of the authors declared that:
1) The Danish RheumaƟsm AssociaƟon has supported the primary
invesƟgator (T.P.K.) with a 6-month grant. 2) The Musculoskeletal
StaƟsƟcs Unit at the Parker InsƟtute is sponsored by the Oak
FoundaƟon. 3) Biomet Biologics Inc. has provided the Recover
GPS II Platelet Concentrate SeparaƟon Kit and donated an
unrestricted grant to the Region Hospital Silkeborg in Denmark. No
No
Funding
(eg, commercial
research suppor
The study was sponsored by Biomet, Dordrecht, The Netherlands. (There was no involvement in the trial phases by the sponsor). - The Danish RheumaƟsm AssociaƟon
- Oak FoundaƟon
- Biomet Biologics Inc. None stated
Not menƟoned
IntervenƟons
used
- PRP(1) injecƟons
- CS(2) injecƟons
- corƟcosteroid injecƟons
- PRP injecƟons
- Saline
- corƟcosteroid injecƟons
- PRP injecƟons
- corƟcosteroid injecƟons
- PRP injecƟons
Post-
intervenƟon
treatments
- Acetaminophen only if necessary. - Standardized stretching protocol to follow for 2 weeks under the supervision of a
physiotherapist. - A formal eccentric muscle- and tendon-strengthening program was iniƟated aŌer this
stretching protocol. Internal validity Items - Acetaminophen when necessary. - Ice packs or Paracetamol rather
than non-steroidal anƟ-
inflammatory drugs. - Acetaminophen only for pain,
but not to use non-steroidal
anƟ-inflammatory medicaƟon. any re-
intervenƟons or
other treatments
given to paƟents
aŌer the trial
- Yes (18 parƟcipants received
re-intervenƟons):
- IntervenƟons in CS group: 6 operaƟons, 1
re-injecƟon CS, 6 injecƟons with PRP. - IntervenƟons in PRP group: 3 operaƟons, 2
injecƟons with CS. (P: 0.970)
Yes (20 parƟcipants received re-
intervenƟons). IntervenƟons in CS group: 6 operaƟons, 1 re-
injecƟon CS, 7 injecƟons with PRP. IntervenƟons in PRP group: 3 operaƟons, 3
injecƟons with CS. Yes (42 paƟents received further treatments). 26 paƟents chose PRP, 12 paƟents received CS, 2 had surgical
treatment and 2 received sclerosing injecƟons. (Unclear what their originaƟng group was). Not menƟoned
Not menƟoned
Was the study
completed (All
stages)
Yes
Yes
No (Priori calculaƟons were originally made on an anƟcipated 12-
month results, which weren’t establish because of high level of
paƟents' drop-out at 12-month follow-up). Study was ended at 3-
month follow-up. Yes
Yes
Drop-out rate
among
parƟcipants
- CS group: 1 patient. - PRP group: 3 patients lost to follow-up
temporarily. (P: 0.700)
- 3 patient from each group lost to follow-
up temporarily. 44/60 dropped out & chose alternative treatments. Not menƟoned
Not menƟoned W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 6 Table 3. Population characteristics of the review studies. Study
Number of
parƟcipants
Mean age of parƟcipants
(± SD) (1)
ParƟcipants’ Gender raƟo (M:F)
Ethnic
groups of
parƟcipants
Clinical
problem
(disease)
Disease
duraƟon
Co-
morbidiƟes
in
Previous
treatments
received
CS (2)
PRP (3)
Saline
CS
PRP
Saline
Peerbooms
et al. (2010)
100
47.3
(±7.6)
46.9
(±8.4)
IR (4)
25:26
23:26
IR
NM (5)
LE (6)
≥ 6 M
(7)
NM
- Cast immobilizaƟon
- CS InjecƟons
- Physiotherapy
(All were received ≥ 6 M before study
intervenƟons). (P: 0.797)
(P: 0.840)
Gosen et al. (2011)
100
47.3
(±7.8)
46.8
(±8.5)
IR
23: 26
23: 28
IR
NM
LE
≥ 6 M
NM
The same as above. (P: 0.780)
(P: 0.712)
Krogh et al. (2013)
60
(Only 40
trialled for
PRP Vs CS)
43.9 (+
8.7)
47.6
(+ 7.1)
44.7
(+ 7.9)
11:9
9:11
9:11
NM
LE
23.1 M
(±38.)
NM
- CS for 35 out of 60 paƟents
( Given ≥ 3M before the intervenƟon)
Gautam et
al. Internal validity Items (2015)
30
NM
NM
IR
NM
NM
IR
NM
LE
≥ 6 M
NM
- Oral medicaƟons. - Non-invasive treatments: (not menƟoned what
they were specifically). paƟents’ ages were between (18-
60)
Omar et al. (2012)
60
(only 30
trialled for
LE)
37.5 (±17.5)
40.5 (±
15.5)
IR
5:10
6:9
IR
NM
LE
NM
NM
No treatments were given at least 4 weeks
before the intervenƟon. (1) SD: Standard deviation. (2) CS: corticosteroid. (3) PRP: platelet-rich plasma. (4) IR: Irrelevant. (5) NM: Not mentioned. (6) LE: Lateral Epicondylitis. (7) M: Month. (P: P W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 7 7 Table 4. Details of steroid and PRP preparations used by the included studies. Steroid preparaƟons
Study
Type of corƟcosteroid
used
Dose and
concentraƟon
Type, dose & concentraƟon of
local anaestheƟc used
Peerbooms et al. (2010)
Triamcinolone
acetonide
1 ml (40 mg/ml)
Bupivacaine hydrochloride 0.5%
+ Epinephrine (1:200000)
Gosens et al. (2011)
Triamcinolone
acetonide
1 ml (40 mg/ml)
Bupivacaine hydrochloride 0.5%
+ Epinephrine (1:200000)
Krogh et al. (2013)
Triamcinolone
1 ml (40 mg/ml)
2 ml lidocaine 10 mg/mL
Gautam et al. (2015)
Methylprednisolone
2 ml (40mg/ml)
NIG (1)
Omar et al. (2012)
NIG
NIG
NIG
PRP preparaƟons
Study
volume
Platelets
concentraƟon
(compared to platelets
concentraƟon in blood)
DuraƟon
between blood
collecƟon and
PRP injecƟon
Exogenous
platelets
acƟvaƟon
Buffering agent
Peerbooms
et al. (2010)
3 ml
NIG
Around 30
minutes
No
8.4% sodium
bicarbonate +
bupivacaine
hydrochloride 0.5%
+ epinephrine
(1:200000)
Gosens et
al. (2011)
3 ml
NIG
Around 30
minutes
No
As above
Krogh et
al. (2013)
3- 3.5
ml
8 folds
Injected directly
aŌer preparaƟon
NIG
8.4% sodium
bicarbonate
Gautam et
al. (2015)
2 ml
NIG
NIG
NIG
NIG
Omar et al. (2012)
NIG
At least 2 folds
NIG
NIG
NIG Table 4. Details of steroid and PRP preparations used by the included studies. (1) NIG: No information given has slower but longer-term clinical effect with no
recurrence of symptoms over the follow-up periods
included. This was almost the opposite with the use of
corticosteroid injections. in this systematic review After reading full-text papers. The quality appraisal of the included studies is summar-
ised in Table 1. In addition, Studies’ characteristics,
demographic details and details of treatment preparations
have also been listed (Tables 2–4). (1) NIG: No information given Internal validity Items Safety of interventional materials used Besides
some of the known local adverse effect [49,50], no other
complications
or
systematic
effects
were
reported
(Table 8). Except for one study [29], The lack of details
regarding the rate/number of complications incidence (
)
Analytic data of the included studies With
careful observation of the studies’ outcome results
(Tables 5–7), it can be concluded that there was a
general trend by most of the studies showing that PRP W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 8 Table 5. Values for Visual analogue scale (VAS) & Disabilities of the Arm, Shoulder and Hand (DASH) as reported by the included
studies. (1) SD: Standard deviation; (2) CS: Corticosteroid; (3) PRP: Platelet-rich plasma. (P: P value). Visual analogue scale (VAS)
Follow-up intervals
study name &
intervenƟons used
Baseline
Value
±SD(1)
2 Weeks
1 month
(4weeks)
6 weeks
8 weeks
3 months
(12
weeks)
6 months
(26
Weeks)
52 weeks
104
weeks
Scale
used
Peerbooms
et al. (2010)
CS(2)
65.8 ±13.8
44.2 ±26.4
42.9 ±29.2
44.2 ±27.1
56.6 ±23.2
50.1 ±28.1
(0 - 100)
PRP(3)
70.1 ±15.1
55.4 ±24.2
46.9 ±24.9
38.7 ±27.2
32.6 ±31.5
25.3 ±31.2
Gosen et al. (2011)
(Data
according to
intenƟon-to-
treat
analyses)
CS
66.2
±
14.0
(P: 0.340)
44.3
±
26.3
( P: 0.023)
43.4
±
28.9
( P: 0.411)
45.5
±
27.1
( P: 0.319)
55.8
±
24.1
( P: < 0.001)
48.8
±27.0
( P: < 0.001)
42.4
±26.8
( P: < 0.001)
(0 - 100)
PRP
69.0
±
15.9
55.7
±
24.1
47.7
±
25.0
40.2
±
27.5
32.9
±
30.8
25.9
±
30.6
21.3
±28.1
Krogh et al. (2013)
CS
N o t a p p l i c a b l e
PRP
Gautam et al. (2015)
CS
7.0
±0.8
(P: 0.650)
2.1
±0.7
(P: 0.000)
1.4
±0.5
(P: 0.000)
1.7
±0.5
(P: 0.493)
2.9
±1.2
(P: 0.001)
(0 - 10)
PRP
7.1
±0.8
4.5
±1.1
2.7
±0.8
1.8
±0.6
1.6
±0.5
Omar et al. (2012)
CS
8.6 ±1.6
4.3 ±2.1
(P:<0.001)
(0 - 10)
PRP
8.0 ±1.4
3.8 ±1.9
(P:<0.001)
DisabiliƟes of the Arm, Shoulder, and Hand QuesƟonnaire (DASH)
Follow-up Intervals
Study
name and
interventions used
Baseline
±SD
2
Weeks
1 month
(4
weeks)
6 weeks
8 weeks
3 months
(12
weeks)
6 months
(26
Weeks)
52 weeks
104
weeks
Peerbooms
et al. (1) SE: Standard error. (2) CS: Corticosteroid. (3) PRP: Platelet-rich plasma. (2) CS: Corticosteroid. (3) PRP: Platelet-rich plasma. Internal validity Items (2010)
CS
131.2 ±
58.2
97.4 ± 69.0
84.7 ± 73.4
92.2 ± 68.7
117.3 ± 75.6
108.4 ± 82.2
PRP
161.3 ±
62.4
135.9 ±
78.0
113.4 ± 79.6
92.0 ± 78.8
79.5 ± 80.3
54.7 ± 73.2
Gosen et al. (2011)
(Data according
to intenƟon-to-
treat analysis)
CS
43.3
± 16.1
( P: 0.002)
31.2
±
20.8
( P: 0.005)
28.3 ±
22.2
(P: 0.060)
32.3 ±
21.7
(P: 0.813)
37.6 ±
23.1
(P: 0.037)
36.8
±
24.0
(P: < 0.001)
36.5 ±
23.8
(P: < 0.001)
PRP
54.3 ±
19.5
43.1
±
21.6
37.2 ±
24.7
21.3 ±
22.0
27.8 ±
24.7
20.0
±
23.5
17.6 ±
24.0
Krogh et al. (2013)
CS
PRP
Gautam et al. (2015)
CS
67.5
±6.9
( P: 0.378)
39.7
±6.7
(P: 0.000)
32.7
±4.1
(P: 0.003)
34.3
±3.3
(P: 0.675)
39.6
±1.0
(P: 0.012)
PRP
69.7
±6.1
51.6
±6.8
38.6
±5.7
33.6
±5.1
32.0
±4.5
Omar et al. (2012)
CS
57.3 ±10.3
20.2
±14.0
(P<0.001)
PRP
58.9 ±10.5
19.9
±12.9
(P<0.001)
1) SE: Standard error. (2) CS: Corticosteroid. (3) PRP: Platelet-rich plasma. 8
W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 Table 5. Values for Visual analogue scale (VAS) & Disabilities of the Arm, Shoulder and Hand (DASH) as reported by the included
studies. (1) SD: Standard deviation; (2) CS: Corticosteroid; (3) PRP: Platelet-rich plasma. (P: P value). Table 5. Values for Visual analogue scale (VAS) & Disabilities of the Arm, Shoulder and Hand (DASH) as r
studies. (1) SD: Standard deviation; (2) CS: Corticosteroid; (3) PRP: Platelet-rich plasma. (P: P value). W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 9 Table 6. Outcome values for Patient-Rated Tennis Elbow Evaluation (PRTEE), Oxford elbow score, Modified Mayo score and Hand
grip strength at different follow-up intervals. es for Patient-Rated Tennis Elbow Evaluation (PRTEE), Oxford elbow score, Modified Mayo score and Hand
t follow-up intervals. Table 6. Outcome values for Patient-Rated Tennis Elbow Evaluation (PRTEE), Oxford elbow score, Modifi
grip strength at different follow-up intervals. Outcome measure used
Patient-Rated Tennis Elbow Evaluation (PRTEE) Krogh et al. tandard error. (2) CS: Corticosteroid. (3) PRP: Platelet-rich plasma. (4) SD: Standard deviation. (P:P value Internal validity Items (2015)
Corticosteroid
injections
Platelet-
rich
plasma
Corticosteroid
injections
Total:15 Patients
Platelet-rich
plasma
Total:15 Patients
DA (1)
(Mean Value) ±SD
(α)
0 (A)
(3.2) ± 0.9
(3.8) ± 0.4
Δ DA (2)
(Mean Value) SE (β)
3 M (B)
(- 3.0) 0.2
(- 0.4) 0.2
6 M (C)
TT (3)
(Mean Value) ±SD
0
(5.1) ± 0.8
(5.4) ± 0.6
Δ TT (4)
(Mean Value) SE
3 M
(- 0.2) 0.1
(0.3) 0.1
6 M
No. & (%) TCET
(5)
0
5 (33)
10 (67)
3 M
4 (27)
8 (53)
6 M
5 (33)
4 (27)
No. & (%) OCET
(6)
0
7 (47)
7 (47)
3 M
3 (20)
6 (40)
6 M
2 (13)
1 (7)
No. & (%) RTCET
(7)
0
2 (13)
3 (20)
3 M
4 (27)
2 (13)
6 M
12 (80)
1 (7)
No. & (%) PIT(8)
0
15 (100)
15 (100)
3 M
9 (60)
10 (67)
6 M
10 (67)
6 (40)
No. & (%) CELE
(9)
0
9 (60)
14 (93)
3 M
11 (73)
14 (93)
6 M
11 (73)
14 (93)
A D
l
i i
(G
d
4) ( ) D DA Ch
i
D
l
i i
( ) TT Thi k
f
d
(i (1) DA: Doppler activity (Grades 0–4); (2) D DA: Change in Doppler activity; (3) TT: Thickness of common extensor tendon (in
millimetres); (4) D TT: Change in Thickness of common extensor tendon; (5) No. and (%) TCET: Number and percentage of patients
who had tears in the common extensor tendon; (6) No. and (%) OCET: Number and percentage of patients who had oedema in the
common extensor tendon; (7) No. and (%) RTCET: Number and percentage of patients who had reduced thickness of the common
extensor tendon; (8) No. and (%) PIT: Number and percentage of patients who had probe-induced tenderness; (9) No. and (%) CELE:
Cortical erosion at the lateral epicondyle. (a) SD: Standard deviation; (b) SE: Standard error. (A) 0: baseline assessment; (B) 3M: 3
month; (C) 6M: 6 months. the two interventions at 2 years, by Gosens and co-
researchers. However, and as the two studies were not
conducted by exactly the same researchers, and with two among the participants prevents any quantitative analysis
to assess the prevalence of these complications among
studies’ participants. Internal validity Items (2013)
Follow-up intervals
Baseline
1 month
3 months
Outcome measure
elements
(Mean Pain
score: 0-50)
& SE (1)
(Mean
FuncƟonal
score: 0-
100)
& SE
(Mean
Change in
Pain)
& SE
(Mean
Change in
Disability)
& SE
(Mean
Change in
Pain)
& SE
(Mean Change
in in Disability)
& SE
Interventions
CS (2)
(28.0) 8.0
(51.1) 22.3
(- 9.8)
2.2
(- 21.9)
4.3
(- 7.1)
2.2
(- 13.8)
4.3
PRP (3)
(27.5) 7.5
(51.5) 19.1
(- 0.5)
2.2
(- 5.2)
4.3
(- 6.0)
2.2
(- 16.6)
4.3
Saline
(25.0) 7.3
(47.1) 22.3
(- 1.7)
2.2
(- 3.4)
4.3
(- 3.3)
2.2
(- 7.6)
4.3
Outcome measure used
Oxford Elbow Score (Value) ±SD (4) Gautam et al. (2015)
Follow-up intervals
Baseline
2 weeks
6 weeks
3 months
6 months
Interventions
CS
(31.2) ±4.1
(P: 0.015)
(39.7) ±3.4
(P: 0.001)
(41.5) ±2.5
(P: 0.045)
(41.7) ±2.4
(P: 0.029)
(36.3)
±5.9
(P: 0.007)
PRP
(27.4) ±3.9
(34.7) ±4.3
(39.3) ±3.1
(39.3) ±3.3
(41.2)
±2.7
Outcome measure used
Modified Mayo Score (Value) ±SD Gautam et al. (2015)
Follow-up intervals
Baseline
2 weeks
6 weeks
3 months
6 months
Interventions
CS
(56.8) ±5.4
(P: 0.770)
(68.5) ±3.9
(P: 0.000)
(70.4) ±3.2
(P: 0.017)
(69.6) ±3.5
(P: 0.578)
(61.5)
±5.8
(P: 0.000)
PRP
(56.1) ±6.9
(61.3) ±3.1
(67.7) ±2.6
(70.2) ±2.2
(70.7)
±3.0
Outcome measure used
Hand grip strength (Value) ±SD Gautam et al. (2015)
Follow-up intervals
Baseline
2 weeks
6 weeks
3 months
6 months
Interventions
CS
(19.2) ±4.6
(P: 0.683)
(25.5) ±4.9
(P: 0.159)
(25.5) ±6.0
(P: 0.976)
(25.8) ±6.7
(P: 0.884)
(23.3)
±6.5
(P: 0.258)
PRP
(18.5) ±5.1
(22.5) ±6.6
(25.5) ±6.3
(25.5) ±5.6
(25.9)
±6.2
SE: Standard error. (2) CS: Corticosteroid. (3) PRP: Platelet-rich plasma. (4) SD: Standard deviation. (P:P value) W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 10 Table 7. Ultrasonographic outcome data reported by 2 studies. Table 7. Ultrasonographic outcome data reported by 2 studies. 7. Ultrasonographic outcome data reported by 2 studies. Study name &
Interventions used
The
Outcome
measures used
(at 3 different
follow-up periods)
Krogh et al. (2013)
Gautam et al. Internal validity Items distinctive articles with study titles, it appears that the
authors in the first study did not originally plan to
measure the patient’s outcomes at 2 years. This have
entailed some changes in the studies settings, including
the use of two different variants of one of the main
outcome measures used (DASH). This was declared by
one of the authors who stated that all DASH items were
summed in the first study, yet the core elements only
were used to calculate the DASH score in the second
study. Secondly, the two studies were conducted at two
teaching hospitals, with no details about which patients Discussion Five primary studies were included in this systematic
review (Table 2). All these studies were high in the
hierarchy of evidence and followed the randomised
controlled trials (RCTs) design [51]. Two of these studies
[45,46] were merely a two-stage trial rather than two
distinctive studies. Initially, Peerbooms and colleagues
investigated the effect of corticosteroid versus PRP
injections, with an end-point assessment at 1 year, which
was followed by an assessment of the longer-term effect of W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 11 Table 8. Adverse effects of injection therapies as reported by the review studies. Study
Adverse effects
Peerbooms et al. (2011)
Local inflammation at the injection site, which caused
increased pain 3 to 4 weeks after the injection
Gosens et al. (2011)
Initial worsening of localised pain (1 - 2 weeks). Krogh et al. (2013)
Generic
adverse
effects
PRP
Corticosteroid
post
injection
pain
- persistent pain
(4 subjects)
- Reduced elbow
movement (3
subjects)
- localised skin rash
(1 subject)
- local skin atrophy
(3 subjects)
- loss of skin
pigmentation (1
subject)
- Persistent pain (1
subject)
Gautam et al. (2015)
NIG
Omar et al. (2012)
NIG
(NIG: No information given) (NIG: No information given) were assessed in each hospital, and how much the
hospital settings were similar, and thus more details
could have eliminated any potential bias. [52,53], the variation in this demographic element
(smoking) can be considered as a confounding factor
especially if we observe the comparable therapeutic
effect noted in saline (placebo) group compared to
corticosteroid and PRP groups. While consecutive sampling was adopted by two
studies [45,46], no details provided by the other studies
[29,47,48]. Population’s details, which are important to
assess the external validity, varied widely among the
included studies (Table 3). Age and gender ratios of
participants were generally representative of the popu-
lation, although no demographic data were provided by
one study [47]. Also, and while Krogh and co-researchers
(2013) [29] provided the most extensive details of
patient’s demographic data including smoking history,
the percentage of patients who were current smokers
varied widely among the three study arms, which ranged
between 10 % in saline group, to around one-third in
PRP
and
glucocorticoid
groups. W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 Additionally, and although Krogh and co-
researchers avoided this potentially confounding element
by using saline as a placebo injection, the saline material
itself could be considered more than an inactive compara-
tor substance, with its possible beneficial/harmful effects
as well. Regarding the cost-effectiveness of the injection
treatments used, PRP injections may not be of interest
to healthcare providers with an overall cost of around ($
840–1000), compared to approximately one-third of this
for corticosteroid injections [45,54]. However, the higher
rate of symptoms recurrence among the relatively cheap
corticosteroid injections means that the expenses of new
treatments, including the higher costs of surgical treat-
ments for example [54], which might be required after
these cheap modalities fail, may entail higher cost than a
single PRP injection. The use of different types, doses and concentrations of
the interventional steroid preparations (Table 4) should
be noted, especially with the known difference of the
therapeutic effect of these preparations [55,56]. Addi-
tionally, the combined use of local anaesthetics with
steroid preparations in some studies [29,45] can be a
source of bias as well, as these anaesthetic agents add
volume to the injectate and help distributing the
corticosteroid substance within the affected tissues
[57], with possible changes in the effect of corticosteroid
injections. This kind of heterogeneity can be seen more
profoundly with the use of PRP injections, where
different
preparation
methods,
various
commercial
centrifugation/separation systems as well as different
concentrations of platelets, growth factors and other
blood products in the PRP preparations were used [58–
63] (Table 4). With no ideal concentrations of platelets
and growth factors in the PRP preparations has been
established yet, these sources of variability were clearly
observed in the included studies. This in addition to
technical details including centrifugation speed, buffer-
ing agents used to achieve physiological pH, and the time
needed for the centrifugation of blood. In all of the included studies, the participants were
provided with some forms of post-injection therapies,
including
different
physiotherapeutic
modalities
[29,45,46]. It should be noted here that these modalities
are known for their therapeutic effect in lateral epicondyl-
opathy [66,67], and again this might have created a source
of uncertainty when assessing the real effect of the injected
treatments. Outcome measures used by the review studies
It is clearly seen the degree of heterogeneity when
observing the various outcome measures used by the
included studies (Table 5). W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 12 have resulted in more pain in the early follow-ups
compared to those who received single-injection cortico-
steroid. This difference in injection technique may have
distributed the injectates differently in the study arms,
with possible subsequent different therapeutic effects in
vivo. Additionally, the same study [29] adopted the
practice of injecting local anaesthetic in the peritendon
(the connective tissue sheath around the tendon) before
administering the interventional treatments, with possible
subsequent detrimental effects on tendon cells viability
and thus change in treatment effect [64]. Also, the possible
mix of lidocaine with PRP at nearly the same site, could
have reduced the therapeutic effect of PRP, as it is shown
that local anaesthetics reduce the positive effects of PRP
[65]. It is unclear how much these details could have
affected the treatment delivery and subsequently the
therapeutic effect of the used treatments. Finally, and
with the inter-personal bias in mind, Peerbooms and co-
researchers [45] reported that treatments were given by
more than one person of different level of experience
(consultant and resident doctors), which might entail that
treatments could have been given differently. This
confounding factor has not been ruled out by two of
included studies [47,48]. Treatments received by the participants before start-
ing the trials is another important element. Although most
studies reported that patients received no treatments in
the last 3–6 months before the interventions, there is still a
potential long-term effect of any treatments given before
that, this in addition to the possible treatment-treatment
interactions after receiving the new interventional treat-
ments. This might be of real significance if we noted, for
instance, that steroid injection was received by as high as
58 % of patients received corticosteroid injections in the
trial by Krogh and co-researchers [29]. Additionally, and
with the considerable variation of the disease duration
before the treatment interventions, it is unclear how far
this kind of heterogeneity has a role when assessing the
comparative effect of the treatments given. Injection materials used While the use of cortico-
steroid as a control injection can be acceptable to compare
the effect of PRP, the use of such medicinal therapeutic
agent itself may undermine the accurate assessment of the
effect of PRP as a treatment option. For example, the
probable deleterious effects of corticosteroid [17] may
falsely
exaggerate
the
beneficial
effect
of
PRP
in
comparison. Discussion With
the
known
correlation between smoking and delayed tissue healing The randomisation process and allocation of study
population was not reported by two studies [47,48], which
entails a potentiality of selection bias, and questions the
quality of these two high-level evidence RCTs. Addition-
ally, the lack of blinding of outcome assessors was one of
the main limitations faced by most of the included studies. Krogh and co-researchers [29] were the best who reported
these elements. Peerbooms and co-researchers [45] only
reported the blinding of patients to treatment interven-
tions, Gautam and colleagues [47] only reported the
blinding of ultrasound operator(s), while the study by
Omar and co-researchers [48] lacked the information
about all the elements of the blinding process. W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 When assessing the outcome
measures used by the first two studies [45,46], and while
there were minimal variations in the baseline VAS values
between corticosteroid and PRP study arms, significant
differences in the DASH baseline scores were noted
(Table 5). This disparity necessitates careful interpreta-
tion of the DASH outcome results by the two studies. While PRTEE has sufficient psychometric properties
[68,69], it may lack the objectivity and thus the accurate
assessment, as it depends solely on the subjective
patient-rated evaluation, beside the fact that this was
the only clinical outcome measure used by Krogh and Although all of the included studies used one-phase of
treatment delivery of one injection, different injection
sites, techniques and methods were used. While peppering
technique (multiple tendon perforations) was adopted by
two studies [45,47], Krogh and colleagues used 1 direct
injection for corticosteroid while adopted peppering
technique for PRP and saline injections [29], which could W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 13 colleagues [29] (Table 6). This also demonstrates the
difficulty in comparing the clinical outcomes of this trial
[29] with the other 4 studies which used VAS and DASH
in their outcome measurement. corticosteroid injections could be attributed to the fact
that these patients usually experience relatively rapid
improvement of symptoms which encourage them to rely
on the affected limb prematurely, with the possible re-
injury of the newly treated tendon and subsequently a
recurrence of symptoms [29,46]. The role of this factor
can be investigated in the future, by observing the
participant’s return to daily activities and assessing/
restricting their degree of reliance on the affected limb
after receiving the treatment. Finally, and despite the
strong argument which states that LE is a degenerative
non-inflammatory process, the rapid therapeutic effect
provided by corticosteroid injections needs to be justified
here, which could hypothesise that the disease itself
could be caused by a combined pathophysiological
mechanisms, which could involve both inflammatory
and degenerative processes. Drop-out of participants Variable drop-out rates
were reported by the included studies. While (6%)
reported by Gosens and colleague [46] by the end of their
trials at 102 weeks, Krogh and colleagues [29] reported a
0% of drop-out at 3 months. W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 However, considering the high
drop-out which faced Krogh and colleagues at 12 months
(around 73%), it can be argued here that this could be the
main reason for these researchers to limit the study follow-
up to 3-month only. Interestingly, and with careful
observation of this study, it can be seen that the drop-
out rate among patients who received PRP was signifi-
cantly less than that in corticosteroid group beyond the 6-
month period. This could have been caused by the late
ongoing therapeutic effect of PRP, which might have
encouraged the patients to adhere to their original
interventional treatment option (PRP). Limitations of this review The relatively small
number of included studies, the ambiguity of some studies’
methodological details, and the potential biases noted in
many of the trial’s aspects are the main limitations of this
review. Additionally, and while the quantitative analysis
(meta-analysis) was considered in this review, the varied
outcome measures used and the different timings of follow-
ups have hindered the chance to conduct a meta-analysis
to draw a large-scale summative conclusion. Finally, and with the above reported data in mind, a
few final observations should be elaborated here. Firstly,
although most of the included studies followed the trend
of rapid reduction then a regaining increase in the VAS
and DASH values for the use of corticosteroid, there was a
unique trend observed at 52 weeks and 104 weeks by the
two consecutive studies [45,46]. This trend shows that the
VAS and DASH values in the corticosteroid group started
to fall again, which means that there were improvements
in pain and function at these two follow-ups. Although
this might be interpreted as an ongoing long-term
improvement using corticosteroid, it should be remem-
bered that this could have resulted from the fact that 13
out of 20 patients on the corticosteroid group have
received further treatments (re-intervention) after the
original treatments. Secondly, Krogh and co-researchers
[29] reported that no superior outcomes can be observed
with the use of PRP compared to saline and corticosteroid
injections. This can be attributed to the fact that the
study did not assess the longer term effects of PRP beyond
3 months, which could have shown improvement in the
PRP group compared to corticosteroids group, especially
that more clinical improvement started to appear in the
PRP group in the period between the 1- and 3-month
follow-ups. Funding for this research work No funding, whether from commercial or non-com-
mercial bodies, was received to support this research work. ( )
16. Wong MW, Tang YY, Lee SK, Fu BS, Chan BP, Chan CK
(2003)
Effect
of
dexamethasone
on
cultured
human
tenocytes and its reversibility by platelet-derived growth
factor. J Bone Jt Surg (American Volume) 85-A(10) 1914–
1920. Conclusion 8. Küçüksen S, Yilmaz H, Salli A, Ugurlu H (2013) Muscle
energy technique versus corticosteroid injection for man-
agement
of
chronic
lateral
epicondylitis:
randomized
controlled trial with 1-year follow-up. Arch Phys Med
Rehabil 94(11), 2068–2074. Platelet-rich plasma demonstrated better, although
delayed, therapeutic effects compared to corticosteroid
injections, with no subsequent regression of their positive
clinical findings for up to 2 years, and with no reported
adverse effects except for local discomfort felt at the
injection site, as report by a single study. This conclusion
can be also supported by the fact that no long gaps can be
observed between the follow-ups conducted by all of the
included studies, and thus minimal chance of missed data,
which might demonstrate different change in effect, can be
assumed. 9. Osborne H (2010) Stop injecting corticosteroid into patients
with tennis elbow, they are much more likely to get better by
themselves!. J Sci Med Sport 13(4), 380–381. 10. Walz DM, Newman JS, Konin GP, Ross G (2010)
Epicondylitis: Pathogenesis, Imaging, and Treatment 1. Radiographics 30(1), 167–184. ( )
11. Ahmad
Z,
Siddiqui
N,
Malik
SS,
Abdus-Samee
M,
Tytherleigh-Strong G, Rushton N (2013) Lateral epicondy-
litis: a review of pathology and management. Bone Jt J 95-B
(9), 1158–1164. ( ),
12. Chesterton LS, Mallen CD, Hay EM (2011) Management of
tennis elbow. Dovepress J 2011(2), 53–59. W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 14 Conflict of interest ( )
13. Carofino B, Chowaniec DM, McCarthy MB, Bradley JP,
Delaronde S, Beitzel K, Mazzocca AD (2012) Corticoste-
roids and local anesthetics decrease positive effects of
platelet-rich plasma: an in vitro study on human tendon
cells. Arthroscopy: J Arthroscopic Related Surg 28(5), 711–
719. The two authors of this systematic review declare no
conflict of interest. Data sharing statement Additional details about this review, including the full
original manuscript, are available on request by contact-
ing the corresponding author. 17. Coombes BK, Bisset L, Vicenzino B (2010) Efficacy and
safety of corticosteroid injections and other injections for
management of tendinopathy: a systematic review of
randomised controlled trials. Lancet 376(9754) 1751–1767. (
)
18. Ciccotti MC, Schwartz MA, Ciccotti MG (2004) Diagnosis
and treatment of medial epicondylitis of the elbow. Clin
Sports Med 23(4) 693–705. Ethical approval 14. Han SH, An HJ, Song JY, Shin DE, Do Kwon Y, Shim JS,
Lee SC (2012) Effects of corticosteroid on the expressions of
neuropeptide and cytokine mRNA and on tenocyte viability
in lateral epicondylitis. J Inflamm 9(40), 1–9. As a systematic review, no ethical approval was needed
to conduct this research work, as there was no involvement
or contact with human subjects. (
)
15. Liu D, Ahmet A, Ward L, Krishnamoorthy P, Mandelcorn
ED, Leigh R, Kim H (2013) A practical guide to the
monitoring and management of the complications of
systemic corticosteroid therapy. Allergy, Asthma & Clin
Immunol 9(1), 1–25. W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 Additionally, the selective reporting by the
same researchers [29] is difficult to be excluded here,
especially that the researchers who clearly reported the
higher clinical improvement in corticosteroid groups in
the period up to 1-month follow-up, did not similarly
acknowledge that this trend has reversed in favour of PRP
in
the
subsequent
follow-up. Thirdly,
despite
the
comparatively longer-term therapeutic effects of PRP,
the self-limited nature of LE should always be remem-
bered, as this might have a role in the condition
improvement on the mid- to long-term [70], although
this effect was not observed in the corticosteroid groups
who experienced deterioration in their symptoms on the
mid- to long-term. Fourthly, the short-term therapeutic
effect and high rate of symptoms recurrence following Recommendations for future research With the
wide variation in using the injection treatments, there is a
growing need to carefully assess and compare many of
these treatments settings, aiming to adopt a unified
preparation/technique, especially with PRP injections,
where different separation systems, concentrations of
blood components and injection techniques are used. For
example, with the different injection methods used by
various studies, the adoption a computer-guided injection
technique [54] can be utilised to enhance needle placement
and to avoid potential variability during the injection
process. It can be also suggested that a unified injection
protocol can possibly be used to standardise the mode of
injection, including for instance, the specific use of
peppering or single-dose injection, agreeing on a pre-
determined injection site, defining specific numbers of
injections, and to determine whether local injections will
be used before/with corticosteroid and PRP injection. Furthermore, the use of post-injection therapeutic modal-
ities should be regulated, by possibly avoiding or at least
adopting a unified post-injection protocol. That also
applies to the outcome measures used, as this can
considerably help comparing the clinical outcomes more
efficiently. These variables, if considered, can eliminate
any potential variations and thus help towards unbiased
assessment of the effect of the injection treatments. Finally, it should be noted here that none of the included
studies reported any details about the ethnicity of the
studies’ population, where different ethnic groups might
have different response to different treatments. Investi-
gating this variable in the future could reveal further
information about the disease pathophysiology and its
management. W. Ben-Nafa and W. Munro: SICOT-J 2018, 4, 11 Gautam VK, Verma S, Batra S, Bhatnagar N, Arora S
(2015) Platelet-rich plasma versus corticosteroid injection
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Nutritional status and food intake of patients with systemic psoriasis and psoriatic arthritis associated
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cc-by
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Nutritional status and food intake
of patients with systemic psoriasis and
psoriatic arthritis associated Marina Yazigi Solis1, Nathalia Stefani de Melo2, Maria Elisa Moschetti Macedo2, Fabiana Prata Carneiro2,
Cid Yazigi Sabbag3, Antonio Hebert Lancha Junior4, Vera Silvia Frangella5 ORIGINAL ARTICLE ORIGINAL ARTICLE ABSTRACT an increased risk for chronic diseases related to obesity, worsening
of the psoriatic lesions, and poor quality of life. Objective: To identify the nutritional status and food intake of
individuals with systemic psoriasis and psoriatic arthritis associated. Methods: This is an exploratory and cross-sectional study with 34
men aged between 19 and 60 years seen at a Psoriasis Center. Participants were divided into systemic psoriasis group and arthritic-
systemic psoriasis associated group. For nutritional assessment we
used anthropometry, bioelectrical impedance analysis and whole-
body plethysmography. Clinical and nutritional information were
assessed using the clinical and nutritional history-taking, and the
24-hour dietary recall. For statistics the general linear model test
(p < 0.05) was used. Results: According to the body mass index
29.4% patients (n = 10) were eutrophic, 41.2% (n = 14) overweight
and 29% (n = 10) obese. Almost all individuals (60%; n = 21) had
body fat percentage above normal levels (> 25%) and a high risk for
metabolic complications according to the waist circumference and
the obesity index, however, there were no statistically significant
differences between groups. The mean food intake, total fat, calories
and protein were above recommended levels, being 58.8% for lipids
(319.17 ± 241.02 mg of cholesterol and 17.42 ± 11.4 g saturated fatty
acids); 29.4% for calories and 67.6% for proteins. Thus, regardless of
the psoriasis type, an excessive consumption of calories, lipids, fatty
acids, cholesterol and a higher incidence of overweight were found. Conclusion: The sample showed an abnormal nutritional condition, Keywords: Psoriasis/complications; Obesity/complications; Feeding;
Food consumption; Nutritional status; Risk factors Study carried out at the Brazilian Psoriasis Study Center – CBEP, São Paulo (SP), Brazil.
1 Nutrition and Metabolism Applied to Motor Activities Laboratory – São Paulo (SP), Brazil; Physical Education and Sports School, Universidade de São Paulo – USP, São Paulo (SP), Brazil; Brazilian Psoriasis
Study Center – CBEP, São Paulo (SP), Brazil.
2 Centro Universitário São Camilo – São Paulo (SP), Brazil.
3 Brazilian Psoriasis Study Center – CBEP, São Paulo (SP), Brazil; Hospital Ipiranga – São Paulo (SP), Brazil.
4 Physical Education and Sports School, Universidade de São Paulo – USP, São Paulo (SP), Brazil; Nutrition and Metabolism Applied to Motor Activities Laboratory – São Paulo (SP), Brazil.
5 Centro Universitário São Camilo – São Paulo (SP), Brasil; Metabolism and Nutrition Institute – IMeN, São Paulo (SP), Brazil.
Corresponding author: Marina Yazigi Solis – Praça Amadeu Amaral, 47 - 4º andar – Conjunto 47 – Paraíso – Zip code: 01327-010 – São Paulo (SP), Brazil – Phone: (11) 3285-1273 – E-mail: ma_yazigi@hotmail.com
Received on: Jun 22, 2011 – Accepted on: Jan 20, 2011
Conflict of interests: None Study carried out at the Brazilian Psoriasis Study Center – CBEP, São Paulo (SP), Brazil. 2 Centro Universitário São Camilo – São Paulo (SP), Brazil. y Center – CBEP, São Paulo (SP), Brazil.
Activities Laboratory – São Paulo (SP), Brazil; Physical Education and Sports School, Universidade de São Paulo – USP, São Paulo (SP), Brazil; Brazilian Psoriasis RESUMO Objetivo: Identificar o estado nutricional e o consumo alimentar
de indivíduos com psoríase sistêmica e artropática associada. Métodos: Pesquisa exploratória e transversal, na qual avaliaram-se
34 homens, de 19 a 60 anos, atendidos em um Centro de Psoríase,
separando-os em Grupo PS (com psoríase sistêmica) e Grupo PAS
(com sistêmica mais artropática). A avaliação nutricional deu-se
pelo emprego da antropometria; bioimpedância e plestimografia de
corpo inteiro. Aspectos clínicos e nutricionais foram investigados
pela anamnese clínica, nutricional e recordatório de 24 horas. Empregou-se o teste General Linear Model (p < 0,05) para avaliação
estatística. Resultados: Segundo Índice de Massa Corporal, 29,4%
(n = 10) apresentaram-se eutróficos; 41,2% (n = 14) com sobrepeso
e 29% (n = 10) com obesidade. A maioria dos avaliados (60%;
n = 21) apresentou valor da porcentagem de gordura (avaliada pela
antropometria, bioimpedância e plestimografia de corpo inteiro)
acima da normalidade (> 25%) e com risco alto para complicações einstein. 2012;10(1):44-52 Nutritional status and food intake of patients with systemic psoriasis 45 metabólicas segundo CC e índice de obesidade, sem diferença
estatística significativa entre os grupos. O consumo alimentar médio
de lipídio, calorias e proteína apresentou-se acima do recomendado,
sendo 58,8% para os lípides (319,17 ± 241,02 mg de colesterol e
17,42 ± 11,4 g de ácidos graxos saturados); 29,4% para as calorias
e 67,6% para as proteínas. Assim, independentemente do tipo de
psoríase, encontrou-se consumo excessivo de calorias, lípides,
colesterol e ácidos graxos, além de maior ocorrência de excesso
de peso. Conclusão: A amostra apresentou estado nutricional
comprometido, aumento do risco para doenças crônicas relacionadas
à obesidade, agravamento das lesões e má qualidade de vida. psoriasis, observed that 30% of them also presented
joint problems. The PsA is characterized by T and B
cell infiltration leading to increased concentrations
of IL-1, IL-6, IL-12, IL-15, IL-17, IL-18, interferon-γ
and TNF-α in tissues that line the inner part of the
joints and are known as synovial tissue. Because of the chronic inflammation provoked
by the skin and joint disease, it is believed that
individuals with psoriasis are subjected to systemic
changes in the organism, such as insulin resistance,
changes in the lipid profile, obesity and increased
cardiovascular risk(7). Many studies point out a wide
relationship between psoriasis and the development
of associated chronic disease as dyslipidemia,
hypertension, non-alcoholic fatty liver disease and
type 2 diabetes mellitus, besides that, a high risk
for heart diseases and metabolic syndromes. INTRODUCTION Psoriasis is a demartosis that affects about 2 to 3% of the
world population and occurs in both sexes. Although it
can happen at any age, studies indicate a high prevalence
between 20 and 30 years, and also between 50 and
60 years of age. The incidence of psoriasis is higher
in countries such as Finland, Iceland, Norway and
Germany, and less common in dark-skinned persons of
East Africa, Indians and Eskimo people. In the United
States it is estimated that 8 million of persons have
psoriasis, in Brazil approximately 2% of population has
this disease(1). The literature reports that nutritional treatment
applied to psoriasis patients (associated to the
control of anthropometric and biochemical variables)
gives more clinical stability, preventing related non-
transmissible chronic diseases (NTCD), and provides
long-term quality of life. In other words, the weight
control improves the prognosis of psoriasis(2). On the
other side, other studies indicate that dietary pattern
established associated to life style may contribute to
the development of psoriasis. Therefore, nutrition can
influence psoriasis in two different ways, as the cause
of the metabolic disorder or as the treatment and
prevention. This disorder can affect the whole skin surface, but
it is most common on the extensor surface of the limbs,
scalp, nails, sacral and palmo-plantar regions. Psoriasis
is classified according to the site of the lesion. When it
appears as a chronic disease in plaque forms it is known
as psoriasis vulgaris, if presented as drops on the skin
it is called psoriasis guttata, which is usually found in
young people. In addition, there is the inverse psoriasis
found in the folds of the skin; the palmo-plantar
psoriasis found in palms and sole of feet; and there is
also the erythrodermic psoriasis that affects most of the
body(2,3). However, despite the fact that nutrition is considered
a tool for the treatment of psoriasis there are no
national or international guidelines that recommend an
adequate diet for such patients. Some authors suggest
that several active compounds in the human diet
perform an important role in the physiopathology of
psoriasis, having the same impact as the monitorization
of the diet energy supply and the total fat and saturated
fat intake on the control of NTCDs. RESUMO The
prevalence of metabolic syndrome in people with PsA
or other psoriasis type was high when compared with
those without psoriasis as suggested by Raychaudhuri
et al.(8) and Cohen et al.(9). Descritores: Psoríase/complicações; Obesidade/complicações; Alimentação;
Consumo alimentar; Estado nutricional; Fatores de risco INTRODUCTION Psoriasis can also be considered an autoimmune
disease mediated by defense cells known as T
lymphocytes (CD4 and CD8) that provoke a high
proliferation of proinflammatory cytokines, such
as interferon-γ, interleukins (IL) 1 and 6 and tumor
necrosis factor-alpha (TNF-α), which increase skin
lesions and causing also a chronic inflammatory
status(4). Among these nutrients some vitamins and minerals
(vitamins A, E, C and D and folic acid), omega 3
polyunsaturated fatty acids besides low-energy diets(10)
are mentioned. It is believed that some vitamins (A, E and
C), carotenoids and minerals (iron, copper, manganese,
zinc and selenium) have antioxidants ability, which
decrease oxidative stress and the production of reactive
oxygen species, mainly in systemic inflammation like
psoriasis(10). In 1818 a study by Alibert verified that psoriasis
patients with skin lesions could also develop joint
problems, which he named as psoriatic arthritis (PsA)
(5). An epidemiologic study done in 2002 by Zachariae
et al.(6) with 5,000 patients from Denmark, Finland,
Norway and Sweden who had different types of einstein. 2012;10(1):44-52 Solis MY, Melo NS, Macedo ME, Carneiro FP, Sabbag CY, Lancha Junior AH, Frangella VS 46 In addition, food fibers also play an important role
in systemic inflammation, decreasing oxidative stress
produced by large amounts of sugar, also improving
glycemic, insulin and lipidemic control. of health risk, assurance of anonymity and privacy of
personal information; the possibility of leaving the
study whenever they decided to do so without any
kind of uneasiness or punishment; they were informed
that their participation did not imply in expenses or
payment; if leaving the study at any time as desired
would have no influence in the treatment or loss of any
of the acquired benefits. All individuals who decided
to join the study signed the Informed Consent Form
(ICF), which included a statement authorizing the
authors to report results in congresses or scientific
journals if applicable. Population, sample, inclusion and exclusion criteria The center where the research was carried out receives
about 30 to 60 new cases of psoriasis each month of
both sexes aged between 20 and 65 years. The sample included men with systemic psoriasis
associated with psoriatic arthritis and who fit the
following inclusion criteria: aged between 19 and 60
years, male gender (to evaluate the natural cycle of
psoriasis without the changes of the aging process, of
gestation hormones or of menstrual cycle), be diagnosed
with psoriasis for at least three years because at this
time the clinical features will be well defined, using
laser therapies, topical medication, oral medicines like
methotrexate, cyclosporine and biological medicines,
being diagnosed with systemic psoriasis associated with
psoriatic arthritis. -
cultural and socioeconomic background, such as
name, age, city of birth, place of birth, address,
occupation, marital status, use of medicines and
practice of physical activity; - family background: parents, grandparents and uncles
health status; - clinical background: (prior and current disease):
prior health problems, allergies, medicine currently
used, test results, clinical or surgical treatment
performed; - nutriticional background (food or dietary): weight
gain or loss, anthropometric measures as weight,
height, circumference and skinfolds; We excluded women, those patients aged less than
19 years and more than 60 years, who had mild psoriasis
forms, and did not fit psoriasis classification as defined
in the study. - 24-hour dietary recall: it enabled to define and
measure food and drink intake in the 24 hours
before the interview. To do so, we provided booklets
with examples of utensils and portions to help
volunteers. OBJECTIVE To define, identify and associate the nutritional status
and food intake of patients with systemic psoriasis
and associated psoriatic arthritis seen at a Psoriasis
Center. Design and Setting The psoriasis area severity index (PASI) was used by
the responsible physician to classify and describe the
disease. The total score of the PASI was measured
based on the evaluation of four body surface areas
(head, chest, upper and lower limbs) using the method
described by Harari et al(11). The selected sample
had only individuals who presented PASI ≥ 18,
which is classified as a high severity. After selecting
the patients, all participants were interviewed to
gather clinical and nutritional history containing the
following issues: This is cross-sectional, quantitative, analytical study of
exploratory approach performed during 1 year and 5
months that started on December 2009. Data collection
was done at the Brazilian Psoriasis Study Center in São
Paulo city. einstein. 2012;10(1):44-52 Ethical issues We also evaluated the lack or the use of medicines
regarding dosage and administration hours before the
appointment at the center. The study was approved by the Research Ethics
Committee of the Centro Universitário São Camilo,
registered under the number 193/09, which addresses
the resolution 196 of October 10, 1996 stated by the
Brazilian National Health Council that supervises
researches using human beings. To evaluate body composition three different methods
of nutritional assessment were applied, and made
available in distinct moments to the subjects, mainly
those that did not show an uncontrolled risk or danger
to the participants. All participants were informed of the study
purpose, as well as the procedures to be done, the lack Nutritional status and food intake of patients with systemic psoriasis 47 -
Anthropometry: the volunteers’ weight was checked
using a digital balance (Plenna®) with 150 kg
weighting capacity and division of 100 g placed on
the floor level. For height (m) we used a metric
stadiometer (Sanny®) of 150 cm placed on the
wall. These measures used to calculate the body
mass index (BMI) given by weigh/height², and the
result was classified according to the World Health
Organization (WHO)(12). Using the techniques
described in the literature we measured the patients’
right hemi body, the arm circumference, waist
and hip circumference with an inelastic tape from
Sanny®(13). After gathering these measures the
relations hip waist/hip circumference was calculated
and the results classified as stated by WHO(14). To evaluate the regional body fat or topographic
evaluation the method proposed by Lohmann
et al. was used(15). Finally, the skinfold measures
were taken by a fat caliper from Lange® of 1.0 mm
of precision and maximum amplitude of 65 mm. The body areas measured followed the standards
proposed in the literature, and were taken by a
skilled evaluator. Each fold was measured in three
different times and the final results determined
by the mean values. We assessed skinfolds at
biceps, triceps, subscapular and suprailiac. From the
obtained results we calculated the fat percentage
based on the equation of Durnin et al.(16). model® was used. In the analysis the participants
were positioned on the stretcher in dorsal decubitus
without watches or any metallic objects. Before
placing skin-electrodes, we cleaned points of
contact with alchool 70%. Patients rested for 3
minutes before the measurement was performed. waist circumference (m) C index = C index = Ethical issues The equipment report provided: (1) fat mass
(fat percentage and body fat (Kg)); (2) thin mass
(muscles, bones and viscera); (3) whole body water
(liters and water percentage in thin mass); (
)
Whole body plethysmography (Air Displacement
Plethysmography, BOD POD®, body composition
system; Life Measurement Instruments, Concord,
CA). The evaluation was done using the criteria
described in the manual of Fields et al.(19). After the
equipment calibration, participants were evaluated
in a seated position, wearing minimal clothes, a
swimming cap, and without any metallic objects like
earrings, rings, necklace, etc. Variations between
pressure and volume were measured to define body
density of each subject using the equation of Siri(20). When the data obtained were insufficient, the
software itself performed a new evaluation until it
got the adequate screening. Statistical analysis
i i
l
l
i Statistical analysis was performed using the General
Linear Model test (GLM) for multivariate and
unbalanced analysis comparing PS group (systemic
psoriasis) with SPA group (systemic psoriatic arthritis
associated). A significance level of p<0.05 was used. In
addition, Shapiro-Wilk test was adopted to verify the
normal sample distribution. The conicity index (C index) was calculated to
determine obesity, body fat distribution and risk to
diseases associated to excessive weight by the equation
of Valdez et al.(17): C index =
waist circumference (m)
0109 × √ body weight (kg) ÷ height (m) RESULTS Relationship between length of time of psoriasis and related diseases
Group
Length of time of psoriasis
(years)
Evidence of related
diseases (%)
PS group
14.96 ± 7.24
36 yes
64 no
SAP group
16.75 ± 9.59
44.4 yes
55.6 no
Total
15.41 ± 7.93
38.2 yes
61.8 no
PS: systemic psoriasis; PAS: systemic arthropathy associated. Figure 1. Patients’ distribution by length of time of psoriasis and related Table 1. Relationship between length of time of psoriasis and related diseases
Group
Length of time of psoriasis
(years)
Evidence of related
diseases (%)
PS group
14.96 ± 7.24
36 yes
64 no
SAP group
16.75 ± 9.59
44.4 yes
55.6 no
Total
15.41 ± 7.93
38.2 yes
61.8 no
PS: systemic psoriasis; PAS: systemic arthropathy associated. Figure 1. Patients’ distribution by length of time of psoriasis and related Table 1. Relationship between length of time of psoriasis and related diseases
Group
Length of time of psoriasis
(years)
Evidence of related
diseases (%)
PS group
14.96 ± 7.24
36 yes
64 no
SAP group
16.75 ± 9.59
44.4 yes
55.6 no
Total
15.41 ± 7.93
38.2 yes
61.8 no
PS: systemic psoriasis; PAS: systemic arthropathy associated. Table 1. Relationship between length of time of psoriasis and related diseases
Group
Length of time of psoriasis
Evidence of related Table 1. Relationship between length of time of psoriasis and related diseases The mean BMI was 28.01± 4.42 kg/m². In the PS
group it was 27.86 ± 4.55 kg/m² and in the PAS group it
was 28.42 ± 3.99 kg/m² without significant differences
regarding BMI in both groups (p> 0.05). Based on the WHO20 classification, 10 patients
(29.4%) showed eutrophy using the BMI as a
parameter, 14 (41.2%) were overweight, 7 (20.6%)
were obese class I, 2 (5.9%) obese class II, and 1 (2.9%)
obese class III. These results showed that 24 patients
(70.6% of the sample) had excessive weight. In the PS
group 7 individuals (28%) had eutrophy, 12 (48) were
overweight, 4 (16%) were obese class I, 1 (4%) obese
class II, and 1 (4%) obese class III. The SAP group had
3 individuals (33.3%) with eutrophy, 2 (22.2%) were
overweight, 3 were (33.3%) were obese class and 1
(11.1%) obese class II (Table 2). Figure 1. Patients’ distribution by length of time of psoriasis and related Figure 1. RESULTS The sample had 34 patients divided into two groups. The PS group had 25 individuals (73.5%) and the SAP
group had 9 (26.5%) patients. The participants mean
age was 40.94± 11.19 years. The PS group mean age was
38.84 ± 10.57 years and in the SAP group it was 46.78
± 10,77 years. No statistical significance was observed
between groups as related to age (p>0.05). In both
groups, a total of 25 individuals (73.5%) were sedentary
and 9 (26.5%) did some physical activity. Both groups
had sedentary individuals, which comprised 76% of
PS group and 66.7% of SAP group. Therefore, in our
study sedentarism was a characteristic of the psoriasis
patients, which constituted 73.5% of all patients of the
sample. The index of central obesity (ICO), a new parameter
for obesity, was also calculated. Some studies have
considered the waist circumference is not sufficient to
measure the total body fat, mainly because height differs
in each race and sex. Therefore, the ICO is considered
the best parameter to determine central obesity, which
is calculated by dividing the waist circumference (m) by
the height (cm). This study cutoff point took the weight mean as
used in many countries and the waist circumference
as suggested by the International Diabetes Federation
(IDF) for metabolic syndrome(18), being 1.18 for women
and 1.25 for men. -
Bioelectrical impedance (BIA): this methods enabled
to obtain body water composition and fat mass
of the participants. To do so a Biodynamics 310 -
Bioelectrical impedance (BIA): this methods enabled
to obtain body water composition and fat mass
of the participants. To do so a Biodynamics 310 A total of 12 patients (32.4%) used biological
medicines (Stelara®, Humir®a, Enbrel®), being 8 (26.5%) einstein. 2012;10(1):44-52 einstein. 2012;10(1):44-52 Solis MY, Melo NS, Macedo ME, Carneiro FP, Sabbag CY, Lancha Junior AH, Frangella VS 48 32.4% high risk and 41.2% no risk, no significant
differences between groups was found (p > 0.05). immunosuppressive agents (methotrexate). Fourteen
patients (41.2%) did not use oral medicines. Among the individuals from the PS group 7
participants (28%) used biological medicines, 8 (32%)
immunosuppressive agents and 10 (40%) did not take
medicines. In the SAP group 4 individuals (44.4%) used
biological medicines, 1 (11.1%) immunosuppressive
agents and 4 (44.4%) did not take medicines. The waist-hip ratio for all participants, PS group,
SAP group was 0.91 ± 0.07; 0.90 ± 0.08 and 0.92 ± 0.06,
respectively. einstein. 2012;10(1):44-52 RESULTS Among all individuals evaluated, 24
(70.6%) did not present risk for cardiovascular disease
and 10 (29.4%) had risk based on WHO classification(14). The same risk proportion was found in the PS and SAP
groups, however, in the SAP group, using the same
parameters, no patient had risk for heart disease. A total of 13 patients (38.2%) presented associated
diseases like hypertension, diabetes mellitus, dyslipidemia,
heart disease and cancer. From this total, 9 individuals
(36%) were in PS group and 4 (44.4%) in SAP group
(Table 1). According to ICO the total mean of the sample was
0.55 ± 0.08 (0.54 ± 0.08 in PS group and 0.57 ± 0.07 in
SAP group). Considering this mean based on the
ICO value proposed by IDF, both groups had no risk
to metabolic syndrome. We did not find statistical
differences between the groups (p > 0.05). For C
index, the total mean for the sample was 25 ± 0.08, PS
group and SAP group had 1.24 ± 0.09 and 1.28 ± 0.07,
respectively, without statistical differences (p > 0.05). The mean length of time of psoriasis was 15.41 ± 7.93
years. Those in PS and SAP group had mean length
of disease time of 14.96 ± 7.24 years and 16.75 ± 9.59
years, respectively. There was no statistical significance
on length of time of the disease between the groups
(p>0.05). We stratified the sample into two other categories
considering the length of the disease for individuals
with up to 20 years of psoriasis and those with more
than 20 years with the disease. We observed among
the participants that 24 had psoriasis for up to 20
and 10 had the disease for more than 20 years, and
5 of these (50%) had one or more associated disease
(Figure 1). Hence, there were more comorbidities
associated to psoriasis in the group affected longer
by the disease. We found by the anthropometric technique after
summing the measurement values of the biceps, triceps,
subscapular and suprailiac skinfolds that body fat was
32.07 ± 6.52%, although more than 2/3 (n=25) of the Table 1. RESULTS Patients’ distribution by length of time of psoriasis and related Besides BMI we considered other central measures
to verify the possible risks. For example, the waist
circumference mean was 95 ± 13.45 cm (94.6 ± 14.45 cm
in PS group and 95.9 ± 10.27 cm in SAP group). As a result based on the metabolic syndrome council
classification(18), 23.5% of all participants presented
extremely high risk to develop metabolic complications, re 1. Patients’ distribution by length of time of psoriasis and relate Figure 1. Patients’ distribution by length of time of psoriasis and related Nutritional status and food intake of patients with systemic psoriasis 49 sample had fat percentage above the normal range with
risk to acquire associated disease. The fat percentage
in the PS group and SAP group was 30.85 ± 5.96% and
35.44 ± 6.81%, respectively. Finally, when fat percentage was measured using
plethysmography (BOD POD®) the mean values
were 27.43 ± 8.99% in the sample (27.02 ± 9.49% in
the PS group and 28.49 ± 7.48% in the SAP group). In addition, 24 participants had fat percentage above
normal ranges, being 17 in the PS group. In statistical
analysis no significant differences were found (p>0.05)
for this parameters between groups. A total of 17 patients (68%) from the PS group
and 9 from the SAP group had fat percentage risk of
associated diseases to obesity. We did not find statistical
differences between groups regarding fat percentage
(p > 0.05). Generally, we verified that fat percentage was above
normal ranges using the anthropometric methods in the
PS (72%) and SAP (88%) groups. According to BIA
and BOD POD®, the PS group had 64% and the SAP
group 78%, without significant differences. In addition,
we could observe that both groups had fat percentage
above normal ranges, and it was present in more than
64% of the participants (n=21) (Table 3). The mean of body fat using the BIA technique
was 24.14 ± 5.91% (23.68 ± 6.2% in PS group and
25.32 ± 4.86% in SAP group). Among the 34 patients,
67% were above normal range with associated risk for
disease, in which 16 were from PS group and 7 from
SAP group. In the comparison using this technique no
statistical significant differences was found between
groups (p > 0.05). Patients’ food intake was distributed in calories
and macronutrients. PS: systemic psoriasis; PAS: systemic arthropathy associated. einstein. 2012;10(1):44-52 RESULTS The mean caloric consumption
was 2,031.87 ± 728.96 Kcal (2,196.62 ± 565.28 Kcal in
PS group and 1,578.81 ± 913.27 Kcal in SAP group). In the SAP group 4 (44.4%) individuals had caloric
consumption lower than recommended. We verified
significant statistical differences between groups related
to caloric consumption (p < 0.05). Table 2. Classification of body composition, according to BMI related to
psoriasis Table 2. Classification of body composition, according to BMI related to
psoriasis
Diagnosis for
BMI
PS group
SAP group
Total
n
%
n
%
n
%
Eutrophy
7
28
3
33.3
10
29.4
Overweight
12
48
2
22.2
14
41.2
Class I obesity
4
16
3
33.3
7
20.6
Class II obesity
1
4
1
11.1
2
5.9
Class III obesity
1
4
0
0
1
2.9
BMI: body mass index; PS: systemic psoriasis; PAS: systemic arthropathy associated. The sample presented consumption of 43.3 ± 12.24% of
carbohydrates; 19.4 ± 5.43% of proteins and 35.8 ± 10.02%
of lipids. The mean consumption in the PS group
was 42.8 ± 12.53% of carbohydrates, 20.1 ± 5.78% of
proteins, 35.4 ± 9.6% of lipids. The PAS group presented Table 3. Comparison of body composition of psoriasis patients using different methods
Methods
Diagnosis (%)
PS group
SAP group
Total
n
%
n
%
n
%
Anthropometry
Morbid obesity (> 40)
2
8
3
33.3
5
14.7
High obesity (35-40)
4
16
1
11.1
5
14.7
Moderate obesity (30-35)
7
28
1
11.1
8
23.5
Mild obesity (25-30)
5
20
3
33.3
8
23.5
Above mean (16-24)
4
16
0
0
4
11.8
Mean (15)
0
0
0
0
0
0
Bioeletrical impedance
Risk (> 30)
4
16
2
22.2
6
17.6
Excessive adiposity (20-30)
12
48
5
55.6
17
50
Moderate thin mass (12-20)
7
28
2
22.2
9
26.5
Adequate (8-12)
0
0
0
0
0
0
Too thin (5-8)
0
0
0
0
0
0
Whole body plethysmography
Risk (> 30)
10
40
4
44.4
14
41.2
Excessive adiposity (20-30)
7
28
3
33.3
10
29.4
Moderate thin mass (12-20)
4
16
2
22.2
6
17.6
Adequate (8-12)
2
8
0
0
2
5.9
Too thin (5-8)
0
0
0
0
0
0
PS: systemic psoriasis; PAS: systemic arthropathy associated. Table 3. Comparison of body composition of psoriasis patients using different methods DISCUSSION The number of patients with joint problems resembled
the total observed in an epidemiological study done
by Zachariae et al.(6) with individuals from Demark,
Finland, Norway and Sweden. The number of patients with joint problems resembled
the total observed in an epidemiological study done
by Zachariae et al.(6) with individuals from Demark,
Finland, Norway and Sweden. Obesity cases observed followed by hypertension
and type 2 diabetes mellitus were almost the same as
those described in the study by Cohen et al.(9), which
evaluated 340 Israeli patients with psoriasis vulgaris. Another study by Altobelli(21) also determined the
occurence of NTCD in more than 1,300 individuals
with psoriasis from 21 dermatology departments in
Italy. These studies stated that the risk for NTCD
increases according to age, more specifically between
35 and 50 years, which seems to be due to exposition to
chemical pollution and bad life habits as to food intake
and by the lack of physical activity. A chronic inflammatory status, common in psoriasis,
inappropriate food intake and the lack of physical activity
may trigger the development of psoriais and related
NTCD, as well as increase the inflammatory process(23). However, it should be remembered interestingly, that
nutrition can influence psoriais in two different ways:
as cause of metabolic disorders or as treatment and
prevention. The literature reports that nutritional
treatment of psoriasis patients, associated to the control
of biochemical variables and anthropometrical control,
assures more clinical stability for individuals with
psoriasis, preventing commonly related NTCD and
providing long-term quality of life(24). A cohort study done in 1996 by Naldi et al.(22), evolving
around 78,000 nurses from the Nurse’s Health Study II
between 1991 and 2005 assessed the relationship among
BMI, waist and hip circumference and waist-hip ratio,
and revealed that high BMI represent high relative risk
(RR) to develop psoriasis. Therefore, patients with
BMI between 23.0 e 24.9 kg/m² have a RR of 1.19, those
with 25.0 to 29.9 kg/m² have a RR of 1.40, patients with
30.0 to 34.9 kg/m² have RR of 1.48, and those with
≥ 35.0 kg/m² a RR of 2.69. They also found a tendency
in the relationship between the increase of waist and hip
circumference and the appearance of psoriasis. Methods einstein. 2012;10(1):44-52 Solis MY, Melo NS, Macedo ME, Carneiro FP, Sabbag CY, Lancha Junior AH, Frangella VS 50 50 occurrence of obesity was two times higher than the
expected for the general population. consumption of 44.8 ± 11.28% of carbohydrates;
17.8 ± 3.83% of proteins; and 36.8 ± 11.02% of lipids. There were no statistic differences regarding carbohydrates
and lipids consumption between groups (p > 0.05), but
the PS group had a significantly higher consumption of
proteins (p = 0.006). The three methods of body composition used pointed
to excessive adiposity, without significant statistical
difference between groups, indicating the efficacy of
these methods to evaluate fat percentage of psoriasis
patients. We found after the analysis the lipid intake quality
that 40% of the sample (n=14) had an inappropriate
consumption of cholesterol (39.17 ± 241.02 mg),
saturated fat (17.42 ± 11.4 g) mono (22.12 ± 15.11 g)
and polysaturated (14.71 ± 10.71 g). Besides that, the
analyzed subjects had low intake of fibers (13.47 ± 8.91 g),
low consumption of fruits and vegetables, which helped
to reduce in both groups the amounts of vitamins (A,
B, C, D and B-complex), minerals (manganese and
selenium) in almost 50% for all micronutrients. The excessive weight was the most common
nutritional diagnosis in both studied groups, being
obesity more frequent in SAP group. It could be
explained by the fact that the individuals in the
SAP group had a systemic inflammation and joint
inflammation. However, such hypothesis could not be
supported by the literature and more studies should be
conduct to confirm this finding. Evidences in the literature and the findings of this
study confirm the positive relationship between psoriasis
and obesity. It is important to note that first one is
also considered an inflammatory disease and obesity
presents high production of pre-inflammatory cytokines
like TNF-α, IL-1, IL-6 e IL-8 by the adipose tissue. So, psoriasis and obesity can induce hyperglycemia,
decreased sensibility to insulin and hypertension, which
enables the occurrence of the metabolic syndrome
when the individual had central adiposity(1,7), that we
also found in this study. einstein. 2012;10(1):44-52 REFERENCES 1. Kremers HM, McEvoy MT, Dann FJ, Gabriel SE. Heart disease in psoriasis. J
Am Acad Dermatol. 2007;57(2):347-54. 2. Wolters M. Diet and psoriasis: experimental data and clinical evidence. Br J
Dermatol. 2005;153(4):706-14. 3. Christophers E. Psoriasis--epidemiology and clinical spectrum. Clin Exp
Dermatol. 2001; 26(4):314-20. This study results enabled to create a nutritional
profile of the participants, to identify the occurrence
of excessive weight no matter the psoriasis type. Therefore, these findings seem to be consistent with
other results reported in the literature regarding the
relationship between psoriasis and obesity, and the
risk to develop other comorbidities, like hypertension,
diabetes mellitus, dyslipidemia and heart diseases. 4. Zhang X, Wang H, Te-Shao H, Yang S, Wang F. Frequent use of tobacco and
alcohol in Chinese psoriasis patients. Int J Dermatol. 2002;41(10):659-62. 5. Henseler T, Christophers E. Disease concomitance in psoriasis. J Am Acad
Dermatol. 1995;32(6):982-6. 6. Zachariae H, Zachariae R, Blomqvist K, Davidsson S, Molin L, Mork C, et al. Quality of life and prevalence of arthritis reported by 5,795 members of the
Nordic Psoriasis Associations. Data from the Nordic Quality of Life Study. Acta Derm Venereol. 2002;82(2):108-13. 7. Sterry W, Strober BE, Menter A. Obesity in psoriasis: the metabolic, clinical
and therapeutic implications. Report of an interdisciplinary conference and
review. Br J Dermatol. 2007;157(4):649-55. In addition, when the psoriasis types were analyzed
trying to understand possible metabolic differences
between them, we verified that both had the same
behavior as to metabolic changes and increased risk to
develop related diseases. 8. Raychaudhuri SK, Chatterjee S, Nguyen C, Kaur M, Jialal I, Raychaudhuri SP. Increased prevalence of the metabolic syndrome in patients with psoriatic
arthritis. Metab Syndr Relat Disord. 2010;8(4):331-4. 9. Cohen AD, Sherf M, Vidavsky L, Vardy DA, Shapiro J, Meyerovitch J. Association between psoriasis and the metabolic syndrome. A cross-sectional
study. Dermatology. 2008;216(2):152-5. This study also indicated different methods and
tools that could be used in clinical practice to evaluate
and follow-up psoriasis patients. 10. Voutilainen S, Nurmi T, Mursu J, Rissanen TH. Carotenoids and cardiovascular
health. Am J Clin Nutr. 2006;83(6):1265-71. 11. Harari M, Shani J, Hristakieva E, Stanimirovic A, Seidl W, Burdo A. Clinical
evaluation of a more rapid and sensitive Psoriasis Assessment Severity Score
(PASS), and its comparison with the classic method of Psoriasis Area and
Severity Index (PASI), before and after climatotherapy at the Dead-Sea. Int J
Dermatol. 2000;39(12):913-8. CONCLUSION The psoriasis patients showed an abnormal nutritional
condition, an increased risk for chronic diseases
related to obesity, worsening of the psoriatic lesions,
and poor quality of life. There were no significant
differences between PS and SAP groups as related to
dietary intake. The psoriasis patients showed an abnormal nutritional
condition, an increased risk for chronic diseases
related to obesity, worsening of the psoriatic lesions,
and poor quality of life. There were no significant
differences between PS and SAP groups as related to
dietary intake. DISCUSSION Hence,
these results corroborate the findings of the present
study as regarding to enlarged waist circumference and
elevated BMI, that almost all participants evaluated
showed as well as having excessive weight, indicating
any degree of overweight and obesity. Besides that, the Results of food intake showed a heterogeneous
consumption between groups, however without statistical
differences. Also, there is the possibility of under report
in this population. According to Scagliusi et al.(25), an
overweight and obese population have higher under
reporting when describing its nutrition. The WHO(26) suggests, as values of percentile
distribution of macronutrients, proteins (10 to 15%)
and lipids (15 to 30%), although, in this study the
mean of protein and fat intake were higher among the
participants. On the other hand, carbohydrates were
lower than the recommended intake, between 50%
and 60%. This study results corroborate the findings einstein. 2012;10(1):44-52 Nutritional status and food intake of patients with systemic psoriasis 51 must be stimulated, following the recommendations
proposed by Brazilian Dietary Guidelines of daily
intake of such foods(29). of Willet(27), which verified a direct relation between
hyperlipidemic diet and the worsening of chronic
disease, like obesity and dyslipidemia. We think the high intake of lipids, cholesterol
and saturated fat was due to the high consumption
of animal proteins as reported by the participants. Besides, these individuals had low intake of fibers,
low consumption of fruits and vegetables, which also
contributed to reduced amount of vitamins (A, C,
D and B-complex), minerals (manganese, zinc and
selenium) in both groups. In a study with 316 psoriasis
patients done by Naldi et al.(22) that assessed food
intake using a semi-quantitative questionnaire, it was
found a low consumption of vitamins and minerals
in all participants. The same authors suggested
that consuming food with carotenoids, flavonoids,
selenium, vitamins A, C and E is extremely relevant to
psoriasis patients because it can reduce the production
of reactive oxygen species and tissue inflammation,
providing stability of cell membrane and recovery of
skin lesions. Food inadequacies according to Mobbs et
al.(28) might be explained by the increase of industrial
and urbanization process in Brazil and around the
world, increasing the consumption of fat/sugar rich
diets, sugar drinks and refined food, besides reducing
the consumption of complex carbohydrates and fibers. ACKNOWLEDGMENT We thank all volunteers of the research to be part of
this analysis, the Brazilian Psoriasis Study Center and
Instituto Vita to provide installations to perform the
investigation. REFERENCES These results indicate that nutritionists should
closely follow the amount and quality of food intake
of these patients. Particularly regarding the intake of
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et al. Comparisons of two-, three-, and four-compartment models of body
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238-45. 16. Durnin JV, Womersley J. Body fat assessed from total body density and its
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as an indicator of risk for cardiovascular disease. A cross-population study. Int
J Obes Relat Metab Disord. 1993;17(2):77-82. 24. Bjorneboe A, Smith AK, Bjorneboe GE, Thune PO, Drevon CA. Effect of dietary
supplementation with n-3 fatty acids on clinical manifestations of psoriasis. Br J Dermatol. 1988;118(1):77-83. 18. I Diretriz Brasileira de Diagnóstico e Tratamento da Síndrome Metabólica. Arq
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Bras Cardiol. 2005;84(1):3-28. 25. Scagliusi FB, Polacow VO, Artioli GG, Benatti FB, Lancha AH Jr. Selective
underreporting of energy intake in women: magnitude, determinants, and
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of air-displacement plethysmography. Curr Opin Clin Nutr Metab Care. 2004;7(1):11-4. REFERENCES 26.Organização Mundial da Saúde (OMS). Necessidades de energia e proteína. São Paulo: Roca, 1998. Série de relatos técnicos, 724. 20. Siri G. [Mental pathology]. Policlinico Prat. 1961;68:1722-4. Italian. 27. Willett WC. Invited commentary: comparison of food frequency questionnaires. Am J Epidemiol. 1998;148(12):1157-9; discussion 62-5. 21. Altobelli E, Petrocelli R, Maccarone M, Altomare G, Argenziano G, Giannetti
A, et al. Risk factors of hypertension, diabetes and obesity in Italian psoriasis
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carbohydrate diets cause obesity, low-carbohydrate diets reverse obesity: a
metabolic mechanism resolving the paradox. Appetite. 2007;48(2):135-8. 22. Naldi L, Parazzini F, Peli L, Chatenoud L, Cainelli T. Dietary factors and the risk
of psoriasis. Results of an Italian case-control study. Br J Dermatol. 1996;
134(1):101-6. 29. Philippi ST, latterza AR, Cruz AT, Ribeiro lC. Pirâmide alimentar adaptada: guia
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Ituglanis agreste, a new catfish from the rio de Contas basin, northeastern Brazil (Siluriformes: Trichomycteridae)
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Neotropical ichthyology/Neotropical Ichthyology
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Neotropical Ichthyology, 11(3):513-524, 2013
Copyright © 2013 Sociedade Brasileira de Ictiologia Neotropical Ichthyology, 11(3):513-524, 2013
Copyright © 2013 Sociedade Brasileira de Ictiologia 1Universidade Federal do Rio Grande do Norte, Laboratório de Ictiologia Sistemática e Evolutiva, Departamento de Botânica, Ecologia e
Zoologia. 59978-970 Natal, RN, Brazil. smaialima@gmail.com (SMQL), cinarapinheironeves@hotmail.com (CPN)
2Rua Professor Joaquim Louzada, 300, Casa 120, Bairro Camaquã, 91920-050 Porto Alegre, RS, Brazil. rafamcpaiva@gmail.com Ituglanis agreste, a new catfish from the rio de Contas basin,
northeastern Brazil (Siluriformes: Trichomycteridae) Sergio M. Q. Lima1, Cinara P. Neves1 and Rafael M. Campos-Paiva2 Ituglanis agreste, a new species of catfish, is described from a tributary stream of the rio Gongogi drainage, rio de Contas
basin, Bahia State, northeastern Brazil, from a transition area between the Atlantic Rain Forest and the semi-arid Caatinga
savanna. This species is distinguished from its congeners by the following characters: elongate interopercle plate with high
number of odontodes (26-30), high number of ribs (5-6), fewer vertebrae (36), number of branchiostegal rays (7), number of
pectoral-fin rays (i,6) and absence of s1 pore. Comparisons with other Ituglanis species and putative plesiomorphic characters
are presented. Some comments about conservation of Ituglanis species from northeastern Brazil are also made. Ituglanis agreste, espécie nova, é descrita de um tributário da drenagem do rio Gongogi, bacia do rio de Contas, Bahia,
nordeste do Brasil, de uma área de transição entre dois biomas, a Caatinga e a Mata Atlântica. Esta espécie distingue-se de
todas as demais espécies do gênero pelas seguintes características: placa interopercular alongada com elevado número de
odontódeos (26-30), número elevado de costelas (5-6), número reduzido de vértebras (36), número de raios branquiostégios
(7), número de raios da nadadeira peitoral (i,6), e ausência do poro s1. Comparações com outras espécies de Ituglanis e a
presença de supostos caracteres plesiomórficos são apresentados. Alguns comentários sobre conservação das espécies de
Ituglanis do nordeste do Brasil também são feitos. Key words: Atlantic Forest, Biome transition, Semi-arid Caatinga, Serra da Ouricana, Taxonomy. Introduction Copionodontinae, and Trichomycterinae) as well as the
specialized forms that comprise the TSVSG clade
(Tridentinae, Stegophilinae, Vandelliinae, Sarcoglanidinae,
and Glanapteryginae), which all share a smaller number of
ribs (2-7) (e.g., I. passensis Fernandez & Bichuette). The genus Ituglanis Costa & Bockmann is a
monophyletic assemblage of small trichomicterid catfishes,
previously included in Trichomycterus Valenciennes
(Trichomycterinae) due to its superficial similarity with the
members of this genus. Thus, a new taxon was proposed to
accommodate species that share three synapomorphies
present in the neurocranium: sphenotic directed anteriorly,
parieto-supraoccipital fontanel reduced to a small orifice or
completely closed (e.g., in I. macunaima Datovo & Landim,
and some specimens of I. epikarsticus Bichuette & Trajano
and I. mambai Bichuette & Trajano; Bichuette & Trajano,
2008), and autopalatine with a deep concavity in the inner
medial portion (Costa & Bockmann, 1993). The Ituglanis
assemblage has been considered essential for understanding
the emergence of the morphological and ecological
adaptations in the trichomycterids (de Pinna, 1998). This is
due to the intermediary phylogenetic position between both
the
more
generalized
forms
(Trichogeninae, Currently comprised of twenty nominal valid species
(Sarmento-Soares et al., 2006; Campos-Paiva & Costa, 2007;
Bichuette & Trajano, 2008; Wosiacki et al., 2012), and there-
fore the second most speciose trichomycterid genus,
Ituglanis has a wide distribution in the cisandine Neotropi-
cal region, from the Guianas to Uruguay (de Pinna & Keith,
2003), including five troglobitic species found in Central
Brazil (Bichuette & Trajano, 2008) and miniaturized forms as
I. macunaima from Araguaia Basin (Datovo & Landim, 2005). However, only recently two species of coastal basins of
northeastern Brazil were described: I. cahyensis Sarmento-
Soares, Martins-Pinheiro, Aranda & Chamon and I. paraguassuensis Campos-Paiva & Costa, respectively, from
the rio Cahy and rio Paraguaçu coastal basins, in Bahia state
(Sarmento-Soares et al., 2006, Campos-Paiva & Costa, 2007). 513 514 New Ituglanis from Brazil Laboratório de Sistemática e Evolução de Peixes Teleósteos,
Universidade Federal do Rio de Janeiro, Rio de Janeiro;
UFRN, Universidade Federal do Rio Grande do Norte, Natal,
and USNM, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. The coastal watersheds of eastern Brazil north of the
rio Pardo have their upper reaches within the temporary
rivers of the Caatinga biome, while the lower reaches are in
the Atlantic Forest biome. In contrast, watersheds south
of the rio Jequitinhonha are entirely in the Atlantic Forest
biome (Langeani et al., 2009). Material and Methods Paratypes. MNRJ 40197, 5, 32.4-40.8 mm SL; UFBA 7134, 5,
33.4-41.7 mm SL; UFRN 29, 6, 32.8-41.9 mm SL (3 c&s), all
collected with holotype. MZUSP 102535, 6., 38.4-44.1 mm SL,
same locality as holotype, 10 Feb 2009, S. M. Q. Lima, R. M. Campos-Paiva, P. Hollanda Carvalho & D. F. Almeida. Measurements and counts follow Costa (1992), and are
presented as percentages of standard length (SL), except
for subunits of head, which are presented as percentage
of head length (HL). Terminology for osteology follows
Adriaens et al. (2010), cephalic and lateral line systems
follow Arratia & Huaquin (1995), and caudal-skeleton
structures according to Bockmann et al. (2004). Counts of
procurrent caudal-fin rays, vertebrae, branchiostegal rays,
teeth and odontodes were made only in cleared and stained
specimens (c&s) prepared according to Taylor & Van Dyke
(1985). Vertebral counts do not include the Weberian
complex or the compound caudal centrum. Numbers in
parentheses indicate the number of specimens. Detailed
osteological description focus on traits informative for
future phylogenetic analyses of trichomycterid groups. Illustrations were prepared using a stereomicroscope with
a camera lucida attachment. Morphological data of
Ituglanis bambui Bichuette & Trajano; I. epikarsticus; I. guayaberensis Dahl; I. herberti (Miranda Ribeiro); I. ina
Wosiacki, Dutra & Mendonça; I. macunaima; I. mambai;
I. nebulosus de Pinna & Keith; I. parkoi (Miranda Ribeiro);
I. passensis Fernández & Bichuette; I. ramiroi Bichuette
& Trajano, were obtained in the original descriptions or in
detailed redescriptions (Bichuette & Trajano, 2004, 2008;
Canto, 2009; Dahl, 1960; Datovo & Landim, 2005;
Eigenmann, 1918; Fernández & Bichuette, 2002; Miranda
Ribeiro, 1940; de Pinna & Keith, 2003; Wosiacki et al.,
2012). Data from caudal skeleton of I. proops, I. nebulosus,
count of vertebrae and ribs of I. proops, I. nebulosus and
I. laticeps were based on digital images of x-ray
photographs available from the image base homepage of
the All Catfish Species Inventory project (http://
acsi.acnatsci.org/base/image_list.html?mode=genus&
genus=Pygidium, 26 December 2012) and the research tool
of the online collection homepage of USNM (http://
collections.mnh.si.edu/search/fishes/, 16 January 2013). Diagnosis. Ituglanis agreste is distinguished from all
species of the genus, except I. paraguassuensis by the
reduced number of vertebrae 36 (vs. 38 or more in all other
Ituglanis, except in I. paraguassuensis, I. nebulosus, I. bambui, I. epikarsticus, I. ramiroi, and I. passensis;
unknown for I. guayaberensis); number of ribs (5-6) (vs. 2
or 3 in I. amazonicus, I. eichorniarum, I. gracilior, I. Introduction While Ituglanis
paraguassuensis was only recorded in the semi-arid
Caatinga, I. cahyensis is restricted to the pluvial Atlantic
forest. This paper describes Ituglanis agreste, a new
species of catfish from the rio de Contas basin, found in a
transition area between the Atlantic Forest and the
Caatinga savanna. Ituglanis agreste, new species
Fig. 1 Holotype. MNRJ 40196, 40.5 mm SL, Brazil, Bahia,
municipality of Boa Nova; rio de Contas basin, rio Gongogi
drainage, rio Tarugo, tributary of rio Uruba, 14º22’08.95”S
40º11’45.33”W, 09 Jul 2008, S. M. Q. Lima, R. M. Campos-
Paiva, P. Hollanda Carvalho & H. Lazzarotto. Material and Methods ina,
I. macunaima, I. nebulosus, and I. parkoi ; 4 in I. laticeps
and I. cahyensis; 7 in I. passensis); number of pectoral-fin
rays i,6 (vs. i,4 in I. cahyensis, I. macunaima, and I. parahybae ; i,5 in I. amazonicus, I. eichorniarum, I. metae,
and I. nebulosus; i,7 in I. bambui, I. epikarsticus, I. mambai, I. passensis; iii,5 in I. guayaberensis; i,8 in I. ramiroi); and skin covered by irregular brown blotches
(vs. distinct color pattern in all other Ituglanis; absence in
I. ina and the subterranean species, I. bambui, I. passensis,
I. epikarsticus, and I. ramiroi). Ituglanis agreste is also easily distinguished from the
subterranean species by large eyes (vs. minute eyes) and
intense pigmentation (vs. pigmentation absent or almost
lack). Ituglanis agreste is distinguished from I. paraguassuensis, its geographically closest species, by a
higher number of odontodes in an elongate interopercle
patch (26-30) (vs. reduced with 14-15 odontodes); seven
branchiostegal rays (vs. 8) and pore s1 lacking (vs. s1
present). Description. Morphometric data for holotype and paratypes
are given in Table 1. Body elongated, subcylindrical about
to dorsal-fin origin, and gradually compressed in caudal
peduncle. Dorsal and ventral side view straights, except in
dorsal part of the head, which is slightly convex. Head
depressed, longer than wide, rounded in dorsal view. Eyes
rounded and small, without free orbital margin, covered by
a thin translucid membrane, lightly located on anterior half Abbreviations for institutions are: LIRP, Laboratório
de Ictiologia de Ribeirão Preto, Ribeirão Preto; MNRJ,
Museu Nacional, Rio de Janeiro; MZUSP, Museu de
Zoologia da Universidade de São Paulo, São Paulo; UFBA,
Universidade Federal da Bahia, Salvador; UFRJ, 515 S. M. Q. Lima, C. P. Neves & R. M. Campos-Paiva S. M. Q. Lima, C. P. Neves & R. M. Campos Paiva
515
f h
d B
b l
d h
d
d b
i
ill
fi
13 (i 11 i)
d
l
13 (2)
14 (1)
d
Fig. 1. Ituglanis agreste, new species, holotype, MNRJ 40196, 40.5 mm SL, Brazil, Bahia State, municipality of Boa Nova, rio de
Contas basin. a - lateral; b - dorsal; c - ventral views. of head. Barbels and head covered by minute papillae. fin rays 13 (i,11,i), procurrent dorsal rays 13 (2) or14 (1) and
Fig. 1. Ituglanis agreste, new species, holotype, MNRJ 40196, 40.5 mm SL, Brazil, Bahia State, municipality of Boa Nova, rio de
Contas basin. Material and Methods Maxilla curved with developed ventral
process, smaller than premaxilla (Fig. 2a). Dentary almost
straight with two regular rows of conical teeth and with
pronounced coronoid process (Fig. 3). Holotype
Paratypes
Range
Mean
SD
Standard length (mm)
40.5
32.9-41.9
38.0
3.35
Percentage of standard length
Body depth
12.4
11.6-15.1
13.5
1.1
Caudal peduncle depth
10.8
8.9-11.1
10.1
0.9
Body width
7.5
6.4-9.3
7.4
1.9
Caudal peduncle width
4.2
3.7-4.7
4.7
0.6
Dorsal-fin base length
10.9
8.6-11.8
10.3
0.8
Anal-fin base length
7.3
6.2-11.6
8.3
1.4
Pectoral-fin length
12.8
11.0-15.1
13.5
1.1
Predorsal length
69.1
64.1-73-3
69.1
1.9
Head length
16.9
15.9-19.6
17.8
1.1
Percentage of head length
Head depth
77.5
41.8-77.5
53.9
8.4
Head width
98.7
83.5-107.9
93.9
6.8
Interorbital width
18.9
14.4-22.8
19.7
2.4
Preorbital length
32.5
30.4-39.6
33.2
4.5
Eye diameter
9.8
8.1-14.5
9.9
1.9
Table 1. Morphometric data of holotype and 17 paratypes of
Ituglanis agreste. SD = standard deviation. p
p
(
g
)
Hyomandibula with an anterior laminar projection, a
deep depression in dorsolateral portion and pores on an-
terior and dorsomedian regions (Fig. 4). Anterior portion
of quadrate laminar and pronounced. Metapterygoid large,
laminar and somewhat trapezoidal, articulating to quadrate
by a cartilaginous block and to hyomandibula by bone
contact area. Interopercular plate broad and elongate bear-
ing conspicuous posterior projection, with 26-30
odontodes placed in two rows, including in anteroventral
projection (Fig. 4). Opercle with 16-17 odontodes. Odontodes conical, curved on the opercular patch of
odontodes and thinner on interopercular patch of
odontodes. Fig. 2. Neurocranium and Weberian capsule of Ituglanis agreste, UFRN 38, 41.8 mm SL; a, dorsal view; b, ventral view. Abbreviations: AOR, antorbital; APA, autopalatine; BAO+EXO, basioccipital-exoccipital bone; EPO, epioccipital; FRO, frontal;
FSO, parieto-supraoccipital fontanel; i1-i11, infraorbital pores; ll1-ll3, lateral line pores; LET, lateral ethmoid; MAX, maxilla;
MET, mesethmoid; ORB, orbitosphenoid; PAS, parasphenoid; PMX, premaxilla; po1, preopercular pore; PSO, parieto-
supraoccipital; PSC, posttemporo-supracleithrum; PTE, pterotic; s1-s6, supraorbital pores; SPH+POT+PSF, sphenotic-prootic-
pterosphenoid; SSO, sesamoid supraorbital; VOM, vomer; WEB, Capsule of Weberian apparatus. Scale bar = 1 mm. Fig. 2. Neurocranium and Weberian capsule of Ituglanis agreste, UFRN 38, 41.8 mm SL; a, dorsal view; b, ventral view. Material and Methods a - lateral; b - dorsal; c - ventral views. Fig. 1. Ituglanis agreste, new species, holotype, MNRJ 40196, 40.5 mm SL, Brazil, Bahia State, municipality of Boa Nova, rio de
Contas basin. a - lateral; b - dorsal; c - ventral views. fin rays 13 (i,11,i), procurrent dorsal rays 13 (2) or14 (1) and
ventral 10 (2) or14 (1). Caudal fin subtruncate. of head. Barbels and head covered by minute papillae. Mouth subterminal. Tip of nasal barbel reaching posterior
edge of opercular patch of odontodes; tip of maxillary barbel
reaching pectoral-fin base and tip of rictal barbel reaching
posterior edge of interopercular patch of odontodes or the
anterior edge of pectoral-fin. Mesethmoid with the anterior portion and shaft nearly
straight, gradually tapering at proximal tip (Fig. 2a). Lateral
ethmoid without lateral projections. Anterior fontanel
restricted to small pit with small enlargement in posterior
third of frontals. Posterior fontanel as small round opening
on posterior portion of parieto-supraoccipital. Sesamoid
supraorbital elongate, slender and curved, without lateral
process, slightly longer than autopalatine; sphenotic-
prootic-pterosphenoid narrow, with anterior portion
anteriorly directed (Fig. 2a). Pterotic with posterolateral
projection. Total vertebrae 36, 12-13 precaudal and 23-24 caudal. Ribs 5 (2) or 6 (1). Origin of dorsal-fin in a vertical through
the 22nd vertebra. Origin of anal-fin in a vertical through
the 23rd vertebra and through the base of the 7th ray of
dorsal fin. Origin of pelvic fin in a vertical through the 17th-
18th vertebra. Pectoral-fin rays i,6, pectoral fin triangular,
first ray simple and pectoral filament absent (holotype plus
one specimen) or ranging from 5-25% (16) greater than the
other rays of the pectoral fin. Pelvic-fin rays 5 (i,4). End of
pelvic fin overreach the urogenital papilla. Dorsal-fin rays
11-12 (v-vi,6), anal-fin rays 10-11 (v-vi,5). Principal caudal- Autopalatine with a deep concavity on the internal
medial border; posterior process moderate, about 50% of
the length of the autopalatine (Fig. 2a). Posttemporo-
supracleithrum with anterior and posterior processes. New Ituglanis from Brazil 516 Table 1. Morphometric data of holotype and 17 paratypes of
Ituglanis agreste. SD = standard deviation. Vomer arrow-shaped, with thin lateral projections and long
posterior process (Fig. 2b). Parasphenoid with two anterior
and one posterior processes. Weberian capsule fused to
basioccipital and exoccipital and with small lateral opening
on each side. Premaxilla trapezoidal and curved with two
rows of conical teeth. Material and Methods Abbreviations: AOR, antorbital; APA, autopalatine; BAO+EXO, basioccipital-exoccipital bone; EPO, epioccipital; FRO, frontal;
FSO, parieto-supraoccipital fontanel; i1-i11, infraorbital pores; ll1-ll3, lateral line pores; LET, lateral ethmoid; MAX, maxilla;
MET, mesethmoid; ORB, orbitosphenoid; PAS, parasphenoid; PMX, premaxilla; po1, preopercular pore; PSO, parieto-
supraoccipital; PSC, posttemporo-supracleithrum; PTE, pterotic; s1-s6, supraorbital pores; SPH+POT+PSF, sphenotic-prootic-
pterosphenoid; SSO, sesamoid supraorbital; VOM, vomer; WEB, Capsule of Weberian apparatus. Scale bar = 1 mm. S. M. Q. Lima, C. P. Neves & R. M. Campos-Paiva 517 P
h
l
ith
i
t
l f
d l t
Fig. 3. Left mandible of Ituglanis agreste, paratype, UFRN
38, 41.8 mm SL. Medial view. Abbreviations: AAR, angulo-
articulo-retroarticular; CPR, coronoid process; DEN, dentary. Scale bar = 1 mm. Fig. 5. Left hyoid arch of Ituglanis agreste, paratype, UFRN 38,
41.8 mm SL. Ventral view. Abbreviations: ACH, anterior ceratohyal;
BRR, branchiostegal rays; PCH, posterior ceratohyal; PUH,
h
l VHH
t lh
h
l S
l b
1 Fig. 5. Left hyoid arch of Ituglanis agreste, paratype, UFRN 38,
41.8 mm SL. Ventral view. Abbreviations: ACH, anterior ceratohyal;
BRR, branchiostegal rays; PCH, posterior ceratohyal; PUH,
parurohyal; VHH, ventral hypohyal. Scale bar = 1 mm. Fig. 3. Left mandible of Ituglanis agreste, paratype, UFRN
38, 41.8 mm SL. Medial view. Abbreviations: AAR, angulo-
articulo-retroarticular; CPR, coronoid process; DEN, dentary. Scale bar = 1 mm. Fig. 3. Left mandible of Ituglanis agreste, paratype, UFRN
38, 41.8 mm SL. Medial view. Abbreviations: AAR, angulo-
articulo-retroarticular; CPR, coronoid process; DEN, dentary. Scale bar = 1 mm. Fig. 5. Left hyoid arch of Ituglanis agreste, paratype, UFRN 38,
41.8 mm SL. Ventral view. Abbreviations: ACH, anterior ceratohyal;
BRR, branchiostegal rays; PCH, posterior ceratohyal; PUH,
parurohyal; VHH, ventral hypohyal. Scale bar = 1 mm. Parurohyal with conspicuous central foramen and lat-
eral process laminar and elongated, gradually tapering from
base to tip, with about same length of ceratohyal (Fig. 5). Ventral hypohyal with deep depressions for articulation
with parurohyal condyles. Seven branchiostegal rays. anterior process; second with two small alternated pro-
cess; third with one rounded posterior process; fourth flat-
tened, somewhat rectangular; fifth absent or not evident. First and second pharyngobranchials absent; third and
fourth stem-like, this last attached to tooth plate. Upper
pharyngeal tooth plate with conical teeth arranged in two
rows in ventromedial region (Fig. 6). Material and Methods Abbreviations: CLE,
cleithrum; PTR, pectoral rays; PR1, proximal radial 1
(cartilaginous); PR2, proximal radial 2; SCO, scapulocoracoid. Scale bar = 1 mm. Fig. 6. Branchial skeleton of Ituglanis agreste, paratype,
UFRN 38, 41.8 mm SL. Dorsal view (left dorsal elements and
gill rakers not shown). Abbreviations: BB2-4, basibranchials
2 to 4; CB1-5, ceratobranchials 1 to 5; EB1-4, epibranchials 1
to 4; HB1-3, hypobranchials 1 to 3; PB3-4, pharyngobranchials
3 to 4; TPL, tooth plate. Scale bar = 1 mm. Fig. 6. Branchial skeleton of Ituglanis agreste, paratype,
UFRN 38, 41.8 mm SL. Dorsal view (left dorsal elements and
gill rakers not shown). Abbreviations: BB2-4, basibranchials
2 to 4; CB1-5, ceratobranchials 1 to 5; EB1-4, epibranchials 1
to 4; HB1-3, hypobranchials 1 to 3; PB3-4, pharyngobranchials
3 to 4; TPL, tooth plate. Scale bar = 1 mm. defined. Chromatophores more concentrated in dorsolat-
eral region of head and opercular patch of odontodes. Nasal, maxillary, and rictal barbels with chromatophores
irregularly distributed throughout its entire length. Ven-
tral region of head with chromatophores forming conspicu-
ous spot on lower lip and with irregular and scattered
blotches in anterior region to isthmus and interopercle (Fig. 1c). Fins with few chromatophores, more often at base of
these. Some individuals exhibit lighter color than others,
but following same color pattern. Fig. 7. Left Pectoral girdle of Ituglanis agreste, paratype,
UFRN 38, 41.8 mm SL. Ventral view. Abbreviations: CLE,
cleithrum; PTR, pectoral rays; PR1, proximal radial 1
(cartilaginous); PR2, proximal radial 2; SCO, scapulocoracoid. Scale bar = 1 mm. Etymology. The specific name “agreste” (from latim agrestis,
which means relative to land, field, wild, or rustic) refers to
a semi-humid narrow strip parallel to the coast in north-
eastern Brazil, encompassing the area between the Rio
Grande do Norte State to the middle section of rio de Contas
basin in Bahia State (Forattini et al., 1981), that marks the
transition between two distinct biomes, the Atlantic For-
est and the semi-arid Caatinga (Prado, 2003), where the
new species was discovered. A noun in apposition. Distribution. Ituglanis agreste is only known from the type
locality, in rio Tarugo, municipality of Boa Nova, Bahia, north-
eastern Brazil. This river is a tributary of the rio Uruba, that
flows into rio Gongogi, principal drainage of right margin on
the middle section of rio de Contas basin (Fig. 10). Material and Methods First basibranchial absent, second and third
basibranchials ossified with anterior and posterior carti-
laginous tips; and fourth basibranchial consisting of elon-
gated, flattened and rounded cartilage (Fig. 6). First hypo-
branchial 1 ossified and stem-like; second elongate some-
what trapezoidal, mostly cartilaginous except for antero-
lateral process; third flattened cartilaginous with ossified
anterolateral process. First, second, third, and fourth
ceratobranchials ossified with cartilaginous tips, and with
posterolateral laminar expansions; fifth one curved with
small teeth on anterior half. First epibranchial with long Cleithrum flatenned, curved and slightly triangular. Scapulocoracoid trapezoidal, ossified with cartilaginous
tip (Fig. 7). Pelvic girdle slender and mostly ossified, with
cartilage restricted to posteromedial margins (Fig. 8). Upper caudal plates composed of pleurostyle and
hypural plates (hypurals 3-5) fused in a single triangular
element (1), or with hypural 3 and hypurals 4-5 not fused
(2) (Fig. 9). Parhypural partially fused to hypurals 1-2, with
distal portion forming a spine-like structure (Fig. 9). Fig. 4. Right suspensorium and opercular series of Ituglanis
agreste, UFRN 38, 41.8 mm SL. Lateral view. The arrow indicates
the interopercle plate broad and elongate. Abbreviations: HYO,
hyomandibula; IOP, interopercle; MPT, metapterygoid; OPE,
opercle; POP, preopercle; QUA, quadrate. Scale bar = 1 mm. Cephalic sensory and lateral line channels composed
by simple tubes ending in single pore, continuous and
interconnected on pterotic, sphenotic-prootic-
pterosphenoid, and lateral border of frontals. All cephalic
pores paired (18): supraorbital 2-3 (s2, s3, and s6; one speci-
men with s2 absent), infraorbital 4-5 (i1, i3, i10, and i11; one
specimen with i9), preopercular 1 and lateral line 3 (ll1, ll2,
and ll3). Coloration. Body with diffuse and irregular blotches on
yellow background. Chromatophores more concentrated
in dorsal and lateral region of body, forming large and ir-
regular blotches. In lateral region, chromatophores less
concentrated and blotches more dispersed. Ventral region
spotless from isthmus to pelvic-fin base, region posterior
to pelvic fin with diffuse, irregular and scattered blotches. Caudal penducle with smaller irregular blotches, but more Fig. 4. Right suspensorium and opercular series of Ituglanis
agreste, UFRN 38, 41.8 mm SL. Lateral view. The arrow indicates
the interopercle plate broad and elongate. Abbreviations: HYO,
hyomandibula; IOP, interopercle; MPT, metapterygoid; OPE,
opercle; POP, preopercle; QUA, quadrate. Scale bar = 1 mm. 518 New Ituglanis from Brazil Fig. 7. Left Pectoral girdle of Ituglanis agreste, paratype,
UFRN 38, 41.8 mm SL. Ventral view. Material and Methods Most Ituglanis species
have a low rib count (2-3; Sarmento-Soares et al., 2006),
but all cave species (I. bambui, I. epikarsticus, I. mambai,
I. passensis, and I. ramiroi; Fernández & Bichuette, 2002;
Bichuette & Trajano, 2004, 2008) and at least some other
species (I. parahybae, I. paraguassuensis, and I. proops)
present a large number of ribs (5-7) (de Pinna & Keith,
2003; Datovo & Landim, 2005). Trichomycterinae (Sarmento-Soares et al., 2006) and the
large interopercular plate bearing numerous odontodes (de
Pinna, 1998; Adriaens et al., 2010). Most Ituglanis species
have a low rib count (2-3; Sarmento-Soares et al., 2006),
but all cave species (I. bambui, I. epikarsticus, I. mambai,
I. passensis, and I. ramiroi; Fernández & Bichuette, 2002;
Bichuette & Trajano, 2004, 2008) and at least some other
species (I. parahybae, I. paraguassuensis, and I. proops)
present a large number of ribs (5-7) (de Pinna & Keith,
2003; Datovo & Landim, 2005). drifts. Substrate consists of rocks or sand. The most com-
mon depth observed was about 0.5 m, but some parts had
about 1.7 m. Some sites had underwater aquatic macro-
phytes and riparian vegetation. The backwaters area had
vegetation composed of Typha sp. The margins were de-
forested and the environment was mainly composed of
pastures. A reservoir without a lake formed by the dam of
the river was observed above the collection site. Other
species collected with Ituglanis agreste were: Astyanax
bimaculatus (Linnaeus), Callichthys callichthys
(Linnaeus), Geophagus brasiliensis (Quoy & Gaimard),
Rhamdia quelen (Quoy & Gaimard) and Poecilia reticulata
Peters, this last an exotic species. Ituglanis valid species usually show interopercle plate
reduction and less odontodes (10-17), a derived character
shared with the most specialized forms of the family, as-
sembled in the large clade TSVSG (de Pinna, 1998). At least
I. proops presents an elongate interopercle plate (de Pinna
& Keith, 2003; Datovo & Bockmann, 2010), a character
also observed in I. agreste. Ituglanis agreste also pre-
sents some others putative plesiomorphic characters, ob-
served in most species of Trichomycterus and Trichogenes
Britski & Ortega: the few number of vertebrae (36) and
anal-fin origin inserted on a vertical at 23rd vertebra. Both
features are also present in I. paraguassuensis (Campos-
Paiva & Costa, 2007), which suggests a basal position to
these species within the genus. Material and Methods Nevertheless, no formal
phylogenetic analysis among Ituglanis species has been
made yet, and the relationships among them remain un-
known (Datovo & Landim, 2005; Wosiacki et al., 2012). Material and Methods This is
the transition zone of the Serra da Ouricana, where small
remnants of a formerly extensive humid forest that delin-
eates the limits of the Atlantic Forest domain in the region
(Gonzaga et al., 1995). Just a few kilometers to the west, the
landscape gives way to the semi-arid Caatinga region, typi-
cally known for regular droughts. Thus, it is possible that
the rio Tarugo represents the western geographical limit of
distribution for this species. During a two year sampling
(2008 - 2009) along a 325 km transect of the rio de Contas
(between the municipalities of Ibuaçussê and Uruçuca),
Ituglanis agreste was found only at one site. Ecological notes. Ituglanis agreste was found along a
stretch of about 200 m in a mid-small size river with width
up to 8 m, in a moderate slope with clear and cold water,
with transparency of about 2 m (Fig. 7). The water flow
was mainly turbulent, with alternating areas of currents
and pools, though a predominance of moderate to strong 519 S. M. Q. Lima, C. P. Neves & R. M. Campos-Paiva Fig. 9. Caudal skeleton of Ituglanis agreste, paratype, UFRN
38, 41.8 mm SL. Left lateral view. Abbreviations: HS, hemal
spines; H3, hypural 3; H4+H5, hypurals 4 and 5 fused; NS,
neural spine; PC1+UR, pleural 1 and ural fused; PC2, pleural
centrum 2; PH+HU1+HU2, parhypural partially fused to
hypurals 1-2; UN, uroneural. Scale bar = 1 mm. Fig. 8. Pelvic girdle of Ituglanis agreste, paratype, UFRN 38,
41.8 mm SL. Dorsal view. Abbreviations: APS, anterior processes;
AMP, anterior medium processes; BPT, basipterygium; PSP,
pelvic splint; PVR, pelvic rays. Scale bar = 1 mm. Fig. 9. Caudal skeleton of Ituglanis agreste, paratype, UFRN
38, 41.8 mm SL. Left lateral view. Abbreviations: HS, hemal
spines; H3, hypural 3; H4+H5, hypurals 4 and 5 fused; NS,
neural spine; PC1+UR, pleural 1 and ural fused; PC2, pleural
centrum 2; PH+HU1+HU2, parhypural partially fused to
hypurals 1-2; UN, uroneural. Scale bar = 1 mm. Fig. 8. Pelvic girdle of Ituglanis agreste, paratype, UFRN 38,
41.8 mm SL. Dorsal view. Abbreviations: APS, anterior processes;
AMP, anterior medium processes; BPT, basipterygium; PSP,
pelvic splint; PVR, pelvic rays. Scale bar = 1 mm. Trichomycterinae (Sarmento-Soares et al., 2006) and the
large interopercular plate bearing numerous odontodes (de
Pinna, 1998; Adriaens et al., 2010). Discussion The new species is recognized as a member of Ituglanis
because it possesses all three synapomorphies proposed
for the genus by Costa & Bockmann (1993). However,
among the genus, Ituglanis agreste has some characters
that are apparently a plesiomorphic condition, character-
istic of the most basal trichomycterid taxa: the high num-
ber of ribs (5-6), since it is present in a highest number
(more than eight) in Copionodontinae, Trichogeninae, and 520 New Ituglanis from Brazil Fig. 10. Geographic distribution of the three Ituglanis species in northeastern Brazil. The locality of Ituglanis agreste (rio
Tarugo) represents the type locality. Fig. 10. Geographic distribution of the three Ituglanis species in northeastern Brazil. The locality of Ituglanis agreste (rio
Tarugo) represents the type locality. If corroborated, these results indicate that the genus
Ituglanis could be another example of the “biogeographic
pattern B” proposed by Ribeiro (2006) for several freshwa-
ter fish groups in the coastal drainages of eastern Brazil,
including
another
trichomycterid
genus,
the
Sarcoglanidinae Microcambeva Costa & Bockmann. Such
pattern makes sister-group relationships found in the
coastal basins of Brazil and subsequent irradiation with
the most inclusive groups located inside the Brazilian
Shield and the rio Amazonas basin (Ribeiro, 2006). paraguassuensis, from the rio Paraguaçu basin adjacent in
the north; and I. cahyensis from rio Cahy basin in the
south, the diagnostic features of the new species of rio de
Contas basin gives enough evidence to differentiate from
these two species formally described, confirming the re-
stricted distribution of most species of the genus. The most conspicuous differences between I. agreste
and I. paraguassuensis are found in interopercle patch of
odontodes (elongate with 26-30 odontodes vs. reduced with
14-15 odontodes), and should not be related to ontoge-
netic development once individuals of similar range size
were observed (3 c&s, 32.8-41.9 mm SL vs. 3 c&s, 30.3-39.6
mm SL). According to Arratia et al. (1990) these integu-
mentary teeth arise very early during ontogeny (Datovo &
Bockmann, 2010), indicating that Ituglanis from rio de
Contas and rio Paraguaçu are not conspecific. They can
be also distinguished by the number of branchiostegal rays,
(7 vs. 8) and lack of s1 pore (vs. presence). Nevertheless,
they share some diagnostic features used to distinguish
other Ituglanis, as 36 vertebrae, seven pectoral-fin rays
(i,6) and six pair of ribs. Discussion (2010), in a study of freshwater fish promoting the expan-
sion and implementation of conservation units in south-
ern Bahia, conduct ichthyological surveys in the rio
Cachoeira, Colônia, Contas, and Jequié basins, including
these new protected areas in Boa Nova, near the type lo-
cality (about 9 km east). These authors only recorded the
presence of Ituglanis sp. about 120 km south of the type
locality of I. agreste, in the rio Cachoeira basin. Because
the taxonomic status of Ituglanis sp. collected in the rio
Cachoeira basin remains uncertain, it could belong to a
previously described species or a distinct new species. Regardless, I. agreste most likely occurs in the new con-
servation areas in Boa Nova (PARNA and/or REVIS), since
these protected areas are situated near the type locality
where the new species was collected. The color pattern of Ituglanis agreste also differs
clearly from I. cahyensis that presents longitudinal rows
of coalescing blotches (Sarmento-Soares et al., 2006). They
also differs by several features, as the higher number of
pectoral fin-rays (i,6 vs. i,4), higher number of opercular
(17 vs. 9-12) and interopercle (26-30 vs. 16) odontodes, fewer
vertebrae (36 vs. 38 or more), higher number of
branchiostegal rays (7 vs. 4-6), higher number of ribs (5-6
vs. 4), 13 caudal-fin rays (vs. 10-11), 10-11 anal-fin rays (vs. 7-8), the position of the pelvic fin, with its origin inserted
on a vertical at 17th and 18th vertebra (vs. 20th) and
metapterygoid large in contact with the quadrate and the
hyomandibular (vs. metapterygoid short, laminar, almost
rectangular joined only with the quadrate; Sarmento-Soares
et al., 2006). Despite of that, they shared at least one char-
acter, the absence of s1 pore. The type locality of Ituglanis agreste is located in the
coastal strand of the Serra da Ouricana, which holds im-
portant small remnants of a formerly humid forest (Gonzaga
et al., 1995) and delineates the westernmost limits of the
Atlantic Forest domain bordering with the Caatinga biome. In northeastern Brazil these transition zone are referred as
“agreste”, which corresponds to a semi-humid climate (Fig. 10). The three species of Ituglanis described to the north-
eastern Brazil were found in streams placed in distinct cli-
mate and vegetation domains: I. paraguassuensis in the
Caatinga, I. cahyensis in the Atlantic Forest and I. agreste
was collected in a transition area between these two
biomes. Discussion The number of vertebrae shared
by them, was referred as an unusual condition among the
genus (Campos-Paiva & Costa, 2007), but it is shared at
least by other five species (I. nebulosus, I. passensis, I. ramiroi, I. epikarstikus and I. bambui; Fernández &
Bichuette, 2002; de Pinna & Keith, 2003; Bichuette &
Trajano, 2004). The troglomorphic species from Central The caudal skeleton of Ituglanis agreste presents two
conditions: upper hypural 3, 4, and 5 fused (one specimen)
or hypural 3 detached from 4 and 5 (two specimens). Both
attributes occurring within the same species was also re-
ported for I. ina (Wosiacki et al., 2012), Trichomycterus
tupinamba Wosiacki & Oyakawa (Wosiacki & Oyakawa,
2005), and vary among the genus in the subfamily
Trichomycterinae (Arratia et al., 1978; comparative mate-
rial). Within Ituglanis, some species has the hypural 3 de-
tached from 4 and 5 (I. amazonicus and I. eichorniarum),
but at least eight species (I. paraguassuensis, I. parahybae, I. proops, I. macunaima, I. nebulosus, I. bambui, I. epikarticus, and I. ramiroi have the hypurals
fused, also seen in the specialized species of the TSVSG
clade (Bichuette & Trajano, 2004; Datovo & Landim, 2005). Although situated between two coastal basins of Ba- Although situated between two coastal basins of Ba-
hia with Ituglanis species previously described: I. S. M. Q. Lima, C. P. Neves & R. M. Campos-Paiva S. M. Q. Lima, C. P. Neves & R. M. Campos-Paiva 521 Fig. 11. Type locality of Ituglanis agreste, Brazil, Bahia State,
Boa Nova municipality, rio Tarugo, rio de Contas basin. Brazil and a few specimens of I. ina also presents a high
number of interopercle odontodes (more than 24, except in
I. epikarsticus) but differ in some aspects from I. agreste
(pectoral-fin rays i,7 or i,8 and some degree of reduction of
eyes and pigmentation in subterranean species or usually
i,5 and homogeneous color pattern with dark vertical bar
in caudal peduncle in I. ina vs. i,6 and spotted color pat-
tern; Bichuette & Trajano, 2008; Wosiacki et al., 2012). Although the color pattern of I. agreste is similar to I. paraguassuensis, both with a yellow background body
covered by blotches, in I. agreste the spots are more dif-
fuse and irregular than I. paraguassuensis, that presents
blotches more conspicuous (Campos-Paiva & Costa, 2007). The concentration of chromatophores is more intense in I. Discussion agreste, covering the dorsal and lateral region of the body,
which gives a darker appearance to I. agreste, if compared
to I. paraguassuensis, in which the pigmentation is slightly
more concentrated near the eyes and opercular region. Fig. 11. Type locality of Ituglanis agreste, Brazil, Bahia State,
Boa Nova municipality, rio Tarugo, rio de Contas basin. agriculture and cattle (Fig. 11) in Boa Nova adjacencies. According to Gonzaga et al. (1995) the forest environments
in Serra da Ouricana are among the most neglected habi-
tats for conservation. Only recently this region was recog-
nized of a high importance for conservation (MMA, 2007),
which led to the creation of new conservation units in 2010:
Boa Nova National Park (PARNA) and Boa Nova Wildlife
Refuge (REVIS), with the aim to preserve the natural eco-
systems of the transition area between Atlantic forest and
Caatinga, protect endangered species and maintain water
sources/springs and streams (Brasil, 2010). Cetra et al. (2010), in a study of freshwater fish promoting the expan-
sion and implementation of conservation units in south-
ern Bahia, conduct ichthyological surveys in the rio
Cachoeira, Colônia, Contas, and Jequié basins, including
these new protected areas in Boa Nova, near the type lo-
cality (about 9 km east). These authors only recorded the
presence of Ituglanis sp. about 120 km south of the type
locality of I. agreste, in the rio Cachoeira basin. Because
the taxonomic status of Ituglanis sp. collected in the rio
Cachoeira basin remains uncertain, it could belong to a
previously described species or a distinct new species. Regardless, I. agreste most likely occurs in the new con-
servation areas in Boa Nova (PARNA and/or REVIS), since
these protected areas are situated near the type locality
where the new species was collected. agriculture and cattle (Fig. 11) in Boa Nova adjacencies. According to Gonzaga et al. (1995) the forest environments
in Serra da Ouricana are among the most neglected habi-
tats for conservation. Only recently this region was recog-
nized of a high importance for conservation (MMA, 2007),
which led to the creation of new conservation units in 2010:
Boa Nova National Park (PARNA) and Boa Nova Wildlife
Refuge (REVIS), with the aim to preserve the natural eco-
systems of the transition area between Atlantic forest and
Caatinga, protect endangered species and maintain water
sources/springs and streams (Brasil, 2010). Cetra et al. Acknowledgments We are grateful to Henrique Lazzarotto by helping to
photograph the holotype and Erica Caramaschi for provid-
ing logistical support. To Daniel Almeida, Henrique
Lazzarotto, and Pedro Hollanda-Carvalho for helping on
fieldwork. To Wilson Costa (UFRJ), Marcelo R. Britto
(MNRJ) and Michel D. Gianeti (MZUSP) that provided the
comparative material. To Ricardo Rosa, Telton Ramos and
Robson Ramos (UFPB) by loan the steromicroscope with
camera lucida. To Ali Ger that reviewed the English of a
previous version of the manuscript. This study was sup-
ported by CNPq (Conselho Nacional de Desenvolvimento
Científico e Tecnológico/ Ministério de Ciências e
Tecnologia), Biodinâmica LTDA and Bahia Mineração
LTDA. Collections were made under permits 010/2008
NUBIO/IBAMA - BA from Instituto Brasileiro do Meio
Ambiente e Recursos Naturais Renováveis/ Ministério
Brasileiro do Meio Ambiente. Comparative material examined. Trichogenes longipinnis,
UFRJ 894, 9, 29.7-46.5 mm SL, Brazil, São Paulo State, Ubatuba
municipality, affluent of rio Camburi; UFRJ 682, 1 c&s, 48.4
mm SL, São Paulo, Ubatuba, affluent of rio Camburi. Trichomycterus potschi, UFRJ 719, 2 c&s 33.4-47.6 mm SL,
Brazil, Rio de Janeiro State, Itaguaí municipality;
Trichomycterus giganteus, UFRJ 5999, holotype, 120.6 mm SL,
Brazil, Rio de Janeiro State, Campo Grande municipality, rio
Guandu-Mirim; UFRJ 5730, paratypes, 10, 116.2-138.7 mm
SL, same locality. Ituglanis amazonicus, MNRJ 35401, 3, 38.8-
56.6 mm SL, Brazil, Pará State, Uruará municipality, tributary
of rio Uruará; USNM 305468, 2, 31.6-31.7 mm SL, Bolivia,
Beni, Ballivián, 40 km east San Borja; USNM 317738, 1 c&s,
42.3 mm SL, same locality; USNM 300990, 1, 33.2 mm SL,
Peru, Madre de Dios, Manu National Park; USNM 263929, 1,
43.4 mm SL, Madre de Dios, Reserva Natural de Tambopata, on
trail to laguna Cocococha; USNM 263930, 1, 34.0 mm SL, Madre
de Dios, rio Tambopata. Ituglanis cahyensis, MNRJ 32080, 1,
39.30 mm SL, Brazil, Bahia State, Prado municipality, rio Cahy. Ituglanis eichorniarum, UFRJ 5474, 1, 24.3 mm SL, Brazil,
Mato Grosso State, Porto Cercado, Corixo Santa Rosa in rio
Cuiabá; UFRJ 5608, 1 c&s, 29.4 mm SL, same locality. Ituglanis
gracilior, USNM 272287, 1, 50.8 mm SL, Venezuela, Amazonas,
caño Temblador, in the road between San Carlos do Rio Negro
and Solano; USNM 272288, 1 c&s, 56.2 mm SL, Amazonas,
caño Loro, in the road between San Carlos do Rio Negro and
Solano; USNM 272289, 1, 50.0 mm SL, Amazonas, caño Urami,
upriver of Santa Lucia. Discussion Unlike rio Cahy basin, which is a small coastal
basin fully inserted in the Atlantic forest domains, the rio
Paraguaçu and Contas basins are perennial middle size riv-
ers, with some intermittent tributaries in the upper and
middle stretches in the Caatinga region, and lower stretches
in the Atlantic Forest (Rosa et al., 2003). The protection of water courses and their riparian mar-
gins through the creation of conservation units is among
the most important measures concerning the future and
diversity of fish species of the coastal basins of Brazil
(Menezes et al., 2007). Fortunately, the three species of
Ituglanis known from Bahia (including the one described
herein), are found in hydrographic basins that pass through
integral protection conservation units: I. paraguassuensis
in the rio Paraguaçu basin that passes through the Chapada
Diamantina National Park, I. cahyensis in the rio Cahy ba-
sin that has part of it course protected by the The collection site was highly degraded due to anthro-
pogenic impacts, mainly removal of riparian vegetation for 522 New Ituglanis from Brazil Paraguaçu; LIRP 5834, paratypes, 5, 34.8-41.3 mm SL; UFRJ
7209, paratype, 1, 36.5 mm SL; UFRJ 7282, paratype, 1 c&s,
39.6 mm SL; USNM 301016, paratypes, 5 (2 c&s), 33.7-41.6
mm SL, same locality. Ituglanis proops, MZUSP 39027, 2 c&s,
34.8-40.7 mm SL, Brazil, São Paulo State, Iporanga municipal-
ity; Ituglanis sp., UFRJ 7237, 8, 31.6-40.7 mm SL, Brazil, Pará
State, Primavera municipality, small stream on the road to
Bragança; UFRJ 7238, 2 c&s, 29.8-32.6 mm SL; UFRJ 7239, 2
c&s, 28.3-33.4 mm SL, same locality. Ituglanis sp., MNRJ 37539,
4, 46.6-54.0 mm SL, Brazil, Mato Grosso State, Aripuanã mu-
nicipality, rio Aripuanã, upstream Dardanelos waterfall. Microcambeva ribeirae, MZUSP 78617, paratypes, 5, 41.3-
48.1 mm SL, Brazil, São Paulo State, Pedro de Toledo munici-
pality, rio São Lourencinho; MZUSP 68169, paratypes, 3 c&s,
37.7-41.8 mm SL, Brazil, São Paulo State, Juquiá, ribeirão Poço
Grande. Listrura picinguabae, UFRJ 6111, 1, holotype, 48.6
mm SL, Brazil, São Paulo State, Ubatuba municipality, small
tributary of rio da Fazenda, near Km 11 of the road BR-101 in
Serra do Mar state Park; UFRJ 5951, 4 c&s, 31.3-35.5 mm SL,
São Paulo State, Ubatuba municipality, small tributary of rio da
Fazenda in Serra do Mar state Park. Descobrimento National Park and I. Discussion agreste in the rio de
Contas basin with the type locality close to the recently
created Boa Nova National Park and Boa Nova Wildlife
Refuge. The discovery of this new species in the rio de Contas
basin reinforces the necessity for ichthyofaunal invento-
ries in northeastern Brazil, especially in the numerous iso-
lated coastal watersheds south of rio São Francisco, where
more undescribed species are expected to exist (Langeani
et al., 2009). In the last three years, six new fish species
were described for the rio de Contas basin: Cyphocharax
pinnilepis Vari, Zanata & Camelier, Gymnotus interruptus
Rangel-Pereira, Hasemania piatan Zanata & Serra,
Hyphessobrycon brumado Zanata & Camelier, a distinct
species from H. negodagua that inhabits the rio Paraguaçu
basin, Leporinus brinco Birindelli, Britski & Garavello,
Trichomycterus tete Barbosa & Costa (Vari et al., 2010;
Zanata & Camelier, 2010; Zanata & Serra, 2010; Barbosa &
Costa, 2011; Rangel-Pereira, 2012; Birindelli et al., 2013). Hence, the case of Ituglanis agreste, together with the
recently discovered endemic fish above, suggests that the
rio de Contas basin contains several endemic species
(Zanata & Camelier, 2010). Acknowledgments Ituglanis laticeps, USNM 163907, 2,
56.7-73.2 mm SL, Ecuador, Pastaza, rio Bobonaza. Ituglanis
metae, USNM 100765, 1, 54.3 mm SL, Colombia, Villavivencio. Ituglanis parahybae, UFRJ 761, 1, 43.8 mm SL, Brazil, Rio de
Janeiro State, Silva Jardim municipality, rio São João, close to
Gaviões; UFRJ 702, 6, 41.4-51.4 mm SL; UFRJ 704, 1, 42.1 mm
SL, same locality. Ituglanis paraguassuensis, LIRP 5780, 1, 41.2
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Cyphocharax (Ostariophysi: Characiformes: Curimatidae)
from the Rio de Contas drainage, Bahia, Brazil. Copeia, 2010:
382-387. Wosiacki, W. B., G. M. Dutra & M. B. Mendonça. 2012. Description of a new species of Ituglanis (Siluriformes:
Trichomycteridae) from Serra dos Carajás, rio Tocantins
basin. Neotropical Ichthyology, 10: 547-554. Wosiacki, W. B. & O. T. Oyakawa. 2005. Two new species of
catfish
genus
Trichomycterus
(Siluriformes:
Trichomycteridae) from the rio Ribeira de Iguape Basin,
Southeastern Brazil. Literature Cited Neotropical Ichthyology, 3: 465-472. Zanata, A. M. & P. Camelier. 2010. Hyphessobrycon brumado: a
new characid fish (Ostariophysi: Characiformes) from the
upper rio de Contas drainage, Chapada Diamantina, Bahia,
Brazil. Neotropical Ichthyology, 8: 771-777. Zanata, A. M. & J. P. Serra. 2010. Hasemania piatan, a new
characid species (Characiformes: Characidae) from
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Ichthyology, 8: 21-26. Submitted October 17, 2012
Accepted June 6, 2013 by Marcelo Britto
Published September 30, 2013
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Cross-cultural similarity in relationship-specific social touching
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Cross‑cultural similarity in relationship‑specific
social touching Suvilehto, Juulia T.; Nummenmaa, Lauri; Harada, Tokiko; Dunbar, Robin I. M.; Hari, Riitta;
Turner, Robert; Sadato, Norihiro; Kitada, Ryo Suvilehto, Juulia T.; Nummenmaa, Lauri; Harada, Tokiko; Dunbar, Robin I. M.; Hari, Riitta;
Turner, Robert; Sadato, Norihiro; Kitada, Ryo 2019 Suvilehto, J. T., Nummenmaa, L., Harada, T., Dunbar, R. I. M., Hari, R., Turner, R., . . . Kitada,
R. (2019). Cross‑cultural similarity in relationship‑specific social touching. Proceedings of
the Royal Society B : Biological Sciences, 286(1901), 20190467‑. doi:10.1098/rspb.2019.0467 https://hdl.handle.net/10356/83249 © 2019 The Author(s). Published by the Royal Society under the terms of the Creative
Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits
unrestricted use, provided the original author and source are credited. Downloaded on 24 Oct 2024 13:21:33 SGT Research 1Department of Neuroscience and Biomedical Engineering, 2Department of Computer Science, and 3Department
of Art, Aalto University, Espoo, Finland
4Turku PET Centre, and 5Department of Psychology, University of Turku, Turku, Finland
6Institute of Biomedical and Health Sciences, Hiroshima University, Hiroshima, Japan
7Department of Experimental Psychology, University of Oxford, Oxford, UK
8Department of Neurophysics, Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany
9National Institute for Physiological Sciences, Okazaki, Japan
10Division of Psychology, School of Social Sciences, Nanyang Technological University, 14 Nanyang Avenue,
637332 Singapore 8Department of Neurophysics, Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany
9National Institute for Physiological Sciences, Okazaki, Japan 10Division of Psychology, School of Social Sciences, Nanyang Technological University, 14 Nanyang Avenue,
637332 Singapore 10Division of Psychology, School of Social Sciences, Nanyang Technological University, 14 Nanyang Avenue,
637332 Singapore royalsocietypublishing.org/journal/rspb Juulia T. Suvilehto1,4, Lauri Nummenmaa4,5, Tokiko Harada6,
Robin I. M. Dunbar2,7, Riitta Hari3, Robert Turner8, Norihiro Sadato9
and Ryo Kitada10 Juulia T. Suvilehto1,4, Lauri Nummenmaa4,5, Tokiko Harada6, Research
Cite this article: Suvilehto JT, Nummenmaa
L, Harada T, Dunbar RIM, Hari R, Turner R,
Sadato N, Kitada R. 2019 Cross-cultural
similarity in relationship-specific social
touching. Proc. R. Soc. B 286: 20190467. http://dx.doi.org/10.1098/rspb.2019.0467 Research
Cite this article: Suvilehto JT, Nummenmaa
L, Harada T, Dunbar RIM, Hari R, Turner R,
Sadato N, Kitada R. 2019 Cross-cultural
similarity in relationship-specific social
touching. Proc. R. Soc. B 286: 20190467. http://dx.doi.org/10.1098/rspb.2019.0467 LN, 0000-0002-2497-9757; RIMD, 0000-0002-9982-9702; RK, 0000-0001-7446-4033 Received: 25 February 2019
Accepted: 4 April 2019
Subject Category:
Behaviour
Subject Areas:
behaviour, evolution, neuroscience
Keywords:
social touch, cultural differences, emotion,
bonding
Author for correspondence:
Ryo Kitada
e-mail: ryokitada@ntu.edu.sg
Electronic supplementary material is available
online at http://dx.doi.org/10.6084/m9. figshare.c.4464836. Received: 25 February 2019
Accepted: 4 April 2019 Many species use touching for reinforcing social structures, and particularly,
non-human primates use social grooming for managing their social
networks. However, it is still unclear how social touch contributes to the
maintenance and reinforcement of human social networks. Human studies
in Western cultures suggest that the body locations where touch is allowed
are associated with the strength of the emotional bond between the person
touched and the toucher. However, it is unknown to what extent this
relationship is culturally universal and generalizes to non-Western cultures. Here, we compared relationship-specific, bodily touch allowance maps
across one Western (N ¼ 386, UK) and one East Asian (N ¼ 255, Japan)
country. In both cultures, the strength of the emotional bond was linearly
associated with permissible touch area. However, Western participants
experienced social touching as more pleasurable than Asian participants. These results indicate a similarity of emotional bonding via social touch
between East Asian and Western cultures. Subject Category:
Behaviour
Subject Areas:
behaviour, evolution, neuroscience
Keywords:
social touch, cultural differences, emotion,
bonding 1. Introduction Author for correspondence:
Ryo Kitada
e-mail: ryokitada@ntu.edu.sg Interpersonal touch is a critical part of human social communication. It contri-
butes to cognitive and socioemotional development in childhood [1,2] and
promotes relational, psychological and physical well-being in adulthood [3,4]. Given its importance, there has been growing interest in the effects of interper-
sonal touch on human social behaviour and in the resulting social relationships. Non-human primates spend remarkable amounts of time in grooming others,
well beyond the necessity of removing parasites or vegetation debris from the
fur [5]. Social grooming thus plays a particularly important role in social bond-
ing, and the psychological experience of increased social closeness is reflected in
prosocial behaviours [5,6]. In individual female primates, the social grooming
patterns are explained by factors such as attraction to high-ranking individuals,
attraction to kin and competition for grooming partners [7,8], implying that
variations in the relationship specificity of social touch might be correlated
with differences in social structure. In our previous study, we asked 1368 people from Western countries (Finland,
France, Italy, Russia and the UK) to indicate where on their body they would
allow relatives, friends and strangers to touch them [9]. We also measured the
emotional bond between them since such bonds are the best predictor for Electronic supplementary material is available
online at http://dx.doi.org/10.6084/m9. figshare.c.4464836. & 2019 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution
License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original
author and source are credited. 1Department of Neuroscience and Biomedical Engineering, 2Department of Computer Science, and 3Department
of Art, Aalto University, Espoo, Finland
4Turku PET Centre, and 5Department of Psychology, University of Turku, Turku, Finland
6Institute of Biomedical and Health Sciences, Hiroshima University, Hiroshima, Japan
7Department of Experimental Psychology, University of Oxford, Oxford, UK
8Department of Neurophysics, Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany
9National Institute for Physiological Sciences, Okazaki, Japan
10Division of Psychology, School of Social Sciences, Nanyang Technological University, 14 Nanyang Avenue,
637332 Singapore Physiological Sciences, Japan. The English version of the online
rating tool (emBODY, [9,22]; https://version.aalto.fi/gitlab/egler-
ean/embody) was first translated into Japanese by a professional
translator (ZENIS Co., Ltd, Tokyo Japan). Then two researchers
(R.K. and T.H.) revised the translated materials by conducting
back-translation and translation until both were satisfied with
the result. engaging in social contact with someone and as they track
the position of different individuals in one’s social network
[10–12]. In each country, the topographic map of body areas
that one is allowed to touch was associated with the strength
of the emotional bond between the participant and the toucher. Thus, the relationship-specific patterns of social touch seem
to be related to the establishment and maintenance of social
structures and affective relationships among human adults. 2 Participants first provided background information about
themselves and members of their social network. They were
given a list of categories of relationship which may be found in
one’s social network (e.g. ‘aunt’, ‘female friend’) and participants
then indicated if they had one or more individuals from these
categories in their own social network. If participants had
multiple individuals in their social network fitting one category,
they were instructed to pick one individual and to answer all
the subsequent questions regarding this individual. We also
added ‘female stranger’ and ‘male stranger’ to the list of social
categories, to probe the acceptable social touch with strangers. However, these results cannot resolve whether bonding by
social touch is a culturally universal phenomenon or specific to
Western Europe which our previous study was limited to. Cross-cultural studies have shown that deeply embedded
differences between Western and Eastern cultures strongly
influence emotional processing, together with ideas regarding
experiences such as emotions [13,14], the facially expressed
emotions of others [15–17] and their integration with emotion-
al voices [18]. Indeed, social touch varies according to culture
[19–21]. For instance, North American students have more fre-
quent physical contact than Japanese students with their
friends and their parents [19]. Moreover, the US students’
social touch patterns in regard to their fathers and mothers
were similar, whereas Japanese students were physically
closer to their mothers than their fathers. However, this
study did not examine the relationship between the touching
pattern and the strength of the emotional bonding. Thus, it
remains unclear to which extent this relationship is culturally
universal and generalizes to non-Western cultures. Participants were then asked to give information about the sex
(only for spouses) and ages of the selected members of their social
network, as well as estimates of how long ago they last met them. We assumed that participants would encounter strangers on a
daily basis and set time since the last meeting to 0 days for the
strangers. We used participants’ own age for the age of the stran-
gers, and in subsequent tasks asked partners to respond with
respect to ‘a woman/man of your age whom you do not know’. Next, the participants reported their emotional bond with each net-
work member (scale from 1 representing no emotional bond to 10
representing the strongest possible emotional bond) and gave an
estimate of how pleasant they would find being touched by each
member of their social network (scale from 1, not pleasant at all,
to 10, extremely pleasant). Here we compared relationship-specific social touching
patterns between one East Asian and one European culture. We used a high-resolution self-reporting tool (emBODY) to
quantify relationship-specific maps of bodily regions where
social touch is allowed. Participants in Japan and UK evalu-
ated their emotional bonds with and drew touchable body
regions for different social network members. We hypoth-
esized that patterns of allowed interpersonal touch differ in
Japan and in the UK but vary in a similar way as a function
of social bond in both countries. y p
)
After completing background questions, the participants were
presented with the colouring task with the emBODY tool. They
were asked to consider where on their bodies they would find it
acceptable for different social network members to touch them
in everyday situations. For each member of their social network,
the participants were shown a body outline in both front and
back view (electronic supplementary material, figure S1a) and
asked to colour the bodily regions where each social network
member would be allowed to touch them. The participants
coloured the bodies using a computer mouse or, if they were com-
pleting the task using a mobile device, using their finger. Painting
was additive so that multiple strokes on the same region increased
opacity; this information was, however, not considered in the
analyses (see below). (c) Data analysis
(i) Data preprocessing
Th d
fi
h
k The data were first checked for completeness. Data from the colour-
ing tasks were then converted to Matlab (R2015b) two-dimensional
matrices, where each cell represented a pixel on the screen. The size
of the figures was 522 342 pixels (for front and back combined),
out of which 89 129 pixels fell within the body outlines. The diam-
eter of the painting tool was set to 17 pixels. The coloured images
were binarized so that the amount of time a participant spent on
colouring an area would not impact the results. Each participant
completed between two and 15 individual touch area maps
(TAMs), depending on the size of their social network. As the aim of the study was to compare data between different
cultures, cultural background criteria were used to exclude some
participants. Japanese participants were asked about their parents’
ethnicity, and whether they had spent an extended period of time
abroad. We excluded any participants who did not report both of
their parents’ ethnicity as ‘Japanese’ or who reported having spent
more than a yearabroad. Altogether, 17 Japanese participants were
excluded based on the background questions. In the English (a) Participants
l
h (a) a t c pa ts
Altogether, 309 Japanese individuals and 622 British individuals
participated in the study. The British sample was collected through
Maximiles (data reported in [9]) and the Japanese sample was
collected via MyVoice Communications, Inc., in 2016. As the popu-
lation-level effect size was unknown, we aimed to have the sample
in line with the first study reporting topographies of acceptable
touch (altogether 1368 participants, average 274 per culture, [9]),
which in turn were targeted to double the sample sizes from the
earliest studies using this technology (total of 773 participants
over seven separate experiments, [22]). After cultural background
had been validated and quality control established (described in
detail in the ‘Data analysis’ section), the data of 255 Japanese indi-
viduals (124 male, 40.1+14.6 years) and 386 British individuals
(214 male, 46.0+12.6. years) were analysed. The British sample
was chosen for comparison [9], (1) because it was collected via a
paid service similar to the Japanese sample and (2) because the
age distributions were similar in both groups. royalsocietypublishing.org/journal/rspb
Proc. R. Soc. B 286: 2019046
3 (ii) Comparing the samples using two-proportion z-test
We compared the acceptable touch areas of the two cultural
samples by comparing each pixel in each image using a two-
proportion z-test, with a two-tailed alternative hypothesis, with
a ¼ 0.05, corrected for false detection rate with no correlation
assumptions. To test the association between emotional bond
and touchable body area, we first calculated a ‘touchability
index’ (TI) as the proportion of coloured pixels within the body
outline for each TAM [9]. To better quantify the differences in
the topographies of acceptable touch, we then defined 13 anatom-
ical regions of interest (ROIs) and calculated ROI-specific TIs as the
proportion of coloured pixels within the ROI. We conducted mul-
tiple linear regression analysis on these TIs for each social network
member in both countries, with the emotional bond and cultural
background as explanatory variables. This analysis was conducted
using group mean data and individual data. pleasantness and touchability index pleasantness and touchability index Figure 2 depicts the correlations between TI, pleasantness and
emotional bond, with TI as the proportion of pixels in the body
that a particular member of the participant’s social network
was allowed to touch (electronic supplementary material,
figure S4). Linear multiple regression analysis with emotional
bond, pleasantness of touch and culture as explanatory vari-
ables revealed that together these variables explained 20% of
the variance in the total touchable body area (adjusted R2 ¼
0.20, F3,7375 ¼ 626, p , 10210). Bond (b ¼ 0.18, p , 10210), plea-
santness (b ¼ 0.31, p , 10210) and culture (b ¼ 0.16, p , 10210)
all predicted TI (see also electronic supplementary material,
table S1 for a linear mixed effects model with subject as
random effect). Partitioning R2 assigned 11.3% of the explana-
tory power to pleasantness of touch, 8.6% to emotional bond
and only 0.4% of explanatory power to the cultural background
(electronic supplementary material, figure S5). For the data
averaged such that each social network member was rep-
resented by the average of responses about that person in
each culture (figure 2), the adjusted R2 was 0.85, F3,26 ¼ 58,
p , 10210. (a) Touch area maps for Japanese and British Figure 1 shows the mean TAMs for different social members
in Japanese and British samples. The relationship-specific TAMs in the Japanese and UK
populations were generally consistent. Specifically, the partner
was allowed to touch basically anywhere on the body, and clo-
sest acquaintances and relatives were allowed to touch over the
head and upper torso. By contrast, strangers were restricted to
touch only the hands. Direct comparison of TAMs from British
and Japanese participants by two-proportion z-test revealed
that Japanese allowed more touching from their female rela-
tives than did British, especially in the lower extremities and
the bottom (figure 1c). On the other hand, more British partici-
pants allowed their partners to touch their bodies on the torso,
face and legs. Moreover, British participants allowed their
mother, aunt, female cousin and female friend to touch their
heads more and male strangers to touch their hands more. Due to the high correlation between pleasantness and
emotional bond, we also ran partial correlation tests. When
controlling for pleasantness of the relationship, the association
between emotional bond and TI was still significant, rs7377 ¼
0.15, p , 10210, 95% CI [0.13, 0.17]. Similarly, when controlling
for emotional bond, the association between pleasantness and
TI remained significant, rs7377 ¼ 0.22, p , 10210, 95% CI [0.20,
0.24]. Thus, both pleasantness and emotional bond contributed
independently to the relationship-specific TI. Altogether, the analysis showed that emotional bond and
perceived pleasantness of touch explained around 20% of the
variance in TI, with only a negligible (0.4%) contribution
from the culture. A supplementary analysis with emotional
bond treated as categorical variable confirmed that emotional
bond explains the variance in the total touchable body area
(electronic supplementary material, figure S8). Finally, we
confirmed this result by conducting the same regression ana-
lyses for both sexes in each culture (mean adjusted R2 ¼ 0.21,
range [0.17, 0.28]). Patterns of TI as a function of emotional
bond were consistent regardless of the culture or toucher’s
sex (electronic supplementary material, figure S6). (i) Emotional bond and pleasantness ratings In both countries, an individual’s emotional bond was the
strongest with their partner, followed by their closest family
members and relatives. By contrast, participants reported the
least emotional bond with strangers (electronic supplementary
material, figure S2). The strengths of the reported emotional
bonds with friends were between those for primary and
extended family members in both samples. A Mann–Whitney
U test (after Holm–Bonferroni correction) on emotional
bond yielded a significant difference only for the partner
(U ¼ 13 917, p 8 1029, Holm–Bonferroni corrected), such
that the emotional bond with the partner was lower in Japanese
than in British participants. (b) Data acquisition The data were collected online. Before beginning the study, partici-
pants gave informed consent online. The Japanese study protocol
was approved by the ethics committee at the National Institute for The participants reported that being touched by their
partner elicited most pleasantness, followed by their close
relatives
(electronic
supplementary
material,
figure
S3). Touch pleasantness and emotional bond were significantly
correlated in both cultures, with Spearman correlation coeffi-
cient rs4402 ¼ 0.69 in the British sample and rs2977 ¼ 0.74 in the
Japanese sample ( ps , 10210), The two cultures differed sig-
nificantly in the degree of pleasantness of being touched by
others; with the exception of sister and male stranger, the
British reported finding social touch as more pleasant than
did the Japanese (Mann–Whitney U test, ps in range [3
10216, 0.015], Holm–Bonferroni corrected). version, the sample consisted of only British nationals. The partici-
pants were asked about their native (first) language and their
cultural identity. We excluded all participants who did not report
their first language as being English, or who did not identify them-
selves as ‘British’, ‘English’, ‘Irish’, ‘Welsh’ or ‘Scottish’. Altogether,
159 British participants were excluded based on the background
questions. Finally, we visually inspected the TAMs and excluded
38 participants from Japanese dataset and 77 participants from
British dataset due to inappropriate drawing (e.g. doodling or
excessive colouring). The remaining samples, consisting of 386
British and 255 Japanese participants, were used for analysis. version, the sample consisted of only British nationals. The partici-
pants were asked about their native (first) language and their
cultural identity. We excluded all participants who did not report
their first language as being English, or who did not identify them-
selves as ‘British’, ‘English’, ‘Irish’, ‘Welsh’ or ‘Scottish’. Altogether,
159 British participants were excluded based on the background
questions. Finally, we visually inspected the TAMs and excluded
38 participants from Japanese dataset and 77 participants from
British dataset due to inappropriate drawing (e.g. doodling or
excessive colouring). The remaining samples, consisting of 386
British and 255 Japanese participants, were used for analysis. 3 (c) Sex differences We next examined if sex influences touch acceptance similarly
in the UK and Japan. Figure 3 shows the relationship between
touchable body area and sex of the toucher with respect to male d female participants (blue and red dots) in the UK
d
ll
l
h
ff
f
partners, the sex of the partner was determined by whethe
h
l
f
l
d h
d ff
partner
back
front
back
front
back
front
father
brother
uncle
mother
sister
aunt
acq. acq. friend
friend
cousin
cousin
stranger
stranger
partner
father
brother
uncle
mother
sister
aunt
acq. acq. friend
friend
cousin
cousin
stranger
stranger
partner
father
brother
uncle
mother
sister
aunt
acq. acq. friend
friend
cousin
cousin
stranger
stranger
(a) Japan
(b) UK
(c) difference
proportion of subjects
1.0
0.8
0.6
0.4
0.2
0
proportion of subjects
1.0
0.8
0.6
0.4
0.2
0
10
5
0
–5
–10
UK
Z score
Japan
gure 1. Relationship-specific TAMs in (a) Japan and (b) the UK. The colouring displays the proportion of the sample reporting that being touched by this person
s area is acceptable to them. (c) Comparison of the proportion of participants per culture who allow touching in different areas. Red colour in the maps indicate
t Japanese participants reported that area more acceptable, blue colour indicates that British participants reported that area more acceptable. The data in (c) a
esholded at p , 0.05, FDR corrected. After FDR correction, Z threshold with no correlation assumptions varied from 3.22 to 5.98, depending on the number o
ticipants who had that particular individual in their social network. Red and blue labels indicate female and male members of the social network, respectivel partner
back
front
front
father
brother
uncle
mother
sister
aunt
acq. acq. friend
friend
cousin
cousin
stranger
stranger
partner
father
brother
uncle
mother
sister
aunt
acq. acq. friend
friend
cousin
cousin
stranger
stranger
(a) Japan
(b) UK
proportion of subjects
1.0
0.8
0.6
0.4
0.2
0
ion of subjects
1.0
0.8
0.6
0 4 (c) difference back
front
partner
father
brother
uncle
mother
sister
aunt
acq. acq. friend
friend
cousin
cousin
stranger
stranger
(c) difference
10
5
0
–5
–10
UK
Z score
Japan front back Figure 1. Relationship-specific TAMs in (a) Japan and (b) the UK. The colouring displays the proportion of the sample reporting that being touched by this person in
this area is acceptable to them. (c) Sex differences 0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
1
2
3
4
5
6
7
8
9 10
1
2
3
4
5
6
7
8
9 10
emotional bond
pleasantness
touchability index (TI)
touchability index (TI)
UK
Japan
country 0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
1
2
3
4
5
6
7
8
9 10
emotional bond
touchability index (TI)
UK
Japan
country 0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
1
2
3
4
5
6
7
8
9 10
pleasantness
touchability index (TI) emotional bond variance in TI explained by 1
2
3
4
5
6
7
8
9
10
emotional bond
10
9
8
7
6
5
4
3
2
1
pleasantness pleasantness Figure 2. Correlations between touchability index (TI), emotional bond and pleasantness. Each dot represents the average response for one member of the social
network in each culture (e.g. mother), with linear regression line and confidence interval for the regression fitted separately for each culture. Bottom right panel
presents the relative importance of regressors in determining the TI in a linear model. 0.3
0.2
0.1
0
female
male
female
male
female
male
sex of toucher
sex of
subject
mean TI
UK
Japan
Figure 3. Interaction plot of the average TI for male and female participants (blue and red dots, respectively) with respect to male and female touchers for each
culture. Error bars depict the 95% confidence interval. Red and blue lines indicate the interaction between toucher sex and the sex of the participant. Note: partners
are excluded from the analyses, as the sex of the partner differs between the male and female participants. male
sex of
subject mean TI Figure 3. Interaction plot of the average TI for male and female participants (blue and red dots, respectively) with respect to male and female touchers for each
culture. Error bars depict the 95% confidence interval. Red and blue lines indicate the interaction between toucher sex and the sex of the participant. Note: partners
are excluded from the analyses, as the sex of the partner differs between the male and female participants. (c) Sex differences (c) Comparison of the proportion of participants per culture who allow touching in different areas. Red colour in the maps indicates
that Japanese participants reported that area more acceptable, blue colour indicates that British participants reported that area more acceptable. The data in (c) are
thresholded at p , 0.05, FDR corrected. After FDR correction, Z threshold with no correlation assumptions varied from 3.22 to 5.98, depending on the number of
participants who had that particular individual in their social network. Red and blue labels indicate female and male members of the social network, respectively. and female participants (blue and red dots) in the UK
and Japan. To statistically evaluate the effect of sex on TI,
we conducted an ANOVA on the TIs of male and female par-
ticipants and male and female touchers in both cultures. For partners, the sex of the partner was determined by whether
the participant was male or female, and hence it was difficult
to compare the effect of sex on TI between them. For this
reason, we excluded the partner data from this analysis. 0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9 10
1
2
3
4
5
6
7
8
9 10
emotional bond
emotional bond
10
9
8
7
6
5
4
3
2
1
pleasantness
pleasantness
variance in TI explained by
pleasantness 43.5%
bond 36.1%
country 7.4%
total R2 87.0%
touchability index (TI)
touchability index (TI)
UK
Japan
country
Figure 2. Correlations between touchability index (TI), emotional bond and pleasantness. Each dot represents the average response for one member of the social
network in each culture (e.g. mother), with linear regression line and confidence interval for the regression fitted separately for each culture. Bottom right panel
presents the relative importance of regressors in determining the TI in a linear model. 0.019, such that the parameter estimates were larger for
female touchers (M ¼ 0.18, s.d. ¼ 0.24) than for male touchers
(M ¼ 0.12, s.d. ¼ 0.19, t6739.3 ¼ 11.7, p , 10210, 95% CI [0.050
0.071], d ¼ 0.085). Thus, touch by female members of the
social network was generally considered more acceptable
across the whole social network (electronic supplementary
material, figure S7). The effect of participant sex was not
significant (F1,6945 ¼ 0.72, p ¼ 0.39). (c) Sex differences The grey intercept and slope terms mean no significant difference in the regression lines between the cultures. The coloured terms signal which culture
ad higher intercept (a) or slope (b) values, with blue and red colours indicating the British and the Japanese, respectively. The visualization is presented with
veraged data in the interest of clarity, but the significance of intercept and slope are calculated from the full (un-averaged) data. Proc. R. Soc. B 286: 20190467 differences in
a (intercept) and b (slope):
greater in Japan differences in
a (intercept) and b (slope):
greater in Japan emotional bond Figure 4. Visualization of ROI-specific cultural differences in TI versus emotional bond. Linear regression lines were fitted to each culture separately. Blue and red
colours depict UK and Japan, respectively. Each dot represents the average response for one member of the social network (e.g. ‘British partner’ or ‘Japanese
mother’). The grey intercept and slope terms mean no significant difference in the regression lines between the cultures. The coloured terms signal which culture
had higher intercept (a) or slope (b) values, with blue and red colours indicating the British and the Japanese, respectively. The visualization is presented with
averaged data in the interest of clarity, but the significance of intercept and slope are calculated from the full (un-averaged) data. All interactions were significant at the p , 0.05 significance
level. Of the two-way interactions, the interaction between the
toucher sex and participant sex was significant at F1,6945 ¼ 18.5,
p 2 1025, h2 ¼ 0.003, such that the difference between TI
for female touchers and male touchers was larger for female
participants (M ¼ 0.08, s.d. ¼ 0.11) than for male participants
(M ¼ 0.04, s.d. ¼ 0.09, t592.61 ¼ 5.7, p 2 1028, d ¼ 0.46). The
interaction
between
the
sex
of
the
participant
and the country was significant (F1,6945 ¼ 22.6, p 2 1026,
h2 ¼ 0.003). The TIs reported by Japanese male participants
(M ¼ 0.18, s.d. ¼ 0.26) were higher on average than those
reported by Japanese women (M ¼ 0.15, s.d. ¼ 0.23), British
men (M ¼ 0.14, s.d. ¼ 0.20), or British women (M ¼ 0.16,
s.d. ¼ 0.20), two sample t-test t-scores t2712.5 ¼ 3.8, p ¼ 0.0002,
d ¼ 0.14; t2329.3 ¼ 5.3, p 1027; and t2496.4 ¼ 3.4, p ¼ 0.0006,
d ¼ 0.13
respectively. (c) Sex differences Three-way ANOVA (2 levels of cultures 2 levels of tou-
cher sex 2 levels of participant’s sex) on the TI revealed a
significant main effect of culture F1,6945 ¼ 12.6, p 4 1025,
h2 ¼ 0.002, such that the TIs in the Japanese sample (M ¼
0.17, s.d. ¼ 0.24) were larger than the TIs in the British
sample (M ¼ 0.15, s.d. ¼ 0.20, t5316.8 ¼ 3.38, p ¼ 0.0007, 95%
CI [0.008 0.029], d ¼ 0.085). The main effect of the sex of the
toucher was also significant F1,6945 ¼ 137.9, p , 10210, h2 ¼ 0.019, such that the parameter estimates were larger for
female touchers (M ¼ 0.18, s.d. ¼ 0.24) than for male touchers
(M ¼ 0.12, s.d. ¼ 0.19, t6739.3 ¼ 11.7, p , 10210, 95% CI [0.050
0.071], d ¼ 0.085). Thus, touch by female members of the
social network was generally considered more acceptable
across the whole social network (electronic supplementary
material, figure S7). The effect of participant sex was not
significant (F1,6945 ¼ 0.72, p ¼ 0.39). 1.0
0.8
0.6
0.4
0.2
0
1.0
0.8
0.6
0.4
0.2
0
1.0
0.8
0.6
0.4
0.2
0
1.0
0.8
0.6
0.4
0.2
0
1 2 3 4 5 6 7 8 9 10
1 2 3 4 5 6 7 8 9 10
1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10
emotional bond
Japan
UK
hair
face
shoulder (front)
shoulder (back)
shoulder (back)
shoulder (front)
torso (back)
torso (front)
arm
hand
crotch
bottom
foot
leg (front)
leg (back)
a b
a b
a b
a b
a b
a b
a b
a b
a b
a b
a b
a b
a b
ROI definitions
hair
arms
face
torso
(front)
torso
(back)
crotch
bottom
hands
legs (front)
legs (back)
feet
differences in
a (intercept) and b (slope):
greater in Japan
greater in the UK
no difference
country
touchability index per ROI
igure 4. Visualization of ROI-specific cultural differences in TI versus emotional bond. Linear regression lines were fitted to each culture separately. Blue and red
olours depict UK and Japan, respectively. Each dot represents the average response for one member of the social network (e.g. ‘British partner’ or ‘Japanese
mother’). (c) Sex differences The
interaction
between
the
tou-
cher sex and country was significant (F1,6945 ¼ 5.0, p ¼ 0.025,
h2 ¼ 0.001). The TIs for Japanese female touchers (M ¼ 0.20,
s.d. ¼ 0.26) were on average higher than those for British
female touchers (M ¼ 0.17, s.d. ¼ 0.22) (t2758.4 ¼ 3.7, p ¼ 0.0003, d ¼ 0.13). The three-way interaction was also significant
(F1,6945 ¼ 25.0, p 6 1027, h2 ¼ 0.003). This result seems to
be mostly driven by the different responses of Japanese
men and women with respect to male touchers (figure 3). Simi-
lar results were obtained using a mixed-effects model with
subject as a random effect (electronic supplementary material,
table S2). (d) Region-of-interest analysis (d) Region-of-interest analysis Whole-body TAM analyses revealed cultural differences in
touchability of specific body areas, such as face, hand, and
arm. To further examine for area-specific cultural differences,
we next conducted linear regression analyses on the regional
TIs for each culture, with emotional bond as the explanatory
variable (figure 4). Adjusted R2 values of fitted linear func-
tions averaged across body regions ranged between 0.04
and 0.15. Comparing linear models fitted to ROI-wise TI
data showed significantly higher baseline acceptability (inter-
cept) of touch on hair (b ¼ 20.02, t7376 ¼ 22.96, p ¼ 0.003), feet (b ¼ 20.05, t7376 ¼ 27.3, p , 10210), legs (both front,
b ¼ 20.03, t7376 ¼ 25.3, p 9 1028, and back, b ¼ 20.03,
t7376 ¼ 25.7, p 2 1028), crotch (b ¼ 20.01, t7376 ¼ 23.0,
p ¼ 0.003) and bottom (b ¼ 20.03, t7376 ¼ 25.1, p 3
1027) in the Japanese sample. The same test showed higher
baseline acceptability of touch on arms (b ¼ 0.06, t7375 ¼ 3.4,
p ¼ 0.0006), hands (b ¼ 0.14, t7375 ¼ 7.2, p , 10210) and face
(b ¼ 0.07, t7376 ¼ 9.3, p , 10210) in the British sample. The
same test showed a different rate of increase in TI for each
unit of emotional bond (slope) on the arms (b ¼ 20.01,
t7375 ¼ 23.5, p ¼ 0.0004) and hands (b ¼ 20.02, t7375 ¼ 27.3,
p , 10210). In both of these ROIs, the TI was more responsive
to changes in emotional bond (steeper slope) in the Japanese
sample. However, a cross-sectional design cannot reveal whether
pleasure derived from being touched by someone enhances
the emotional bond to the toucher, or whether liking someone
makes their touch feel more pleasant. 7 Social touch is important for group cohesion of non-human
primates, and grooming relationships predict the level of
support during conflict situations and when in need of help
[8,31–34]. It is hypothesized that grooming-induced relaxed
and pleasurable feelings could constitute the psychological
mechanism for an individual’s willingness to offer subsequent
help, thus forming the basis of mutual exchanges of social sup-
port
[5,34]. Because social touching increases prosocial
behaviour [35–37], it is possible that human social touch
increases emotional bonding with the individual being
touched, enhancing their prosocial behaviour. (a) Gender difference in both cultures One possible explanation for the cross-cultural similarities
in social touching is the effect of globalization. For instance,
Japanese culture may be influenced by social customs in
e.g. Western movies that are shown widely in Japan. How-
ever, the relationship specificity of touchable areas (found
here in European and Japanese cultures) is consistent with
both our earlier study with Western cultures [9] as well as
an older study looking at Japanese and North American
cultures [19]. Thus, it is unlikely that the role of recently
shared cultural elements plays a major role in the cross-cultu-
rally similar association between social touch and emotional
bond. Participants in both cultures allowed women to touch their
bodies more than men, which accords with prior findings in
Western cultures [9,47]. The preference to touch by women
was comparable in British men and women. By contrast,
responses of the Japanese participants showed a clear gender
difference: Japanese men did not show a statistically significant
preference for touch by women, whereas Japanese women
showed a remarkably strong preference. Earlier studies have found opposite-sex touch to be prefer-
able to same-sex touch [19,23,48–50]. Preference for female
touch in the female participants might therefore seem to con-
tradict earlier findings. On closer inspection, this seeming
contradiction is caused by a difference in terminology. Most
of the earlier studies discussing gender effects have inspected
the difference of body accessibility for ‘same-sex friend’ and
‘opposite-sex friend’. Unfortunately, most of the older exper-
iments [23,48–50] use the euphemism ‘opposite-sex friend’ to
indicate a romantic partner (a usage made explicit in [51]),
whereas ‘same-sex friend’ seems to refer to a platonic friend. This practice biases the comparison, as touching romantic part-
ners is both qualitatively and quantitatively different from
touching other people [9]. Social touch induces positive feelings and improves
interpersonal evaluation [24–27]. For example, participants
evaluate even a stranger more positively, if that person
has inconspicuously touched them during an interaction
[25,27–29]. This close connection between social touching and
preference toward the source of touch suggests a causal role
of touch on social bonding. There is some evidence of touching
causally impacting bonding in romantic relationships [30]. However, it is not clear if this extends to other relationships
and this cannot be directly addressed from the current cross-
sectional data. We also found that emotional bonds are
correlated with experienced pleasantness of social touch. (d) Region-of-interest analysis Neurophysiological studies have revealed that the un-
myelinated afferent c-tactile nerve fibres are selectively
responsive to light and slow touch [38–40]. The signals car-
ried out by these fibres are indirectly transmitted to the
insula [38], a part of the neural network responsible for affec-
tive tactile interactions [40,41]. This pathway could thus
support culturally universal hedonic nature of interpersonal
touch. However, as shown in the present study, relationship
information imposes top-down influences on this circuitry
(e.g. how the individuals are touched and by whom they
are touched, electronic supplementary material, figure S3). It is also known that physical contact with members of
one’s social network attenuates brain activity associated
with both the threat of physical pain stimulus [42] and aver-
sive images [43]. Processing interpersonal aspects of touch
may recruit additional brain networks [44,45], including the
action observation network (AON) [46]. Thus, the positive
and calming effect of interpersonal tactile interchanges may
be generated by the interaction of these brain networks, med-
iating the relationship between touchable area and emotional
bonding, regardless of the different cultural norms. 4. Discussion The results show that although the touchable areas differed
between the UK and Japan, they were similarly dependent
on the strength of the emotional bond. Emotionally close
individuals in
the inner layers of the social
network
were allowed to touch larger bodily areas, whereas touching
by strangers was primarily limited to the hands. The ROI
analyses confirmed that most of the ROIs were similarly
sensitive to emotional bond in both countries. This result
suggests that the use of social touch for bonding purposes
serves a similar function in East Asian and European
cultures, rather than being merely culture-based normative
behaviour. This
interpretation
accords
with
previous
studies [9,19,23] suggesting cultural invariance in social
touching. We have previously reported consistent social touching pat-
terns in a large sample of European cultures [9]. Additionally,
American and Japanese college students were asked where
they had touched and been touched by their parents, a same-
sex friend and an opposite-sex friend [19]. The relative
frequencies of touch by these different members of the social
network followed the same pattern in both cultures [19]. (c) Limitations
O
d (c) Limitations Our study was conducted online. Because the participants did
not meet the experimenter, they could answer the questions
without the sense of invasion of privacy (e.g. feeling embar-
rassed that experimenters in front of them know their
touching behaviour). While there is some experimental evi-
dence
suggesting
that
attitudes
towards
touch
impact
touching behaviour [55], it is possible that the data may not
directly translate to real-life touching behaviour. Therefore, it
will be important to validate the current findings by conduct-
ing observational research of real-life touching in the future. Second, although the currently available evidence suggests
that social touching is concordant across a wide range of Wes-
tern, Orthodox and Japanese cultures (see also [9]), these data
do not generalize to the other major cultures of the world [56]. However, already the present data demonstrate marked cross-
cultural consistency in touch-dependent bonding across a wide
range of cultures and geographical locations. Second, we also observed three notable differences in the
topographies of acceptable touch. The British participants
allowed their partners to touch their bodies more than the
Japanese (figure 1). This finding accords with previous research
showing that touch between partners is more frequent in most
body areas in the Western (North American) culture than in
Japan [19]. The British participants in the present study also
reported stronger emotional bonds with their partners than
did the Japanese participants, suggesting that the difference
between touch allowances for the partner might not be merely
a cultural norm in behaviour but indicative of a wider difference
in the intimate relationships in the two cultures. Moreover, British participants allowed female family mem-
bers and female friends to touch their faces more than did the
Japanese participants. A previous study demonstrated that
the British tend to touch their own faces more frequently than
the Japanese do [53]. It is possible that such ‘accessibility’ to
face may originate from gestures that are more specific to British
than Japanese and their self-touching behaviour. By contrast,
more Japanese participants reported that their female relatives
are allowed to touch them on their legs and bottoms. Female
members in the Japanese culture often take care of children in
their network and have physical contact with them. It is con-
sidered that British culture is more individualistic than Japan
and that individuals in such culture assume responsibility
only for themselves and their immediate family [54]. (c) Limitations
O
d Thus,
the observed difference may be explained by the roles of the
female relatives in child rearing in the two cultures. (b) Cultural differences between the Japanese
and the British While the overall association of touching and social bonding
was concordant across cultures, some differences were also
found. First, the Japanese participants reported the overall
pleasantness of being touched to be lower than the British
did (electronic supplementary material, figure S3), although
both cultures showed similar changes in emotional bond
across different touchers (figure 2). This difference may be in
part due to differences in wording (‘kokochiyoi’ in Japanese
versus pleasant in British), but it can also reflect differences
in daily non-verbal communication: Japanese conduct social
touching (e.g. handshakes and hugs) much less frequently
than Americans [19] and they do likely less than British. Funding. This work was supported by Emil Aaltonen Foundation grant
and Alfred Kordelin Foundation grant to J.T.S.; by Academy of
Finland grant (no. 294897) to L.N. and European Research Council
grants to L.N. (no. 313000) & R.I.M.D. (no. 295663), by MEXT/JSPS (a) Gender difference in both cultures Taking this difference of nomenclature into account, our
findings would in fact seem to be in line with earlier studies. For example, a similar preference for female touch has pre-
viously been found with respect to parents: participants
touch and are touched more by their mothers than fathers,
both in the United States [23,52] and in Japan [19]. Similar
to the current findings, both male and female Japanese
participants touch and are touched by their mothers more
than by their fathers, but the difference is greater for female
participants [19]. Conversely, in the American sample, the
preference for mother over father was of comparable size
between male and female participants [19]. such as handshakes and hugs, are more common in Western
countries than in East Asian countries, such as Japan. This is
supported by the higher intercepts in arm and hand for the
UK sample in the ROI-wise analysis. Thus, more emotional
closeness may be necessary for the Japanese to engage in
this type of social touching. 8 Finally, the difference between the touch allowances for
female touchers and male touchers was larger for Japanese
women than for Japanese men or British participants of
either sex (culture sex of toucher sex of participant inter-
action, figure 3). This culture-specific gender difference can
be also associated with the above-mentioned points. More
specifically, Japanese women had much smaller TIs for all
males in their social network than for women with similar
formal relationship (e.g. mother and father, see electronic sup-
plementary material, figure S7). Collectively, although we
found some differences, the associations between emotional
bond, pleasantness and TI were concordant in both cultures. Competing interests. The authors declare no conflict of interest. Acknowledgements. We thank Enrico Glerean for his help in collecting
the data. We acknowledge the computational resources provided
by the Aalto Science-IT project. 5. Conclusion Relationship-specific emotional bonds account for the magni-
tude of social touching from social network members
similarly between Western and East Asian (Japanese) cul-
tures. Pleasure derived from touch, however, depended on
the culture. Because the relation-specific social touching pat-
terns of Japanese and Western respondents are consistent,
the relationship between emotional bonding and touchable
body area may be largely biologically determined. Human
social touch, like social grooming of non-human primates,
may provide a scaffold for social bonding with the members
of one’s social network. This supports the role of somatosen-
sation and emotional feelings in the maintenance of social
bonds in humans [57]. We also observed cultural differences in the relationship
between the emotional bond and the touchable area in
specific body parts such as the arms and hands. In these
regions, the strength of the emotional bond with a social net-
work member increased touchability more in the Japanese
versus British sample, i.e. the Japanese had a steeper slope
in the fitted linear function (figure 4). This difference might
be
explained
in
terms
of
cultural
difference
in
daily
gestures. For instance, gestures involving physical contacts, Data accessibility. The data are available from the Dryad Digital Reposi-
tory: https://doi.org/10.5061/dryad.f0f2642 [58] and the code to
produce all the figures and numbers are available at https://ver-
sion.aalto.fi/gitlab/jtsuvile/cultural-similarity-touch. Funding. This work was supported by Emil Aaltonen Foundation grant
and Alfred Kordelin Foundation grant to J.T.S.; by Academy of
Finland grant (no. 294897) to L.N. and European Research Council
grants to L.N. (no. 313000) & R.I.M.D. (no. 295663), by MEXT/JSPS Acknowledgements. We thank Enrico Glerean for his help in collecting
the data. We acknowledge the computational resources provided
by the Aalto Science-IT project. 9 KAKENHI (grant no. 26244031; 15H01846) to N.S. and (grant no.
16H01680; 25135734) to R.K.; and by an NAP start-up grant of
Nanyang Technological University to R.K. 1177/0956797611403) 19. Barnlund DC. 1975 Public and private self in Japan
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РОЗРОБКА ЗАВДАНЬ МАЙБУТНІМИ ВЧИТЕЛЯМИ МАТЕМАТИКИ ЯК ЗАПОРУКА ЇХ ПРОФЕСІЙНОГО ЗРОСТАННЯ
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Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя
Випуск 21
УДК 373.51:377.1(075)
Олександр Манченко
здобувач другого (магістерського) рівня вищої освіти
Харківський національний педагогічний університет імені Г.С. Сковороди
https://orcid.org/0000-0002-5378-3841
Євгеній Войтов
здобувач другого (магістерського) рівня вищої освіти
Харківський національний педагогічний університет імені Г.С. Сковороди
https://orcid.org/0000-0001-5259-7863
Оксана Жерновникова
доктор педагогічних наук, професор,
завідувач кафедри математики
Харківський національний педагогічний університет імені Г.С. Сковороди
https://orcid.org/0000-0002-5383-4493
РОЗРОБКА ЗАВДАНЬ МАЙБУТНІМИ ВЧИТЕЛЯМИ
МАТЕМАТИКИ ЯК ЗАПОРУКА ЇХ ПРОФЕСІЙНОГО
ЗРОСТАННЯ
Анотація. Статтю присвячено проблемі розробки та створення
математичних
завдань
майбутніми
вчителями
математики.
Проаналізовано провідний досвід підгоовки майбутніх вчителів
математик з Сполучених Штатів Америки щодо створення завдань на
уроках математики. Виділені характерні особливості створення та
підготовки математичних завдавань для підвищення ефективності
занять.
Ключові слова: математичні завдання, учні, вчителі математики Випуск 21 удосконалення професійної підготовки майбутнього вчителя РОЗРОБКА ЗАВДАНЬ МАЙБУТНІМИ ВЧИТЕЛЯМИ
МАТЕМАТИКИ ЯК ЗАПОРУКА ЇХ ПРОФЕСІЙНОГО
ЗРОСТАННЯ Анотація. Статтю присвячено проблемі розробки та створення
математичних
завдань
майбутніми
вчителями
математики. Проаналізовано провідний досвід підгоовки майбутніх вчителів
математик з Сполучених Штатів Америки щодо створення завдань на
уроках математики. Виділені характерні особливості створення та
підготовки математичних завдавань для підвищення ефективності
занять. Ключові слова: математичні завдання, учні, вчителі математики Олександр Манченко Олександр Манченко
здобувач другого (магістерського) рівня вищої освіти
Харківський національний педагогічний університет імені Г.С. Сковороди
https://orcid.org/0000-0002-5378-3841 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Науково-дослідна робота студентів як чинник Ключові слова: математичні завдання, учні, вчителі математики Математичні завдання є важливою частиною навчання, оскільки
вони є інструментами, які вчителі використовують, щоб допомогти
учням засвоїти певні математичні концепції [7]. Завдання «визначають
не тільки те, що (учні) вивчають, але й те, як вони починають думати
про математику, розвивати, використовувати та розуміти її» [9]. Крім
того, завдання, які здобувачі виконують на своїх курсах математики,
«передають знання про те, що таке математика і що передбачає заняття 95 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього Випуск 21 у
р
у
удосконалення професійної підготовки майбутнього вчителя з математики» (Національна рада викладачів математики) [5, с. 24]. Оскільки «те, що учні вивчають, фундаментально пов’язане з тим, як
вони це вивчають» [6, с. 18], типи завдань, які вчителі пропонують
своїм учням для виконання, визначають навчальні можливості учнів. Таким чином, завдання, які вчителі використовують під час навчання,
повинні бути вартими того, щоб витрачати навчальний час, не тільки
тому, що вони впливають на те, як учень повинен мислити [10], що
впливає на те, як учні навчатимуться, але також тому, що вони
впливають на те, як учні сприймають природу математики. Розробка математичних завдань є одним із найважливіших
обов’язків учителів математики [5]. Математичні завдання можуть бути
більш-менш корисними для розвитку математичного розуміння залежно
від характеристик, якими вони володіють. Проте дослідники виявили,
що багато вчителів включають у свої програми завдання, які, на думку
дослідників, мають меншу цінність, ніж хотілося б. Наприклад, Jacobs,
J. та ін. [3] проаналізували відеодослідження TIMSS 1995 та 1999 років,
щоб побачити, якою мірою вчителі математики у восьмих класах США
викладали або не викладали згідно з стандартів, встановленими
національною агенцією у 1995, 2000 роках. Ці дослідники вважали, що
математичні завдання та те, як вони використовувалися під час уроків, є
важливим показником того, чи вчителі викладають на основі стандартів. Більшість вчителів у своєму дослідженні використовували завдання, які
не були складними і які можна було розв’язати за допомогою звичайної
процедури рішення, що означає, що завдання були вправами, а не
завданнями, які варті уваги. Багато вчителів, які брали участь у
дослідженні, вірили, що вони викладають згідно з духом реформи
освіти, але це не так. Отже, ймовірно, що багато вчителів зараз не
розробляють варті уваги завдання, щоб включити їх у навчання, навіть
якщо вони думають, що вони це роблять. Оскільки ми знаємо, що вчителі-практики рідко включають варті
уваги математичні завдання у свої уроки, ми очікуємо, що вони не
навчилися розробляти варті уваги завдання під час свого навчання. Ключові слова: математичні завдання, учні, вчителі математики Розглядаючи приклад Сполучених Штатів Америки можна зазначити,
що на допомогу майбутнім вчителям математики прийшли курси
підвищення кваліфікації. На цих курсах майбутні вчителі математики 96 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбут Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчител Випуск 21 удосконалення професійної підготовки майбутнього вчителя покращували свої навички розробки математичних завдань та
навчалися, як розробляти різнорівневі завдання для учнів з різною
підготовкою. Цей досвід став першим кроком до вдосконалення їхніх
навичок проектування завдань на якісно новому рівні. покращували свої навички розробки математичних завдань та
навчалися, як розробляти різнорівневі завдання для учнів з різною
підготовкою. Цей досвід став першим кроком до вдосконалення їхніх
навичок проектування завдань на якісно новому рівні. Мета цього дослідження полягає в тому, щоб описати, розробку
математичних завдань, як один з факторів підвищення професійного
рівня майбутніх учителів математики. Дізнатися, на що звертають увагу майбутні учителі математики,
коли вони намагаються розробити важливі математичні завдання, є
цінним, оскільки це дає педагогам конкретну інформацію, яку вони
можуть використати для роботи над покращенням свого професійного
рівня. Здатність розробляти важливі математичні завдання стане
неоціненним надбанням для майбутніх учителів математики, коли вони
почнуть викладати, оскільки ця важлива навичка дасть їхнім учням
доступ до знань, які дозволять їм розвинути глибоке розуміння
математики. Математичне завдання – це «заняття в класі, метою якого є
зосередити увагу учнів на певній математичній ідеї» [9]. Прикладами
математичних завдань є проекти, задачі, програми та вправи [5]. Є три способи, якими майбутні учителі математики можуть
розробляти математичні завдання. По-перше, вчителі можуть вибрати
завдання з доступних ресурсів. Ці ресурси можуть включати
підручники, які вони використовують, інші навчальні матеріали або
Інтернет. По-друге, вчителі можуть адаптувати вже наявні завдання. Часто вчителі можуть знайти завдання, дослідити його та змінити
завдання, щоб воно відповідало їхнім потребам. По-третє, вчителі
можуть створити завдання з нуля як унікальний продукт який поєднує
найкращі практики математичного проектування. У цьому дослідженні
розробка завдання стосуватиметься всіх цих трьох підходів: вибору,
адаптації та створення. Релевантні
математичні
завдання
знайомлять
студентів
з
важливими математичними ідеями та залучають їх до них. У
професійних стандартах зазначено, що варте уваги математичне
завдання має ґрунтуватися на «надійній та значущій математиці» [5, с. 25], а Hiebert J. та ін. [2] стверджував, що відповідні завдання повинні 97 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Випуск 21 вимагати від студентів роздумів над важливими математичними ідеями
і тільки тоді вони зможуть творчо імплементувати їх у свою майбутню
професійну діяльність вчителя математики. Ключові слова: математичні завдання, учні, вчителі математики Окрім змісту завдання,
головна увага в літературі також приділяється міркуванню, яке вимагає
від учнів чіткого розуміння поставленої математичної цілі завдання. Незважаючи на те, що автори можуть описувати це міркування,
використовуючи іншу термінологію, таку як вирішення проблем або
когнітивна діяльність, очевидно, що життєво необхідно, щоб здобувачі
думали, щоб вивчати математику з розумінням. Професійні стандарти
стверджують, що завдання мають бути інтелектуально захоплюючими
та вимагати від здобувачів міркування та вирішення проблем у різний
спосіб [5]. Hiebert та ін. [2] стверджував, що відповідні завдання
повинні бути справжньою проблемою для здобувачів, оскільки вони ще
не знають, як їх вирішити. На мислення, яке потрібне учням для виконання завдання,
впливають їхні попередні знання та досвід. Завдання мають бути
складними, але доступними. Smith M. та Stein M. [8] припустили, що
вчитель повинен враховувати вік, рівень класу, попередні знання та
досвід своїх учнів, вирішуючи які завдання надавати для роботи у класі. Сміт і Штейн пояснили, що якби вчитель дав шестикласнику завдання,
де він повинен скласти п’ять двозначних чисел і пояснив, як він це
зробив, учень, швидше за все, використав би алгоритм і перерахував
кроки, через які він пройшов, щоб його вирішити. Однак, якби вчитель
дав задачу другокласнику, який не знав алгоритму та мав доступні
знання, учень, швидше за все, досліджував би математичні поняття. Варте завдання вимагає від здобувачів використання наявних знань для
вирішення нового завдання [2] і «створює потребу» [1, с. 318] у новій
концепції, вимагаючи від них побудови нових знань. Є й інші характеристики, які допомагають зробити завдання
вартим уваги. Професійні стандарти підготовки у США включали такі
визначення: гідне завдання має передати, що математика є постійною
діяльністю людини, вимагати від учнів спілкування про математику,
допомагає учням встановлювати зв’язки між математичними ідеями та
розвивати навички учнів разом із розумінням цих явишь та процесів. Професійні стандарти [5] (також запропонували, щоб вчителі зважали 98 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Науково-дослідна робота студентів як чинник
ф
ій
ї і
йб Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Випуск 21 на рівень здобувачів при розробці завдань, рекомендуючи, щоб варті
завдання базувалися на «знанні розуміння, інтересів і досвіду учнів» і
«знанні ряду способів, за допомогою яких різні учні вивчають
математику. ‖ (с. 25), і що корисні завдання повинні розвивати в учнів
схильність до математики через розуміння, а не заучування. Smith M. та
Stein M. Ключові слова: математичні завдання, учні, вчителі математики [8] також заявили, що завдання повинні вимагати від учнів
глибокого мислення, але не повинні бути надто складними для учнів,
щоб учні відчули, що вони можуть залучитися до завдання. Важливим фактором підвищення якості освіти є впровадження
креативності у завдання, що значно підвищує зацікавленість здобувачів
у вивченні різних предметів [4]. Таким чином, можна констатувати, що розробка математичних
завдань є важливою складовою підвищення професійного рівня
майбутніх учителів математики в умовах постійного розвиваючихся
педагогічних технологій навчання. Математичні завдання це комплекс
дій у класі які направлені на розуміння математичних догм та знань
через сприйняття інформації у посильній мисленевій діяльності учнів. У майбутніх наукових розвідках планується дослідити питання
впровадження диференційованих математичних завдань у процесі
опанування основ математики. Список використаних джерел 1. Confrey J. Learning to see children's mathematics: Crucial challenges
in constructivist reform. In K. Tobin (Ed.), The practice of
constructivism in science education (pp. 299- 321). 1993. Hillsdale,
NJ: Lawrence Erlbaum Associates. 2. Hiebert J., Carpenter T. P., Fennema E., Fuson K. C., Wearne D.,
Murray H., . . . Human P. Making sense: Teaching and learning
mathematics with understanding. 1997. Portsmouth, NH: Heinemann. 3. Jacobs J. K., Hiebert J., Givvin K. B., Hollingsworth H., Garnier H., &
Wearne D. Does eighth-grade mathematics teaching in the United
States align with the NCTM Standards? Results from the TIMSS 1995
and 1999 video studies. Journal for Research in Mathematics
Education. 2006. 37. P. 5-32. 99 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя
Випуск 2 Науково-дослідна робота студентів як чинник
удосконалення професійної підготовки майбутнього вчителя Випуск 21 4. Nalyvaiko O., & Ronzhes O. Creativity in digital environment. possibilities of application in the educational space. Electronic
Scientific Professional Journal ―Open Educational E-Environment of
Modern
University‖. 2021. (11). P. 106-120. https://doi.org/10.28925/2414-0325.2021.119 4. Nalyvaiko O., & Ronzhes O. Creativity in digital environment. possibilities of application in the educational space. Electronic
Scientific Professional Journal ―Open Educational E-Environment of
Modern
University‖. 2021. (11). P. 106-120. https://doi.org/10.28925/2414-0325.2021.119 5. National Council of Teachers of Mathematics. Professional standards
for teaching mathematics. 1991. Reston, VA: Author. 5. National Council of Teachers of Mathematics. Professional standards
for teaching mathematics. 1991. Reston, VA: Author. 6. National Council of Teachers of Mathematics. Mathematics teaching
today (2nd ed.). 2007. Reston, VA: Author. 6. National Council of Teachers of Mathematics. Mathematics teaching
today (2nd ed.). 2007. Reston, VA: Author. 7. Simon M. A. & Tzur R. Explicating the role of mathematical tasks in
conceptual learning: An elaboration of the hypothetical learning
trajectory. Mathematical Thinking and Learning. 2004. 6. P. 91-104 8. Smith M. & Stein M. Selecting and creating mathematical tasks: From
research to practice. Mathematics Teaching in the Middle School. 1998. 3. P. 344-350. 9. Stein M., Grover B. & Henningsen M. Building student capacity for
mathematical thinking and reasoning: An analysis of mathematical
tasks used in reform classrooms. American Educational Research
Journal. 1996. 33. P. 455-488. 10. Zhernovnykova O. A., Nalyvaiko O. O. & Chornous N. A. Intelectual competence: essence, components, levels of formation. Pedagogy
and
Psychology. 2017. 58. P. 32-41. doi:
10.5281/zenodo.1116977. O. Manchenko, Y. Voitov, O. Zhernovnykova Annotation. The article is devoted to the problem of developing
and creating mathematical tasks by future teachers of mathematics. The leading experience of training future teachers of mathematics
from the United States of America regarding the creation of tasks in
mathematics lessons was analyzed. Characteristic features of 100
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O estágio supervisionado nos cursos de graduação em Educação Física: um desafio presente nesta formação
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Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. O estágio supervisionado nos cursos
de graduação em Educação Física:
um desafio presente nesta formação Enny Vieira Moraes
Professora assistente – UESB;
Doutoramento em História Social – PUC. São Paulo – SP[Brasil]
ennymoraes@hotmail.com
Alantiara Peixoto Cabral
Graduada em Educação Física – Universidade
Estadual do Sudoeste da Bahia. Luanda Nogueira Souza
Graduada em Educação Física – Universidade
Estadual do Sudoeste da Bahia. Manuela Soares de Alcântara
Graduada em Educação Física – Universidade
Estadual do Sudoeste da Bahia. Alantiara Peixoto Cabral
Graduada em Educação Física – Universidade
Estadual do Sudoeste da Bahia. Luanda Nogueira Souza
Graduada em Educação Física – Universidade
Estadual do Sudoeste da Bahia. Manuela Soares de Alcântara
Graduada em Educação Física – Universidade
Estadual do Sudoeste da Bahia. Neste artigo, discute-se o estágio supervisionado, como discipli
na curricular obrigatória, nas licenciaturas em Educação Física,
além dos desafios presentes na relação entre formação profissio
nal e ética na escola pública. Essa proposta amplia-se quando
a ótica adotada para discutir tais desafios envolve não apenas a
visão de docentes, mas também a visão dos discentes. Concluiu-
se que o estágio, um momento desafiador na formação, produz a
consciência da importância do profissional da educação e de sua
participação como agente questionador da realidade. Palavras-chave: Estágio supervisionado. Formação profissional. História da Educação Física Palavras-chave: Estágio supervisionado. Formação profissional. História da Educação Física Artigos 199 Outros autores, no século XIX, já escreviam
sobre os exercícios físicos ou ginástica, como afirma
Cunha Júnior (1998, p. 26): A prática de ensino nos cursos de licen
ciatura em Educação Física traz à tona
a dificuldade dos graduandos – que, por
vezes, reflete a de alguns professores – inte
grarem prática e teoria que, apesar de não
serem idênticas, possuem interdependência
(GONÇALVES JÚNIOR e RAMOS, 1998)1. Os professores civis também atuaram na
escrita de manuais e livros sobre Educação
Física/Ginástica, durante o século XIX. A
formação profissional do grupo ocorreu
em suas vivências nas escolas e colé
gios em que haviam freqüentado aulas
de Educação Física/Ginástica. Eram ex-
alunos que, ao se destacarem nas aulas,
recebiam convites de seus mestres e passa
vam posteriormente a ministrá-las. Os professores civis também atuaram na
escrita de manuais e livros sobre Educação
Física/Ginástica, durante o século XIX. A
formação profissional do grupo ocorreu
em suas vivências nas escolas e colé
gios em que haviam freqüentado aulas
de Educação Física/Ginástica. Eram ex-
alunos que, ao se destacarem nas aulas,
recebiam convites de seus mestres e passa
vam posteriormente a ministrá-las. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 1
Introdução A dissociação entre teoria e prática na
Educação Física brasileira foi, durante anos, um pro
blema que se refletia e, talvez se reflita até hoje, mais
profundamente, na prática pedagógica dos profissio
nais de ensino. Como conseqüência dessa ruptura,
identificamos problemas em relação ao aspecto pe
dagógico da área, que consistem num desafio que se
apresenta e que se evidencia, primordialmente, nas
disciplinas de estágio curricular obrigatório, presen
te nas licenciaturas desses cursos. Também nesse período, publicações começa
ram a surgir no País, principalmente em formato
de manuais que “ensinavam” atividades físicas, pu
blicados por autores nacionais e internacionais, que
auxiliaram a despertar o interesse da população pela
prática das atividades físicas, como afirma Cunha
Júnior (1998) sobre Arthur Higgins, ex-jornalista
que passou a se dedicar aos exercícios ginásticos, na
condição de autor e professor. É nesse contexto que
surge a Educação Física no Brasil. Assim, até hoje responder a questões, tais
como o que é o estágio em Educação Física? Qual seu
conteúdo específico? Quais as orientações e discus
sões pedagógicas que baseiam o estágio curricular
em Educação Física? São desafios não apenas para
o profissional que está presente no espaço escolar,
mas também para aquele em processo de formação. É nesse sentido que, com base em algumas expe
riências de docentes e discentes no âmbito formal,
buscamos sugerir direcionamentos introdutórios
para essa discussão. Assim, a Educação Física surge, como uma
prática corporal, impulsionada pela atuação de
diferentes atores sociais – civis e mesmo imigran
tes –, que não tinham, necessariamente, qualquer
tipo de formação e, cuja prática tinha como base a
sua própria experiência e, normalmente, ocorriam
em diferentes locais, com objetivos e com base em
elementos distintos para sustentar essas mesmas
práticas. Pode-se dizer, então, que o surgimento da
Educação Física brasileira tem como reflexo a po
larização entre o conhecimento teórico e prático e
que essa dicotomia caracterizou historicamente essa
área por muitos anos. Desde que foi introduzida obrigatoriamen
te nas escolas brasileiras, no fim dos anos 1920, a
Educação Física caracterizou-se pela fragmentação
entre o conhecimento da produção da medicina hi
giênica e a prática dos profissionais que atuavam
nas escolas e em outros espaços (MELO, 1996). Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 200 É sabido, entretanto, que a produção teórica
da área dará um salto qualitativo no Brasil, durante
os anos 1970 e 1980. 1
Introdução Entretanto, como mostra
Daolio (1995), os professores de Educação Física
que atuavam nas escolas durante esse período, não
percebiam claramente a importância dessa disci
plina no currículo escolar. Apesar de a atuação
desses docentes parecer diferenciada em relação
às demais disciplinas, em sua visão, eles mesmos
ainda não apresentam uma argumentação precisa
para definir sua área ou mesmo sua ação peda
gógica. Essa indefinição se reflete também nos
currículos dos cursos de graduação em Educação
Física, ou melhor, na ausência de clareza sobre
essa formação, demonstrando também a distância
entre as discussões teóricas que se apresentavam e
sua implementação no espaço escolar. fissionais que tinham como característica basilar a
fragmentação de seu conhecimento. Realmente, existe um grande desafio a ser
enfrentado na formação desses profissionais no que
diz respeito a uma efetiva união entre o fazer pe
dagógico, a clareza de sua opção metodológica e a
reflexão filosófica desse fazer – complexidade que se
apresenta, de forma mais transparente, nas discipli
nas de estágio curricular ou estágio supervisionado. A Resolução CNE/CP 1, de 19 de fevereiro de
2002, publicada em 4 de março de 2002, contribuiu
para que as disciplinas de estágio e as licenciatu
ras, de modo geral, sofressem importantes altera
ções tanto em relação à ampliação da carga horária
quanto à definição dos espaços de atuação dos es
tágios curriculares que, anteriormente, não eram
definidos claramente. De acordo com a nova resolução e em con
sonância com o Parecer 109/2002 do Conselho
Nacional de Educação, homologado em 9 de maio
de 2002 e publicado em 13 de maio de 2002 (sessão
1, p.21), no que se refere ao estágio curricular: Embora esse não seja o foco de nossa dis
cussão, é importante refletir sobre as diversas
concepções (higienista, militarista, tecnicista,
crítico-superadora, entre outras) que marcaram a
formação dos professores de Educação Física. Essas
concepções nortearam, durante anos, a formação
desses profissionais, e contribuíram para criar as
lacunas pedagógicas dessa prática, o que se reflete
no cotidiano do espaço escolar. Neste estudo, o
objetivo não é afirmar que essas correntes histori
camente não foram importantes; entretanto, é pos
sível observar que, pedagogicamente, não deram
respostas suficientes a questões que se encontram
na base e no surgimento da Educação Física brasi
leira – no caso específico deste trabalho, à prática
pedagógica dessa formação. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 1
Introdução Cada Instituição de Ensino Superior, por
tanto, deverá incluir no seu projeto pedagó
gico como componente curricular obrigató
rio, o estágio curricular supervisionado de
ensino como um momento de capacitação
em serviço de 400 horas, que deverá ocorrer
em unidades escolares onde o estagiário, ao
final do curso, assuma efetivamente, sob su
pervisão, o papel de professor. Portanto, ficam definidos a carga horária
mínima para os estágios e os espaços em que deverão
ocorrer. No caso da Educação Física, como tal for
mação, mesmo em licenciatura, incluía, em alguns
cursos de nível superior, experiência nos espaços A ausência de definição do compromisso po
lítico-social dos profissionais da área desnudou um
problema central: a necessidade de unir o conheci
mento teórico com a prática pedagógica desses pro Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. Artigos Artigos 201 cenário exige dos discentes soluções ou alternativas de
soluções, permeadas de dúvidas (científicas e políti
cas), constituindo-se num momento importante para
qualquer formação profissional. não-formais, esse aspecto também sofreu alteração. Antes dessa resolução legal a docência poderia ser
exercitada nas escolas e também em outros espaços
como academias de ginástica, clubes etc. o que muda
com a Lei é que os estágios, em Educação Física, vão
se resumir apenas aos espaços formais. O futuro profissional deve ser levado a perce
ber no estágio a possibilidade de se ver não como um
manipulador de instrumentos ou simples executor
de atividades, mas como construtor de propostas que
contribuirão no processo de formação daqueles que
se encontram no espaço escolar. Com base na refle
xão concebida teoricamente, são abertas perspecti
vas para o futuro educador exercer conscientemente
posturas mais críticas no exercício de sua prática. Além de tudo, é importante que busque extrapolar
a repetição, o que permitirá a identificação dos pro
blemas que permeiam suas atividades e a fragilida
de da ação no exercício da prática pedagógica. Para os estágios, o grande desafio na for
mação é ser eixo articulador entre a capacitação
teórico-científica e a atuação político-pedagógica
dos futuros profissionais. No entanto, é importante
lembrar que essa responsabilidade, especialmen
te em relação à qualidade da formação, não deve
nem pode se limitar apenas a uma ou duas disci
plinas de estágio soltas no currículo, mas precisa
estar presente em todos os momentos da graduação;
caso contrário, apenas os estágios responderiam às
demandas para essa formação, especificamente em
se tratando das licenciaturas. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 1
Introdução O estágio supervisionado realizou-se de 23 de
outubro a dezembro de 2007 nas turmas de 1° ano
do ensino médio, do período vespertino, formadas,
em média, por 35 alunos cada uma, com os alunos
e alunas de uma faixa etária entre 15 a 16 anos. Convém salientar que, no fluxograma do
curso de licenciatura plena em Educação Física da
Universidade Estadual do Sudoeste da Bahia (UESB),
campus Jequié, as disciplinas que oferecem um
contato com o futuro local de trabalho, analisam e
questionam os conhecimentos adquiridos durante o
processo de ensino-aprendizagem, na busca de uma
possível transformação do ambiente, são as discipli
nas de Estágio Supervisionado I, II e III, com as te
máticas: I (Atividades Físicas e Saúde), II (Educação
Física Escolar) e III (Educação Física e Lazer). No momento do estágio, o acompanhamento,
a supervisão e a orientação – atividades exercidas
pelos professores orientadores – são fundamen
tais. Esses profissionais, que exercem eticamente
seu papel, servem de referência, não no sentido da
cópia, mas pelo norte inicial de uma prática pro
fissional pautada na seriedade e no compromisso No momento do estágio, o acompanhamento,
a supervisão e a orientação – atividades exercidas
pelos professores orientadores – são fundamen
tais. Esses profissionais, que exercem eticamente
seu papel, servem de referência, não no sentido da
cópia, mas pelo norte inicial de uma prática pro
fissional pautada na seriedade e no compromisso
– elementos que, infelizmente, nem sempre estão
presentes no setor educacional, de um modo geral. Entretanto, o reconhecimento dessa disciplina na
formação é inquestionável. – elementos que, infelizmente, nem sempre estão
presentes no setor educacional, de um modo geral. Entretanto, o reconhecimento dessa disciplina na
formação é inquestionável. É importante frisar que os Estágios
Supervisionados I e III eram realizados em espaços
não-formais, porém, com a nova resolução, os con
teúdos dessas disciplinas passaram a ser trabalha
dos também na escola. Mesmo com inúmeros problemas e dificul
dades que precisam ser superadas, o estágio é im
portante no currículo tanto dos discentes quanto
dos docentes (SOUZA; BONELA; PAULA, 2007). Por
isso, abriu-se espaço, neste estudo, para a expla
nação de relato de experiência de três discentes do
curso de licenciatura plena em Educação Física da
Universidade Estadual do Sudoeste da Bahia (Uesb),
campus Jequié. Lembrando sempre que, para a efe
tivação do estágio, de fato, é necessário que ele seja
construído no cotidiano da escola, ouvindo todos
os atores. 1
Introdução Nesse sentido, concor
damos com Piconez (1991, p. 25), ao afirmar: Sabe-se que as dificuldades são muitas, prin
cipalmente porque não há um fazer pedagógico
pronto, completo. Essa disciplina tem como carac
terística “o não ser/estar pronta”, ou seja, “o fazer-se
constante na constância e na dinâmica do cotidiano
do espaço escolar”. É nessa dinâmica, que é aquilo
que trava, interrompe, amadurece, cria e recria pos
sibilidades de um refazer e refazer-se constantemen
te, que o estágio deve nortear-se. Acreditamos que a problematização da
prática desenvolvida coletivamente pelas di
ferentes disciplinas do currículo, portanto,
articuladas, podem assegurar a unidade,
favorecer a sistematização coletiva de novos
conhecimentos e preparar o futuro profes
sor para compreender os determinantes
mais profundos de sua prática, com vistas a
sua possível transformação. Além dos problemas que envolvem o fazer pe
dagógico nas disciplinas de estágio, outros, de ordem
prática, também se apresentam, entre os quais o
número insuficiente de docentes na supervisão e
no acompanhamento dos discentes no campo de
estágio. Essa ausência transformada em impessoali
dade gerou uma série de dificuldades, como afirma
Silva (2005, p. 37): Nesse processo, é importante que as disciplinas
do currículo levem o futuro profissional a questionar
sua formação, a ter dúvidas e assumi-las de forma
mais concreta. Já nas disciplinas que enfocam prática
de ensino e o estágio supervisionado, esse questio
namento surge, principalmente, no contato com a
realidade contextual dinâmica, plural e contraditó
ria que se instala, propondo desafios concretos. Esse Já se tornou tradicional em nosso país
que a orientação de estágio seja encarada
pelas IES como um procedimento mera Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 202 mente burocrático, cuja ação do profes
sor-orientador é a cobrança e o controle
de entrega de cartas de credenciamento
de instituições concedentes de estágio,
fichas de seguro, cômputo de horas
de estágio e relatórios de estágio […]
Paradoxalmente, a impessoalidade acaba
tornando-se a tônica de um processo ide
alizado para que o graduando construa
sua futura identidade profissional, onde a
personalização do acompanhamento da
ação e reflexão sobre a realidade vivida
deveria vir em primeiro lugar. – reflexão – ação, num movimento constante, pois
a dinâmica da realidade exige esse esforço. A experiência relatada ocorreu no Colégio da
Rede Pública Estadual, Luiz Eduardo Magalhães2,
situado na cidade de Jequié (BA), e que oferece
ensino médio nos três turnos: noturno, vesperti
no e matutino. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 1
Introdução Nesse processo, os discentes e docentes
precisam estar em constante diálogo sobre a ação Após essa discussão, propôs-se a inclusão,
neste trabalho, de um relato da experiência dos
discentes no encerramento da disciplina Estágio
Supervisionado I, cujo conteúdo é atividade física
e saúde. Esse relato foi uma sugestão de trabalho
com o conteúdo programático dessa disciplina. Como foi produzido depois de finalizada a discipli
na, não houve aferição de nota ou do conceito dado
aos discentes e nem foi atribuído sentido avaliativo. Essa produção está associada ao compromisso do Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. Artigos Artigos 203 Desse modo, a disciplina de Estágio
Supervisionado I, do curso de Licenciatura Plena
em Educação Física da UESB, que será tratada no
relato de experiência, tem como finalidade propor
cionar aos discentes experiências pedagógicas, que
lhes possibilitarão planejar, executar e avaliar situa
ções de ensino-aprendizagem, durante as atividades
físicas e de saúde, buscando focalizá-las como um
importante conteúdo a ser trabalhado nas aulas de
educação física. grupo em produzir conhecimento sobre o próprio
fazer pedagógico. Assim, o relato que consiste na experiência
de três discentes do curso de Educação Física da
UESB foi resultado de um esforço que envolveu a
participação de todos e, certamente, servirá de re
ferência na formação dessas futuras licenciadas em
Educação Física. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 2
Relato de experiência próprio professor regente não dá a devida impor
tância a essa etapa, pulando-a e deixando que os
estagiários se responsabilizem por seus discentes. Acreditamos que esse período seja essencial para a
formação de futuros educadores, sendo necessário o
comprometimento de ambas as partes. As disciplinas de estágio supervisionado no
currículo são divididas em duas etapas: aulas na uni
versidade e no campo de estágio (na escola) que, por
sua vez, subdivide-se em três momentos: observação,
co-participação e regência. No primeiro momento
– observação – os estagiários se familiarizam com
o colégio, conhecem sua estrutura e a(as) turma(as)
a ser(em) trabalhada(s) e, por meio das observações
das aulas do professor, analisam a prática docente
e suas implicações na formação do sujeito. No
segundo – co-participação – são vivenciadas experi
ências docentes, de forma integrada e participativa,
com o professor regente. Por fim, a regência, em que
os estagiários assumem verdadeiramente a respon
sabilidade pela turma, desenvolvendo atividades de
docência adquiridas durante o curso, tais como pla
nejamento, procedimentos metodológicos, seleção e
aplicação de materiais didáticos, além da operacio
nalização dos projetos desenvolvidos durante a dis
ciplina, particularmente nas orientações práticas,
por meio do contato com o docente da disciplina na
universidade e o supervisor externo. Em relação à co-participação, é possível per
ceber seu valor durante todo o processo de estágio,
em que a atuação era conjunta (professor e estagiá
rias) e, em cada momento de interação em que uma
nova aprendizagem ocorria, contribuindo qualitati
vamente para a futura atuação docente. Durante a regência, aprende-se que, ao entrar
em uma sala de aula, é preciso ter segurança e
pensar que, independentemente de qualquer coisa,
não se deve esquecer de que sempre há uma troca
constante de aprendizado entre estagiários e alunos,
ou seja, cada um traz suas experiências pessoais
para serem re-passadas, re-discutidas, re-pensadas. Nesse período também se compreende
quanto é importante relacionar os conhecimentos
a serem transmitidos para os alunos com os adqui
ridos na vida cotidiana, e assim associar a necessi
dade de adquirir esses conhecimentos para usá-los
na vida diária. Nessa perspectiva, entende-se que
é indispensável investigar, no processo educativo,
a situação individual e social do grupo de alunos,
os conhecimentos e experiências que trazem, de
modo que, nas situações didáticas, ocorra a ligação
entre os objetivos e conteúdos propostos pelo pro
fessor e as condições de aprendizagem dos alunos
(LIBÂNEO, 1994). 2
Relato de experiência Logo no início, verificamos algumas difi
culdades nessa disciplina, como os problemas para
encontrar as turmas pela falta de horário, uma vez
que, nas escolas estaduais do município, as de 1° e
2° anos têm apenas duas aulas semanais de educa
ção física, e as de 3° ano, apenas uma. Percebemos,
assim, por essa e outras questões específicas de
nossa realidade, que a educação física é uma dis
ciplina pouco valorizada no currículo escolar, pois,
em algumas escolas, há apenas uma aula de educa
ção física semanal. O curso de Licenciatura Plena em Educação
Física da Universidade Estadual do Sudoeste da
Bahia (UESB), campus de Jequié, tem 11 anos de
existência e propõe uma formação generalista aos
seus estudantes. Em seu fluxograma, apresenta
disciplinas do conhecimento biológico, profissiona
lizante, socioantropológico, filosófico, psico-peda
gógico, lingüístico e científico. Dessas as de estágio
supervisionado tem por objetivo aproximar o gradu
ando da realidade escolar, possibilitando que viven
cie experiências de sua futura profissão, o que lhe
possibilitará conhecer melhor sua área de atuação. Essa prática é de fundamental importância, pois in
troduz o futuro professor no âmbito escolar, com o
intuito de integrar teoria e prática pedagógica, favo
recendo o desenvolvimento profissional. Após encontrar as turmas, outra dificuldade
foi trabalhar com Atividade Física e Saúde, conte
údo a ser abordado no Estágio Supervisionado I,
pois o professor regente – e supervisor externo – já
havia abordado esse conteúdo com suas turmas. Desse modo, foram ministradas poucas aulas com
a temática proposta, uma vez que o professor au
torizou realizar o estágio nas turmas, desde que
fosse dada continuidade às atividades planejadas e
desenvolvidas por ele. Nesse sentido, concordamos com Pimenta
(2004) quando afirma que, […] no estágio dos cursos de formação
de professores, compete possibilitar que os
futuros professores compreendam a com
plexidade das práticas institucionais e das
ações aí praticadas por seus profissionais
como alternativa no preparo para sua in
serção profissional. Em razão disso, foi preciso reorganizar o
estágio e adequá-lo ao planejamento do docente que
estava preparando uma gincana com jogos popula
res, com todas as turmas dos primeiros anos do res
pectivo colégio. Diante dessa situação, observou-se
que a universidade deveria dialogar mais com a co Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 204 munidade externa, neste caso a escola, para tentar
minimizar esses problemas. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 2
Relato de experiência A observação do estágio Supervisionado I foi de
fundamental importância para o processo de ensino-
aprendizagem, pois proporcionou um entrosamento
com a turma e com o professor regente. Além disso,
possibilitou que se criticassem as atitudes do professor,
tanto dentro quanto fora da sala de aula. Por meio
dessa análise, foi possível absorver certos compor
tamentos didáticos, considerados necessários para
um docente, que resultaram em uma reflexão mais
apurada sobre a prática educativa, como também
maneiras de atuar no espaço escolar. De acordo com essa experiência, também
se percebe que o planejamento e a metodologia de
ensino são de suma importância para o proces
so educativo, e que as aulas nem sempre ocorre
rão exatamente da forma planejada. Ainda com
relação ao mesmo autor, concordamos com seu É sabido que, muitas vezes, os estagiários se
restringem a apenas “observar” as aulas, esque
cendo de analisá-las criticamente. Outras vezes, o Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. Artigos Artigos 205 ponto de vista, quando afirma que o planejamen
to tem a função de orientar a prática pedagógica,
porém não pode ser um documento rígido e abso
luto, pois uma das características do processo de
ensino é que está sempre em movimento, sofrendo
modificações ante as condições reais. só as dimensões biológicas dos indivíduos, mas
também, as dimensões sociais e pedagógicas,
sensibilizando as pessoas sobre a importância de
adotar um estilo de vida ativo. Ao longo de sua história, a Educação Física
teve uma relação direta com a saúde, seja voltada
para a educação higiênica e eugênica da popula
ção, ou para o desenvolvimento da aptidão física. Sua inclusão nos currículos escolares ocorreu com
fortes influências médicas, que pretendiam restau
rar a saúde da população, adotando hábitos higiê
nicos com uma visão eminentemente biologicista
do homem. Em seguida, vieram as influências
militar e desportiva, também com uma visão bio
lógica, como afirma Moraes (1996). Diante disso, podemos verificar que todas
as etapas foram de importância incalculável para
o processo de ensino-aprendizagem, e que, a
cada momento, no ambiente escolar, tentávamos
extrair o máximo de cada aluno, contribuindo,
assim, para discussão sobre a formação daqueles
futuros profissionais. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 2
Relato de experiência 206 quantidade mínima de professores supervisores ou
mesmo sua insuficiência em relação ao número de
estagiários, ou até mesmo falta de compromisso de
alguns professores, são algumas das queixas fre
qüentemente apresentadas pelos alunos. na formação dos estudantes, para que possam ser
cidadãos que reivindiquem sobre seus direitos e re
flitam sobre eles buscando melhorias para a popu
lação que integram e são sujeitos. Atualmente há nas escolas, quando muito,
duas aulas de Educação Física de 50 minutos cada
uma. Diante disso, constata-se que, para um estu
dante, é muito difícil obter os benefícios de uma
prática de atividade física apenas em aulas regula
res. Por isso, acredita-se que o principal objetivo do
professor esteja focalizado na reflexão do conteúdo
trabalhado e em seus métodos. Nesse contexto, seria
possível criar situações em que os alunos possam,
com a população, reivindicar seus direitos para a
melhoria de condições de vida, incluindo aí políticas
públicas para uma melhor qualidade de saúde. Nesse sentido, Silva (2005) relata que, quando
a experiência dos estágios curriculares é bem orien
tada, seguramente, gera não apenas benefícios ao
graduando, mas também serve de instrumento de
avaliação, retroalimentação e aperfeiçoamento do
próprio curso de graduação. Dessa forma, percebemos que a presença e o
acompanhamento dos supervisores são fundamen
tais, uma vez que suas orientações e apoio con
tribuíram para a construção de outras formas de
práxis pedagógicas, além de auxiliar nos estudos e
na superação das dificuldades que sempre se apre
sentaram durante todo o processo de estágio. Assim, não se trata de desvirtuar a prática de
atividade física como componente que exerce influ
ência sobre a melhoria do estado de saúde, quando
praticada corretamente. No entanto, é preciso não
se deixar levar por afirmações simplistas como a
de que, atividade física, por si só, é capaz de gerar
saúde. A atividade física deve ser compreendida
como um componente de inúmeros fatores impres
cindíveis (capacidade econômica, habitação, tempo
livre, acesso a escolarização, cultura, nutrição ade
quada etc), que concorrem para promoção de saúde. (SCHNEIDER, 2001). Acredita-se ainda que seja necessário estar
na escola no começo do ano letivo participando
da semana pedagógica e de outras atividades que
fazem parte da prática docente, auxiliando desde
seu planejamento, até o desenvolvimento das ati
vidades sem interromper a proposta do docente. É sabido que, para isso, várias barreiras devem
ser superadas. 2
Relato de experiência Assim, tanto alguns docentes, professores
das universidades e de instituições dos diversos
graus de ensino, quanto os próprios estagiários
precisam mudar sua maneira de ver o processo de
ensino-aprendizagem, buscando aproveitar todas
as oportunidades que surgem, para construir uma
prática educativa realmente comprometida com o
meio social do qual todos nós integramos. No entanto, acreditamos que o professor deva
superar essas visões, abordando a questão da ativi
dade física relacionada à saúde, de forma diferencia
da, não levando em consideração apenas o aspecto
biológico. O professor deve levar em conta que o ser
humano é resultado de inúmeras determinações
que vão desde o aspecto biológico, o sócio-histórico,
e cultural. De outro modo, e como historicamen
te fizemos, estaríamos apenas reproduzindo uma
visão única da realidade. Quanto ao conteúdo relativo à atividade
física e saúde, ou mesmo do lazer e da educação
física escolar, acreditamos ser fundamental que
os profissionais o discutam anteriormente, para
que sua aplicação na escola ocorra efetivamente
– de forma crítica. Especificamente os conteú
dos da área da atividade física e saúde, poucas
pessoas, principalmente as oriundas das classes
desfavorecidas socialmente, não têm muito
acesso a esse tipo de informação, e a escola pode
ser uma instituição social importante para levar
esse conhecimento aos atores presentes no espaço
escolar. Nesse sentido, a Educação Física se torna
a maior responsável pela transmissão desse con
teúdo, pois o tem enquanto conteúdo curricu
lar, e o acesso a esse conhecimento no ambiente
escolar se faz necessário para contemplar não Quanto ao conteúdo relativo à atividade
física e saúde, ou mesmo do lazer e da educação
física escolar, acreditamos ser fundamental que
os profissionais o discutam anteriormente, para
que sua aplicação na escola ocorra efetivamente Diante disso, concordamos com Matsudo et al. (2003), quando afirmam que não basta apenas falar
para as crianças e adolescentes praticarem atividade
física, o que trará vários benefícios para sua saúde,
mas também deixar claro que vários fatores influen
ciam na execução dessa prática, contribuindo para
que os estudantes possam refletir sobre a sociedade,
questionando as informações adquiridas. Portanto,
é importante que, em aulas de Educação Física, os
estudantes possam analisar o sistema em que a so
ciedade vive e suas possíveis formas de intervenção. Acredita-se que o professor, agindo dessa
forma, poderá contribuir, de maneira significativa, Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. The supervised training in the
postgraduate studies in physical
education: a challenge of this process The supervised training in the
postgraduate studies in physical
education: a challenge of this process sibilitaram maior reflexão sobre a prática docente,
além de reforçar nossa percepção de que sua execu
ção vai muito além das salas de aula. É importante ressaltar que essa instituição de
ensino em que se realizou este estágio se diferen
cia das outras e, provavelmente, é essa participação
dos docentes que faz a diferença, não apenas da
Educação Física, mas também de outras áreas, nas
citadas reuniões e em outras atividades da escola. A atuação dos profissionais desta instituição é um
exemplo do interesse e do compromisso de todos,
agindo realmente coletivamente. In this article, it is discussed the supervising
period of probation as a compulsory discipline
in the Physical Education bachelor’s degree,
besides the challenges of the relation between
professional training and ethics in the public
school. This proposal gets bigger when the point
of view to discuss these challenges involves not
only the academician vision, but also the stu
dents’ point of view. Then, it is concluded that
the period of probation, a challenger moment
in the graduation, produces the conscience of
the importance of the professional’s education
and his participation as an agent that analyses
the reality. 3
Tentando concluir… Fica registrada, portanto, com essa experi
ência, que a participação dos futuros professores
em espaços escolares contribui significativamente
para sua identidade profissional, e que, apesar das
indefinidas dificuldades encontradas, o estágio
auxilia expressivamente em sua formação, pois é
só por meio da vivência no meio escolar que conse
guiremos constituir uma base de formação profis
sional e pessoal. O estágio é, como afirma Pimenta
(2004, p. 34), “[…] uma atitude investigativa
que envolve a reflexão e a intervenção na vida da
escola, dos professores, dos alunos e da sociedade.” Key words: History of Physical Education. Professional training. Supervised probation. 2
Relato de experiência Uma delas é levar os estudantes a
entender que o meio escolar ultrapassa as ativi
dades realizadas em salas de aulas, chegando aos
corredores em conversas com professores, alunos
e funcionários. Desse modo, a Educação Física, como dis
ciplina escolar, não pode abrir mão do conteúdo
atividade física e saúde em sua proposta pedagógi
ca nas escolas. No entanto, essa incorporação não
pode ocorrer acriticamente e, muito menos, des
vinculada da realidade. Apesar dos impasses verificados durante o
estágio, muitos aspectos importantes foram deba
tidos nas reuniões de planejamento pedagógico
(aulas de coordenação – AC), que ocorreram sema
nalmente, possibilitando um aprendizado significa
tivo pela troca de conhecimentos entre os estagiários
e os docentes mais experientes, que integravam o
quadro de efetivos da escola. Esses momentos pos Sobre o supervisionamento dos estágios3,
muitas vezes, eles não acontecem de maneira
correta, ficando o estudante, por diversos motivos,
sem orientação e supervisão adequadas. Uma Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. Artigos Artigos 207 Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. Referências CUNHA JÚNIOR, C. F. F. A produção teórica brasileira
sobre educação física/ginástica no século XIX: autores,
mercado e questões de gênero. In: NETO, A. F. (Org.). v. 3. Pesquisa histórica em educação física brasileira. Aracruz, ES: Faculdade de Ciências Humanas de
Aracruz, 1998, p. 26. MORAES, E. V. A presença do ideário higienista no
curso de formação em Educação Física da Universidade
Federal de Alagoas. Dissertação de mestrado,
Universidade Gama Filho, 1996. DAOLIO, J. Da cultura do corpo. Campinas, São Paulo:
Papirus, 1995. Notas 1 Embora de forma sintética, nesse trabalho surgem
importantes questionamentos sobre o estágio
como espaço reconstrutivo da ação pedagógica
em educação física e, por esse motivo, sugerimos
sua consulta. 2 Gostaríamos de agradecer a todos os profissionais
desse estabelecimento de ensino, seus funcionários
e alunos pela forma como nos receberam e nos
trataram durante todo o percurso do estágio -
espaço e local onde todos podemos refletir, discutir,
recriar e aprender. Outro aspecto importante é percebermos que
essa disciplina se faz em seu percurso, embora, an
teriormente, tentemos buscar definir sua estrutura,
seus objetivos e ações. No entanto, outra análise é
possível e necessária: se todos se comprometerem,
será possível construir uma experiência rica, não
apenas para aqueles em formação, mas também
para os docentes, alunos e demais envolvidos, pois é
necessário compreender que somos todos incomple
tos, eternos aprendizes. 3 Como foi citado, existe a figura do supervisor
externo de estágio, aquele que se encontra no
espaço escolar, e o outro, que é o docente professor
da disciplina estágio supervisionado, ocupante
dessa cadeira na universidade. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. 208 MELO, V. A. de. Escola Nacional de Educação Física e
Desportos: um estudo histórico, a ‘história’ de um estudo
e o estudo da história. In NETO, A. F. (Org.). Pesquisa
histórica em educação física brasileira. Vitória: UFES. Centro de Educação Física e Desportos, 1996. Dialogia, São Paulo, v. 7, n. 2, p. 199-209, 2008. DAOLIO, J. Da cultura do corpo. Campinas, São Paulo:
Papirus, 1995. PIMENTA, S.G. L.; LUCENA, M.S. Estágio e docência. São Paulo: Cortez, 2004. FAZENDA; RIBEIRO; BIZZO et al. A prática de ensino e o
estágio supervisionado. 2.ed. Campinas: Papirus, 1994 SILVA, S. A. P. S. Estágios curriculares na formação
de professores de educação física: o ideal, o real e o
possível. Revista Digital, Buenos Aires, ano 10, n. 82,
mar. 2005. GONÇALVES J. L.; RAMOS, G. N. S. A prática de ensino e
o estágio supervisionado nos cursos de licenciatura em
Educação Física. Revista da Unicastelo, São Paulo,
v. 1, n. 1, p.13-15, 1998. GONÇALVES J. L.; RAMOS, G. N. S. A prática de ensino e
o estágio supervisionado nos cursos de licenciatura em Educação Física. Revista da Unicastelo, São Paulo,
v. 1, n. 1, p.13-15, 1998. SOUZA, J. C. A.; BONELA, L.A.; PAULA, A. H. de. A
importância do estágio supervisionado na formação
do profissional de educação física: uma visão docente e
discente. MOVIMENTUM. Revista digital de Educação
Física – Ipatinga; Unileste- MG, v. 2, nº 2, ago, dez. 2007. SCHNEIDER, O. Educação Física como promoção da
saúde: Contradições de um discurso. Revista Brasileira
de Ciências do Esporte, v. 21, n. 1, 2001. SOUZA, J. C. A.; BONELA, L.A.; PAULA, A. H. de. A
importância do estágio supervisionado na formação
do profissional de educação física: uma visão docente e
discente. MOVIMENTUM. Revista digital de Educação
Física – Ipatinga; Unileste- MG, v. 2, nº 2, ago, dez. 2007. LIBÂNEO, J. C. Didática. São Paulo: Cortez, 1994. LIBÂNEO, J. C. Didática. São Paulo: Cortez, 1994. MATSUDO, V. K. R.; ANDRADE, D. R.; MATSUDO, S. M. M.; ARAUJO, T. L.; ANDRADE, E.; OLIVEIRA, L.C.;
BRAGGION, G; RIBEIRO, M. A. “Construindo” saúde
por meio da atividade física em escolares. R. Bras. Ci e
Mov. Brasília, DF, v. 11, n.4, p.111-118, out./dez. 2003. SCHNEIDER, O. Educação Física como promoção da
saúde: Contradições de um discurso. Revista Brasileira
de Ciências do Esporte, v. 21, n. 1, 2001. Artigos
recebido em jun. 2008 / aprovado em ago. 2008
Para referenciar este texto:
MORAES, E. V. et al. O estágio supervisionado
nos cursos de graduação em educação física: um
desafio presente nesta formação. Dialogia, São
Paulo, v. 7, n. 2, p. 199-209, 2008. recebido em jun. 2008 / aprovado em ago. 2008 Para referenciar este texto:
MORAES, E. V. et al. O estágio supervisionado
nos cursos de graduação em educação física: um
desafio presente nesta formação. Dialogia, São
Paulo, v. 7, n. 2, p. 199-209, 2008. Artigos 209
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EFFECT OF INOCULATION OF Acetobacter diazotrophicus (ACN) WITH DIFFERENT LEVELS OF NITROGEN ON THE YIELD OF SUGARCANE VARIETY CoC-92061.
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Distribution of Paramagnetic Fe2O3/SiO2–Core/Shell Nanoparticles in the Rat Lung Studied by Time-of-Flight Secondary Ion Mass Spectrometry: No Indication for Rapid Lipid Adsorption
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Received: 10 July 2018; Accepted: 24 July 2018; Published: 26 July 2018 Abstract: Amorphous silica nanoparticles comprise a class of widely used industrial nanomaterials,
which may elicit acute inflammation in the lung. These materials have a large specific surface to
which components of the pulmonary micro-milieu can bind. To conduct appropriate binding studies,
paramagnetic Fe2O3/SiO2 core/shell nanoparticles (Fe-Si-NP) may be used as an easy-to-isolate
silica surrogate, if several prerequisites are fulfilled. To this end, we investigated the distribution
of Fe, Si, protein and phosphatidylcholine (PC) by Time-of-Flight secondary ion mass spectrometry
(ToF-SIMS) in cryo-sections from the rat lungs to which Fe-Si-NP had been administered for
30 min. Regions-of-interest were identified and analyzed with incident light and enhanced
dark-field microscopy (DFM). Fe-Si-NP particles (primary particle size by electron microscopy:
10–20 nm; aggregate size by tracking analysis: 190 ± 20 nm) and agglomerates thereof were mainly
attached to alveolar walls and only marginally internalized by cells such as alveolar macrophages. The localization of Fe-Si-NP by DFM was confirmed by ToF-SIMS signals from both, Fe and Si
ions. With respect to an optimized signal-to-noise ratio, Fe+, Si+, CH4N+ and the PC head group
(C5H15NO4P+) were the most versatile ions to detect iron, silica, protein, and PC, respectively. Largely congruent Fe+ and Si+ signals demonstrated that the silica coating of Fe-Si-NP remained stable
under the conditions of the lung. PC, as a major lipid of the pulmonary surfactant, was colocalized
with the protein signal alongside alveolar septa, but was not detected on Fe-Si-NP, suggesting that
silica nanoparticles do not adsorb lipids of the lung surfactant under native conditions. The study
shows that ToF-SIMS is a valuable technique with adequate spatial resolution to analyze nanoparticles
together with organic molecules in the lung. The paramagnetic Fe-Si-NP appear well suited to study
the binding of proteins to silica nanomaterials in the lung. Keywords:
ToF-SIMS; dark-field microscopy; lipid adsorption; amorphous silica; core/shell
nanoparticles; lung tissue; protein corona formation; nanotoxicology; rat nanomaterials nanomaterials Lothar Veith 1 ID , Antje Vennemann 2, Daniel Breitenstein 1, Carsten Engelhard 3,
Birgit Hagenhoff 1 and Martin Wiemann 2,* Lothar Veith 1 ID , Antje Vennemann 2, Daniel Breitenstein 1, Carsten Engelhard 3,
Birgit Hagenhoff 1 and Martin Wiemann 2,* 1
Tascon GmbH, Mendelstraße 17, 48149 Münster, Germany; lothar.veith@tascon-gmbh.de (L.V.);
daniel.breitenstein@tascon-gmbh.de (D.B.); birgit.hagenhoff@tascon-gmbh.de (B.H.) daniel.breitenstein@tascon-gmbh.de (D.B.); birgit.hagenhoff@tascon-gmbh.de (B.H.)
2
IBE R&D Institute for Lung Health gGmbH, Mendelstraße 11, 48149 Münster, Germany;
vennemann@ibe-ms.de 3
Department of Chemistry & Biology, University of Siegen, Adolf-Reichwein-Str. 2, 57076 Siegen, Germany
engelhard@chemie.uni-siegen.de 3
Department of Chemistry & Biology, University of Siegen, Adolf-Reichwein-Str. 2, 57076 Siegen, Germany;
engelhard@chemie.uni-siegen.de *
Correspondence: martin.wiemann@ibe-ms.de; Tel.: +49-251-9802340 www.mdpi.com/journal/nanomaterials nanomaterials nanomaterials 1. Introduction Amorphous silica (SiO2) is among the most widely used industrial nanomaterials, which can be
found in products such as tires, plastics, and lacquers. It is also used as a food additive to prevent Nanomaterials 2018, 8, 571; doi:10.3390/nano8080571 www.mdpi.com/journal/nanomaterials 2 of 14 Nanomaterials 2018, 8, 571 clogging or to ensure fluidity [1]. The high abundance of SiO2 nanomaterials in everyday life products
may lead to an increased exposure of humans to this type of nanomaterial. With respect to the
workplace situation, the lung is a major route for a non-intentional uptake of SiO2. In contrast to
crystalline silica (quartz and cristobalite), which elicit inflammation and progressive fibrosis in the
lung, effects of amorphous silica appear to be confined to a transient inflammation of the lung, even at
comparably high doses [2,3]. With respect to the mode of action, it is important that amorphous
silica nanomaterials have a large specific surface of up to several hundred square meters per gram. It has been shown that the inflammatory effect of amorphous silica on the lung and also the cytotoxic
effect on macrophages increases with surface size [4]. Of note, the inflammatory effect of colloidal
SiO2 can be reduced by surface coating with amino or phosponate residues [5,6]. Since these surface
coatings have no major influence on particle size or agglomeration behavior of the SiO2 nanoparticles,
it is highly likely that the inflammatory effects of SiO2 on the lung involve an early interaction with
biomolecules accessible in the lung micro-milieu. The analysis of biomolecules interacting with the
surface of silica nanomaterials under in vivo conditions is, therefore, of pivotal interest. Nanosized SiO2 particles, which are deposited alongside the alveolar wall, most likely bind
to molecules of the lipid-rich pulmonary surfactant and/or the protein containing hypophase
underneath [7]. The pulmonary surfactant covers the inner surface of lung alveoli and reduces
the surface tension. It is a thin layer mainly composed of (phospho) lipids (90% by mass),
with phosphatidylcholine being a major component [8]. The lung surfactant also contains specific
surfactant proteins, some of which (e.g., SP-A) may bind to bacteria and, therefore, contribute to the
first line of defense against invading microorganisms [9]. The lung surfactant is produced by type-2
epithelial cells and are stored in lamellar bodies. Upon secretion of lamellar bodies, the surfactant
spreads and self-organizes as a monolayer whose non-polar lipids chains are directed towards the
alveolar space [8]. 2.2. Sample Preparation and Animal Experiments Fe-Si-NP were diluted with sterile distilled H2O to a concentration of 6 mg/mL and ultrasonicated
with a probe (VibraCellTM, Sonics & Materials, Danbury, CT, USA) adjusted to 50 W (20 kHz) for 10 s. This suspension was used as the final instillation fluid. Animal experiments were conducted at the animal facility of the University Clinics of
Münster, Germany, and ethically approved by LANUV (Dortmund, Germany, Accession No. 84–02.04.2022.A157). Female Wistar rats (Charles River Laboratories, Sulzfeld, Germany),
weighing 200–220 g, were maintained at a 12 h lights-on lights-off cycle; food and water were provided
ad libitum. To administer Fe-Si-NP into the lung, animals were briefly anaesthetized with isoflurane
and intratracheally instilled with 500 µL of instillation fluid containing 3 mg (w/v) Fe-Si-NP using a
Penn Century Microsprayer. After 30 min, rats were deeply anaesthetized with ketamine/xylazine
and bled via the descending aorta. A cannula was inserted into the trachea and the lung was
lavaged five times with 5 mL 0.9% NaCl, to obtain lavage fluid for an accompanying investigation
(data not included). Thereafter, the lung was inflated with 5 mL Cryomatrix (Thermo Shandon
Ltd., Runcorn, UK), resected, snap frozen in liquid nitrogen, and stored at −80 ◦C for histological
studies. Transverse sections were cut from the hilar region of the left lung with a cryo-microtome
(HM 500, MICROM International GmbH, Walldorf, Germany). For ToF-SIMS analysis, 7 µm thick
sections were dried onto ITO-coated glass slides (Sigma Aldrich, Germany) and stored at −20 ◦C to
investigate Fe-Si-NP. Of note, the lavage procedure as carried out here leaves a considerable number
of e.g., cells (>70%) within the lung parenchyma [14]. The same is true for particles attached to the
alveolar walls as shown below. 2.3. Enhanced Dark-Field Microscopy Enhanced dark-field microscopy (DFM) makes use of highly intense white light and allows
to visualize particles and NPs in cells and tissues with strong light scattering properties down to
a size of 20–30 nm. To apply the technique, air-dried cryo-sections were fixed in formaldehyde
for 10 min, washed in phosphate buffer and cover-slipped using an aqueous mounting medium
(Shandon Immu-Mount, Thermo Fisher Scientific, Bremen, Germany). Sections were viewed with an
upright microscope (Olympus BX51) and digital bright-field images were captured using a RETIGA
2000R camera (Q Imaging, Surrey, BC, Canada). Areas of interest were then imaged with a DFM
microscope, equipped with a 40-fold UPlanApo Oil Iris objective (Olympus, Hamburg, Germany),
and a CytoViva enhanced dark-field condenser. All DFM components were purchased from CytoViva,
Inc. (Auburn, AL, USA). 1. Introduction Considering the spatial organization of surfactant components and the fact that nanoparticles
bathed in biological fluids such as plasma rapidly acquire a protein corona [10], it is reasonable to
assume that SiO2 nanoparticles, once deposited inside alveoli, form a corona, which, at least in vitro,
can be composed of surfactant lipids and/or proteins [11]. However, because non-polar (phospho)
lipid residues are facing the alveolar space, the corona of biomolecules formed around polar SiO2
nanomaterials under in vivo conditions has not been investigated and may be different from the corona
attracted by non-polar nanoobjects such as carbon nanotubes [12]. The analysis of the protein corona formed around NP inside the lung is a challenging task. However, isolation of protein-laden nanoparticles (NP) would be easier, if NP could be rapidly isolated
from the broncho-alveolar lavage fluid, freed from cells and non-bound components, and eventually be
analyzed for their protein corona. A tempting tool for this task is paramagnetic Fe2O3/SiO2 core/shell
nanoparticles (Fe-Si-NP). While their maghemite iron (Fe) core allows for a rapid magnetic isolation,
their shell is composed of amorphous SiO2, to which biomolecules can bind. In this investigation,
we explored commercially available paramagnetic Fe-Si-NP for the purpose of analyzing the protein
corona in vivo. The main questions were: (i) How are Fe-Si-NP distributed in the lung if they are
applied via intratracheal instillation? (ii) Is the SiO2 shell stable under the conditions of the lung? (iii) Are there any hints for a binding or accumulation of surfactant (phospho) lipids? To answer all these questions with a single investigation, we applied Time-of-Flight secondary
ion mass spectrometry (ToF-SIMS) to sections of Fe-Si-NP-laden rat lungs. ToF-SIMS is an ideal
technique to detect inorganic as well as organic materials with detection limits in the femtomole range. A lateral resolution down to 30 nm may be reached and the technique has recently been shown to
unambiguously detect SiO2 nanomaterial in lung tissue [13]. Here, we detect Si, Fe and PC by mass
spectrometry in a three-dimensional approach. Regions of interest were identified by light microscopic
techniques such as enhanced dark-field microscopy, and incident light microscopy. The results of this
study confirm our assumption that Fe-Si-NP are a useful tool to describe the early process of protein
corona formation in the lung. 3 of 14 Nanomaterials 2018, 8, 571 2.1. Particle Characterization and Sample Preparation 2.1. Particle Characterization and Sample Preparation The Fe-Si-NP were purchased from Kisker Biotech GmbH, Steinfurt, Germany (order No. PMSI-H.25–5). According to manufacturer, particles are composed of a maghemite Fe2O3 core and an
amorphous SiO2 shell and had a surface area of 50 m2/g, according to the Brunauer–Emmet–Teller
(BET) method. To further characterize particle size and shape by transmission electron microscopy
(TEM), we dried 0.5 µL of the aqueous suspension, as used for intratracheal instillation (6 mg/mL
H2O), onto carbon-coated copper grids. TEM analysis was carried out with a Tecnai G2 (ThermoFisher
Scientific, Waltham, MA, USA). The size distribution of these aggregates (in H2O) was measured with
a NanoSight LM10 instrument equipped with a green laser (532 nm), an Andor CCD camera, and NTA
software 2.1 (Malvern Instruments GmbH, Herrenberg, Germany). 2.5. ToF-SIMS Analysis
y
software using the ex
S
S
ft To confirm the chemical identity of the nanoparticles and their Fe/Si core/shell structure a
droplet of the aqueous Fe-Si-NP nanoparticle suspension was dried onto aluminum foil. ToF-SIMS
spectra were acquired in the spectrometry mode at a TOF.SIMS5 (IONTOF, Münster, Germany) for the
pre-characterization of the nanoparticles. The raster size was 120 × 120 µm2 at 128 × 128 pixels. 25 keV
Bi3 ions were applied at an ion dose of 1.5 × 108 ions and a cycle time of 200 µs. The corresponding
mass range reached from 1 Da up to 1650 Da. SensoScan software (Version 3.5.2). 2.5. ToF-SIMS Analysis
To confirm the chemical identity of the nanoparticles and their Fe/Si core/shell structure a
droplet of the aqueous Fe-Si-NP nanoparticle suspension was dried onto aluminum foil. ToF-SIMS
spectra were acquired in the spectrometry mode at a TOF.SIMS5 (IONTOF, Münster, Germany) for
the pre-characterization of the nanoparticles. The raster size was 120 × 120 µm² at 128 × 128 pixels. Measurements using O2 sputtering were performed using Bi3 at 25 keV with an ion dose of
about 2.2 × 1010 ions applied to an analysis field-of-view of 300 × 300 µm2 with a pixel raster
of 512 × 512 pixels. O2+ was used as a sputter ion at an energy of 1 keV. A total sputter dose of
8.0 × 1014 ions was applied to a sputter raster size of 700 × 700 µm2. The delayed extraction mode in
combination with non-interlaced sputtering was used at a cycle time of 80 µs resulting in a mass range
from 1 Da to 550 Da. Three replicate analyses were conducted to assure the validity of the obtained data. p
p
µ
p
25 keV Bi3 ions were applied at an ion dose of 1.5 × 108 ions and a cycle time of 200 µs. The
corresponding mass range reached from 1 Da up to 1650 Da. Measurements using O2 sputtering were performed using Bi3 at 25 keV with an ion dose of about
2.2 × 1010 ions applied to an analysis field-of-view of 300 × 300 µm² with a pixel raster of 512 × 512
pixels. O2+ was used as a sputter ion at an energy of 1 keV. A total sputter dose of 8.0 × 1014 ions was
applied to a sputter raster size of 700 × 700 µm². 2.6. Correlation Analysis
Da. Three replicate an Image processing was carried out with the FIJI distribution for ImageJ (National Institute of Health,
Bethesda, MD, USA) [16,17]. The colocalization threshold plugin was used for the colocalization
analysis [18]. The image registration was executed with the Turboreg [19] plugin (Biomedical Imaging
Group, EPFL Lausanne, Switzerland). 2.6. Correlation Analysis
Image processing was carried out with the FIJI distribution for ImageJ (National Institute of
Health, Bethesda, MD, USA) [16,17]. The colocalization threshold plugin was used for the
colocalization analysis [18]. The image registration was executed with the Turboreg [19] plugin
d
l I
E
l
d 2.5. ToF-SIMS Analysis
y
software using the ex
S
S
ft The delayed extraction mode in combination with
non-interlaced sputtering was used at a cycle time of 80 µs resulting in a mass range from 1 Da to 550 2.4. Topographic Analysis DFM imaged slides were cautiously immersed and rinsed in H2O to remove the coverslip and
embedding medium, respectively. The sections were air-dried again and analyzed with an optical profiler 4 of 14 Nanomaterials 2018, 8, 571 instrument (PLu neox, Sensofar-Tech, Barcelona, Spain). A 50-fold EPI-objective (numerical aperture: 0.80)
was used to acquire images with a resolution of 768 × 576 pixels for the selected area. The corresponding
dimensions of a single pixel are 330 × 330 µm2, which corresponds to a pixel density of 3.03 pixels/mm. Up to 33 consecutive images were acquired at a focal distance of 0.2 µm by an automated routine. The image stack was converted into a single sharp image with ImageJ software using the extended depth
of field plugin [15] or used to generate a topographic image using SensoScan software (Version 3.5.2). Nanomaterials 2018, 7, x FOR PEER REVIEW
4 of 14
aperture: 0.80) was used to acquire images with a resolution of 768 576 pixels for the selected area. The corresponding dimensions of a single pixel are 330 330 µm², which corresponds to a pixel
density of 3.03 pixels/mm. Up to 33 consecutive images were acquired at a focal distance of 0.2 µm
by an automated routine The image stack was converted into a single sharp image with ImageJ 3. Results and Discussion
(
g
g
3 Results and Discussio The
e e
e of Fe u o
i
a y io
bo
ba d
e t u
e t
that the ToF SIMS i fo
atio
While the findings illustrate the need for an extensive pre-characterization of nanoparticles before
their use in toxicology studies, they also indicate several characteristic secondary ions (28Si+, 30Si+,
54Fe+, 56Fe+, and 56FeOH+) that can be used for the detection of the nanoparticles in the tissues. While the findings illustrate the need for an extensive pre-characterization of nanoparticles
before their use in toxicology studies, they also indicate several characteristic secondary ions (28Si+,
30Si+, 54Fe+, 56Fe+, and 56FeOH+) that can be used for the detection of the nanoparticles in the tissues. The presence of Fe upon primary ion bombardment suggests that the ToF SIMS information
While the findings illustrate the need for an extensive pre-characterization of nanoparticles before
their use in toxicology studies, they also indicate several characteristic secondary ions (28Si+, 30Si+,
54Fe+, 56Fe+, and 56FeOH+) that can be used for the detection of the nanoparticles in the tissues. The presence of Fe upon primary ion bombardment suggests that the ToF-SIMS information
depth might be greater than the thickness of the Si shell. This finding can be explained by the highly
energetic primary ion beam, which induces significant amounts of damage to nanoparticles and
sometimes may even disrupt the particle. Sandoval et al. found an increased secondary ion yield for
nanoparticles compared to bulk materials [21]. Increases in the secondary ion yield are always to be
expected from nanoparticles depending on their sizes, because they have a larger surface area than
bulk materials. However, the performance of the collision cascade is affected, if the size of the
particles is in the same order of magnitude as the collision cascade. Obviously, this can also lead to
increased intermixing of the particle components. Yang et al. observed a deformation of nanoparticles
upon ion bombardment [22], which could also explain the simultaneous detection of core and shell
components observed here for Fe-Si-NP. In any case, the simultaneous detection of the Si and Fe
signals without the need for sputtering was beneficial for the detection of the core/shell particles in
the tissue
The presence of Fe upon primary ion bombardment suggests that the ToF-SIMS information
depth might be greater than the thickness of the Si shell. 3. Results and Discussion
(
g
g
3 Results and Discussio 3.1. Electron Microscopic and ToF-SIMS Characterization of the Nanoparticles
3 1 El t
Mi
i
d T F SIMS Ch
t i
ti
f th N
ti l TEM analysis showed that Fe-Si-NP consisted of nanoparticles with a size of ca. 10–20 nm,
which mainly formed larger aggregates (Figure 1, inset). Fe-Si-NP dispersed in H2O showed two
maxima at 168 nm and 228 nm, as analyzed with optical tracking analysis; mean and mode values
amounted to 217.5 ± 7.8 nm and 190.3 ± 19.6 nm, respectively (Figure 1). 3.1. Electron Microscopic and ToF SIMS Characterization of the Nanoparticles
TEM analysis showed that Fe-Si-NP consisted of nanoparticles with a size of ca. 10–20 nm, which
mainly formed larger aggregates (Figure 1, inset). Fe-Si-NP dispersed in H2O showed two maxima at
168 nm and 228 nm, as analyzed with optical tracking analysis; mean and mode values amounted to
217.5 ± 7.8 nm and 190.3 ± 19.6 nm, respectively (Figure 1). Figure 1. Size distribution of Fe-Si nanoparticles (Fe-Si-NP) in aqueous suspension as used for
intratracheal instillation. Measurements were carried out by NanoSight optical tracking analysis. The
inset shows electron microscopic images of Fe-Si-NP as typically found in the instilled suspension. Figure 1. Size distribution of Fe-Si nanoparticles (Fe-Si-NP) in aqueous suspension as used for
intratracheal instillation. Measurements were carried out by NanoSight optical tracking analysis. The inset shows electron microscopic images of Fe-Si-NP as typically found in the instilled suspension. Figure 1. Size distribution of Fe-Si nanoparticles (Fe-Si-NP) in aqueous suspension as used for
intratracheal instillation. Measurements were carried out by NanoSight optical tracking analysis. The
inset shows electron microscopic images of Fe-Si-NP as typically found in the instilled suspension. Figure 1. Size distribution of Fe-Si nanoparticles (Fe-Si-NP) in aqueous suspension as used for
intratracheal instillation. Measurements were carried out by NanoSight optical tracking analysis. The inset shows electron microscopic images of Fe-Si-NP as typically found in the instilled suspension. 5 of 14
5 of 14 Nanomaterials 2018, 8, 571
Na o ate ial 2018 7
FO The mass spectrum, as revealed by ToF-SIMS, shows several prominent Si- and Fe-containing
species (Figure 2), from which 28Si+, 30Si+, 54Fe+, 56Fe+ and 56FeOH+ were selected as relevant
secondary ions for the detection of Fe-Si-NP in lung tissue sections. 3. Results and Discussion
(
g
g
3 Results and Discussio Besides the often detected
hydrocarbons [20], B+, Na+ and Ca+ were detected with notable intensities and these substances are
most likely associated with the particle production process. The mass spectrum, as revealed by ToF-SIMS, shows several prominent Si- and Fe-containing
species (Figure 2), from which 28Si+, 30Si+, 54Fe+, 56Fe+ and 56FeOH+ were selected as relevant secondary
ions for the detection of Fe-Si-NP in lung tissue sections. Besides the often detected hydrocarbons
[20], B+, Na+ and Ca+ were detected with notable intensities and these substances are most likely
associated with the particle production process. Figure 2. Mass spectrum of the Fe-Si-NP with a Fe/Si core/shell structure. Particles were deposited on
an aluminum substrate and analyzed by ToF-SIMS. The particle components SixOyHz+ and FexOyHz+
and their respective isotopes are detected along with hydrocarbons (CH3+ and C2H3+), substrate
components (Al+), and other residues possibly linked to the production process (B+, Na+, and Ca+). Figure 2. Mass spectrum of the Fe-Si-NP with a Fe/Si core/shell structure. Particles were deposited on
an aluminum substrate and analyzed by ToF-SIMS. The particle components SixOyHz+ and FexOyHz+
and their respective isotopes are detected along with hydrocarbons (CH3+ and C2H3+), substrate
components (Al+), and other residues possibly linked to the production process (B+, Na+, and Ca+). Figure 2. Mass spectrum of the Fe-Si-NP with a Fe/Si core/shell structure. Particles were deposited on
an aluminum substrate and analyzed by ToF-SIMS. The particle components SixOyHz+ and FexOyHz+
and their respective isotopes are detected along with hydrocarbons (CH3+ and C2H3+), substrate
components (Al+), and other residues possibly linked to the production process (B+, Na+, and Ca+). Figure 2. Mass spectrum of the Fe-Si-NP with a Fe/Si core/shell structure. Particles were deposited on
an aluminum substrate and analyzed by ToF-SIMS. The particle components SixOyHz+ and FexOyHz+
and their respective isotopes are detected along with hydrocarbons (CH3+ and C2H3+), substrate
components (Al+), and other residues possibly linked to the production process (B+, Na+, and Ca+). While the findings illustrate the need for an extensive pre-characterization of nanoparticles
before their use in toxicology studies, they also indicate several characteristic secondary ions (28Si+,
30Si+, 54Fe+, 56Fe+, and 56FeOH+) that can be used for the detection of the nanoparticles in the tissues. 3. Results and Discussion
(
g
g
3 Results and Discussio This finding can be explained by the highly
energetic primary ion beam, which induces significant amounts of damage to nanoparticles and
sometimes may even disrupt the particle. Sandoval et al. found an increased secondary ion yield for
nanoparticles compared to bulk materials [21]. Increases in the secondary ion yield are always to be
expected from nanoparticles depending on their sizes, because they have a larger surface area than
bulk materials. However, the performance of the collision cascade is affected, if the size of the particles
is in the same order of magnitude as the collision cascade. Obviously, this can also lead to increased
intermixing of the particle components. Yang et al. observed a deformation of nanoparticles upon ion
bombardment [22], which could also explain the simultaneous detection of core and shell components
observed here for Fe-Si-NP. In any case, the simultaneous detection of the Si and Fe signals without
the need for sputtering was beneficial for the detection of the core/shell particles in the tissue. 3 2 Inspection of Fe Si NP Laden Lung Tissue by Light Microscopy
3.2. Inspection of Fe-Si-NP-Laden Lung Tissue by Light Microscopy 3.2. Inspection of Fe Si NP Laden Lung Tissue by Light Microscopy
Light microscopic techniques are increasingly used to detect nanoparticles in tissue sections [23]. Especially enhanced dark-field microscopy can detect light scattering particles down to a size of 20–
30 nm [24]. Furthermore, metallic or metal oxidic nanoparticles often have plasmonic, reflective and
diffractive properties, which allow for their differentiation against translucent tissue [25], or, in ideal
cases, for identification via hyperspectral imaging [26]. Here, bright-field microscopy combined with
DFM was used to localize Fe Si NP in the lung tissue prior to ToF SIMS analysis
Light microscopic techniques are increasingly used to detect nanoparticles in tissue sections [23]. Especially enhanced dark-field microscopy can detect light scattering particles down to a size of
20–30 nm [24]. Furthermore, metallic or metal oxidic nanoparticles often have plasmonic, reflective and
diffractive properties, which allow for their differentiation against translucent tissue [25], or, in ideal
cases, for identification via hyperspectral imaging [26]. Here, bright-field microscopy combined with
DFM was used to localize Fe-Si-NP in the lung tissue prior to ToF-SIMS analysis. 6 of 14 Nanomaterials 2018, 8, 571 Although the microscopic detection of SiO2 nanomaterials in tissue sections is hampered by
low optical contrast, the iron core of Fe-Si-NP caused a “rust”-like color facilitating the identification
of larger agglomerates in bright-field images (Figure 3A–F). DFM confirmed this distribution but
additionally revealed smaller yellow or blue dots in, or attached to, the hardly light scattering tissue
structure (Figure 3C,D). Although the detection of nanoparticles in tissues by DFM is generally
a straightforward approach, it is obvious that any light scattering inhomogeneity might lead
to misinterpretation. Nanomaterials 2018, 7, x FOR PEER REVIEW
6 of 14
larger agglomerates in bright-field images (Figure 3A–F). DFM confirmed this distribution but
additionally revealed smaller yellow or blue dots in, or attached to, the hardly light scattering tissue
structure (Figure 3C,D). Although the detection of nanoparticles in tissues by DFM is generally a
straightforward approach, it is obvious that any light scattering inhomogeneity might lead to
misinterpretation Figure 3. Light microscopic analyses of a Fe-Si-NP-laden cryo-section from rat lung. Overview
(A,C,E,G) and detail (B,D,F,H) of the boxed area shown on the left. (A–D) Routine images of the
coverslipped section taken with bright-field optics (A,B) and DFM (C,D); Fe-Si-NP agglomerates
appear as brownish patches at alveolar septa or in macrophages. 3 2 Inspection of Fe Si NP Laden Lung Tissue by Light Microscopy
3.2. Inspection of Fe-Si-NP-Laden Lung Tissue by Light Microscopy With DFM, additional small light-
scattering particles are seen alongside alveolar septa. (E–H) Same tissue area after removal of the
coverslip, rinsing and drying. (E,F) Microscopic image of the dry section without coverslip, taken with
a reflective light microscope. Prominent Fe-Si-NP agglomerates are still identifiable as yellow patches. (G,H) Optical profilometry analysis showing the relief structure of the dry lung section as subjected
to ToF-SIMS; the topographic map reveals an amplitude of several micrometers. Arrows in (A,C,E,G)
point to a particle-laden macrophage (A,C), which became lost in the course of tissue preparation. Similarly, arrows in (B,D,F,H) point to small Fe-Si-NP (seen in (B,D)), which were absent after drying. Bright-field microscopy and DFM demand that the tissue section is immersed in ringer solution,
co erslipped and
iewed with oil immersion lenses As ToF SIMS analysis is carried out under
5.75 µm
0
Figure 3. Light microscopic analyses of a Fe-Si-NP-laden cryo-section from rat lung. Overview (A,C,E,G)
and detail (B,D,F,H) of the boxed area shown on the left. (A–D) Routine images of the coverslipped
section taken with bright-field optics (A,B) and DFM (C,D); Fe-Si-NP agglomerates appear as brownish
patches at alveolar septa or in macrophages. With DFM, additional small light-scattering particles
are seen alongside alveolar septa. (E–H) Same tissue area after removal of the coverslip, rinsing and
drying. (E,F) Microscopic image of the dry section without coverslip, taken with a reflective light
microscope. Prominent Fe-Si-NP agglomerates are still identifiable as yellow patches. (G,H) Optical
profilometry analysis showing the relief structure of the dry lung section as subjected to ToF-SIMS;
the topographic map reveals an amplitude of several micrometers. Arrows in (A,C,E,G) point to a
particle-laden macrophage (A,C), which became lost in the course of tissue preparation. Similarly,
arrows in (B,D,F,H) point to small Fe-Si-NP (seen in (B,D)), which were absent after drying. 5.75 µm
0 5.75 µm
0 Figure 3. Light microscopic analyses of a Fe-Si-NP-laden cryo-section from rat lung. Overview
(A,C,E,G) and detail (B,D,F,H) of the boxed area shown on the left. (A–D) Routine images of the
coverslipped section taken with bright-field optics (A,B) and DFM (C,D); Fe-Si-NP agglomerates
appear as brownish patches at alveolar septa or in macrophages. With DFM, additional small light-
scattering particles are seen alongside alveolar septa. (E–H) Same tissue area after removal of the
coverslip, rinsing and drying. (E,F) Microscopic image of the dry section without coverslip, taken with
a reflective light microscope. 3 2 Inspection of Fe Si NP Laden Lung Tissue by Light Microscopy
3.2. Inspection of Fe-Si-NP-Laden Lung Tissue by Light Microscopy Prominent Fe-Si-NP agglomerates are still identifiable as yellow patches. (G,H) Optical profilometry analysis showing the relief structure of the dry lung section as subjected
to ToF-SIMS; the topographic map reveals an amplitude of several micrometers. Arrows in (A,C,E,G)
point to a particle-laden macrophage (A,C), which became lost in the course of tissue preparation. Similarly, arrows in (B,D,F,H) point to small Fe-Si-NP (seen in (B,D)), which were absent after drying. Bright-field microscopy and DFM demand that the tissue section is immersed in ringer solution,
Figure 3. Light microscopic analyses of a Fe-Si-NP-laden cryo-section from rat lung. Overview (A,C,E,G)
and detail (B,D,F,H) of the boxed area shown on the left. (A–D) Routine images of the coverslipped
section taken with bright-field optics (A,B) and DFM (C,D); Fe-Si-NP agglomerates appear as brownish
patches at alveolar septa or in macrophages. With DFM, additional small light-scattering particles
are seen alongside alveolar septa. (E–H) Same tissue area after removal of the coverslip, rinsing and
drying. (E,F) Microscopic image of the dry section without coverslip, taken with a reflective light
microscope. Prominent Fe-Si-NP agglomerates are still identifiable as yellow patches. (G,H) Optical
profilometry analysis showing the relief structure of the dry lung section as subjected to ToF-SIMS;
the topographic map reveals an amplitude of several micrometers. Arrows in (A,C,E,G) point to a
particle-laden macrophage (A,C), which became lost in the course of tissue preparation. Similarly,
arrows in (B,D,F,H) point to small Fe-Si-NP (seen in (B,D)), which were absent after drying. 7 of 14 Nanomaterials 2018, 8, 571 Bright-field microscopy and DFM demand that the tissue section is immersed in ringer solution,
coverslipped and viewed with oil-immersion lenses. As ToF-SIMS analysis is carried out under
vacuum conditions, the coverslip has to be cautiously removed and the sections needs to be dried. Mild shearing forces and washing steps are indispensable during this procedure and may influence
the signals gathered from particles in the subsequent ToF-SIMS analysis. To show these effects,
reflective light microscopic images were taken at the same position of the dried section. Reflective light microscopic images have a lower optical resolution but are highly useful to
visualize the structure of a section prepared for ToF-SIMS analysis. 3 2 Inspection of Fe Si NP Laden Lung Tissue by Light Microscopy
3.2. Inspection of Fe-Si-NP-Laden Lung Tissue by Light Microscopy Here, it is shown that numerous
large yellow spots (up to 15 µm) previously seen in the bright-field or DFM image were still in
place, whereas others structures, which were interpreted as Fe-Si-NP filled macrophages, were no
longer visible (Figure 3E,F; white arrows). Similarly, smaller particles (<3 µm), which are marked by
arrows in the magnified bright-field and DFM images (Figure 3B,D), were not seen on reflective light
microscopy images. py
g
As ToF-SIMS analysis makes use of sputtering and ion bombardment, which is applied under a
certain angle, the relief structure of the dried tissue section will unavoidably influence image resolution. Figure 3G,H shows the relief of the dried tissue on the indium tin oxide (ITO)-coated glass substrate,
as analyzed by profilometry. The height of the formalin-fixed and afterwards dried alveolar septa on
the substrate ranged 2.5–5.25 µm, which was 36–75% of the initial thickness of the cryo-section (7 µm)
and may be explained by evaporation of H2O prior to immersion fixation. Although the nature of the
highest peaks remains unknown, these sites do not contain agglomerated Fe-Si-NP, which were mainly
found in areas ranging in height from 2–3.5 µm. In summary, the combination of several microscopic methods showed that Fe-Si-NP were
distributed alongside the alveolar wall or were contained in macrophage-like structures. The successive
observation by DFM and ToF-SIMS may lead to a loss of Fe-Si-NP containing structures and this has to
be considered when both images are compared (see below). 3.3. ToF-SIMS Detection of Nanoparticles in Tissue—Localization However, due to the preparation steps
of the lung prior to fixation (filling with cryomatrix, freezing, sectioning, and drying) and to the
relief structure of the tissue (Figure 3G), a precise cellular localization of particles cannot be made by
light microscopy or ToF-SIMS. A study by Shon et al. carried out on air-dried unfixed macrophages
laden with dimercaptosuccinic acid-coated Fe3O4 nanoparticles succeeded in localizing particles in
the cytoplasm [27]. Phosphatidylcholine components in that study were detected as specific C4H7+
ions only. Nanomaterials 2018, 7, x FOR PEER REVIEW
8 of 14
tissue (Figure 3G), a precise cellular localization of particles cannot be made by light microscopy or
ToF-SIMS. A study by Shon et al. carried out on air-dried unfixed macrophages laden with
dimercaptosuccinic acid-coated Fe3O4 nanoparticles succeeded in localizing particles in the
cytoplasm [27]. Phosphatidylcholine components in that study were detected as specific C4H7+ ions
only. Figure 4. ToF-SIMS signal distributions of organic and inorganic components in a Fe/Si core/shell
nanoparticle-containing lung tissue section 30 min post intratracheal instillation of particles. (A)
CH4N+ indicative of proteins to show tissue distribution and lung structure. (B) C5H15NPO4+ is the
phosphocholine (PC) head group fragment showing the distribution of lipids of lung tissue and
surfactant; PC signal seen in the alveolar space is interpreted as to be artificially dislocated during
filling of the lungs with cryomatrix. (C) Overlay of 54Fe+ (green) and CH4N+ (red) signal showing Fe-
Si-NP distribution in lung tissue. (D) 56Fe+, and (E) 54Fe+ distributions of Fe-Si-NP particles. Small and
large agglomerates appear alongside the alveolar walls. (F) The overlay of 54Fe+ (D, red) and 28Si+ (G,
green) reveals the colocalization of both particle-related signals. (G) ToF-SIMS signal distribution of
Si species from the topmost layers as indicated by the horizontal green lines in the X-Y slice of the
ToF-SIMS dataset in (H). (I) 28Si+ signal distribution integrated for the complete depth including the
indium tin oxide-coated glass slide. 3.3. ToF-SIMS Detection of Nanoparticles in Tissue—Localization 3.3. ToF-SIMS Detection of Nanoparticles in Tissue—Localization In the next step, we tested the intactness of the Fe/Si core/shell in the tissue using both the
Fe- and Si-signal distributions in the ToF-SIMS analysis. A sputter erosion with 1 keV O2+ ions was
necessary to analyze the complete depth of the sample section. The tissue structure of the lung with all its alveolar septa, as shown in Figure 3, was properly
reflected by the CH4N+ signal, which is a typical fragment of amino acids and, thus, indicates the
position of fixed proteins (Figure 4A). Due to the relatively harsh sputtering conditions of 1 keV O2+
ions, only few organic molecular species were preserved. Nevertheless, the lateral distribution of the
signal at m/z 184.07 could be summed up over the whole depth of the analysis. This fragment
is indicative of the phosphocholine head group C5H15NPO4+ and most likely attributable to
phospholipids previously located in cell membranes and/or in the pulmonary surfactant. A minor
fraction of the signal was found in alveolar spaces (Figure 4B), suggesting that it was transferred onto
the substrate during the preparation process. With respect to Fe, several relevant signals (besides the main isotopes 54Fe+ and 56Fe+) were
identified in the spectrum of both, the isolated and the lung-incorporated Fe-Si-NP (Figure 2). Of note,
further iron-related species with the same lateral distribution were found, e.g., 56FeOH+, 54FeOH+,
and 56Fe2O+, proving that the signals were correctly assigned to all Fe species. In any case, the lateral
signal distribution of the main Fe isotopes 54Fe+ and 56Fe+ (Figure 4D,E) was largely congruent with
the distribution of Fe-Si-NP in the tissue (Figure 3C,E). This is particularly obvious for some larger
areas with diameters of up to 15 µm (Figure 4D,E), but also for numerous smaller and less intense
Fe patches. As most of these areas have a diameter larger than 200 nm, they most likely represent
Fe-Si-NP agglomerates. Colocalization with the CH4N+ signal (Figure 4C) shows that the majority of Fe signals (green)
in the range of 3–10 µm was localized alongside the alveolar septa (red), which is in accordance 8 of 14 Nanomaterials 2018, 8, 571 with the microscopic results of Figure 3 and was expected due to the short incubation time of only
30 min. Some of the smaller Fe-signals (green) were colocalized with the epithelium and might
be interpreted as to be incorporated by the epithelial cell. 3.3. ToF-SIMS Detection of Nanoparticles in Tissue—Localization 300
250
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MC: 17; TC: 7.854e+005
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MC: 126; TC: 1.820e+006
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MC: 148; TC: 2.462e+007
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MC: 12; TC: 6.369e+004
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MC: 60; TC: 9.601e+005
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Overlay of: 54Fe+, 28Si+ (Scan 1-20)
Overlay of: 54Fe+, 28Si+ (Scan 1-20)
Figure 4. ToF-SIMS signal distributions of organic and inorganic components in a Fe/Si core/shell
nanoparticle-containing lung tissue section 30 min post intratracheal instillation of particles. (A) CH4N+
indicative of proteins to show tissue distribution and lung structure. (B) C5H15NPO4+ is the
phosphocholine (PC) head group fragment showing the distribution of lipids of lung tissue and
surfactant; PC signal seen in the alveolar space is interpreted as to be artificially dislocated during
filling of the lungs with cryomatrix. (C) Overlay of 54Fe+ (green) and CH4N+ (red) signal showing
Fe-Si-NP distribution in lung tissue. (D) 56Fe+, and (E) 54Fe+ distributions of Fe-Si-NP particles. Small and large agglomerates appear alongside the alveolar walls. (F) The overlay of 54Fe+ (D, red)
and 28Si+ (G, green) reveals the colocalization of both particle-related signals. (G) ToF-SIMS signal
distribution of Si species from the topmost layers as indicated by the horizontal green lines in the X-Y
slice of the ToF-SIMS dataset in (H). (I) 28Si+ signal distribution integrated for the complete depth
including the indium tin oxide-coated glass slide. 3.3. ToF-SIMS Detection of Nanoparticles in Tissue—Localization 300
250
200
150
100
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μm
CH4N+
MC: 17; TC: 7.854e+005
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MC: 7; TC: 4.348e+005
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MC: 126; TC: 1.820e+006 00
50 56Fe+
MC: 126; TC: 1.820e+006 28Si+ (Scan 1-20)
MC: 7; TC: 2.048e+004
Overlay of: 54Fe+, 28Si+ (Scan 1-20) MC: 12; TC: 6.369e+004
300
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MC: 60; TC: 9.601e+005 300
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0 Si
MC: 148; TC: 2.462e+007 Figure 4. ToF-SIMS signal distributions of organic and inorganic components in a Fe/Si core/shell
nanoparticle-containing lung tissue section 30 min post intratracheal instillation of particles. (A)
CH4N+ indicative of proteins to show tissue distribution and lung structure. (B) C5H15NPO4+ is the
phosphocholine (PC) head group fragment showing the distribution of lipids of lung tissue and
surfactant; PC signal seen in the alveolar space is interpreted as to be artificially dislocated during
filling of the lungs with cryomatrix. (C) Overlay of 54Fe+ (green) and CH4N+ (red) signal showing Fe-
Si-NP distribution in lung tissue. (D) 56Fe+, and (E) 54Fe+ distributions of Fe-Si-NP particles. Small and
large agglomerates appear alongside the alveolar walls. (F) The overlay of 54Fe+ (D, red) and 28Si+ (G,
green) reveals the colocalization of both particle-related signals. (G) ToF-SIMS signal distribution of
Si species from the topmost layers as indicated by the horizontal green lines in the X-Y slice of the
ToF-SIMS dataset in (H). (I) 28Si+ signal distribution integrated for the complete depth including the
indium tin oxide-coated glass slide. Figure 4. 3.4. Colocalization of Fe and Si in Lung Tissue by ToF-SIMS 3.4. Colocalization of Fe and Si in Lung Tissue by ToF-SIMS With respect to the simultaneous identification of Fe and Si, isobaric interferences may occur
due to the similarity of the theoretical mass of 56Fe+ (m/z 55.934939) and 28Si2+ (m/z 55.953854). This hampers a reliable distinction by mass, and only an improvement of mass resolution from the
current R = 1500 to R > 3000 could solve this problem. However, the 54Fe+ signal, although relatively
weak, is not influenced by any Si cluster and, therefore, allows analyzing the Fe distribution unbiased
from Si signals. Thus, adopting 54Fe+ as a reference, it turned out that 56Fe+ signals virtually showed
the same distribution as did 54Fe+. In addition, the 56Fe+ signal was hardly influenced by 28Si2+ from
any non-particle source and showed a low background intensity in tissue-free areas (Figure 4D,G). Due to the obvious correlation of the signal distributions for the 54Fe+ and 56Fe+ isotopes, we decided
to use the 56Fe+ ion to measure the distribution of Fe in lung sections. g
To identify Si on the paramagnetic Fe core, any Si signal from the ITO-covered glass substrate
had to be avoided. Therefore, the 28Si signal was reconstructed from only the first 20 sputter/analysis
cycles (i.e., scans). The signal distribution obtained by this was highly similar to the Fe distributions
(Figure 4G), indicating that the ITO layer had successfully shielded the underlying glass substrate
against the analysis beam. In contrast, if the signal was summed up over all 81 cycles (compare X–Z
slice in Figure 4H), the sputter cycles had eroded the ITO layer and the apparent 28Si+ distribution
reflected the underlying substrate in tissue-free regions (Figure 4I). Based on selected data evaluation
and an isotopic signal distribution, the presence of Si on nanoparticles could be confirmed even in the
positive ion polarity upon oxygen sputtering, as opposed to earlier studies using Cs sputtering and
the negative ion polarity [13]. Consequently, the identification by ToF-SIMS of pure, unlabeled SiO2
nanoparticles in tissue sections should be possible with this protocol, provided that any contamination,
e.g., with the widespread polysiloxanes, can be ruled out. Another option to identify correlating distributions of Si and Fe species is the comparison of
Fe isotope distributions with 30Si, as oxide clusters of this ion cannot form signals with the same
mass-to-charge-ratio as the Fe isotopes in the observed mass range. 3.4. Colocalization of Fe and Si in Lung Tissue by ToF-SIMS However, as 30Si is also a part of
the ITO shielded substrate, the same limitations as for 28Si apply. Of note, the reconstruction of the less
abundant 30Si+ isotope ruled out that the 28Si+ signal merely originated from 56Fe+ (m/z 27.967470),
which would partially interfere with the 28Si+ (m/z 27.976927), since both, 28Si and 30Si show the same
signal distributions (not shown here). Based on the comparison of Figure 4D,E (for Fe) and Figure 4G (for Si), the most important result
is that the 28Si+ distribution derived from the first 20 sputter/analysis cycles was properly colocalized
with the Fe distributions, as is shown by the numerous yellow spots in Figure 4F, which result from
red 54Fe pixels overlaid with green 28Si pixels. This demonstrates that the Si shell of the paramagnetic
Fe particles is stable under in vivo conditions. 3.3. ToF-SIMS Detection of Nanoparticles in Tissue—Localization ToF-SIMS signal distributions of organic and inorganic components in a Fe/Si core/shell
nanoparticle-containing lung tissue section 30 min post intratracheal instillation of particles. (A) CH4N+
indicative of proteins to show tissue distribution and lung structure. (B) C5H15NPO4+ is the
phosphocholine (PC) head group fragment showing the distribution of lipids of lung tissue and
surfactant; PC signal seen in the alveolar space is interpreted as to be artificially dislocated during
filling of the lungs with cryomatrix. (C) Overlay of 54Fe+ (green) and CH4N+ (red) signal showing
Fe-Si-NP distribution in lung tissue. (D) 56Fe+, and (E) 54Fe+ distributions of Fe-Si-NP particles. Small and large agglomerates appear alongside the alveolar walls. (F) The overlay of 54Fe+ (D, red)
and 28Si+ (G, green) reveals the colocalization of both particle-related signals. (G) ToF-SIMS signal
distribution of Si species from the topmost layers as indicated by the horizontal green lines in the X-Y
slice of the ToF-SIMS dataset in (H). (I) 28Si+ signal distribution integrated for the complete depth
including the indium tin oxide-coated glass slide. Figure 4. ToF-SIMS signal distributions of organic and inorganic components in a Fe/Si core/shell
Figure 4. ToF-SIMS signal distributions of organic and inorganic components in a Fe/Si core/shell 9 of 14 9 of 14 Nanomaterials 2018, 8, 571 3.5. Correlation and Comparison of Dark-Field Microscopy and ToF-SIMS Enhanced dark-field microscopy (DFM) detects NP down to a size of 15 nm [28] and was, therefore,
used as a sensitive tool to visualize the Fe-Si-NP in tissue sections prior to ToF-SIMS analysis. Overall,
the ToF-SIMS 54Fe+ signal and the DFM signal showed a high degree of similarity (Figure 5A,B). Colocalization analysis of both signals using Pearson’s correlation coefficient confirmed this high
degree of co-localization. As this coefficient ranges from −1 (perfect anti-correlation) to +1 (perfect
correlation), a value of 0.541 confirmed that a considerable number of pixels (54Fe+ positive by
ToF-SIMS) correlate with the bright spots in the DFM image (Figure 5C). Of note, this result strongly
suggests that ToF-SIMS had not detected false positive signals. However, the correlation was not perfect and differences were observed for some larger dots
(marked in Figure 5), most likely corresponding to particle-laden macrophages, but also for smaller
particles seen in DFM. As outlined above, especially inappropriately fixed material may have become
lost during preparation. 10 of 14 10 of 14 Nanomaterials 2018, 8, 571 Other reasons may also account for such differences: Fe-Si-NP might not have been detected
by ToF-SIMS due to insufficient sample erosion (false negative ToF-SIMS signals). However, as the
sputtering with O2+ and subsequent analysis were extended into the depth of the substrate, it appears
unlikely that the soft lung tissue shielded Fe-Si-NP and prevented the accumulation of supra-threshold
signal intensities in the ToF-SIMS analysis. Rather, the accessible mass (i.e., particle size and/or
concentration) was too small, and a low signal did not reach the limit of detection. This appears
unlikely for most of the Fe-Si-NP aggregates larger than 100 nm but may account for the 10–50 nm
fraction of Fe-Si-NP, which is visible by TEM (Figure 1) and, at least in part, should be visible by DFM. Nanomaterials 2018, 7, x FOR PEER REVIEW
10 of 14
50 nm fraction of Fe-Si-NP, which is visible by TEM (Figure 1) and, at least in part, should be visible
by DFM. y
g
p
y
Finally, unlike the large brown agglomerates (Figure 3A,B), small objects can scatter light as well
and may have been erroneously designated as Fe-Si-NP by DFM. Hyperspectral microscopy may be
used to further identify such questionable structures. y
Finally, unlike the large brown agglomerates (Figure 3A,B), small objects can scatter light as well
and may have been erroneously designated as Fe-Si-NP by DFM. 3.5. Correlation and Comparison of Dark-Field Microscopy and ToF-SIMS Hyperspectral microscopy may be
used to further identify such questionable structures. In summary, the ion distribution measured by ToF-SIMS explains most of the DFM signals,
thereby serving as a validation technique. However, DFM images contained several signals without
any ToF-SIMS equivalent. These DFM signals were most likely caused by salt residues or tissue
material and indicate that it is necessary to confirm the chemical identity of nanoparticles in DFM
imaging studies. In summary, the ion distribution measured by ToF-SIMS explains most of the DFM signals,
thereby serving as a validation technique. However, DFM images contained several signals without
any ToF-SIMS equivalent. These DFM signals were most likely caused by salt residues or tissue
material and indicate that it is necessary to confirm the chemical identity of nanoparticles in DFM
imaging studies. Figure 5. Comparison of ToF-SIMS 54Fe+ signal distribution and dark-field microscopy (DFM) images. Image taken from Figure 3. (A) 54Fe+ ToF-SIMS signal (red); (B) DFM signal (green) taken before
washing and ToF-SIMS analysis; and (C) Colocalization of (A,B). A high degree of correlation is
evident (white overlay). The Pearson correlation coefficient for this image was 0.541. Arrows point to
green spots (B) which were detected by DFM, but not by ToF-SIMS. 6 Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings
Figure 5. Comparison of ToF-SIMS 54Fe+ signal distribution and dark-field microscopy (DFM) images. Image taken from Figure 3. (A) 54Fe+ ToF-SIMS signal (red); (B) DFM signal (green) taken before
washing and ToF-SIMS analysis; and (C) Colocalization of (A,B). A high degree of correlation is evident
(white overlay). The Pearson correlation coefficient for this image was 0.541. Arrows point to green
spots (B) which were detected by DFM, but not by ToF-SIMS. Figure 5. Comparison of ToF-SIMS 54Fe+ signal distribution and dark-field microscopy (DFM) images. Image taken from Figure 3. (A) 54Fe+ ToF-SIMS signal (red); (B) DFM signal (green) taken before
washing and ToF-SIMS analysis; and (C) Colocalization of (A,B). A high degree of correlation is
evident (white overlay). The Pearson correlation coefficient for this image was 0.541. Arrows point to
green spots (B) which were detected by DFM, but not by ToF-SIMS. Figure 5. Comparison of ToF-SIMS 54Fe+ signal distribution and dark-field microscopy (DFM) images. Image taken from Figure 3. (A) 54Fe+ ToF-SIMS signal (red); (B) DFM signal (green) taken before
washing and ToF-SIMS analysis; and (C) Colocalization of (A,B). 3.5. Correlation and Comparison of Dark-Field Microscopy and ToF-SIMS A high degree of correlation is evident
(white overlay). The Pearson correlation coefficient for this image was 0.541. Arrows point to green
spots (B) which were detected by DFM, but not by ToF-SIMS. .6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings
.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings 3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Finding
3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings It appears conceivable that this sequential arrangement of biomolecules and ions
in the lung orchestrates the formation of the protein corona as well as agglomeration of NP. A protein
corona, which can be artificially generated with different preparations of lung surfactant [11] may,
therefore, differ from the native corona. To monitor the putative binding of (phosho) lipids to silica nanoparticles in the lung, we To monitor the putative binding of (phosho) lipids to silica nanoparticles in the lung, we compared
the m/z 184.07 signal (representing phosphatidylcholine as the main constituent of pulmonary
surfactant and cell membranes), with the distribution of Si, which forms the outer shell of the Fe-Si-NP:
While the m/z 184.07 signal delineated the alveolar septal structure, and was also found on the ITO
substrate at lower intensities, volumes occupied by the Si signal in the first 20 scans were completely
free of PC (Figure 6, arrows). Although we are aware that the sensitivity of analytical methods can be
improved, e.g., by well-adapted sputtering methods, we conclude that there is no major accumulation
of phospholipids on the polar surface of Si nanoparticles. We furthermore suggest that phospholipids,
as constituents of the lung surfactant, are not involved in the formation of particle agglomerates. p
g
(p
)
p
p
g
compared the m/z 184.07 signal (representing phosphatidylcholine as the main constituent of
pulmonary surfactant and cell membranes), with the distribution of Si, which forms the outer shell
of the Fe-Si-NP: While the m/z 184.07 signal delineated the alveolar septal structure, and was also
found on the ITO substrate at lower intensities, volumes occupied by the Si signal in the first 20 scans
were completely free of PC (Figure 6, arrows). Although we are aware that the sensitivity of analytical
methods can be improved, e.g., by well-adapted sputtering methods, we conclude that there is no
major accumulation of phospholipids on the polar surface of Si nanoparticles. We furthermore
suggest that phospholipids, as constituents of the lung surfactant, are not involved in the formation
of particle agglomerates. Figure 6. Comparison of the Si and phosphocholine distribution in a cryo-section of the rat lung
30 min post administration of Fe-Si-NP. All images contain information of the uppermost 20 scans
only. (A) 28Si+; (B) C5H15NPO4+, which is the 184.07 fragment of the phosphocholine (PC) head group
showing the distribution phosphatidylcholine; and (C) the overlay of (A,B). 3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Finding
3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings In this study, most of the nanomaterial was found closely related to the alveolar septa by both
optical and mass spectrometric techniques (Figures 3 and 4G). In addition, a partial uptake into
macrophages was observed despite the relatively short incubation time of 30 min. The obviously
persistent macrophage activity suggests that the intratracheal instillation of H2O, which was used to
In this study, most of the nanomaterial was found closely related to the alveolar septa by both
optical and mass spectrometric techniques (Figures 3 and 4G). In addition, a partial uptake into
macrophages was observed despite the relatively short incubation time of 30 min. The obviously 11 of 14 Nanomaterials 2018, 8, 571 persistent macrophage activity suggests that the intratracheal instillation of H2O, which was used
to favor the dispersion of the polar NP, has no negative impact on the micro-milieu of the lung. Nevertheless, larger agglomerates of Fe-Si-NP with diameters of several micrometers were found in
the lung. These agglomerates may have formed upon contact with the lung lining fluid and/or during
lymphatic retrieval of the aqueous suspension fluid. As such, an agglomeration of nanoparticles in
lung instillation studies is common [29], although, e.g., citrate-stabilized gold nanoparticles do not
agglomerate under these conditions [30]. Nanomaterials 2018, 7, x FOR PEER REVIEW
11 of 14
In any case, it appears plausible that Fe-Si-NP, which had a size of less than 200 nm (see Figure gg
In any case, it appears plausible that Fe-Si-NP, which had a size of less than 200 nm (see Figure 1)
when they were administered to the lung, may have come into direct contact with several well
organized layers of biomolecules such as the lung surfactant, the hypophase, and/or the membrane of
(epithelial) cells. It appears conceivable that this sequential arrangement of biomolecules and ions in
the lung orchestrates the formation of the protein corona as well as agglomeration of NP. A protein
corona, which can be artificially generated with different preparations of lung surfactant [11] may,
therefore, differ from the native corona. In any case, it appears plausible that Fe Si NP, which had a size of less than 200 nm (see Figure
1) when they were administered to the lung, may have come into direct contact with several well
organized layers of biomolecules such as the lung surfactant, the hypophase, and/or the membrane
of (epithelial) cells. 3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Finding
3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings (A) 28Si+; (B) C5H15NPO4+, which is the 184.07 fragment of the phosphocholine (PC) head group
showing the distribution phosphatidylcholine; and (C) the overlay of (A,B). Arrows in all images
point to the position of larger agglomerates of Fe-Si-NP; note that in (B) PC is not detected at the
indicated sites. Figure 6. Comparison of the Si and phosphocholine distribution in a cryo section of the rat lung
30 min post administration of Fe-Si-NP. All images contain information of the uppermost 20 scans
only. (A) 28Si+; (B) C5H15NPO4+, which is the 184.07 fragment of the phosphocholine (PC) head group
showing the distribution phosphatidylcholine; and (C) the overlay of (A,B). Arrows in all images
point to the position of larger agglomerates of Fe-Si-NP; note that in (B) PC is not detected at the
indicated sites. Figure 6. Comparison of the Si and phosphocholine distribution in a cryo section of the rat lung
30 min post administration of Fe-Si-NP. All images contain information of the uppermost 20 scans
only. (A) 28Si+; (B) C5H15NPO4+, which is the 184.07 fragment of the phosphocholine (PC) head group
showing the distribution phosphatidylcholine; and (C) the overlay of (A,B). Arrows in all images
point to the position of larger agglomerates of Fe-Si-NP; note that in (B) PC is not detected at the
indicated sites. This finding seems to be in contrast to a recent investigation on lipid binding to SiO2, ZrO2, or
AlOOH in vitro [11]. Using two different preparations of pulmonary surfactant, it was found that
polar nanoparticles bind indeed to phospholipids, if the mixture (Curoserve®) contained surfactant
protein A. However, the role of surfactant proteins for an in-situ opsonization of nanoparticles is still
not known. As mentioned above, the incubation of nanoparticles in artificial or isolated pulmonary
surfactant may be profoundly different from binding conditions in the lung. While in the lung the
air–liquid interface carries non-polar lipid residues facing the gas phase, lipids in isolated lung
surfactant may form micelles with their polar residues directed outwards. The current results provide for the first time new and more realistic insights in the formation
This finding seems to be in contrast to a recent investigation on lipid binding to SiO2, ZrO2,
or AlOOH in vitro [11]. Using two different preparations of pulmonary surfactant, it was found that
polar nanoparticles bind indeed to phospholipids, if the mixture (Curoserve®) contained surfactant
protein A. 3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Finding
3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings Arrows in all images
point to the position of larger agglomerates of Fe-Si-NP; note that in (B) PC is not detected at the
indicated sites. 300
250
200
150
100
50
0
300
200
100
0
μm
Phosphatidylcholine
MC: 30; TC: 2.287e+006
25
20
15
10
5
0
300
250
200
150
100
50
0
300
200
100
0
μm
28Si+ (scan 1-20)
MC: 5; TC: 3.396e+004
4
3
2
1
300
250
200
150
100
50
0
300
200
100
0
μm
Overlay of Phosphatidylcholine,
28Si (scan1-20)
Figure 6. Comparison of the Si and phosphocholine distribution in a cryo-section of the rat lung
30 min post administration of Fe-Si-NP. All images contain information of the uppermost 20 scans
only. (A) 28Si+; (B) C5H15NPO4+, which is the 184.07 fragment of the phosphocholine (PC) head group
showing the distribution phosphatidylcholine; and (C) the overlay of (A,B). Arrows in all images
point to the position of larger agglomerates of Fe-Si-NP; note that in (B) PC is not detected at the
indicated sites. 300
250
200
150
100
50
0
300
200
100
0
μm
Phosphatidylcholine
MC: 30; TC: 2.287e+006
25
20
15
10
5
0 300
250
200
150
100
50
0
300
200
100
0
μm
Overlay of Phosphatidylcholine,
28Si (scan1-20) 300
250
200
150
100
50
0
300
200
100
0
μm
28Si+ (scan 1-20)
MC: 5; TC: 3.396e+004 Phosphatidylcholine
MC: 30; TC: 2.287e+006 (scan 1 20)
5; TC: 3.396e+004 Figure 6. Comparison of the Si and phosphocholine distribution in a cryo-section of the rat lung
Figure 6. Comparison of the Si and phosphocholine distribution in a cryo-section of the rat lung Figure 6. Comparison of the Si and phosphocholine distribution in a cryo-section of the rat lung
30 min post administration of Fe-Si-NP. All images contain information of the uppermost 20 scans
only. (A) 28Si+; (B) C5H15NPO4+, which is the 184.07 fragment of the phosphocholine (PC) head group
showing the distribution phosphatidylcholine; and (C) the overlay of (A,B). Arrows in all images
point to the position of larger agglomerates of Fe-Si-NP; note that in (B) PC is not detected at the
indicated sites. Figure 6. Comparison of the Si and phosphocholine distribution in a cryo-section of the rat lung
30 min post administration of Fe-Si-NP. All images contain information of the uppermost 20 scans
only. 3.7. Methodical Considerations It is well known that any type of tissue preparation may have an influence on bio-imaging
results. As will be described below (3.2–3.4), tissue sections analyzed by ToF-SIMS were taken from
lavaged lungs, which were filled with a cryomatrix and immediately snap-frozen in liquid nitrogen. Although filling with a cryomatrix preserves the tissue architecture of the lung, the compound does
not fix subcellular structures, such that a reliable distinction between cell-attached and intracellular
NP cannot be made. Furthermore, sections were cut, dried onto ITO slides, stored in a frozen state,
and eventually fixed with formalin on the slide. We are aware that this procedure cannot prevent
a partial loss of NP, especially if they are not in contact with fixable tissue components. However,
numerous particles and agglomerates proved to be firmly attached, because even the withdrawal of
the coverslip prior to ToF-SIMS analysis left the majority of particles in place, as shown in Figure 5. There may also be a partial loss or wash-out of lipids, as these molecules are not fixable with
formalin. However, neither ethanol nor other organic solvents were used, such that PC molecules
remained highly abundant in all tissue structures. Nevertheless, there was a low background signal of
PC in alveolar areas, which may be interpreted as an artificial displacement of PC. However, as levels
of PC were low in the vicinity of the alveolar septa (Figure 6B), we assume that PC found within alveoli
was not displaced from cellular structures. Rather, it may reflect a mixture of lung surfactant and
liquid cryomatrix which dried onto the ITO carrier. Considering all these facts, we cannot completely
rule out that the lack of PC on Fe-Si-NP was partially artificial, but the complete absence of PC over all
Si signals (see Figure 6C) makes this interpretation highly unlikely. Finally, the lack of PC on Fe-Si-NP might also hint at an inappropriate sensitivity of the ToF-SIMS
method. However, in a previous investigation on the composition of Langmuir–Blodgett films, PC and
other phospholipids were successfully identified in a lung surfactant-like monolayer [32]. Thus, although some artificial displacement of particles and/or a loss of PC cannot completely
be ruled out, we suggest that the absence of PC on the silica surface of the Fe-Si-NP in the
lung, as measured by the highly sensitive ToF-SIMS method, is not an artifact and, therefore,
of biological significance. 3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Finding
3.6. Lack of Colocalization of Phophatidylcholine with Fe-Si-NP and General Impact of Findings However, the role of surfactant proteins for an in-situ opsonization of nanoparticles is still
not known. As mentioned above, the incubation of nanoparticles in artificial or isolated pulmonary
surfactant may be profoundly different from binding conditions in the lung. While in the lung
the air–liquid interface carries non-polar lipid residues facing the gas phase, lipids in isolated lung
surfactant may form micelles with their polar residues directed outwards. 12 of 14 Nanomaterials 2018, 8, 571 The current results provide, for the first time, new and more realistic insights in the formation of
the lipid/protein corona around silica nanoparticles under native conditions of the lung. Findings may
be representative for pulverulent silica nanomaterials, unless they are coated, e.g., by hydrophobic
molecules. These findings have been achieved with high-resolution ToF-SIMS, a state-of-the-art
bio-imaging method, which is universally applicable and circumvents the need to isolate particles
from tissues. Future developments of the technique such as an ultra-high mass resolution detector are
in sight, which may further increase the performance of the method [31]. This will allow us to obtain
more detailed analyses of molecules bound to NP under in vivo conditions. References 1. Chaudhry, Q.; Scotter, M.; Blackburn, J.; Ross, B.; Boxall, A.; Castle, L.; Aitken, R.; Watkins, R. Applications
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interaction on short-term inhalation toxicity. Nanotoxicology 2016, 10, 970–980. [CrossRef] [PubMed] 12. Nanomaterials 2018, 8, 571 Nanomaterials 2018, 8, 571 Author Contributions: L.V., D.B., C.E. and M.W., conceived and designed the experiments. L.V. and D.B. performed the ToF-SIMS analyses. B.H., D.B., and L.V. analyzed the ToF-SIMS data. A.V. and M.W. performed the
animal experiments and contributed the tissue thin sections. L.V. and M.W. wrote the paper. Author Contributions: L.V., D.B., C.E. and M.W., conceived and designed the experiments. L.V. and D.B. performed the ToF-SIMS analyses. B.H., D.B., and L.V. analyzed the ToF-SIMS data. A.V. and M.W. performed the
animal experiments and contributed the tissue thin sections. L.V. and M.W. wrote the paper. Funding: This research was funded by German Federal Ministry of Education and Research (BMBF), grant number
[03X0146A and 03X0146C]. BMBF also covered the costs for Open Access publishing. Funding: This research was funded by German Federal Ministry of Education and Research (BMBF), grant number
[03X0146A and 03X0146C]. BMBF also covered the costs for Open Access publishing. Acknowledgments: Authors would like to thank Christian Schechtmann for excellent technical assistance. Acknowledgments: Authors would like to thank Christian Schechtmann for excellent technical assistance. Conflicts of Interest: The authors declare no conflict of interest. The founding sponsors had no role in the design
of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, and in the
decision to publish the results. Conflicts of Interest: The authors declare no conflict of interest. The founding sponsors had no role in the design
of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, and in the
decision to publish the results. 4. Conclusions In this study on paramagnetic silica nanoparticles (Fe-Si-NP) administered to the lung via
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of the Fe-Si-NP remains intact under these conditions, such that Fe-Si-NP may now be used to study
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enhanced dark-field microscopy. Toxicol. Pathol. 2018, 46, 28–46. [CrossRef] [PubMed] 29. Creutzenberg, O.; Bellmann, B.; Korolewitz, R.; Koch, W.; Mangelsdorf, I.; Tillmann, T.; Schaudien, D. Change
in agglomeration status and toxicokinetic fate of various nanoparticles in vivo following lung exposure in
rats. Inhal. Toxicol. 2012, 24, 821–830. [CrossRef] [PubMed] 30. Gosens, I.; Post, J.A.; de la Fonteyne, L.J.; Jansen, E.H.; Geus, J.W.; Cassee, F.R.; de Jong, W.H. Impact of agglomeration state of nano- and submicron sized gold particles on pulmonary inflammation. Part. Fibre Toxicol. 2010, 7, 37. [CrossRef] [PubMed] 31. Passarelli, M.K.; Pirkl, A.; Moellers, R.; Grinfeld, D.; Kollmer, F.; Havelund, R.; Newman, C.F.; Marshall, P.S.;
Arlinghaus, H.; Alexander, M.R.; et al. The 3d orbisims—Label-Free metabolic imaging with subcellular
lateral resolution and high mass-resolving power. Nat. Methods 2017, 14, 1175. [CrossRef] [PubMed] g
g p
32. Breitenstein, D.; Batenburg, J.J.; Hagenhoff, B.; Galla, H.J. Lipid specificity of surfactant protein b studied by
time-of-flight secondary ion mass spectrometry. Biophys. J. 2006, 91, 1347–1356. [CrossRef] [PubMed] © 2018 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access
article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/).
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Catchment Soil Properties Affect Metal(loid) Enrichment in Reservoir Sediments of German Low Mountain Regions
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Applied sciences
| 2,022
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cc-by
| 538
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License: CC BY 4.0 International - Creative Commons, Attribution Supplementary material Table S1. Precautionary values according to the German Federal Soil Protection Ordinance (mg kg‐1). Texture
As
Cd
Cr
Cu
Fe
Hg
Mn
Ni
Pb
V
Zn
Sand
-
0.4
30
20
-
0.1
-
15
40
-
60
Silt or Loam
-
1.0
60
40
-
0.5
-
50
70
-
150
Clay
-
1.5
100
60
-
1
-
70
100
-
200
1 The application on arable land requires that the metal concentrations of the applied material do not exceed 70%
of the precautionary values. Table S1. Precautionary values according to the German Federal Soil Protection Ordinance (mg kg‐1). 1 The application on arable land requires that the metal concentrations of the applied material do not exceed 70%
of the precautionary values. Table S2. Rotated component matrix of the three analyzed components (EV: Eigen value; Vt (%): percentage of
total variance; Vc (%): cumulative percentage of variance). Table S2. Rotated component matrix of the three analyzed components (EV: Eigen value; Vt (%): percentage of
total variance; Vc (%): cumulative percentage of variance). Table S2. Rotated component matrix of the three analyzed components (EV: Eigen value; Vt (%): percentage of
total variance; Vc (%): cumulative percentage of variance). total variance; Vc (%): cumulative percentage of variance). Comp. 1
2
3
Cr
.942 .156 ‐.051
Al
.905 .096 ‐.021
V
.903 .229 .026
Fe
.854 .333 ‐.173
As
.784 .215 .049
CN
‐.536 ‐.349 .454
Zn
.478 .803 .130
Cd
.244 .762 .447
pH
.167 .756 ‐.274
Ni
.561 .727 .000
Mn
.135 .654 ‐.007
Pb
.012 .463 .328
TOC
‐.156 ‐.030 .957
Nt
.068 .180 .925
EV
4.83 3.37 2.42
Vt (%) 34.5 24.0 17.2
Vc (%) 34.5 58.5 75.8 Table S3. Percentage of easily extractable trace metal concentrations on the aqua regia soluble concentrations of
air‐dried reservoir sediments (RS). Table S3. Percentage of easily extractable trace metal concentrations on the aqua regia soluble concentrations of
air‐dried reservoir sediments (RS). As
Cd
Cr
Fe
Mn
Ni
Pb
V
Zn
Mean
1.26 10.9 0.07 0.02 62.1
4.41
0.19
0.03 2.93
Median
0.59 7.17 0.06 0.01 71.2
3.30
0.06
0.02 0.67
SD
1.92 9.61 0.03 0.02 36.8
5.43
0.29
0.02 4.87
Min
0.20 0.88 0.02 0.00 0.73
0.28
0.01
0.01 0.13
Max
9.31 38.1 0.15 0.06
100
20.5
1.13
0.13 19.4 License: CC BY 4.0 International - Creative Commons, Attribution Table S4. Supplementary material The variability of the median values of the element‐specific enrichment factors (EF) illustrated by the
example of the Breitenbach, Obernau and Bitburg reservoirs (RS: reservoir sediment; SS: stream sediment; CS:
catchment soil). As
Cd
Cr
Fe
Mn
Ni
Pb
V
Zn
Breitenbach
RS
SS
CS
4.3
7.1
6.3
10.2
13.9
2.4
2.8
3.7
3.5
5.3
8.9
5.2
40.5
19.0
3.1
8.9
20.9
3.9
21.2
42.6
36.2
3.0
4.0
4.1
10.4
14.0
3.3
Obernau
RS
SS
CS
3.0
3.8
3.9
8.2
2.2
2.5
3.2
3.2
3.1
4.2
4.9
4.3
27.6
7.6
4.4
7.4
6.5
3.9
6.0
4.7
6.2
3.2
2.4
3.0
6.4
3.5
2.9
Bitburg
RS
SS
CS
2.9
4.7
2.9
2.5
1.7
1.1
2.8
3.4
2.7
3.9
5.4
4.0
3.5
6.0
5.9
4.9
5.8
4.2
8.9
3.5
2.4
3.0
2.9
2.4
4.1
3.3
2.5
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English
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Adaptive Motion Estimator Based on Variable Block Size Scheme
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Zenodo (CERN European Organization for Nuclear Research)
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cc-by
| 5,202
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World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 Authors are with Electronics and Micro-Electronics Laboratory (LAB-
IT06), Faculty of Sciences of Monastir, Monastir, 5000, Tunisia.
the image and the handled video. The choice of the research
strategy to be used in the stage of motion estimation is International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 In order to efficiently reduce the computational cost while
achieving better visual quality with low cost power, the proposed
motion estimator is based on a Variable Block Size (VBS) scheme
that uses only the 16x16, 16x8, 8x16 and 8x8 modes. The computational cost of a motion estimation algorithm
is greatly affected by the number of candidate blocks to be
computed. In order to compute the Somme of Absolute
Difference (SAD), the FBMA evaluates every possible
motion vector inside the search area, find the best matched
block and give the highest PSNR. Other strategies are based
on the so called fast block motion estimation algorithms
group which includes the Three-Step Search algorithm
(TSS), the Modified Log Search algorithm (LS), the Four
Steep Search (FSS) and the Alternative Pixel-Decimation
Search algorithm (APDS) [3, 4, 5, 6, 7, 8, 9]. These
algorithms take into account a reduced group of available
candidate blocks at the expense of a reduction in term of
PSNR. Experimental results show that the adaptive motion estimator
allows better results in term of Peak Signal to Noise Ratio
(PSNR), computational cost, FPGA occupied area, and dissipated
power relatively to the most popular variable block size schemes
presented in the literature. Keywords—H264, Configurable Motion Estimator, Variable
Block Size, PSNR, Dissipated power. Adaptive Motion Estimator Based on Variable
Block Size Scheme S. Dhahri, A. Zitouni, H. Chaouch, and R. Tourki decisive in the total execution and on the video quality. One
of the principal strategies to obtain an effective
compression is to use a technique which exploits the space
and temporal redundancy. Two main techniques are used
for motion estimation: pixel recursive algorithms and block
matching algorithm (BMA). The first one estimates the
motion between successive frames on a pixel by pixel base,
whereas the BMA estimates motion on a block by block
basis. In general, due to its simple hardware realization, the
block-matching approach is the most suitable. Abstract—This paper presents an adaptive motion estimator
that can be dynamically reconfigured by the best algorithm
depending on the variation of the video nature during the lifetime
of an application under running. The 4 Step Search (4SS) and the
Gradient Search (GS) algorithms are integrated in the estimator in
order to be used in the case of rapid and slow video sequences
respectively. The Full Search Block Matching (FSBM) algorithm
has been also integrated in order to be used in the case of the
video sequences which are not real time oriented. A. Four Step Search International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 munication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 The Four-Step Search (4SS) is an algorithm that builds
upon the TSS, while being more center-based. The 4SS
differs in that it maintains a more regular search pattern
with half-stop techniques employed in the algorithm [6]. The 4SS offers a similar best-case scenario for
computational complexity when compared to the TSS (17
searches). The worst-case situation for the 4SS is 27 search
points compared to 33 for the TSS, which is an
improvement. Experimental studies performed on multitude
standard video sequences show that this algorithm is well
adapted to the fast sequences by giving the better PSNR
and compression rates relatively to other algorithms. International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Pu 4 SS
GS
Adaptive motion estimator
Rapid
Slow
Two motion types
Fig. 1 Principal of the adaptive motion estimator Rapid Fig. 2 Four Step Search paths Fig. 2 Four Step Search paths Adaptive motion estimator Two motion types Fig. 1 Principal of the adaptive motion estimator This paper is organized as follow: Section 2 presents
brief overview of the Block Matching Algorithms (BMA)
which are integrated in the adaptive motion estimator (4SS,
GS, FS). In section 3, the architecture of the proposed
motion estimator is presented. The proposed VBS scheme
is described in section 4. Section 5 presents some
experimental results and Section 6 concludes the paper. Fig. 2 Four Step Search paths Fig. 2 Four Step Search paths World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 split [12], and low complexity motion estimation [13], etc. (
)
∑∑
=
=
+
+
−
−
=
16
1
i
16
1
j
)
l
j,
k
i(
1
tP
j,i
tP
)
l,
k
(
SAD split [12], and low complexity motion estimation [13], etc. In general, since, a video application can contain many
sequences that are different in terms of motion nature
(slow, stationary, fast), an adaptive motion estimation
scheme is very suitable to improve performances of a video
Codec (Fig. 1). Where (k, l) is the location in the search window, Pt(i, j) is
a pixel at (i, j) in the current frame, and Pt−1(i, j) is a pixel
in the previous frame. When the value SAD(k,l) is
minimum, (k,l) is the motion vector of the macro block. This paper presents the proposed adaptive motion
estimator that can be dynamically configured by three
algorithms (FSBM, 4SS, GS) depending on the variation of
the video nature during the lifetime of a running
application. This configuration is based on a recognition
step that collects motion information of each block from its
same block in the previous images and their neighbor’s
blocks in the current image. Contrary to many solutions
proposed in the literature, the proposed approach is based
on an efficient technique for decomposing the motion
block. In fact, a new VBS motion estimation algorithm
based on 16x16, 16x8, 8x16 and 8x8 only has been
proposed, which can efficiently reduce the computational
cost while achieving similar or better visual quality with
low cost power on different architectures. The advantages of implementing FSBM as the motion
estimation algorithm include both the guaranteed optimality
of the solution and the regularity of a hardware
implementation. This regularity stems from the fact that
consecutive motion vectors share many of the pixel values
in calculation. nce Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 B. Gradient Search The block-based gradient descent search (BBGDS)
algorithm has been proposed by literature [5]. Based on the
observation that global minimum distribution is centralized
in real video sequence, this algorithm uses a very center-
based search pattern of nine checking points in each step
with a step size of one. It does not restrict the number of
searching steps but it stops when the minimum checking
point of the current step is the central one or it reaches the
search window boundary. This algorithm has advantages:
very low complexity in terms of candidate to evaluate, good
regularity in terms of motion vector generation and only I.
INTRODUCTION IDEO communication is a rapidly evolving field for
several applications which include video telephony,
videoconference, remote surveillance, remote working and
learning, etc. It is also a key feature for the upcoming
information and communication technologies based on
residential digital lines (VDSL, ADSL and ISDN) and the
3rd generation of mobile telephony system (UMTS). In this
scenario, video image compression plays a fundamental
role in reducing the enormous bit-rate for transmission and
storage (approximately hundreds of Mbits/s for main image
formats). To this objective, the ISO and the ITU-T
committees have worked on several compression standards
such as JPEG, MPEG (versions 1, 2, 4), H.261, H.263 and
H.26L [1, 2]. V V Experimental results performed in [10] show that the full
search algorithm is very greedy in computing times, but
always gives the best results in compression ratio and
PSNR (Fig. 1). This work shows also that in the case of fast
sequences (ex. Forman) the 4SS algorithm gives better
PSNR and compression rates relatively to other algorithms. In the case of slow sequences (ex. Miss America), [10]
shows that the GS algorithm is more advantageous since the
best block are always very close to the fetched block. By
studying the sequences that contain the two types of motion
(fast and slow) (ex. News and Carphone), [10] shows that
the results are very close and speed can be improved by
considering methods of research adapted for each type of
block. Motion estimation is regarded as one of the most
effective technique to reduce the flow required by a video
codec. This stage is the most expensive in computing times
and power consumption for the standard H264. Indeed, the
method of estimation influences enormously the quality of The motion estimation for each 16x16 macro-block in
the emerging H.264/AVC can be divided into 16x16, 16x8,
8x16, and 8x8 modes. If the 8x8 mode is chosen, each 8x8
block can be independently coded into 8x4, 4x8, and even
4x4 blocks. Consequently, many VBS motion estimation
algorithms has been proposed in literature, such as bottom-
up merge and top-down split [11], prediction by merge and scholar.waset.org/1307-6892/14948 International Scholarly and Scientific Research & Innovation 3(2) 2009 International Scholarly and Scientific Research & Innovation 3(2) 2009 218 International Scholarly and Scientific Research & Innovation 3(2) 2009 C. Full Search This algorithm defines a rectangular geometry for the
image areas, consequently, the research window is limited
to [- p, p] which is the area of research around the origin of
the block in the reference image (Fig. 4). For an image with
a size of (2p+1) × (2p+1) and for a window with a size of
M*M and a block with a size of N ×N, the number of
operations that are treated by the FSBM algorithm for each
block is Nb = (2p+1)2×N2. The number of operations for
each window is Nf = (2p+1)2×M2. Since this algorithm is
the slowest in term of computing times, it is generally used
for the applications with high image quality. It allows the
sweeping of all the blocks of the research window. This
makes it possible to reach the adequate motion vector in
this window. 4 -Phase Bus
Reference
Memory
SAD
FSM
FSBM
FSM
4SS
FSM
GS
Configuration
Module
Current
Memory
Fig. 5 Optimal architecture of the estimator Configuration
Module Fig. 4 Method of search for the full search algorithm Fig. 5 Optimal architecture of the estimator In our realization, each algorithm is specified by its
control part that is described as a Finite State Machine
(FSM). The architecture is optimized in order to make the
redundant blocks reusable by the three algorithms (Fig. 5). According to the algorithm to be implemented, the
corresponding FSM communicates with the other blocks
via a 4-Phase handshaking bus. According to the
application to be treated, the estimator is configured by the
suitable algorithm. This allows the use of a single estimator
for various applications, which makes it possible to
guarantee an effective video compression. The estimator is
composed of a set of modules which are configured with
the proposed VBS scheme. The calculated SAD is stored in
a memory. If the scanning of the search window is finished,
then the memorized SAD is compared and the minimal
SAD will be detected. Then a new research can be started. The optimal SAD with its position is sent towards a buffer Fig. 4 Method of search for the full search algorithm This algorithm selects the minimum among the possible
vectors inside the research window (the motion vector
belongs to the area [- p, p]). This approach is inadaptable
by the majority of real time applications because of the high
number of operations that is performed. II. BLOCK MATCHING ALGORITHMS The FSBM algorithm is the most straightforward motion
estimation operation. The process of block-matching is
found in a search window of previous frames of the macro
blocks most similar to the macro blocks in the current
frame. The accuracy of ME depends on the matching
criteria and one of the most popular criteria is the sum of
absolute difference (SAD) given by: scholar.waset.org/1307-6892/14948 International Scholarly and Scientific Research & Innovation 3(2) 2009 219 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 portion of search area memory can be accessed. Experimental studies performed on multitude standard
video sequences show that this algorithm is well adapted to
the slow sequences by giving the better PSNR and
compression rates relatively to other algorithms algorithm suffers from greatest data-processing complexity. For example, considering the research area of [- 7, +7],
there are 226 calculations steps of the SAD which must be
carried out for each macro block in a window. III. ADAPTIVE MOTION ESTIMATOR MODELING Fig.3 Gradient Search paths The motion estimation suggested integrates three
algorithms: FSBM, 4SS and GS. The 4 modes (16x16,
16x8, 8x16 and 8x8) defined by the proposed VBS
technique are integrated in this estimator. Knowing that all
the motion estimators use memory to store the pixels of the
block under running as well as block of reference and carry
out the SAD calculation, these modules were defined only
once inside the proposed architecture. Thus, each technique
of motion estimation is defined only by its controller. Once
configured, the selected controller communicates with the
common SAD module inside architecture by using the
memory common resources and ensures the estimation
according to the algorithm of estimation in question. Such a
strategy allows the estimator to be configured according to
the nature of the application under running. Generally, the
new Codec use motion estimation which integrates only
one algorithm for each application [14, 15, 16, 17, 18]. For
an application which handles video sequences of variable
types, several estimators are normally necessary in order to
guarantee an acceptable video quality for each type of
sequence. nce Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/P Fig.3 Gradient Search paths memory. 6 Algorithmic description of the Full Search algorith
International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 dex, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 For (k = -p; k ≤ (p- bloc_col_v+1); k++)
For l = -p; l≤ (p- bloc_line_v +1); l++)
req <= '1';
For (r = 0; r≤ length_v-1; r++)
data_cour <= var_cour(r);
data_ref <= var_ref(r);
End;
For v = 0; v≤length_v-1; v++)
var_cour (v) <= "00000000";
End;
wait until ack ='1';
motion_vec (z) <= sad_in;
motion_vec2 (z, 0) <= k;
motion_vec2 (z, 1) <= l;
Req <= '0';
z: = z+1;
wait until ack ='0';
For (m = k; m≤k+ (bloc_col_v-1) ; m++)
For (n = l; n ≤ l+ (bloc_line_v-1); n++)
var_cour (v) <= memory (m, n);
v: =v+1;
End;
End;
End;
End;
For (h = 0; h≤ nsad_v -1; h++)
If (motion_vec (h) <= sad_min) then
sad_min:= motion_vec (h);
posit_x:= motion_vec2 (h, 0);
posit_y:= motion_vec2 (h, 1);
Else
sad_min:= sad_min;
End if;
sum_sort0 <= sad_min;
x_sort0 <= posit_x;
y_sort0 <= posit_y;
End;
Fig. 6 Algorithmic description of the Full Search algorithm
International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 Fig. 7 The window read/write protocol memory. The proposed motion estimator has been designed by
Register Transfer Level (RTL) level by using the VHDL
language. After being calculated by the chosen algorithm,
the calculated SAD values are memorized in a memory in
order to define for each block the starting point in the
research window. All the calculated SAD values of the
window are stored in a memory buffer and the minimal
value is determined by trying. Then the output of the
corresponding algorithm search block is the value of the
minimal SAD and their positions. In other terms, the
outputs are the motion vectors which will be used for the
reconstruction of the image by the coder. Fig. 7 The window read/write protocol For (k = -p; k ≤ (p- bloc_col_v+1); k++)
For l = -p; l≤ (p- bloc_line_v +1); l++)
req <= '1';
For (r = 0; r≤ length_v-1; r++)
data_cour <= var_cour(r);
data_ref <= var_ref(r);
End;
For v = 0; v≤length_v-1; v++)
var_cour (v) <= "00000000";
End;
wait until ack ='1';
motion_vec (z) <= sad_in;
motion_vec2 (z, 0) <= k;
motion_vec2 (z, 1) <= l;
Req <= '0';
z: = z+1;
wait until ack ='0';
For (m = k; m≤k+ (bloc_col_v-1) ; m++)
For (n = l; n ≤ l+ (bloc_line_v-1); n++)
var_cour (v) <= memory (m, n);
v: =v+1;
End;
End;
End;
End;
For (h = 0; h≤ nsad_v -1; h++)
If (motion_vec (h) <= sad_min) then
sad_min:= motion_vec (h);
posit_x:= motion_vec2 (h, 0);
posit_y:= motion_vec2 (h, 1);
Else
sad_min:= sad_min;
End if;
sum_sort0 <= sad_min;
x_sort0 <= posit_x;
y_sort0 <= posit_y;
End;
Fig. A. Variable Block Size 1 (VBS1) Unlike previous standards, H.264/AVC adopts a tree-
based decomposition to partition the macro-block (MB)
into smaller sub-blocks of specified sizes. The quad-tree
structure enables the possibility of a MB being coded in
four different modes illustrated in Fig. 8, with partitions
sizes of 16x16, 16x8, 8x16 and 8x8, while in the 8x8
partition mode, each 8x8 partition can be further split into
8x4, 4x8, and 4x4 sub-partitions. The availability of smaller
ME blocks improves prediction in general, and in
particular, the small blocks improve the ability of the model
to handle fine motion detail and result in better subjective
viewing quality because they do not produce large blocking
artifacts. Fig. 8 Multiple motion estimation modes defined in H.264/AVC Fig. 8 Multiple motion estimation modes defined in H.264/AVC Fig. 8 Multiple motion estimation modes defined in H.264/AVC C. Full Search Obviously, this
algorithm always finds the optimal motion vector for the
given research window. It should also be obvious, that this scholar.waset.org/1307-6892/14948 International Scholarly and Scientific Research & Innovation 3(2) 2009 220 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 and reference). The setting to '0' of this signal allows the
beginning of the sweeping of the window. For a new
window the same operation will be repeated. An
algorithmic description of the Full Search algorithm is
represented by Fig. 6. memory. V. EXPERIMENTAL RESULTS To evaluate the performances of the proposed adaptive
motion
estimator,
both
software
and
hardware
implementations have been performed. The impact of the
new VBS3 motion scheme on the encoding process, along
with several analyses on three CIF (352x288) sequences
test with different characteristics is proposed. At the
beginning, the performances evaluation of the VBS3
algorithm compared to the VBS1 and VBS2 on a SW DSP
(TMS320C64) design platform are presented. The used
parameters consists on a QP =38 and a search window
(Horizontal: [-15, 15] and Vertical:[-15, 15]). Finally, the
logic synthesis and the power estimation results of the
proposed estimator with the VBS3 scheme are presented
and compared with the VBS1 and VBS2 schemes. These
designs target the Altera FPGA technology and performed
by the ISE and the X-Power tools. ndex, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 B. Variable Block Size 2 (VBS2) Serious experiments on the test video sequences used in
JVT Test Model Ad Hoc Group [11] show that there is an
average of 35 per cent homogeneous area in a typical video
frame, and these areas are suitable for larger size inter mode
coding. Therefore, several cost calculation of small size
modes can be saved. Based on this consideration, several
mode decision algorithms were proposed to reduce the
number of candidate modes [11, 12]. In [19], many Fast
VBS Motion Estimation algorithms were tested before
extracting the Zoom Motion Estimation (ZME) algorithm
[19] based on 3 partitions sizes 16x16, 8x8 and 4x4
arranged as shown in Fig. 9. Fig. 6 Algorithmic description of the Full Search algorithm In the proposed architecture, the presence of a new
window is indicated by a command signal (cmd_fen). This
signal allows the initialization of the memory. After
initialization, the algorithm begins the treatment by a signal
called wr_ena (Fig. 7). The setting to '1' of the signal
wr_ena makes it possible to charge a new window (current International Scholarly and Scientific Research & Innovation 3(2) 2009 scholar.waset.org/1307-6892/14948 221 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 Fig. 9 Multiple motion estimation modes defined in H.264/AVC
org/Publication/14948 A. Subjective, Objective and Complexity Analysis International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Pu Fig. 9 Multiple motion estimation modes defined in H.264/AVC Table I and Table II show that the elimination of 4x8,
8x4 and 4x4 block size (performed by our VBS3 scheme)
from the mode selection does not affect the video quality
(PSNR and SSIM [20]). Thus, the VBS3 algorithm
provides better subjective and objective quality compared
to the VBS2 scheme with complexity reduction in terms of
clock cycles up to 44 per cent (Table III). C. Variable Block Size 3 (VBS3) C. Variable Block Size 3 (VBS3) The proposal of the new typical based VBS scheme is to
extract, using the SAD criteria, the block-size introducing
the minimum computational complexity with minimum
memory requirement. In this algorithm, only 16x16, 16x8,
8x16, and 8x8 block-sizes for motion estimation are
considered. This approach will reduce extensively the
computational complexity since the 3 other modes are
eliminated, without affecting the overall visual video
quality. For better understanding, the following flowchart
explains in detail our proposed algorithm steps Fig. 10. Fig. 11 Number of use modes for different variable block decision
algorithm International Science Index, Electronics and Communication En Fig. 10 Fast block size selection algorithm FBSA
International Science Index, Electronics and Communica Fig. 11 Number of use modes for different variable block decision
algorithm Fig. 11 Number of use modes for different variable block decision
algorithm Fig. 10 Fast block size selection algorithm FBSA International Scholarly and Scientific Research & Innovation 3(2) 2009 scholar.waset.org/1307-6892/14948 222 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 World Academy of Science, Engineering and Technology
International Journal of Electronics and Communication Engineering
Vol:3, No:2, 2009 motion estimator would be critical for inclusion in a
portable or other power-conscious device. These results demonstrate the fact that from one hand,
VBS3 outperforms the other variable block size techniques
used in this study, such as VBS1 and VBS2 in terms of MB
sizes (bit) distortion with a major loss in complexity. On the
other hand, VBS3 is very similar to the other VBS
predicting algorithms in terms of subjective quality with a
major gain in the computational complexity and a better
image quality depending on the motion characteristic of the
video. In fact, the visual image quality with a major
reduction in the computational effort complexity has been
objectively improved. However, the penalized is performed
only for the low motion sequence since the extra search will
not be necessary (Fig. 11). The gain in terms of the FPGA equivalent area designed
with our VBS and the VBS2 schemes is about +4,35%
relatively to the FPGA equivalent area designed by the
standard VBS1. Also the gain in terms of the power
dissipated by our VBS scheme is about +5,17% and
+1,79% relatively to the power dissipated by the VBS1 and
the VBS2 schemes (Table V). TABLE V
COMPARISON OF THE VBS SCHEMES IN TERMS OF FPGA EQUIVALENT
AREA AND DISSIPATED POWER TABLE V
COMPARISON OF THE VBS SCHEMES IN TERMS OF FPGA EQUIVALENT
AREA AND DISSIPATED POWER AREA AND DISSIPATED POWER
Equivalent area:
VBS2 vs. VBS1
)%
100
1
VBS
2
VBS
100
(
×
−
+4,35%
Equivalent area:
VBS3. vs. VBS1
)%
100
1
VBS
3
VBS
100
(
×
−
+4,35%
Dissipated power:
VBS2 vs. VBS1
)%
100
1
VBS
2
VBS
100
(
×
−
+3,45%
Dissipated power:
VBS3. vs. VBS1
)%
100
1
VBS
3
VBS
100
(
×
−
+5,17%
Dissipated power:
VBS3. vs. VBS2
)%
100
2
VBS
3
VBS
100
(
×
−
+1,79% TABLE II
SUBJECTIVE QUALITY PERFORMANCE (SSIM) OF BLOCK MODE
SELECTION SCHEMES TABLE II
SUBJECTIVE QUALITY PERFORMANCE (SSIM) OF BLOCK MODE
SELECTION SCHEMES TABLE III
SPEED PERFORMANCE (MILLIONS CYCLES) OF BLOCK MODE SELECTION
SCHEMES
B. RTL Design Results TABLE III
SPEED PERFORMANCE (MILLIONS CYCLES) OF BLOCK MODE SELECTION
SCHEMES VI. CONCLUSION In this paper a configurable motion estimator for the
codec H264 has been presented. This estimator integrates
three algorithms (FSBM, 4SS and GS) and implements a
VBS scheme. Depending on the nature of the application
under running this algorithm will be reconfigured
dynamically by the adequate algorithm. Since, the motion
estimator is responsible for nearly 70% of the power
dissipated and computation cost in certain video encoders,
any amelioration of the motion estimator performances
would be critical for inclusion in a portable or other power-
conscious device. Experimental results lead us to conclude
that the proposed estimator integrating our VBS scheme
allows better image quality with less computing time, less
FPGA occupied area and less dissipated power relatively to
the other VBS schemes. As a perspective, further design
targeting ASIC technology for proposed adaptive motion
estimator show that substantial improvements, in terms of
encoding speed and dissipated power, can be obtained
through optimizing heavily. This research work is ongoing,
and the results will be presented in future publications. C. Variable Block Size 3 (VBS3) TABLE IV
FPGA EQUIVALENT AREA AND POWER ESTIMATION RESULTS
Used VBS
Equivalent area
Dissipated power
(mW)
VBS1
23%
58
VBS2
22%
56
VBS3
22%
55 International Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 TABLE I
OBJECTIVE QUALITY PERFORMANCE (PSNR) OF BLOCK MODE SELECTION
SCHEMES
TABLE II
SUBJECTIVE QUALITY PERFORMANCE (SSIM) OF BLOCK MODE
SELECTION SCHEMES
TABLE III
SPEED PERFORMANCE (MILLIONS CYCLES) OF BLOCK MODE SELECTION
SCHEMES
B. RTL Design Results
In order to evaluate the occupied area and the dissipated
Science Index, Electronics and Communication Engineering Vol:3, No:2, 2009 waset.org/Publication/14948 TABLE I TABLE I
OBJECTIVE QUALITY PERFORMANCE (PSNR) OF BLOCK MODE SELECTION
SCHEMES REFERENCES [1]
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Jae Hun Lee, and Al, “Variable block size motion estimation
algorithm and its hardware architecture for H.264 /AVC”, IEEE Inter. Symp. on Circuits and Sys., May 2004. [10] A. Djeffal, and Z. Baarir, “Video coding adaptive block matching “,
8th African Conference on Research in Computer Science, November
2006, pp.1-8. [11] Y.K. Tu, J.F. Yang and M.T. Sun, Fast Variable-size Block Motion
Estimation Using Merging Procedure with an Adaptive Threshold,
IEEE International Conference on Multimedia and Expo. Baltimore,
July 2003, p.II-789-792. [12] Z. Zhou, M.T. Sun and Y.F. Hsu, Fast variable block-size motion
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H.264/MPEG-4 AVC, IEEE International Symposium on Circuits
and Syst., ISCAS. Vancouver, British Columbia, Canada, May 23-26,
2004.II-789-792. [13] Y. Jiang,S. Li and S. B. RTL Design Results In order to evaluate the occupied area and the dissipated
power of the proposed motion estimator, the ISE and the X-
Power FPGA design tools have been used. Table IV
presents the logic synthesis and the power estimation
results of the adaptive motion estimator configured with the
FSBM algorithm and integrating the VBS1, VBS2, and the
VBS3 schemes targeting the Virtex2pro FPGA technology. Experimental results show that the FPGA equivalent area
and the dissipated power obtained by our VBS scheme are
slightly inferior to those obtained by the VBS1 and the
VBS2 schemes. But since the motion estimator is
responsible for nearly 70% of the power dissipated in
certain video encoders, any reduction in power in the scholar.waset.org/1307-6892/14948 International Scholarly and Scientific Research & Innovation 3(2) 2009 223 World Academy of Science, Engineering and Technology
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Motion Estimation Algorithm for Video Telephony Communication,
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[14] M. G. Xavier, “Optimizing performance of an encoder following the
standard Advanced Video Coding for a vector machine,” Master
Memory, Faculty of Sciences libre University of Bruxcelle, 2006. [15] S. Yalcin, H.F. Ates, I.Hamzaoglu, “A high performance hardware
architecture for an SAD reuse based hierarchical motion estimation
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Memory, Faculty of Sciences of Orsay, November 2005. [17] LIU Hao, ZHANG Wen-jun, CAI Jun, “A fast block-matching
algorithm based on variable shape search”, Journal of Zhejiang
University Science A, Mars 2005. [18] Tiago Miguel Braga da Silva Dias, “High-Performance VLSI Motion
Estimation Processors: Data Reuse and Sub-Pixel Accuracy”, Master
Memoir, Univ. de Tec. de Lisboa, Inst. Sup. Tec., Sep. 2004. [19] M.A. Ben Ayed,A. Samet and N. Masmoudi, Toward an Optimal
Block Motion Estimation Algorithm for H.264/AVC, International
Journal of Image and Graphics (IJIG). 2006. [20] Z. Wang,A.C. Bovik,H.R. Sheikh and E.P. Simoncelli, Image quality
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ORIGINAL RESEARCH
published: 09 September 2021
doi: 10.3389/fvets.2021.688656 ORIGINAL RESEARCH
published: 09 September 2021
doi: 10.3389/fvets.2021.688656 Stefan Deleuze 1*†, Fany Brotcorne 2†, Roland Polet 2, Gede Soma 3, Goulven Rigaux 4,
Gwennan Giraud 2, Fanny Cloutier 2, Pascal Poncin 5, Nengah Wandia 3 and
Marie-Claude Huynen 2 Stefan Deleuze 1*†, Fany Brotcorne 2†, Roland Polet 2, Gede Soma 3, Goulven Rigaux 4,
Gwennan Giraud 2, Fanny Cloutier 2, Pascal Poncin 5, Nengah Wandia 3 and
Marie-Claude Huynen 2 Edited by: Reviewed by:
Paolo Martelli,
Ocean Park, Hong Kong, SAR China
Joanne Paul-Murphy,
University of California, Davis,
United States Worldwide, primates, and humans increasingly share habitats and often enter in conflict
when primates thrive in human-dominated environments, calling for special management
measures. Reproductive control is increasingly used to manage population growth but
very few monitoring data are available. Therefore, the efficiency and implications of such
programs require a careful examination. In the context of a contraception program in wild
female long-tailed macaques in Ubud, Bali, conducted over four successive campaigns
between 2017 and 2019, including 140 females (i.e., 41.9% of the reproductive females
of the population in 2019), modifications of an endoscopic tubectomy procedure, a
permanent sterilization method, clinical evaluation of this method, and the post-operative
monitoring results of the neutered females after release are described. This surgical
approach was applicable for pregnant females: 28.6% of the treated females were
pregnant at the time of the surgery. The procedure used a single lateral port to reach
and cauterize both oviducts in non-pregnant as well as in early to mid-term pregnant
females. Pregnant females nearer to term required a second lateral port to access both
oviducts masked by the size of the gravid uterus. Moreover, bipolar thermocauterization
was utilized successfully without resection to realize the tubectomy. The average duration
of the laparoscopic surgery was 14 min for non-pregnant females and 22 min for pregnant
females. Animals were released 3 h 22 min in average following their capture. This
short holding time, recommended for free-ranging primates, was made possible by
the minimal invasiveness of the sterilization approach. A laparoscopic post-operative
evaluation conducted on two patients during the following campaign confirmed that the *Correspondence:
Stefan Deleuze
s.deleuze@uliege.be *Correspondence:
Stefan Deleuze
s.deleuze@uliege.be †These authors share first authorship Specialty section:
This article was submitted to
Zoological Medicine,
a section of the journal
Frontiers in Veterinary Science
Received: 31 March 2021
Accepted: 09 August 2021
Published: 09 September 2021 Specialty section:
This article was submitted to
Zoological Medicine,
a section of the journal
Frontiers in Veterinary Science Received: 31 March 2021
Accepted: 09 August 2021
Published: 09 September 2021 1 Research Unit FARAH, Equine and Companion Animal Reproduction Pathologies Clinic, Veterinary Medicine Faculty,
University of Liège, Liège, Belgium, 2 Research Unit SPHERES, Department of Biology, Ecology and Evolution, Sciences
Faculty, University of Liège, Liège, Belgium, 3 Primate Research Center, Veterinary Medicine Faculty, Udayana University,
Denpasar, Indonesia, 4 Pan Anima, Tournai, Belgium, 5 Research Unit FOCUS, Department of Biology, Ecology and Evolution,
University of Liège, Liège, Belgium Edited by:
Kathryn Christine Gamble,
Lincoln Park Zoo, United States INTRODUCTION urges conducting systematic studies during and in the aftermath
of the fertility control interventions to monitor the impacts,
efficiency, and adequacy of the methods adopted with primates
(10, 19). As for laparoscopic tubectomy, clinical intra-, and post-
operative evaluations have been described in a small number
of rhesus (17) and Formosan macaques (M. cyclopis) (20) in
captive settings, and more recently in a long term program in field
conditions in free-ranging rhesus cross long-tailed macaques
(14). However, the application of laparoscopic tubectomy to
M. fascicularis in free-ranging settings, its clinical evaluation
and possible modification to optimize the technique as well
as a long-term post-surgery monitoring remain lacking and
need to be documented. Reasons for selecting this technique
with urban long-tailed macaques are manyfold but most of all,
the species’ reproductive biology [i.e., a multimale/multifemale
mating system where a female is likely to mate with multiple
males; (21)] and social dynamics [i.e., a female philopatry system
of resident matrilineal groups wherein demographic recruitment
is a function of births and male migrations; (22)], naturally orient
the choice toward female-based sterilization. Worldwide, a growing proportion of primate populations of
various species thrive in human-modified habitats (1). As a
consequence of the rapid land-use change, these populations are
increasingly forced to cohabit with humans (2). The macaques
(Macaca sp.) in particular share one of the widest spatial
overlaps with humans, and sometimes proliferate, even in urban
environments (3). This is the case for the long-tailed macaque
(Macaca fascicularis) which increasingly gravitates toward
human environments throughout south-east Asia, becoming
sometimes locally overabundant within human-dominated lands
(4). Uncontrolled increase of populations in urban or peri-
urban settings often causes conflicts between macaques and
people, threatens public health, multiplies nuisance problems
for people, and has in turn significant impacts for macaques
(2, 5). Regarding sanitary risk, non-human primates present a
high potential for zoonotic pathogen exchange with humans by
their close phylogenetic relationship (6), especially when they
expand their range in anthropogenic landscape and increase in
abundance (7). The risk of infectious disease outbreaks increases
in human-macaque interface zones with intensified inter-species
contacts, with implications for both public health and primate
conservation (8). The recent COVID-19 pandemic events have
recalled for a pressing need to monitor these populations at the
interface with humans (9). Citation: Deleuze S, Brotcorne F, Polet R,
Soma G, Rigaux G, Giraud G,
Cloutier F, Poncin P, Wandia N and
Huynen M-C (2021) Tubectomy of
Pregnant and Non-pregnant Female
Balinese Macaques (Macaca
Fascicularis) With Post-operative
Monitoring. Front. Vet. Sci. 8:688656. doi: 10.3389/fvets.2021.688656 September 2021 | Volume 8 | Article 688656 Frontiers in Veterinary Science | www.frontiersin.org Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. oviducts were definitely disrupted and no longer patent. Moreover, no new pregnancies
in sterilized females were recorded during the 3-year observation period. The survival
rate of the treated females 6 months after sterilization was high (96.3%) with no
major post-operative complications clinically recorded. Among females that were
pregnant during surgery, 81.1% were confirmed to experience term delivery. This study
demonstrates the safety and efficiency of endoscopic tubectomy, even for pregnant
females, as a mean of wild macaques’ population control. Keywords: bipolar thermocautery, laparoscopy, monkey, population control, salpingectomy, sterilization, wildlife Keywords: bipolar thermocautery, laparoscopy, monkey, population control, salpingectomy, sterilization, wildlife INTRODUCTION Managing these escalating conflicts
has to consider altogether macaque population sustainability,
public health implications, as well as cultural and economic
concerns of neighboring local communities (10). The first objective of this study is to document the
modification of the endoscopic tubectomy procedure used in
the long-tailed macaque, to assess its applicability with gravid
females, and to provide a clinical evaluation of the method
and its potential associated complications. The second objective
is to assess the suitability of this sterilization technique in
a free-ranging setting via a long-term post-op monitoring of
the contracepted females after release as regards to survival
and births. In several places in Asia, measures have been carried on
to limit the expansion of monkey overpopulation associated
with human-macaque conflicts (3). Contraception represents
an ethical alternative to culling and translocation and is
increasingly used in free-ranging populations. Over the years,
various approaches have been used including male castration
(11), progestin-based contraceptive treatment administered to
females (12), programs combining immune-contraception on
females and neutering chemical vasectomy on males (13), or
endoscopic tubectomy as a permanent sterilization method for
females. The latter technique has since been used either alone
(14) or combined with vasectomies (15–17) on a large-scale on
rhesus macaques (Macaca mulatta) in different sites in India and
Hong Kong. Frontiers in Veterinary Science | www.frontiersin.org Keywords: bipolar thermocautery, laparoscopy, monkey, population control, salpingectomy, sterilization, wildlife Study Population and Local Issue Study Population and Local Issue
Despite the limited size of available habitat (∼21 ha in 2018), the
population of macaques in the Ubud Monkey Forest dramatically
grew over 30 years, with a 10% mean annual growth rate (24). In 2017, the population counted 754 individuals split into seven
social groups (C, X, S, N, H, T, L) and kept increasing up
to 1,059 individuals in 2019, reaching the highest population
density (50 individuals/ha) reported for that species in the
semi-wild (25). Reasons for this constant growth and high
density include the combined effects of forest loss and limitation
to emigration imposed by the recent development of human
barriers (26), the absence of natural predators, the cultural
protection the macaques granted in the temple forest site, and
the regular large-scale food provisioning of the monkeys which
otherwise would not find enough resources in the forest to
sustain themselves (27). Therefore, the site is now overcrowded
and this situation imposes an array of detrimental consequences
and costs to the macaques, including growing social tensions,
increased aggressions and fading social group cohesiveness (28). Concurrently, the stakeholders of the sanctuary report increasing
complaints from the neighborhood about macaque nuisances as
well as aggressive interactions between macaques and visitors of
the site. The capture protocol complies with the ethical recommendations
of Jolly et al. (35) for capturing free-ranging habituated primates. The capture-release process was conducted by the trained local
staffof the Ubud Monkey Forest. Four giant trapping cages (5 mL
× 2.4 mW × 2 mH) located within the groups’ home range were
baited daily with food since 2016 to habituate the monkeys to
use the cage (Supplementary Figure 1). The cages were equipped
with a trapdoor that had to be manually maneuvered by an
operator at distance. On capture days, once an acceptable number
of reproductive females had entered the cage, the trap door was
closed and another operator slid a movable partition on the back
side of the cage to lead the monkeys into smaller squeeze cages for
anesthesia (Supplementary Figure 2). Darting using blowpipe
was occasionally used to increase numbers of females and reach a
minimum of 8–10 surgeries per day. The first dose administrated
was based on the visual assessment of the age class. Based on
previously collected data, estimated bodyweights of adult and
subadult females were 5.5 and 3.5 kg, respectively. Study Site
h
h This research was conducted in the Ubud Monkey Forest
(Mandala Wisata Wenara Wana), a forest sanctuary located in
Ubud in south-central Bali (8◦31′S-155◦15′E), Indonesia. It is a
highly frequented tourist site, visited by more than one million of
tourists per year, generating a plethora of commercial activities
and economic incomes (23). The site consists of a Hindu temple
complex within a 21 ha of secondary mixed forest enclosed in
an urban landscape (Figure 1). This sanctuary shelters both a
long-term residential and free-ranging population of long-tailed
macaques. The village committee has managed this site since
1980 and provisions the macaques on a daily basis with sweet
potatoes and various fruits and vegetables. Although wildlife fertility control has gained influence over
the past decades (18), experimental data are critically lacking in
this field when it comes to free-ranging primates. This situation September 2021 | Volume 8 | Article 688656 Frontiers in Veterinary Science | www.frontiersin.org 2 Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. females were sterilized. A demographical monitoring and further
projections are in progress to check the model suitability and
calibrate the needed adjustment campaigns (31). FIGURE 1 | The study site and location of the four trapping cages and the
surgery room (modified from www.monkeyforestubud.com). Frontiers in Veterinary Science | www.frontiersin.org Study Animals Over successive sterilization campaigns between 2017 and 2019,
140 reproductive females underwent an intervention for tubal
cauterization. Only reproductive females (identified by the
presence of nipples development and sexual skin swelling) were
selected for surgery. These reproductive females were either full
adults or subadults, identified on a combination of morphological
characteristics including dental eruption patterns (32) and
development stage of sexual organs (33, 34). Subadults (2.5–
3.5 years) are nulliparous females characterized by buttonlike
nipples and 24–28 permanent teeth. Adults are primiparous
or multiparous females with elongated nipples and 28–32
permanent teeth. Out of 140 females, 129 (92.1%) were adult and
11 (7.9%) were subadults. Characteristics of the treated animals
are documented in Supplementary Table 1. FIGURE 1 | The study site and location of the four trapping cages and the
surgery room (modified from www.monkeyforestubud.com). Study Population and Local Issue Animals were
induced using intramuscular injection of 5 mg/kg of ketamine
(Ketamine 10% R⃝, Kepro, Holland) and 1.33 mg/kg of xylazine
(Xyla R⃝, Interchemie, Holland). Once induced all animals were
weighed to allow adjusted dosage of all drugs administered
thereafter. For animals that underwent surgery, anesthesia
was then maintained with intermittent intravenous boluses of
1% propofol (Recofol R⃝, 1 mg/kg, Primex, Switzerland). Vital
parameters, respiratory and heart rate, pulse oximetry (with the
probe placed either on the cheek, tongue, or tip of the index
finger) and rectal temperature were monitored all along the
procedure. Given that the capture method used a food-baiting
strategy, the animals were not fasted before their anesthesia. After
anesthesia, monkeys were equipped with a pediatric bracelet
marked with an individual code for ID tracking and were
transported to an air-conditioned operation building located
inside the sanctuary (Figure 1). In the aforementioned conditions, the question of population
control has become crucial and a sterilization program has
been initiated in 2017 as a means of controlling the density
of monkeys in an ethical way. This first required the careful
estimation of the magnitude of the sterilization needed. A
stochastic matrix population model (29) based on a 10-year
census dataset of the population was implemented to estimate
the sterilization proportion of reproductive females required
to achieve population stability (i.e., a null population growth). Simulations suggested that 47% of the reproductive females in
2017 (i.e., 135 individuals) had to be sterilized in order to reach
population stability (30). By 2019, 41.9% of the reproductive September 2021 | Volume 8 | Article 688656 3 Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. Marking and Release A permanent double marking for sterilized females was used
with a microchip transponder inserted in the left antebrachium
and a tattoo on their chest to permit remote identification
(Supplementary Figure 4). After
marking,
the
identifying
pediatric
bracelet
was
removed,
a
reversal,
atipamezole
hydrochloride (Antimedin R⃝, 0.02 mg/kg, Dong Bang Co.,
Korea) was administrated intramuscularly (using a volume
consistent with that of the previously administered xylazine)
and animals were positioned in lateral recumbency in individual
holding cages for recovery (Supplementary Figure 5). Animals
were monitored for signs of pain during the recovery period. When they showed clear signs of full recovery from anesthesia
(i.e., strong grip and climbing reflex, visual fixation, and
following reflexes), individuals were transported in their holding
cage to the capture site and released in their group. Post-operative Monitoring p
g
This study has been conducted over 3 years (2017–2020) during
four successive sterilization campaigns (16 days in total: 3
days in July 2017 and July 2018, 5 days in February 2019
and August 2019). In parallel with the campaigns, a long-term
monitoring of the contracepted females was conducted. Those
females were monitored over 27 months split in several field
observation periods between 2017 and 2020 (i.e., July 2017–
September 2018, February 2019–September 2019, and December
2019–Mars 2020). Within 6 months after their surgery, the
general body condition of contracepted females in the field, as
well as any injury on sutures or elsewhere, disabilities, or cues
indicating major unhealthy conditions were assessed. For the
females pregnant at the time of the surgery, information about
the pregnancy process and births was collected. Delivery failure
in contracepted pregnant females was defined as abortion and
perinatal mortality (36), based on clinical observation of (non-
)occurrence of births after the intervention. Finally, survival of
the sterilized females was defined as individuals still present at
least 6 months after surgery. Clinical Exam and Preparation cavity were carefully evaluated for any sign of bleeding or
complication. The telescope and cannulae were removed and the
abdomen was deflated by gentle massage. The two abdominal
incisions were closed using 3–0 polyglecaprone (Monocryl R⃝,
Ethicon, USA) in simple interrupted pattern. An intradermal
suture with the same material was used for the skin incisions
(Supplementary Figure 3). A general clinical exam was conducted to care for potential
injuries, remove food material from the oral cavity and cheek
pouches to reduce the risk of aspiration pneumonia, and assess
the position and size of the uterus that enlarged cranially with
pregnancy for subjective staging via abdominal palpation (early
stage defined as <1.5 months, mid-term as 1.5–3 months, and
advanced stage as >3 months). Females selected for surgery were
catheterized in the cephalic vein for fluidotherapy with saline
perfusion (NaCl 0.9%) and received a dose of antibiotic (Betamox
LA R⃝, amoxicillin, 15 mg/kg, I.M., Norbrook, United Kingdom),
non-steroidal anti-inflammatory (Tolfedine R⃝CS, 4 mg/kg,
subcutaneous, Vétoquinol, France), and eye lubricant (Visine R⃝,
Johnson & Johnson) to protect the cornea from desiccation. The
abdomen was shaved from the xiphoid appendix to the pubis and
as far as the lower line of the flank. Patients were placed in dorsal
recumbency with the pelvis placed on a cushion to slightly elevate
it and allow movement of the abdominal organs cranially in the
abdominal cavity (Trendelenburg position). The surgical site was
scrubbed with antiseptic soap and 70% alcohol and draped. Anesthesia charts including time of drug administration,
duration of surgery from incision to suture, and from induction
to release, as well as clinical remarks were recorded. Surgery The tubectomy procedure routinely consisted in a laparoscopic
two-5 mm-ports approach. Briefly, a 15 mm skin incision
was performed with a scalpel blade ∼1–2 cm cranially to the
umbilicus to allow placement of a Veress needle (150 mm,
CovidienTM, USA), taking care to avoid the underlying organs
(e.g., the gravid uterus) or any blood vessel. The abdomen
was then insufflated with CO2 to a pressure of 10–12 mmHg
(Insufflator, WISAP R⃝, Germany) until a satisfactory abdominal
distension was achieved. The Veress needle was then removed
and a 5 mm trocar was inserted through the same skin incision. The rigid telescope (Hopkins R⃝straight 0◦, 5 mm, 29 cm, Karl
Storz, Germany), connected to an endoscopic camera (Sopro-
Comeg 368 HD, Germany) and light source (LED Nova 150,
Karl Storz, Germany) was introduced in the abdomen via the
cannula. The abdominal cavity was evaluated for topography of
the reproductive organs, gravidity and presence of adhesions. The site for the insertion of a second 5 mm trocar was localized
in the left lower quadrant, just cranially to the oviduct. After a
skin incision of ∼5 mm, the second trocar was inserted under
visual endoscopic control, guided by transillumination thus easily
avoiding large blood vessels. The ampulla of the left oviduct
was easily recognized as flexuous, and paler than the rest of the
organ. Using a bipolar Kelly grasping forceps (RoBi R⃝38151,
Karl Storz, Germany) connected to a thermo-cauterization unit
(Autocon R⃝II 80, Karl Storz, Germany), it was grasped and
coagulated until blanching of the entire oviductal segment was
observed. A second cautery was performed proximally to the first. The ampulla of the right side was accessed through the same
port and cauterized in a similar way (Supplementary Video 1),
except for pregnant females nearer to term for which a
third access in the right flank was performed to reach the
right oviduct (Supplementary Video 2). The sites of thermo-
cauterization were checked for adequate cauterization of the
sections of the oviduct as they were not resected and were left
in place. The reproductive organs and the rest of the abdominal Complications The major complication encountered during the surgery was
caused by the extent of fibrous adhesions, that significantly
interfered with the exploration of the abdominal cavity. These
adhesions were recorded in 24/140 (17.1%) of the treated females
(Supplementary Table 1) and were mostly located in the caudal
abdomen. A biopsy, obtained from a lesion appearing as a small
mass located at the surface of the uterus of the patient (X=) with
such adhesions and for which the laparoscopy was converted
to a laparotomy, confirmed a diagnosis of endometriosis. Five
cases of herniation were recorded, usually associated with
dense adhesions. For exhaustive data, see Supplementary Table 1. Intraoperative Evaluation
Success of Tubectomy and Pregnancy Stage-Related
Feasibility proportion (%) of age class (adult, subadult), pregnancy stage at the time of the
intervention, pregnancy outcome, female survival at least 6 months following
surgery, and intraoperative complications. Parameter
Category
Occurrence
%
Age
Adult
129
92.1
Subadult
11
7.9
Pregnancy stage
Not
gravid
100
71.4
Early
11
7.9
Mid-term
10
7.1
Advanced
19
13.6
Pregnancy outcome
Delivery
30
83.3
Abortion
6
16.7
Unknown
4
10
Survival (6 month)
Yes
131
96.3
No
5
3.7
Intraoperative complication
Tubectomy
failure
4
2.9
Adhesions
24
17.1
For exhaustive data, see Supplementary Table 1. y
Among the 140 treated females, 136 (97.1%) were successfully
tubectomized. Forty of these females (28.6%) were at various
stages of pregnancy (early stage: 7.9%, mid-term: 7.1%, advanced
stage: 13.6%). Among these 40 pregnant females, 3 (7.5%)
could not be successfully sterilized due to various intraoperative
complications. All three were at an advanced stage of pregnancy
at the time of surgery. In two cases (C6, H8), the size of the uterus
was such that the oviducts could not be adequately visualized and
accessed, and the procedure was interrupted to avoid prolonged
manipulations and anesthesia. The third case (S14) was a female
at an advanced stage of gestation that died during the surgery
for undetermined cause. In a fourth case, that was tattooed
with a mark “X=,” a non-pregnant female presented abundant
adhesions in the uterine zone, making it impossible to access the
ovaries despite a conversion to an open laparotomy approach. The mean surgery duration was 14 ± 3 min for non-pregnant
females and 22 ± 18 min for pregnant females. The overall whole
capture-release process lasted in average 3 h 22 min, and ranged
from 1 h 11 min to 5 h 52 min. Post-operative Clinical Evaluation The
potential
post-operative
complications
by
direct
observations of the released females within days after the
surgery were monitored (Supplementary Figure 4). No issues
apart from dehiscence of the skin suture, due to intensive
scratching by the monkey, seen in 8 cases (5.7%) without major
infection cues or systemic impact, no major complication and no
pain-associated behavior associated with surgery in non-human
primates (37, 38) have been observed after release. To monitor the efficacy of the laparoscopic tubectomy
approach, a laparoscopic post-operative assessment was made
on two patients (L3, T4) randomly re-captured 1 or 2 years
after their intervention. Capture, anesthesia, analgesia and
laparoscopy procedures were the same as previously described. Both laparoscopic post-operative evaluations confirmed that the
oviducts were definitely disrupted and no longer patent. The
ovaries looked normal and were clearly no longer connected
to the uterus as part of the oviduct was missing. There was
no sign of local reaction or adhesions on the surgical site
(Supplementary Video 3). Long-Term Field Monitoring The 3-year monitoring has revealed a 100% reliability rate of the
sterilization method since no new birth has been recorded so
far in females which underwent surgery, and were not pregnant
at that time. 96.3% of the sterilized females survived at least 6
months after the surgery and 91.9% until 3 years after surgery. This survival rate is similar to the annual survival rate of all
reproductive females of this population (92.3%) calculated via a
matrix population model based on 10-year life table dataset (30). Most bodies of deceased sterilized females could not be found
but their behavior and condition before their disappearance
suggested causes unrelated to surgery (e.g., falls from trees,
human-related hazards such as electric shock in poles). Finally,
the pregnancy outcome for 36 of the 40 gestating females that
underwent surgery was confirmed: delivery failure (including
abortion and perinatal mortality) was observed in 6 (16.7%) of
these females. Relevant results are summarized in Table 1. Intraoperative Evaluation
Success of Tubectomy and Pregnancy Stage-Related TABLE 1 | Data summary of the treated females (N = 140) with frequency and
proportion (%) of age class (adult, subadult), pregnancy stage at the time of the
intervention, pregnancy outcome, female survival at least 6 months following
surgery, and intraoperative complications. Frontiers in Veterinary Science | www.frontiersin.org RESULTS Over the four sterilization campaigns, 140 long-tailed female
macaques (129 adult and 11 subadult females) have been treated
and monitored (Supplementary Table 1). The average weight of
these females was 5.2 kg (N = 140). Adult females weighed more
(5.3 kg, N = 129) than subadults (4.0 kg, N = 11). Over the four sterilization campaigns, 140 long-tailed female
macaques (129 adult and 11 subadult females) have been treated
and monitored (Supplementary Table 1). The average weight of
these females was 5.2 kg (N = 140). Adult females weighed more
(5.3 kg, N = 129) than subadults (4.0 kg, N = 11). September 2021 | Volume 8 | Article 688656 Frontiers in Veterinary Science | www.frontiersin.org 4 Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. DISCUSSION Contraception is increasingly used as a tool for population
control of primates living in anthropogenic habitats in Asia. Despite an increasing interest in using this method, their
implications and suitability remain poorly understood (39). Very few systematic studies have indeed evaluated the outputs
of such measures, which constitutes a major research gap
calling for robust assessments (40). This study aimed to September 2021 | Volume 8 | Article 688656 Frontiers in Veterinary Science | www.frontiersin.org 5 Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. provide an evaluation of the laparoscopic tubectomy by bipolar
thermocautery as a mean of fertility control in free-ranging long-
tailed macaques. It also aimed at adjusting the procedure for
pregnant patients and provide a clinical evaluation of the method
and its potential associated complications. supports the unlikelihood of recanalization and reversal with
this technique in long-tailed macaques as supported by the
100% efficacy of the method (absence of recorded pregnancy
or birth) for all the patients that survived over the 3-year-
period monitoring we report. These results confirm that a bipolar
thermocautery alone, without further resection of the fallopian
tubes is a safe and simpler approach to achieve an effective
hemostasis and permanent disruption of the oviducts. Previous studies have shown various undesirable side effects
associated with castration, ovariectomy or contraceptive agents
in primates (41–45). Conversely, while impeding fecundation,
vasectomy and tubectomy keep intact the steroid hormonal
functions
underpinning
normal
sexual
activity
(20,
46). Preservation of the gonadal hormones represents the best option
when sterilizing wild female macaques as it allows them to keep
normal sexual and social behaviors in their group. It has also
been shown to have beneficial effects on bone metabolism both
in women (47) and monkeys (48). Laparoscopic tubectomy
is a minimally-invasive intervention highly recommended
as a substitute for conventional ovariectomy for permanent
contraception of macaques as it does not induce observable
hormonal alterations (49). It is associated with a lower incidence
of intra- and post-operative complications (17) and significantly
reduces the surgical and post-operative recovery times in
primates (50). This fact is of paramount importance with social
primates since prolonged separation of individuals is likely
to jeopardize their reintroduction to social group. Common
adverse consequences of prolonged isolation are fighting,
increased aggressions toward the reintroduced animals and
socio-behavioral changes including loss of the social status
after
reintroduction
(51). The
maximum
holding
period
recommended is 24 h (52). DISCUSSION In this study, all animals were
reintroduced to their group within 6 h after capture, as reported
on large cohorts (14), and the release process monitoring showed
no substantial fights during release. Previous reports of laparoscopic tubectomy in macaques
accessed each ovary from its own side (17, 20, 50) resulting in a
three-port technique. More recently, both ovaries were accessed
from the same side but maintained a three-port approach (14). Although, the original plan had been to access each ovary from
its own side, it became immediately obvious that the first port
in the left flank could easily be used to access the right oviduct
as well with the bipolar forceps, at least for non-pregnant and
early to mid-term pregnant females. The two-port approach was
therefore preferred as easier to perform and avoiding the opening
and closing of the third access, thus reducing the duration of
the procedure. p
Capture is a keypoint of free-ranging animal sterilization
programs. Although less stressful for the animals, and giving
access to higher number of them (35), trapping in cages does
not allow selection of the captured animals. Individuals that may
not be contracepted have to be released with reduced chances
to be trapped again in a short interval. These include females
with palpable pregnancies which were not usually considered
suitable candidates for surgery (14). In order to avoid having
to release animals without neutering them, the technique was
adjusted to allow surgery on pregnant females. The placement
of the first trocar was adjusted based on gestational age and
the relative position of the gravid uterus evaluated by palpation
as commonly described in humans (55). While the two-port
approach proved to grant easy access to both oviducts in early
to mid-term pregnancies, in females nearer to their term the
size of the gravid uterus precluded the adequate visualization
of the contralateral oviduct. For those cases, following cautery
of the left oviduct, patients were slightly tilted to the left and a
third access in the right flank is performed as described above. This position is routinely recommended in human laparoscopy
(55, 56) and it has been shown that it is unnecessary to place the
patient in a strict lateral decubitus position (57). Not only does it
allow better visualization of the right oviduct but it also reduces
aortocaval compression. Frontiers in Veterinary Science | www.frontiersin.org September 2021 | Volume 8 | Article 688656 DISCUSSION In humans, one of the major concerns
with laparoscopy on pregnant patients was originally the effect of
the CO2 pneumoperitoneum on the fetal physiology. However,
it has been shown that no substantial adverse effects are observed
when the duration of the procedure is limited (under 60 min) and
the intraabdominal pressure is kept under 10–12 mmHg (58, 59). While the insufflator is set at 12 mmHg as default in this approach
for non-pregnant monkeys, the intraabdominal pressure was
limited as possible to 10 mmHg in advanced pregnancies as
long as a sufficient visualization of the oviducts can be achieved. Accidental puncture of the gravid uterus with the Veress needle
is considered the most serious complication in pregnant women
(58, 60). This complication has not been observed in any of the 40
pregnant females that underwent surgery during the campaigns. The technique we describe here is adapted from the
endoscopic salpingectomy in wild macaques first demonstrated
by P. Martelli in a DVD entitled “Endoscopic Tubectomy in
Macaques” for a German company selling endoscopic equipment
(Karl Storz-Endoskope) in 2009. Endoscopic salpingectomy has
since widely been used for neutering campaigns in Formosan
(20), rhesus (14, 17) macaques and quite recently, on a
chimpanzee (Pan troglodytes) (53). Here the choice of cautery, a two-port-approach, the inclusion
of pregnant females and the complications as well as the
modifications to the original technique are discussed. While all previous reports combined coagulation of the
oviducts followed by resection using endoscopic scissors of a
section of each tube, that was removed from the abdominal
cavity through the cannula (17, 20, 50, 53), this study opted for
thermocautery only. The oviduct is grasped and cauterized twice
but no section is resected and removed. The severed tissues are
left to aseptically necrotize in situ, which was considered safe
and sufficient to ensure sterilization of the animal as it has been
shown to be as safe and effective as partial salpingectomy in
women (54). Although that information is limited to two patients
that underwent a second laparoscopy for control purposes, the
anatomical disruption of the portion of the oviduct that had been
severed by thermocautery and the consequent loss of connection
between the ovary and the uterus was confirmed visually. This September 2021 | Volume 8 | Article 688656 6 Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. DISCUSSION Laparoscopic tubal cautery during pregnancy was also safe
and feasible. However, the procedure feasibility was limited by
the gestational stage: females near term (≥4 months) could not
easily be treated because the size of the uterus reduced the access
to oviducts. Provided that this limitation is considered, this new
insight is of great interest for sterilization of animals living in
free-ranging settings where captures are very challenging. That
is particularly true for non-seasonal breeding species, where
no optimal timing of capture can be identified because of
year-round distribution of births. Captures can therefore be
optimized if most of the gestating females can be included and
undergo tubectomy. Post-operatively, 8 cases of partial wound dehiscence were
observed which did not impact the general condition of the
animals and was self-limited. These delayed wound healing
came unsurprisingly as grooming behavior, wound, or stitching
material picking are common in macaques (66). However, this
complication has not been mentioned in other studies (17) and
more particularly in a large scale study (14) where the skin
was closed using surgical glue rather than absorbable suturing
material, which should be considered to further improve our
technique. No other complications were recorded. Systematic
post-operative observations are not feasible with animals living
in a semi-wild habitat, and although no pain scoring observations
were performed, females that were seen in the days following
surgery showed no apparent signs of pain and displayed normal
behavior of feeding and grooming. These observations support
the fact that, laparoscopic tubectomy being a minor surgical
procedure, one single administration of NSAIDs for analgesia
was sufficient to allow swift release of the animals back to their
social groups. Although this duration may need to be reduced for smaller
animals (<2 kg) because of their propensity to hypoglycemia,
it is usually considered advisable to withhold food for 12–16 h
and water for 2 h before induction of anesthesia of non-human
primates (67). This reduces the risk of gastrointestinal bloat
and aspiration pneumonia as macaques may vomit on induction
of anesthesia. Fasting the animals—or even administration
of metoclopramide 30–60 min before induction which is also
sometimes recommended (67)—was unfortunately not possible
under the conditions of this study. However, like others (14),
no such complications were observed during our sterilization
campaigns and thorough emptying of the cheek pouches was
considered sufficient to prevent aspiration pneumonia. DISCUSSION involved fibrous adhesions, especially between the omentum
and the uterus, liver, spleen, and urinary bladder (64). The same
paper also describes cases where one or both ovaries could not be
identified and were suspected to be incorporated or effaced by the
fibrous adhesions. Although, it would be hazardous to conclude
that all cases with intra-abdominal adhesions would necessarily
be due to endometriosis, this is exactly what was observed in
the patient [“(X=)” where endometriosis was confirmed. In
non-human primates, decidualization of endometriotic tissue
most often occurs under the influence of exogenous progestins
or during pregnancy (65). In this study, none of the 24 animals
with such lesions received exogenous progestins and 16 were not
pregnant, supporting the hypothesis of that endometriotic tissue
may decidualize under the influence of elevated endogenous
progesterone (64). It may be suspected that the size of the patients, allowing a
good transabdominal palpation and localization of the uterus,
permitted readjustment of the site of insertion more cranially to
avoid the uterine fundus. It is worth noticing that while all the
previous reports based on the initial description of the technique
place the first trocar caudal to the umbilicus (14, 17, 20, 50, 53),
the first access is located just cranially to the umbilicus in this
study. This may contribute to an easier access to the contralateral
oviduct even in pregnant patients and limit conflicts between
instruments inside the abdominal cavity. y
As laparoscopic oviduct cautery does not induce observable
hormonal alterations (49), it is therefore not expected to interfere
with pregnancy maintenance. By comparison, delivery failure
was observed in 16.7% females. This proportion of abortion in
gravid sterilized females was similar to or even lower than the
one reported for long-tailed macaques in captivity (23.7%) (61). Out of a very large cohort, where pregnant females were normally
excluded, 9 cases of tubectomy on females suspected of being
pregnant based on the presence of mildly rounded congested and
enlarged uteri, among which 3 completed a gestation successfully
have been reported. However, as these findings were incidental
and pregnancy had not been confirmed, the authors preferred
not to draw conclusions (14). These results establish that the
laparoscopic tubectomy did not interfere with gestation and
is therefore a safe and reliable sterilization technique even for
advanced pregnant female long-tailed macaques. Laparoscopic
procedures for various surgical interventions during pregnancy
have been considered effective and safe in women and macaques
(62). Frontiers in Veterinary Science | www.frontiersin.org DISCUSSION Complications
of
endoscopic
salpingectomy,
including
difficult entry into the abdominal cavity using the Veress needle,
intraoperative hemorrhage and inadvertent cauterization of the
uterus have been reported in Formosan macaques (20). As in
other reports (14), very few intraoperative complications were
observed during this study’s campaigns. The main intraoperative
limitation was due to the presence of adhesions, observed in
24/140 (17%) of the operated females out of which 16 were not
pregnant. In one case, these adhesions were such that access
to the ovaries could not be achieved even after conversion
to a standard laparotomy. A biopsy obtained from this
patient confirmed the presumptive diagnosis of endometriosis
associated with these cases of adhesions mainly located in the
caudal abdomen. The incidence of spontaneous endometriosis
in a breeding colony of Macaca fascicularis has been reported
to be 28.7% (63) in aging females. Lesions of endometriosis
have recently been characterized in long-tailed macaques and In this study, the mean surgery duration of the laparoscopic
tubectomy was 14 ± 3 min for non-pregnant females and 22 ±
18 min for pregnant females. This is longer than the 3–10 min
reported in other papers (14, 17) that included non-pregnant
females only. Possible explanations for this difference may be
found in the degree of experience of the operator but also in
at least two technical aspects. First, females with pre-existing
severe adhesions that sometimes seriously interfered with the
access to the oviducts were included and thus prolonged the
duration of the procedure. Second, 5 mm trocars instead of 3 mm September 2021 | Volume 8 | Article 688656 Frontiers in Veterinary Science | www.frontiersin.org 7 Endoscopic Tubectomy in Long-Tailed Macaques Deleuze et al. of NSAID for analgesia and a holding period as short as
possible). This research has been approved by the Animal Ethics
Commission of Udayana University (No. 282/KE-PH/I/2017)
and has been granted with permission of the Provincial
Office of Conservation in Bali (BKSDA, No. KT49/BKSDA
BL-1/KK/2017;
Capture
license:
#29/PPSP/XII/2017). Moreover,
this
research
was
conducted
under
research
permission from the Indonesian Ministry of Research and
Technology
(No. 186/SIP/FRP/E5/Dit.KI/VII/2017,
No. 46/SIP/FRP/E5/Dit.KI/II/ 2018, No. 10/E5/E5.4/ SIP.EXT/2019,
No. 83/E5/E5.4/SIP.EXT/2019). Universal precautions (gloves,
masks, and overshoes/caps specifically for surgery) against
hazards and zoonotic pathogen transmission were carefully
taken during the whole process from capture, clinical exam,
surgery and postoperative procedures. ones were used, which led to suture the abdominal wall rather
than leaving it unsutured. ACKNOWLEDGMENTS We thank the Indonesian Ministry of Research and Technology,
the Ubud Monkey Forest Management Committee, Mr. Nyoman
Buana and Mr. Wayan Buda for support and permission to
conduct this research in Indonesia. We are very grateful to the
Ubud Monkey Forest staff, the field assistants (from ULiege,
UCL, and UNUD) and the senior colleagues from Udayana
University for their assistance with the sterilization campaigns
and data collection. We also thank Dr. Nicolas Schtickzelle for
his help in matrix population modelling, Dr. Paolo Martelli and
Mrs. Femke den Haas for fruitful advices and logistic support
regarding the sterilization campaigns. ETHICS STATEMENT The
whole
protocol
of
this
study
followed
the
ethical
recommendations for animal welfare (36). All procedures
have been chosen in order to cause the least stress, pain
and discomfort to the monkeys (e.g., collective trapping
allowing to achieve a maximal number of captures while
causing the least stress and discomfort to the monkeys,
limiting handling when animals are not sedated, minimally
invasive
endoscopic
tubectomy
procedure,
administration DATA AVAILABILITY STATEMENT The original contributions presented in the study are included
in the article/Supplementary Material, further inquiries can be
directed to the corresponding author/s. AUTHOR CONTRIBUTIONS FB, SD, and M-CH: conceived and designed the study. SD, RP, and GS: undertook the laparoscopic tubectomy
and collected clinical data. SD, GR, M-CH, FB, NW, GS,
GG,
and
FC:
veterinary
care
and
logistics
during
the
sterilization campaigns. GG, FB, and FC: data collection
for
field
monitoring. FB:
data
analyses. NW
and
PP:
facilitation to fieldwork. SD and FB: wrote the manuscript. M-CH:
provided
in-depth
editorial
advice. All
authors
reviewed
and
edited
the
manuscript
and
approved
the
submitted version. Our follow-up over a 3-year period on patients’ clinical
evolution, pregnancy outcome, and later infertility demonstrates
the safety and efficiency of the modifications brought to the
procedure. Moreover, as controlling effects of fertility control
on wild animal life is of paramount importance, a long-term
monitoring of this population is still ongoing and further
focuses on the potential impact of reproduction suppression
on behavior and social dynamics (68). Effects of reproduction
suppression on the health, welfare, social behavior, and ranging
patterns of primates are often only superficially considered. This influence needs to be examined to further complete the
data on the technique and its impacts beyond the population
control aspects. FUNDING This study was carried out with the financial support of the
Padangtegal village leadership committee, the Belgian National
Fund for Scientific Research, the Erasmus + International
Credit Mobility Programme, the University of Liège, and
Wallonie-Bruxelles International. This study was carried out with the financial support of the
Padangtegal village leadership committee, the Belgian National
Fund for Scientific Research, the Erasmus + International
Credit Mobility Programme, the University of Liège, and
Wallonie-Bruxelles International. DISCUSSION Finally, to limit wound picking, the
skin was apposed with an intradermal suture instead of surgical
glue which is obviously faster. Laparoscopic vessel sealant tools,
not yet available in 3 mm, were shown not necessary and that
standard bipolar forceps, that are available both in 3- and 5-
millimeter size, were sufficient to efficiently sterilize the patients. Smaller size instruments should therefore probably be preferred
for surgeries on animals the size of a macaque. As a conclusion, laparoscopic oviduct cautery with standard
bipolar forceps can be adjusted to safely sterilize barren and
pregnant long-tailed macaques even at late stages of pregnancy. Use of a single lateral port is feasible in non-gestating and
early to mid-term pregnant females, while females nearer to
their term are best operated using a three-port entry. The
idea that including pregnant females and patients with some
degree of adhesions will optimize the trapping efforts and the
overall success of population control programs. The absence
of significant complications with the technique, which allows
release of free-ranging animals very shortly after procedure and
avoids complications related to prolonged isolation from the
group. The technique could be further optimized by combining
a 3 mm bipolar forceps via a two-port approach and the use of
surgical glue for the closure of the skin, as minuscule accesses
allow suture of the linea alba only thus making the entire
procedure shorter again. REFERENCES Primates and the ecology
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Wandia and Huynen. This is an open-access article distributed under the terms
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copyright owner(s) are credited and that the original publication in this journal
is cited, in accordance with accepted academic practice. No use, distribution or
reproduction is permitted which does not comply with these terms. 56. Guterman S, Mandelbrot L, Keita H, Bretagnol F, Calabrese D, Msika S. Laparoscopy in the second and third trimesters of pregnancy for abdominal
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Gene Co-expression Network Reveals Potential New Genes Related to Sugarcane Bagasse Degradation in Trichoderma reesei RUT-30
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ORIGINAL RESEARCH
published: 22 October 2018
doi: 10.3389/fbioe.2018.00151 Gustavo Pagotto Borin 1,2, Marcelo Falsarella Carazzolle 3,
Renato Augusto Corrêa dos Santos 4, Diego Mauricio Riaño-Pachón 5 and
Juliana Velasco de Castro Oliveira 1,2* 1 Laboratório Nacional de Ciência e Tecnologia do Bioetanol (CTBE), Centro Nacional de Pesquisa em Energia e Materiais
(CNPEM), Campinas, Brazil, 2 Programa de Pós-Graduação em Genética e Biologia Molecular, Instituto de Biologia,
Universidade de Campinas (UNICAMP), Campinas, Brazil, 3 Laboratório de Genômica e Expressão (LGE), Departamento de
Genética, Evolução, Microbiologia e Imunologia, Instituto de Biologia, Universidade Estadual de Campinas (UNICAMP),
Campinas, Brazil, 4 Faculdade de Ciências Farmacêuticas de Ribeirão Preto, Universidade de São Paulo (USP), Ribeirão
Preto, Brazil, 5 Centro de Energia Nuclear na Agricultura, Universidade de São Paulo (USP), Piracicaba, Brazil Edited by:
Roberto Silva,
Universidade de São Paulo, Brazil The biomass-degrading fungus Trichoderma reesei has been considered a model for
cellulose degradation, and it is the primary source of the industrial enzymatic cocktails
used in second-generation (2G) ethanol production. However, although various studies
and advances have been conducted to understand the cellulolytic system and the
transcriptional regulation of T. reesei, the whole set of genes related to lignocellulose
degradation has not been completely elucidated. In this study, we inferred a weighted
gene co-expression network analysis based on the transcriptome dataset of the T. reesei RUT-C30 strain aiming to identify new target genes involved in sugarcane
bagasse breakdown. In total, ∼70% of all the differentially expressed genes were
found in 28 highly connected gene modules. Several cellulases, sugar transporters,
and hypothetical proteins coding genes upregulated in bagasse were grouped into
the same modules. Among them, a single module contained the most representative
core of cellulolytic enzymes (cellobiohydrolase, endoglucanase, β-glucosidase, and lytic
polysaccharide monooxygenase). In addition, functional analysis using Gene Ontology
(GO) revealed various classes of hydrolytic activity, cellulase activity, carbohydrate binding
and cation:sugar symporter activity enriched in these modules. Several modules also
showed GO enrichment for transcription factor activity, indicating the presence of
transcriptional regulators along with the genes involved in cellulose breakdown and sugar
transport as well as other genes encoding proteins with unknown functions. Highly
connected genes (hubs) were also identified within each module, such as predicted
transcription factors and genes encoding hypothetical proteins. In addition, various hubs
contained at least one DNA binding site for the master activator Xyr1 according to our
in silico analysis. The prediction of Xyr1 binding sites and the co-expression with genes
encoding carbohydrate active enzymes and sugar transporters suggest a putative role Reviewed by:
Kentaro Inokuma,
Kobe University, Japan
Andrei Steindorff,
Joint Genome Institute (JGI),
United States *Correspondence:
Juliana Velasco de Castro Oliveira
juliana.velasco@ctbe.cnpem.br Specialty section:
This article was submitted to
Bioenergy and Biofuels,
a section of the journal
Frontiers in Bioengineering and
Biotechnology Specialty section:
This article was submitted to
Bioenergy and Biofuels,
a section of the journal
Frontiers in Bioengineering and
Biotechnology Received: 10 July 2018
Accepted: 03 October 2018
Published: 22 October 2018 Gene Co-expression Network
Reveals Potential New Genes Related
to Sugarcane Bagasse Degradation
in Trichoderma reesei RUT-30 Gustavo Pagotto Borin 1,2, Marcelo Falsarella Carazzolle 3,
Renato Augusto Corrêa dos Santos 4, Diego Mauricio Riaño-Pachón 5 and
Juliana Velasco de Castro Oliveira 1,2* INTRODUCTION These genes represent interesting new targets, because several
were found either activated or repressed in culture media having
complex carbon sources (Häkkinen et al., 2012; Borin et al.,
2017; Daly et al., 2017; Horta et al., 2018). In this context,
the gene co-expression network analysis has become a valuable
bioinformatic toolkit for data integration and the identification
of new candidate genes related to a biological process of
interest (Gonzalez-Valbuena and Treviño, 2017). Only a few
studies have inferred gene networks from the transcriptome
of T. reesei. T. reesei RUT-C30 is a well-known industrial
strain able to abundantly produce and secrete cellulases, and
it has been used as a genetic background to develop other
industrial strains. RUT-C30 was isolated following three rounds
of mutagenesis and screening from the ancestral QM6a strain,
and its hypercellulolytic phenotype is attributed in part to the
truncation of cre1, the main player of carbon catabolic repression. Since then, RUT-C30 strain has been the target of numerous
studies on the conversion of biomass into biofuels and other
high-value products (Marx et al., 2013; Mello-De-Sousa et al.,
2014; Druzhinina and Kubicek, 2017). 2G ethanol technology primarily consists of three major steps:
pretreatment, enzymatic hydrolysis and sugar fermentation,
being
pretreatment
and
enzymatic
hydrolysis
the
major
limitations for the economic feasibility of 2G ethanol. In this
context, a cost-effective process is required to make this biofuel
more attractive and competitive (Bornscheuer et al., 2014; Gupta
and Verma, 2015). Trichoderma reesei is the primary fungal source of the
industrial
cellulases
present
in
enzymatic
cocktails
for
lignocellulose degradation and the production of 2G ethanol. Due to its remarkable capacity to produce and secrete enzymes
that are active on carbohydrates, T. reesei has been used as a
microbial factory for the breakdown of lignocellulose and a
host for heterologous expression (Schmoll et al., 2016). These
enzymes and other accessory proteins are collectively named
as Carbohydrate-Active enZymes (CAZymes) (Lombard et al.,
2014) and their genes are under control of several transcriptional
regulators, which are activated or repressed according to sugar
(xyr1 and cre1, for example) and nitrogen (areA) availability, pH
alteration (pacC), light (env1), and other factors (Stricker et al.,
2006; Rassinger et al., 2018; Schmoll, 2018a). In this context,
some of them have been target of studies of genetic manipulation
involving
deletion
and
overexpression
of
transcriptional
regulators in order to enhance the cellulolytic phenotype of T. INTRODUCTION In the last few years, several advances have been achieved
in
different
research
fields
and
contributed
to
a
better
understanding of the physiology and key features behind the
T. reesei hypercellulolytic capacity. Such advances include the
development of new tools for genetic manipulation (Derntl et al.,
2015; Liu et al., 2015), transcriptomic and proteomic studies
using lignocellulosic residues (Dos Santos Castro et al., 2014;
Borin et al., 2015, 2017; Daly et al., 2017; Ellilä et al., 2017;
Cologna et al., 2018), the discovery of new transcription factors
(TFs) and regulatory elements (Derntl et al., 2016, 2017; Stappler
et al., 2017; Zheng et al., 2017b; Benocci et al., 2018), promoter
characterization (Zheng et al., 2017a; Kiesenhofer et al., 2018)
and structural studies of cellulases (Li et al., 2015; Bodenheimer
and Meilleur, 2016; Eibinger et al., 2016; Ma et al., 2017; Borisova
et al., 2018). Over the previous years, the development of new sustainable
alternatives to fossil fuels has become critical to mitigate
greenhouse gas emissions and to avoid the exhaustion of natural
sources. Among the biofuels, second generation (2G) ethanol
has emerged as one of the most promising substitutes for
gasoline since it can be produced from lignocellulosic feedstocks,
including agroindustrial residues and municipal waste (Lynd
et al., 2017). The United States and Brazil are the world’s largest 1G ethanol
producers and are responsible for 57 and 27% of its global
production, respectively (Renewable Fuels Association, 2017). In
Brazil, the 1G ethanol production system is well established and
is based on sugarcane milling and sugar-rich juice fermentation
with generation of bagasse and straw as byproducts. Currently
bagasse is used to produce steam and energy in the sugar-ethanol
biorefineries, and the straw is left on the soil to prevent erosion
and enhance the organic carbon content (da Rosa, 2013; Pereira
et al., 2015; Lisboa et al., 2017). These residues have already
been used as feedstocks for 2G ethanol in Brazilian industrial
plants (GranBio and Raízen), and they have a high potential to
be explored more thoroughly in an integrated process with 1G
ethanol technology (Junqueira et al., 2017; Lynd et al., 2017). However, despite all of the studies conducted and the
knowledge acquired, the biotechnological potential of T. reesei
has not been completely explored since various genes identified
by different “omic” approaches have still not been characterized. Citation: Borin GP, Carazzolle MF,
dos Santos RAC, Riaño-Pachón DM
and Oliveira JVdC (2018) Gene
Co-expression Network Reveals
Potential New Genes Related to
Sugarcane Bagasse Degradation in
Trichoderma reesei RUT-30. Front. Bioeng. Biotechnol. 6:151. doi: 10.3389/fbioe.2018.00151 October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org Borin et al. Gene Co-expression Network of Trichoderma reesei of these hubs in bagasse cell wall deconstruction. Our results demonstrate a vast range
of new promising targets that merit additional studies to improve the cellulolytic potential
of T. reesei strains and to decrease the production costs of 2G ethanol. Keywords: Trichoderma reesei, sugarcane bagasse, 2G ethanol, enzymatic cocktail, gene co-expression network,
Xyr1-binding site Fungal Strain and Culture Conditions Fungal Strain and Culture Conditions
Gene co-expression network analysis was performed based on
a RNA-Seq dataset from a previous study conducted by our
research group (Borin et al., 2017). Briefly, T. reesei RUT-
C30 spores were first cultivated in potato dextrose agar for
7–10 days at 29◦C, and harvested in sterile distilled water. The spore suspensions were inoculated to a final concentration
of 1 × 106 spores per 30 mL of basic culture medium
(BCM) (pH 5.5) composed of 0.05% yeast extract (w/v), 50
mL/L salt solution (6 g/L NaNO3, 1.5 g/L KH2PO4, 0.5 g/L
KCl, and 0.5 g/L MgSO4), 200 µL/L trace elements (10 g/L
ethylenediaminetetraacetic acid, 4.4 g/L ZnSO4·7H2O, 1.0 g/L
MnCl2·4H2O, 0.32 g/L CoCl2·6H2O, 0.315 g/L CuSO4·5H2O,
0.22 g/L (NH4)6Mo7O24·4H2O), 1.47 g/L CaCl2·2H2O, and 1 g/L
FeSO4·7H2O), with 1% fructose (w/v) as the carbon source at
29◦C, 200 rpm for 48 h. The pre-grown mycelia were transferred
to fresh BCM (without yeast extract) with 0.5% bagasse (w/v) as
the carbon source for 6, 12, and 24 h, and to 1% fructose (w/v)
for 24 h. Fructose was used as control condition in the RNA-
Seq experiment as it is an inert sugar, which neither induces
nor suppresses overall expression of lignocellulolytic enzymes
(Amore et al., 2013). The cultures were grown under continuous
light exposure, as it influences positively cellulase gene expression
(Schmoll, 2018b). Mycelia were harvested by filtration, washed
with sterile water and immediately ground into powder in liquid
nitrogen. Frozen material was then used for the RNA extraction. The MCODE (v1.4.2) plugin (Bader and Hogue, 2003) of
Cytoscape was used to identify subclusters of genes densely
connected within each module with the parameters set to
default (degree cutoff= 2, node score cutoff= 0.2, K-core
= 2, maximum depth = 100, haircut method). Genes with
high connectivity within the modules, i.e., nodes with degree
values higher than 90% of the entire degree distribution, were
considered to be hubs (Liang et al., 2012; Bi et al., 2015). The individual betweenness centrality of the nodes generated
by Cytoscape was also compared to the corresponding degree
value in order to confirm the centrality of the hubs. Hub
genes tend to demonstrate a positive correlation between these
two parameters (Potapov et al., 2005; Lee, 2006; Li et al.,
2014). INTRODUCTION reesei (Meng et al., 2018; Rassinger et al., 2018; Zhang et al.,
2018). Previously, Borin et al. (2017) investigated the transcriptome
of T. reesei RUT-C30 grown on steam-exploded sugarcane
bagasse (referred to as “bagasse” from this point on) in a
time course of 6, 12, and 24 h as well as on fructose after
24 h. Interestingly, a set of cellulase, hemicellulase, TF, and
sugar transporter coding genes were activated in bagasse along
with genes encoding hypothetical or uncharacterized proteins. Based on the assumption that co-expressed genes tend to share
similar expression patterns and that they could be co-regulated
by the same elements, such as the carbon source and pH
(van Dam et al., 2017), this study attempts to identify new Frontiers in Bioengineering and Biotechnology | www.frontiersin.org October 2018 | Volume 6 | Article 151 2 Gene Co-expression Network of Trichoderma reesei Borin et al. Portal (9,852 gene models) (Le Crom et al., 2009; Nordberg
et al., 2014) using TopHat2 (Kim et al., 2013). The mapped reads
were counted with the featureCounts function from the Rsubread
v1.12.6 package (Liao et al., 2014). Low abundance genes were
filtered out, keeping only genes with a cpm ≥1 in at least
three samples. RPKM values were calculated following TMM
normalization using the edgeR package v3.12.1 (Bioconductor)
(Robinson et al., 2010) within the R environment (R Core
Team, 2015). Only differentially expressed genes (DEGs) with
two-fold change cutoff(SEB 6, 12 or 24 h vs. fructose 24 h)
were considered, i.e., log2-fold change ≥1 (upregulated) or ≤-
1 (downregulated). The identification of the genes from T. reesei
was based on RUT-C30 strain retrieved from JGI database. genes related to the T. reesei lignocellulose degradation response
using a Weighted Correlation Network Analysis (WGCNA)
(Langfelder and Horvath, 2008). WGCNA estimates the co-
expression similarity between the genes and constructs a
weighted correlation matrix following a scale-free topology. In
scale-free networks, a few nodes (genes) have a high degree
(links), while most nodes have a small number of interactions
(edges). These highly connected genes (hubs) play a central
role in the network stability against perturbations, and they are
very important in diverse cellular processes (Han et al., 2004;
Luscombe et al., 2004). INTRODUCTION In addition, an in silico prediction of the DNA binding
sites for Xyr1, the master activator of cellulases (Stricker et al.,
2006), was performed using the promoters to find genes of
unknown function that could be regulated by this TF. To our
knowledge, this is the first report of the use of a network approach
combined with regulatory motif analyses to reveal new genes
of biotechnological interest in T. reesei RUT-C30. In this study,
an extensive number of genes were found to be co-expressed in
bagasse, and they could be the target of new studies to evaluate
their role in lignocellulose degradation. Gene co-expression network analysis was then performed
using the RPKM values and the WGCNA package v1.51
(Langfelder and Horvath, 2008). Briefly, a softpower β was
chosen using the function pickSoftThreshold to fit the signed
network to a scale-free topology. Next, an adjacency matrix was
generated as follows: adj = (0.5 ∗(1+cor))β, where adj, cor
and β are adjacency, pairwise Pearson correlation and softpower
value, respectively. Topological Overlap Matrix (TOM) was
used as an input in the function hclust (“average” method)
to construct a hierarchical clustering tree (dendrogram). TOM
is a measure that quantifies the topological similarity between
the genes within a network, i.e., it evaluates whether two or
more nodes share links within the network and groups them
into the same module (Ravasz et al., 2002; Langfelder, 2013). A threshold of 0.15 (correlation > 85%) was chosen to merge
similar modules, and only modules having at least 30 genes were
kept. Network visualization and analyses for the highly connected
genes (absolute Pearson correlation > 0.8, adj = 0.064) were
carried out in Cytoscape v3.3.0. The entire R script used in the
WGCNA analysis is available in Data Sheet 1. Fungal Strain and Culture Conditions Biological process and molecular function annotation
from the Gene Ontology (GO) consortium was obtained from
the JGI database (https://genome.jgi.doe.gov/TrireRUTC30_1/
TrireRUTC30_1.home.html) and used for the enrichment
analysis using Cytoscape’s plugin BiNGO v3.0.3 (Maere et al.,
2005) configured to perform hypergeometric test and adjust p-
values for multiple testing using the Benjamini & Hochberg’s
false discovery rate (FDR) method (p ≤0.05). To completely
annotate the function of the T. reesei proteins, KEGG and
KOG annotations were retrieved from the JGI database. In
addition, manually curated annotation of the QM6a strain based Gene Co-expression Network Analysis Gene Co-expression Network Analysis
The reads obtained by Borin et al. (2017) (BioProject accession
PRJNA350272) were size-filtered (minimum of 40 bp) and
selected by quality (Q > 20) using AlienTrimmer software
(Criscuolo and Brisse, 2013). The filtered reads were mapped to
the T. reesei RUT-C30 v1.0 genome available in the JGI Genome October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 3 Gene Co-expression Network of Trichoderma reesei Borin et al. with secretion signal peptides were gathered in a few modules. Most of the upregulated genes of these functional classes were
found in four modules: coral1, darkorange, black, and darkred
(Figure 1A, Table S1), while most of the downregulated genes
were in additional four modules: brown, darkolivegreen4, grey60,
and lightcyan1 (Figure 1B, Table S1). The first four modules will
be designated the “up set” and the latter four the “down set.”
These different classes of DEGs are of major importance to the
identification of new targets related to bagasse deconstruction
and other processes associated, as they cover enzymes for the
lignocellulose breakdown, transcriptional regulators of the gene
expression, transport of inducer-acting molecules and other
unknown players. Taken together, these modules represented
more than half of all the DEGs in the T. reesei transcriptome. For
this reason, further analyses focused only on them. on the trichoCODE pipeline (Druzhinina et al., 2016) was also
transferred to the RUT-C30 strain using clusters of 1:1 orthologs
generated by applying OrthoMCL pipeline (Li et al., 2003). Identification of Subclusters and Go
Enrichment reesei network and gathered a significant share of
the CAZyme and predicted sugar transporter genes (Figure 1A). Therefore, it is not surprising that the enrichment of these GO
terms were identified for these modules and shows that our
analyses were directed toward the identification of the genes of
unknown function, which were co-expressed with known players
in lignocellulose degradation and/or sugar transport. Prediction of the Xyr1-Binding Sites The promoter region of the T. reesei RUT-C30 genes was
searched for Xyr1-binding sites (XBS) according to a modified
pipeline developed by Silva-Rocha et al. (2014). The 1.5 kb
sequences immediately upstream from the start codon ATG
of 22 cellulase genes regulated directly by Xyr1 (Castro et al.,
2014) were retrieved from the JGI database according to an
in-house Biopython script available online (https://github.com/
SantosRAC/UNICAMP_RACSMaster/tree/master/GFFTools). For motif discovery, these sequences were used as input in
the MEME program (Bailey et al., 2009) using the following
parameters: (i) zero or one occurrence per sequence at the
forward and reverse strand, (ii) a minimum of 10 motifs, and (iii)
a minimum and maximum motif width of 6 and 10, respectively. The frequency matrix of the motif most similar to the Xyr1
consensus sequence 5
′-GGC(A/T)3-3
′ (Rauscher et al., 2006)
was chosen to be sought in the promoter of the network genes
using the matrix-scan tool from the RSAT server (p-value ≤
1.00E-04)
(http://rsat-tagc.univ-mrs.fr/rsat/matrix-scan_form. cgi). KOG annotation was used to identify the functional groups
of genes that have the predicted XBS, and a hypergeometric test
was applied to enrich the statistically significant KOG groups
(p-value ≤1.00E-03) using KOG annotation for all the genes
as background. P-values were adjusted for multiple testing
corrections using Benjamini & Hochberg’s false discovery rate
(FDR) method (p ≤0.05). Identification of Subclusters and Go
Enrichment Enrichment
Subclusters of co-expressed genes were classified using the
MCODE algorithm to find protein coding genes acting together
during the process of lignocellulose deconstruction from bagasse. Each of the modules previously identified was partitioned, and a
few subclusters were formed for each module from the up and
down sets (Table S2). Next, a GO enrichment was conducted
with the genes within each subcluster. Only subclusters having
at least 10 genes were considered for GO enrichment (Table S3). Overall,
subclusters
of
coral1,
black,
darkorange,
and
darkred modules presented 26 GO terms enriched, including
cellulose
binding
(GO:0030248)
and
hydrolase
activity
(GO:0004553,
0016798),
transferase
activity
(GO:0016758,
0016740), carbohydrate transport (GO:0008643) and regulation
of transcription (GO:0006355), and transcription factor activity
(GO:0003700), respectively (Table S3). As already described,
these modules had the largest number of upregulated genes of
the whole T. reesei network and gathered a significant share of
the CAZyme and predicted sugar transporter genes (Figure 1A). Therefore, it is not surprising that the enrichment of these GO
terms were identified for these modules and shows that our
analyses were directed toward the identification of the genes of
unknown function, which were co-expressed with known players
in lignocellulose degradation and/or sugar transport. Subclusters of co-expressed genes were classified using the
MCODE algorithm to find protein coding genes acting together
during the process of lignocellulose deconstruction from bagasse. Each of the modules previously identified was partitioned, and a
few subclusters were formed for each module from the up and
down sets (Table S2). Next, a GO enrichment was conducted
with the genes within each subcluster. Only subclusters having
at least 10 genes were considered for GO enrichment (Table S3). during the process of lignocellulose deconstruction from bagasse. Each of the modules previously identified was partitioned, and a
few subclusters were formed for each module from the up and
down sets (Table S2). Next, a GO enrichment was conducted
with the genes within each subcluster. Only subclusters having
at least 10 genes were considered for GO enrichment (Table S3). g
(
)
Overall,
subclusters
of
coral1,
black,
darkorange,
and
darkred modules presented 26 GO terms enriched, including
cellulose
binding
(GO:0030248)
and
hydrolase
activity
(GO:0004553,
0016798),
transferase
activity
(GO:0016758,
0016740), carbohydrate transport (GO:0008643) and regulation
of transcription (GO:0006355), and transcription factor activity
(GO:0003700), respectively (Table S3). As already described,
these modules had the largest number of upregulated genes of
the whole T. Frontiers in Bioengineering and Biotechnology | www.frontiersin.org Construction of a Weighted Gene
Co-expression Network Recently, the transcriptome of T. reesei RUT-C30 grown on
bagasse after a time course of 6, 12 and 24 h as well as on
fructose after 24 h was investigated (Borin et al., 2017). Several
DEGs were identified, including CAZymes, genes encoding sugar
transporters and uncharacterized TFs and proteins. Using this
dataset, a gene co-expression network was inferred to identify
new targets related to the cell wall degradation of bagasse. g
g
g
p
Alternatively, the down set demonstrated only 15 GO
terms enriched, such as threonine-type endopeptidase activity
(GO:0004298)
(module
brown),
carbohydrate
biosynthetic
process (GO:0016051) (darkolivegreen4), DNA conformation
change (GO:0071103) and electron carrier activity (GO:0009055)
(grey60),
and
anion
binding
(GO:0043168)
(lightcyan1)
(Table S3). The difference between the GO terms enriched in
the up and down sets could be explained by the carbon sources
used in the culture media. Bagasse is a complex recalcitrant
structure that must be broken down by the combined action of
CAZymes. The sugars released in this process are transported
to the intracellular environment, where they are utilized in
carbohydrate catabolism to produce energy. These sugars,
including cellobiose and xylose, can also act as inducers of
cellulase and hemicellulase expression (Mach-Aigner et al., From 9,852 genes, 8,402 were kept for further analyses after
gene expression filtering and normalization. The data were
imported into the WGCNA package, and a softpower β of 26 (R²
= 0.85) was chosen to fit the scale independence to a scale-free
topology and taking into account the connectivity between the
genes based on their expression (Data Sheet 2A). Genes having
similar expression patterns were grouped into modules, and
highly connected modules were merged (Data Sheet 2B). In total, 28 different modules were formed with the genes
highly co-expressed (Pearson correlation > 0.8). The DEGs
identified and annotated by Borin et al. (2017) were sought
within each module generated using the WGCNA package, and
∼70% of all of the up and downregulated genes were identified
in the T. reesei network (Table 1, Table S1). Genes encoding
CAZymes, TFs, sugar transporters and uncharacterized proteins October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 4 Gene Co-expression Network of Trichoderma reesei Borin et al. TABLE 1 | Modules of co-expressed genes in the T. reesei network and number of DEGs found within each module. Construction of a Weighted Gene
Co-expression Network Set
Module
Nodes
Edges
Up (% total)8
Down (% total) 8
Down
Brown
1676
1149157
–
672 (44.8)
Up
Coral1
1173
482242
386 (26.2)
–
Up
Black
875
335415
157 (10.6)
2 (0.1)
Up
Darkorange
683
156692
270 (18.3)
–
Down
Darkolivegreen4
615
136748
–
144 (9.6)
Up
Darkred
607
124171
170 (11.5)
–
Coral2
375
45234
16 (1.1)
10 (0.7)
Down
Grey60
370
53283
–
130 (8.7)
Down
Lightcyan1
332
35644
–
27 (1.8)
Brown2
291
28624
–
6 (0.4)
Navajowhite2
159
8570
5 (0.3)
8 (0.5)
Bisque4
135
6022
10 (0.7)
–
Royalblue
117
4724
–
22 (1.5)
Darkgreen
116
4704
–
1 (0.1)
Yellowgreen
91
895
–
–
Plum1
85
1537
2 (0.1)
3 (0.2)
Ivory
79
1641
–
–
Brown4
72
743
6 (0.4)
–
Darkslateblue
71
1258
–
–
Thistle2
66
1552
–
4 (0.3)
Lavenderblush3
62
1464
–
1 (0.1)
Darkseagreen4
59
1464
–
–
Orangered3
52
1143
4 (0.3)
–
Lightsteelblue
49
579
2 (0.1)
3 (0.2)
Lightcoral
48
470
3 (0.2)
–
Firebrick4
45
548
–
–
Blue2
43
378
3 (0.2)
–
Darkviolet
41
671
2 (0.1)
–
Total
8387
2585573
1036 (70.1)
1033 (68.9)
Genes were up or downregulated in at least one time point of T. reesei grown on sugarcane bagasse. Nodes and edges represent genes and pairwise interactions, respectively. 8 The percentage was calculated based on the total number of up (1475) and downregulated (1500) genes expressed in the transcriptome of T. reesei (Borin et al., 2017). co-expressed genes in the T. reesei network and number of DEGs found within each module. Genes were up or downregulated in at least one time point of T. reesei grown on sugarcane bagasse. Nodes and edges represent genes and pairwise interactions, respectively. 8 The percentage was calculated based on the total number of up (1475) and downregulated (1500) genes expressed in the transcriptome of T. reesei (Borin et al., 2017). amino acid transporters; CAZymes; TFs; proteins of chromatin
remodeling; enzymes from carbohydrate, amino acid, lipid and
nucleotide metabolism; chaperones and hypothetical proteins
without annotated function (Table S5). 2010; Zhou et al., 2012; Zhang et al., 2013). In contrast, the
disaccharide fructose is a readily assimilable sugar that does not
require cellulase enzymes to be utilized by the fungus. Therefore,
it is evident that T. reesei adapts its metabolism according to the
available carbon source. Construction of a Weighted Gene
Co-expression Network For simplification purposes, only a few hubs are shown in
Table 2. In the up set, 3 CAZymes, 9 sugar transporters, 6 TFs and
5 genes encoding unknown proteins with a predicted secretion
signal peptide were found to be hub genes. A few TFs and
other proteins having a putative regulatory function in gene
transcription were identified as hub genes, and most were only
characterized in their corresponding fungal orthologs, including
the PRO1 Zn2Cys6 transcriptional regulator (jgi|136533, module
darkorange) (Masloffet al., 1999), the PRO41 protein (jgi|8730,
module coral1) (Nowrousian et al., 2007) and the AMA1
activator (jgi|114362, module coral1) (Diamond et al., 2009)
(Table 2). Interestingly, two genes encoding methyltransferases
(jgi|79832, module darkred; jgi|72465, module coral1) and one
gene encoding a GCN5-acetyltransferase (jgi|133861, module
coral1) were also identified as hub genes. Among the hub genes
encoding proteins of unknown function with signal peptides, the Hub Genes Identification Important genes for a biological process tend to be central
in a gene co-expression network and share a high number of
co-expressed neighbors having a lower degree (Villa-Vialaneix
et al., 2013). In this study, hubs were defined as the genes at
the top of the degree distribution and those that demonstrated
a positive correlation between the degree and betweenness
centrality (Table S4). In total, 321 and 294 hubs were identified
in the up and down sets, respectively (Table S5). Among the
321 hubs, 129 (40%) were upregulated in bagasse in at least one
time point (6, 12 or 24 h), and 191 (65%) out of 294 genes
were downregulated in the down set. Various genes encoding
proteins that have different functions were found to be hubs
in all eight modules investigated, including sugar, ion, and October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 5 Gene Co-expression Network of Trichoderma reesei Borin et al. Borin et al. FIGURE 1 | DEGs identified in the modules of the T. reesei RUT-C30 network encoding CAZyme, sugar transporter, transcription factor and unknown proteins with
signal peptide. Most of the upregulated (A) and downregulated (B) genes in sugarcane bagasse were found in a few modules. work encoding CAZyme, sugar transporter, transcription factor and unknown proteins with
i
b
f
d i
f
d l FIGURE 1 | DEGs identified in the modules of the T. reesei RUT-C30 network encoding CAZyme, sugar transporter, transcription factor and unknown proteins with
signal peptide. Most of the upregulated (A) and downregulated (B) genes in sugarcane bagasse were found in a few modules. (Rauscher et al., 2006) and the motif used in the study of Silva-
Rocha et al. (2014). As a result, only one putative XBS of 10
nucleotides resembling the Xyr1 motifs was chosen for further
analyses (Figure 2). The frequency matrix of the motif chosen
was then retrieved from the MEME server and used as a model
in the identification of XBS predicted within the promoter of the
genes from the up and down sets. The complete list of the motifs
predicted in the promoter of the 22 CAZymes and the frequency
matrix is available in Table S6. (Rauscher et al., 2006) and the motif used in the study of Silva-
Rocha et al. (2014). Hub Genes Identification As a result, only one putative XBS of 10
nucleotides resembling the Xyr1 motifs was chosen for further
analyses (Figure 2). The frequency matrix of the motif chosen
was then retrieved from the MEME server and used as a model
in the identification of XBS predicted within the promoter of the
genes from the up and down sets. The complete list of the motifs
predicted in the promoter of the 22 CAZymes and the frequency
matrix is available in Table S6. genes jgi| 124417, 128655 were co-expressed in the module coral1
and darkorange, respectively, and were upregulated in bagasse. The trichoCODE annotation indicated that their orthologous
genes in T. reesei QM6a encode SSCRPs, and therefore they
could be acting in the extracellular environment (Table 2,
Table S5). However, the true evidence that they are pivotal to
T. reesei response when grown on bagasse still needs to be
evaluated. Similar to the up set, various hub genes encoding different
proteins were found in the down set. In total, 7 CAZymes, 5
sugar transporters, 8 TFs and 4 proteins of unknown functions
that have predicted secretion signal peptide were found to
be downregulated in bagasse. In addition to the identification
of the hub genes that have a putative regulatory role in the
gene expression, such as TFs, chromatin remodeling and signal
transduction proteins, several genes encoding unknown proteins
significantly downregulated were also found, especially in the
brown module (Table 2, Table S5). To validate the motifs found in silico, they were compared
with the motifs already characterized in the promoters of the
CAZyme genes (cbh1 and xyn1) of other T. reesei strains. Cellobiohydrolase 1 (Cbh1/Cel7a) is one the most produced and
secreted enzymes from T. reesei (Kiesenhofer et al., 2018), and
endo-β-1,4-xylanase 1 (Xyn1) is a hemicellulase with important
role in xylan deconstruction (Liu et al., 2017). The comparison
of the motifs showed that our pipeline could predict XBS that
had been previously described and characterized (Figures S3,
S4) (Rauscher et al., 2006; Furukawa et al., 2009; Ries et al.,
2014; Kiesenhofer et al., 2018), similarly to the prediction of
Silva-Rocha et al. (2014). Frontiers in Bioengineering and Biotechnology | www.frontiersin.org Prediction of the Xyr1-Binding Sites in the
Promoter Region Next, the predicted XBS found in the promoter of the genes
from the up and down sets were investigated (Table S7). In
total, 245 upregulated and 210 downregulated genes had at least
one XBS predicted in the promoter region. This represented
24 and 22% of the entire set of DEGs present in the up and
down sets, respectively. Considering only the number of DEGs
in each module, coral1 (45.5%) and grey60 (40.4%) were the
modules having the largest percentage of genes with predicted
XBS (Table S1). Previously, Castro et al. (2014) showed that 22 genes encoding
cellulases and hemicellulases of the T. reesei QM9414 strain are
regulated directly by the master activator Xyr1 (jgi|98788). This
TF is essential for the induction of cellulase and hemicellulase
genes, and it has been the target of various studies aiming to
improve the hypercellulolytic phenotype of T. reesei using genetic
manipulation (Wang et al., 2013; Lv et al., 2015; Zhang et al.,
2017). To predict the presence of a specific DNA binding motif for
the RUT-C30 strain, the promoter regions (1.5 kb immediately
upstream the start codon ATG) of the orthologous genes between
the RUT-C30 and QM9414 strains were used to seek motifs that
were similar to the Xyr1 consensus sequence 5′-GGC(A/T)3-3′ KOG functional annotation of the genes having predicted
XBS from up and down sets was also investigated (Figure 3). The hypergeometric test and multiple testing correction showed
that the KOG classes metabolism, carbohydrate transport and
metabolism, and information storage and processing
were October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 6 Gene Co-expression Network of Trichoderma reesei Borin et al. TABLE 2 | Hub genes found in the up and down sets that were differently expressed in sugarcane bagasse (Borin et al., 2017). Prediction of the Xyr1-Binding Sites in the
Promoter Region In the down set, 33 (17.3%) hub genes had XBS predicted in this
study, including two CAZymes (chitin synthase and glucan endo-
1,3-β-glucosidase), one ABC transporter family protein, one
putative nonribosomal peptide synthase (NRPS), one putative
SWI-SNF chromatin-remodeling complex protein and several
uncharacterized proteins (Table S5). FIGURE 2 | Putative XBS predicted in the promoter of 22 genes encoding
cellulases and hemicellulases and chosen to be sought in the gene promoter
regions of T. reesei. Only this XBS was found to resemble the Xyr1 consensus
sequence 5
′-GGC(A/T)3-3
′ and the motif used by Silva-Rocha et al. (2014). Prediction of the Xyr1-Binding Sites in the
Promoter Region RUT-C30 ID1
QM6a ID2
Description3
Module
Degree
log2FC4
B6h
B12h
B24h
Up sets
23456
122048
Sec61 beta subunit of ER translocase
Black
859
–
–
1.09
103979
81884
Translocon-associated protein (TRAP)
Black
859
–
–
1.06
133861
111236
GCN5-related N-acetyltransferase
Coral1
1106
3.29
3.18
4.12
8730
57737
PRO41 protein
Coral1
1086
2.03
1.27
2.69
114362
67408
AMA1 activator
Coral1
1097
–
–
1.11
139402
121107
Zn2Cys6 transcriptional regulator
Coral1
1104
–
–
1.38
95791
120698
C2H2 transcriptional regulator PacC
Darkorange
652
2.28
2.15
1.94
136533
76590
Zn2Cys6 transcriptional regulator Pro1
Darkorange
650
2.01
2.00
2.09
79832
5366
S-adenosyl-L-methionine-dependent Methyltransferase
Darkred
558
1.93
1.71
–
38522
103158
Zn2Cys6 transcriptional regulator
Darkred
572
2.12
1.99
–
Down sets
98900
78688
Heat shock factor-type DNA-binding domain-containing protein
Brown
1642
−1.69
−1.53
−2.29
140865
106171
HET protein
Brown
1640
−4.10
−2.82
−4.99
124027
122943
SWI-SNF chromatin-remodeling complex protein
Brown
1639
–
−1.10
−1.28
77229
106259
Zn2Cys6 transcriptional regulator
brown
1643
−2.18
−1.86
−2.86
139518
124228
GT2 chitin synthase
Darkolivegreen4
572
–
–
−1.28
86800
66606
Zn2Cys6 transcriptional regulator
Darkolivegreen4
577
–
–
−1.11
139776
21255
bHLH transcriptional regulator
Grey60
347
−1.12
−1.05
–
131957
53893
Hypothetical protein
Grey60
353
−2.17
−2.79
−1.55
105289
82667
ThrB Homoserine kinase
Lightcyan1
307
−1.22
−1.17
–
96918
120953
Unknown secreted protein
Lightcyan1
310
−3.29
−3.04
–
1Trichoderma reesei RUT C30 v1.0 database from JGI was used to recover the T. reesei proteins ID; 2QM6a ortholog genes; 3Functional annotation according to KEGG, KOG and
Druzhinina et al.’s work (2016); 4Log2 fold change (FC), B: sugarcane bagasse. Red and blue colors indicate gene expression of genes up and downregulated in sugarcane bagasse,
respectively. Major Facilitator Superfamily (MFS) and one putative calcium
transporter, one Ras GTPase, one putative SWI-SNF chromatin-
remodeling complex protein, one polyketide synthase (PKS) and
various hypothetical proteins with unknown function (Table S5). In the down set, 33 (17.3%) hub genes had XBS predicted in this
study, including two CAZymes (chitin synthase and glucan endo-
1,3-β-glucosidase), one ABC transporter family protein, one
putative nonribosomal peptide synthase (NRPS), one putative
SWI-SNF chromatin-remodeling complex protein and several
uncharacterized proteins (Table S5). Major Facilitator Superfamily (MFS) and one putative calcium
transporter, one Ras GTPase, one putative SWI-SNF chromatin-
remodeling complex protein, one polyketide synthase (PKS) and
various hypothetical proteins with unknown function (Table S5). Xyr1 and Co-expressed Genes in the
Coral1 Module DISCUSSION transcriptional regulator, one protein with an ankyrin repeat, and
several other genes still not characterized (Figure 4B, Table S8). cbh2), endoglucanases (egl1, egl2, egl3, and egl5), β-glucosidases
(bgl1 and bgl2), endo-β-1,4-xylanases (xyn2, xyn3, xyn4, and
xyn5), and monooxygenases from the AA9 family (cel61a and
cel61b) (Figure 4A, Table S8). Xyr1 and Co-expressed Genes in the
Coral1 Module As already described, Xyr1 is critical for the activation of
cellulases and hemicellulases, and it was expressed in bagasse
more highly than fructose during the entire time course (Log2FC:
6 h: 0.48; 12 h: 0.99; 24 h: 1.65). Along with the other 385
upregulated genes, xyr1 was grouped into the coral1 module
which had the largest number of CAZyme genes upregulated in
bagasse (Figure 1A). Thus, it is worth investigating the genes
that were co-expressed with this activator, since they could be
involved in the fungal response to the lignocellulosic biomass. FIGURE 2 | Putative XBS predicted in the promoter of 22 genes encoding
cellulases and hemicellulases and chosen to be sought in the gene promoter
regions of T. reesei. Only this XBS was found to resemble the Xyr1 consensus
sequence 5
′-GGC(A/T)3-3
′ and the motif used by Silva-Rocha et al. (2014). statistically significant in the up set, while secondary metabolites
biosynthesis, transport and catabolism was the only class enriched
in the down set (Figure 3). This difference stresses the diversity
of genes encoding proteins with several functions that could be
modulated by the activator Xyr1, in addition to other elements. From the pairwise edges of the coral1 module, 858 neighbor
nodes to Xyr1 were retrieved. A total of 338 out of 858 genes were
upregulated in bagasse, and only 115 had at least one XBS in the
promoter. Among these genes, 35 CAZymes were found to be
co-expressed with xyr1, including cellobiohydrolases (cbh1 and y
y
Analyzing the hub genes of the up set, 30 genes (23.3%)
demonstrated at least one XBS in their promoter, including
one Zn2Cys6 transcriptional regulator, transporters from the October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 7 Borin et al. Gene Co-expression Network of Trichoderma reesei FIGURE 3 | KOG function classification of the DEGs identified in the up and down sets that had at least one predicted XBS. KOG classes significantly enriched
(p-value ≤1.00E-03) are shown with an asterisk (*). FIGURE 3 | KOG function classification of the DEGs identified in the up and down sets that had at least one predicted XBS. KOG classes significantly enriched
(p-value ≤1.00E-03) are shown with an asterisk (*). transcriptional regulator, one protein with an ankyrin repeat, and
several other genes still not characterized (Figure 4B, Table S8). DISCUSSION The list of hubs
included 14 genes encoding one MFS transporter, one Zn2Cys6 October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 8 Gene Co-expression Network of Trichoderma reesei Borin et al. FIGURE 4 | (A) Upregulated genes encoding proteins of different functions being co-expressed with the activator xyr1 in the coral1 module. (B) Hub genes identified
in the same module having at least one predicted XBS in the promoter region. The nodes are identified with the T. reesei RUT-C30 ID retrieved from JGI database. The
complete list of co-expressed genes with xyr1 is shown in Table S8. FIGURE 4 | (A) Upregulated genes encoding proteins of different functions being co-expressed with the activator xyr1 in the coral1 module. (B) Hub genes identified
in the same module having at least one predicted XBS in the promoter region. The nodes are identified with the T. reesei RUT-C30 ID retrieved from JGI database. The
complete list of co-expressed genes with xyr1 is shown in Table S8. et al., 2018), clarifying the importance of these putative sugar
transporters in the induction of CAZymes and lignocellulose
degradation. were chosen for further analysis due to the largest number of
up and downregulated genes encoding CAZymes, TFs, sugar
transporters and other genes of unknown function that were
co-expressed (Figure 1). g
Distinct GO terms were enriched in the down set, such
as threonine-type endopeptidase activity (GO:0004298) and
carbohydrate biosynthetic process (GO:0016051) (Table S3). The
latter was enriched in the darkolivegreen4, and interestingly, this
GO term consisted of seven enzymes possibly involved in fungal
cell wall biosynthesis, including three glycosyl transferases, one
α-amylase, one rhamnose reductase, one pyruvate carboxylase,
and one glucose 6-phosphate isomerase (GPI) (Table S3). Fungal cell wall is primarily composed of β-1,3-glucan, β-1,6-
glucan, mixed β-1,3-/ β-1,4-glucan, α-1,3-glucan, chitin, and
glycoproteins. They are organized in a complex backbone, and
it is thought that glycosyl hydrolases and glycosyl transferases are
fundamentally important in cell wall biosynthesis (Free, 2013). Genes encoding enzymes from the gluconeogenesis pathway,
such as pyruvate carboxylase and GPI, could also provide hexose
phosphates as building blocks for the cell wall biosynthesis (Ene
et al., 2012). Considering that fructose is a simpler sugar than
the bagasse lignocellulose, the fungus must be utilizing this
readily assimilable carbon source to grow, and consequently it
has to remodel its cell wall. DISCUSSION Interestingly, swo1 (jgi|104220) and cip1 (jgi| 121449) were
found as neighbor nodes to xyr1, and they were under strong
activation in bagasse during the entire time course (6, 12, and
24 h) (Table S8). Swo1 acts synergistically with xylanases to
remove the hemicellulosic fraction of the lignocellulose (Gourlay
et al., 2013), and Cip1 appears to be important to the hydrolysis of
the lignocellulosic biomass (Lehmann et al., 2016). Both proteins
have a carbohydrate-binding domain that belongs to family 1
(CBM1), and one XBS was predicted in each gene promoter. Therefore, they play a role in bagasse deconstruction and
are transcriptionally regulated by Xyr1, as reported previously
(Reithner et al., 2014; Ma et al., 2016). In addition, 14 putative
sugar transporters, three Zn2Cys6 transcriptional regulators,
one putative GCN5-acetyltransferase, one methyltransferase and
several other genes were also co-expressed with xyr1 and had XBS
in their promoters (Figure 4A, Table S8). The construction of a gene co-expression network allows us to
identify clusters of genes that have a similar expression pattern
and to assess the biological information that is relevant to
a specific phenotype. Hitherto there were only a few studies
investigating gene co-expression network in T. reesei. Dos Santos
Castro et al. (2014), for instance, performed RNA-Seq of T. reesei QM9414 strain grown on sophorose, cellulose and glucose,
and their analysis revealed specific differentially expressed genes
in sophorose and cellulose. More recently, Horta et al. (2018)
examined the transcriptome and exoproteome of T. reesei, T. harzianum, and T. atroviride grown on cellulose and glucose. Based on their co-expression network, they found a set of 80
genes shared between the three Trichoderma species that could
represent a common cellulose degradation system. However, no
research has been reported exploring the gene co-expression
network of T. reesei RUT-C30 grown on sugarcane bagasse. In this study, the recently published transcriptome of T. reesei
RUT-C30 grown on bagasse after 6, 12 and 24 h was used to
determine the modules of co-expressed genes. As a result, 28
modules of co-expressed genes were formed, and only eight Finally, hub genes that were neighbor nodes to the master
activator Xyr1 (represented in the coral1 module) and that had
XBS in their promoter were also identified. Frontiers in Bioengineering and Biotechnology | www.frontiersin.org DISCUSSION Although most of those genes were
downregulated and highly co-expressed, the engagement of these
genes in this process remains elusive. p
g
After the MCODE clustering, the GO enrichment showed
that
several
terms
related
to
lignocellulose
breakdown,
including polysaccharide (GO:0030247) and cellulose binding
(GO:0030248),
and
hydrolase
activities
(GO:0004553
and
GO:0016798), were enriched in the subcluster 1 of the coral1
module (Table S3) where the master activator Xyr1 was also
found. The presence of 50 upregulated CAZyme genes encoding
cellulases, hemicellulases and oxidative enzymes in the coral1
module (Table S1), along with the co-expression of xyr1,
allows us to hypothesize that soon after xyr1 is expressed in
bagasse, its protein product regulates the transcription of the
(hemi)cellulolytic genes. In addition, 36 out of the CAZyme
50 genes demonstrated at least one XBS in their promoter,
which is consistent with previous studies (Castro et al., 2014;
Silva-Rocha et al., 2014) and highlights the ability of our
approach to identify the Xyr1 target genes. In addition to
coral1, the darkorange module had the carbohydrate transport
(GO:0008643) term enriched (Table S3), revealing that several
genes encoding transporters were grouped in this module. Most of them belonged to the MFS family, and its members
are thought to transport a vast array of small molecules, such
as sugars, inorganic ions, siderophores and amino acids (Yan,
2015; Quistgaard et al., 2016). Some studies have also shown that
MFS transporters can still mediate the induction of cellulases
in T. reesei (Zhang et al., 2013; Huang et al., 2015; Nogueira Hub genes were also sought out within the main modules,
and a large number of new targets that were co-expressed
were discovered (Table 2, Table S5). For example, the Pro1
coding gene (jgi|136533) was found as a hub node within the October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 9 Gene Co-expression Network of Trichoderma reesei Borin et al. identity: 33%) encodes a meiosis-specific activator related to
spore morphogenesis (Okaz et al., 2012; Schmoll et al., 2016). One gene encoding the developmental regulatory protein WetA
(jgi|9281) was found in the same module being upregulated
during the entire time course. In Fusarium graminearum, the
putative ortholog WetA (I1S0E2.2; identity: 43%) is necessary for
conidiogenesis and conidial maturation, and the wetA mutants
produced conidia that were more sensitive to oxidative and heat
stress (Son et al., 2014). Intriguingly, Wu et al. DISCUSSION (2017) suggested
that the Aspergillus flavus ortholog WetA (EED47149.1; identity:
57%) could be a global player in the regulation of conidial
development, acting during the fungal cell wall biogenesis and
regulating the secondary metabolic pathways. In addition, the
putative TF (jgi|98900) was one of the hub genes of the brown
module, and its corresponding orthologs encode the heat shock
transcription factor 1 (Hsf1) (Schmoll et al., 2016), the master
stress response regulator in eukaryotes (Zheng et al., 2016) and
the activator of virulence in the pathogen Candida albicans
(Nicholls et al., 2011) (Table 2, Table S5). Another four and seven
putative TF coding genes were also found to be hubs in the
up and down set, respectively (Table S5). The search for their
orthologous genes in other ascomycetes revealed that none had
been characterized, and therefore, their function remains to be
elucidated. The relevant role of the characterized ortholog hub
genes identified in this study highlights the central position of
these nodes in the up and down sets and support the necessity of
investigating their putative regulatory influence on the T. reesei
response in bagasse. darkorange module, and it was upregulated in the three time
points analyzed (6, 12, and 24 h). Pro1 is a member of the
Zn2Cys6 transcription factors, and several orthologs had already
been characterized in ascomycetes. In Neurospora crassa, the
ortholog Adv-1 (NCU07392; identity: 66%) regulates cell-to-cell
fusion and sexual development (Chinnici et al., 2014; Dekhang
et al., 2017), and in Sordaria macrospora (CAB52588.2; identity:
66%), Pro1 has a pivotal role in various different processes,
including the regulation of genes involved in cell wall integrity,
the NADH oxidase pathway and pheromone signaling (Masloff
et al., 2002; Steffens et al., 2016). Steffens et al. (2016) demonstrated that, in addition to Adv-1,
N. crassa requires the pH response transcription regulator PacC
to activate its female development. Intriguingly, its ortholog gene
pac1 (jgi|95791) in T. reesei RUT-C30 was also identified to be a
hub node (Table 2, Table S5). pac1 was upregulated in bagasse
and co-expressed with the pro1 gene in the same module. He
et al. (2014) functionally characterized pac1 in T. reesei, and they
observed increased cellulase transcription and production in the
1pac1 mutants at neutral pH. They also noted that the pac1
deletion impaired fungal growth and development. DISCUSSION In total, the
results point to possible crosstalk between Pro1, Pac1 and other
genes involved in an intricate regulatory network. As already described, two putative methyltransferase coding
genes (jgi|79832, module darkred; jgi|72465, module coral1)
and a GCN5-acetyltransferase coding gene (jgi|133861, module
coral1) were also identified as hubs in the up set (Table S5). These classes of enzymes are thought to be responsible for
DNA methylation and histone modification, respectively, and
epigenetically regulate various important cellular processes, such
as the silencing of transposable elements and the activation
of gene transcription (Xin et al., 2013; Su et al., 2016;
Lyko, 2017). Intriguingly, recent studies have suggested that
methyltransferases are also implicated in lignin degradation
in the basidiomycete Phanerochaete chrysosporium, broadening
the functions of these enzymes (Korripally et al., 2015; Thanh
Mai Pham and Kim, 2016; Kameshwar and Qin, 2017). In
addition, five genes encoding unknown proteins that have
secretion peptide signal were considered to be hubs in the
up set (Table S5). Among them, two (jgi|124417, 128655) had
their QM6a orthologs annotated as annotated as small secreted
cysteine-rich proteins (SSCRPs) using trichoCODE. In general,
SSCRPs are related to interactions between the microorganisms
and the environment, including biocontrol, the induction of
plant resistance and adhesion (Shcherbakova et al., 2015; Qi
et al., 2016). The presence of a predicted secretion signal
peptide allows us to hypothesize that these proteins could be
secreted to support the bagasse-fungal adhesion. In total, the
identification of these hub genes shows that they could be
acting as regulatory players or accessory proteins in the bagasse
degradation response. Finally, other genes encoding proteins of different functions
were also identified as hub nodes in the up and down sets
(Table S5). In the up set, three CAZymes (jgi|97768, 75420,
128705), two SSCRPs (jgi|124417, 128655), the translocon-
associated
protein
(TRAP)
(jgi|103979)
and
the
Sec61
beta subunit (jgi|23456) from the endoplasmic reticulum
(ER)
translocon,
ion
and
amino
acid
transporters,
and
various unknown proteins were found. In the down set,
other
hubs
were
discovered,
including
seven
CAZymes
(jgi|24326, 104519, 103899, 124897, 97721, 139518, 104242),
six
proteases
(jgi|90298,
135507,
75405,
82006,
138263,
87936), a SWI-SNF chromatin-remodeling complex protein
(jgi|124027)
and
unknown
proteins
(Table 2,
Table S5). This vast array of hub genes demonstrates that the great
diversity of genes is central in the co-expressed modules. However, most of them are still not characterized, and
efforts are required to elucidate their function in fungal
physiology. Frontiers in Bioengineering and Biotechnology | www.frontiersin.org DISCUSSION The prediction of XBS based on the promoter of cellulase and
hemicellulase coding genes was used in this study to verify the
genes possibly regulated by Xyr1 in the presence of a complex
lignocellulosic biomass. Xyr1 is the key transcriptional regulator
of the CAZymes involved in cell wall deconstruction, and it
is regulated by the carbon catabolite repressor Cre1, which is
truncated and confers a hypercellulolytic phenotype in the RUT-
C30 strain (Ilmén et al., 1996; Silva-Rocha et al., 2014). To
validate our pipeline of regulatory motif predictions, the XBS
predicted in the cbh1 and xyn1 promoters were compared with
those from other studies. Other putative regulators were also identified as hub nodes in
the up and down sets, and some of them were not characterized
in T. reesei, only in its homologs (Table S5). For example,
the gene encoding the Ama1 activator (jgi|114362) was found
in the coral1 module having an XBS in the promoter, and
its putative ortholog in Saccharomyces cerevisiae (YGR225W; October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 10 Gene Co-expression Network of Trichoderma reesei Borin et al. According to Ries et al. (2014), there are two XBS in the
positions −733 and −320 in relation to the start codon ATG of
the cbh1 gene (Data Sheet S3). The −733 motif (5′-TTTGCC-
3′) was predicted by the pipeline of Silva-Rocha et al. (2014). However, it was not predicted in this study. Alternatively, we
identified four XBS in the promoter region of cbh1 at positions
−508, −748, −771, and −1376 that were not identified in the
study by Silva-Rocha et al. (2014). This was likely to be due
to the differences between the strains and pipelines. Using a
labeled Xyr155−195 probe, Furukawa et al. (2009) found that two
of these predicted sequences (positions−508 and−748) showed
strong binding to the probe. In addition, Kiesenhofer et al. (2018)
replaced the promoter of the glycine oxidase (goxA) reporter
gene with the cbh1 promoter containing deletions in three
different regions and observed a significant decrease in the GoxA
activity when the mutant strains were grown on different carbon
sources, including lactose, carboxymethylcellulose (CMC) and
pretreated wheat straw. Two of these regions spanned the
XBS predicted in this study at positions −748 and −771
(Data Sheet S3). DISCUSSION To verify the putative targets of Xyr1, the direct co-expressed
neighbors of the xyr1 node were retrieved from module1, and
we searched for the XBS in the gene promoters (Table S8). From a total of 115 Xyr1-partner genes having a predicted XBS,
almost half encode CAZymes and putative sugar transporters. Three Zn2Cys6 transcription factors (jgi|139402, 77124, 141251)
were also found, and one (jgi|141251) demonstrated 55 and 72%
identity to the NirA ortholog in F. fujikuroi (KLO85312.1) and
A. nidulans (AN0098), respectively (Table S8). NirA is a nitrate-
specific transcription factor that modulates nitrogen metabolite
repression (NMR), and this TF accumulates in the nucleus in
the presence of nitrate or nitrite and a low concentration of
assimilable nitrogen sources, such as ammonium. The nitrogen
metabolism regulator AreA interacts physically with NirA, and
the complex formed activates the genes for nitrate assimilation
(Gallmetzer et al., 2015; Pfannmüller et al., 2017). The T. reesei
ortholog areA (jgi|140814) was not differentially expressed in
bagasse, and therefore it is curious to note the upregulation of the
T. reesei ortholog nirA in this carbon source and its co-expression
with xyr1. In addition to cbh1, the XBS predicted in the xyn1 promoter
were also compared with the ones previously reported (Rauscher
et al., 2006; Furukawa et al., 2009; Kiesenhofer et al., 2018). Two
XBS were predicted at positions −417 and −621 being the first
one identified and characterized in other studies (Data Sheet S4). For example, Rauscher et al. (2006) investigated a 217-bp region
(−321 to −538) inside the xyn1 promoter and discovered that
Xyr1 was able to bind to two sequences at positions −404 and
−420. Point mutations in each of these two motifs caused a
substantial decrease in the reporter activity of the gene glucose
oxidase and showed that they are critical for the transcriptional
activation of xyn1 in the presence of xylan, one of the primary
sugar inducers of hemicellulases (Rauscher et al., 2006). Some
years later, Furukawa et al. (2009) analyzed three XBS (−80, −420
and −887) in the xyn1 promoter and discovered that only the
XBS at position −420 showed strong binding to the Xyr1 probe. Finally, Kiesenhofer et al. (2018) recently demonstrated that the
deletion of these XBS at positions −404 and −420 abolished
the GoxA activity under control of the xyn1 promoter even in
the presence of 0.5 mM xylose (Data Sheet S4). DISCUSSION In summary, the
previous studies indicate that some of the XBS predicted in this
study could be functional in T. reesei RUT-C30 and therefore, are
important for the transcriptional regulation of CAZyme genes
and probable additional genes. In addition to the cbh1 and xyn1,
the promoter of the cbh2 gene also had two XBS predicted in this
study that were shared with its homologs in other T. reesei strains
(data not shown). Three genes (jgi|126063, 73493, 92240) encoding putative
transporters of non-sugar solutes were co-expressed with xyr1
and had XBS in their promoters (Table S8). The gene jgi|73493
encodes a putative calcium transporter with 10 transmembrane
domains, and it was found as a hub in the coral1 module
(Table S8). It has already been reported that metal ions, such
as Ca2+ and Mn2+, have a positive effect on the mycelial
growth of T. reesei and cellulase production, and this molecular
signaling mechanism is mediated by the Mn2+ transporters
TPHO84-1 and TPHO82-2, a Ca2+/ Mn2+ ATPase (TPMR1)
and the components of the Ca2+/calmodulin signal transduction,
including the TF Crz1 (Chen et al., 2016, 2018). Therefore, it is
worth investigating the role of this putative calcium transporter
in the induction of the genes responsive to lignocellulose
degradation, since it could transport cations that activate gene
expression. In addition to the non-sugar transporters, 15 putative sugar
transporter coding genes were co-expressed with xyr1 and
demonstrated at least one XBS predicted at their promoter
region (Table S7). One of them was considered to be a hub
gene encoding a protein annotated as allantoate permease
(jgi|93149), but its participation in the fungal response toward
the biomass deconstruction remains unclear. Most of these
putative transporters are members of the MFS superfamily, and
therefore, could transport a variety of solutes, including sugar
and amino acids. Sloothaak et al. (2016) developed a Hidden
Markov Model (HMM) to identify new xylose transporters in
T. reesei and A. niger. Several candidates were identified, and
one (RUT-C30: jgi| 7811; QM6a ortholog: jgi| 106330) was found
upregulated and co-expressed with xyr1 in this study (Table S7). Unexpectedly, this putative xylose transporter coding gene had
one XBS predicted in the promoter and showed an increasing
expression profile in bagasse. The prediction of an XBS in the
promoter region of this gene suggests that it could be regulated
by Xyr1 and could be involved in the xylose assimilation after the
hemicellulose breakdown of bagasse. Frontiers in Bioengineering and Biotechnology | www.frontiersin.org REFERENCES Bornscheuer, U., Buchholz, K., and Seibel, J. (2014). Enzymatic degradation
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Trichoderma reesei Cel7A catalytic domain in open and closed states. FEBS Lett. 590, 4429–4438. doi: 10.1002/1873-3468.12464 Cologna, N., Gómez-Mendoza, D. P., Zanoelo, F. F., Giannesi, G. C.,
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enzymes of biotechnological interest. Enzyme Microb. Technol. 109, 1–10. DISCUSSION Based on the assumption that Xyr1 is essential to the induction
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that genes related to carbohydrate transport and metabolism,
such as CAZymes and putative sugar transporters, were the most
abundant upregulated genes that had XBS in the KOG analyses. Alternatively, the XBS were enriched in the genes of secondary
metabolites biosynthesis, transport, and catabolism class in the
down set (Figure 3), which suggests that Xyr1 participates in the
regulation of secondary metabolism. October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 11 Gene Co-expression Network of Trichoderma reesei Borin et al. ACKNOWLEDGMENTS We would like to thank the Laboratório Nacional de Ciência e
Tecnologia do Bioetanol (CTBE) and the Centro Nacional de
Pesquisa em Energia e Materiais (CNPEM) for the computational
facility. FUNDING This study received financial support from the Fundação de
Amparo à Pesquisa do Estado de São Paulo (FAPESP processes
n 2014/15799-7, 2014/11766-7, 2015/08222-8, and 2017/18987-
7), from the Conselho Nacional de Desenvolvimento Científico e
Tecnológico (CNPq process number 141574/2015-1), and also in
part by the Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior - Brasil (CAPES). AUTHOR CONTRIBUTIONS GB performed the analyses, and RdS carried out the data
processing. DR-P and MC supervised the study and performed
the analyses. JO supervised the study and planned the analyses. The Supplementary Material for this article can be found
online
at:
https://www.frontiersin.org/articles/10.3389/fbioe. 2018.00151/full#supplementary-material CONCLUSION All the authors wrote the draft and approved its final
version. The T. reesei gene co-expression network analysis grouped
several differentially expressed genes into modules based on their
expression patterns in steam-exploded sugarcane bagasse. A large
number of interesting genes encoding CAZymes, putative sugar
and ion transporters, as well as TFs and proteins with a putative
regulatory role were highly co-expressed within some modules. The prediction of XBS in the promoters confirmed the influence
of Xyr1 in the CAZyme coding genes regulation and enabled the
identification of new putative targets of this master regulator. Hub nodes were also found within the modules, and many of
them had not been characterized. Several CAZymes, accessory
proteins and uncharacterized protein coding genes were co-
expressed with xyr1. Finally, this study provided an extensive
number of genes that were co-expressed in bagasse. These genes
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and to the development of T. reesei hypercellulolytic strains, and
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with several of the authors, RS and DR, at the time of the review. Copyright © 2018 Borin, Carazzolle, dos Santos, Riaño-Pachón and Oliveira. This
is an open-access article distributed under the terms of the Creative Commons
Attribution License (CC BY). The use, distribution or reproduction in other forums
is permitted, provided the original author(s) and the copyright owner(s) are credited
and that the original publication in this journal is cited, in accordance with accepted
academic practice. No use, distribution or reproduction is permitted which does not
comply with these terms. Wang, S., Liu, G., Wang, J., Yu, J., Huang, B., and Xing, M. (2013). Enhancing
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bridges cellular and chemical development in Aspergillus flavus. PLoS ONE
12:e0179571. doi: 10.1371/journal.pone.0179571 October 2018 | Volume 6 | Article 151 Frontiers in Bioengineering and Biotechnology | www.frontiersin.org 15
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Recent Progress in Osteocyte Research
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Terms of Use This article was downloaded from Harvard University’s DASH repository, and is made available
under the terms and conditions applicable to Other Posted Material, as set forth at http://
nrs.harvard.edu/urn-3:HUL.InstRepos:dash.current.terms-of-use#LAA Permanent link http://nrs.harvard.edu/urn-3:HUL.InstRepos:11879695 Published Version
doi:10.3803/EnM.2013.28.4.255 doi:10.3803/EnM.2013.28.4.255 Citation Divieti Pajevic, Paola. 2013. “Recent Progress in Osteocyte Research.” Endocrinology and
Metabolism 28 (4): 255-261. doi:10.3803/EnM.2013.28.4.255. http://dx.doi.org/10.3803/
EnM.2013.28.4.255. INTRODUCTION blasts that during the process of matrix mineralization remain
entrapped in the mineralizing matrix that they are actively
synthesizing. They reside both in the mineralized matrix and
in the newly formed osteoid. Earlier work of Gaillard et al. [1]
and Rutishauser and Majno [2], described two stages into the
life of an osteocyte; an early stage (young osteocyte) in which
the cell is smaller in size, reside into the osteoid and do not
express alkaline phosphatase and a late stage (mature osteo-
cyte) in which the larger cell re-express alkaline phosphatase
and is deeply embedded in the mineralized bone. The larger
cell will then degenerate and leave an empty lacuna [1]. This
classification has been recently revised to include an addition-
al stage of differentiation: according to their spatial localiza-
tion and gene expression, these cells are now divided into os-
teoid, mineralizing, and the mature osteocytes [3]. The oste-
oid, or nascent osteocyte is characterized by a relative proxim- Our understanding of the function of osteocytes has expanded
dramatically over the last decade primarily due to the identifi-
cation of osteocytes specific markers, such as dentin matrix
protein 1 (DMP1) and SOST/sclerostin, that has allowed, for
the first time a closer look at the biology of these cells. Osteo-
cytes, the cells deeply entrapped into the mineralized matrix,
have emerged as key regulators not only of skeletal and min-
eral homeostasis, but also hematopoiesis. This review will
summarize novel findings in osteocyte biology and future ave-
nues of research. Recent Progress in Osteocyte Research Paola Divieti Pajevic Endocrine Unit, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA The last decade has seen an exponential increase in our understanding of osteocytes function and biology. These cells, once con-
sidered inert by-standers trapped into the mineralized bone, has now risen to be key regulators of skeletal metabolism, mineral
homeostasis, and hematopoiesis. As tools and techniques to study osteocytes improved and expanded, it has become evident that
there is more to these cells than initially thought. Osteocytes are now recognized not only as the key responders to mechanical
forces but also as orchestrators of bone remodeling and mineral homeostasis. These cells are the primary source of several impor-
tant proteins, such as sclerostin and fibroblast growth factor 23, that are currently target as novel therapies for bone loss (as the
case for antisclerostin antibodies) or phosphate disorders. Better understanding of the intricate cellular and molecular mecha-
nisms that govern osteocyte biology will open new avenue of research and ultimately indentify novel therapeutics to treat bone
and mineral disorders. This review summarizes novel findings and discusses future avenues of research. Keywords: Osteocytes; Sclerostin; Mineral homeostasis; Bone homeostasis Corresponding author: Paola Divieti Pajevic
Endocrine Unit, Massachusetts General Hospital and Harvard Medical School,
50 Blossom Street, Boston, MA 02114, USA
Tel: +1-617-726-6184, Fax: +1-617-726-7543, E-mail: divieti@helix.mgh.
harvard.edu Review
Article Endocrinol Metab 2013;28:255-261
http://dx.doi.org/10.3803/EnM.2013.28.4.255
pISSN 2093-596X · eISSN 2093-5978 Endocrinol Metab 2013;28:255-261
http://dx.doi.org/10.3803/EnM.2013.28.4.255
pISSN 2093-596X · eISSN 2093-5978 Share Your Story The Harvard community has made this article openly available. Please share how this access benefits you. Submit a story . Accessibility Keywords: Osteocytes; Sclerostin; Mineral homeostasis; Bone homeostasis THE OSTEOCYTES Primary cilia have been described as important mechanosen-
sor and mice lacking Kif3a or ITF88 (polaris), both key com-
ponents of the cilia, in bone cells, display skeletal abnormali-
ties. Targeted ablation of Kif3a from osteoblasts, and conse-
quently from osteocytes, using the Col1α 2.3 promoter, affects
skeletal anabolic responses to cyclic axial compression, sug-
gesting that primary cilia are needed for proper skeletal mech-
anosensory [14,15]. Similarly, mice with in which PDK1 is
ablated in osteocytes, using the 10kb-DMP1 promoter have
impaired anabolic response to mechanical loading, further
supporting the importance of cilia in skeletal mechanosensa-
tion [16,17]. Marotti et al. [7] and Palumbo et al. [8,9] have extensively
described the morphological changes that accompany the
transformation from a motile osteoblast into an entombed os-
teocyte. They demonstrate, by histological analysis of new-
born rabbit bones, that the formation of osteocyte cytoplasmic
processes is asynchronous and asymmetrical and precede the
mineralization of the organic matrix. Moreover, osteocytes are
evolutionary highly conserved and the organized structure of
these cells within a mineralized matrix is present in bone spec-
imens from Tyrannosaurus rex, dating back more than 80 mil-
lion of years ago, clearly indicating an important role for these
cells in skeletal metabolism [10]. Osteocytes communicate with each other and with cells at
the endosteal and periosteal surface through an extensive and
intricate system of canaliculi. These cells are also in close
proximity of capillary and vessels, raising the hypothesis that
osteocytes might function as an endocrine organ and directly
secrete proteins, such as FGF-23, Phex, or sclerostin into the
circulation [11]. Moreover, the osteocytic network, with its
extensive system, is an ideal structure to sense mechanical
loading and control mineral homeostasis. It has been postulat-
ed that osteocytes can send signals for both bone resorption
and formation and thus orchestrate a proper cycle of remodel-
ing, as discussed in details below. Osteocytes responses to load, both in vivo and in vitro, include
secretion of prostaglandin E2 and nitric oxide and the expression
of several mechanosensitive transcripts such as DMP1, MEPE,
and Phex [18,19]. Another osteocytic gene highly regulated by
mechanical forces is SOST. SOST/sclerostin in suppressed dur-
ing load whereas it is increased during hindlimb unloading [20]. Moreover mice lacking SOST are resistant to disuse induced
bone loss and antisclerostin antibodies can prevent bone loss as-
sociated with hindlimb unloading [21]. THE OSTEOCYTES Osteocytes are postmitotic, terminally differentiated osteo- Copyright © 2013 Korean Endocrine Society Corresponding author: Paola Divieti Pajevic This is an Open Access article distributed under the terms of the Creative Com
mons Attribution Non-Commercial License (http://creativecommons.org/
licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribu
tion, and reproduction in any medium, provided the original work is properly
cited. 255 255 www.e-enm.org Pajevic PD Pajevic PD ity to the endosteal (and possibly periostal) surface and the ex-
pression of transcripts such as E11/gp38/podoplain [4], matrix
extracellular phosphoglycoprotein (MEPE) and phosphate-
regulating gene with homologies to endopeptidases on the X
chromosome (Phex). This cell, opposite to the more mature
osteocyte, is negative for sclerostin and fibroblast growth fac-
tor (FGF)-23 expression but does express DMP1, as elegantly
demonstrated by Kalajzic et al. [5] in a recent review. It is not
clear, however, if osteoid, mineralizing and mature osteocytes
exert different biological functions or if they all work as
mechanosensor and skeletal regulators. Indeed recent studies
suggest that cortical and trabecular osteocytes might have dis-
tinct roles with the former being the sensor of load while the
latter being the controller of bone metabolism. Windahl et al. [6], for example, reported that estrogen receptor-α ablation in
osteocytes differentially affects trabecular and cortical bone
compartments in males mice. for over 30 years that osteocytes are mechanosensor of bone. They are indeed the cells capable of sensing mechanical forces
applied to the skeleton and transform these forces into biologi-
cal stimuli. It is now widely accepted that the cell perceives
forces in the form of shear stress created by the flow of fluid
inside the lacuna-canalicular network. It has been proposed
that the flow derives from both the compression induced by
loading and the extravascular pressure. Efforts to physically
measure and quantify this flow are currently ongoing and re-
cent reports suggest that osteocytes are subjected to forces in
the order of 5 Pa [12]. Although the theory that osteocytes are
the mechanosensor of bone has been around for decades, the
definite demonstration of this theory comes from work of Tat-
sumi et al. [13]. Using a mouse model of targeted osteocytes
ablation, they demonstrated resistance to disuse induces bone
loss highlighting the central role that osteocytes play in mech-
anosensation. How an osteocyte can sense the fluid flow in the
lacuno-canalicular network is still not completely understood. THE OSTEOCYTES This demonstrates that
osteocytes are key orchestrators not only of skeletal responses to
loading but also to unloading, as occurs during paralysis, pro-
longed bed rest or space flight. Frost [22] postulated that osteo-
cytes are the mechanostat of bone, ascribing to these cells the
role of driving skeletal adaptation to mechanical forces. Bone
adapts its form to mechanical demands trough a mechanisms
known as bone modeling. During modeling, bone resorption, OSTEOCYTE FUNCTIONS:
MECHANOTRANSDUCTION What are the functions of an osteocyte? It has been recognized Copyright © 2013 Korean Endocrine Society www.e-enm.org Recent Progress in Osteocytes driven by osteoclasts, and bone formation, osteoblasts-mediated,
occurs on different surfaces of the skeleton such as the bone un-
dergoes reshaping to adapt to different loading conditions. Bone
remodeling, on the other hand, is the continuous and spatially
coupled, resorption and formation of the skeleton to preserve
functional integrity. According to Frost [22], the mechanostat
distinguishes between bone modeling (shape change) and re-
modeling (replacement only). Examples of bone adaptation to
loading are evident in professional athletes, where the high load
applied to the skeleton results in stronger and bigger bones,
whereas in astronauts or in paralyzed patients there is severe
bone loss due to unloading. As described above, compelling evi-
dences ascribe to sclerostin an important role in this disuse in-
duced bone loss. duced periosteal circumference and have a blunted response to
mechanical forces. When IGF-1 conditional KO mice were
subjected to mechanical loading, there was a significant reduc-
tion in osteoblastic bone formation response, indicating that
IGF-1 is an important determinant of both bone size and bone
strength [30]. Pioneering studies on osteocytes involving histological
analysis of bone specimens raised the possibility that osteo-
cytes could reabsorb bone and directly contribute to mineral-
ion homeostasis. The finding that in various diseases (renal
osteodystrophy, hyperparathyroidism, and immobilization) the
size of the osteocytic lacunae was increased led to the hypoth-
esis that these cells were indeed capable of modifying the
perilacunar mineralized matrix. This theory was quickly aban-
doned when isolated osteocytes failed to reabsorb bone is a
classical osteoclast pit-forming assay and few decades later,
the concept that osteocytes can indeed reabsorb their perilacu-
nal matrix has been revisited. The current osteocytic osteolysis
concept, as proposed by Cullinane [31], suggests that osteo-
cytes do not remove substantial amount of bone (as osteo-
clast), but rather modify the matrix minerals in the perilacunar
areas. This perilacunal remodeling is evident during continu-
ous infusion with PTH [32], treatment with prednisone [33] or
during lactation [34]. As described above, osteocytes express
receptor for PTH, a known regulator of calcium homeostasis,
and they are the ideal candidates for systemic homeostasis
regulation. Mice lacking the PTH1R specifically in osteocytes
have indeed an impaired calcium homeostasis when subjected
to a low calcium diet, indicating these cells also control calci-
um homeostasis [35]. OSTEOCYTE FUNCTIONS:
MECHANOTRANSDUCTION Lastly, when the vitamin D receptor is
ablated from osteocytes, the animals are unable to mobilize
calcium from bone under a vitamin D stimulus, indicating that
vitamin D signaling in osteocytes is critical for moving calci-
um from skeletal stores to the blood [36]. In light to recent re-
port indicating that osteocytes are the major source of RANKL
and therefore bone remodeling [26,27], we can speculate that
the calcemic effect of vitamin D on bone is via osteocyte-de-
rived RANKL although evidences for this pathway are still
missing. BONE AND MINERAL HOMEOSTASIS Osteocytes are not only the sensor of mechanical forces, they
are also master controller of bone and mineral homeostasis. These cells are, postnatally, the main source sclerostin and re-
ceptor activation of nuclear factor-κB ligand (RANKL). Sclerostin, the product of the gene SOST, has emerged as a
powerful inhibitor of bone formation. The protein binds to
LRP5 and the related LRP6 and four receptors and block the
canonical Wnt-ßcatenin signaling pathway. Lack (or mutation)
of sclerostin induces sclerosteosis whereas its over expression
causes severe osteopenia (for review on Wnts) [23]. Osteo-
cytes are also the main source of FGF-23, a key regulator, to-
gether with parathyroid hormone (PTH) of phosphate homeo-
stasis [24,25]. Thus, it is not surprising that genetic manipula-
tion of osteocytes induced both skeletal and mineral defects. Mice lacking RANKL in osteocytes have increased bone min-
eral density and reduced bone remodeling [26,27], as do mice
lacking the receptor of PTH (PTH1R). Mice lacking Atg7, an
autophagy gene, from osteocytes, display similar skeletal phe-
notype as aging mice, indicating that these cells are key regu-
lator of skeletal metabolism [28]. Both DMP1-null mice and
individuals with inactivating mutation of this gene have rickets
and osteomalacia and are hypophostatemic as a consequence of
high levels of FGF-23 [29]. On the contrary, mice lacking
FGF-23 are hyperphosphatemic, osteopenic, and dye prema-
turely. It has been reported that osteocytes are also the major
source of insulin-like growth factor (IGF)-1 and IGF-1 pro-
duced by these cells is required for proper anabolic responses
to mechanical loading. Deletion of IGF-1 in osteocytes, (using
the DMP1-Cre promoter) impairs skeletal growth, have re- www.e-enm.org 2 www.e-enm.org 257 HEMATOPOIESIS Hematopoiesis, the process that continuously gives rise blood
cells, is tightly regulated not only by intrinsic factors but also
by extrinsic clues derived from various cells within the bone
marrow microenvironment. The bone marrow is indeed the pri- Copyright © 2013 Korean Endocrine Society www.e-enm.org 257 www.e-enm.org 257 Pajevic PD Pajevic PD mary site for hematopoiesis that is highly orchestrated by inter-
actions between the hematopoietic stem cells (HSCs) and their
niches on the endosteal bone surface. The HSC niche consists
of cells from the osteolineage, sinusoidal endothelial cells,
mesenchymal stromal and stem cells, sympathetic neurons, and
the extracellular matrix [37,38]. In these niches, the HSC un-
dergo self-renewal and/or differentiate into hematopoietic lin-
eage with cues from the niche-supporting cells, which provide
structural as well as signaling support for the hematopoietic
microenvironment. Osteoblasts have been shown as critical
regulators of HSC maintenance, proliferation, and maturation
[39,40] via various cell-surface proteins such as Notch1 ligand
Jagged1 and via secreted cytokines such as CXCL12. Osteo-
blasts also provide a niche for B-lymphopoiesis in part via in-
terleukin-7 expression through Gsα-signaling [41]. Hypoxic
osteoblasts are also a major producer of erythropoietin through
which they regulate erythropoiesis [42]. Moreover, osteoclasts
has been shown to be potential HSC regulators through degra-
dation of endosteal components and promotion of mobilization
of HSCs [43]. In contrast, the role of osteocytes in HSC and
progenitor regulation, and hence hematopoiesis, has remained
completely unexplored despite the fact that these cells com-
prise 90% to 95% of all bone cells in the adult skeleton and
they produce an extensive canalicular network that reaches the
endosteal bone surface, a preferred site for the HSC niche. Os-
teoblasts and osteocytes express several G-protein coupled re-
ceptor (GPCR) and signaling trough these receptor s has been
shown to control the niche. Interestingly constitutive activation
of the PTH/PTHrP receptor in osteoblasts increased the num-
bers of HSCs [39] whereas its expression in osteocytes does
not [44], suggesting that cells early in the osteoblast lineage are
critical for maintaining an intact HSC niche. To examine the
role of osteocytes and GPCR signaling in regulation of hema-
topoiesis we recently generated mice lacking Gsα in osteocytes
(Ocy-GsαKO). Surprisingly, ablation of Gsα from these cells in-
duces a marked neutrophilia, thrombocytopenia and spleno-
megaly. CONFLICTS OF INTEREST No potential conflict of interest relevant to this article was re-
ported. FUTURE DIRECTIONS It is evident that there is a need for further investigations
aimed at better understand osteocytes function. Several ques-
tion remain unanswered and areas in need of better under-
standing are numerous. For example, which are the molecular
cues that signal an osteoblast to become an osteocyte? How
can osteocytes sense mechanical forces applied to the skeleton
and what is the role of primary cilia? What control the in-
creased in SOST/sclerostin induced by unloading? How can
an osteocyte sense phosphate? What is the role of osteocytes
in hematopoiesis? These are many more other unanswered
questions will require better tools and in vitro models to study
and analyze these cells and advance our understanding of their
complex functions even further. HEMATOPOIESIS Ocy-GsαKO mice displayed a profound myeloprolifera-
tive phenotype characterized by a dramatic increase in myeloid
cells in bone marrow, spleen, and peripheral blood and marked
[45]. Using a novel ex vivo coculture system using osteocyte-
enriched bone explants we identified granulocyte colony-stim-
ulating factor as the principal cytokine regulating granulopoie-
sis in these mice Interestingly PTH1R signaling in osteocytes SOST/sclerostin, a Wnt inhibitor and suppressor of osteoblast
proliferation and functions, was increased significantly in these
mice. Treatment with antisclerostin antibody partially restored
the number of osteoblasts in Ocy-GsαKO mice without rescuing
the bone marrow abnormalities demonstrating that the hemato-
poietic abnormalities present in these mice are independent on
osteoblasts or increased SOST/sclerostin expression [45]. On
the other hand, SOST/sclerostin can directly control the fate of
B-cells, as recently reported [46]. Cain et al. [46] showed that
SOST-/- mice, despite the high bone mass and increased num-
ber of osteoblasts, have no differences in the frequency or ab-
solute number of HSCs, common lymphoid progenitors, com-
mon myeloid/megakaryocyte erythroid progenitors, or granu-
locyte/monocyte progenitors, confirming the findings in mice
overexpressing the constitutive PTH/PTHrP receptor in osteo-
cytes [44]. Interestingly, in these mice, B cells are significantly
reduced in both their frequency and cell number in the bone
marrow and the reduction was a consequence of increased
apoptosis due to a reduction in Cxcl12 expression in the stro-
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Outcomes in patients with lacrimal gland carcinoma treated with definitive radiotherapy or eye-sparing surgery followed by adjuvant radiotherapy
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RESEARCH Open Access Abstract Background: The optimal treatment for lacrimal gland cancer remains unclear. Eye-preserving surgery, as opposed
to exenteration, followed by adjuvant radiotherapy (RT), has recently been reported to deliver satisfactory outcomes,
but evidence is sparse. The aim of the present study was to evaluate outcomes in patients with lacrimal gland
cancer treated at two tertiary medical centers. Methods: We retrospectively examined data from patients with lacrimal gland cancer who had received eye-
preserving surgical treatment followed by adjuvant RT with or without chemotherapy, or (if the tumor was
inoperable) needle biopsy with definitive RT with or without chemotherapy. Baseline clinical and pathological
characteristics were considered. Outcomes of interest included post-treatment complications, overall survival (OS),
locoregional progression-free survival (LPFS), and distant metastasis-free survival (DMFS). Results: Eighteen patients were included. Two-year OS, LPFS, and DMFS rates were 69.0, 76.7, and 71.4%,
respectively. Patients with early-stage (T1–T2) lacrimal gland cancer had significantly better outcomes than those
with advanced-stage disease (T3–T4). Two-year OS, LPFS, and DMFS rates were each 100% in patients with disease
stages T1–T2, and 37.5, 50, and 37.5%, respectively, in those with disease stages T3–T4 (P < 0.05). Orbital
complications were well tolerated. Conclusions: Eye-sparing surgery with adjuvant RT can achieve satisfactory results in patients with T1–T2 lacrimal
gland carcinoma. Disease stage T3 and above was associated with poor outcomes even with post-operative RT,
likely due to distant metastasis. Adding neoadjuvant chemotherapy or adjuvant chemotherapy to current treatment
strategies might be a suitable choice for this group of patients. Keywords: Lacrimal gland carcinoma, Orbital tumors, Radiotherapy, Orbital exenteration * Correspondence: ocean@ntut.edu.tw
†Yun-Hsuan Lin and Shih-Ming Huang contributed equally to this work. 1Department of Electro-Optical Engineering, National Taipei University of
Technology, 1, Sec. 3, Chung-Hsiao E. Rd, Taipei 10608, Taiwan, ROC
Full list of author information is available at the end of the article © The Author(s). 2020 Open Access This article is licensed under a Creative Commons Attribution 4.0 International License,
which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give
appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if
changes were made. The images or other third party material in this article are included in the article's Creative Commons
licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons
licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain
permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the
data made available in this article, unless otherwise stated in a credit line to the data. RESEARCH
Open Access
Outcomes in patients with lacrimal gland
carcinoma treated with definitive
radiotherapy or eye-sparing surgery
followed by adjuvant radiotherapy
Yun-Hsuan Lin1,2,3†, Shih-Ming Huang3,4,5†, Wing-Keen Yap5, Ju-Wen Yang2,3, Ling Yeung2,3, Din-Li Tsan4,
Joseph Tung-Chieh Chang5 and Lung-Chien Chen1*
Lin et al. Radiation Oncology (2020) 15:156
https://doi.org/10.1186/s13014-020-01601-8 RESEARCH
Open Access
Outcomes in patients with lacrimal gland
carcinoma treated with definitive
radiotherapy or eye-sparing surgery
followed by adjuvant radiotherapy
Yun-Hsuan Lin1,2,3†, Shih-Ming Huang3,4,5†, Wing-Keen Yap5, Ju-Wen Yang2,3, Ling Yeung2,3, Din-Li Tsan4,
Joseph Tung-Chieh Chang5 and Lung-Chien Chen1*
Lin et al. Radiation Oncology (2020) 15:156
https://doi.org/10.1186/s13014-020-01601-8 Lin et al. Radiation Oncology (2020) 15:156
https://doi.org/10.1186/s13014-020-01601-8 Outcomes in patients with lacrimal gland
carcinoma treated with definitive
radiotherapy or eye-sparing surgery
followed by adjuvant radiotherapy Yun-Hsuan Lin1,2,3†, Shih-Ming Huang3,4,5†, Wing-Keen Yap5, Ju-Wen Yang2,3, Ling Yeung2,3, Din-Li Tsan4,
Joseph Tung-Chieh Chang5 and Lung-Chien Chen1* © The Author(s). 2020 Open Access This article is licensed under a Creative Commons Attribution 4.0 International License,
which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give
appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if
changes were made. The images or other third party material in this article are included in the article's Creative Commons
licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons
licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain
permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the
data made available in this article, unless otherwise stated in a credit line to the data. Page 2 of 10 Page 2 of 10 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Radiation Oncology Precis Eye-sparing surgery with adjuvant radiotherapy can
achieve satisfactory results in patients with T1–2 lacri-
mal gland carcinoma. Disease stage T3 and above was
associated with poor outcomes even with post-operative
RT, likely due to distant metastasis. This retrospective study included patients with lacrimal
gland cancer who underwent either eye-preserving surgi-
cal treatment followed by adjuvant RT with or without
chemotherapy, or (in cases involving an inoperable
tumor), needle biopsy followed by definitive RT with or
without chemotherapy. Patients diagnosed with recur-
rent tumors at their first visit, and patients diagnosed
with lacrimal duct cancer or other type of orbital cancer,
were excluded. Disease stage was reviewed by a radiolo-
gist and ophthalmologist based on the American Joint
Committee on Cancer (AJCC) staging system, eighth
edition. All patient records included detailed patho-
logical reports with data on cell type, margin status,
perineural invasion (PNI), and lymphovascular invasion
(LVSI), if the tumor had undergone resection. Treatment Radiotherapy was recommended for patients at high risk
of recurrence, including those with an advanced stage
lacrimal
gland
cancer
and/or
multiple
unfavorable
pathological factors such as bony structure invasion,
positive surgical margins, LVSI, or PNI. RT was deliv-
ered in the form of three-dimensional conformal RT,
intensity modulated RT, volumetric modulated arc ther-
apy, or proton beam therapy. The RT field included the
tumor bed and the residual gross tumor volume. The
median prescribed dose for adjuvant RT was 50–60 Gy
and 66–70 Gy for the definitive setting, with 1.8–2 Gy
daily fractions administered over 5–7 weeks. Whether or
not chemotherapy was given was at the attending physi-
cian’s discretion. The regimens used were platinum-
based chemotherapy. Lacrimal gland carcinomas are associated with poor
local control and significant morbidity and mortality
rates [9]. To date, no guidelines on standard treatment
for lacrimal gland carcinomas have been developed due
to the rarity of these diseases, which are conventionally
treated with orbital exenteration followed by radio-
therapy (RT) [5, 10]. However, locoregional recur-
rence, distant relapse, and cancer-related mortality
risks are high after orbital exenteration [5, 11, 12]. Thus, eye-preserving surgery followed by adjuvant RT
has recently gained popularity, but the optimal ap-
proaches to surgery and RT in these patients remain
subject to debate [5, 9, 13, 14]. Statistical analyses
ll
l ( In this study, we examined clinical outcomes associ-
ated with either eye-preserving surgery followed by
adjuvant RT, or needle biopsy followed by definitive
RT, in a cohort of 18 patients with lacrimal gland
carcinoma. Clinical outcomes such as local control,
survival, and ocular complications were considered. The clinical and pathological factors that affected the
outcomes were also examined. Overall survival (OS) was defined as the time from the
date of surgical treatment or biopsy until death or the
last follow-up appointment. Local progression-free sur-
vival (LPFS) was defined as the time between the date of
surgical treatment or biopsy and the detection of local
tumor progression. Data on the first site(s) of distant
metastasis were collected, and distant metastasis-free
survival (DMFS) was determined. The Kaplan–Meier
method was used to estimate the OS, LPFS, and DMFS
rates, while the statistical significance of between-group
differences in clinical characteristics (including sex, age,
RT dose, cell types, and pathological features) and OS,
LPFS, DMFS was determined using the log-rank test. All
statistical analyses were performed using SPSS v. 23.0
(IBM Corp., New York, NY; formerly SPSS Inc., Chicago,
IL). All P-values were two-sided, and a P-value < 0.05
was considered statistically significant. Introduction Lacrimal gland tumors are among the rarest types of
head and neck cancers, accounting for approximately
10% of all orbital tumors [1]. Previous studies have esti-
mated annual incidence for these tumors at 0.19–1 per
1,000,000 people [1–3]. Epithelial lesions, including be-
nign pleomorphic adenoma and malignant tumors, ac-
count
for
20–30%
of
all
lacrimal
gland
tumors. Malignant epithelial tumors include adenoid cystic car-
cinoma (ACC) (60%), carcinoma ex pleomorphic aden-
oma or adenocarcinoma (20–30%), and mucoepidermoid
carcinoma (5%) [1, 4, 5]. In contrast, non-epithelial le-
sions, such as inflammation, lymphoid tumors, plasma-
cytoma, histiocytoma, lipoma, and hemangioma account
for 70–80% of all lacrimal gland tumors [2, 4, 6–8]. Survival analysis For the entire cohort, 2-year OS, LPFS, and DMFS rates
were 69.0, 76.7, and 71.4%, respectively (Fig. 2). Asso-
ciations between clinicopathological factors and 2-year
OS, LPFS, and DMFS rates are shown in Table 3. Pa-
tients
with
early-stage
(T1–T2)
lacrimal
cancer
achieved significantly better 2-year OS, LPFS, and
DMFS than those with advanced-stage disease (T3–
T4). Two-year OS was 100% in patients with disease
stage T1–T2, and 37.5% in those with disease stage
T3–T4. Moreover, 2-year LPFS was 100% in patients
with disease stage T1–T2 and 50% in those with dis-
ease stage T3–T4. Finally, 2-year DMFS was 100% in
patients with disease stage T1–T2 and 37.5% in those
with disease stage T3–T4 (Fig. 3). Other factors, in-
cluding sex, age (< 54 yrs. vs. ≥54 yrs), RT dose (< 66
vs. ≥66 Gy), cell types (ACC vs. others), margin (posi-
tive vs. negative), PNI (positive vs. negative), and
LVSI (positive vs. negative) had no significant impact
on OS, LPFS, or DMFS. Materials and methods This study protocol was reviewed and approved by
the institutional review board of Chang Gung Me-
morial Hospital (Reference No.: 202000138B0). All
data were stored in the hospital database and ex-
tracted for research. The participants’ informed con-
sent requirement was waived due to the retrospective
nature of this study. Page 3 of 10 Page 3 of 10 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Radiation Oncology Post-therapy surveillance RT was 6000 cGy (range 3600–7000 cGy) in 1.8–2 Gy
per fraction, and the definitive RT dose was 6000
cGy/25 fractions, 7200 cGy/36 fractions, and 7200
cGy/36 fractions for three patients, respectively. Follow-up in the form of radio–oncologic and ophthal-
mic clinic appointments was arranged every 3 months
during the first 2 years, every 4–6 months during the
third and fourth year, and every 6–12 months thereafter. Imaging examination was performed at specific intervals:
chest radiography every 3 months, and computed tom-
ography or orbital magnetic resonance imaging every 3–
6 months, until symptoms indicating recurrence were
noted. Ocular complication evaluation Follow-up in the form of ophthalmic clinic appoint-
ments was arranged every 3 months during the first 2
years and every 6–12 months thereafter. Detailed ocular
examination, including visual acuity, intraocular pres-
sure, slit lamp examination, and fundus examination was
performed at every ophthalmologic visit. Medical or sur-
gical treatment was administered if an ocular complica-
tion was noted. Patient characteristics During 2000–2018, a total of 18 patients from the Lin-
kou and Keelung branches of Chang Gung Memorial
Hospital were included in the present study. The median
follow-up time was 39 months (range 5–161 months) for
the patients who were alive at the end of the study
period, and 26 months (range 5–161 months) for the
whole cohort. The patients’ demographic and clinical
characteristics are shown in Table 1 and Table 2. Fifteen
patients (83.3%) underwent eye-sparing surgery and the
remaining patients had needle biopsy alone due to inop-
erable tumor invading brain tissue or extending to the
infratemporal fossa. ACC was found in nine, carcinoma
ex pleomorphic adenoma in seven, and poorly differenti-
ated adenocarcinoma in two patients. In five patients
(27.8%), negative margins were achieved during surgery. Six patients (33.3%) had pathologically confirmed LVSI
and eight patients (44.4%) had pathologically confirmed
PNI. Patients who underwent eye-sparing surgery also
received adjuvant RT or concurrent platinum-based che-
moradiotherapy (CCRT) due to the presence of risk fac-
tors, such as positive surgical margin, confirmed LVSI or
PNI, or initial advanced tumor size and invasion status. Three patients who underwent needle biopsy received
definitive RT or CCRT, also platinum-based. Ocular toxicity analysis In our study subjects, one patient was lost follow up at
the ophthalmic clinic 3 months after surgery and three
patients 6 months after surgery. During their follow-up
period, no ocular complications were reported. In the
remaining 14 patients who were followed up by the oph-
thalmologist for more than 12 months, the administered
treatment was well tolerated. Treatment-related acute
and late ocular toxicity effects are shown in Table 4. Ocular toxicities, including punctate keratitis, devel-
oped in five patients (27.78%) 1.14–7.9 months after
radiotherapy (RT total/fraction dose: 5000–7000/200
cGy). Cataract developed in four patients (22.22%)
and occurred 2.13–112.7 months after the RT (RT
total/fraction dose: 6000–6600/200 cGy). Dry eye de-
veloped in two patients (11.11%) and occurred 6.29–
11.05 months after the RT (RT total/fraction dose:
6600/200 cGy). Blepharitits, which developed in two
patients (11.11%), occurred 2.06–8.00 months after the
treatment
(RT
total/fraction
dose:
6600/200 cGy). Other complications, including trichiasis, acute con-
junctivitis, corneal epithelial defect, filamentary kera-
titis, radiation retinopathy, and vitreous hemorrhage
were each reported in one patient (5.56%). The pa-
tient with radiation retinopathy received an intravitre-
ous injection of 1.25 mg Avastin (bevacizumab) for
retinal neovascularization. Regarding RT technique, 2 patients received three-
dimensional conformal radiotherapy (3DCRT), 14 re-
ceived intensity-modulated radiotherapy (IMRT), one
received proton beam therapy alone, and one received
combined proton–photon RT. The dosimetry com-
parison between photon plan and proton plan is pre-
sented in Fig. 1. The median RT dose for adjuvant Page 4 of 10 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Ocular toxicity analysis Radiation Oncology Table 1 Baseline characteristics of patients with lacrimal gland carcinoma
Total (n = 18)
Patient number
Percent
Median age (years)
54 (20–85)
Sex
Male
8
44.4
Female
10
55.6
T stage
1
1
5.6
2
9
50.0
3
3
16.7
4
5
27.7
Types of surgery
Excision
15
83.3
Needle biopsy
3
16.7
RT treatment
Adjuvant RT alone
13
72.2
Adjuvant CCRT
2
11.1
Definitive RT
2
11.1
Definitive CCRT
1
5.6
RT type
3DCRT
2
11.1
IMRT
14
77.8
Proton
2
11.1
RT dose
< 6600 cGy
9
50.0
≥6600 cGy
9
50.0
Cell type
Adenoid cystic carcinoma
9
50.0
Carcinoma ex pleomorphic adenoma
7
38.9
Poorly differentiated adenocarcinoma
2
11.1
Margin
positive
13
72.2
negative
5
27.8
LVSI
Positive
6
33.3
Negative
12
66.7
PNI
Positive
8
44.4
Negative
10
55.6
RT radiotherapy, CCRT concurrent chemoradiotherapy, 3DCRT 3-dimensional conformal radiotherapy, IMRT intensity-modulated radiotherapy, cGy centigray,
lymphovascular invasion, PNI perineural invasion. ncurrent chemoradiotherapy, 3DCRT 3-dimensional conformal radiotherapy, IMRT intensity-modulated radiotherapy, cGy centigray, LVSI
PNI perineural invasion. RT radiotherapy, CCRT concurrent chemoradiotherapy, 3DCRT 3-dimensional conformal radiotherapy, IMRT intensity-modulated radiotherapy, cGy centigray, LVSI
lymphovascular invasion, PNI perineural invasion. Discussion treatment followed by adjuvant radiotherapy with or with-
out chemotherapy. In the present cohort, 2-year OS, LPFS,
and DMFS rates were 69.0, 76.7, and 71.4%, respectively. These outcomes were inferior to those reported in previ-
ously, which was probably due to differences in patient This study reported outcomes in patients with lacrimal
gland carcinoma who underwent either needle biopsy
followed by definitive radiotherapy with or without chemo-
therapy due to inoperable tumor, or eye-preserving surgical Page 5 of 10 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Discussion Page 6 of 10 (2020) 15:156 Lin et al. Radiation Oncology (2020) 15:156 Lin et al. Radiation Oncology Fig. 1 Patient number 7, cT2bN0M0 lacrimal gland cancer, pre-surgery T1-MRI and the photon or proton beam dosimetry of adjuvant
radiotherapy. Total RT dose was 70 Gy in 33 fractions. Dose constraints for organs at risk: Brain stem: Dmax < 54 Gy. Left eye and left optic nerve:
Dmax ≤59.5Gy (85% dose). Right eye and right optic nerve: Dmax ≤50 Gy. Optic chiasm: Dmax < 54 Gy. Right len: Dmax ≤18Gy. No ocular
complication was noted during our study period. a. Axial view of the tumor. b. Sagittal view of the tumor. c. Photon beam dosimetry. 1d. Proton
beam dosimetry Fig. 1 Patient number 7, cT2bN0M0 lacrimal gland cancer, pre-surgery T1-MRI and the photon or proton beam dosimetry of adjuvant
radiotherapy. Total RT dose was 70 Gy in 33 fractions. Dose constraints for organs at risk: Brain stem: Dmax < 54 Gy. Left eye and left optic nerve:
Dmax ≤59.5Gy (85% dose). Right eye and right optic nerve: Dmax ≤50 Gy. Optic chiasm: Dmax < 54 Gy. Right len: Dmax ≤18Gy. No ocular
complication was noted during our study period. a. Axial view of the tumor. b. Sagittal view of the tumor. c. Photon beam dosimetry. 1d. Proton
beam dosimetry carcinoma (three patients with T3 [16.7%], and five
patients with T4 [27.7%]). In addition, the majority of
patients were at high risk of locoregional or distant
recurrence, with a positive surgical margin in 72.2%
of the patients, and confirmed LVSI and PNI in 33.3
and 44.4% of patients, respectively. The differences in
patient baseline characteristics may explain the rela-
tively lower OS, LPFS, and DMFS rates reported in
our study. characteristics. Esmaeli et al. [15] studied the eye-sparing
approach to treating lacrimal gland carcinoma in 11 pa-
tients during 2007–2014. All 11 patients were disease-free
at the last follow-up visit after the eye-sparing surgery, with
a median follow-up time of 33 months (range, 14–64
months). A significant proportion of patients recruited by
Esmaeli et al. had early-stage disease (one patient with T1,
six patients with T2, one with T3, and three with
T4). In addition, Han et al. Discussion Radiation Oncology Table 2 Detailed patient, tumor, treatment characteristics and outcomes
Patient Age Gender Cell
type
AJCC
stage
laterality Surgery
type
Margin PNI LVSI Adjuvant
treatment
Post
treatment
status
Time to local
progression
(months)
Time to distant
metastasis (months)
Metastasis site
Total follow up
time (months)
Final
status
1
45
Female
CEPA
cT1aN0M0
L
Excision
+
–
–
RT alone
CR
–
–
161
alive
2
20
Female
ACC
cT2aN0M0
L
excision
–
+
–
RT alone
CR
–
–
28
dead
3
58
Female
ACC
cT2aN0M0
L
excision
+
–
–
RT alone
CR
–
–
20
alive
4
55
Male
CEPA
cT2aN0M0
R
excision
+
–
+
RT alone
CR
–
–
39
alive
5
77
Male
CEPA
cT2aN0M0
R
excision
+
–
–
RT alone
CR
–
–
26
alive
6
53
Female
CEPA
cT2aN0M0
L
excision
–
+
+
RT alone
(proton)
CR
–
–
5
alive
7
45
Male
ACC
cT2bN0M0 L
excision
+
–
–
CCRT (proton
+ photon)
CR
–
–
14
alive
8
35
Male
ACC
cT2cN0M0
L
excision
+
+
+
RT alone
CR
–
–
158
alive
9
43
Female
ACC
cT2cN0M0
R
excision
+
+
–
RT alone
CR
–
26
mediastinal LNs,
liver, bone, lungs
26
dead
with
disease
10
53
Female
ACC
cT2cN0M0
R
excision
+
+
+
RT alone
CR
–
–
40
alive
11
85
Male
ACC
cT3cN0M0
L
needle
biopsy
–
–
–
RT alone
PR
10
34
lungs
43
alive with
disease
12
62
Male
CEPA
cT3cN0M0
L
excision
+
–
+
RT alone
CR
50
51
lungs
92
dead
with
disease
13
80
Male
CEPA
cT3cN0M0
R
needle
biopsy
–
–
–
RT alone
CR
–
–
17
dead
14
36
Female
ACC
cT4bN0M0 L
excision
+
+
–
RT alone
CR
10
6
liver
16
dead
with
disease
15
30
Female
ACC
cT4bN0M0 L
excision
+
+
–
RT alone
CR
5
10
bone and lungs
28
dead
with
disease
16
58
Female
CEPA
cT4bN0M0 R
excision
+
+
+
RT alone
CR
6
5
parotid, neck
nodes, lungs
12
dead
with
disease
17
60
Male
PD
adeno
cT4cN0M0
L
needle
biopsy
–
–
–
CCRT
PR
–
5
bone
20
dead
with
disease
18
71
Female
PD
adeno
cT4cN0M0
R
excision
+
–
–
CCRT
CR
–
5
liver and bone
15
dead
with
disease
AJCC the American Joint Committee on Cancer, PNI perineural invasion, LVSI lymphovascular invasion, ACC adenoid cystic carcinoma, CEPA carcinoma ex pleomorphic adenoma, PD adeno poorly
differentiated adenocarcinoma, L left, R right, RT radiotherapy, CCRT concurrent chemoradiotherapy, CR=, LN lymph node. Discussion [16] reported outcomes
following eye-sparing surgery and adjuvant radiother-
apy in 10 patients with adenoid cystic carcinoma of
the lacrimal gland, treated during 1998–2012 (one pa-
tient with T1, seven with T2, and two with T3 stage
disease), at a median follow-up time of 89.5 months
(range 37–217 months). Reported OS was 90%, with a
single
patient
who
died
58 months
after
surgery. Moreover, DMFS was 100%, while local recurrence
occurred in one patient. In our series, almost half of
the
patients
had
advanced-stage
lacrimal
gland The present study is among a small number that
have examined factors affecting outcomes in patients
with lacrimal gland carcinoma, which is an aim that
is difficult to achieve due to the rarity of this disease. In a previous report by Woo et al., the only tumor
characteristic associated with outcome was T stage. In
the same study, adjuvant radiotherapy was strongly
associated with a better 5-year recurrence-free sur-
vival rate [17]. Friedrich and Bleckmann also found
that
lower
T
stage
was
associated
with
a
better Lin et al. Radiation Oncology (2020) 15 Page 7 of 10 Lin et al. Radiation Oncology Fig. 2 The overall survival (OS), local progression-free survival (LPFS), and distant metastasis-free survival (DMFS) graphs of the whole cohort. For
the entire cohort, 2-year OS, LPFS, and DMFS rates were 69.0, 76.7, and 71.4%, respectively Fig. 2 The overall survival (OS), local progression-free survival (LPFS), and distant metastasis-free survival (DMFS) graphs of the whole cohort. For
the entire cohort, 2-year OS, LPFS, and DMFS rates were 69.0, 76.7, and 71.4%, respectively Fig. 2 The overall survival (OS), local progression-free survival (LPFS), and distant metastasis-
the entire cohort, 2-year OS, LPFS, and DMFS rates were 69.0, 76.7, and 71.4%, respectively outcomes even when treated with surgical resection
followed by adjuvant RT, which occurred most likely
due to distant metastasis. Adding neoadjuvant chemo-
therapy or adjuvant chemotherapy to current treat-
ment strategies might be a suitable choice for this
group of patients to improve the control of distant
disease. prognosis [18]. Among factors reported as negatively
affecting patient outcomes were older age, basaloid or
solid pattern ACC, perineural invasion, and the pres-
ence of a macroscopic tumor on imaging scans before
RT. [12, 19–22] In the present study, among patients
who had received adjuvant radiotherapy, T stage and
tumor size emerged as the most important factors
that determined patient outcome. LPFS local progression free survival, DMFS distant-metastasis free survival, RT radiotherapy, ACC adenoid cystic carcinoma, PNI perineu
ular invasion g
Y year, OS overall survival, LPFS local progression free survival, DMFS distant-metastasis free survival, RT radiotherapy, ACC adenoid cy
invasion, LVSI lymphovascular invasion Log rank test
Y year, OS overall survival, LPFS local progression free survival, DMFS distant-metastasis free survival, RT radiotherapy, ACC adenoid cystic carcinoma, PNI perineural
invasion, LVSI lymphovascular invasion Discussion Specifically, patients
with tumor at stage T1–2 treated with eye-sparing
surgery followed by adjuvant radiotherapy achieved
100% 2-year LPFS, DMFS, and OS rates. However,
tumor stage T3 and higher was associated with poor Proton therapy or even heavy ion therapy applied to
lacrimal gland cancer is considered promising for re-
ducing low dose delivered outside the treated fields
[3]. We have observed better dosimetry associated
with the proton plan compared to the photon plan. Table 3 Two-year overall survival, local progression-free survival, and distant-metastasis free survival rates for patients with lacrimal
gland carcinoma, including factors affecting survival
2-Y OS (%)
P-value*
2-Y LPFS (%)
P-value*
2-Y DMFS (%)
P-value*
Sex (male vs. female)
87.5 vs. 67.5
0.533
87.5 vs. 67.5
0.533
87.5 vs. 57.1
0.353
Age (< 54 vs. ≧54)
85.7 vs. 55.6
0.529
76.2 vs. 77.8
0.621
75.0 vs. 66.7
0.331
T stage (T1–2 vs. T3–4)
100.0 vs. 37.5
0.013
100.0 vs.50.0
0.006
100.0 vs. 37.5
0.002
RT dose (< 66 vs. ≧66 Gy)
87.5 vs. 50.8
0.114
63.5 vs. 88.9
0.464
75.0 vs. 66.7
0.344
Cell types (ACC vs. others)
87.5 vs. 50.0
0.517
66.7 vs. 87.5
0.581
77.8 vs. 66.7
0.914
Margin (positive vs. negative)
75.5 vs. 50.0
0.43
76.9 vs. 75.0
0.944
69.2 vs. 80.0
0.76
PNI (positive vs. negative)
66.7 vs. 71.4
0.438
58.3 vs. 90.0
0.264
58.3 vs. 80.0
0.583
LVSI (positive vs. negative)
80.0 vs. 75.0
0.879
80.0 vs. 75.0
0.879
83.3 vs. 66.7
0.292
*Log-rank test
Y year, OS overall survival, LPFS local progression free survival, DMFS distant-metastasis free survival, RT radiotherapy, ACC adenoid cystic carcinoma, PNI perineural
invasion, LVSI lymphovascular invasion survival, local progression-free survival, and distant-metastasis free survival rates for patients with lacrimal
ng factors affecting survival Lin et al. Radiation Oncology (2020) 15:156 Page 8 of 10 Lin et al. Radiation Oncology Fig. 3 Kaplan-Meier survival curves showing overall survival (a), local
progression free survival (b), and distant-metastasis free survival (c)
in patients with lacrimal gland carcinoma according to T category as
determined using the eighth edition of the American Joint
Committee on Cancer staging system. * Log-rank test. Discussion (a) The 2-year
OS rates were 100 and 37.5% in T1–2 and T3–4 groups (p = 0.013),
with (b) the 2-year DMFS rates were 100 and 37.5% (p = 0.002), and
(c) the 2-year LPFS rates of 100 and 50%, respectively (p = 0.006) At a high dose (> 70% dose), proton beam dosimetry
is equivalent to photon therapy; however, low-dose
distribution is better for proton therapy, whereby the
20% isodose line can spare the opposite eye. Ensuring
low dose spread to the nearby healthy tissue is critical
in certain cases. Lower dose delivered to the regional
organ at risk can reduce the likelihood of secondary
malignancy and long-term complications, and there-
fore should be considered for patients with early-stage
lacrimal gland cancer who are likely to achieve good
local
control
and
long-term
survival
after
the
treatment. Several ocular toxicities were recorded among our pa-
tients. These complications were expected as the radi-
ation doses used were mostly above the dose that is well
tolerated by ocular structures other than the sclera. Sen-
sitivity to radiation of the ocular structures has been
shown to vary [23–25]. The lens has been reported as
the most radiosensitive ocular structure, with a mini-
mum cataractogenic dose of 5.5 Gy. The lacrimal gland,
cornea, and conjunctiva can each tolerate up to 50 Gy of
radiation. The retina tolerates doses > 55 Gy. The sclera
is the most resistant ocular structure and is believed to
tolerate doses > 1000 Gy. The majority of ocular compli-
cations reported in the present study were effectively
managed by ophthalmologists. Only one patient received
a punctal plug due to intolerable dry eye and another
patient required intravitreous injection of Avastin (beva-
cizumab) at 1.25 mg due to radiation-associated retinop-
athy
with
retinal
neovascularization. Therefore,
we
recommend that all patients who have received RT for
lacrimal gland carcinoma visit an ophthalmologic clinic
regularly. This study has some limitations. First, this was a retro-
spective study based on medical records, which makes
selection bias inevitable. Moreover, some of the patients
did not attend the ophthalmologic clinic as recom-
mended, making it likely that the rate of ocular compli-
cations was underestimated. Second, due to the rarity of
lacrimal gland cancer, the small sample size meant that
the analysis was unlikely to yield statistically significant
results. Fig. 3 Kaplan-Meier survival curves showing overall survival (a), local
progression free survival (b), and distant-metastasis free survival (c)
in patients with lacrimal gland carcinoma according to T category as
determined using the eighth edition of the American Joint
Committee on Cancer staging system. * Log-rank test. (a) The 2-year
OS rates were 100 and 37.5% in T1–2 and T3–4 groups (p = 0.013),
with (b) the 2-year DMFS rates were 100 and 37.5% (p = 0.002), and
(c) the 2-year LPFS rates of 100 and 50%, respectively (p = 0.006) Acknowledgements
Not applicable. 3. Hayashi K, Koto M, Ikawa H, Ogawa K, Kamada T. Efficacy and safety of
carbon-ion radiotherapy for lacrimal gland carcinomas with extraorbital
extension: a retrospective cohort study. Oncotarget. 2018;9(16):12932–40. extension: a retrospective cohort study. Oncotarget. 2018;9(16):12932–40. 4. Ni C, Kuo PK, Dryja TP. Histopathological classification of 272 primary
epithelial tumors of the lacrimal gland. Chin Med J. 1992;105(6):481–5. 5. Wright JE, Rose GE, Garner A. Primary malignant neoplasms of the lacrimal
gland. Br J Ophthalmol. 1992;76(7):401–7. Ethics approval and consent to participate 10. Henderson JW, Neault RW. En bloc removal of intrinsic neoplasms of the
lacrimal gland. Am J Ophthalmol. 1976;82(6):905–9. This study protocol was reviewed and approved by the institutional review
board of Chang Gung Memorial Hospital (Reference No.: 202000138B0). This study protocol was reviewed and approved by the institutional review
board of Chang Gung Memorial Hospital (Reference No.: 202000138B0). 11. Esmaeli B, Golio D, Kies M, DeMonte F. Surgical management of locally
advanced adenoid cystic carcinoma of the lacrimal gland. Ophthalmic Plast
Reconstr Surg. 2006;22(5):366–70. Authors’ contributions
d y
4. Ni C, Kuo PK, Dryja TP. Histopathological classification of 272 primary
epithelial tumors of the lacrimal gland. Chin Med J. 1992;105(6):481–5. 4. Ni C, Kuo PK, Dryja TP. Histopathological classification of 272 primary
epithelial tumors of the lacrimal gland. Chin Med J. 1992;105(6):481–5. 5. Wright JE, Rose GE, Garner A. Primary malignant neoplasms of the lacrimal y
y
epithelial tumors of the lacrimal gland. Chin Med J. 1992;105(6):481–5. 5. Wright JE, Rose GE, Garner A. Primary malignant neoplasms of the lacrimal
gland. Br J Ophthalmol. 1992;76(7):401–7. SMH, YHL and LCC conceived the study; SMH and YHL collected data; SMH
analyzed and interpreted data; YHL, SMH and WKY wrote manuscript; LCC
provided statistical analyses; JWY, LY, DLT, and JTCC provided critical review
and revision of manuscript; All authors reviewed the results and approved
the final version of the manuscript. p
g
5. Wright JE, Rose GE, Garner A. Primary malignant neoplasms of the lacrimal
gland. Br J Ophthalmol. 1992;76(7):401–7. 6. Alkatan HM, Al-Harkan DH, Al-Mutlaq M, Maktabi A, Elkhamary SM. Epithelial
lacrimal gland tumors: a comprehensive clinicopathologic review of 26
lesions with radiologic correlation. Saudi J Ophthalmol. 2014;28(1):49–57. 6. Alkatan HM, Al-Harkan DH, Al-Mutlaq M, Maktabi A, Elkhamary SM. Epithelial
lacrimal gland tumors: a comprehensive clinicopathologic review of 26
lesions with radiologic correlation. Saudi J Ophthalmol. 2014;28(1):49–57. 7. Gao Y, Moonis G, Cunnane ME, Eisenberg RL. Lacrimal gland masses. AJR
Am J Roentgenol 2013 201(3) W371 81 lesions with radiologic correlation. Saudi J Ophthalmol. 2014;28(1):49 57. 7. Gao Y, Moonis G, Cunnane ME, Eisenberg RL. Lacrimal gland masses. AJR
Am J Roentgenol. 2013;201(3):W371–81. Funding 7. Gao Y, Moonis G, Cunnane ME, Eisenbe
Am J Roentgenol. 2013;201(3):W371–81 Funding
The authors state that this work has not received any funding. The authors state that this work has not received any funding. 8. Zeng J, Shi JT, Li B, Sun XL, An YZ, Li LQ, et al. Epithelial tumors of the
lacrimal gland in the Chinese: a clinicopathologic study of 298 patients. Graefes Arch Clin Exp Ophthalmol. 2010;248(9):1345–9. Discussion Finally, the patients receiving proton treatment
were only followed for 1–2 years, as our hospital began
proton beam therapy in 2017. A longer follow-up period Lin et al. Radiation Oncology (2020) 15:156 Page 9 of 10 Lin et al. Radiation Oncology Table 4 Frequency of ocular complications after radiotherapy reported in the current study and the corresponding radiotherapy
dose
Complications
Patient number
Percentage (%)
Time of occurrence (m)
RT total dose (cGy)
RT fraction dose (cGy)
Punctate keratitis
5
27.78
1.14–7.9
5000–7000
200
Cataract
4
22.22
2.13–112.7
6000–6600
200
Dry eye
2
11.11
6.49–11.05
6600
200
Blepharitis
2
11.11
2.06–8.00
6000
200
Trichiasis
1
5.56
16.36
6000
200
Acute conjunctivitis
1
5.56
0.68
5000
250
Corneal epithelial defect
1
5.56
23.38
7200
200
Filamentary keratitis
1
5.56
6.65
7000
200
Radiation retinopathy
1
5.56
74.39
6600
200
Vitreous hemorrhage
1
5.56
10.92
6840
180
RT radiotherapy, cGy centigray Table 4 Frequency of ocular complications after radiotherapy reported in the current study and the corresponding radiotherapy
d Table 4 Frequency of ocular complications after radiotherapy reported in the current study and the corresponding radiotherapy Keelung, Taiwan. 5Department of Radiation Oncology, Chang Gung Memorial
Hospital, Linkou, Taiwan. is required to evaluate the effects of proton therapy on
patient survival. is required to evaluate the effects of proton therapy on
patient survival. In conclusion, eye-sparing surgery with adjuvant RT
can achieve satisfactory results in patients with T1–2
lacrimal gland carcinoma. Disease stage T3 and above
was associated with poor outcomes even with post-
operative RT, probably due to distant metastasis. Adding
neoadjuvant or adjuvant chemotherapy to current treat-
ment strategies may be a suitable choice for this group
of patients. Received: 1 May 2020 Accepted: 16 June 2020 Received: 1 May 2020 Accepted: 16 June 2020 Consent for publication All data were stored in the hospital database and extracted for research. The
participants’ informed consent requirement was waived due to the
retrospective nature of this study. 12. Esmaeli B, Ahmadi MA, Youssef A, Diba R, Amato M, Myers JN, et al. Outcomes in patients with adenoid cystic carcinoma of the lacrimal gland. Ophthalmic Plast Reconstr Surg. 2004;20(1):22–6. Competing interests 13. Pommier P, Liebsch NJ, Deschler DG, Lin DT, McIntyre JF, Barker FG 2nd,
et al. Proton beam radiation therapy for skull base adenoid cystic
carcinoma. Arch Otolaryngol Head Neck Surg. 2006;132(11):1242–9. 13. Pommier P, Liebsch NJ, Deschler DG, Lin DT, McIntyre JF, Barker FG 2nd,
et al. Proton beam radiation therapy for skull base adenoid cystic
carcinoma. Arch Otolaryngol Head Neck Surg. 2006;132(11):1242–9. The authors declare that they have no competing interests. The authors declare that they have no competing interests. Availability of data and materials The datasets used during the current study are available from the
corresponding author on reasonable request. The datasets used during the current study are available from the
corresponding author on reasonable request. 9. Woo KI, Yeom A, Esmaeli B. Management of Lacrimal Gland Carcinoma:
lessons from the literature in the past 40 years. Ophthalmic Plast Reconstr
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Heegaard S. Lacrimal gland lesions in Denmark between 1974 and 2007. Acta Ophthalmol. 2013;91(4):349–54. Author details
1 Thariat J, Racadot S, Pointreau Y, Boisselier P, Grange JD, Graff P, et al. Intensity-
modulated radiotherapy of head and neck cancers: dose effects on the ocular,
orbital and eyelid structures. Cancer Radiother. 2016;20(6–7):467–74. 24. Marchand V, Dendale R. Normal tissue tolerance to external beam radiation
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1 14. Meldrum ML, Tse DT, Benedetto P. Neoadjuvant intracarotid chemotherapy
for treatment of advanced adenocystic carcinoma of the lacrimal gland. Arch Ophthalmol. 1998;116(3):315–21. 14. Meldrum ML, Tse DT, Benedetto P. Neoadjuvant intracarotid chemotherapy
for treatment of advanced adenocystic carcinoma of the lacrimal gland. Arch Ophthalmol. 1998;116(3):315–21. Author details
1Department of Electro-Optical Engineering, National Taipei University of
Technology, 1, Sec. 3, Chung-Hsiao E. Rd, Taipei 10608, Taiwan, ROC. 2Department of Ophthalmology, Chang Gung Memorial Hospital, Keelung,
Taiwan. 3College of Medicine, Chang Gung University, Taoyuan, Taiwan. 4Department of Radiation Oncology, Chang Gung Memorial Hospital, 15. Esmaeli B, Yin VT, Hanna EY, Kies MS, William WN Jr, Bell D, et al. Eye-sparing
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understanding. Arch Ophthalmol. 2009;127(8):1016–28. 22. Chawla B, Kashyap S, Sen S, Bajaj MS, Pushker N, Gupta K, et al. Clinicopathologic review of epithelial tumors of the lacrimal gland. Ophthalmic Plast Reconstr Surg. 2013;29(6):440–5. 23. Thariat J, Racadot S, Pointreau Y, Boisselier P, Grange JD, Graff P, et al. Intensity-
modulated radiotherapy of head and neck cancers: dose effects on the ocular,
orbital and eyelid structures. Cancer Radiother. 2016;20(6–7):467–74. 23. Publisher’s Note Springer Nature remains neutral with regard to jurisdictional claims in
published maps and institutional affiliations. Springer Nature remains neutral with regard to jurisdictional claims in
published maps and institutional affiliations. Springer Nature remains neutral with regard to jurisdictional claims in
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Leaky pipeline, gender bias, self-selection or all three? A quantitative analysis of gender balance at an international palliative care research conference
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Citation for published version (APA):
Sleeman, K. E., Koffman, J., & Higginson, I. J. (2019). Leaky pipeline, gender bias, self-selection or all three? A
quantitative analysis of gender balance at an international palliative care research conference. BMJ Supportive
and Palliative Care, 9(2), 146-148. https://doi.org/10.1136/bmjspcare-2016-001211 Citation for published version (APA):
Sleeman, K. E., Koffman, J., & Higginson, I. J. (2019). Leaky pipeline, gender bias, self-selection or all three? A
quantitative analysis of gender balance at an international palliative care research conference. BMJ Supportive
and Palliative Care, 9(2), 146-148. https://doi.org/10.1136/bmjspcare-2016-001211 Citing this paper
Pl
h C t
g t
s pape
Please note that where the full-text provided on King's Research Portal is the Author Accepted Manuscript or Post-Print version this may
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volume/issue, and date of publication details. And where the final published version is provided on the Research Portal, if citing you are
again advised to check the publisher's website for any subsequent corrections. Citation for published version (APA):
Sleeman, K. E., Koffman, J., & Higginson, I. J. (2019). Leaky pipeline, gender bias, self-selection or all three? A
quantitative analysis of gender balance at an international palliative care research conference. BMJ Supportive
and Palliative Care, 9(2), 146-148. https://doi.org/10.1136/bmjspcare-2016-001211 ABSTRACT Department of Palliative Care,
Policy and Rehabilitation, King’s
College London—Cicely
Saunders Institute, London, UK The gender balance in academia is the
subject of considerable debate.1–3 The
progress of women through academia has
been
described
as
a
‘leaky
pipeline’,
where there is attrition of women at each
step of the career ladder. Initiatives to
redress this are gaining momentum, for
example the UK Athena SWAN Charter
which was established in 2005 to encour-
age and recognise commitment to advan-
cing the careers of women in science,
technology, engineering, maths and medi-
cine (STEMM). Objectives The ‘leaky pipeline’ in academia is a
clearly described phenomenon, but has not been
examined in palliative care. We analysed the
gender balance of speakers at the 9th World
Research Congress of the European Association
of Palliative Care (EAPC) to test the null
hypothesis that there is no difference in the
proportion of women and men with senior
academic visibility in palliative care conference
programmes. Correspondence to
Dr Katherine E Sleeman,
Department of Palliative Care,
Policy and Rehabilitation, King’s
College London—Cicely
Saunders Institute, London, UK;
katherine.sleeman@kcl.ac.uk Correspondence to
Dr Katherine E Sleeman,
Department of Palliative Care,
Policy and Rehabilitation, King’s
College London—Cicely
Saunders Institute, London, UK;
katherine.sleeman@kcl.ac.uk Received 28 July 2016
Revised 22 November 2016
Accepted 26 January 2017 Received 28 July 2016
Revised 22 November 2016
Accepted 26 January 2017 Methods The final programme of the 2016
EAPC World Congress was examined, and the
gender of each speaker was recorded. Presentations were assessed using a three-tier
hierarchy of senior academic visibility: Free
Communication sessions, Themed sessions and
invited Plenaries (low to high). As there was only
one Invited Plenary at EAPC 2016, we examined
the gender balance at EAPC Plenaries from 2012
to 2016. Visibility, for example through confer-
ence talks and plenaries, is an important
dimension of gender equality. A paucity
of women conference speakers has been
demonstrated in several academic disci-
plines.4 However, it is unclear if the same
phenomenon exists in palliative care, a
specialty
where
women
traditionally
make up the majority of the workforce.5 Results Overall, the majority of speakers at
EAPC 2016 (96/130, 73.8%) were women. The
proportion of women was highest in the Free
Communication sessions (84/107, 78.5%). In the
Themed sessions, women made up just over half
of speakers (12/22, 54.5%). In 2016, there was
1 invited Plenary speaker, a man. General rights General rights
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the work immediately and investigate your claim. Download date: 24. Oct. 2024 Short report To cite: Sleeman KE,
Koffman J, Higginson IJ. BMJ
Supportive & Palliative Care
Published Online First: [please
include Day Month Year]
doi:10.1136/bmjspcare-2016-
001211 Katherine E Sleeman, Jonathan Koffman, Irene J Higginson Katherine E Sleeman, Jonathan Koffman, Irene J Higginson Received 28 July 2016
Revised 22 November 2016
Accepted 26 January 2017 ABSTRACT From 2012 to
2016, just 6 of 23 invited Plenary speakers at
EAPC conferences have been women (26.1%)
(χ2=25.4, p<0.001). In June 2016, 1200 delegates attended
the 9th World Research Congress of the
European Association of Palliative Care
(EAPC) in Dublin, making it one of the
largest research conferences in palliative
care globally. The programme featured
17 Free Communication sessions where
researchers were chosen on the quality of
anonymous (and gender-blind) abstracts
to present their work in 8 min talks. In
addition, there were seven Themed ses-
sions in which speakers were invited on
the basis of their level of authority on a
specific subject to give 20 min talks. The
most prestigious speaking opportunity is
the invited Plenary where high-profile Conclusions These data reject our null
hypothesis and suggest that there is attrition of
women along the academic pipeline in palliative
care. Other factors such as self-selection (that
women decline invitations to give talks) and
unconscious gender bias need further
exploration, as well as actions to address the
imbalance. To cite: Sleeman KE,
Koffman J, Higginson IJ. BMJ
Supportive & Palliative Care
Published Online First: [please
include Day Month Year]
doi:10.1136/bmjspcare-2016-
001211 1 Sleeman KE, et al. BMJ Supportive & Palliative Care 2017;0:1–3. doi:10.1136/bmjspcare-2016-001211 Short report academics are invited to give longer talks (30 min) on
the main conference stage. gender balance of invited Plenary speakers for a
5-year period (figure 1). At EAPC 2015, there were
eight invited Plenary speakers (including a debate), of
whom two were women (25%). At EAPC 2014, there
were two Plenary speakers, both men. At EAPC 2013,
there were nine Plenary speakers, including three
women (33.3%). At EAPC 2012, one of three Plenary
speakers
was
a
woman
(33.3%). Taking
5 years
together, 6 of 23 invited Plenary speakers at EAPC
conferences have been women (26.1%) (χ2=25.4,
p<0.001). We aimed to analyse the gender balance of speakers
at the 2016 EAPC World Congress to test the null
hypothesis that there is no attrition of women in
senior academic positions in palliative care, using the
three-tiered hierarchy of Free Communication ses-
sions, Themed sessions and Plenaries as a proxy for
seniority. DISCUSSION
P lli i Palliative care is a profession where women make up
the majority of the workforce,5 and this is reflected in
the overall gender balance of those giving talks at
EAPC 2016. However, we identified that the propor-
tion of women is lower for invited talks compared to
those chosen anonymously from abstracts: whereas
around
three
quarters
of
speakers
in
the
Free
Communication sessions were women, this proportion
reduced to roughly half for the Themed sessions, and
to only a quarter for the invited Plenaries (using
pooled data from 2012 to 2016). Given the large number of women working in pal-
liative care, it is of concern that the most prestigious
invited Plenaries are so infrequently given by women,
and it is noteworthy that two EAPC conferences over
the past 5 years have included no female Plenary
speakers at all (2014 and 2016). These data suggest
that there is attrition of women along the ‘academic
pipeline’ in palliative care. However, the relative con-
tribution of other potential factors such as self-
selection (that women decline invitations to give talks)
and unconscious gender bias (that women are not
invited to give talks) is unclear. In addition, palliative
care is multiprofessional, with gender unequally repre-
sented within each of the professions. The potential
contribution of discipline should also be explored. METHODS The final programme of the 2016 EAPC World
Congress was examined, and the gender of each
speaker was recorded. Speaker gender was determined
either by direct observation of their talk or by infer-
ence from their first name. In eight cases, Google was
used for more information. Where there was a substi-
tution of speaker at the last minute, the gender of the
speaker originally planned was used. At the 2016
EAPC, five top scoring abstracts were presented as
15 min Plenaries on the main stage. Since the decision
to feature these presentations was made on the basis
of an anonymous abstract, these were analysed with
the Free Communication sessions. Since there was
only one invited Plenary speaker at 2016 EAPC, we
examined the gender balance of invited EAPC Plenary
speakers
over
5 years
(2012–2016)
by
searching
through relevant scientific programmes online. χ2
tests were used to assess differences. We did not
analyse Meet the Expert sessions or Poster Discussion
sessions. Sleeman KE, et al. BMJ Supportive & Palliative Care 2017;0:1–3. doi:10.1136/bmjspcare-2016-001211 RESULTS At EAPC 2016, there were a total of 17 Free
Communication sessions comprising 102 talks. An
additional five top scoring abstracts were presented on
the main stage. There were seven Themed sessions
which included 22 speakers. There was a single
invited Plenary, the Ventafridda Lecture. Given that there was a single Plenary speaker in
2016, we examined the gender balance of Plenary
speakers over 5 years. This was based on the assump-
tion that variation from conference to conference is
likely to outweigh any temporal trends. The propor-
tion of female Plenary speakers was one-third or
fewer for each of the 5 years examined. There was no
indication that the proportion of female Plenary
speakers improved over time. Indeed, the 2012 and
2013 conferences had the greatest proportion of
female Plenary speakers (one-third). y
Overall, the majority of speakers at EAPC 2016
were women. The proportion of women was highest
in the Free Communication sessions. In the Themed
sessions, women made up just over half of speakers. The Ventafridda Lecture was given by a man (table 1). g
y
Since there was only one invited Plenary speaker at
EAPC 2016, we examined previous EAPC conference
programmes to obtain a more representative view of Table 1
Men and women giving talks at EAPC 2016, according
to session type
Men (%)
Women (%)
Invited plenary
1 (100)
0 (0)
Themed session
10 (45.5)
12 (54.5)
Free communication
23 (21.5)
84* (78.5)
Total
34 (26.2)
96 (73.8)
*Includes five top scoring abstracts presented on the main stage. Table 1
Men and women giving talks at EAPC 2016, according
to session type Speaking at academic conferences is important for
career advancement. Such opportunities not only
facilitate networking and collaboration, but are used
as
a
marker
of
quality
by
promotions
panels.6
Strategies to
improve
the
proportion
of
women
invited to speak, particularly those invited to give
Plenaries, should be explored. These include consider-
ation of the gender balance on scientific organising 2 Sleeman KE, et al. BMJ Supportive & Palliative Care 2017;0:1–3. doi:10.1136/bmjspcare-2016-001211 Figure 1
Number of men and women giving invited Plenaries, 2012–2016. Short report Short report Figure 1
Number of men and women giving invited Plenaries, 2012–2016. Competing interests IJH was Chair of the Scientific Organising
Committee of the 2015 EAPC World Congress in Copenhagen
and gave an invited Plenary at the 2012 EAPC Research
Congress in Trondheim. RESULTS KES, JK and IJH are members of the
King’s College London Department of Palliative Care Athena
SWAN Self-Assessment Team. committees, as the proportion of women on conven-
ing panels has been shown to be correlated with the
proportion of women speakers.4 6 The issue of self-
selection is important: previous studies have shown
that women are more likely than men to decline invi-
tations to speak at conferences,7 and less likely than
men to put themselves forward for talks.8 These
issues should be explored and addressed, for example
through routine collection and analysis of the gender
of all authors who submit abstracts, not only for
EAPC Congresses but for all major palliative care con-
ferences. Scientific organising committees should scru-
tinise information on invitations, acceptances and
refusals for high-profile speaking positions. Where
women decline invitations to speak, the reasons for
this should be explored, and steps to increase future
acceptance rates identified. These initiatives will help
us to understand the relative contributions of lack of
assertiveness and lack of recognition. In addition, rou-
tinely collecting feedback from delegates on the issue
of gender equality might identify hidden issues. Last,
we suggest that explicit acknowledgement of low visi-
bility of female speakers by conference organisers may
itself act as an intervention to redress the balance. committees, as the proportion of women on conven-
ing panels has been shown to be correlated with the
proportion of women speakers.4 6 The issue of self-
selection is important: previous studies have shown
that women are more likely than men to decline invi-
tations to speak at conferences,7 and less likely than
men to put themselves forward for talks.8 These
issues should be explored and addressed, for example
through routine collection and analysis of the gender
of all authors who submit abstracts, not only for
EAPC Congresses but for all major palliative care con-
ferences. Scientific organising committees should scru-
tinise information on invitations, acceptances and
refusals for high-profile speaking positions. Where
women decline invitations to speak, the reasons for
this should be explored, and steps to increase future
acceptance rates identified. These initiatives will help
us to understand the relative contributions of lack of
assertiveness and lack of recognition. In addition, rou-
tinely collecting feedback from delegates on the issue
of gender equality might identify hidden issues. RESULTS Last,
we suggest that explicit acknowledgement of low visi-
bility of female speakers by conference organisers may
itself act as an intervention to redress the balance. Provenance and peer review Not commissioned; externally
peer reviewed. Provenance and peer review Not commissioned; externally
peer reviewed. Open Access This is an Open Access article distributed in
accordance with the terms of the Creative Commons
Attribution (CC BY 4.0) license, which permits others to
distribute, remix, adapt and build upon this work, for
commercial use, provided the original work is properly cited. See: http://creativecommons.org/licenses/by/4.0/ REFERENCES 1
Moss-Racusin CA, Dovidio JF, Brescoll VL, et al. Science
faculty’s subtle gender biases favor male students. Proc Natl
Acad Sci USA 2012;109:16474–9. 2
Edmunds LD, Ovseiko PV, Shepperd S, et al. Why do
women choose or reject careers in academic medicine? A
narrative review of empirical evidence. Lancet
2016;388:2948–58. 3
Handley IM, Brown ER, Moss-Racusin CA, et al. Quality of
evidence revealing subtle gender biases in science is in the
eye of the beholder. Proc Natl Acad Sci USA
2015;112:13201–6. Failure of promotion on merit is deleterious for aca-
demia and ultimately for patient care. Whether these
data represent a leaky pipeline, gender bias, self-
selection or all three, it is essential that as a specialty
we take this issue seriously. We hope that the data pre-
sented here provide a starting point for more in-depth
consideration of this issue and appropriate strategies
to tackle it. 4
Isbell LA, Young TP, Harcourt AH. Stag parties linger:
continued gender bias in a female-rich scientific discipline. PLoS ONE 2012;7:e49682. 5
NCPC Specialist Palliative Care (SPC) Workforce Survey 2013. http://www.ncpc.org.uk/sites/default/files/NCPC_SPC_Workforce_
Survey_2013.pdf (accessed 6 February 2017) 6
Casadevall A. Achieving speaker gender equity at the American
Society for Microbiology General Meeting. mBio 2015;6:e01146. 7
Schroeder J, Dugdale HL, Radersma R, et al. Fewer invited
talks by women in evolutionary biology symposia. J Evol Biol
2013;26:2063–9. 7
Schroeder J, Dugdale HL, Radersma R, et al. Fewer invited
talks by women in evolutionary biology symposia. J Evol Biol
2013;26:2063–9. Contributors KES conceived and designed this study, performed
the analysis and wrote the manuscript, with help from IJH and
JK throughout. 8
Jones TM, Fanson KV, Lanfear R, et al. Gender differences in
conference presentations: a consequence of self-selection? PeerJ
2014;2:e627. Funding KES is funded by an NIHR Clinician Scientist
Fellowship and has received funding from Cicely Saunders
International. No specific funding was received for this study. 3 Sleeman KE, et al. BMJ Supportive & Palliative Care 2017;0:1–3. doi:10.1136/bmjspcare-2016-001211
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Fabrication of a Double Core–Shell Particle-Based Magnetic Nanocomposite for Effective Adsorption-Controlled Release of Drugs
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Polymers
| 2,022
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cc-by
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Article
Fabrication of a Double Core–Shell Particle-Based Magnetic
Nanocomposite for Effective Adsorption-Controlled Release
of Drugs Manzoor Hussain 1, Touseef Rehan 2, Khang Wen Goh 3, Sayyed Ibrahim Shah 4,*, Abbas Khan 1
,
Long Chiau Ming 5,*
and Nasrullah Shah 1,* 1
Department of Chemistry, Abdul Wali Khan University Mardan, Mardan 23200, Pakistan; 1
Department of Chemistry, Abdul Wali Khan University Mardan, Mardan 23200, Pakistan;
manzoor14ics@gmail.com (M.H.); abbas053@gmail.com (A.K.) 1
Department of Chemistry, Abdul Wali Khan University Mardan, Mardan 23200, Pakistan;
manzoor14ics@gmail.com (M.H.); abbas053@gmail.com (A.K.) 2
Department of Food and Nutrition, Shaheed Benazir Bhutto Women University, Peshawar 25000, Pakistan;
touseefnasr@gmail.com 2
Department of Food and Nutrition, Shaheed Benazir Bhutto Women University, Peshawar 25000, Pakistan;
touseefnasr@gmail.com 3
Faculty of Data Science and Information Technologies, INTI International University, Nilai 78100, Malaysia
khangwen.goh@newinti.edu.my 3
Faculty of Data Science and Information Technologies, INTI International University, Nilai 78100, Malaysia;
khangwen.goh@newinti.edu.my g
g
y
4
Department of Pharmacy, Abdul Wali Khan University Mardan, Mardan 23200, Pakistan 4
Department of Pharmacy, Abdul Wali Khan University Mardan, Mardan 23200, Pakistan
5
PAP Rashidah Sa’adatul Bolkiah Institute of Health Sciences, Universiti Brunei Darussalam,
Gadong BE1410, Brunei Department of Pharmacy, Abdul Wali Khan University Mardan, Mardan 23200, Pakistan
5
PAP Rashidah Sa’adatul Bolkiah Institute of Health Sciences, Universiti Brunei Darussalam,
Gadong BE1410, Brunei *
Correspondence: ibrahimshah@awkum.edu.pk (S.I.S.); long.ming@ubd.edu.bn (L.C.M.);
nasrullah@awkum.edu.pk (N.S.) Abstract: There has been very limited work on the control loading and release of the drugs aprepitant
and sofosbuvir. These drugs need a significant material for the control of their loading and release
phenomenon that can supply the drug at its target site. Magnetic nanoparticles have characteristics
that enable them to be applied in biomedical fields and, more specifically, as a drug delivery system
when they are incorporated with a biocompatible polymer. The coating with magnetic nanoparticles
is performed to increase efficiency and reduce side effects. In this regard, attempts are made to
search for suitable materials retaining biocompatibility and magnetic behavior. In the present study,
silica-coated iron oxide nanoparticles were incorporated with core–shell particles made of poly(2-
acrylamido-2-methylpropane sulfonic acid)@butyl methacrylate to produce a magnetic composite
material (MCM-PA@B) through the free radical polymerization method. The as-prepared compos-
ite materials were characterized through Fourier-transform infrared (FTIR)spectroscopy, scanning
electron microscopy (SEM), X-ray diffraction analysis (XRD), energy-dispersive X-Ray Analysis
(EDX), and thermogravimetric analysis (TGA), and were further investigated for the loading and
release of the drugs aprepitant and sofosbuvir. Citation: Hussain, M.; Rehan, T.;
Goh, K.W.; Shah, S.I.; Khan, A.; Ming,
L.C.; Shah, N. Fabrication of a Double
Core–Shell Particle-Based Magnetic
Nanocomposite for Effective
Adsorption-Controlled Release of
Drugs. Polymers 2022, 14, 2681. https://doi.org/10.3390/
polym14132681
Academic Editors: Faisal Raza,
Muhammad Ovais and
Carmine Coluccini
Received: 21 May 2022
Accepted: 27 June 2022
Published: 30 June 2022 Citation: Hussain, M.; Rehan, T.;
Goh, K.W.; Shah, S.I.; Khan, A.; Ming,
L.C.; Shah, N. Fabrication of a Double
Core–Shell Particle-Based Magnetic
Nanocomposite for Effective
Adsorption-Controlled Release of
Drugs. Polymers 2022, 14, 2681. https://doi.org/10.3390/
polym14132681
Academic Editors: Faisal Raza,
Muhammad Ovais and
Carmine Coluccini
Received: 21 May 2022
Accepted: 27 June 2022
Published: 30 June 2022 Citation: Hussain, M.; Rehan, T.;
Goh, K.W.; Shah, S.I.; Khan, A.; Ming,
L.C.; Shah, N. Fabrication of a Double
Core–Shell Particle-Based Magnetic
Nanocomposite for Effective
Adsorption-Controlled Release of
Drugs. Polymers 2022, 14, 2681. https://doi.org/10.3390/
polym14132681 Publisher’s Note: MDPI stays neutral
with regard to jurisdictional claims in
published maps and institutional affil-
iations. Publisher’s Note: MDPI stays neutral
with regard to jurisdictional claims in
published maps and institutional affil-
iations. Keywords: magnetic core–shell particles; composite material; thermogravimteric analysis; controlled
release; adsorption kinetic model Article
Fabrication of a Double Core–Shell Particle-Based Magnetic
Nanocomposite for Effective Adsorption-Controlled Release
of Drugs The maximum loading capacity of 305.76 mg/g for
aprepitant and 307 mg/g for sofosbuvir was obtained at pH 4. Various adsorption kinetic models
and isotherms were applied on the loading of both drugs. From all of the results obtained, it was
found that MCM-PA@B can retain the drug for more than 24 h and release it slowly, due to which it
can be applied for the controlled loading and targeted release of the drugs. polymers polymers 1. Introduction Among the
list of essential drugs, it has been included in basic health systems by the World Health
Organization. By combining it with the NS5A inhibitor ledipasvir, sofosbuvir is used to
treat genotypes 1, 2, 3, 4, 5, and 6, as well as hepatitis C infections [15–17]. Aprepitant and
sofosbuvir have little water solubility, resulting in erratic bioavailability. This is because the
active substance present in the drugs should be in dissolved form in aqueous form in the
gastrointestinal tract, such that it crosses the luminal wall to make its therapeutic effects
more significant. Moreover, higher solubility allows the active form of the drug to easily
enter the blood and reach its target site to show its therapeutic effects. The loading and
release of these drugs into the carrier molecules is also of great concern due to limited data
available for them. Copolymers of 2-acrylamido-2-methyl propane sulfonic acid (AMPS) with other poly-
mers have been synthesized and applied for several purposes. Molecularly imprinted
polymer of N-maleoyl chitosan with AMPS have been prepared and applied for the recog-
nition and delivery of bovine serum albumin [18]. A copolymer of AMPS with acrylamide
has been applied for adsorption of cadmium from aqueous solution and for the loading
and release of 5- fluorouracil, as well as the antiviral drug indinavir sulfate [19–21]. An-
other copolymer of AMPS with polyvinyl alcohol and acrylamide has been applied for
the controlled loading and release of ibuprofen [22]. Hydrogel prepared from chitosan
and poly(AMPS) has been applied for the removal of methylene blue, acid red dye, Cd(II),
and Cr(II) from wastewater [23]. Hydrogel prepared from poly(AMPS) and pectine has
been applied for the controlled loading and release of captopril [24]. Different types of
nanomaterials prepared from poly(α-L-glutamic acid) have been prepared and applied
for the loading and controlled release of various types of drugs [25]. In one of the recent
studies, hollow polymeric nanosphere (HPN)-supported imidazolium-based ionic liquids
were shown to be efficient materials against multidrug-resistant bacteria, due to which they
can also be applied in the biomedical field [26]. The fabrication of magnetic core–shell composites for drug loading and controlled
release is of great value due to their efficiency and ability of controlled targeted drug
delivery. The controlled loading and release of curcumin has also been demonstrated by the
application of core–shell particles comprising chitosan with magnetic iron oxide nanoparti-
cles (IONPs) and gold nanoparticles [27]. 1. Introduction Copyright:
© 2022 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article
distributed
under
the
terms
and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/). Magnetic nanoparticles have recently been synthesized and applied in a large number
of fields, including loading and controlled release of drugs, separation of cells, enhancing
magnetic resonance imaging, the treatment of cancer, and many more [1]. Nanoparticles
based on iron oxide are chemically stable, easily diffusible, less toxic, and biocompatible,
due to which they are highly significant [2]. The use of bare iron oxide nanoparticles has
problems of low accumulation, low separation yield by magnets, and the possibility that
their atoms may oxidize, decreasing the properties of dispersion and magnetism [3]. The https://www.mdpi.com/journal/polymers Polymers 2022, 14, 2681. https://doi.org/10.3390/polym14132681 Polymers 2022, 14, 2681 2 of 20 property of having a high surface-to-volume ratio of iron oxide nanoparticles provides room
for the adsorption of plasma proteins, followed by the activation of clearance mechanisms
before they reach the target site. The time for the circulation of nanoparticles and decrease in
the risk of adsorption of plasma proteins is also enhanced by the modification of the surface
with some biocompatible polymers with a hydrophilic nature [4,5]. Iron oxide nanoparticles
are mostly encapsulated by the silica, because it is inert and has ease of functionalization,
and after encapsulation these nanoparticles become water-compatible [6,7]. The use of bare
magnetic nanoparticles is usually prohibited for the application of drug delivery, because
of the compromised biocompatibility and they do not possess various functionalities
facilitating the coating of polymers on their surfaces. For this reason, they are coated with
silica so that their surfaces are ready for coating and enhanced interaction with drugs [8–10]. y
g
g
Aprepitant is an orally active neurokinin receptor antagonist used for chemotherapy-
induced nausea and vomiting (CINV) [11]. It is the first FDA-approved drug that is used
to treat CINV that works for more than 24 h after chemotherapy treatment [12]. It has a
very low affinity for serotonin (5-HT3), corticosteroid, and dopamine receptors. Various
drugs have been developed to prevent acute and delayed emesis, but aprepitant has been
approved for PONV and nausea treatment during cancer therapy [13]. Sofosbuvir is a drug
that is used in combination with other drugs in order to treat hepatitis C [14]. 2.1. Materials Sodium hydride 60%, magnesium turnings, and potassium persulfate were pur-
chased from Daejung, S. Korea. Dimethylformamide (DMF), 4-vinylbenzyl chloride
(VBC), 2-acrylamido-2-methylpropane sulfonic acid (AMPS), 4,4′-azobis(4-cyanovaleric
acid) (ABCA), dimethyl sulfoxide (DMSO), butyl methacrylate (BMA), bromobenzene,
and iodine were purchased from Sigma-Aldrich, Schnelldorf, Germany. Pyrroles and
carbon disulfide were purchased from Daejung, Korea, diethyl ether from Sigma-Aldrich,
Schnelldorf, Germany, magnesium sulfate and petroleum ether were from Fisher Scien-
tific, Chicago, IL, USA, and tetrahydrofuran (THF) was purchased from Daejung, Korea. Silica-gel, FeCl2.4H2O, and FeCl3.6H2O were purchased from UniChem USA. n-Hexane
was purchased from Sigma-Aldrich, Germany. Aprepitant and sofosbuvir were donated by
Ferozsons pharmaceutical company, Nowshera, Pakistan. 2.2. Methods 2.2.1. Preparation of Poly(2-acrylamido-2-methylpropane sulfonic acid) (poly(AMPS)) 1. Introduction The core-shell structure made of IONPs@Au has
been applied as an anticancer agent [28]. Iron oxide magnetic nanoparticles have been com-
bined with large number of polymers for vast applications, e.g.,polylactide-co-glycolide
combined with iron oxide nanoparticles has been applied for the encapsulation of both
hydrophilic and hydrophobic drugs [29]. AMPS incorporated with pectin followed by the
loading of IONPs has been applied for the controlled release of diclofenac sodium [30]. Polymers 2022, 14, 2681 3 of 20 Poly-(sodium-2-acrylamido-2-methyl propane sulfonic acid-co-styrene)/magnetite have
been used for the purpose to inhibit corrosion from steel [31]. Poly-(sodium-2-acrylamido-2-methyl propane sulfonic acid-co-styrene)/magnetite have
been used for the purpose to inhibit corrosion from steel [31]. p
p
Generally, there is a large number of drug delivery systems used, including liposomes,
graphene, gold, carbon nanotubes, and magnetic nanoparticles [32–36]. These aforemen-
tioned materials have little ability to deliver the drug in a controlled way. Bare magnetic
nanoparticles have limitations to their application, due to which they are incorporated
in some polymeric materials to enhance their applicability. Current research is focused
on site-specific drug delivery systems, by releasing the drug in response to an external
trigger, i.e., physiological pH and temperature [37–39]. In order to overcome the problem
of controlled drug delivery, we here focus on the synthesis of a novel drug delivery system
based on the combination of magnetic nanoparticles (silica-coated IONPs) and core–shell
particles (poly(2-acrylamido-2-methylpropane sulfonic acid)@butyl methacrylate). The
as-prepared magnetic composite material (MCM-PA@B) was thoroughly characterized by
various analytical techniques, and was applied for loading and controlled release using
aprepitant and sofosbuvir as model drugs. 2.2.2. Preparation of Core–Shell Latex of Poly(AMPS)@BMA (PA@B) 2.2.2. Preparation of Core–Shell Latex of Poly(AMPS)@BMA (PA@B) The fabricated Poly(AMPS) (1 g) was taken and dissolved in DI water in a reactor, and
then BMA (25 mL) was added, followed by the addition of K2S2O8 (1 g in 15 mL of DI
water). An overhead condenser was installed over the flask, and the mixture was stirred at
60 ◦C for 24 h in a water bath at 900 rpm using a magnetic stirrer. A milky, white-colored
suspension of CS was obtained, which was transferred to a reagent bottle and stored for
further use. 2.2.3. Synthesis of Silica-Coated IONPs (IONPs@SiO2) The synthesis of silica-coated IONPs was carried out in two consecutive steps: In
the first step, a co-precipitation method was used to synthesize the magnetic IONPs, as
reported by Veisi et al. [41]. Fe2+ and Fe3+ were mixed in 40 mL of double-DI water at a
molar concentration of 0.15 mol and 0.3 mol, respectively. Sodium hydroxide solution (2 M)
was added dropwise to keep the pH above 10, and temperature was maintained at 80 ◦C
for 6 h. Then, the solution was cooled and filtered, followed by washing three times with
distilled water. The black-colored particles were then dried in an oven to remove moisture,
and then stored in an inert environment. In the second step, the synthesized IONPs were coated with SiO2 using theStöber
method [10]. From the synthesized IONPs, 1.6 g was taken in a flask. Then, ethanol (80 mL),
deionized water (16 mL), 10% NaOH solution (16 mL), and TEOS (5 mL) were added to
it, followed by sonication for 1 h at 30 ◦C. Blackish grey-colored particles were produced,
which were further separated by using an external magnet. These synthesized silica-coated
magnetic nanoparticles were further washed with distilled water and ethanol, followed by
drying at 80 ◦C in an oven for six hours. 2.2.1. Preparation of Poly(2-acrylamido-2-methylpropane sulfonic acid) (poly(AMPS)) The as-prepared poly(AMPS) revealed the
following NMR data: g
1H NMR (400 MHz (D2O), ppm): 7.6 2H, (br s, N-pyrrole–H), 7.1 (br m, –Ar–), 6.3 (2H,
br s, N-pyrrole–H), 3.3 (2H, br, m, C-CH2-C), 2.1 (C-CHC-C) 1.4 (6H, br m, C-CH3)
13C NMR (400 Mhz (D2O) ppm): 58.1 (RCH2SO3H), 52 (RCHS), 38.6 (RCH2), 26.6 (RCH3) g
1H NMR (400 MHz (D2O), ppm): 7.6 2H, (br s, N-pyrrole–H), 7.1 (br m, –Ar–), 6.3 (2H,
br s, N-pyrrole–H), 3.3 (2H, br, m, C-CH2-C), 2.1 (C-CHC-C) 1.4 (6H, br m, C-CH3)
13C NMR (400 Mhz (D2O) ppm): 58.1 (RCH2SO3H), 52 (RCHS), 38.6 (RCH2), 26.6 (RCH3) 2.2.1. Preparation of Poly(2-acrylamido-2-methylpropane sulfonic acid) (poly(AMPS)) First of all, the RAFT agent 4-vinylbenzyl pyrrole carbodithioate (VP) was synthesized
as reported in our previous study [40]. Briefly, 160 mL of DMF was taken, in which sodium
hydride (6.03 g) was dissolved to produce a grey-colored suspension. Pyrrole solution was
prepared by dissolving 10.02 g in DMF to make 20 mL of solution, which was further added
to the mixture, forming a yellow-colored product. The temperature of this mixture was
dropped to 0 ◦C, and CS2 solution already prepared from 9.01 mL in 20 mL of DMF was
added to the mixture through continuous stirring for 30 min. Then, 4-VBC (22.2 g) solution
in 20 mL of DMF was added with continuous stirring to produce a dark-red-colored mixture
that was stirred overnight at room temperature (25 ◦C). This product was separated from
its mixture by using a separating funnel that contained a mixture of deionized water and
diethyl ether (1:1) and was further dried with 20 g of solid MgSO4. The product was filtered,
followed by vacuum distillation, and then, using silica gel as stationary phase, the product
was separated through the column. The obtained product was again vacuum-distilled,
producing a yellow-colored product thatwas kept in a nitrogen-purged environment at
−18 ◦C. For the preparation of poly(AMPS), 1 g of the RAFT agent 4-VP was added to 25 mL of
DMSO in a two-neck flask reactor. Then, 25 g of AMPS monomer was added to it, followed
by the addition of 0.5 g of 4,4′-azobis(4-cyanovaleric acid) (ABCA) initiator. The flask
containing the mixture was made oxygen-free by purging with N2 several times. Then, the
mixture was heated to 60 ◦C for 12 h. After heating, the mixture converted to brown-colored
poly(AMPS). The obtained product was stored at −5 ◦C in a nitrogen atmosphere, and was Polymers 2022, 14, 2681 4 of 20 used further for the synthesis of core–shells. The as-prepared poly(AMPS) revealed the
following NMR data: used further for the synthesis of core–shells. 2.2.4. Fabrication of Magnetic Composite Material (MCM-PA@B) Next, IONPs@SiO2wastaken in different amounts (0.1 g to 0.5 g) and dispersed in
deionized (DI) water by sonication. PA@B was added to it in different amounts (1 mL to
5 mL) and sonicated for 15 min. Then, 4,4′-azobis(4-cyanovaleric acid)(0.015 g) in 1.5 mL
of DI water was added to it and again sonicated for 15 min, followed by the addition of
IONPs@SiO2 suspension with continuous sonication for 10 min (Table 1). The as-prepared
mixture was then stirred for 2 h under nitrogen-purged conditions at 60 ◦C toproduce
the composite materials. The synthesized magnetic composite material (MCM-PA@B)
was then transferred to a beaker and kept in the oven until dried and converted to a
black-colored material. Table 1. Different ratios of IONPs@SiO2 and PA@B in the synthesis of MCM-PA@B. S. No. IONPs@SiO2:PA@B
IONPs@SiO2 (g)
PA@B (mL)
ABCA (mL)
1
IONPs@SiO21:PA@B1
0.1
1
2
2
IONPs@SiO21:PA@B3
0.1
3
2
3
IONPs@SiO21:PA@B5
0.1
5
2
4
IONPs@SiO23:PA@B5
0.3
5
2
5
IONPs@SiO25:PA@B5
0.5
5
2 Table 1. Different ratios of IONPs@SiO2 and PA@B in the synthesis of MCM-PA@B. 2.2.6. Application of MCM-PA@B for Drug Loading and Release Studies The as-prepared composite materials were further applied for the loading and release
of the drugs aprepitant and sofosbuvir. A stock solution of eachdrug (100 ppm) was
prepared in 100 mL of methanol through sonication at room temperature. From the stock
solution, five separate sample solutions were further prepared through dilution with a
concentration of 20 ppm. The composite material was added to each of the sample solutions
at neutral pH and STP. All of the drug solutions containing the composite were shaken
overnight. From each of the samples, aliquots were taken out after a certain interval of time
and checked through the UV spectrophotometer to find the loading amount of the drug. To optimize the conditions for maximum loading of the drugs, both of the drugs were
tested for their loading at different pH and drug concentrations. For pH optimization, drug
solutions were prepared, and their pH was set to2 by the addition of Britton–Robinson
buffer, after which 0.05 g of composite was added. This solution was shaken for 2 h, and
its absorbance was checked using the UV spectrophotometer. A similar procedure was
followed for pH 4, 6, 8, and 10. For optimization of the drug concentration used, drug solutions from 20 ppm to
100 ppm were prepared, and their pH was set to 4 by adding buffer, followed by the
addition of 0.05 g of composite materials. These solutions were shaken for 2 h and then
their absorbance was checked. Pseudo-2nd-Order Kinetic Model Pseudo-2nd-Order Kinetic Model A second-order kinetic equation was used for the interpretation of the data (Equation (3)): t/qt = t/qe + 1/k2 qe2
(3) (3) where k2 shows the rate constant for the second order, while qe and qt show the amount of
loaded drug at equilibrium and at any time. 2.2.5. Characterization of the Synthesized MCM-PA@B FTIR Spectrometer (Varian 640-IR, USA) was used to obtain the FTIR spectra of the
prepared MCM-PA@B in the wavenumber region of 4000–400 cm−1. A thermogravimetric
analyzer (TGA-50H Shimadzu, Kyoto, Japan) was used to study the thermal stability of Polymers 2022, 14, 2681 5 of 20 5 of 20 the prepared composite materials. The machine was run from 25 to 600 ◦C ata heating rate
of 30 ◦C/min. An X-ray diffractometer (D-2 Phraser, Bruker, Denver, CO, USA) was used
to analyze the XRD patterns by applying a scan rate of 20◦/min, and a Cu Kα radiation
source was used. A field-emission scanning electron microscope (SEM) (model JSM5910,
JEOL. Kyoto, Japan,) was used with an acceleration voltage of 30 KV to analyze the surface
morphology of the composite materials. A Thermo Electron Co., USA, Helios β UV–Vis
Spectrometer (Lambda 25, Perkin Elmer, Waltham, MA, USA) was used to determine the
UV–Vis absorption spectra of the samples. 2.2.7. Kinetic Study of Adsorption
Pseudo-1st-Order Kinetic Model Drugs’ loading data were obtained and, using a pseudo-1st-order kinetic model, were
used to interpret the obtained results (Equations (1) and (2)): Log (qe −qt)= log qe −k1t/2.303
(1)
qe or qt = (Ci −Ce)V/m
(2) Log (qe −qt)= log qe −k1t/2.303
(1) (1) qe or qt = (Ci −Ce)V/m
(2) (2) where qt is the amount of drug loaded at any time, while qe shows the amount of loaded
drug at equilibrium time, k1 is the constant for the first order, Ce and Ci are the drug’s
concentration at equilibrium and initial use, respectively, V is the volume of the drug
solution, and m is the mass of the composite materials (0.1 g). Intra-Particle Diffusion Kinetic Model The data were also interpreted using an intra-particle diffusion model (Equation (4)) qt = Kintt1⁄2 + C
(4) (4) Polymers 2022, 14, 2681 6 of 20 where C is the intercept and is associated withthe width of the border layer i.e., the higher
the value of C, the further is its boundary layer outcome. where C is the intercept and is associated withthe width of the border layer i.e., the higher
the value of C, the further is its boundary layer outcome. 2.2.8. Equilibrium Studies 3.1. Scanning Electron Microscope Study of MCM-PA@B 3.1. Scanning Electron Microscope Study of MCM-PA@B The morphological, internal, and topographical structure of the as-prepared MCM-
PA@B was evaluated by SEM analysis. It can be seen in Figure 1A,B that the IONPs and
IONPs@SiO2 had a small particle size; however, the incorporation of silica onto the IONPs
converted them into rough and heterogeneous surface particles [42]. The incorporation
of IONPs@SiO2 into the PA@B to produce MCM-PA@B can be seen in Figure 1C–G. The
IONPs@SiO21: PA@B1 sample contained an equal ratio of both IONPs@SiO2 and PA@B, due
to which its morphological structure did not look uniform. However, as the IONPs@SiO2
were incorporated with a higher amount of PA@B, uniformity in the structure enhanced
and made it more recognizable, as shown in Figure 1D–F. Moreover, the core–shell polymer
of PA@B was coated over the silica-coated IONPs to form double core–shell nanocompos-
ites. The surface polymer molecules covered the nanoparticles from all sides and, hence,
prevented their agglomeration, due to which these double core–shell nanocomposites were
distinguishable and clear. The similar composition of PA@B, with changes only to its
quantity, did not cause significant changes in the morphology of the nanocomposites [41]. As the ratio of IONPs@SiO2 increased, the surface again shifted towards non-uniformity. From the SEM results, it can be seen that using the IONPs@SiO2 and PA@B latex at ratios
of 1:3, 1:5, and 3:5 gave us composite materials with similar morphological structures. The
magnetic behavior of the synthesized magnetic nanoparticles and MCM-PA@B is presented
in Figure 2. 2.2.8. Equilibrium Studies Langmuir and Freundlich isotherms were used to determine the equilibrium study of
the drug incorporated into the synthesized composite materials. The Langmuir equation
shows monolayer attachment of the drug with the composite materials by providing many
sites for linking the drug with the composite materials. The Langmuir equation in presented
in Equation (5): Ce/qe = 1/KL + aL Ce/KL
(5) (5) where KL shows the equilibrium adsorption constant, while qe is the monolayer loading
capacity of the drug. p
y
g
The Freundlich isotherm is given in Equation (6): Log qe = log KF +1/n log Ce
(6) (6) The values for the constants (KF and N) of Freundlich and Langmuir (Q and KL) as
well as the regression coefficient R2 were calculated. 3.3. Thermogravimetric Analysis
3.3. Thermogravimetric Analysis By looking into the spectra of TGA analysis given in Figure 4, we can observe two
degradation stages for IONPs and IONPs@SiO2. In the first stage, the weight loss of IONPs
at 125 °C is due to the loss of water molecules attached tothe surface of the IONPs, while
in the second stage the weight loss at 170 °C and 360 °C is due to the transformation of
crystal from Fe3O4 to γ-Fe2O3 caused by heating,and only a 5% decrease in the weight of
the composite occurs [47]. In the case of IONPs@SiO2, in the first stage, 2% weight loss
occurs up to 230 °C due to the loss of any remaining water and volatile molecules, and
then the weight loss decreases by 12% due to the conversion of Fe3O4 to γ-Fe2O3 at 360 °C,
followed by no significant weight loss, indicating a higher degree of thermal stability [48]. All of the composites of CS and IONPs@SiO2 contain different amounts of poly(AMPS)
and IONPs@SiO2. The graph shows the TGA of all of these composites, and indicates that
each of the composite has three degradation stages. For the composites
IONPs@SiO21:PA@B1, IONPs@SiO21:PA@B3, IONPs@SiO21:PA@B5; the first degradation
stage occurs below 150 °C, showing a 10% loss of water and volatile molecules contents
[46], followed by degradation up to 300 °C, showing the loss of sulfonate groups of the
polymers up to 30%, and then 85% weight loss occurs at 450 °C due to the degradation of
the polymer on the IONPs@SiO2. Above 450 °C, there is no significant change in the
weight, confirming the presence of IONPs, as these particles have high thermal stability. Moreover, for the composites IONPs@SiO23:PA@B5 and IONPs@SiO25:PA@B5, both have
high amounts of IONPs as compared to previous composites, so they have relatively high
thermal stability, and a maximum of 70% weight loss occurs at 450 °C, followed by no
significant decrease in the weight of the samples. As IONPs@SiO25:PA@B5 contains more
IONPs@SiO2, it gives more thermal stability to the composite, due to which its graph is
slightly above that of the IONPs@SiO23:PA@B5 [31,46]. From the above discussion it can
By looking into the spectra of TGA analysis given in Figure 4, we can observe
two degradation stages for IONPs and IONPs@SiO2. 3.2. FTIR Study of the Composite The synthesis of IONPs is confirmed by the presence of peak at 565 cm−1, which is
due to the presence of stretching of Fe–O bonds in Fe2O3 in IONPs [43], while the band at
1084 confirms the silica coating on the IONPs to produce Fe2O3@SiO2 [44]. The bands at
1353 cm−1 are due to the presence of the sulfonate group of the poly(AMPS) [45].The peaks
at 2872 and 1452 cm−1 are due to CH and CH2 as well as C-N stretching vibrations [46]. The broad peak at 3100–3500 cm−1 indicates OH stretching and the presence of the NH
group. The peak at 941 cm−1 is due to S–O–C symmetrical stretching vibrations of the
poly(AMPS) shell, while that at 1735 cm–1 assures the presence of a CO group, and that at
1249 cm−1 shows the presence of a BMA core [20] (Figure 3). 7 of 20 Polymers 2022, 14, 2681
Polymers 2022, 14, x Figure 1. SEM images of IONPs (A), IONPs@SiO2 (B), MCM-PA@B (C–G), IONPs@SiO21:PA@B1
(C), IONPs@SiO21:PA@B3 (D), IONPs@SiO21:PA@B5 (E), IONPs@SiO23:PA@B5 (F), and
IONPs@SiO25:PA@B5 (G). Figure 1. SEM images of IONPs (A), IONPs@SiO2 (B), MCM-PA@B (C–G), IONPs@SiO21:PA@
(C),
IONPs@SiO21:PA@B3
(D),
IONPs@SiO21:PA@B5
(E),
IONPs@SiO23:PA@B5
(F),
a
IONPs@SiO25:PA@B5 (G). rs 2022, 14, x
8 of 22 Figure 1. SEM images of IONPs (A), IONPs@SiO2 (B), MCM-PA@B (C–G), IONPs@SiO21:PA@B1
(C), IONPs@SiO21:PA@B3 (D), IONPs@SiO21:PA@B5 (E), IONPs@SiO23:PA@B5 (F), and
IONPs@SiO25:PA@B5 (G). Figure 1. SEM images of IONPs (A), IONPs@SiO2 (B), MCM-PA@B (C–G), IONPs@SiO21:PA@B1
(C),
IONPs@SiO21:PA@B3
(D),
IONPs@SiO21:PA@B5
(E),
IONPs@SiO23:PA@B5
(F),
and
IONPs@SiO25:PA@B5 (G). 8 of 22 Figure 1. SEM images of IONPs (A), IONPs@SiO2 (B), MCM-PA@B (C–G), IONPs@SiO21:PA@B1
(C), IONPs@SiO21:PA@B3 (D), IONPs@SiO21:PA@B5 (E), IONPs@SiO23:PA@B5 (F), and
IONPs@SiO25:PA@B5 (G). Figure 1. SEM images of IONPs (A), IONPs@SiO2 (B), MCM-PA@B (C–G), IONPs@SiO21:PA@B1
(C),
IONPs@SiO21:PA@B3
(D),
IONPs@SiO21:PA@B5
(E),
IONPs@SiO23:PA@B5
(F),
and
IONPs@SiO25:PA@B5 (G). 8 of 22 Figure 2. Magnetic behavior of the synthesized magnetic nanoparticles (A,B) and MCM-PA@B
(C,D). Figure 2. Magnetic behavior of the synthesized magnetic nanoparticles (A,B) and MCM-PA@B (C,D). Figure 2. Magnetic behavior of the synthesized magnetic nanoparticles (A,B) and MCM-PA@B
(C,D). Figure 2. Magnetic behavior of the synthesized magnetic nanoparticles (A,B) and MCM-PA@B (C,D). 8 of 20
9 of 22 Polymers 2022, 14, 2681
Polymers 2022, 14, x Figure 3. FTIR spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. Figure 3. 3.2. FTIR Study of the Composite FTIR spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. Figure 3. FTIR spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. Figure 3. FTIR spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. 3.3. Thermogravimetric Analysis
3.3. Thermogravimetric Analysis 0
100
200
300
400
500
600
Weight loss (%)
0
20
40
60
80
100
120
IONPs
IONPs@SiO2
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 composite materials with high concentrations of IONPs@SiO2 are more thermally stable
than those with low amounts of IONPs@SiO2. composite materials with high concentrations of IONPs@SiO2 are more thermally stable
than those with low amounts of IONPs@SiO2. Temperature
0
100
200
300
400
500
600
Weight loss (%)
0
20
40
60
80
100
120
IONPs
IONPs@SiO2
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
Figure 4. Thermogravimetric analysis of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B
with different ratios of IONPs@SiO2 and PA@B. Figure 4. Thermogravimetric analysis of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B
with different ratios of IONPs@SiO2 and PA@B. Weight loss (%) 0 Figure 4. Thermogravimetric analysis of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@
with different ratios of IONPs@SiO2 and PA@B. Figure 4. Thermogravimetric analysis of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B
with different ratios of IONPs@SiO2 and PA@B. 3.4. XRD Analysis
3.4. XRD Analysis The XRD analysis results of the synthesized composite materials are given in Figure
5. The data from the graph show us that the IONPs have some sharp peaks that confirm
their crystalline structure. Moreover, the crystalline form of the nanoparticles is confirmed
by the sharp peaks at 30.99°, 36.3°, 44.1°, 58°, and 63.6° [46]. The graph of the IONPs@SiO2
also shows similar peaks that ensures its crystalline nature, with no loss of any of the en-
tity. This shows that after coating of of silica on the IONPs, the crystalline structure of the
nanoparticles was not disturbed, and showed the peaks in the similar range. By incorpo-
ration of the core–shell latex with IONPs@SiO2, the intensity of these peaks decreased and
became very low, ensuring their combination. However, due to the presence of the
IONPs@SiO2, the peaks are still present with low intensity, confirming the presence of
both the PA@B and IONPs@SiO2. The XRD analysis results of the synthesized composite materials are given in Figure 5. The data from the graph show us that the IONPs have some sharp peaks that confirm their
crystalline structure. Moreover, the crystalline form of the nanoparticles is confirmed by the
sharp peaks at 30.99◦, 36.3◦, 44.1◦, 58◦, and 63.6◦[46]. The graph of the IONPs@SiO2 also
shows similar peaks that ensures its crystalline nature, with no loss of any of the entity. This
shows that after coating of of silica on the IONPs, the crystalline structure of the nanoparti-
cles was not disturbed, and showed the peaks in the similar range. By incorporation of the
core–shell latex with IONPs@SiO2, the intensity of these peaks decreased and became very
low, ensuring their combination. However, due to the presence of the IONPs@SiO2, the
peaks are still present with low intensity, confirming the presence of both the PA@B and
IONPs@SiO2. 3.5. Zero-Point Charge (pHzpc) on the Composite 3.3. Thermogravimetric Analysis
3.3. Thermogravimetric Analysis In the first stage, the weight loss of
IONPs at 125 ◦C is due to the loss of water molecules attached to the surface of the IONPs,
while in the second stage the weight loss at 170 ◦C and 360 ◦C is due to the transformation
of crystal from Fe3O4 to γ-Fe2O3 caused by heating, and only a 5% decrease in the weight
of the composite occurs [47]. In the case of IONPs@SiO2, in the first stage, 2% weight loss
occurs up to 230 ◦C due to the loss of any remaining water and volatile molecules, and
then the weight loss decreases by 12% due to the conversion of Fe3O4 to γ-Fe2O3 at 360 ◦C,
followed by no significant weight loss, indicating a higher degree of thermal stability [48]. All of the composites of CS and IONPs@SiO2 contain different amounts of poly(AMPS) and
IONPs@SiO2. The graph shows the TGA of all of these composites, and indicates that each
of the composite has three degradation stages. For the composites IONPs@SiO21:PA@B1,
IONPs@SiO21:PA@B3, IONPs@SiO21:PA@B5; the first degradation stage occurs below
150 ◦C, showing a 10% loss of water and volatile molecules contents [46], followed by
degradation up to 300 ◦C, showing the loss of sulfonate groups of the polymers up to
30%, and then 85% weight loss occurs at 450 ◦C due to the degradation of the polymer on
the IONPs@SiO2. Above 450 ◦C, there is no significant change in the weight, confirming
the presence of IONPs, as these particles have high thermal stability. Moreover, for the
composites IONPs@SiO23:PA@B5 and IONPs@SiO25:PA@B5, both have high amounts of
IONPs as compared to previous composites, so they have relatively high thermal stability,
and a maximum of 70% weight loss occurs at 450 ◦C, followed by no significant decrease
in the weight of the samples. As IONPs@SiO25:PA@B5 contains more IONPs@SiO2, it
gives more thermal stability to the composite, due to which its graph is slightly above that
of the IONPs@SiO23:PA@B5 [31,46]. From the above discussion it can be concluded that Polymers 2022, 14, 2681 9 of 20
re more 9 of 20
re more composite materials with high concentrations of IONPs@SiO2 are more thermally sta
than those with low amounts of IONPs@SiO2. 3.5. Zero-Point Charge (pHzpc) on the Composite The PA@B contains a sulfonate group from the poly(AMPS) shell, which is incorpo-
rated with the IONPs@SiO2. Due to the presence of these acidic sulfonate groups, the
pHzpc of the composite material was expected to be in the acidic range, as confirmed by
the results obtained, with a value of 2.5. Below the pHzpc value, the composite has positive
charge, while increasing above this value it has negative charge (Figure 6). 10 of 20 10 of 20 Polymers 2022, 14, 2681 Figure 5. XRD spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. 3.5. Zero-Point Charge (pHzpc) on the Composite
The PA@B contains a sulfonate group from the poly(AMPS) shell, which is incorpo-
rated with the IONPs@SiO2. Due to the presence of these acidic sulfonate groups, the
pHzpc of the composite material was expected to be in the acidic range, as confirmed by
the results obtained, with a value of 2.5. Below the pHzpc value, the composite has posi-
tive charge, while increasing above this value it has negative charge (Figure 6). Figure 5. XRD spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. Polymers 2022, 14, x
12 of 22
Figure 6. pHzpc of the fabricated composite materials (MCM-PA@B). 3.6. EDX Analysis
Figure 7 shows the EDX analysis of the synthesized IONPs. The incorporation of sil-
i
ith th IONP i
fi
d b
th
f it
k i
Fi
7
M
th
pH
0
2
4
6
8
10
12
14
ΔpH
-1
0
1
2
3
4
5
6
7
Figure 6. pHzpc of the fabricated composite materials (MCM-PA@B). Figure 5. XRD spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. Figure 5. XRD spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. olymers 2022, 14, x
12 of 22 Polymers 2022, 14, x
12 of 22 Figure 5. XRD spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. Figure 5. XRD spectra of pure IONPs, IONPs@SiO2, and the fabricated MCM-PA@B with different
ratios of IONPs@SiO2 and PA@B. 3.5. Zero-Point Charge (pHzpc) on the Composite
The PA@B contains a sulfonate group from the poly(AMPS) shell, which is i
rated with the IONPs@SiO2. 3.5. Zero-Point Charge (pHzpc) on the Composite Due to the presence of these acidic sulfonate gro
pHzpc of the composite material was expected to be in the acidic range, as confir
the results obtained, with a value of 2.5. Below the pHzpc value, the composite h
tive charge, while increasing above this value it has negative charge (Figure 6). Figure 6. pHzpc of the fabricated composite materials (MCM-PA@B). 3 6 EDX A
l
i
pH
0
2
4
6
8
10
12
14
ΔpH
-1
0
1
2
3
4
5
6
7
Figure 6. pHzpc of the fabricated composite materials (MCM-PA@B). 3.5. Zero-Point Charge (pHzpc) on the Composite
The PA@B contains a sulfonate group from the poly(AMPS) shell, which is
rated with the IONPs@SiO2. Due to the presence of these acidic sulfonate gr
pHzpc of the composite material was expected to be in the acidic range, as con
the results obtained, with a value of 2.5. Below the pHzpc value, the composite
tive charge, while increasing above this value it has negative charge (Figure 6). pH
0
2
4
6
8
10
12
14
ΔpH
-1
0
1
2
3
4
5
6
7 pH Figure 6. pHzpc of the fabricated composite materials (MCM-PA@B). Figure 6. pHzpc of the fabricated composite materials (MCM-PA@B). 11 of 20 Polymers 2022, 14, 2681 11 of 20 3.6. EDX Analysis 3.6. EDX Analysis y
Figure 7 shows the EDX analysis of the synthesized IONPs. The incorporation of
silica with the IONPs is confirmed by the presence of its peaks in Figure 7a. Moreover, the
presence of the peaks for carbon and sulfur also shows the presence of the core–shell latex
linked with IONPs@SiO2. The presence of peaks for Fe confirms the magnetic nature of all
of the prepared MCM-PA@B (Figure 7). 13 of 22 Figure 7 shows the EDX analysis of the synthesized IONPs. The incorporation of
silica with the IONPs is confirmed by the presence of its peaks in Figure 7a. Moreover, the
presence of the peaks for carbon and sulfur also shows the presence of the core–shell latex
linked with IONPs@SiO2. The presence of peaks for Fe confirms the magnetic nature of all
of the prepared MCM-PA@B (Figure 7). 13 of 22 Figure 7 shows the EDX analysis of the synthesized IONPs. The incorporation of
silica with the IONPs is confirmed by the presence of its peaks in Figure 7a. Moreover, the
presence of the peaks for carbon and sulfur also shows the presence of the core–shell latex
linked with IONPs@SiO2. The presence of peaks for Fe confirms the magnetic nature of all
of the prepared MCM-PA@B (Figure 7). 13 of 22 linked with IONPs@SiO2. The presence of peaks for Fe confirms the magnetic nature of a
of the prepared MCM-PA@B (Figure 7). Figure
7. EDX
analysis
of
the
IONPs@SiO21:PA@B1
(a),
IONPs@SiO21:PA@B3
(
IONPs@SiO21:PA@B5 (c), IONPs@SiO23:PA@B5 (d), and IONPs@SiO25:PA@B5 (e). Figure
7. EDX
analysis
of
the
IONPs@SiO21:PA@B1
(a),
IONPs@SiO21:PA@B3
(b
IONPs@SiO21:PA@B5 (c), IONPs@SiO23:PA@B5 (d), and IONPs@SiO25:PA@B5 (e). Figure
7. EDX
analysis
of
the
IONPs@SiO21:PA@B1
(a),
IONPs@SiO21:PA@B3
(b
IONPs@SiO21:PA@B5 (c), IONPs@SiO23:PA@B5 (d), and IONPs@SiO25:PA@B5 (e). Figure
7. EDX
analysis
of
the
IONPs@SiO21:PA@B1
(a),
IONPs@SiO21:PA@B3
(b)
IONPs@SiO21:PA@B5 (c), IONPs@SiO23:PA@B5 (d), and IONPs@SiO25:PA@B5 (e). 3.7. Drug Loading and Release Study
3.7. Drug Loading and Release Study 3.7. Drug Loading and Release Study
3.7. Drug Loading and Release Study The loading of the drugs aprepitant and sofosbuvir at various times, pH values, and
concentrations is given in Figures 8–10, respectively. It can be observed from the figures
that the MCM-PA@B with a lower ratio of PA@B has lower loading, while it was found to
be higher in PA@B in the MCM-PA@B. This might be due to the presence of higher attach-
ment sites (functional groups) to which the drug(s) can attach, ultimately leading to higher
drug loading. Moreover, as the amounts of IONPs@SiO2 are increased, there is no observ-
able increase in the loading capacity of the drugs, which might be due to the incorporation
of IONPs@SiO2 with the core–shell latex molecules. As shown in the figure, maximum
loading of the drugs occurs at a quantity of 229 mg/g for aprepitant and 203 mg/g for
sofosbuvir at a time of 3 h. After 3 h, there was no increase in the loading capacity of the
drug, showing that all of the attachment sites were occupied and, hence, equilibration had
occurred
The loading of the drugs aprepitant and sofosbuvir at various times, pH values, and
concentrations is given in Figures 8–10, respectively. It can be observed from the figures
that the MCM-PA@B with a lower ratio of PA@B has lower loading, while it was found to be
higher in PA@B in the MCM-PA@B. This might be due to the presence of higher attachment
sites (functional groups) to which the drug(s) can attach, ultimately leading to higher drug
loading. Moreover, as the amounts of IONPs@SiO2 are increased, there is no observable
increase in the loading capacity of the drugs, which might be due to the incorporation
of IONPs@SiO2 with the core–shell latex molecules. As shown in the figure, maximum
loading of the drugs occurs at a quantity of 229 mg/g for aprepitant and 203 mg/g for
sofosbuvir at a time of 3 h. After 3 h, there was no increase in the loading capacity of the
drug, showing that all of the attachment sites were occupied and, hence, equilibration
had occurred. 3.7. Drug Loading and Release Study
3.7. Drug Loading and Release Study 12 of 20 12 of 20 Polymers 2022, 14, 2681 in Time (min)
0
50
100
150
200
250
300
Aprepitant drug loaded (mg/g)
0
50
100
150
200
250
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
A
Time (min)
0
50
100
150
200
250
300
Sofosbuvir drug Loaded (mg/g)
0
50
100
150
200
250
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
B
Figure 8. Loading of drugs into the fabricated MCM-PA@B at different time points: aprepitant (A);
sofosbuvir (B). Time (min)
0
50
100
150
200
250
300
Aprepitant drug loaded (mg/g)
0
50
100
150
200
250
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
A Time (min)
0
50
100
150
200
250
300
Sofosbuvir drug Loaded (mg/g)
0
50
100
150
200
250
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
B Figure 8. Loading of drugs into the fabricated MCM-PA@B at different time points: aprepitant (A);
sofosbuvir (B). As previously shown in Figure 4, the synthesized composite materials have a pHzpc
of 2.5. Above this pH, MCM-PA@B has positive charge, while below this pH it has negative
charge. According to Bedi et al. [49], aprepitant has a pKa value of 9.7, while Sofosbuvir
has a pKa value of 9.3 [50]. These pKa values show that both drugs are in an anionic form
below pH 9. Maximum loading of the drugs was found to be 307 mg/g for aprepitant
and 306 mg/g for sofosbuvir at pH 4 due to the presence of opposite charges on the drugs
and composite materials. The loading capacity decreases as the pH is increased from 4 to
10, due to conversion of the drugs to a cationic form, which causes repulsion at higher
pH. The lesser amount of drug loading is attributed to the free space available for the
accommodation of drug molecules. The loading of both drugs was also studied for different concentrations of the drugs. From Figure 10 (upper), it can be observed that the loading of aprepitant reached a max-
imum of 490 mg/g for composite materials with a lower quantity of core–shells and
501 mg/g for composites with higher quantities of PA@B. The increase in the loading of
the drug occurred until 80 ppm, and then there was no significant change in the loading
amount of the drug. 3.7. Drug Loading and Release Study
3.7. Drug Loading and Release Study The loading of sofosbuvir increased to maximum values of 515 mg/g
for composite materials with lower quantities of core–shells and 525 mg/g for composites Polymers 2022, 14, 2681 13 of 20
i
(A) 13 of 20
i
(A) with higher quantities of core–shell latex. The loading of sofosbuvir into the composite
materials is given in Figure 10 (lower). with higher quantities of core shell latex. The loading of sofosbuvir into
materials is given in Figure 10 (lower). pH
0
2
4
6
8
10
12
Aprepitant drug loaded (mg/g)
0
50
100
150
200
250
300
350
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 materials is given in Figure 10 (lower). pH
0
2
4
6
8
10
12
Aprepitant drug loaded (mg/g)
0
50
100
150
200
250
300
350
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
pH
0
2
4
6
8
10
12
Sofosbuvir drug Loaded (mg/g)
50
100
150
200
250
300
350
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
Figure 9. Loading of drugs into the fabricated MCM-PA@B at different pH values:aprepita
per); sofosbuvir (lower). Figure 9. Loading of drugs into the fabricated MCM-PA@B at different pH values: aprepitant (u
sofosbuvir (lower). pH
0
2
4
6
8
10
12
Sofosbuvir drug Loaded (mg/g)
50
100
150
200
250
300
350
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 pH Figure 9. Loading of drugs into the fabricated MCM-PA@B at different pH values:aprepitant (up-
per); sofosbuvir (lower). Figure 9. Loading of drugs into the fabricated MCM-PA@B at different pH values: aprepitant (upper);
sofosbuvir (lower). 14 of 20 14 of 20 Polymers 2022, 14, 2681 Concentration (ppm)
0
20
40
60
80
100
120
Aprpitant drug Loaded (mg/g)
0
100
200
300
400
500
600
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
Concentration (ppm)
0
20
40
60
80
100
120
Sofosbuvir Loaded (mg/g)
0
100
200
300
400
500
600
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
Figure 10. Loading of drugs into the fabricated MCM-PA@B at different concentrations:
(upper); sofosbuvir (lower). Figure 10. Loading of drugs into the fabricated MCM-PA@B at different concentrations: a
(upper); sofosbuvir (lower). The release of the drugs aprepitant and sofosbuvir at various time intervals
in Figure 11. 3.7. Drug Loading and Release Study
3.7. Drug Loading and Release Study C
t
ti
(
)
0
20
40
60
80
100
120
Aprpitant drug Loaded (mg/g)
0
100
200
300
400
500
600
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 Concentration (ppm)
0
20
40
60
80
100
120
Sofosbuvir Loaded (mg/g)
0
100
200
300
400
500
600
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 Figure 10. Loading of drugs into the fabricated MCM-PA@B at different concentrations:aprepitant
(upper); sofosbuvir (lower). Figure 10. Loading of drugs into the fabricated MCM-PA@B at different concentrations: aprepitant
(upper); sofosbuvir (lower). The release of the drugs aprepitant and sofosbuvir at various time intervals is shown
in Figure 11. The release of the drugs aprepitant and sofosbuvir at various time intervals is shown
in Figure 11. 15 of 20 Polymers 2022, 14, 2681 Time (hr)
0
5
10
15
20
25
30
Drug Release(%)
0
10
20
30
40
50
60
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
Time (h)
0
5
10
15
20
25
30
Drug Release(%)
0
10
20
30
40
50
60
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5
igure 11. Controlled release study of drugs from the fabricated MCM-PA@B:aprepitant (upper);
ofosbuvir (lower). Figure 11. Controlled release study of drugs from the fabricated MCM-PA@B: aprepitant (upper);
sofosbuvir (lower). Time (hr)
0
5
10
15
20
25
30
Drug Release(%)
0
10
20
30
40
50
60
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 Time (h)
0
5
10
15
20
25
30
Drug Release(%)
0
10
20
30
40
50
60
IONPs@SiO21:PA@B1
IONPs@SiO21:PA@B3
IONPs@SiO21:PA@B5
IONPs@SiO23:PA@B5
IONPs@SiO25:PA@B5 1. Controlled release study of drugs from the fabricated MCM-PA@B:aprepitant (upper);
ir (lower). Figure 11. Controlled release study of drugs from the fabricated MCM-PA@B: aprepitant (upper);
sofosbuvir (lower). orption Kinetics Models
3.8. Adsorption Kinetics Models orption Kinetics Models
3.8. Adsorption Kinetics Models arious adsorption kinetics models were applied tothe loading of both drugs onto
ricated MCM-PA@B. Table 2A,B show the data of the kinetic model applied. As we
that first-order kinetics is concentration-dependent, first-order kinetics explains
e incorporation of drugs into the MCM-PA@B is directly dependent on the amount
drug incorporated and its saturation concentration. From the table, we can see that
maximum values of 0.92 and 0.94 for MCM-PA@B, informing us that this first-order
model is not enough to explain the loading of both drugs. Various adsorption kinetics models were applied tothe loading of both drugs onto
the fabricated MCM-PA@B. Table 2A,B show the data of the kinetic model applied. As we
know that first-order kinetics is concentration-dependent, first-order kinetics explains that
the incorporation of drugs into the MCM-PA@B is directly dependent on the amount of the
drug incorporated and its saturation concentration. From the table, we can see that R2 has
maximum values of 0.92 and 0.94 for MCM-PA@B, informing us that this first-order kinetic
model is not enough to explain the loading of both drugs. Polymers 2022, 14, 2681 16 of 20 16 of 20 Table 2. (A). Kinetic models for loading of aprepitant into the fabricated IONPs@SiO2-PA@B com-
posite materials. (B). Kinetic models for the loading of sofosbuvir into the IONPs@SiO2-PA@B
composite materials. orption Kinetics Models
3.8. Adsorption Kinetics Models (A)
IONPs@SiO21:
PA@B1
IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25:
PA@B5
qe (exp)
mg/g
193.82
193.903
194.02
194.02
194.06
Pseudo-first-order kinetic model
K1(min−1)
−0.035
−0.18
−0.042
−0.015
−0.042
qe (mg/g)
8.18
9.79
9.86
4.56
9.98
R2
0.92
0.91
0.91
0.92
0.91
Pseudo-second-order kinetic model
K2(min−1)
0.001
0.001
0.001
0.001
0.001
qe (mg/g)
197.08
197.08
197.08
197.08
197.08
R2
1
1
1
1
1
Intra-particle diffusion model
Kint
0.336
0.338
0.342
0.342
0.345
C
189.18
189.25
189.33
189.34
189.32
R2
0.901
0.898
0.897
0.893
0.898
(B)
IONPs@SiO21:
PA@B1
IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25:
PA@B5
qe (exp)
mg/g
194.29
194.43
194.53
194.57
194.57
Pseudo-first-order kinetic model
K1(min−1)
−0.027
−0.026
−0.025
−0.027
−0.029
qe (mg/g)
7.05
6.92
6.73
7.24
7.57
R2
0.93
0.94
0.94
0.93
0.93
Pseudo-second-order kinetic model
K2(min−1)
0.001
0.001
0.001
0.001
0.001
qe (mg/g)
196.07
196.07
196.07
196.07
196.07
R2
1
1
1
1
1
Intra-particle diffusion model
Kint
0.366
0.366
0.366
0.366
0.370
C
187.37
187.37
189.37
189.41
189.42
R2
0.895
0.899
0.902
0.898
0.893 The second-order kinetics equation is dependent on the loading equilibrium capacity,
which assumes that the rate of loading over the sites of attachment is proportional to the
square of unavailable sites for attachment. The rate of loading is related to the concentration
of the activated sites on the surface of MCM-PA@B. According to Table 2A,B, second-order
kinetics for loading of both drugs fits well due to its higher R2 value of 1. This high R2
value shows that in the loading of both of the drugs, the rate-controlling step could be
chemisorption. The formation of a link between the hydrogel and the drug takes place
through exchange of electrons, which is chemisorption [51]. g
g
p
The diffusion process plays a very crucial role in the loading of drugs into the com-
posite materials. According to Table 2A,B, the obtained data show good correlation for
the second-order kinetics model, but cannot explain the drug diffusion mechanism. The
values obtained initially for the loading of drugs can be used to evaluate the intra-particle
diffusion model of kinetics. Inside the active sites of MCM-PA@B, the loading of both of
the drugs may be controlled by mass transfer through pores, liquid-phase external mass
transfer, or both [52]. 3.9. Application of Equilibrium Isotherms on the Loading of Drugs
osite materials. (B). Equilibrium isotherms and their values for the loadin
Ps@SiO2 PA@B composite materials
e interaction between the drugs and composite materials was eva
therms. The loading of drugs over the uniform surface of the com Langmuir and Freundlich equilibrium isotherms were used to evaluate the loading of
drugs into the fabricated composite materials. The data for equilibrium isotherms are given
in Table 3A,B. The interaction between the drugs and composite materials was evaluated
through these isotherms. The loading of drugs over the uniform surface of the composite
materials through monolayer incorporation is shown by the Langmuir adsorption isotherm. Both of the drugs follow Langmuir isotherms, for which the R2 value is 1, showing us that
the loading of both drugs is favorable, due to the presence of many numbers of active sites
as well as the uniform surface of the composite materials. The data for the Freundlich
isotherms for both of the drugs show that the loading of the drugs does not follow them
satisfactorily, with n values of less than 1 as compared to the Langmuir isotherms [54]. s@SiO2 PA@B composite materials. 1: IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25
: PA@B5
−25
−25.31
−25.31
−25.38
216.27
215.97
216.02
215.87
0.99
0.99
0.99
0.99
−250
−256.41
−256.41
−256.41
−1.35
−1.38
−1.38
−1.38
h monolayer incorporation is shown by the Langmuir adsorption
ugs follow Langmuir isotherms, for which the R2 value is 1, show-
of both drugs is favorable, due to the presence of many numbers
s the uniform surface of the composite materials. The data for the
or both of the drugs show that the loading of the drugs does not
ly, with n values of less than 1 as compared to the Langmuir iso-
m isotherms and their values for the loading of aprepitant on the
osite materials. (B). Equilibrium isotherms and their values for the loading Table 3. (A). Equilibrium isotherms and their values for the loading of aprepitant on the IONPs@SiO2-
PA@B composite materials. (B). Equilibrium isotherms and their values for the loading of sofosbuvir
on the IONPs@SiO2-PA@B composite materials. 185.18
185.18
185.18
185.18
1
1
1
1
Ps@SiO2 PA@B composite materials. orption Kinetics Models
3.8. Adsorption Kinetics Models The loading of drugs over the uniform surface of the com 3.9. Application of Equilibrium Isotherms on the Loading of Drugs
osite materials. (B). Equilibrium isotherms and their values for the loadin
Ps@SiO2 PA@B composite materials
e interaction between the drugs and composite materials was eva
therms. The loading of drugs over the uniform surface of the com 1: IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25
PA@B5 (A)
IONPs@SiO21:
PA@B1
IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25:
PA@B5
N
−24.69
−25
−25.31
−25.31
−25.38
Kf
216.57
216.27
215.97
216.02
215.87
R2
0.99
0.99
0.99
0.99
0.99
KL (L/g)
−243.9
−250
−256.41
−256.41
−256.41
AL (L/mol)
−1.317
−1.35
−1.38
−1.38
−1.38
IONPs@SiO21:
PA@B1
IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25
: PA@B5
N
−25.83
−26.24
−26.52
−26.73
−26.80
Kf
215.42
215.07
214.78
214.58
214.53
R2
0.99
0.99
0.99
0.99
0.99
KL (L/g)
−270.27
−277.77
−277.77
−285.71
−285.71
AL
(L/mol)
−1.45
−1.5
−1.5
−1.54
−1.54
Qͦ
185.18
185.18
185.18
185.18
185.18
R2
1
1
1
1
1
185.18
185.18
185.18
185.18
185.18
R2
1
1
1
1
1
(B)
IONPs@SiO21:
PA@B1
IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25:
PA@B5
N
−25.83
−26.24
−26.52
−26.73
−26.80
Kf
215.42
215.07
214.78
214.58
214.53
R2
0.99
0.99
0.99
0.99
0.99
KL (L/g)
−270.27
−277.77
−277.77
−285.71
−285.71
AL (L/mol)
−1.45
−1.5
−1.5
−1.54
−1.54
N
−24.69
−25
−25.31
−25.31
−25.38
Kf
216.57
216.27
215.97
216.02
215.87
R2
0.99
0.99
0.99
0.99
0.99
KL (L/g)
−243.9
−250
−256.41
−256.41
−256.41
AL
(L/mol)
−1.317
−1.35
−1.38
−1.38
−1.38
Qͦ
185.18
185.18
185.18
185.18
185.18
R2
1
1
1
1
1
(B)
IONPs@SiO21:
PA@B1
IONPs@SiO21:
PA@B3
IONPs@SiO21:
PA@B5
IONPs@SiO23:
PA@B5
IONPs@SiO25
: PA@B5
N
−25.83
−26.24
−26.52
−26.73
−26.80
Kf
215.42
215.07
214.78
214.58
214.53
R2
0.99
0.99
0.99
0.99
0.99
KL (L/g)
−270.27
−277.77
−277.77
−285.71
−285.71
AL
(L/mol)
−1.45
−1.5
−1.5
−1.54
−1.54
Qͦ
185.18
185.18
185.18
185.18
185.18
R2
1
1
1
1
1
185.18
185.18
185.18
185.18
185.18
R2
1
1
1
1
1 (A) N orption Kinetics Models
3.8. Adsorption Kinetics Models The rate-determining step of the loading of drugs may be pore Polymers 2022, 14, 2681
3 9 Application
(mg/g)
R2 17 of 20 17 of 20 diffusion or film diffusion. However, we can conclude from the data in Table 2A,B that
the intra-particle diffusion step was not involved in the loading of either drug. The results
of the intra-particle diffusion models shows that three steps are involved in the loading
of drugs: In the initial step, drug molecules are loaded at quicker rate from the solution,
with active sites of composite materials on their surface [53]. During the second stage, the
drug molecules go inside the space present in composite materials at a quicker rate. In the
third stage, the drug molecules diffuse inside the composite materials through very tiny
pores—where the diffusion takes place very slowly—and reach their maximum quantity,
after which no further loading takes place. From all of these results obtained for the loading
of both of the drugs we can conclude that this kinetic model is not the rate-controlling step
for the loading of both of the drugs, for which this model does not pass through the origin. ated composite materials. The data for equilibrium isotherms are
e interaction between the drugs and composite materials was eval-
therms. The loading of drugs over the uniform surface of the com-
h monolayer incorporation is shown by the Langmuir adsorption
ugs follow Langmuir isotherms, for which the R2 value is 1, show-
of both drugs is favorable, due to the presence of many numbers
the uniform surface of the composite materials. The data for the
or both of the drugs show that the loading of the drugs does not
ly, with n values of less than 1 as compared to the Langmuir iso-
m isotherms and their values for the loading of aprepitant on the
196.07
196.07
196.07
196.07
1
1
1
1
Intra-particle diffusion model
0.366
0.366
0.366
0.370
187.37
189.37
189.41
189.42
0.899
0.902
0.898
0.893
brium Isotherms on the Loading of Drugs
undlich equilibrium isotherms were used to evaluate the loading
ated composite materials. The data for equilibrium isotherms are
i
i
b
h d
d
i
i l
l 3.9. Application of Equilibrium Isotherms on the Loading of Drugs
site materials. (B). Equilibrium isotherms and their values for the loadin
@SiO2 PA@B composite materials
interaction between the drugs and composite materials was eval
herms. 4. Conclusions In this study, IONPs@SiO2 and CS were successfully synthesized and incorporated
with one another to produce composite materials. The synthesis of these composite ma-
terials was confirmed by the FTIR, TGA, XRD, and SEM analysis techniques. From the
TGA, it was found that the composite materials with high ratios of IONPs@SiO2 were more Polymers 2022, 14, 2681 18 of 20 18 of 20 thermally stable. All of the synthesized composite materials were further applied for the
loading and release of aprepitant and sofosbuvir. Maximum loading was found at pH
4 and at a time of 2 h; moreover, this loading amount increased with the increase in the
concentration of the drugs. These loaded composite materials were further investigated for
their release at pH 7.4 and 37 ◦C. The release of the drugs reached a maximum of 50% after
24 h. From all of the data obtained, we can conclude that increasing the amount of CS and
IONPs@SiO2 in the composite materials can enhance the loading efficiency of both drugs,
while slow release of the drugs occurs in these composite materials. All of our results
show that the composite materials of IONPs@SiO2 with CS can be used as efficient drug
delivery systems. Author Contributions: Conceptualization, M.H. and N.S.; methodology, L.C.M. and T.R.; software
S.I.S. and A.K.; validation, S.I.S. and N.S.; formal analysis, K.W.G., L.C.M. and M.H.; investigation,
N.S.; resources, K.W.G. and L.C.M.; data curation, L.C.M.; writing—original draft preparation, M.H. and N.S.; writing—review and editing, L.C.M., N.S., S.I.S. and A.K.; visualization, K.W.G. and S.I.S.;
supervision, N.S.; project administration, N.S.; funding acquisition, N.S., K.W.G. and L.C.M. All
authors have read and agreed to the published version of the manuscript. Funding: This research work was fully supported by the HEC Pakistan under the NRPU project
number 20-5201, and partially supported by another project number NRPU-20-3868 awarded to
Nasrullah Shah. Acknowledgments: The authors are indebted to the R&D section of the Ferozsons Pharmaceutical
Industry, Nowshera, Pakistan for providing the standard drugs for use in the present study. Acknowledgments: The authors are indebted to the R&D section of the Ferozsons Pharmaceutical
Industry, Nowshera, Pakistan for providing the standard drugs for use in the present study. Conflicts of Interest: The authors declare no conflict of interest. Conflicts of Interest: The authors declare no conflict of interest. Conflicts of Interest: The authors declare no conflict of interest. References 1. Shah, S.A.; Majeed, A.; Rashid, K.; Awan, S.U. PEG-coated folic acid-modified superparamagnetic MnFe2O4 nanoparticles for
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Design Approaches to Improve Organic Solar Cells
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Journal of Technology Innovations in Renewable Energy, 2014, 3, 63-71 Journal of Technology Innovations in Renewable Energy, 2014, 3, 63-71 63 Design Approaches to Improve Organic Solar Cells Fahmi Fariq Muhammad* t of Physics, Faculty of Science and Health, Koya University, Koya, Kurdistan Region, Iraq Abstract: Organic solar cells (OSC)s are valuable economical and environmental friendly devices capable to generate
electricity from sunlight. This is because of their simple fabrication process and minimal material usage in comparison to
the inorganic solar cells. OSCs have not been widely commercialized yet because of their relatively low power
conversion efficiency and stability problem. It is expected that these obstacles will be surmounted in a foreseeable future
upon rigorous research studies performed in the field. This paper is devoted to reviewing the design related strategies
that can be taken to enhance OSC performance. These strategies can be accomplished through modulating the
architecture of the devices and by considering the alignment of the molecular energy levels between their active layer
components. Keywords: Organic solar cell, Nanostructure material, OSC performance, OSC efficiency, OSC improvemen Keywords: Organic solar cell, Nanostructure material, OSC performance, OSC effi *Address correspondence to this author at the Department of Physics, Faculty
of Science and Health, Koya University, Koya, Kurdistan Region, Iraq;
Tel: + 964-7701935331; E-mail: fahmi982@gmail.com 1. INTRODUCTION conjugated
polymers,
fullerenes
(C60
and
its
derivatives), and carbon nanotubes, CNTs [11-17]. However, due to their low efficiencies of about 9% [18, 19]
and stability problem [20], they are still far from being
commercialized widely, while at least 10% of stable
power conversion efficiency is required for practical
applications [21, 22]. The unsolved issues of low
efficiency, and short life time, necessitate rigorous
research studies to be performed covering from
materials analysis to the devices fabrication and
characterization. Nevertheless,
relatively
little
information is known about the exact correlation
between the devices parameters and their materials
performance, still some of the OSCs parameters are
not fully understood [23, 24]. Figure 1 shows a
prototype structure of OSC device. Indium-tin-oxide
(ITO) is normally utilized as first electrode. The most
commonly used second electrode is one of these
metals; aluminum, calcium, magnesium, or gold. The continuous demand for energy and the limiting
supply of its today’s main sources (petroleum, natural
gas, and coal) with their detrimental long term effects
on the environment necessitates a rapid development
into the clean and renewable energy sources. The only
sustainable source that can supply electric power with
its capability of protecting our environment from being
polluted is thought to be solar energy [1-3]. Solar
energy can be exploited to generate electricity by
means of solar cell devices in a technology known as
photovoltaic, PV technology. The most widely used
material in PV fabrication is crystalline silicon, c-Si
representing over 90% of the global commercial PV
module
production
in
its
various
forms
[4]. Manufacturing of inorganic based SC is expensive due
to strong requirements to the high purity of this material
and the techniques used in the devices fabrication [5]. Besides, emissions to the environment are mainly
occurred from using fossil-fuel based energy in
producing the materials for inorganic solar cells [6]. Figure 1: Prototype structure of organic solar cells. Over the last two decades great efforts have been
made to develop the low cost and ambient temperature
processing solar cells based on organic and polymeric
materials [7-10]. Despite easy fabrication of these
devices, they are able to show extra important features,
such as flexibility, light weight, and degradability for
recycling purposes. OSC is a green technology, which
is basically made from those materials that carbon
atoms are participating in their chemical structures. 2. BULK HETEROJUNCTION STRUCTURES The idea behind bilayer heterojunction, which
consists of two layers of D-A, is to use two materials
with different electron affinities, LUMOs and ionization
potentials, HOMOs [26]. By this, favorable exciton
dissociation is obtained; the electron will be accepted
by the material with the higher electron affinity while the
hole by the material with the lower ionization potential
[27]. The main drawback of this concept resides in the
rather short diffusion length of excitons (5–15 nm) [28]. Indeed, only those excitons that are created within a
distance from the D-A interface shorter than their
diffusion length may contribute to the photocurrent
generation [25]. This limits the photocurrent and hence
the overall performance of bilayer organic solar cells. To overcome this limitation, the surface area of the D-A
interface needs to be increased. This can be achieved
by creating a mixture of donor and acceptor materials
with a nanoscale phase separation resulting in a three
dimensional interpenetrating network called the bulk
heterojunction
like
shown
in
Figure
2. Bulk
heterojunction concept suggests the goal of achieving
a larger interfacial area between the electron and hole
transporting materials. The photocurrent achieved by
bulk
heterojunction
devices
are
up
to
several
milliamperes
per
square
centimeter,
improving
drastically the efficiencies over that of bilayer strucutres
[24, 29]. It has been proved that solution-processed
bulk
heterojunction
structures
with
D-A
blends
sandwiched between the anode and cathode are the One major obstacle in front of OSCs is how to make
excitons dissociate effectively into free charge carriers. Another challenge, which is common in both of organic
and inorganic solar cells, is the full collection of
photogenerated charge carriers by the correspondence
electrodes (negative and positive electrodes). In the
organic materials, especially disordered ones, carrier
mobility is several orders of magnitude smaller than
that in crystalline inorganic semiconductors. This
imposes restriction on the maximum thickness of
organic photovoltaic devices and makes them to have
very
thin
active
layers
(in
nanometers
scale). Furthermore, D-A organic components suffer from an
imbalance transport of electron and hole carriers. Accumulation of less mobile charge carriers in the
active layer will hamper charge collection at the
electrodes and thereby drastically reduce the solar cell
efficiency. Another problem is the difficulties in an
efficient harvest of excitons. Because of a large binding
energy, intrinsic dissociation of excitons into free
carriers is virtually impossible. 1. INTRODUCTION OSCs are seen to be the most plausible candidates
with
substantial
future
prospects
consisting
of Figure 1: Prototype structure of organic solar cells. It is clear that upon the absorption of light by OSCs,
excitons (bound electron-hole) are formed in the donor-
acceptor (D-A) active layer, followed by the exciton
diffusion and dissociation, which occurs at the D-A
interfaces
via
an
ultra-fast
charge
transfer. via E-ISSN: 1929-6002/14 © 2014 Lifescience Global © 2014 Lifescience Global Journal of Technology Innovations in Renewable Energy, 2014, Vol. 3, No. 2 Fahmi Fariq Muhammad 64 Subsequently, the separated free electrons and holes
transport through individual percolating pathways, and
then they are
extracted by the corresponding
electrodes. Hence, one might conclude that the device
performance can be approached by improving factors
contributing in the processes of light absorption,
exciton diffusion/dissociation, charge transport, and
charge collection. bottlenecks and hence to produce efficient OSC
fabrication. The next sections will provide an interesting
overview of these approaches. 2. BULK HETEROJUNCTION STRUCTURES Due to this, diffusion of
excitons towards either charge transfer centers or D-A
interfaces is a prerequisite for charge photogeneration
[25]. Therefore, different practical approaches are
needed
to
be
undertaken
to
overcome
these Figure 2: Bulk heterojunction structure for OSCs. Figure 2: Bulk heterojunction structure for OSCs. Journal of Technology Innovations in Renewable Energy, 2014, Vol. 3, No. 2 Design Approaches to Improve Organic Solar Cells 65 most promising alternative to realize large-scale solar
cell production [30, 31]. most promising alternative to realize large-scale solar
cell production [30, 31]. the charge transport plays a crucial role in the device
performance. As a result, the optimized thickness for
most of OSCs is less than 100 nm [13]. Apart from bulk
heterojunction structure that was discussed previously,
several materials and device structures have been
developed to obtain high short circuit current density,
Jsc. The concept of tri-layer organic p-i-n junction [34],
in which i-layer is a co-deposited layer of two different
organic semiconductors is another approach, as shown
in Figure 3a. Co-deposited layers have a vast number
of heteromolecular (donor–acceptor) contacts acting as
efficient photocarrier generation sites [28]. Additionally,
tandem structure as an effective approach to enhance
the light harvesting by means of stacking multiple cells
with complementary absorption spectra has also been
proposed (see Figure 3b). The limits to power
conversion
efficiency
and
photovoltage
can
be
breached through fabrication of tandem solar cells [35-
37]. A tandem solar cell consists of two stacked solar
cells made from materials with different optical gaps. Initially, light is absorbed by the higher-gap cell hence
the lower energy photons pass through the higher gap
device and then photons are absorbed by the second
cell. There is a conductive layer connecting the two
cells accordingly, which works as a site for charge
recombination. However, in terms of device fabrication,
there are difficult tasks such as optimizing the layers
thickness,
selection
of
suitable
materials,
and
recombination sites that have to be treated with a great
caution. Figure 3b shows a representative tandem
solar cell with two stacked cell structure having a gold
metal, Au layer as the conductive layer between the
two stacked cells to provide the recombination site. The bulk heterojunction device is similar to the
bilayer device with respect to the D-A concept, but it
exhibits a vastly increased interfacial area dispersed
throughout the bulk. 2. BULK HETEROJUNCTION STRUCTURES The bulk heterojunction requires
percolated pathways for both phases throughout the
volume, i.e., a bicontinuous and interpenetrating
network. Therefore, the nanoscale morphology in the
bulk D-A active layer is more complicated to be
controlled well in terms of the phase separation
between the donor and acceptor regions and hence
possibly the leakage current persists due to the
insufficient contact of the acceptor phase with the
cathode electrode and donor phase with the anode
electrode [32]. The bulk heterojunction devices used
today consist, in general, of a poly(4-tyrenesulfonate)
(PEDOT-PSS) covered ITO substrate, coated with a
single photoactive bulk heterojunction layer (e.g.,
polymer–fullerene mixtures) closed by a low work
function cathode such as Al [33]. 3. MULTILAYER STRUCTURES It is known that each organic material has a unique
optical band gap and a broad absorption spectrum of
sunlight is not easy to be achieved by a single organic
layer. Therefore, multiple layers and stacked tandem
cell structure, in which each layer absorbs a different
light wavelength, have been proposed to mostly
resolve the limited absorption problem. Organic
materials have higher absorption coefficient than that of
inorganic materials. Therefore, about 300 nm film is
thick enough to absorb the most incident light [13]. However, the thickness is ultimately limited by the short
exciton diffusion length and low charge carrier mobility
[17] since the balance between the light absorption and 5. DOUBLE CABLE POLYMER It was understood that the control of nanoscale
morphology in bulk heterojunction structure was not
easy. The degree of the phase separation between D-A
and domain size depend on solvent choice, speed of
evaporation, materials solubility, miscibility of the donor
and acceptor components, annealing temperature, etc. It was understood that the control of nanoscale
morphology in bulk heterojunction structure was not
easy. The degree of the phase separation between D-A
and domain size depend on solvent choice, speed of
evaporation, materials solubility, miscibility of the donor
and acceptor components, annealing temperature, etc. One strategy towards increasing the D-A phase control
is to covalently link donor and acceptor molecules to
get some sort of polymers called the double cable
polymers. Researchers [41-43] have synthesized a
pendant
fullerene
moieties
and
polythiophene
backbone
with
covalently
bound
tetracyanoan-
thraquino-dimethane, TCAQ moieties (donor–acceptor
double-cable polymer), respectively, aiming at utilizing
these structures in organic solar cells. Even though the
synthesized soluble cable polymers have shown some
promising results, the fully optimized double-cable
polymers are still not achieved. As the complexity of
the designed systems increase, the more critical it
becomes
to
optimize
design
parameters. The
realization of effective double cable polymers will bring
the D-A heterojunction at a molecular level. Figure 6a
shows a schematic representation of this system, while
Figure 6b shows an alternative approach to double
cables polymer, known as block copolymers, consisting
of donor and acceptor blocks linked side by side
together. In general, block copolymers [44, 45] are well
recognized for phase separation and ordered domains
formation, similar to those of the double cable
polymers. Stalmach et al. [46] synthesized a block
copolymer consisting of an electron acceptor block and Figure 4: An OSC device incorporating exciton blocking layer
of Alq3 between the cathode electrode and acceptor material. Figure 4: An OSC device incorporating exciton blocking layer
of Alq3 between the cathode electrode and acceptor material. Similar to the cathode interface, atoms from the
anode electrode can react with the organic material. Indium atoms from indium tin oxide, ITO electrode were
found to diffuse into the organic layer, where it acts as
trapping site for the charge carriers [27]. One strategy
that is used to minimize indium and oxygen diffusion is
to put an interfacial hole-transporting layer, such as
poly(3,4-ethylenedioxythiophene):poly(4-styrenesulfo-
nic) acid, PEDOT-PSS between ITO and the active
material. 4. EXCITON BLOCKING LAYER The active layer of bulk heterojunction structure is
sandwiched
between
the
anode
and
cathode Figure 3: The structure of (a) multilayer organic p-i-n solar cells and (b) organic tandem solar cells. Figure 3: The structure of (a) multilayer organic p-i-n solar cells and (b) organic tandem solar cells. Journal of Technology Innovations in Renewable Energy, 2014, Vol. 3, No. 2 66 Fahmi Fariq Muhammad nm) between the active layer and anode electrode, the
efficiency of the device was pronouncedly increased
[24]. electrodes, in which both donor and acceptor materials
are in direct contact with the electrodes. So it is
possible for the acceptor material to transfer electrons
to the hole-collecting anode and for the donor to
transfer holes to the electron-collecting cathode,
thereby producing a large leakage current and lowering
the performance of the device. In order to tackle these
problems, interfacial buffer layers can be inserted
between the active layer and electrodes to enhance the
collection of the photogenerated charges and to reduce
the leakage current [31]. In a study in which an exciton
blocking layer, EBL of bathocuproine, BCP was
incorporated, photovoltaic properties of devices based
on pentacene/PCBM were investigated [38]. It was
seen that a thin layer of BCP has improved the EQE
and charge carrier collection in the devices. More
recently, tris (8-hydroxyquinolinate) aluminium, Alq3
was also used between the cathode electrode and
acceptor materials [39, 40] (see Figure 4). It was
observed that this has led to increase in both efficiency
and stability of the OSC devices. Figure 5: An OSC device incorporating exciton blocking layer
of PEDOT:PSS between the anode electrode and acceptor
material. 6. ENERGY BAND ALIGNMENT The discovery of conjugated polymers stimulated
the research field of organic electronics, thereby
developing a variety of organic based devices such as,
solar cells, light-emitting diodes, field-effect transistors,
and memory devices. The emergence of the fields can
be tracked back to the mid-1970 when Shirakawa
reportedly prepared the first polymer (polyacetylene) by
accident,
subsequently
Heeger
and
MacDiarmid
discovered that the polymer would undergo an increase
in conductivity by 12 orders [47]. The most important
functionality of the organic materials is the large
polarizability of their extended -conjugated electron
systems formed by the delocalization of the pz orbitals
of the carbon atoms [14]. Due to this basic functionality,
upon the absorption of sun light, these materials can
show the photo-induced charge carriers and transport
properties by hopping process along their conjugated
backbone. Hence, most of the organic materials have
attracted considerable attention to be exploited in the
fabrication of electronic and optoelectronic devices
[48]. In particular, two types of materials are usually
selected as photovoltaic active layers in the fabrication
of OSCs. The first layer must be conductive to holes. This is referred to as donor, while the second layer is
conductive to electrons and known as acceptor. Regardless of whether the device is produced in a
bilayer or bulk heterojunction structure, it is of great
importance the HOMO and LUMO energy levels of the
D-A system are matched well to facilitate efficient Step
3:
(D A)* (D + A )*
(polarization
of
excitation: partial charge transfer) Step
3:
(D A)* (D + A )*
(polarization
of
excitation: partial charge transfer) Step
3:
(D A)* (D + A )*
(polarization
of
excitation: partial charge transfer) Step 4: (D + A )* (D•+ A•) (ion radical pair
formation) Step 4: (D + A )* (D•+ A•) (ion radical pair
formation)
Step
5:
(D•+ A•) D•+ + A•
(complete
charge
separation) Table 1 shows some representative donors and
acceptors including their HOMO and LUMO energy
levels [50, 51] with their molecular structures. Fullerenes are considered the best electron acceptors
so far. Step 1: D + A D* + A (photoexcitation of D) Step 1: D + A D* + A (photoexcitation of D) Step 1: D + A D* + A (photoexcitation of D)
Step
2:
D* + A (D A)*
(excitation
delocalized
between D and A) Step
2:
D* + A (D A)*
(excitation
delocalized
between D and A) Step
2:
D* + A (D A)*
(excitation
delocalized
between D and A) 5. DOUBLE CABLE POLYMER This layer serves to smooth out the uneven
surface of ITO and provides larger injection of holes
into the anode electrode. Figure 5 shows a device, in
which after insertion of a thin PEDOT:PSS layer (35 Journal of Technology Innovations in Renewable Energy, 2014, Vol. 3, No. 2 6 Design Approaches to Improve Organic Solar Cells 67 Figure 6: Shows (a) schematic representation of a realistic double cable polymer and (b) self-assembled layered structure of di-
block copolymers. Figure 6: Shows (a) schematic representation of a realistic double cable polymer and (b) self-assembled layered structure of di-
block copolymers. exciton generation, dissociation and charge transport
properties. The basic process of the photo-induced
charge transfer between a donor, D molecule and an
acceptor, A molecule can be described as follows [49]: an electron donor block by means of atom-transfer
radical addition, aiming to enhance the photovoltaic
efficiency of the PPV-C60 system (PPV = poly(p-
phenylenevinylene)). Since the solubility of such
complicated structures is very limited, the practical
handling for device fabrication is cumbersome [27]. 6. ENERGY BAND ALIGNMENT This is because of (i) ultrafast (~50 fs)
photoinduced charge transfer
that is happened
between the donors and fullerenes, (ii) fullerenes
exhibit high mobility, for example, C60 has shown field
effect electron mobility of up to 1 cm2 V-1 s-1 and (iii)
fullerenes show a better phase segregation in the blend
films [13, 49]. Among the organic donor materials
concerned, sexithiophene, 6T shows the highest
mobility when they are used as hole transporting layers
[52, 53]. By improving the solubility of sexithiophenes
through the addition of hexyl side chains to -
sexithiophene (6T) [54], it was possible to synthesize
,
-dihexyl-sexithiophene,
DH6T
semiconductor,
which was characterized by reasonable field-effect
mobility reaching as high as 0.1 cm2/V.s [55, 56]. Journal of Technology Innovations in Renewable Energy, 2014, Vol. 3, No. 2 Fahmi Fariq Muhammad 68 gy
gy,
,
,
q
Table 1: Examples of Donor and Acceptor Organic Materials Including their Basic Structural and Energy Levels
Information
Nomenclature
Molecular structure
HOMO (eV)
LUMO (eV)
Poly [2-methoxy-5-(2’-ethyl-hexyloxy)-
1,4-phenylene vinylene]
(a)
O
O
n
MEH-PPV
5.2
2.8
Poly (3-hexylthiophene)
(a)
S
n
P3HT
4.8
2.7
Donors
Poly[2,6-(4,4-bis-(2-ethylhexyl)-4H-
cyclopenta[2,1-b;3,4-b']dithiophene)-
alt-4,7-(2,1,3-benzothiadiazole)]
(b)
S
S
N
N
S
n
PCPDTBT
4.9
3.5
,-dihexyl-sexithiophene
(c)
S
S
S
S
S
S
C6H13
C6H13
DH6T
5.2
2.9
[6,6]-phenyl-C61 butyric acid methyl
ester (PCBM)
(b)
OMe
O
PCBM, C61
6.0
3.9
[6,6]-phenyl-C70 butyric acid methyl
ester (PCBM)
(b)
OMe
O
PCBM, C70
6.1
4.3
Tris (8-hydroxyquinolinate) aluminium
(d)
N
O
N
O
N
O
Al
Alq3
6.3
3.4
Acceptors
Tris (8-hydroxyquinolinate) gallium
(d)
N
O
N
O
N
O
Ga
Gaq3
5.8
3.0
(a)[57],
(b)[37],
(c)[51],
(d)[50]. Benefited by the high carrier mobility of DH6T and
PCBM, and their appropriate energy band alignment
various architectures of organic solar cells based on
DH6T/PCBM have been investigated [24]. Interestingly,
if a ternary bulk heterjunction structure is considered,
then the right choice of materials based on their energy able 1: Examples of Donor and Acceptor Organic Materials Including their Basic Structural and Energy Levels
Information O
O
n
MEH-PPV S
n
P3HT ,-dihexyl-sexithiophene
(c) OMe
O
PCBM, C61 OMe
O
PCBM, C70 N
O
N
O
N
O
Al
Alq3 N
O
N
O
N
O
Ga
Gaq3 (a)[57],
(b)[37],
(c)[51],
(d)[50]. DH6T/PCBM have been investigated [24]. 7. OUTLINE http://dx.doi.org/10.1016/j.jpowsour.2009.12.082 The simple and low cost fabrication process of
organic solar cells makes them attractive candidates
for
generation
of
electricity. The
performance
enhancement of these devices is of prior request by the
researchers in the field. This can be accomplished
upon rigorous research studies performed through
materials analysis to the devices fabrication and
characterization, thereby realizing valuable strategies
to improve the performance of these devices. The
architecture of the active layer plays a vital role in
defining the overall efficiency of OSCs. In this way,
various approaches can be utilized to modulate the
device
components
including
heterojunction
and
multilayer structures, insertion of excitons blocking
layers, designing the double cable or di-block
copolymers along with the optimum selection for the
HOMO and LUMO energy levels between the cells
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//d d i
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PCBM, and their appropriate energy band alignment
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Muhammad FF, Abdul Hapip AI, Sulaiman K. Study of
optoelectronic energy bands and molecular energy levels of Published on 30-05-2014 Received on 23-04-2014 Received on 23-04-2014 Accepted on 01-05-2014 DOI: http://dx.doi.org/10.6000/1929-6002.2014.03.02.4 DOI: http://dx.doi.org/10.6000/1929-6002.2014.03.02.4
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https://openalex.org/W2806551024
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https://www.e3s-conferences.org/articles/e3sconf/pdf/2018/13/e3sconf_icemee2018_02028.pdf
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English
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Experimental study on flame propagation characteristics of Hydrogen premixed gas in gas pipeline
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E3S web of conferences
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cc-by
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1 Introduction concentration, enough oxygen and ignition source. It is a
process of chemical reaction flow. The typical process of
premixed flame propagation in the constrained space
includes the ignition, the laminar combustion, the fire
perturbation, the transformation from laminar to
turbulent combustion and the final detonation. At the
beginning of the reaction, the laminar flame is unstable. The fold appears caused by the instability of the thermal
diffusion perturbation. The vortex generated to increase
the disturbance of the flame front and to prompt the
turbulent combustion, when the explosion space is
constrained or there are obstacles. Then, if the flame
continues to accelerate, the explosion occurs and even
transforms to detonation. Figure 1 gives a sketch of the
propagation process of the reaction. Flammable gas refers to the gas that can cause
combustion or explosion and release a large amount of
energy under the ignition energy excitation within the
environment
full
of
combustion
improver. The
flammable gases play a very important role in the fields
of petrochemical industry, metallurgical industry and gas
production and supply industry, such as methane and
hydrogen. By far, hydrogen is the cleanest high-energy
gas fuel, and also is the main industrial material. However, hydrogen is more explosive and more
powerful than conventional gas fuels due to the lower
ignition energy, the wider explosion limitation and the
larger laminar burning and diffusion velocity. The
hydrogen explosion accidents occur frequently. The
research on the combustion and flame propagation
characteristics of hydrogen premixed system has a great
practical significance. The propagation of gas explosion
wave, the kinetic theory of laminar flame and the
constant flame theory were discussed in the early 1950s
[1-2]. The propagations of gas explosion wave in the
closed system were also focused as follow [3-6]. But the
investigations about the hydrogen premixed system are
not enough. It has very important practical significance
and is the key of whether hydrogen can be used as a
conventional energy. Fig. 1. A sketch of the propagation process of the reaction. The following combustion chemical reaction occurs,
when premixed hydrogen is ignited. Fig. 1. A sketch of the propagation process of the reaction. Fig. 1. A sketch of the propagation process of the reaction. The following combustion chemical reaction occurs,
when premixed hydrogen is ignited. The following combustion chemical reaction occurs,
when premixed hydrogen is ignited. Experimental study on flame propagation characteristics of
Hydrogen premixed gas in gas pipeline 1Liaoning Shihua University, Fushun, Liaoning 113001, China Abstract. Hydrogen is the cleanest high-energy gas fuel, and also is the main industrial material. However,
hydrogen is more explosive and more powerful than conventional gas fuels, which restricts its application. In particular, the expansion of premixed combustion under a strong constraint is more complicated, the
reaction spreads faster. The flame propagation characteristics of premixed hydrogen/air were investigated
by experiment. The mechanism of reaction acceleration is discussed, and then the speed of the flame
propagation and the reaction pressure were tested and analysed. * Corresponding author: danzhuma@163.com 1 Introduction 2
2
2
2
2
H +0.5O +1.88N
H O+1.88N
→ In this paper, the deflagrating characteristic of
premixed hydrogen was concerned, based on the theory
of flame dynamics and pressure change. The speed of
flame propagation and the pressure fluctuation were
studied and analysed. In this paper, four different hydrogen concentrations
were considered for test, they are 13.5%, 18%, 24%,
28.5% and 38.5%. © The Authors, published by EDP Sciences. This is an open access article distributed under the terms of the Creative Commons Attribution
License 4.0 (http://creativecommons.org/licenses/by/4.0/). https://doi.org/10.1051/e3sconf/20183802028 https://doi.org/10.1051/e3sconf/20183802028 E3S Web of Conferences 38, 02028 (2018)
ICEMEE 2018 2 Hydrogen premixed combustion and
its mechanism in the constrained space The experimental setup is shown as in figure 2, which
consists of an automatically gas supply system, a high
explosive
pressure
test
system,
a
high-speed
photography system, a high-voltage ignition system and
a pressure relief channel. The simulated pipe used in the The combustion and explosion of premixed gas needs
the conditions, such as the suitable flammable gas E3S Web of Conferences 38, 02028 (2018)
ICEMEE 2018 https://doi.org/10.1051/e3sconf/20183802028 experiment was made by polymathic methacrylate tube. The inner diameter of the tube was 108 mm, with a
uniform length of 2 m. Two relief channels are set on the
side of the pipeline to ensure the safety. The ignitions
were charged by high voltage spark discharge. The
reaction pressures and the flame propagation were
recorded by the pressure sensors and high-speed camera. with the pipe wall, the acceleration is terminated into the
plane flame phase, the propagation speed of the flame
front is beginning to decline. Then the flame forward
reverses, gradually forming the tulip flame, which
increases the surface area of the flame, the flame forward
speed increases again. It can be seen from figure 4 that
the tulip flame appears at
2
H
18%
X
=
. But, at the higher Fig. 2. Schematic of the experimental setup. Fig. 2. Schematic of the experimental setup. hydrogen concentration, the premixed flame is in the
accelerated propagation period, there is no feature of
tulip flame in the length of the experimental pipeline. hydrogen concentration, the premixed flame is in the
accelerated propagation period, there is no feature of
tulip flame in the length of the experimental pipeline. Fig. 4. The flame propagation speed curves of pipelines wit
different hydrogen concentration. Fig. 2. Schematic of the experimental setup. 4.1 The speed of flame propagation of premixed
hydrogen in the pipeline The reaction wave fronts were recorded by the high-
speed camera, and then the speed of the flame
propagation could be calculated by the image processing
method. The position of flame front is calculated and the
speed of propagation is the ratio between the distance
and the time difference between two positions, as shown
in figure 3. In this paper, the speed of flame propagation
under different hydrogen volume fraction is calculated
by using the above method. y
g
Fig. 3. The sketch of the flame propagation in the pipeline. y
g Fig. 3. The sketch of the flame propagation in the pipeline. Fig. 3. The sketch of the flame propagation in the pipeline. The effect of different hydrogen volume fractions on
the speed of flame propagation was discussed. The
results are shown in figure. 4. The propagation speed of
the flame in the pipe increased with the flame front
propelling,
and
then
decreased
after
the
peak
for
2
H
13.5%
X
=
and
2
H
18%
X
=
. The speed of flame
propagation decreased with the irregular flame front. But
the decrease of the speed was not tested in the higher
hydrogen content. Fig. 4. The flame propagation speed curves of pipelines with
different hydrogen concentration. References gas washes out of the pipe. At the same time, the gas in
the pipeline continues to participate in the reaction and
consumed, the pressure drops in the pipe, resulting the
negative pressure. The negative pressure in the tube
causes the ejected gas to be sucked back into the reaction,
causing the pressure rise again, thus causing the pressure
oscillation. With the constant consumption of unburned
gas in the tube, the absolute value of the pressure
oscillation decreases and finally returns to the initial
value. 1. W. Bartknecht, Explosions: Course, Prevention,
Protection (Berlin, Springer, 1981) 2. D. Bjerketvedt, J. R. Bakke, K. Wingerden, Gas
Explosion Handbook (Journal
of
Hazardous
Materials, 1997) 3. K. K. Kuo, Principles of Combustion (Wiley-
Interscience, Switzerland, 2005) 4. Mustafa
Ilbas,
Serhat
Karyeyen,
Ismail
Ozdemir, Int. J Hydrogen energy, E 26, 1 (2015) The results of the pressure test show that the pressure
propagation of hydrogen-air in the pipeline is divided
into three stages. (1) After the input of the ignition
energy, the premixed gas is ignited, the hydrogen-air
continue to react, the pressure in the pipe continuously
accumulates to reach the first crest value. (2) Due to the
high temperature of the premixed combustion, when the
water vapor which generated by the reaction contact the
lower temperature environment to condense into droplets. This process causes the negative pressure in the pipe. And then, the unreacted hydrogen-air are sucked back to
the pipe to join in the reaction, the pressure goes up
again. This causes the pressure shock in the pipeline. (3)
The pressure amplitude reduces to the initial value with
the complete reaction of the premixed gas. The peak
pressure and the intensity of the pressure oscillation
increase with the increasing of hydrogen content, while
the time of peak pressure decrease. 5. Xianzhong Hu, Qingbo Yu, Nan Sun, Qin Qin, Int. J Hydrogen energy, E 39, 9527 (2014) 6. Hongfa
Huo,
Xingjian Wang,
Vigor
Yang,
Combustion and Flame, E 161, 3040 (2014) p
p
Fig. 5. Typical reaction pressure curve. p
p
Fig. 5. Typical reaction pressure curve. Fig. 5. Typical reaction pressure curve. 4.2 The reaction pressure The peak speed of flame propagation and the location
of the peak are all related to the hydrogen concentration. The higher the concentration of hydrogen, the more
violent the reaction is, the faster the flame spread. The
propagation speed of the premixed gas in the restricted
pipeline is closely related to the shape of the flame front. In the early stage, the flame is not restrained by the wall,
it is semi-spherical and rapidly evolves into a fingertip
flame, the surface area of the flame is accelerated by
exponential growth. When the flame comes into contact The reaction pressure was recorded by the pressure
sensor, which was lie in the front of the pipe. The typical
curves were shown in figure 5. As shown in the figure,
the reaction pressure of premixed hydrogen-air has
obvious pressure oscillation and appears negative
pressure. At the beginning of the reaction, the pressure
increases rapidly with the reaction progresses. Then the
film on the other side of the pipeline breaks, the pipe
becomes to a semi-enclosed space, part of the premixed 2 2 https://doi.org/10.1051/e3sconf/20183802028 E3S Web of Conferences 38, 02028 (2018)
ICEMEE 2018 5 Summary The combustion characteristics of premixed hydrogen is
the foundation of its safe application as a high-energy
fuel. The experiment results show that the peak speed of
flame propagation and the location of the peak depend
on the concentration of hydrogen. The higher hydrogen
concentration, the more violent the reaction is, and the
faster the flame spread. The reaction pressure develops
through three stages: the pressure rises to the first peak;
the pressure continues to oscillate and the pressure
reduces to the initial value. Acknowledgements This work was financially supported by the Scientific
Research Project of the Department of Education of
Liaoning Province (L2015303) and the Breeding
program of Liaoning Shihua University (2016[177]-19). 3 3
|
https://openalex.org/W2905290538
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https://europepmc.org/articles/pmc6343587?pdf=render
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English
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Lactobacillus Dominate in the Intestine of Atlantic Salmon Fed Dietary Probiotics
|
Frontiers in microbiology
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cc-by
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Lactobacillus Dominate in the
Intestine of Atlantic Salmon Fed
Dietary Probiotics Shruti Gupta 1, Adriána Fe ˇckaninová 2, Jep Lokesh 1, Jana Koš ˇcová 3, Mette Sørensen 1,
Jorge Fernandes 1 and Viswanath Kiron 1* Shruti Gupta 1, Adriána Fe ˇckaninová 2, Jep Lokesh 1, Jana Koš ˇcová 3, Mette Sørensen 1,
Jorge Fernandes 1 and Viswanath Kiron 1* 1 Faculty of Biosciences and Aquaculture, Nord University, Bodø, Norway, 2 Department of Food Hygiene and Technology,
University of Veterinary Medicine and Pharmacy in Košice, Košice, Slovakia, 3 Department of Microbiology and Immunology,
University of Veterinary Medicine and Pharmacy in Košice, Košice, Slovakia Probiotics, the live microbial strains incorporated as dietary supplements, are known to
provide health benefits to the host. These live microbes manipulate the gut microbial
community by suppressing the growth of certain intestinal microbes while enhancing the
establishment of some others. Lactic acid bacteria (LAB) have been widely studied as
probiotics; in this study we have elucidated the effects of two fish-derived LAB types (RII
and RIII) on the distal intestinal microbial communities of Atlantic salmon (Salmo salar). We employed high-throughput 16S rRNA gene amplicon sequencing to investigate the
bacterial communities in the distal intestinal content and mucus of Atlantic salmon fed
diets coated with the LABs or that did not have microbes included in it. Our results
show that the supplementation of the microbes shifts the intestinal microbial profile
differentially. LAB supplementation did not cause any significant alterations in the alpha
diversity of the intestinal content bacteria but RIII feeding increased the bacterial diversity
in the intestinal mucus of the fish. Beta diversity analysis revealed significant differences
between the bacterial compositions of the control and LAB-fed groups. Lactobacillus
was the dominant genus in LAB-fed fish. A few members of the phyla Tenericutes,
Proteobacteria, Actinobacteria, and Spirochaetes were also found to be abundant in the
LAB-fed groups. Furthermore, the bacterial association network analysis showed that
the co-occurrence pattern of bacteria of the three study groups were different. Dietary
probiotics can modulate the composition and interaction of the intestinal microbiota of
Atlantic salmon Edited by:
Konstantinos Papadimitriou,
Agricultural University of Athens,
Greece Reviewed by:
Atte Von Wright,
University of Eastern Finland, Finland
Carmen Wacher,
National Autonomous University of
Mexico, Mexico *Correspondence:
Viswanath Kiron
kiron.viswanath@nord.no Specialty section:
This article was submitted to
Systems Microbiology,
a section of the journal
Frontiers in Microbiology Received: 06 October 2018
Accepted: 14 December 2018
Published: 11 January 2019 Keywords: fish, Salmo salar, feed additive, probiotics, intestinal bacteria, Lactobacillus, microbiota, amplicon
sequencing ORIGINAL RESEARCH
published: 11 January 2019
doi: 10.3389/fmicb.2018.03247 Test Probiotics, Feed Type, and Design Test Probiotics, Feed Type, and Design
Two species of Lactobacillus (RII and RIII) that were previously
isolated from the intestinal content of farmed healthy juveniles
of rainbow trout (commercial fish farm–Rybárstvo PoŽehy s.r.o.,
Slovak Republic) were employed in this study. Antimicrobial
susceptibility of the microorganisms was assessed based on
the “Guidance on the assessment of bacterial susceptibility to
antimicrobials of human and veterinary importance” provided
by the European Food Safety Authority. Sensitivity or intrinsic
resistance of the isolated organisms to a recommended set of
antibiotics make them safe for use as probiotics in aquaculture. Both RII and RIII showed antagonistic activity against salmonid
pathogens Aeromonas salmonicida subsp. salmonicida CCM
1307 and Yersinia ruckeri CCM 6093 (Feˇckaninová, 2017). Furthermore, high level of tolerance to different pH, bile,
temperature, and high growth properties of the two species
were confirmed through in vitro studies (Feˇckaninová, 2017). The test probiotics were coated on commercial salmon feeds. Briefly, a pure culture of probiotic bacteria that were grown on de
Man, Rogosa and Sharpe agar (MRS) plates (HiMedia, Mumbai,
India) for 48 h were inoculated into 1,000 ml of MRS broth and
incubated for 18 h at 37◦C. The culture was centrifuged at 4,500
rpm for 20 min at 4◦C in a cooling centrifuge (Universal 320 R,
Hettich, Germany). The resulting cell pellets were washed twice
and resuspended in 30 ml of 0.9% (w/v) sterile saline. The feed
(batches of 1,800 g) was thoroughly coated with the bacterial
suspensions (Spirit Supreme, Skretting AS, Norway) using a
vacuum coater (Rotating Vacuum Coater F-6-RVC, Forberg
International AS, Norway). The bacterial counts on feeds were
∼108 cells.g−1 (RII/RIII), as determined by spread plating on
MRS agar plates and incubating for 48 h at 37◦C. The control
feeds were coated with 0.9% of sterile saline alone. The coated
feeds were stored at 4◦C until they were offered to Atlantic
salmon. Lactic acid bacteria (LAB) maintain intestinal health by
producing lactic acid that can be utilized by short-chain fatty
acids (SCFAs)-producing microorganisms. SCFAs (particularly
acetate, propionate and butyrate) contribute to host health
maintenance; for example, butyrate is used as energy source by
the intestinal epithelial cells and also have anti-inflammatory
effects on the host cells (Louis et al., 2014). Test Probiotics, Feed Type, and Design LAB that is generally
found in the GIT of endothermic animals have been extensively
investigated and their benefits have been reviewed by many
researchers (Pavan et al., 2003; Masood et al., 2011; Yang et al.,
2015; Karamese et al., 2016). The importance of fish gut-
dwelling LAB in aquaculture has been described in other reviews
(Ringø and Gatesoupe, 1998; Gatesoupe, 2008). Lactobacillus
that colonize the intestinal regions of fish are able to evoke
immune responses and impart protection against diseases (He
et al., 2017). Feeding diets supplemented with beneficial bacteria such
as LAB is being considered as an alternative approach to
control diseases in farmed fish (Martínez Cruz et al., 2012;
Feˇckaninová et al., 2017; Rodriguez-Nogales et al., 2017). Not many studies in fish have employed high-throughput
sequencing techniques to understand the changes in bacterial
communities following LAB feeding. In this study, we examined
the ability of Lactobacillus to modulate the distal intestinal
microbiota of Atlantic salmon, a farmed salmonid fish. In addition, we describe the differences in the topology
of
co-occurrence
networks
associated
with
the
intestinal
bacteria of Atlantic salmon offered feeds with and without
Lactobacillus. Citation: Gupta S, Fe ˇckaninová A, Lokesh J,
Koš ˇcová J, Sørensen M, Fernandes J
and Kiron V (2019) Lactobacillus
Dominate in the Intestine of Atlantic
Salmon Fed Dietary Probiotics. Front. Microbiol. 9:3247. doi: 10.3389/fmicb.2018.03247 The ecological community of microorganisms that reside (Marchesi and Ravel, 2015) in the
gastrointestinal tract (GIT) of an organism is referred to as the gut microbiota (Lozupone et al.,
2012). The GIT of a healthy human harbors a dense (Kelsen and Wu, 2012; Marchesi et al.,
2016) and diverse population (Lozupone et al., 2012) of commensal microorganisms, which offer
many benefits to the host, including immune homeostasis and health maintenance (Sommer and
Bäckhed, 2013). These commensal gut bacteria are also known to aid in amino-acid production The ecological community of microorganisms that reside (Marchesi and Ravel, 2015) in the
gastrointestinal tract (GIT) of an organism is referred to as the gut microbiota (Lozupone et al.,
2012). The GIT of a healthy human harbors a dense (Kelsen and Wu, 2012; Marchesi et al.,
2016) and diverse population (Lozupone et al., 2012) of commensal microorganisms, which offer
many benefits to the host, including immune homeostasis and health maintenance (Sommer and
Bäckhed, 2013). These commensal gut bacteria are also known to aid in amino-acid production January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. MATERIALS AND METHODS (Lin et al., 2017), nutrient metabolism and absorption (Morowitz
et al., 2011; Semova et al., 2012), vitamin and bioactive metabolite’
synthesis (Cummings and Macfarlane, 1997; LeBlanc et al.,
2013), and pathogen displacement (Kamada et al., 2013). An
imbalance in the gastrointestinal microbial composition can lead
to immune-mediated diseases (Petersen and Round, 2014). A
healthy gut bacterial assembly is essential for the well-being of
the host organisms including fish, the microbiome of which is
shaped by environment- and host-related factors (Wong and
Rawls, 2012; Eichmiller et al., 2016; Lokesh et al., 2018). Ethics Statements This
study
was
approved
by
the
Norwegian
Animal
Research
Authority,
FDU
(Forsøksdyrutvalget
ID-7898). Fish handling and sampling procedures were in compliance
with the description in LOVDATA. The rearing water was
treated with UV rays to remove substances that could be harmful
to the fish. Optimum values for water salinity, oxygen and
nitrogen concentration were maintained in the rearing tanks. The temperature of the fish rearing hall was kept stable during
the entire feeding experiment. Probiotics
are
“living
bacteria,”
and
when
they
are
administered as supplements in the right amount they can
confer health benefits to humans (FAO and WHO, 2006), by
targeting, among others intestinal health through stimulation
of intestinal epithelial cell proliferation and differentiation,
fortification
of
intestinal
barrier
and
immunomodulation
(Gareau et al., 2010; Thomas and Versalovic, 2010; Hemarajata
and Versalovic, 2013). Probiotics also have both direct and
indirect
effects
on
the
intestinal
microbial
composition
and diversity, and global host metabolic functions (Scott
et
al.,
2015). These
live
bacteria
produce
antimicrobial
compounds that suppress the growth of other microorganisms
and compete for their receptors and binding sites (Spinler
et al., 2008; O’Shea et al., 2012); thus altering the gut
microbiota (Collado et al., 2007). Members of the genera
Lactobacillus and Bifidobacterium are the most commonly
used probiotic organisms for humans (O’Toole and Cooney,
2008). Frontiers in Microbiology | www.frontiersin.org Collection of the Intestinal, Tank Biofilm,
and Rearing Water Samples First, the fish were euthanized using 160 mg/L of MS222 tricaine
methanesulfonate (Argent Chemical Laboratories, Redmond,
WA, USA). Thereafter, the body surface of the fish was swiped
with 70% ethanol. The fish were then dissected to aseptically
remove the GIT from the abdominal cavity. The distal intestinal
(DI) region was separated from the GIT and the content and
surface mucus samples from the DI were collected (n = 18 for
each group; 6 fish/tank) using sterile forceps and sterile glass
slides, respectively. In addition to these fish samples, we collected
environmental samples: water from the main inlet to the rearing
hall (inlet water, n = 1), water from the rearing tanks (n = 3)
and biofilm from the walls of the rearing tanks (n = 3). From
the 3 tanks of each group, one liter of rearing water was filtered
using 0.2 µm pore-size filters (Pall Corporation, Hampshire,
United Kingdom) and the filter paper was stored at −80◦C. The
biofilm samples were scraped from the walls of the 3 tanks of each
group. The fish and biofilm samples were collected in cryotubes,
snap-frozen in liquid nitrogen and stored at −80◦C. The sample abbreviations reported in this article are: (i)
fish samples–Control distal intestine content (CDC), RII distal
intestine content (RIIDC), RIII distal intestine content (RIIIDC),
Control distal intestine mucus (CDM), RII distal intestine
mucus (RIIDM), RIII distal intestine mucus (RIIIDM); (ii)
environmental samples– Control tank water (CW), RII tank
water (RIIW), RIII tank water (RIIIW), inlet water (IW), Control
tank biofilm (CB), RII tank biofilm (RIIB), RIII tank biofilm
(RIIIB). 16S rRNA Gene Sequence Data Processing
Sequence data quality check, processing and analyses: The
sequence quality of the raw reads generated from the Illumina
MiSeq machine was checked using FastQC (Andrews, 2010). The
forward reads (R1) corresponding to V3 region were employed
for subsequent analyses because they were of better quality than
the reverse reads (R2) corresponding to V4 region [Phred quality
score (Q) ≤15]. Sequence processing was performed using the
UPARSE (USEARCH version 9.2.64) software by Edgar (2013);
this step included quality filtering and operational taxonomic
units OTU clustering. FastQ files were used as the input file for
the UPARSE pipeline. The raw reads were truncated to 240 bp
and quality-filtered. The reads were truncated to remove the low-
quality base pairs at the 3
′-end and to make all samples of same
sequence length. Collection of the Intestinal, Tank Biofilm,
and Rearing Water Samples Furthermore, chimeric sequences were removed
using the UCHIME algorithm (Edgar et al., 2011). The quality-
filtered sequences were clustered into OTUs at 97% sequence
similarity level. For taxonomy prediction, we employed the
16S rRNA Ribosomal Database Project (RDP) training set with
species names v16. This RDP training set was used as a reference
database because the large 16S databases like SILVA, Greengenes,
or the full RDP database may give unreliable annotations of short
16S rRNA tags (Edgar, 2018). Taxonomic ranks were assigned
to the OTUs using the SINTAX algorithm (Edgar, 2016) using a
bootstrap cutoffvalue of 0.5. Afterwards, OTUs with a confidence
score <1 at the domain level and the OTUs belonging to the
phyla Cyanobacteria and Chlorophyta were removed to exclude
the plant-related sequences from the microbiota analysis. After Experimental Fish, Feeding Regime, and
Environmental Parameters The water flow rate, temperature,
salinity and O2 levels in the tanks were 800 L/h, 6.7–7.1◦C, 33
ppt, >85% saturation measured at the outlet, respectively. A
photoperiod of 24:0 LD was maintained throughout the feeding
trial. Experimental Fish, Feeding Regime, and
Environmental Parameters Atlantic salmon of average weight 522 ± 68 g were maintained in
800 L tanks in a flow-through seawater system, earlier described
in Sørensen et al. (2017). A 20-day feeding trial was conducted
at the research station, Nord University, Bodø, Norway. Three
groups of fish (n = 45 fish/tank; 3 replicate tanks per group) were
offered feeds with (RII ∼108 cells.g−1-RII; RIII ∼108 cells.g−1-
RIII) or without probiotics (Control—C). The fish were fed ad January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 2 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. Biosystems, Woburn, USA), 1.5 µl of each forward and reverse
primer (at a final concentration of 100 nM), 3.5 µl of DNAse
and nuclease free water (Merck, Darmstadt, Germany) and 6
µl of DNA template and/ or 6 µl of negative PCR control. The
thermocycling conditions included initial denaturation at 95◦C
for 5 min, followed by 35 cycles of denaturation at 98◦C for 30 s,
annealing at 58◦C for 30 s, extension at 72◦C for 45 s, and the
final extension performed at 72◦C for 2 min. After performing
the PCR, the resulting amplicon triplicates were pooled and
visualized on 1.2% (w/v) agarose gel stained with SYBR R⃝Safe
(Thermo Fisher Scientific, Rockford IL, USA), and the amplicon
size was compared to a 1 kb DNA ladder (Thermo Fisher
Scientific, Inc.). No amplification was observed in the negative
PCR control. Only the amplicons (∼550 bp) with clear visible
bands were selected, purified using the ZR-96 ZymocleanTM Gel
DNA Recovery Kit (Zymo Research) and eluted in 15 µl of
elution buffer. The eluted amplicon library (sequencing library)
was quantified by qPCR using the KAPA Library Quantification
Kit (KAPA Biosystems). After quantification, each amplicon
library was normalized to an equimolar concentration (3 nM)
and validated on the TapeStation (Agilent Biosystems, Santa
Clara, USA), prior to sequencing. The normalized library pool
was further diluted to 12 pM, spiked with equimolar 10% Phix
control and then paired-end sequencing was performed using
the 600 cycle v3 sequencing kit on the Illumina MiSeq Desktop
sequencer (Illumina, San Diego, CA, United States) in 2 runs
with inter-run calibrators to reduce eventual differences between
sequencing runs. libitum; the feeds were dispensed two times a day, between
08.00–09.00 and 14.00–15.00, using automatic feeders (Arvo-
Teck, Huutokoski, Finland). Frontiers in Microbiology | www.frontiersin.org DNA Extraction and PCR Amplification of
Bacterial 16S rRNA Gene for Illumina
MiSeq Amplicon Sequencing Genomic DNA was extracted from the content, mucus and
biofilm samples using the Quick-DNATM Fecal/Soil Microbe
96 kit (Zymo Research, Irvine, CA, USA) following the
manufacturer’s protocol. Metagenomic DNA Isolation kit for
water (Epicenter Biotechnologies, Madison, WI, USA) was
employed to extract the genomic DNA from the water samples. The quality of the extracted DNA was checked on 1.2% (w/v)
agarose gel. Qubit 3.0 fluorometer (Life Technologies, Carlsbad,
USA) was employed to quantify the concentration of DNA. To describe the changes in the intestinal bacteria under
the influence of LAB, we amplified the V3–V4 region of the
bacterial 16S rRNA gene employing a dual-index sequencing
strategy described by Kozich et al. (2013). The PCR reactions
were carried out in triplicates, each reaction (25 µl) volume
contained 12.5 µl of Kapa HiFi Hot Start PCR Ready Mix (KAPA January 2019 | Volume 9 | Article 3247 3 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. constructing the OTU table, the counts were rarefied to the lowest
number of sequences per sample to get an even sampling depth to
facilitate comparisons between the treatment groups. The OTU
count data was divided into 4 sets based on the sample type,
namely the DI content, DI mucus, tank water and tank biofilm
samples. The downstream analyses were performed separately on
these 4 sets. Furthermore, to ensure that we employ content and
mucus data from the same fish, only 14 fish from each group were
considered for the downstream analyses. In total 103 samples
were used for the downstream analyses, including the tank water
and biofilm samples. The raw 16S rRNA gene sequence data from
this study has been deposited in the European Nucleotide Archive
(ENA) under the accession number ERP110004. interaction networks to be sparse (Kurtz et al., 2015). In this
study, we employed the neighborhood selection (MB) method
on the sequenced 16S rRNA gene (V3 region) data of both
DI content and mucus samples to understand the community
organization. We explored the co-occurrence networks to uncover the
probable biological interactions occurring within the microbial
communities. We used the top 200 OTUs for network
construction, since it is advised to avoid extremely rare OTUs
or OTUs with a large number of zeros (Banerjee et al., 2018). The co-occurrence microbial networks were constructed and
analyzed using the functions of the R package “igraph” v1.2.1
and customized ggplot2 commands. Statistical Analysis of the Bacterial 16S
rRNA Gene Amplicon Data The differences in the DI bacterial communities of the LAB-
fed fish compared to the control fish are explained based on
the following diversity metrics: overall microbial richness (i.e.,
counts of individual OTUs), effective number of OTUs (counts of
common and dominant OTUs), taxonomic composition, relative
abundances of the bacterial taxa. Furthermore, we present
the significant and relevant bacterial communities of the DI
microbiota. We also describe the topology of the networks of the
bacterial communities in the 3 fish groups. Statistical analysis was also performed in RStudio v3.5.0. Kruskal-Wallis test followed by Dunn’s test was employed to
detect differences in alpha diversity, and we report statistically
significant differences at p < 0.05 and statistical trends at
p ≤0.15. Betadisper was used to check the assumption of
heterogeneity in dispersions; after that Adonis (PERMANOVA)
followed
by
pairwise
comparisons
was
employed
(999
permutations) to understand the significant dissimilarities
of the communities. “ANCOM” v1.1–3 (Mandal et al., 2015)
was used to detect the differentially abundant OTUs in the
treatment groups, and “Boruta” v5.3.0 R package (Kursa and
Rudnicki, 2010) was employed to find the relevant OTUs that
caused the differences in the intestinal bacteria of the three fish
groups. Differences in the Microbial Diversity and
Composition of the Intestinal and
Environmental Microbiota LAB feeding did not affect the species richness of the bacterial
community in the DI content (Figure 2A). However, this was
not the case for bacteria in the DI mucus; the species richness
was found to be higher in the mucus of the RIII-fed group (p =
0.004 for RII vs. RIII and p = 0.071 for RIII vs. C) (Figure 3A). We observed differences in the effective number of common and
dominant OTUs in the mucus of LAB-fed groups, (p = 0.109 and
p = 0.146 for RII vs. RIII; Figures 2B, 3B and Figures 2C, 3C). Comparison of the Faith’s phylogenetic diversity (PD) of the DI
content did not reveal any significant differences (Figure 2D). RESULTS We analyzed the V3 region amplicons of the 16S rRNA
gene that was sequenced on our high-throughput sequencing
platform. A total of 28,747,884 high-quality reads were clustered
into 1,823 OTUs at 97% identity threshold. These reads were
rarified based on sample-size to 12,855 reads/sample; this
allowed us to assess most of the underlying microbial diversity
(Supplementary Figures 1A,B). The feeding design, sample processing and sequencing, and
analyses are shown in Figure 1. DNA Extraction and PCR Amplification of
Bacterial 16S rRNA Gene for Illumina
MiSeq Amplicon Sequencing A network consists of a set
of vertices (commonly called as nodes) and set of edges. The
degree of a node is the number of connections it has with the
other nodes in the network. Betweenness estimates the number
of shortest paths that pass through the nodes in the network
and assortativity coefficient quantifies the extent of the selectively
connected labeled pair of nodes (Kolaczyk and Gábor, 2014). We compared the topology of the networks of the content and
mucus samples separately by analyzing the node degrees and
betweenness of the control and LAB-fed groups using Kruskal-
Wallis test followed by Dunn’s test. Analyses of microbial diversity and composition: R codes
were executed in RStudio v3.5.0 (RStudio Team, 2016) and
the functions of the R packages “iNEXT” v2.0.12 (Hsieh et al.,
2016), “phyloseq” v1.22.3 (McMurdie and Holmes, 2013) and
“ggplot2” v2.2.1 (Wickham, 2016) were used to make the
rarefaction curves for the species richness, to calculate and
visualize diversity indices, and to prepare the abundance plots. Another R package called “microbiome” v1.0.2 (Lahti et al., 2017)
was used to make core and rare microbiota (relative abundance
of core taxa) plots. Alpha diversities were calculated based on
the formula suggested by Jost (2006); for Shannon diversity
(effective number of common OTUs) and Simpson diversity
(effective number of most abundant OTUs). Beta diversity was
examined by conducting weighted UniFrac distance metric (for
fish samples)-based PCoA and double principal coordinates
analysis (DPCoA, for water and biofilm samples) (Fukuyama
et al., 2012). Microbial Network Construction and
Comparison of Topology g
We used “SPIEC-EASI” v0.1.4 R package (SParse InversE
Covariance Estimation for Ecological Association Inference) for
generating the single-domain bacterial network. SPIEC-EASI
is a statistical method that assumes the underlying microbial January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 4 Gupta et al. Lactobacilli Modulate Salmon Intestinal Microbiota FIGURE 1 | Illustration of the study design, sampling and sequence analyses. Although we did not observe any significant differences
between
the
bacterial
communities
of
the
tank
biofilm
and
the
intestinal
mucus
bacteria
of
the
LAB-fed
fish
(Supplementary Figures 3B–D,F–G),
the
biofilm
and
mucus
bacteria
of
the
control
group
were
different
(Supplementary Figure 3E, F statistic = 16.29, R2 = 0.520,
p = 0.003). For the DI mucus, differences were observed between the PD
associated with the three fish groups (p = 0.004 for RII vs. RIII
and p = 0.079 for RIII vs. C; Figure 3D). It is noteworthy that
the median alpha diversities of RII lies below the corresponding
values of C although we did not detect a trend or statistically
significant difference between the feed groups. PCoA based on
the weighted UniFrac distance matrix revealed the beta diversity
of the bacterial communities; the differences between the control
and LAB-fed groups were statistically significant (Figure 4A: F
statistic = 9.215, R2 = 0.320, p < 0.001; and Figure 4B: F statistic
= 3.114, R2 = 0.137, p < 0.002). Frontiers in Microbiology | www.frontiersin.org Intestinal Bacterial Composition Under the
Influence of LAB fujisawaense),
and
Sphingomonas were also the shared core taxa in the content
(Figure 7A). In the DI mucus, the genera that had higher
abundance in the RIII-fed fish viz. Brevinema and Pelomonas
(B. andersonii, P. saccharophila) were observed among the core
bacterial members. Photobacterium, Ralstonia, Aquabacterium,
Bradyrizhobium,
Methylobacterium,
Phyllobacterium,
(P. phosphoreum, R. pickettii, A. parvum, B. jicamae, M. fujisawaense,
P. myrsinacearum), Sphingomonas, and Mycoplasma were also
the shared core taxa of the intestinal mucus (Figure 7B). At the genus level, Lactobacilli (Lactobacillus fermentum
and Lactobacillus paraplantarum) were found to be the
most dominant bacteria in the content and mucus of LAB-
fed fish (Figure 5B, and Supplementary Figures 4B–D) and
Mycoplasma was also found to be dominant in the DI mucus of
LAB-fed fish (Figure 6B). The DPCoA indicated differences in the core members of the
LAB-fed and the control group (content: F-statistic: 3.879, R2 =
0.165, p = 0.004; mucus: F-statistic: 5.844, R2 = 0.219, p = 0.001;
Supplementary Figures 6A,B). Intestinal Bacterial Composition Under the
Influence of LAB Bacteria belonging to 23 phyla were present in the DI
content and mucus (Figures 5A, 6A). Firmicutes, Proteobacteria,
Spirochaetes, Tenericutes, and Actinobacteria were found to
be dominant in the intestine of the three study groups
(Supplementary Figures 4A,C). Firmicutes were found to be
more abundant than the rest, in both the content and mucus
of the LAB-fed fish (Figures 5A,B and Figures 6A,B). The
abundance of the phylum Tenericutes (content and mucus)
was higher in RII-fed fish, than in the RIII-fed fish group
(Figures 5A,B and Figures 6A,B). Proteobacteria (content and The beta diversity of the bacterial communities in the
rearing tank water and biofilm samples were also analyzed. The bacterial communities in the water of the 3 study groups
were
not
different
(Supplementary Figure 2A,
F-statistic
= 0.753, R2 = 0.273, p = 0.684), as was the case with
the bacteria in the biofilm (Supplementary Figure 3A, F
statistic = 0.681, R2 = 0.185, p = 0.574). On the other hand,
the bacterial communities in the water were significantly
different from those of the fish (Supplementary Figures 2B–G). January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 5 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. FIGURE 2 | Diversity of the bacterial communities of the intestinal content. Boxplots show the species richness (A), Shannon index (B), Simpson index (C), and
Faith’s phylogenetic diversity (D). The feed group codes are as follows: Control, CDC; RII, RIIDC; RIII, RIIIDC. FIGURE 2 | Diversity of the bacterial communities of the intestinal content. Boxplots show the species richness (A), Shannon index (B), Simpson index (C), and
Faith’s phylogenetic diversity (D). The feed group codes are as follows: Control, CDC; RII, RIIDC; RIII, RIIIDC. mucus) decreased in the LAB-fed groups compared to the control
group (Figures 5A,B; Figures 6A,B and Table 1). The abundance
of Spirochaetes was higher in the DI mucus of RIII-fed fish
and lower in the RII-fed fish (Figures 6A,B). The abundant
phyla in water is shown in Supplementary Figure 5A. The
dominant phyla in water were Bacteriodetes and Proteobacteria
(Supplementary Figure 5B). The changes in the abundance of
most bacterial taxa in both DI content and mucus of the LAB-fed
groups compared to the control group is shown in Table 1. Brevinema, Methylobacterium (B. jicamae, P. phosphoreum,
P. myrsinacearum,
B. andersonii,
M. Core Bacterial Communities of the
Intestinal Microbiota Significantly Abundant and Relevant
Bacterial Taxa of the Intestinal Microbiota
ANCOM analysis detected the significantly abundant bacterial
OTU in the DI content, which turned out to be L. fermentum in
RIII-fed fish (Table 1). However, this bacterium was not detected
as a significant feature in the DI mucus. We identified the core microbiota, i.e., the members of the
bacterial communities that were commonly shared among 99%
of the samples.The common core taxa–at prevalence (bacterial
community population frequency) of 99% and abundance
detection threshold of 20%–are shown in Figures 7A,B. In
the DI content, the abundant genera in the LAB-fed fish,
namely Lactobacillus, Ralstonia (L. paraplantarum, R. pickettii)
and Mycoplasma were noted to be among the core bacterial
members. Bradyrizhobium, Photobacterium, Phyllobacterium, Boruta analysis gave 9 and 8 relevant OTUs in the intestinal
content and mucus, respectively. In the DI content, L. fermentum,
L. paraplantarum, Streptococcus sobrinus, Corynebacterium
simulans, Lactococcus plantarum, W. cibaria, C. amphilecti, and
bacterial taxa belonging to Streptococcus and Xanthomonodales January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 6 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. FIGURE 3 | Diversity of the bacterial communities of the intestinal mucus. Boxplots show the species richness (A), Shannon index (B), Simpson index, (C) and Faith’s
phylogenetic diversity (D). Different letters indicate statistically significant differences (P < 0.05) between the study groups. The feed group codes are as follows:
Control, CDM; RII, RIIDM; RIII, RIIIDM. FIGURE 3 | Diversity of the bacterial communities of the intestinal mucus. Boxplots show the species richness (A), Shannon index (B), Simpson index, (C) and Faith’s
phylogenetic diversity (D). Different letters indicate statistically significant differences (P < 0.05) between the study groups. The feed group codes are as follows:
Control, CDM; RII, RIIDM; RIII, RIIIDM. assortativity (assortativity coefficient ca) of the phylum-level
network associated with the three groups (control, RII- and RIII-
fed fish) were 0.09, 0.19, and 0.10, respectively. The significantly
abundant and relevant OTUs belonged to different phyla and
modules (Figures 8A–C and Supplementary Figures 9 A–C). The degree distribution of the microbial network (for all OTUs)
of the study groups (Supplementary Figure 11A) revealed that
there are many highly connected hub nodes for the bacterial
network of the RII-fed fish and the hubs of the control group
have more node degrees. were the relevant bacteria. L. paraplantarum was found to be
abundant in the RII-fed group, whereas L. Core Bacterial Communities of the
Intestinal Microbiota fermentum and
Xanthomonodales were found to be abundant in the RIII-fed
group. S. sobrinus, C. simulans, L. plantarum, W. cibaria, C. amphilecti were reduced in abundance in the LAB-fed groups. In
the mucus, Lewinella antarctica, L. paraplantarum, L. fermentum,
Salinisphaera, Colwellia aestuarii and bacteria belonging to-
Gammaproteobacteria, Rhodobacteraceae, and Clostridiales were
found to be the relevant bacterial taxa (most of them were
abundant in the mucus of the RIII-fed fish–Table 1). DISCUSSION FIGURE 4 | Beta diversity of the intestinal bacterial communities. Principal
coordinate analysis plot (A) shows the differences in the composition of the
bacterial communities in the intestinal content (Control, CDC; RII, RIIDC; RIII,
RIIIDC). Principal coordinate analysis plot (B) shows the differences in the
composition of the bacterial communities in the intestinal mucus (Control,
CDM; RII, RIIDM; RIII, RIIIDM). Probiotics are live microbes that can impart health-benefiting
effects on host organisms. For instance, feeding of some species
belonging to genera Lactobacillus and Bifidobacterium can elicit
positive effects on host health (Wang et al., 2015; Bagarolli
et al., 2017). Probiotics alter the gut microbiota and interact
with them to produce several types of metabolites, vitamins, and
antimicrobial agents that affect the host physiology (Saulnier
et al., 2011; O’Shea et al., 2012; LeBlanc et al., 2017). In
the present study, we investigated the intestinal microbiota
changes in Atlantic salmon after feeding them with dietary
supplements of two Lactobacillus spp., named RII and RIII. To understand the differences in the microbial community
associated with the content and mucus of the DI, the bacteria in
the two samples were analyzed separately because the microbial
niche in the DI mucus is distinct compared to the intestinal
contents. Feeding LAB to the fish may facilitate their establishment
in the intestine, although significant difference was noted for
the abundance of only one of the two LAB species. The feed-
delivered organisms also altered the diversity and composition
of the DI bacteria differently. RIII supplementation caused a
significant increase in the species richness and phylogenetic
diversity of the bacterial community in DI mucus. Furthermore,
both RII and RIII caused a shift in the community composition;
bacteria belonging to different genera were altered in the two
feed groups. The co-occurrence networks indicated differential
bacterial associations in the control and LAB-fed groups. FIGURE 4 | Beta diversity of the intestinal bacterial communities. Principal
coordinate analysis plot (A) shows the differences in the composition of the
bacterial communities in the intestinal content (Control, CDC; RII, RIIDC; RIII,
RIIIDC). Principal coordinate analysis plot (B) shows the differences in the
composition of the bacterial communities in the intestinal mucus (Control,
CDM; RII, RIIDM; RIII, RIIIDM). Water bacterial communities may have an effect on the
microbiota of fish. To clarify this, we compared the microbial
community composition in the intestinal and environmental
samples. DISCUSSION Notwithstanding the fact that different extraction
methods can cause small variations in the microbial profile
(Wagner Mackenzie et al., 2015) studies have shown that
rearing water has a minor effect on the GI microbiota in fish
(Giatsis et al., 2015; Uren Webster et al., 2018). Betiku et al. (2018) have demonstrated that recirculating water systems have
more diverse microbial composition compared to the flow-
through system. However, similar to other reports (Yan et al.,
2016, Lokesh et al., 2018, Gupta et al., under review) water
bacterial communities might not have affected the intestinal
bacterial profile in our study. Also, none of the dominant
OTUs of water were detected in the DI of fish, suggesting
that host-specific gut microbial species selection is modulated
by the host gut habitat and host’s genotype (Giatsis et al.,
2015). The connectivity pattern of the significantly abundant and
relevant OTUs in the phylum-level co-occurrence network is
shown in Supplementary Figures 10 A–C. The average node
degrees were 4.12 (SD: 2.20), 2.29 (SD: 2.09), 2.74 (SD: 1.19)
for the control, RII- and RIII-fed fish, respectively. The values
for betweenness of the control, RII- and RIII-fed fish were
505 (SD: 664), 481 (SD: 596), 613 (SD: 766), respectively. Dunn’s test identified significant differences between the LAB-
fed groups, and between control and RIII-fed fish; for node
degree, but not for edge betweenness; p = 0.0002, p =
0.003 and p = 0.08, p = 0.07, respectively. The degree
of assortativity (assortativity coefficient ca) of the phylum-
level network for the three groups (control, RII- and RIII-
fed fish) were −0.01, −0.07, and 0.13, respectively. The
degree distribution of the microbial network (for all OTUs)
of the three groups is shown in Supplementary Figure 11B. The node degree histogram showed that the hubs of the
RII-fed groups have higher node degrees than the other
groups. The DI Mucus Bacteria The single-domain bacterial (SDB) network derived from the
DI content of the 3 groups comprised of one giant connected
component
(Supplementary Figure 7). The
significantly
abundant and relevant OTUs were labeled based on their
membership
in
different
modules
(Figures 8A–C). The
connectivity pattern of the significantly abundant and relevant
OTUs in the phylum-level co-occurrence network is shown
in Supplementary Figures 9 A–C. The average node degrees
were 4.27 (SD: 3.44), 3.71 (SD: 1.52), 4.06 (SD: 2.48) for the
control, RII- and RIII-fed fish, respectively. Similarly, the
values for betweenness were 370 (SD: 369), 396 (SD: 351), 388
(SD- 391). The average node degrees and betweenness of the
three groups were not significantly different. The degree of The SDB network derived from the DI mucus of the control,
RII, and RIII groups comprised of one giant connected
component
(Supplementary Figure 8). In
the
bacterial
network
of
RII-fed
fish,
we
observed
a
singleton
(C. aestuarii), a dyad (2 OTUs of Mycoplasma), and a triad (L. paraplantarum, W. cibaria, and P. piscicola) with no connection
to the main network (Supplementary Figure 8). As for the
RIII-fed group, there were 3 dyads (Sphingobacteriales +
Myxococcales, 2 OTUs of Mycoplasma, and Xanthomonadales
+ Gammaproteobacteria) with no connection to the main
network (Supplementary Figure 8). The significantly abundant
and relevant OTUs were labeled based on their membership in
different modules (Figures 9A–C). January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 7 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. Frontiers in Microbiology | www.frontiersin.org LAB Increases the Microbial Diversity in
the Intestinal Mucus Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—orchid, and Tenericutes—dark
orange LAB Increases the Microbial Diversity in
the Intestinal Mucus Corresponding to our observation on the content bacteria, a few
previous studies have also shown that LAB supplementation does
not alter the intestinal bacterial diversity (Chao1 and Shannon
diversities); in humans (Van Zanten et al., 2014) and in mice
with colon cancer (Mendes et al., 2018). On the other hand,
species richness, Shannon and Simpson diversities, and PD
of the bacteria in the DI mucus were higher in the RIII-fed g
The main results of this study are summarized in Figure 10. January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 8 Gupta et al. Lactobacilli Modulate Salmon Intestinal Microbiota FIGURE 5 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B), in the intestinal content. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from greatest to smallest, and separated by a thin black border line. Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—orchid, and Tenericutes—dark
orange. sh. In the case of mucus bacteria of RII-fed fish, we noted
li h d
(
) i
l h
di
i
d
LAB Promotes the Abundance and
D
i
f I t
ti
l L
t b
ill
d FIGURE 5 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B), in the intestinal content. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from greatest to smallest, and separated by a thin black border line. Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—orchid, and Tenericutes—dark
orange. FIGURE 5 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B), in the intestinal content. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from greatest to smallest, and separated by a thin black border line. Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—orchid, and Tenericutes—dark
orange. FIGURE 5 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B), in the intestinal content. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from greatest to smallest, and separated by a thin black border line. LAB Promotes the Abundance and
Dominance of Intestinal Lactobacillus and
Other Firmicutes fish. In the case of mucus bacteria of RII-fed fish, we noted
a slight decrease (p > 0.05) in alpha diversity compared to
the control fish. Previous studies have shown that Lactobacillus
can increase the bacterial PD in the gut of mice (Usui et al.,
2018) and weaning piglets (Zhao et al., 2016). On the contrary,
offering LAB in combination with Bifidobacterium breve and
Bifidobacterium longum did not result in greater bacterial species
diversity (Chao1, Shannon index and PD) in mice that received
antibiotics (Grazul et al., 2016). fish. In the case of mucus bacteria of RII-fed fish, we noted
a slight decrease (p > 0.05) in alpha diversity compared to
the control fish. Previous studies have shown that Lactobacillus
can increase the bacterial PD in the gut of mice (Usui et al.,
2018) and weaning piglets (Zhao et al., 2016). On the contrary,
offering LAB in combination with Bifidobacterium breve and
Bifidobacterium longum did not result in greater bacterial species
diversity (Chao1, Shannon index and PD) in mice that received
antibiotics (Grazul et al., 2016). L. paraplantarum (LP) is related to L. plantarum (Curk
et al., 1996). It was dominant in the RII-fed group and L. fermentum (LF) was found dominant in the RIII-fed group. Lactobacilli are a group of gram-positive ubiquitous LAB that
produce organic acids as end products of their metabolic
activity linked to carbohydrate fermentation (Bernardeau et al., January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org Frontiers in Microbiology | www.frontiersin.org 9 Gupta et al. Lactobacilli Modulate Salmon Intestinal Microbiota FIGURE 6 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B) in the intestinal mucus. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from highest to smallest, and separated by a thin black border line. Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—shades of orchid, and
Tenericutes—dark orange. FIGURE 6 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B) in the intestinal mucus. The height of each bar segment represents
he abundance of individual operational taxonomic units (OTUs) stacked in order from highest to smallest, and separated by a thin black border line. Frontiers in Microbiology | www.frontiersin.org LAB Promotes the Abundance and
Dominance of Intestinal Lactobacillus and
Other Firmicutes Sample type
Intestinal content
Intestinal mucus
XXXXXXXX
Taxa
Groups
RII
RIII
RII
RIII
Acidobacteria
↑
↑
↑
↑
Actinobacteria
↑
↑
↓
↓
Fusobacteria
↓
↑
↓
↓
Deinococcus-Thermus
↑
↑
↓
↑
SR1
↑
–
–
↑
Chloroflexi
↑
↑
–
–
Parcubacteria
↓
↓
↑
↑
Planctomycetes
↓
↓
↑
–
Lactobacillus fermentum
↑
↑
↓
↑
Lactobacillus paraplantarum
↑
↓
↑
↑
Colwellia aestuarii
↑
↓
↓
↑
Streptococcus sobrinus
↓
↓
↓
↑
Lewinella antarctica
↓
↑
↑
↑
Lactobacillus plantarum
↓
↓
↓
↓
Acinetobacter radioresistens
↓
↑
↓
↓
Novosphingobium sediminicola
↓
↑
↓
↓
Phyllobacterium myrsinacearum
↓
↑
↓
↓
Ralstonia pickettii
↓
↑
↓
↓
Stenotrophomonas maltophilia
↓
↑
TND
TND
Undibacterium oligocarboniphilm
↓
↑
TND
TND
Micrococcus luteus
↓
↑
↓
↓
Enterococcus cecorum
↑
–
TND
TND
Mycoplasma
↑
↓
↑
↑
Aquabacterium
↓
↓
↓
↓
Bradyrizhobium
↓
↓
↓
↓
Brevinema
↓
↓
↓
↑
Delftia
↓
↓
↓
↑
Methylobacterium
↓
↓
↓
↓
Aquabacterium parvum
↓
↓
↓
↓
Pelomonas
↓
↓
↓
↑
Photobacterium
↓
↓
↓
↓
Sphingomonas
↓
↓
↓
↓
Weissella
↓
↓
TND
TND
Brevinema andersonii
↓
↓
↓
↑
Pelomonas saccharophila
↓
↓
↓
–
Bradyrizhobium jicamae
↓
↓
↓
↓
Methylobacterium fujisawaense
↓
↓
↓
↓
Photobacterium phosphoreum
↓
↓
↓
↓
Aliivibrio logei
TND
TND
↓
↓
Caulobacter segnis
TND
TND
↓
↓
Cornybacterium
↓
↓
↓
↓
Propionibacterium acnes
TND
TND
↓
↓
Arrows indicate changes in abundance (blue arrow: increase, red arrow: decrease, bold
black line: no change, TND: taxon not dominant). L. plantarum (LbOG1 strain) exhibit in vitro antibacterial
activities against fish pathogens in Clarias gariepinus (Adenike
and Olalekan, 2009). The higher abundance of intestinal Enterococcus cecorum, was also found to be dominant
in
the
content
of
the
RII-fed
group
compared
to
the
control group (Table 1). Enterococcus spp. isolated from the
intestine of rainbow trout (Oncorhynchus mykiss) are used
as probiotics due to their antimicrobial activity against fish
pathogens (Carlos et al., 2015). The functional potential of
E. cecorum in Atlantic salmon has not yet come to light
although one particular strain is known to cause infections in
broilers (Herdt et al., 2009). Clostridiales (belonging to Firmicutes) were higher in
the mucus of salmon offered diets with RIII. LAB Promotes the Abundance and
Dominance of Intestinal Lactobacillus and
Other Firmicutes Commensal
Clostridiales are known to promote gut health by modulating
gut homeostasis and taking part in immune activation (Lopetuso
et al., 2013). LAB Favors Certain Members of
Tenericutes, Spirochaetes, and
Actinobacteria LAB significantly aided in altering the abundance of the genus
Mycoplasma (Tenericutes) and B. andersonii (Spirochaetes) in
the mucus, which are the common core members in the
DI content of Atlantic salmon (Figure 7A). Mycoplasma has
consistently been isolated from salmon intestine (Holben et al.,
2002; Zarkasi et al., 2014) and its presence as a core microbiota
suggests that it may be a commensal organism in the intestinal
ecosystem. B. andersonii has been reported in the intestinal
microbiota of flatfish, Solea senegalensis (Tapia-Paniagua et al.,
2010). Although B. andersonii is known to digest lignocellulose
and fix nitrogen in termite guts (Kudo, 2009), their functional
importance needs to be elucidated. The abundance of the genus
Micrococcus (M. luteus), a member of Actinobacteria, was higher
in the DI content of the RIII-fed group (Table 1). Though M. luteus is known to be a pathogen for rainbow trout (Salmo trutta
L.) and brown trout (Oncorhynchus mykiss) (Pkala et al., 2018)
an in vivo feeding study has suggested that they can enhance
the growth and health of Nile tilapia (Abd El-Rhman et al.,
2009). LAB Promotes the Abundance and
Dominance of Intestinal Lactobacillus and
Other Firmicutes TABLE 1 | Changes in abundances of the bacterial taxa by LAB feeding. Sample type
Intestinal content
Intestinal mucus
XXXXXXXX
Taxa
Groups
RII
RIII
RII
RIII
Acidobacteria
↑
↑
↑
↑
Actinobacteria
↑
↑
↓
↓
Fusobacteria
↓
↑
↓
↓
Deinococcus-Thermus
↑
↑
↓
↑
SR1
↑
–
–
↑
Chloroflexi
↑
↑
–
–
Parcubacteria
↓
↓
↑
↑
Planctomycetes
↓
↓
↑
–
Lactobacillus fermentum
↑
↑
↓
↑
Lactobacillus paraplantarum
↑
↓
↑
↑
Colwellia aestuarii
↑
↓
↓
↑
Streptococcus sobrinus
↓
↓
↓
↑
Lewinella antarctica
↓
↑
↑
↑
Lactobacillus plantarum
↓
↓
↓
↓
Acinetobacter radioresistens
↓
↑
↓
↓
Novosphingobium sediminicola
↓
↑
↓
↓
Phyllobacterium myrsinacearum
↓
↑
↓
↓
Ralstonia pickettii
↓
↑
↓
↓
Stenotrophomonas maltophilia
↓
↑
TND
TND
Undibacterium oligocarboniphilm
↓
↑
TND
TND
Micrococcus luteus
↓
↑
↓
↓
Enterococcus cecorum
↑
–
TND
TND
Mycoplasma
↑
↓
↑
↑
Aquabacterium
↓
↓
↓
↓
Bradyrizhobium
↓
↓
↓
↓
Brevinema
↓
↓
↓
↑
Delftia
↓
↓
↓
↑
Methylobacterium
↓
↓
↓
↓
Aquabacterium parvum
↓
↓
↓
↓
Pelomonas
↓
↓
↓
↑
Photobacterium
↓
↓
↓
↓
Sphingomonas
↓
↓
↓
↓
Weissella
↓
↓
TND
TND
Brevinema andersonii
↓
↓
↓
↑
Pelomonas saccharophila
↓
↓
↓
–
Bradyrizhobium jicamae
↓
↓
↓
↓
Methylobacterium fujisawaense
↓
↓
↓
↓
Photobacterium phosphoreum
↓
↓
↓
↓
Aliivibrio logei
TND
TND
↓
↓
Caulobacter segnis
TND
TND
↓
↓
Cornybacterium
↓
↓
↓
↓
Propionibacterium acnes
TND
TND
↓
↓
Arrows indicate changes in abundance (blue arrow: increase, red arrow: decrease, bold
black line: no change, TND: taxon not dominant). TABLE 1 | Changes in abundances of the bacterial taxa by LAB feeding. LAB Promotes the Abundance and
Dominance of Intestinal Lactobacillus and
Other Firmicutes Color codes for
he dominant genera: Proteobacteria
shades of green Spirochaetes
dark blue Firmicutes
shades of yellow Actinobacteria
shades of orchid and FIGURE 6 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B) in the intestinal mucus. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from highest to smallest, and separated by a thin black border line. Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—shades of orchid, and
Tenericutes—dark orange. FIGURE 6 | Barplots showing the abundance of all bacterial phyla (A), and dominant genera (B) in the intestinal mucus. The height of each bar segment represents
the abundance of individual operational taxonomic units (OTUs) stacked in order from highest to smallest, and separated by a thin black border line. Color codes for
the dominant genera: Proteobacteria—shades of green, Spirochaetes—dark blue, Firmicutes—shades of yellow, Actinobacteria—shades of orchid, and
Tenericutes—dark orange. acid-producing bacterial populations in mouse suffering from
colitis (Rodriguez-Nogales et al., 2017). 2006). LP is known to produce bacteriocins, which are
antimicrobial peptides produced as a defense response (Tulini
et al., 2013). A Lactobacillus isolate (LP 11-1) stimulated
the innate immune system and induced tolerance against
the pathogenicity of Pseudomonas aeruginosa in silkworm
(Nishida et al., 2017). LF has been found to restore the
expression of markers associated with the maintenance of
intestinal barrier function, and recover the SCFAs- and lactic acid-producing bacterial populations in mouse suffering from
colitis (Rodriguez-Nogales et al., 2017). Lactobacillus is part of the normal intestinal flora of fish
(Ringø et al., 1995; Spanggaard et al., 2000; Ringø and
Olsen, 2003). In zebrafish, probiotic Lactobacillus helps to
overcome infection (He et al., 2017). In Nile tilapia (Oreochromis
niloticus), LF is known to improve fish immune response
(Nwanna and Bamidele, 2014). LF (LbFF4 strain) along with Lactobacillus is part of the normal intestinal flora of fish
(Ringø et al., 1995; Spanggaard et al., 2000; Ringø and
Olsen, 2003). In zebrafish, probiotic Lactobacillus helps to
overcome infection (He et al., 2017). In Nile tilapia (Oreochromis
niloticus), LF is known to improve fish immune response
(Nwanna and Bamidele, 2014). LF (LbFF4 strain) along with January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 10 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. LAB Largely Decreased the Abundance of
Proteobacteria Proteobacteria is the most abundant phylum in many marine
and freshwater fishes (Yan et al., 2016; Lokesh et al., 2018) and
it is also known to dominate the gut microbiota of Atlantic
salmon (Gajardo et al., 2016; Lokesh et al., 2018). Therefore, it
was surprising to find this taxon in low abundance in the LAB-
fed and the control fish. A general decrease in the abundance
of intestinal Proteobacteria has also been reported in farmed
Atlantic salmon that were transferred to seawater (Rudi et al.,
2018). Taxa belonging to Proteobacteria are involved in metabolic
pathways that participate in carbon and nitrogen fixation and in
the stress response regulatory system (Vikram et al., 2016). They
are also important in the digestive process in fish (Romero et al.,
2014). P. phosphoreum, a known gut symbiont of marine fish,
helps in chitin digestion and use luciferase- reoxidize reduced
coenzymes and other molecules for metabolism (Nealson and
Hastings, 1979). N. sediminicola and P. myrsinacearum are Arrows indicate changes in abundance (blue arrow: increase, red arrow: decrease, bold
black line: no change, TND: taxon not dominant). L. plantarum (LbOG1 strain) exhibit in vitro antibacterial
activities against fish pathogens in Clarias gariepinus (Adenike
and Olalekan, 2009). The higher abundance of intestinal
Lactobacillus members and the altered bacterial abundance in the
LAB-fed fish confirms that LAB feeding can change the intestinal
microbial composition. January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 11 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. Gupta et al. FIGURE 7 | Abundance of the core bacterial taxa in the intestinal content (A) and mucus (B). Color codes: Shades of green—Proteobacteria, yellow—Firmicutes,
dark blue—Spirochaetes, and dark orange—Tenericutes. FIGURE 7 | Abundance of the core bacterial taxa in the intestinal content (A) and mucus (B). Color codes: Shades of green—Proteobacteria, yellow—Firmicutes,
dark blue—Spirochaetes, and dark orange—Tenericutes. FIGURE 7 | Abundance of the core bacterial taxa in the intestinal content (A) and mucus (B). Color codes: Shades of green—Proteobacteria, yellow—Firmicutes,
dark blue—Spirochaetes, and dark orange—Tenericutes. connectivity. The node degree histograms also communicate
interesting information about the network; the mucus bacteria
of RII-fed group had hubs with more node degree. However,
the lower average node degree and lower selective linking
of the RII-fed group indicate less interactions among the
gut bacteria. Cooperative microbial communities are known
to provide microbiome stability because of their functional
dependence. LAB Largely Decreased the Abundance of
Proteobacteria Studies have shown that the stability declines with
an increase in microbial diversity and proportion of cooperative
interactions (Coyte et al., 2015). However, higher cooperating
microbial communities can cause a runaway effect that can
collapse the competing microbial population due to over-
representation of the most stable community (McNally and
Brown, 2016). known as nitrogen-fixing bacteria (Gonzalez-Bashan et al., 2000;
Muangthong et al., 2015). On the other hand, R. pickettii
formerly known as Burkholderia pickettii has genes to biodegrade
aromatic hydrocarbons (Ryan et al., 2007). In the current and
in our recent (Gupta et al, under review) studies we found
that P. myrsinacearum and R. pickettii are part of the core
gut microbiota of Atlantic salmon; N. sediminicola was also
significantly abundant in the intestinal mucus of the fish fed
oligosaccharide. Functions of the aforementioned bacteria are
not yet reported in fish. LAB Affects the Microbial Association We inferred single-domain networks using the SPEIC-EASI
framework, and highlighted the significantly abundant and
relevant OTUs in the intestinal microbiota. For DI mucus, the
inferred SDB network for RII-fed fish showed lower overall The dyads in the mucus bacterial networks of LAB-fed
fish were different, the exception being the one constructed January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 12 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. 8 | Network graphs showing the significantly abundant and relevant OTUs of the intestinal content in different modules of the network. Bacterial networks of
(A), RII (B), and RIII (C) fish. Nodes represent OTUs and specific colors of the modules reveal the membership of the significantly abundant and relevant
e left graph shows the location of the OTUs and in the right graph, the significantly abundant and relevant OTUs that belong to the same module are shown FIGURE 8 | Network graphs showing the significantly abundant and relevant OTUs of the intestinal content in different modules of the network. Bacterial networks of
the control (A), RII (B), and RIII (C) fish. Nodes represent OTUs and specific colors of the modules reveal the membership of the significantly abundant and relevant
OTUs. The left graph shows the location of the OTUs and in the right graph, the significantly abundant and relevant OTUs that belong to the same module are shown
in callouts FIGURE 8 | Network graphs showing the significantly abundant and relevant OTUs of the intestinal content in different modules of the network. Bacterial networks of
the control (A), RII (B), and RIII (C) fish. Nodes represent OTUs and specific colors of the modules reveal the membership of the significantly abundant and relevant
OTUs. The left graph shows the location of the OTUs and in the right graph, the significantly abundant and relevant OTUs that belong to the same module are shown
in callouts. January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 13 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. ork graphs showing the significantly abundant and relevant OTUs of DI mucus in different modules based on their membership for control fish (A),
h (C). Nodes represent OTUs and specific colors of the modules reveal the membership of the significantly abundant and relevant OTUs. CONCLUSION with 2 OTUs of Mycoplasma which had higher abundance
in the RII-fed fish and lower abundance in the RIII-fed fish. This result could be suggesting that intestinal Mycoplasma in
the LAB-fed fish was not associated with other gut bacterial
communities. In the content of LAB-fed fish, most of the labeled
OTUs (except OTU 8) were existing in their respective modules
(Figures 8B, C). In the mucus of RIII-fed fish OTUs belonging
to C. aestuarii, L. paraplantarum and Clostridiales were found
to exist in one module. Clostridiales and Rhodobacteraceae,
which had same module membership in the network of
the control fish were no longer closely associated after LAB
feeding. So was the case with L. fermentum and C. aestuarii. Members affiliated to Rhodobacteraceae are known for their
denitrification properties, and Kraft et al. (2014) have shown
that Clostridiales indirectly participates in nitrate respiration
by providing fermentation substrates (e.g., acetate, formate, or
hydrogen) to Rhodobacteraceae-like denitrifiers. Our findings
suggests that the taxa belonging to the same module can be
functionally dependent but the alteration of their membership
after LAB feeding has to be further investigated. In
summary,
LAB
feeding
promoted
the
dominance
of
intestinal Lactobacillus (Firmicutes) and certain members
of the phyla Tenericutes, Spirochaetes, and Actinobacteria. Although the abundances of many members of Proteobacteria
were decreased, the phylum remained dominant in the
distal intestine of Atlantic salmon. Dietary supplementation
with the two LAB strains shifted the intestinal bacterial
community
composition. Furthermore,
the
co-occurrence
networks of the intestinal bacteria were also different for
the LAB-fed fish. Taken together, our results show that the
LAB influences the gut microbiota of Atlantic salmon. This
information will help in future studies that explore the microbial
interactions between LAB-modulated gut microbiota and the
host. AUTHOR CONTRIBUTIONS MS and VK procured the funding for the study. VK, MS, JK,
AF, and SG designed the study. JK provided the probiotics. AF
and SG conducted the feeding experiment. SG performed the 16S
rRNA sequencing studies. SG, VK, and JF analyzed the data. SG
wrote the manuscript with the guidance of VK. All authors read,
revised and approved the manuscript. The mucus bacteria of RIII-fed fish had higher species richness
and PD, and the significantly abundant and relevant OTUs
belonged to different modules. For the RIII-associated network,
2 OTUs each belonging to two modules (Rhodobacteraceae
and L. fermentum; C. aestuarii, and Clostridiales) had higher
abundances compared to the control group. In addition,
significantly
abundant
and
relevant
bacteria
had
higher
abundance in the RIII-fed fish compared to the control group. This abundance pattern does not indicate negative feedback
loops (Coyte et al., 2015). These results of bacterial networks
have to be validated through culture-based studies. LAB Affects the Microbial Association The left
ocation of the OTUs and in the right graph the significantly abundant and relevant OTUs that belong to the same module are shown in callouts. twork graphs showing the significantly abundant and relevant OTUs of DI mucus in different modules based on their membership for control fish (A) FIGURE 9 | Network graphs showing the significantly abundant and relevant OTUs of DI mucus in different modules based on their membership for control fish (A),
RII (B), and RIII fish (C). Nodes represent OTUs and specific colors of the modules reveal the membership of the significantly abundant and relevant OTUs. The left
graph shows the location of the OTUs and in the right graph the significantly abundant and relevant OTUs that belong to the same module are shown in callouts. FIGURE 9 | Network graphs showing the significantly abundant and relevant OTUs of DI mucus in different modules based on their membership for control fish (A),
RII (B), and RIII fish (C). Nodes represent OTUs and specific colors of the modules reveal the membership of the significantly abundant and relevant OTUs. The left
graph shows the location of the OTUs and in the right graph the significantly abundant and relevant OTUs that belong to the same module are shown in callouts. January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 14 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. FIGURE 10 | Illustration summarizing the salient observations of the study. DI, distal intestine; C, control group; RII and RIII, LAB-fed groups. FIGURE 10 | Illustration summarizing the salient observations of the study. DI, distal intestine; C, control group; RII and RIII, LAB-fed groups. Frontiers in Microbiology | www.frontiersin.org ACKNOWLEDGMENTS = 16.291, R2 = 0.520, P = 0.003 (E), RII intestinal mucus and RII tank water:
F-statistic = 2.934, R2 = 0.163, P = 0.051 (F), RIII intestinal mucus and RIII tank
water: F-statistic = 3.910, R2 = 0.206, P = 0.03 (G). The Lactobacillus strains employed in this study are the
property of The University of Veterinary Medicine and
Pharmacy in Košice, Košice, The Slovak Republic. We thank
Professors
Peter
Popelka
(Department
of
Food
Hygiene
and Technology) and Dagmar Mudroová (Department of
Microbiology and Immunology), The University of Veterinary
Medicine Košice for providing the microorganisms for this
study. We are thankful to Ghana Vasanth for her assistance
in sample collection, Martina Kopp for her technical help in
sequencing the libraries, and Nord University research station
stafffor their help during the period of fish sampling. Special
thanks to Bisa Saraswathy for her support in data analysis,
scientific input, helpful discussions and preparation of the
manuscript. The
authors
acknowledge
the
open
access
publication funding provided by Nord University. Supplementary Figure 3 | Double principal coordinate analysis plots showing
the beta diversity of the bacterial communities. Tank biofilm bacteria (A), Control
intestinal content and control tank biofilm: F-statistic = 2.061, R2 = 0.120, P =
0.082 (B), RII intestinal content and RII tank biofilm: F-statistic = 1.915, R2 =
0.113, P = 0.015 (C), RIII intestinal content and RIII tank biofilm: F-statistic =
4.171, R2 = 0.217, P = 0.043 (D), Control intestinal mucus and control tank
biofilm: F-statistic = 5.807, R2 = 0.1279, P = 0.002 (E), RII intestinal mucus and
RII tank biofilm: F-statistic = 1.476, R2 = 0.09, P = 0.146 (F), RIII intestinal
mucus and RIII tank biofilm: F-statistic = 2.078, R2 = 0.121, P = 0.076 (G). Supplementary Figure 4 | Barplots showing the dominant bacterial phyla and
species in the intestinal content (A,B) and mucus (C,D). Supplementary Figure 5 | Barplots showing the abundance of the bacterial
phyla (A), dominant phyla (B) in the tank water. The height of each bar segment
represents the abundance of individual operational taxonomic units (OTUs)
stacked in order from largest to smallest, and separated by a thin black border
line. Color codes: Proteobacteria—green, Bacteroidetes—light blue. Supplementary Figure 6 | DPCoA showing the differences in the composition of
the core members of the intestinal content (A) and mucus (B) samples of the
control and LAB-fed groups. FUNDING The study was undertaken as part of the project Bioteknologi–
en framtidsrettet næring (FR-274/16), funded by the Nordland
County Council, Norway. January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 15 Lactobacilli Modulate Salmon Intestinal Microbiota Gupta et al. SUPPLEMENTARY MATERIAL The Supplementary Material for this article can be found
online
at:
https://www.frontiersin.org/articles/10.3389/fmicb. 2018.03247/full#supplementary-material Supplementary Figure 7 | The single-domain network graph of the bacteria in
the intestinal content. Nodes represent different phyla shown in different colors. The three panels represent the three feed groups: Control (A), RII (B), RIII (C). Supplementary Figure 8 | The single-domain network graph of the bacteria in
the intestinal mucus. Nodes represent different phyla shown in different colors. The three panels represent the three feed groups: Control (A), RII (B), RIII (C). Supplementary Figure 1 | Sample-size-based rarefaction curves for the reads
obtained from the intestinal content (A) and mucus (B). The shaded portion
around each line represents the 95% confidence interval. Color code for the feed
groups: green lines- control, orange lines- RII, pink lines- RIII. Codes for content
samples: CDM-control, RIIDC-RII, RIIIDC-RIII. Codes for mucus samples:
CDM-control, RIIDM-RII, RIIIDM-RIII. Supplementary Figure 9 | Network association graph showing the connectivity
pattern of the significantly abundant and relevant OTUs in the intestinal content of
the Control (A), RII (B), and RIII (C) groups. Supplementary Figure 2 | Double principal coordinate analysis plots showing
the beta diversity of the bacterial communities. Tank and inlet water (A), control
intestinal content and control tank water: F-statistic = 4.035, R2 = 0.211, P =
0.01 (B), RII intestinal content and RII tank water: F-statistic = 2.375, R2 = 0.136
P = 0.07 (C), RIII intestinal content and RIII tank water: F-statistic = 5.006, R2 =
0.250, P = 0.002 (D), Control intestinal mucus and control tank water: F-statistic Supplementary Figure 2 | Double principal coordinate analysis plots showing
the beta diversity of the bacterial communities. Tank and inlet water (A), control
intestinal content and control tank water: F-statistic = 4.035, R2 = 0.211, P =
0.01 (B), RII intestinal content and RII tank water: F-statistic = 2.375, R2 = 0.136,
P = 0.07 (C), RIII intestinal content and RIII tank water: F-statistic = 5.006, R2 =
0.250, P = 0.002 (D), Control intestinal mucus and control tank water: F-statistic Supplementary Figure 10 | Network association graph showing the connectivity
pattern of the significantly abundant and relevant OTUs in the intestinal mucus of
the Control (A), RII (B), and RIII (C) groups. Supplementary Figure 11 | Histograms showing the degree distribution of the
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10034 Copyright © 2019 Gupta, Feˇckaninová, Lokesh, Košˇcová, Sørensen, Fernandes and
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Lactobacillus plantarum and Lactobacillus casei increased microbial diversity January 2019 | Volume 9 | Article 3247 Frontiers in Microbiology | www.frontiersin.org 19
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Valuation through Narrative Intelligibility
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Valuation through Narrative
Intelligibility Patrycja Kaszynska Valuation Studies 10(1) 2023: 148–166 Valuation Studies 10(1) 2023: 148–166 Abstract Abstract
Narrative intelligibility is central to making sense of valuation. Narrative
intelligibility is a framing device that combines empirical observation and
situated interaction with teleological, purpose-oriented, normative inquiry. Thus understood, narrative intelligibility provides a useful analytical frame to
explain how the phenomenon of valuation is practised. At the same time – and
on the level of research – it bridges synthetically different traditions of
thought, including actor–network theory’s descriptive accounts of valuation
practices and humanities-grounded, normative theories of value. As such,
narrative intelligibility offers a way of avoiding the alleged weakness of
overstating the agency of devices and material actors in actor–network
informed approaches, without however seeking to relocate analysis into the
‘ineffable’ realm of purely theoretical constructs, the way some humanities
scholars are said to have done. The argument shows that the humanities-
derived understanding of values, approached in terms of standards of
justification and norms of criticism, can be combined with the vernacular
concepts of valuation from actor–network theory in a way that promises a
unified research agenda going forward. Keywords: valuation; Valuation Studies; narrative; normativity; humanities;
actor–network theory Introduction If one of the ambitions of the Valuation Studies journal is to
consolidate a field of inquiry under the same name – the question
arises: what fragmented, pre-existing areas have enough proximity and
relevance to enter the gravitational pull? Perhaps an obvious answer is:
those concerned with value and with valuing. And yet, a little
paradoxically, existing accounts of value and those concerned with
valuing do not, at present, overlap in a productive way. The starting – contestable – point of this article is that the approach
to valuation represented, or perhaps over-represented, in the present
journal has not intersected in a sustained and systematic manner with
the humanities’ ‘way’ of thinking about value. As explained in the next
section, to talk about distinctive ‘schools’ is an oversimplification and
possibly a reification. And yet, if one accepts that there is a notable
distinction between descriptive and normative approaches – those
concerned with describing how valuation practices unfold on the one
hand, and those asking why they develop the way they do and how
they should be conducted on the other – one can legitimately insist
that there is a spectrum with two quite distinctive ends. Humanities
literature, in particular, of a philosophical orientation, has traditionally
aimed at articulating theories of value, and has often – but certainly
not always (cf. Dewey 1939) – been conducted in a priori terms and
based on speculative claims concerning which normative concepts
should be applied. The descriptive ‘school’ can be characterised by
empirical engagement with situated contexts and ‘devices’ used in these
contexts. The latter, actor–network theory (ANT)-inspired tradition,
downplays (Munk and Abrahamsson 2012) or eliminates outright
(Roberts 2014; Heinich 2020) the normative concerns that form the
core of the humanities approaches, its focus is on mapping ‘the
systems, devices, instruments and infrastructures that underpin various
kinds of valuation (rating, pricing, ranking, accounting, funding, and
assessing)’ (Helgesson et al. 2017: 3). This article argues that the ANT-informed valuation studies and the
humanities-grounded theories of value need one another: without any
standards of justification or critique, ANT accounts are unable to
characterise the kind of agency that is operative in valuation and thus,
to genuinely act on the recognition that ‘to describe the real is always
an ethically charged act’ (Law 2007: 17). Patrycja Kaszynska is Senior Research Fellow at University of the Arts London. © 2023 The authors
This work is licensed under a Creative
Commons Attribution 4.0 International License. https://doi.org/10.3384/VS.2001-5992.2023.10.1.148-166
Hosted by Linköping University Electronic press
http://valuationstudies.liu.se Valuation Studies
149 Introduction On the other hand,
humanities-grounded accounts of value need to appreciate the material
and the empirical dimensions highlighted by ANT – this in order to
curtail their propensity for abstraction and reification. The key contribution of this article is to present the notion of
narrative intelligibility as a meeting point for the two ‘schools’ (ends of
the spectrum) without compromising their central tenets. That is to
say, narrative intelligibility allows making the most of the socio-
material analysis of the ANT-informed approaches, while at the same Valuation through Narrative Intelligibility
150 time capitalising on the humanities to present valuation as an object
susceptible to critique and thus something that can be analysed in
normative terms. As explained in detail in what follows, narrative
intelligibility is defined here in terms of teleological continuity and
situated interaction and draws on the account of narrative developed
by MacIntyre (2007), Haraway (2016) as well as, crucially, the
Deweyan understanding of situational means-ends adjustment (Dewey
1939). The upshot of this article is a better framework for understanding
valuation which does not rest on metaphysical speculation about value
and values, nor does it reduce valuation to a multitude of patterns. This marks an approach to valuation that is both practical and
normative (Horkheimer 1972; see also Lynch and Fuhrman 1991;
Radder 1992, 2008). This is because the normativity in question is
derived from the ‘know-how’ of everyday practice which is
situationally embedded (Bohman 2004; see also Kaszynska 2021). In
suggesting that the frame of narrative intelligibility should be added to
the ‘toolbox’ of valuation studies, this article makes an important
contribution to consolidating the field that cuts across the socio-
material and humanistic analysis. The frame of narrative intelligibility
is a means of bridging the ends of the spectrum which have remained
too distant up until now. Between theories of value and valuation networks There is a long tradition of thinking about value in the humanities
across axiology, ethics, theology and moral theory, aesthetics and
theory of art, intellectual history and political philosophy and – of
special interest in this article – the theory of value as related to action
and pragmatics. Some canonical names in this lineage include:
Aristotle([340BC] Crisp 2014); Thomas Aquinas ([1265–1274] 2012);
Bentham ([1780] 1996); Kant ([1785] 2005); Marx ([1867–1883]
2010); Nietzsche ([1887] Nietzsche and Hollingdale 1989); Dewey
(1939); Arendt [1958] 2013); Habermas (2015); Foucault (1980);
Rorty (1989); Chang (1997); and Nussbaum (2009); more recently,
Srinivasan (2015); McMullin (2018). The list is necessarily selective
and partial (for one thing, it privileges philosophy over other
humanities disciplines such as literature and history) but serves well
the purpose of illustrating the historical expansiveness of thinking
about value in the humanities. i In comparison, valuation studies as a field of inquiry consolidated
around the journal of the same name established in 2013 – is in its
infancy. Of course, if valuation is understood as ‘any social ... practice ... where ... the ... value ... or values of something are
established, assessed, negotiated, provoked, maintained, constructed
and/or contested’ (Doganova et al. 2014: 87) – an interest in Valuation Studies
151 understanding these practices predates 2013. The sociology of
valuation can easily be traced to the pioneering work of Viviana A. Zelizer (1979) but has perhaps deeper roots in institutional economics
(Veblen [1899] 1973) and Weberian sociology (Weber [1922] 2019). Interest in the social embedding and determination of value has since
found articulation across the pragmatic valuation sociology (Espeland
and Stevens 2008; Stark 2011; Lamont 2012), fed into the
developments of economic sociology (Beckert and Aspers 2011;
Muniesa 2011; McFall and Ossandón 2014) and the economics of
convention (Thévenot 2001; Diaz-Bone 2011), got a stronghold in
science and technology studies (Callon and Muniesa 2005; Haywood
et al. 2014), and informed some strands of management and business
(Ramirez 1999; Chandler and Vargo 2011). There are of course individual scholars working in the sociology of
valuation who have been engaged with the journal and whose work
has presented a sustained involvement with literature, history and
literary criticism (e.g. Czarniawska 1997, 2004; Fourcade 2009, 2011). Valuation through Narrative Intelligibility
152 Valuation through Narrative Intelligibility
152 Valuation through Narrative Intelligibility
152 Guggenheim 2020). The sociology of valuation can be perhaps located
somewhere in-between in that it shows a range of different points of
emphasis, e.g. from Weber’s theoretical reconstructions to Stark’s
empirically-underpinned conceptualisations (for the purposes of this
article, the sociology of valuation will not be considered as a separate
category but rather as redistributed on the spectrum). Also, it worth
noting that there are internal differences within the ‘schools’
themselves, including differences between the ‘founding fathers’ of
ANT: Latour and Law (see Munk and Abrahamsson 2012). Setting the
question of internal unity aside, what is apparent is that the
humanities and the ANT-inspired ‘schools’ do not meet in a productive
way. This, as this article argues, results in partial accounts of valuation
and puts constraints on what the ‘schools’ can individually accomplish. The humanities ‘school’ lacks empirical credibility; the ANT valuation
studies ‘school’ is deprived of much ability to compare across different
contexts and to critique individual accounts, indeed to issue value
judgements in the traditional sense of the word (cf. Doganova et al. 2014). In other words, the ‘Normative– Descriptive’ (Radder 1998:
325) schism or, at very least, a spectrum arises – the theoretical
accounts with standards of justification are set apart from the
descriptive reports with empirical engagement. Between theories of value and valuation networks Indeed, it is worth noting that Latour himself presents an interesting
case when it comes to recognising the cogency of a perspective
spanning a plurality of different discourses in making sense of
valuation. While, on the one hand, his commitment to deflating
abstract constructs makes him suspicious of ‘values’ as construed by
the modern western epistemology, ethics and metaphysics (Latour
2013), his central argument for why We Have Never Been Modern
rests on observing the entanglement of different value registers in
practical situations (Latour 2012) and the need for resolving those as
quasi-ethical matters of concern (Latour 2004). It should also be
acknowledged that the editors of Valuation Studies did openly
recognise that there are different disciplinary ‘positions’ which raise
different questions and give different answers about how the
constructions of values and valuation are or should be studied
(Doganova et al. 2014). In this sense, the seeds of cross-disciplinarity
spanning the spectrum from ANT-informed descriptive accounts and
humanities-grounded normative theories – have been planted. And yet,
this cross-fertilisation has not been pervasive enough to benefit both
‘schools’: ANT approaches to valuation (which have been well
represented in Valuation Studies) and humanities-grounded theories
(which have not). Needless to say – as indeed observed by Helgesson
and Muniesa (2013) in the inaugural editorial address to Valuation
Studies – there is much to be gained from more collaborative
approaches. This raises the question: where and how do these
approaches meet? Oversimplifying, the humanities ‘school’ studies value and values
starting with theoretical presuppositions and concepts; the valuation
studies ‘school’ starts with material and semiotic networks and seeks
to stay ‘local’, ‘situated’ and ‘empirically descriptive’ (Law 2004; Does ANT need normativity? First, a word of explanation about what is meant by ‘normativity’,
since the word is used differently in ANT and in the humanities,
including critical theory. To be normative in the sense of Annemarie
Mol (Mol 1999) or John Law (Law 2004) is to make deliberate
choices: to foreclose some alternatives. To be normative in the Critical
Theory sense is to pay attention to how actors manifest the normative
attitudes implicit in their practical knowledge (Geuss 1981;
Horkheimer 1993; Habermas 2015) and in the everyday ‘know-how’
of human conduct (Bohman 2004). What is sometimes not fully
understood is that normativity in the critical theory sense is not about
proscribing or fixing courses of action, nor even about setting rules. Rather, it is based on capitalising on the fact that action coordination
presupposes that some standards of criticism and justification are
applied and can be collectively ascertained (Kaszynska 2021). pp
y
y
ANT designates a method rather than a theory but it signals a
predilection for a certain mode of inquiry. As John Law put it back in
1992, ‘actor–network theory almost always approaches its tasks
empirically’ (Law 1992: 6). This often translates into the suspicion of
‘occult entities’ and the preference for the ontological ‘desert
landscapes’, to borrow expressions from the philosopher Willard Van
Orman Quine (1948). In other words, the default position of the ANT Valuation Studies
153 sympathisers is a commitment to flatter or flat ‘ontologies’ in the sense
suggested by Quine and notably, in the opposite sense to that intended
by Bhaskar (1978) who first coined the term. The emphasis in this tradition is very much on the mechanism – the
nuts and bolts – of valuation, such as: ‘market technologies’ (Muniesa
2007), ‘political technology’ (Muniesa and Doganova 2020) and other
kinds of ‘the systems, devices, instruments and infrastructures that
underpin various kinds of valuation’ (Helgesson et al. 2017: 3: see also
Orlikowski and Scott 2014). What this amounts to in practice is
studies preoccupied with detailed description of the formation of
socio-material-semantic networks, attentive to contexts and devices as
a way of mapping out the trajectories along which valuation unfolds. Does ANT need normativity? Even though there is a growing interest in generalising from specific
circumstances and in understanding the conditions of critique for the
systems in question (see for instance Muniesa 2017, 2019; Muniesa
and Doganova 2020; also Latour 2004, 2012), the main emphasis
remains on mapping the actants and the circumstances in which they
act. With reference to the Valuation Studies journal specifically, this
can be exemplified by the influential study of the role of algorithmic
valuation devices and verification mechanisms in the tourism and
travel sector by Baka (2015). Forseth et al.’s account of ‘Reactivity and
Resistance to Evaluation Devices’ (2019) in the banking sector is
another example. These ANT-informed approaches are now well established in
valuation studies – and indeed Valuation Studies – but with this
success comes a growing recognition of potential limitations. One such
an indication is the reaction against, or perhaps a growing self-
awareness of, the excessive concentration on the ‘mechanics’ of
valuation (Zuiderent-Jerak and van Egmond 2015). As Hauge points
out, ‘while the many studies of valuation practices have drawn
attention to the pervasive effects of valuation devices, only a few
studies have taken into account the fact that many spaces, including
organizations, are already flooded with practices and ideas that
constitute what is valuable’ (Hauge 2016: 117). (
g
)
Some commentators sympathetic to ANT were explicit about the no
absence of normative considerations and standards of critique (Radder
1992; Marres 2009). An overt concern about ‘flat-land of relativism’
was noted in the editorial address in Valuation Studies back in 2014
(Doganova et al. 2014). The root can be traced back to the early
1990s. The ‘Epistemological Chicken’ controversy involved some of
the prominent figures such as Latour and Callon (Fuller 1991; Lynch
and Fuhrman 1991; Redder 1992) and centred on the claim made by
Collins and Yearley (2010) that the ANT method leads to an infinite
regress because it is unable to fix any parameters of the debate as
standards of comparison and criticism. If there is nothing beyond the
shifting networks of relationships and all actors and actants have the Valuation through Narrative Intelligibility
154 same status – questions such as: what motivates agency in valuation
and what gives it direction? what binds the discrete moments of action
into a valuation situation? and above all, which courses of action are
to be preferred over others? – are difficult to settle. Does ANT need normativity? It should be noted
that Latour’s talk of ‘ontopolitics’ (Latour 2012) and Mol’s
‘ontological politics’ (Mol 1999) reveal that these issues are cogent
even within the ANT discourse. At some risk of overstating the case,
the need to ponder politics in ANT is an admission that the ethical
choices have not gone away. As Law puts it: ‘There is nowhere to hide
beyond the performativity of the webs’: But since our own stories weave further webs, it is never the case that they
simply describe. They too enact realities and versions of the better and the
worse, the right and the wrong, the appealing and the unappealing. There is
no innocence. The good is being done as well as the epistemological and the
ontological. (Law 2007: 16). Think of the existence of racism and racial oppression in the USA. There are multiple accounts – many told by the oppressors, and few
told by the oppressed. The current situation is precarious because of
the polarisation fuelled by culture wars. As Charles Mills (2014)
argues in The Racial Contract, the one constant is that racial contract
is continually being reformatted. Once upon a time it was manifested
as physical enslavement; later, more intangible forms of segregation
and inequality were proposed; now, it is primarily based on immaterial
forms of discrimination by institutional and cultural norms which
maintain the same racialised hierarchy. In Mills’s (2014) opinion, not
much has changed but the reality needs to change. How to achieve
this? One way of handling this is to encourage some open-ended plurality
of ‘webspinning’ without arbitration (cf. Stark 2011). But is this
enough? Surely, justice cannot be achieved just by pitting the opposing
factions against one another the way ANT might recommend. But how
can this prevent the ‘might is right’ scenario? If, in the context of racial
oppression, one finds oneself confronted with the question: What am I
to do here and now? – it is far from obvious that the ‘laissez-faire’
attitude of ANT has a satisfactory answer. The point is that there is ‘no
innocence’: there is the right and the wrong way of acting when it
comes to racial oppression. Does ANT need normativity? pp
Admittedly, as noted above, there have been some attempts in ANT
to offer ‘embedded’ accounts of normativity and to answer the ‘where
to’ question from within the contexts, networks and situations
themselves: be they Situated Intervention and the Ethics of Specificity
(Zuiderent-Jerak 2015), or Care in Practice (Mol et al. 2015), or
Latour’s ‘ontopolitics’ (Latour 2012). These attempts however do not
offer satisfactory answers on a scale capable of changing collective Valuation Studies
155 behaviours. As the next section discusses, it is by relocating the
discussion from the realm of causes and patterns (the how) to the
realm of purposes (the why), that the humanities opens up the
possibility of normative analysis to ANT. This however does not need
to mean betraying the instincts of ANT to stay situated and grounded. Narrative intelligibility is introduced as a meeting point for empirical
social inquiry and normative philosophical argumentation. The notion
of narrative intelligibility is presented as a frame which makes it
possible to cross-fertilise the ANT-informed studies of valuing and the
humanities tradition of thinking about value. Narrative and narrative intelligibility Some have taken MacIntyre to be making a metaphysical claim that
individuals are narrative constructs; others understood him as saying
that narratives are useful as reflective prisms for people to look at their
life (Kupfer 2014; cf. Taylor 2016). Resolving this is not necessary for
the purposes of this article. What matters is that MacIntyre defines
narrative as a form that has a beginning, a middle and an ending, and
crucially, an internal coherence that is instrumentally useful in making
sense of valuation situations. Importantly, what is required for
narrative coherence is more than just being a sequence. The chain of
events has to unfold in a certain purpose- oriented way with parts
interpreted in the light of the whole sequence and with the whole
defined by its teleological orientation. To rephrase, what is at issue is
not just a mechanical pattern where one thing causes the next (cf. Aristotle’s Poetics in Janko 1987) but that events unfold according to a
unifying purpose. Some may at this point be alarmed by the prospect of resurrecting
‘occult entities’ such as the notion of ‘divine causes’, ‘absolute spirits’
and other ontological ‘pathologies’ that ANT has set to abolish. This
alarm would be premature. This can be demonstrated by a quick look
at how narrative – as a form that has a beginning, a middle and an
ending, and crucially, some internal coherence – can be found in
Donna Haraway’s work. Haraway’s account of the power of
‘fabulating’ in the influential Staying with the Trouble (2016) presents
an account not dissimilar to MacIntyre’s. There Haraway sets out a
programme which uses telling stories as a practice for generating
imaginative patterns, something on which she elaborates with the
example of string figures which can be transformed over and over
again. Each moment of transformation is underpinned by its unique
teleological drive. Each stringing moment has some – arguably
purposive, even if implicit – unity, even though there is no overarching,
universal purpose. To be clear, Haraway is concerned with the material
semiotics (Haraway 2013). Even though she agrees that ‘for a material
semiotics teleology may not reside in human intentions’ (Law 2007:
10) she retains the idea of teleology as the means-ends adjustment for
every discrete moment of the string transformation. Narrative and narrative intelligibility Starting in the 1980s, there has been a growing interest in narratives
across a wide range of disciplines. A number of publications set out to
examine the use of narrative structures in different domains: history,
literature, psychology and social science (Polkinghorne 1988) and
literature, philosophy and science (Nash 2005 [1994]; see also Brosch
and Sander 2015) to name but some. The interest in narratives as a
subject matter has gone hand-in-hand with the development of
narrative-based methodologies: narrative inquiry and narrative
analysis (Mishler 1990; Plummer 2001; Riessman 2008). What has
become known as the ‘narrative turn’ is usefully summarised by
Hinchman and Hinchman (1997) as a rejection, among other things,
of the atomistic understanding of social phenomena and the dis-
embedding of individual acts from the ‘web of communication’ out of
which they arise (Hinchman and Hinchman 1997: xiv). Narratives in
this context are presented as a means of retrieving the plurality of
stories that cultures and subcultures tell without submerging them into
some uniform meta-narrative, while at the same time not giving up on
the ambition of finding a common communicative ground. Not
surprisingly, a number of interesting scholars have been developing
different forms of engagement with narratives in relation to valuation
studies (e.g. Czarniawska 1997, 2004; Smith 1988; Fourcade 2009,
2011). More recently, Beckert and Bronk (2018) revived the idea of
narrative as a coordinating device that can be used to deal with the
conditions of radical uncertainty when a stabilising of expectations is
needed. Alasdair MacIntyre was one of the first to take an interest in
the role of narrative in valuation. In broader terms, the key argument of After Virtue can be
reconstructed as saying that without a minimal narrative structure,
lives and actions of individuals will lack intelligibility. Actions and
events need to be formulated as episodes in a narrative to have
meaning. In other words, moral discourse has been cut adrift from the
narrative understanding of purposes, and with this, it was rendered
incoherent. This is a problem because: ‘I can only answer the question, Valuation through Narrative Intelligibility
156 “What am I to do?” if I can answer the prior question, “Of what story
or stories do I find myself a part?’’’(MacIntyre 2007 [1981]: 216) – as
MacIntyre put it. Narrative and narrative intelligibility To be fair, in a
similar vein, MacIntyre recognises that narratives do not occur in a
vacuum, rather they are historically and geographically situated. In this
context, MacIntyre speaks evocatively that ‘we enter upon a stage
which we did not design and we find ourselves part of an action that
was not of our making. Each of us being the main character in his own
drama plays subordinate parts in the dramas of others, and each
drama constrains the others’ (2007 [1981]: 49). Even though the
ontology of socio-material and technological networks does not figure Valuation Studies
157 in MacIntyre’s vocabulary, he admits that narratives are situated and
contextually circumscribed. Taking MacIntyre’s and Haraway’s work together, it can be
suggested that narrative intelligibility is where socio-material networks
and teleology meet (cf. Bevir and Galisanka 2016). That is to say, to
make a narrative intelligible both are needed: an understanding of a
sequence as internally coherent and so directed towards a goal and
also an understanding of how it is situated in a specific context. Accordingly, the notion of narrative intelligibility is defined in this
article in terms of both: goal-oriented, teleological continuity and
contextually circumscribed interaction. But, what kind of entity is
narrative intelligibility? g
y
Narrative intelligibility could be thought a framing construct, to
borrow a term from communication and media studies (cf. Clarke
2003). The framing at issue should not be understood as a
phenomenon known as agenda-setting (Kuypers 2010) nor is it an
attempt to forge uniformity through appealing to some set of universal
values (Lakoff 2014). Rather, the frame is here understood in the sense
presupposed by Kenneth Burke’s concept of terministic screens. Terministic screens are filters for photographs which accentuate
different features of photographs. So, applying different screens to the
same photograph may make different features apparent and prompt
different interpretations. In the same way, Burke (1966) argued that
individuals can represent issues based on the choice of some aspects
fixing their attention – whether consciously or unconsciously – which
they emphasise in language. Narrative intelligibility is a way of
filtering valuation situations through the prism of the chosen goals as
they become adjusted to the available means. This underscores that
what is at issue with narrative intelligibility is not some universalistic
grand narrative. Far from it as the next section demonstrates turning
to a quintessentially Deweyan understanding of valuation. Situational interaction and teleological continuity in
Dewey Dewey approaches valuation as a type of inquiry. Inquiry for Dewey
is a social process that follows the following six steps: 1. Identify the
problem 2. Plan possible solutions 3. Evaluate and test the various
solutions 4. Decide on a mutually acceptable solution 5. Implement the
solution 6. Evaluate the solution. Following this general pattern,
valuation takes the form of a practical judgement, that is – as spelled
out in Dewey’s The Logic of Judgments of Practise (1915) – the
judgement characterised in terms of being geared towards future
actions, based on an assessment of desired outcomes and using means-
ends reasoning, and being grounded in some form of empirical inquiry. Valuation through Narrative Intelligibility 15 158 This, in a nutshell, is tantamount to meeting the conditions of
situational interaction and teleological continuity. g
y
Thus, it could be suggested that narrative intelligibility is the
grounding of Dewey’s valuation theory. According to Dewey, valuation
is best thought of in terms of acts of interpretation and justification
configured by specific socio-material contexts (Dewey 1939). Even
though anchored in individual experiences, as these are shaped by
moral norms and ethical values, value can be seen as constructed and
reconstructed by groups and communities who themselves are shaped
by their historical and geographical, social and cultural, institutional
and technological contexts (cf. Foucault 1980; Latour 2005). This is
consonant with ANT and, not surprisingly, Dewey has been embraced
by key figures in valuation studies (e.g. Muniesa 2011). It is worth
pointing out in this context that Dewey and his fellow George Herbert
Mead were instrumental in the development of the Chicago School of
Sociology. There they played a pivotal role in driving the work to
understand the nature of relationships in regional geography
(Singelmann 1972) later to be associated with the thinking about
‘social worlds’ (Strauss 1978) which, through science and technology
studies, fed into ANT thinking (Clarke and Star 2008) and situational
analysis (Clarke 2003). y
Dewey’s thinking is thus informed by the considerations concerning
‘social worlds’ and ‘site ontologies’ that preoccupied the Chicago
school. However, for Dewey the idea of situated ‘shared discursive
spaces that are profoundly relational’ (Clarke and Star 2008: 120) is
never separated from teleological analysis and never purely discursive. Simply put, according to Dewey, people value things because they are
useful to them. And yet, not all desired outcomes and future actions
are equal. Situational interaction and teleological continuity in
Dewey On the one hand, Dewey admits that there are a number of
normative standards that can in principle apply to any given situation. At the same time, the process of inquiry, as outlined in the six steps
above, rests on the idea of justification. To value is to weigh the means
and ends in specific situations and crucially, to demonstrate that the
proposed course of action is justifiable in collective terms. This is the
basis of normativity in Dewey. y
y
Thus, while faithful to some tenets of what this article dubbed the
ANT-informed valuation, Dewey is also a member of the humanities
tradition with his insistence on normativity. The problematic situations
that trigger valuation are unmistakably a part of the socio-material
networks. Yet, the teleology constitutive of valuation unfolds
according to the means-ends reasoning and in line with normative
constraints. His account brings together MacIntryre and Haraway. Situational interaction and teleological continuity underpinning
narrative intelligibility are the ground and a way of bridging the
descriptive and normative approaches to valuation in Dewey. In such a
way, narrative intelligibility provides a solution to what Dewey dubs Valuation Studies
159 ‘the deepest problem of modern life’: ‘the problem of restoring
integration and cooperation between man’s [sic] beliefs about the
world in which he lives and his beliefs about values and purposes that
should direct his conduct’ (1929: 204). Acknowledgments g
I am grateful for the generous feedback from the two anonymous
reviewers and for the thoughtful, careful, and supportive steer from the
editors of this theme issue. I also wish to thank Christian Nold –
Lecturer in Design at The Open University – for talking with me about
ANT. Valuation through Narrative Intelligibility
160 Valuation through Narrative Intelligibility
160 Valuation through Narrative Intelligibility
160 Acknowledgments Concluding reflections on overcoming ‘the great
divide’ This article argues that the ANT-informed descriptive approach to
valuation and the humanities-grounded normative theories of value
will not succeed in separation from one another. Moreover, they have
much to gain from coming together, individually and collectively. For
the humanities-grounded perspective, unless this tradition finds a way
of keeping itself empirically embedded, the use of reasoning alone is
likely to result in rarefied and reifying accounts of value. As for the
ANT perspective, it too will fail in separation. By its own admission,
ANT sees itself as ‘a toolkit for telling interesting stories about, and
interfering in, those [world] relations’ (Law 2007: 2). But can ANT
really tell stories rather than compose the proverbial shopping lists? Are the networks created just random collections or is there any
orientation to them? Moreover, are some ways of composing these
networks better than others – are some courses of affairs
straightforwardly bad? The argument presented here is that, if ANT
remains stranded with lists and not stories, it will fail in its attempts to
account for valuation as a phenomenon in the world where some ways
of acting are better than others, where norms of criticism and
standards of justifications apply. This article argues that narrative intelligibility is a useful framing
device that brings together the tradition concerned with value in the
humanities and the ANT approach preoccupied with valuing. Narrative intelligibility binds the notions of teleological progression,
orientation and purpose (MacIntyre 2007 [1987]) with those of the
situated, material interconnectedness (Haraway 2013, 2016). Conceived in these terms, it allows valuation to be seen as normatively
grounded and goal oriented, as well as empirically situated and
contextually circumscribed (Dewey 1939). y
y
Narrative intelligibility provides thus a good frame to make sense of
the phenomenon of valuation but also to set an agenda for research
into valuation. Adding narrative intelligibility to the ‘toolbox’ of
valuation studies opens up the possibility of integrating the normative
and descriptive approaches and with this, the option of interrogating
the assumptions and hegemonies present in the understandings around
‘good’ and ‘bad’ valuing (cf. Horkheimer 1972) without relying on
some fixed metaphysical abstractions. This option is, arguably, needed
in valuation studies and in Valuation Studies (Helgesson and Muniesa
2013; Helgesson 2021). Valuation through Narrative Intelligibility
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Studies 3(1): 45–73. Patrycja Kaszynska is Senior Research Fellow at University of the Arts
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166 for the Arts and Humanities Research Council (AHRC) Cultural Value
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Instinto de mejora.Diferencia entre dos movimientos migratorios: el éxodo rural de la década de 1960 y la inmigración actual.
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ISSN: 0214-0314 ISSN: 0214-0314 Cuadernos de Trabajo Social
Vol 15 (2002): 217-234 Cuadernos de Trabajo Social
Vol 15 (2002): 217-234 Abstract The present article tries to show the similitu-
des and differences between two of the most im-
portant and with more consequences migratory
movements that have taken place in the last fifty
years in Spain: the depopulation of the country-
side that laid out and prepared for city develop-
ment and industrialized the countre since 1950
and present immigration surrounded by discrimi-
nation, racism and intolerance. El presente artículo intenta mostrar las simili-
tudes y diferencias entre dos de los movimientos
migratorios más importantes y con más conse-
cuencias que se han producido en los últimos cin-
cuenta años en España: el éxodo rural, que urba-
nizó e industrializó el país a partir de 1950, y la
inmigración actual rodeada de discriminación ra-
cismo e intolerancia. Juan José MUÑOZ ORTEGA* Juan José MUÑOZ ORTEGA* * Sociólogo. Cuadernos de Trabajo Social
Vol 15 (2002): 217-234 Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz Juan José Muñoz mentos más utilizados para entrar en
el país, no es el elemento definitorio
del fenómeno de la inmigración actual,
sino más bien es el avión el medio más
utilizado en conjunto para llegar a
nuestro país. Concretamente, si en
1999 había un total de 801.329 inmi-
grantes (legales) residiendo en Espa-
ña, un 20% eran marroquíes mientras
que un 32% eran europeos del este,
sudamericanos y asiáticos1. Con ello
quiero decir que, aunque los inmi-
grantes marroquíes fueran los más nu-
merosos, la patera no es el vehículo
más utilizado. Son muchos los tipos
de inmigración y, porque la televisión
nos muestre imágenes lamentables de
sucesos ocurridos en las costas an-
daluzas, no vamos a generalizar y a to-
mar por el todo lo que es una las par-
tes de lo que es la inmigración. Sobre
todo cuando los movimientos migra-
torios nos han acompañado y ayuda-
do en el propio devenir histórico de la
sociedad humana. por sus posibles consecuencias po-
tenciales en otros aspectos, como son
las sociales (probables conflictos y di-
ficultades de integración), las políticas
(medidas y acuerdos políticos para la
total integración de los inmigrantes) y
las económicas (el paro estructural que
sufre la sociedad española). La solución del actual problema in-
migratorio viene de la mano de una de
las variables que explican mejor el fe-
nómeno: la legalidad o ilegalidad de los
inmigrantes. Las pregunta es: ¿qué ha-
cemos con los inmigrantes ilegales? Díez Nicolás advierte que España podría
soportar la entrada de 100.000 inmi-
grantes al año para mantener su creci-
miento demográfico y económico. Para
ello, se debería controlar más si cabe los
flujos de entrada a nuestro país. Así pues, según los resultados de
la encuesta del CIS, la inmigración ac-
tual se presenta como un problema
que se ha de resolver a los ojos de los
ciudadanos españoles, aunque desde
una esfera política se piense en la in-
migración como una ventaja en su sen-
tido demográfico, económico, político
y cultural, sobre todo, cuando el nú-
mero de inmigrantes que llegan a nues-
tro país crece, haciendo de España un
país de destino de inmigración. 1 Anuario Estadístico de la Comunidad de Madrid 2001. Instituto de Estadística. Consejería de Presiden-
cia y Hacienda. 1. Introducción por la masiva entrada de inmigrantes
en nuestro país; pero esta entrada es-
tá condicionada por la imagen que la
sociedad española tiene de la inmi-
gración. Me refiero a las imágenes de
la entrada ilegal de africanos, cruzan-
do el Estrecho, alojados en miserables
pateras que, un día sí y otro también,
invaden las costas andaluzas y los no-
ticiarios españoles. Como es sabido,
ésta no es la única inmigración que se
produce, ya que el fenómeno migra-
torio ha existido siempre y no preci-
samente gracias a pateras y norteafri-
canos. Si nos atenemos a los datos,
las pateras, siendo uno de los instru- S
e
( S
egún el barómetro del Centro de
Investigaciones Sociológicas
(CIS) de agosto de 2001, la in-
migración se situaba como el segun-
do problema nacional por detrás de
ETA y por delante del paro aunque,
analizando los datos por Comunida-
des Autónomas, podíamos observar
cómo en determinadas regiones (An-
dalucía y Murcia), la inmigración se
consideraba el principal problema que
ha de ser resuelto. Como vemos, la opinión pública
española está seriamente preocupada 217 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Dicho lo cual, aportando una nue-
va visión sobre la inmigración, intenta-
ré comparar ambos movimientos mi-
gratorios partiendo de la base de que
sus causas pueden ser parecidas: la
búsqueda de un futuro mejor, el cam-
bio de un ámbito rural, pobre, subde-
sarrollado hacia otro urbano, desarro-
llado, más atractivo que la sociedad de
origen, bien sea los crecientes Madrid,
Barcelona o Bilbao de los años 1960 o
los ya consolidados Madrid, Barcelo-
na o Bilbao del recién inaugurado siglo
XXI; pero sin intentar llegar a conclu-
siones explicativas generales que re-
suelvan el problema, sino más bien
aportando aproximaciones hipotéticas,
ampliando el campo de estudio y mos-
trando efectos descriptivos del fenó-
meno de la inmigración. Para ello, he
realizado dos entrevistas en profundi-
dad interesándome por la historia de
vida de dos sujetos que experimenta-
ron la sensación de abandonar su ciu-
dad natal, sus costumbres, y familias
de origen para buscar destino en una
ciudad completamente diferente, bien
sea nacidos en un pueblo de Sala-
manca o en el mismo Marruecos con
el objeto de analizar las expectativas
del viaje, las causas y los modos, así
como las pautas de adaptación y la vi-
da diaria en la sociedad de acogida a
la que llegaron. Instinto de mejora. Diferencias entre dos movimientos migratorios... Tanto es así, que los expertos en de-
mografía e incluso la ONU postulan la
necesidad de acoger y facilitar la en-
trada de inmigrantes para soportar el
creciente ritmo demográfico actual de
la sociedad, tanto española como eu-
ropea (sobre todo en el caso de Espa-
ña cuya tasa de natalidad es una de las
más bajas de Europa). El profesor Díez
Nicolás comentaba en el I Congreso
Profesional de Sociólogos y Politólo-
gos celebrado en Madrid en julio de
2001 que el fenómeno de la inmigración
es interesante, no sólo por sus reper-
cusiones demográficas sino también El proceso migratorio actual ad-
quiere tintes semejantes a aquel otro
que ocurrió en la España de la déca-
da de 1950-1960, cuando centenares
de personas abandonaron sus pueblos
de origen y abastecieron de mano de
obra las recién industrializadas ciuda-
des importantes de España. Me refie- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 218 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz uno quede sorprendido, pero exami-
nando detenidamente ambos movi-
mientos migratorios podemos observar
que, en su génesis y en su desarrollo,
son similares, salvando y procurando
respetar las tres grandes diferencias
que minimizan los efectos de la com-
paración: el momento del viaje —el
éxodo rural se da en torno a 1950-
1960 y la inmigración actual se pro-
duce cuarenta años más tarde—; el te-
rritorio —el éxodo rural se produce al
interno de España, del campo a la ciu-
dad, y la emigración es de un país a
otro, de una cultura e idioma a otros—;
y su consolidación en la sociedad de
acogida —en 1960 el emigrante rural
pudo mantener su nivel de vida, inclu-
so mejorarlo, mientras que en la ac-
tualidad el inmigrante encuentra serias
dificultades para trabajar, formar una
familia y educar a sus hijos en un cli-
ma acorde para su desarrollo y forta-
lecimiento—. ro al masivo éxodo rural que asoló los
campos y urbanizó las ciudades. Po-
demos decir que ambos procesos pre-
sentan características comunes y no
cabe duda de que son fuente de cam-
bio social en el sentido que modifican
las bases culturales de la sociedad. 2. Aproximaciones
teóricas sobre
las migraciones El fenómeno de la migración se
puede definir como cualquier cambio
permanente de residencia, que impli-
que la interrupción de las actividades
en un lugar y su reorganización en otro
(Ravenstein, 1885). Si eso es así, po-
demos ver que las migraciones han si-
do un recurso muy importante para
asegurar la subsistencia humana. A su
vez, la movilidad social asociada a la
geográfica ha sido durante siglos un
ansia en el sentimiento de gran parte
de los seres humanos, ya que el ins- A primera vista, puede parecer és-
ta una comparación ficticia y más de Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 219 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz de las migraciones —a decir de R. E. Park— que centra la atención en quien
emprende el viaje. “El protagonista de
la migración es el migrante, la perso-
na que experimenta y que sufre todo
el proceso” (Park, 1950). Enfatiza las
particularidades que conforman la per-
sonalidad del migrante, como si tuvie-
ra características especiales: una ini-
ciativa superior, una capacidad para el
trabajo y de adaptación mayor a los de
su entorno; todo ello sumado a los
agentes externos. tinto de mejora es innato en el ser hu-
mano, es el motor de progreso y, cuan-
do el hombre percibe la existencia de
una posible mejora en sus condiciones
de vida, a través de los desplazamien-
tos geográficos se ponen en juego una
serie de factores, cuya conjunción
puede determinar que una persona
emprenda la migración. Una vez producido el desplaza-
miento, el individuo experimenta un
proceso de adaptación que puede
acarrear conflictos, en el sentido de
confrontar dos concepciones de vida
totalmente diferentes. Los valores
aprendidos en la niñez y los nuevos
valores asumidos en la nueva socie-
dad deben confluir adecuadamente en
el hábito de vida de la persona que mi-
gra, aunque ello no impida que el sen-
timiento de desarraigo y la pérdida de
la antigua identidad traigan consigo
particulares características que mar-
quen al sujeto en cuestión. Otra concepción la ofrece Manga-
lam, al entender la migración como un
proceso adaptativo cuyo principal ob-
jetivo es mantener el equilibrio diná-
mico de la organización social con mo-
vimientos de cambios destinados a
proveer a sus miembros de recursos
para superar sus privaciones. 2.1. Teorías sobre las migraciones Los primeros estudios sobre las mi-
graciones datan de finales del siglo XIX,
cuando las migraciones se veían como
un proceso de expulsión-atracción (Ra-
venstein, 1885, y Haddon, 1912), por el
cual todo individuo se halla sometido a
una serie de fuerzas que tienden a ex-
pulsar a las personas de la comunidad
en la que viven y, a su vez, en la mente
de las mismas se desarrolla una idea
acerca de la posible sociedad a la que
dirige su migración. Esta idea es atra-
yente y muestra las ventajas que se po-
dían obtener con el cambio. 2. Aproximaciones
teóricas sobre
las migraciones De esta
manera, presta más atención al pro-
ceso de adaptación que a la persona
y al proceso de expulsión o atracción
de la gente (Mangalam, 1968). Por su parte, Kasdan presenta al
migrante no como un sujeto pasivo de
su historia personal, sino como un in-
dividuo voluntarioso que comprende
su entorno y toma sus propias deci-
siones entre las alternativas que se le
ofrecen. Esta cuarta forma de enten-
der el proceso migratorio trata de va-
lorar la propia iniciativa de la persona
(Kasdan, 1970). Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 2.2. Causas de las migraciones Según uno de los expertos en eco-
nomía y demografía española, Ramón
Tamames, los movimientos migrato-
rios se producen por diferentes ten-
siones demográficas en los recursos Años más tarde, a mediados del si-
glo XX, surge una nueva concepción Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 220 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz económicos disponibles. Tamames
pone el dedo en la llaga y lleva la ex-
plicación al terreno que le correspon-
de, ya que, independientemente de los
factores personales, es un hecho que
los movimientos migratorios son pro-
cesos de carácter esencialmente eco-
nómico (Tamames, 1983). Todos los
procesos migratorios que se produje-
ron en España a partir de la década de
los cincuenta y sesenta responden a
la industrialización y a la demanda de
mano de obra. gar de expulsión como en el lugar de
recepción, ya que se crea un desequi-
librio demográfico en ambos lugares,
las diferencias culturales y sociales se
acentúan y la diversidad de etnias, len-
guas y religiones requieren una inte-
gración para aliviar esas tensiones. Los expertos en migraciones han
podido elaborar una serie de caracte-
rísticas comunes de los procesos de
migración: — Las migraciones son selectivas. Es un fenómeno discriminatorio, ya
que no puede migrar todo el mundo,
sino un tipo determinado de personas:
los varones y las mujeres jóvenes prin-
cipalmente. El gran conocedor de migraciones
en España, el profesor Víctor Pérez
Díaz, señala que cuando una crisis ame-
naza con hacer estallar una sociedad
tradicional, el éxodo rural contribuye a
que desaparezca el trabajador de la tie-
rra. La migración —dice— es una solu-
ción particular a los problemas surgidos
en una sociedad particularista y al em-
pobrecimiento de zona rural. En defini-
tiva, “la emigración es la manera más
rotunda que tienen los campesinos de
manifestar su crítica de las condiciones
de vida y de trabajo que les ofrece el
campo” (Pérez Díaz, 1974: 37). — Existen redes de migraciones. Tienden a migrar las personas más ca-
pacitadas para luego comunicar la ex-
periencia. — No son permanentes ni en el
tiempo ni en el espacio. De un sitio se
va a otro para luego buscar otro mejor. 2.2. Causas de las migraciones — Existe una teoría de red por la
cual el que migra necesita tener una
comunidad afín a su propia individuali-
dad con el objeto de obtener un apoyo. — Las migraciones suponen un
flujo de divisas de los emigrantes ha-
cia su lugar natal con las que la fami-
lia que se queda en el lugar de expul-
sión continúa viviendo. — Las migraciones suponen un
flujo de divisas de los emigrantes ha-
cia su lugar natal con las que la fami-
lia que se queda en el lugar de expul-
sión continúa viviendo. 2.3. Características
de las migraciones Como decíamos al principio, los
movimientos migratorios son cualquier
cambio permanente de residencia que
implique la interrupción de las activi-
dades en un lugar y su reorganización
en otro. Su origen está en el desequi-
librio entre la población y los recursos
del lugar en los que se reside y del lu-
gar al que se va. No cabe duda de que
tal necesidad de recursos produce
problemas y tensiones, tanto en el lu- 2.4. Marco teórico propio Parto de la idea de que las migra-
ciones afectan a las personas que las
experimentan y suponen un cambio
radical en sus vidas. Lo más curioso
es que siempre obedecen a una deci-
sión enteramente personal, aunque el
individuo desconozca las consecuen- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 221 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz cisión de migrar: el factor económico. La miseria rural que produjo la Guerra
Civil desoló los campos y proporcionó
mano de obra a las fábricas de las ciu-
dades más importantes de España. cias de su movimiento. El migrante
percibe su entorno socioeconómico al
igual que sus compatriotas, pero el
emigrar responde a una necesidad
personal y raramente percibe que esa
decisión personal obedece a una cri-
sis socioeconómica del entorno en el
que vive y a un auge económico y de
ampliación de posibilidades de elec-
ción de la sociedad a la que se dirige. De esta manera, lo que he estudia-
do es el fenómeno migratorio compa-
rando las similitudes y diferencias entre
dos de los movimientos más importan-
tes ocurridos en España intentando
mostrar las expectativas, los procesos,
las causas y sus consecuencias. Pero,
¿en qué contextos se sitúan ambos fe-
nómenos? Los dos siguientes epígrafes
ofrecen un breve contexto histórico de
ambas situaciones. De este modo, toda migración com-
prende un marco socioeconómico de-
terminado, un individuo, una sociedad
de origen, una comunidad receptora,
unas causas del movimiento, unas ex-
pectativas, un intento de mejora y unos
objetivos deseados. Como vemos,
además de los factores personales que
inciden en la toma de la decisión de mi-
grar, encontramos factores externos
que son los que acaban provocando
los grandes movimientos de masa. Junto a todo ese proceso interno de la
persona migrante no podemos olvi-
darnos del componente clave de la de- 3. Contexto
socio-histórico de la
España de 1950 y 1960 La Guerra Civil supuso un cierto re-
traso en las condiciones de vida de to-
dos los españoles. No sólo por la des- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 222 Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz tantes de España y la precaria situa-
ción de vida y laboral de los campos
provocaron un proceso migratorio de
alto rango, en el que más de dos mi-
llones de campesinos abandonaron los
campos en poco tiempo. La vida rural
se hacía cada vez más difícil y sobre
todo los jóvenes ansiosos de mejoras
vieron con buenos ojos la búsqueda de
otras oportunidades, desembarazán-
dose de las duras tareas agrícolas. aparición de hombres y mujeres que lu-
charon sino por las míseras condicio-
nes en las que se quedaron las familias
que trabajaban en el campo. En este
caso no hay que olvidar que la mayoría
de los activos se dedicaban a la agri-
cultura y las consecuencias de la Gue-
rra Civil redujo su trabajo únicamente a
la supervivencia física y a conseguir ali-
mentos a través de un trabajo mal pa-
gado en una nación bloqueada econó-
micamente, en el interior y en el exterior,
gracias al autárquico régimen político
español que no gustó a las potencias
aliadas de la Segunda Guerra Mundial. En paralelo al proceso de urbani-
zación, se produjo el proceso de in-
dustrialización. Los centros urbanos
más importantes necesitaron mano de
obra para las nuevas industrias recién
creadas con las ayudas económicas
exteriores. Ante ese efecto llamada, las
personas que decidían emigrar a la
ciudad veían compensado su esfuer-
zo con un trabajo estable en la fábri-
ca. En total, entre 1940 y 1960 dejaron
su provincia de origen en torno a
2.743.000 personas (Del Campo y Na-
varro, 1987). Con el objetivo de crear una Espa-
ña autosuficiente se intentó industria-
lizar el país y para ello se crearon mu-
chas industrias nuevas con el apoyo
del gobierno. Se intentó controlar has-
ta el máximo el escaso comercio ex-
terior y, por último, se trató de au-
mentar la producción agraria. Pero el
crecimiento industrial que se produjo
no podía continuar dentro de la políti-
ca restrictiva de autarquía. En años posteriores, el cambio de
sistema llevó consigo una transforma-
ción de la coyuntura económica. 3. Contexto
socio-histórico de la
España de 1950 y 1960 La
ayuda económica de Estados Unidos
en 1951 facilitó la adquisición de bie-
nes y favoreció la expansión econó-
mica que llegó hasta 1959, año en el
que se publicó el Programa de Estabi-
lización cuyo objetivo era alinear la po-
lítica económica española a la inter-
nacional. De esta manera, se ponía fin
al aislamiento económico y el país se
encaminaba a una economía libre de
mercado según las pautas del capita-
lismo occidental. 5. Del pueblo a Madrid Se llama María Inmaculada y nació
en San Muñoz, como dice ella, un pue-
blo pequeñito en la provincia de Sala-
manca. Un día, con catorce años, des-
pués de acabar el último año de
escuela y de pasar todo un verano
ayudando a su padre en las tareas del
campo, sonó uno de los dos teléfonos
de los que disponía el pueblo. Al otro
lado estaba Isabel, su hermana mayor. Le dijo que la señora que vivía arriba
buscaba una muchacha para las ta-
reas del hogar y que la habló de ella. Isabel ya estaba en Madrid, trabajan-
do para los maestros del pueblo. A
partir de aquel día su futuro estaba es-
crito: un día cualquiera se iría a traba-
jar a Madrid al lado de su hermana. En nuestro país el hecho más sig-
nificativo ha sido el aumento del tráfi-
co de pateras entre las costas africa-
nas y el territorio español. Durante el
año 2000, alrededor de 15.000 perso-
nas fueron detenidas y decenas de
ellas perdieron la vida en el intento,
mientras que durante el año 1999 fue-
ron aproximadamente 3.500 los inter-
ceptados. Estos inmigrantes llegaron
en más de 780 pateras, sin contar los
que lograron entrar ni los que entraron
legalmente por la frontera con dinero
suficiente y visado de turista para tres
meses. Tampoco se contabilizaron los
desaparecidos ni las decenas de inmi-
grantes fallecidos en los naufragios. Al
margen de los problemas de atención,
los colapsos de los centro de acogida
y de las comisarías de los puntos de
llegada, la repatriación hacia los paí-
ses de origen se presenta complicada
ya que la expulsión no es posible le-
galmente en menos de 72 horas. San Muñoz no era un pueblo muy
rico y lo único que podían hacer los jó-
venes era buscarse la vida en otro si-
tio o quedarse trabajando en el cam-
po ayudando a sus familias. Como su
padre no disponía de tierras propias,
Isabel primero y Mari después tuvie-
ron que emigrar a Madrid para traba-
jar y buscar un destino. Llegó el día pa-
ra Mari, cogió un autobús en dirección
a Salamanca y otro para Madrid y en
Atocha estaba su hermana esperán-
dola, la llevó a su nueva casa y así em-
pezó la historia de Mari-emigrante en
la capital. 4. Contexto de la
inmigración actual El problema de la inmigración se
centra sobre todo en el equilibrio en-
tre la contención de flujos y la necesi-
dad de trabajadores. En nuestro caso,
España es uno de los países de la
Unión Europea con más inmigrantes
ilegales (cerca de 300.000) y, aunque
el proceso de regulación concluido en
el año 2000 permitió legalizar su si-
tuación en torno a 140.000 inmigran-
tes, otros 60.000 fueron rechazados
sub conditione de revisión en algunos
casos. El desarrollo industrial que se esta-
ba dando en las ciudades más impor- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 223 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz Juan José Muñoz Según los datos de la agencia es-
tadística de la Unión Europea, Euros-
tat, en España hay un inmigrante por
cada 1.000 habitantes, un porcentaje
bastante bajo comparado con el res-
to de los países miembros de la UE,
sólo por encima de Francia y de Fin-
landia (por ejemplo, Holanda tienen 3
por cada 1.000 habitantes, Irlanda 6,
Italia 3, Alemania 7, Bélgica 20 o Lu-
xemburgo 114). Además, general-
mente estos inmigrantes trabajan en
economías sumergidas, se les paga
menos, operan en peores condiciones
y carecen en muchos casos de vivien-
da digna. propio relato de dos mujeres (Mari y
Babía) que experimentaron la sensa-
ción de emigrar a otro lugar especial-
mente diferente. Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 5. Del pueblo a Madrid Actualmente, tiene 53 años,
vive en Leganés (Madrid), está sepa-
rada, trabaja limpiando una oficina por Los siguientes epígrafes del artí-
culo intentan poner de manifiesto to-
do lo dicho anteriormente en base el Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 224 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz cha independiente, etc. hacían que la
única salida fueran las redes sociales
que se formaban alrededor del pueblo. De esta manera, a muchos de los jó-
venes se les presentaba la oportuni-
dad de emigrar amparados por otra
persona que podía ayudarlos en su
consolidación fuera de allí. La gente
se iba a Logroño, San Sebastián, Bil-
bao, Salamanca, Madrid, etc.; todos
utilizaban a otra persona como puen-
te, gracias a la buena amistad entre
iguales y a la buena vecindad entre los
pudientes del pueblo y las familias tra-
bajadoras. las mañanas y tiene dos hijos: Antonio
de 30 años, casado y con un niño, y
Jesús de 28, que vive en un piso al-
quilado con su novia. 5.1. Las raíces del pueblo La familia Ortega vivía en una casa
al lado de la plaza del pueblo, no era
la mejor casa de la zona, pero por lo
menos les daba suficiente cobijo co-
mo para soportar el helado frío inver-
nal de la zona montañosa salmantina. La poca gente que quedaba en el pue-
blo se dedicaba a la agricultura y a la
ganadería. Jesús, el padre de Mari,
trabajaba duro en una finca de los se-
ñores más pudientes del pueblo y
Amelia, su mujer, tenía bastante con
las tareas propias del hogar. Por aquel
entonces, Isabel y Mari eran muy jó-
venes, pensaban en jugar con las ami-
gas y en divertirse, aunque eso no qui-
taba para que tuvieran que aportar
algo a la familia sirviendo y limpiando
en las casas del médico, del cura, del
maestro o del alcalde. Como vemos,
la familia se sustentaba gracias al es-
fuerzo de todos los miembros del ho-
gar, daba igual la edad que tuvieran las
hijas: debían aportar de su parte para
tener de comer al día siguiente. Como vemos, la vida de Mari se vio
sometida a dos aspectos de vital im-
portancia: la falta de recursos econó-
micos y la ruralidad. El análisis que he
hecho se ha basado en estos dos ejes
intentando explicitar la visión de su vi-
da. El siguiente gráfico muestra gros-
so modo la historia de Mari desde su
comienzo en San Muñoz —rodeada de
dificultades económicas y del contex-
to rural— hasta hoy en Leganés —sin
dificultades económicas graves y en
un contexto urbano radicalmente di-
ferente al de sus comienzos—. 5.2. Una decisión tomada La vida en el pueblo transcurría
monótonamente, la rutina y la reacción
negativa a los cambios propiciaban
una vida que giraba alrededor de la
subsistencia económica y de las tare-
as de la tierra y la cría de animales. En San Muñoz, cuando uno aca-
baba la escuela con catorce años, se
planteaba la primera elección impor-
tante; aunque eran muy jóvenes,
los/as chiquillos/as del pueblo hacían
lo mismo tarde o temprano. La impo-
sibilidad de continuar con las tareas
del padre, la inviabilidad de aprender
un oficio diferente al del campo, la in-
alcanzabilidad de comprar tierras, las
dificultades para emprender una mar- Poco a poco, las niñas iban cre-
ciendo y empezaron a divertirse con las
muchas muchachas del pueblo. Entre
tanto un ambiente pesimista les rode-
aba, los pocos mozos que había se Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 225 Juan José Muñoz
Instinto de mejora. Diferencias entre dos movimientos migratorios. Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz una muchacha que se hiciera cargo de
los críos, ya que ambos trabajaban, y
pensaron en Isabel. Después de pedir
permiso a sus padres y obtener su con-
sentimiento, al acabar el curso, em-
prendieron viaje hacia Madrid. marchaban a la capital a trabajar y las
pocas muchachas mayores dejaban la
casa de sus padres para buscar fortu-
na más allá de las fronteras invisibles
del pueblo. A los 14 años se acababa
la escuela y, como pasa cuando se aca-
ba una etapa, había que buscar nuevos
horizontes. Las cohortes de edades
cercanas a los 14 se fueron yendo a
otros lugares con el consentimiento de
sus padres ya que las posibilidades de
hacer algo en el pueblo eran escasas. Unos se fueron primero y otros des-
pués, pero no cabe duda de que la ló-
gica de ser en el pueblo era abando-
narlo para trabajar en otro sitio. Por otro lado, cuando acabó la es-
cuela, Mari tenía unas ganas locas de
salir de aquel ambiente rural, no era
consciente del cambio que podía su-
ponerle una modificación en el estilo
de vida como aquél, pero ella quería
irse. San Muñoz para Mari representa
una buena parte de su personalidad. Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 5.2. Una decisión tomada Aquel ambiente tradicional, sin cam-
bios, sin conflictos, donde la vecindad
y la tranquilidad eran las señas de
identidad de la gente que vivía allí con-
figuraron desde el principio la infancia
de Mari. De similar modo, la pobreza
que debían soportar, no sólo su fami-
lia sino todas aquellas que no dispo-
nían de una fuente de ingresos cons-
tante ni clara, estructuró su infancia en Isabel fue la primera de la familia
que lo hizo. Era una chica muy cono-
cida en el pueblo por lo vivaracha y no
pasó desapercibida para los maestros,
un matrimonio con tres hijos pequeños,
que pidieron el traslado a Madrid para
continuar su profesión y dar una mejor
educación a sus hijos. Necesitaban Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 226 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz el sentido de luchar y trabajar por so-
brevivir. ... pero como ya veníamos con los
maestros... mi hermana fue la casuali-
dad y gracias a eso me vine. Y de he-
cho estabas contenta, estabas bien,
añorando tus raíces y tus cosas... ... Entonces a los 14 años pasaba
eso. O te ibas de allí o te quedabas,
pues eso, a vestir santos o a nada por-
que nosotros, en mi casa, no teníamos
labor, no teníamos nada y como todas
se iban pues tú igual te ibas... La forma de salir del pueblo era
común: la utilización de otras gentes
que hubieran pasado por ese proce-
so. Bien sea por amigos de más edad,
que buscaban acomodo en el lugar
de atracción a los chicos más jóve-
nes, o por recomendaciones de las
gentes pudientes del pueblo. Todo
ello, encaminado a ciudades en auge
por los procesos de industrialización
y de oferta de mano de obra de las
ciudades. ... se veía normal, porque nosotros
no teníamos tierras, no teníamos nada
pero gente que sí las tenía y podía vi-
vir de ello, también se venían... En el pueblo, el modelo estaba cla-
ramente construido, la vida exigía bus-
car una salida a la precariedad y en
aquellos momentos, la única opción
era irse y aumentar así las oportuni-
dades de elección, la niñez en el pue-
blo duraba poco, los 14 eran una edad
en la que ya se podía salir adelante in-
dependientemente de los padres. 5.2. Una decisión tomada Sa-
lir del entorno rural se veía normal, co-
mo un ciclo por el que había que pasar. Los padres querían a sus hijos como
ahora, pero eran conscientes de que
ellos tenían un trabajo, que eran po-
bres, y que no podían supeditar la vi-
da de un hijo a un destino tan poco va-
lorado y con tanto sacrificio. Además,
si hubiera tierras, ganado o alguna otra
industria en donde poder trabajar, la
gente se hubiera podido quedar, pero
éste no era el caso de San Muñoz. Los
padres de entonces sabían que la vi-
da no estaba en el pueblo y las co-
rrientes migratorias anteriores mos-
traban el camino de los mozos
siguientes. ... yo estaba loca por salir del pue-
blo... ... yo estaba loca por salir del pue-
blo... ... por cambiar, porque si nos hu-
biéramos quedado allí no hubiera ha-
bido nada que hacer... ... poco futuro y ningún aliciente,
nada... Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz cién llegados. Según Mari, todo era
grande, las dimensiones de la ciudad
sobrepasaban lo imaginado por ella. Acostumbrada a cuatro calles, a la pla-
za, la única carretera de acceso, los
pisos bajos, la rutina, etc. salir del pue-
blo para vivir en una ciudad de un mi-
llón de habitantes, con edificios de
diez plantas, el metro, los autobuses,
las nuevas tecnologías, oportunida-
des, consumo, etc. Se trataba de un
cambio importante, sobre todo, cuan-
do no había salido nunca de San Mu-
ñoz, ni siquiera para ir a Salamanca
que estaba a 42 kms. hacía compañía a cambio de propor-
cionarle cama, comida y dinero para
que ella pudiera aprovechar la opor-
tunidad. Mari se encontró muy bien
allí, no le faltó de nada, incluso pudo
conocer cosas nuevas que en el pue-
blo nunca podría haber encontrado:
bicicletas, dulces, tebeos, etc. No era
un trabajo tan duro como en el cam-
po, pero exigía más tiempo del que
Mari estaba acostumbrada. Al princi-
pio le costó un poco la adaptación a
la ciudad, ella sólo quería quedarse en
casa, pasar el día conversando con Dª
María o con alguna amiga, leer tebe-
os, comer regaliz, etc. Los dos ámbitos en lo que Mari se
encontraba tenían diferentes caracte-
rísticas: por un lado estaba el pueblo
donde se encontraba con su familia,
con sus amigos de la niñez, con las ra-
íces, con los juegos, con la compañía,
etc.; mientras que en la ciudad dispo-
nía de una segunda familia, la juven-
tud, el desarraigo, la gente conocida,
el trabajo, la soledad, etcétera. La relación con la gente de Madrid
era excelente, contaba con su her-
mana a un palmo de distancia y ella le
ayudó a adaptarse más deprisa. El ca-
riño mutuo que se tenían era y es
constante, no sabrían vivir la una sin
la otra. En los primeros meses, Isabel
se preocupaba mucho por dar aco-
modo a su hermana y, como ella ya
llevaba mucho tiempo allí, le había da-
do tiempo para hacer amistades, pre-
ocupándose de presentárselas a su
hermana. Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 5.3. Un nuevo hogar en Madrid La casa a la que llegó Mari estaba
encima del hogar en que trabajaba su
hermana. Se trataba de dos mujeres,
Dª María, viuda desde hacía muchos
años, y su hija que acababa de que-
darse sola al fallecer a su marido en-
fermo. Ambas habían estudiado en la
Universidad y trabajaban fuera de ca-
sa. Al estar solas necesitaban la ayu-
da doméstica de una chica como Ma-
ri. Al ser muy joven, las señoras la
trataron como si fuera una hija más. Para Mari, aquellas personas signifi-
caban todo. A pesar de ser sus amas,
no miraron a Mari como a una niña que
venía a trabajar para ellas, sino que les ... (hablando de los padres)... esta-
ban conformes, lo primero porque
cuando yo me vine, no me vine a la
aventura, venías con una referencia, un
poco para tranquilizarlos... Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 227 6. España, país de destino chaban para ir a ver a sus padres y a
las amigas. Era un pretexto para vol-
ver, pero también para mostrar lo bien
que les iba en la ciudad, para afian-
zarse moralmente en su elección de
emigrar a Madrid y para mostrar el sta-
tus adquirido a las demás gentes del
pueblo. Por otra parte, Rabía nació en Lara-
che, a pocos kilómetros al sur de Tán-
ger (Marruecos) hace 31 años. Toda su
vida la dedicó a ayudar a sus padres en
el negocio familiar hasta que, en 1993,
tuvo la posibilidad de cruzar el estrecho
y buscarse una nueva vida en España. Por aquel entonces tenía 23 años y tar-
dó poco tiempo en decidirse, puso en
una balanza lo que tenía en Larache (la
familia) y lo que podría tener en España
(trabajo, dinero, su propia familia, etc.)
y con el consentimiento de sus padres
y ayudada por unos vecinos, que iban
y venían de España frecuentemente,
emigró hacia nuestro país. Hoy está ca-
sada, tiene un niño y vive en un piso del
barrio de La Latina en Madrid, en com-
pañía de dos hermanos suyos que han
venido también de Marruecos. ... locas por ir a las fiestas, por lle-
var ropa bonita, por llevar cosas que a
lo mejor en el pueblo no conocían... Entre tanto y tanto, un día cual-
quiera, en el Retiro, intentando apren-
der a montar en bicicleta conoció a un
chico muy simpático llamado Antonio. Alternando el trabajo y el noviazgo
buscaron un piso en las afueras de
Madrid y encontraron uno en Leganés,
una ciudad en auge poblacional. En
poco tiempo se casaron y Mari dejó de
vivir en la casa de Dª María. La vida transcurrió con normalidad,
como ella dice “con etapas buenas y
malas” y actualmente continúa vivien-
do en Leganés, sus hijos son mayores
y tienen la vida resuelta. Ella sigue tra-
bajando y sintiendo algo más que nos-
talgia por aquella infancia en el pue-
blo. Instinto de mejora. Diferencias entre dos movimientos migratorios... El problema era que Mari te-
nía un carácter distinto del de su her-
mana, era más bien triste, su timidez,
su tranquilidad y sencillez le hacía ser
una chica muy formal y muy respeta-
da en su círculo de amistades. Al pa-
recer las características rurales toda-
vía estaban presentes en Mari: el
apego a la familia, la tradición, la reli-
giosidad, etc. El tiempo pasaba con las tareas del
hogar, pero también tenía momentos
libres para divertirse. Era muy hogare-
ña, se sentía bien en casa, con los su-
yos, hablando y permitiéndose los ca-
prichos que en el pueblo nunca tuvo. Pensando en el pueblo, tanto Mari co-
mo Isabel, se compraban ropa, bolsos,
zapatos, se cuidaban, iban a la pelu-
quería, siempre pensando en volver en
las fiestas del pueblo para ver a sus
padres. La vuelta al pueblo era muy espe-
rada por ambas. Significaba volver a
las raíces, al lugar donde habían pa-
sado su tierna infancia. Cada 24 de ju-
nio se celebraban las fiestas de San
Juan en el pueblo y ambas aprove- Por aquel entonces, Madrid estaba
creciendo. La ciudad se llenaba de
gente de todas partes y la zona del
centro se convertía en escaparate de
las vanguardias sociales para los re- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 228 Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz Juan José Muñoz 6. España, país de destino El contraste se presenta cuando
comparamos los espacios en los que
vivió: de la subsistencia que ofrecía el
pueblo, de la imposibilidad de movili-
dad social que representaba un ámbi-
to rural cerrado a la innovación a la po-
sibilidad de ascensión en una ciudad
grande, al trabajo remunerado base de
todo el patrimonio que tiene hoy en
día, al bienestar social y personal en
un piso propio de Leganés represen-
tante la lucha por sacar una familia
adelante. Al igual que en el caso de Mari, la
vida de Rabía se podría estudiar me-
diante dos ejes: uno señalaría el paso
de un ámbito subdesarrollado y de
economía de subsistencia a otro más
desarrollado y de economía de merca-
do y hábitos de consumo; el otro eje
señalaría el paso de una posición so-
cial empobrecida y lejana del bienes-
tar hacia una posición social más aco-
modada y más libre de ataduras
económicas. De este modo, podremos
observar que la trayectoria vital de Ra-
bía mejoró sustancialmente, no sólo
por su coraje al intentar cruzar el Es-
trecho, sino también por haber conse-
guido un trabajo, mantenerlo, apren-
der un idioma, obtener los papeles que
legalizaban su estancia y lograr esta-
bilidad emocional al lado de su familia:
su marido, su hijo y sus dos hermanos. ... te metes en una rueda y la tienes ... te metes en una rueda y la tienes
que seguir... que seguir... Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 229 Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz En relación al propio relato de Ra-
bía, el texto se va estructurar en tres
partes: una dedicada al viaje desde
Larache a Madrid, haciendo referen-
cia a las causas del movimiento y a las
expectativas del viaje; otra dedicada
a los primeros años de vida en la ca-
pital y a su consolidación en una cul-
tura diferente a la suya; y, por último,
una tercera dedicada a la propia visión
que tienen nuestra protagonista sobre
el fenómeno de la inmigración. ta donde pasó por una segunda adua-
na y la vigilancia de perros policía en
busca de droga. Y, por último, ence-
rrada durante dos horas en el trayecto
en barco desde Ceuta hasta Algeciras. Allí, amigos marroquíes nacionalizados
españoles del hombre que la trajo en
el maletero la condujeron hacia la ca-
pital. Una vez en Madrid, Rabía se di-
rigió a la casa de sus vecinos de Ma-
rruecos que la esperaban con un
trabajo para empezar al día siguiente. Los primeros meses fueron muy di-
fíciles para Rabía, se encontraba en un
país diferente, no sabía nada de es-
pañol y debía de ir todos los días a una
casa a cuidar a un niño pequeño. Los
problemas empezaron temprano, al
primer mes de estancia en España,
cuando cobró su primer sueldo, se
compró ropa y sus vecinos, aquellos
que la dieron una cama en su casa, la
acusaron de robo en la casa donde
trabajaba por envidia, según Rabía. El 6.1. La llegada Rabía llegó a España en octubre de
1993 con 23 años alojada en un male-
tero de un coche. Lo pasó muy mal y
fue una experiencia especialmente du-
ra para ella que la marcó en toda su es-
tancia en España. Allí encerrada estu-
vo durante el viaje de Larache hasta
Tetuán pasando por una primera adua-
na. Después, desde Tetuán hasta Ceu- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 230 Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz peles y en 1995 Rabía consiguió lega-
lizar su estancia en España. Poco
tiempo después, se casó con un chi-
co marroquí que conoció en Madrid,
trabajaba como portero en una casa y,
al tener dinero ahorrado, ambos deci-
dieron irse a vivir en un piso alquilado
en el barrio madrileño de La Latina. Como ya no vivía en la casa de aque-
lla señora, respetándola y queriéndo-
la mucho, cambió de trabajo. Ahora
debía cuidar a una niña desde las ocho
de la mañana hasta las ocho de la tar-
de. Las cosas se fueron arreglando ya
que en esos dos años aprendió a uti-
lizar con fluidez el castellano, pudo
disfrutar de la oferta de ocio madrile-
ña, incluso pudo viajar con su marido
por España gracias a la compra de un
coche de segunda mano. Por aquel
entonces, con los papeles bajo el bra-
zo hizo su primer viaje de regreso a
Marruecos, porque a pesar de tener
los papeles arreglados, el viaje fue una
catástrofe; cuando llegó a la aduana
en Marruecos la pidieron el visado de
salida del país, pero como ella había
entrado ilegalmente la primera vez tu-
vo que pasar otra noche en la comi-
saría de la aduana hasta que llegaron
sus padres pagando su libertad. Para
volver a España legalmente tuvo que
ir al consulado español en Tánger y
pedir visado para poder volver nor-
malmente. incidente la llevó a comisaría. Allí pa-
só toda la noche hasta que la señora
del piso donde trabajaba desmintió to-
do y pagó la fianza. La dejó en liber-
tad bajo su tutela, la dio un contrato
de trabajo y como Rabía no tenía pa-
peles prometió a la policía que arre-
glaría su situación. La señora contra-
tó a un abogado y a los dos años ya
tenía en regla sus papeles. 6.1. La llegada No volvió
a casa de sus vecinos y se quedó du-
rante esos dos años interna en la ca-
sa de aquella señora. En aquellos primeros momentos,
empezó a comprobar que lo que le con-
taron de España no se asemejaba a la
realidad. De las cosas que observó, lo
más llamativo fue el choque cultural del
idioma, la religión, el trato de los hom-
bres hacia las mujeres y el trato de los
hijos hacia los padres, muy diferentes
a lo que conocía y practicaba en Ma-
rruecos. Lo pasó muy mal, incluso has-
ta tuvo miedo, ya que, aparte de no ha-
blar castellano y no tener papeles, no
tenía a sus padres ni a sus hermanos,
no tenía a nadie, sólo a la señora y al
niño que cuidaba. Aún pasándolo mal
tenía la esperanza de tener una vida
mejor que la que tenía en su país. ... porque en mi país no tenía ni tra-
bajo ni nada, y por eso pensé venir
aquí, para buscar una vida mejor que
la mía, que la que tenía en mi casa y en
mi país, pero siempre en esta vida te
cuesta ... Pero lo que más echaba en falta en
Madrid, además de su familia, era el
tiempo. A veces no disponía de tiem-
po para ella misma; se levantaba, iba
a trabajar, volvía, preparaba la cena y
a la cama para repetir el proceso al día
siguiente. No podía aguantar así. Unos
años más tarde, en 1998, llegó un in- 6.3. El fenómeno de la inmigración Este último aspecto que hemos
presentado en la vida de Rabía, es de-
cir, las ganas de regresar a Larache no
sólo son porque allí están sus raíces, ni
tampoco porque aquí lo haya pasado
tan mal, sino por las diferencias cultu-
rales y los hábitos de comportamien-
tos tan diferentes entre lo conocido
por Rabía en su estancia en Marrue-
cos y en Madrid, además del senti-
miento de rechazo y discriminación no
manifiesto pero sí latente. Actualmente viven los cinco en un
pisito pequeño intentando aportar los
recursos suficientes para continuar vi-
viendo en Madrid. Pero, hace poco, su
marido se quedó sin trabajo. El edifi-
cio en el que trabajaba fue comprado
por otras personas y se quedó sin em-
pleo. El niño tiene ya tres años y como
su marido está buscando trabajo, Ra-
bía se ha visto obligada a trabajar. Ahora se dedica a limpiar una casa en
Fuencarral. Trabaja de ocho a seis y le
gustaría pasar más tiempo con su ni-
ño. Entre tanto, su marido y su her-
mana, a la vez que buscan trabajo, se
dedican a cuidar al pequeño mientras
ella no está. ... siempre notas que eres dife-
rente de la gente que te rodea, no es
tu gente, no es tu religión, ni el idioma
ni tu país..., con el tiempo noté que to-
dos somos personas... Por ejemplo, al llegar a España le lla-
mó especialmente la atención cómo se
comportaban y cómo eran tratadas las
mujeres aquí. Le parecía bien que des-
de pequeñas, las niñas pudieran elegir
qué hacer y cómo ser, lo que en Ma-
rruecos no sucede. Por el contrario, un
aspecto más negativo que llamó su
atención fue la falta de respeto que tie-
nen los jóvenes españoles con los pa-
dres y mayores, lo que en Marruecos
no sucede ya que un joven no hace na-
da sin el consentimiento de los padres. Todo esto le da más razones a Ra-
bía para pensar en su tierra y en su fa-
milia. Piensa que cuando tenga dine-
ro suficiente, cuando le toque la lotería
volverá a Larache porque es su sitio,
su lugar, sus raíces y porque no se vi-
ve tan mal. 6.2. Su consolidación
y la actualidad Rabía estuvo dos años en casa de
aquella señora cuidando a su hijo. Con
el tiempo, el abogado tramitó los pa- Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 231 Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz Juan José Muñoz de quiere vivir, si en España o en Ma-
rruecos. ... siempre tengo pensado que al-
gún día tengo que volver a mi país. ¡Al-
gún día tengo que volver! No me voy a
quedar toda mi vida aquí, aunque me
va bien, pero siempre echas de menos
tu tierra... vitado, en 1998 nació su primer hijo. Pero, otra vez lo pasó realmente mal,
ya que tuvo que dejar de trabajar pa-
ra cuidar a su pequeño y, a su vez, se
sentía sola, a pesar de la compañía de
su marido, echaba en falta a su pa-
dres, sus hermanos y amigos. Para
ayudarla y aprovechando la situación,
llegaron dos hermanos suyos de La-
rache. Uno tenía 29 años y encontró
trabajo rápidamente en la construc-
ción y otra de 20, la más pequeña de
la familia, que se quedó en casa con
ella, cuidando al niño. El tiempo solu-
cionó los problemas. de quiere vivir, si en España o en Ma-
rruecos. ... siempre tengo pensado que al-
gún día tengo que volver a mi país. ¡Al-
gún día tengo que volver! No me voy a
quedar toda mi vida aquí, aunque me
va bien, pero siempre echas de menos
tu tierra... Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 6.3. El fenómeno de la inmigración Piensa tener su negocio
propio, un trabajo y una casa, además
quiere que su hijo conozca la cultura
marroquí, después cuando sea más
mayor le dará libertad para elegir don- A su vez, pudo comprobar el re-
chazo y la discriminación, no sólo en Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 232 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz su incidente con los vecinos que le ha-
bían acusado de robar en su trabajo,
sino en otras muchas ocasiones cuan-
do la gente se manifestaba en estos
términos a la hora de coger el autobús
o buscar trabajo. En las muchas oca-
siones que iba en autobús, las seño-
ras de alrededor agarraban el bolso y
se apartaban de ella. A su vez, su ma-
rido, cuando dejó de ser conserje y
buscó trabajo, “con lo preparado que
estaba, teniendo conocimientos y ex-
periencia de mantenimiento, jardine-
ría o electricidad, no ha conseguido en
ninguna de las muchas entrevistas a
las que ha ido ningún empleo”. sensación de novedad, de triunfo, de
riqueza que llega a Marruecos se vuel-
ve contra ellos aquí ya que son capa-
ces de abandonar sus familias y su vi-
da tranquila para venir a España, lugar
en el cual no es oro todo lo que relu-
ce, sobre todo si intentan llegar a Es-
paña ilegalmente. Rabía nos contaba como muchos
de sus amigos que han llegado ilegal-
mente se arrepentían de su decisión,
ya que en Marruecos tenían a su fa-
milia y su trabajo digno y que lo deja-
ron todo para mal vivir en España tra-
bajando muchas horas al día. Incluso
nos contaba que lo han intentado mu-
chas veces por patera y que no lo han
conseguido habiéndose gastado aho-
rros de la familia para nada. Lo peor es
que lo volverán a intentar. 6.3. El fenómeno de la inmigración vida que te pasa todo en ella, y pien-
san, a lo mejor, que nada más llegar
aquí van a trabajar, van a tenerlo todo,
pues no es así, todo es difícil y todo te
cuesta, todo... Aunque es verdad que ambos mo-
vimientos están separados por cua-
renta años y que uno sucede en Es-
paña y en el otro media con fuerza un
país diferente, no cabe duda de que el
resultado lo es también, debido a que
todavía la sociedad en la que vivimos
tiene prejuicios acerca del origen y el
modo de vestir de los que llegan a Es-
paña. En definitiva, que el desarrollo
de ambos procesos es idéntico, tiene
el mismo esquema, pero el final reve-
la cómo las barreras culturales entre
unos y otros hacen que fracase la in-
tegración actual de las personas que
llegan a España procedentes de otros
países. ... conozco a un profesor, que es de
Marruecos, ha dejado a su mujer, sus
dos hijos y vino hacia aquí, y nada, tra-
baja en la construcción, trabaja en lo
que sea, en coger butanos y cosas así,
y se arrepintió muchísimo por dejar su
trabajo allí... 6.3. El fenómeno de la inmigración Pero es al-
go lógico, todos los seres humanos in-
tentan mejorar su situación siempre
que tengan expectativas de cumplir-
las y la propia historia reciente dice
que no es muy difícil llegar a España,
otra cosa es que una vez en llegados
salgan las cosas bien para seguir me-
jorando, sobre todo cuando aquí, por
ejemplo, el sueldo que cobraba Rabía
cuidando al niño de la señora que la
ayudó cuando la acusaron de robo, era
el salario de un profesor de escuela de
Larache. Ante estos datos, ¿qué ma-
rroquí no quiere llegar a España? Otro de los aspectos que no en-
tiende de la vida de un inmigrante en
España tiene que ver con la propia re-
lación entre estos colectivos. En este
período de tiempo en que Rabía ha es-
tado viviendo en España ha podido
comprobar cómo la envidia y el rencor
entre los propios inmigrantes crece a
un ritmo arrollador. Como ya hemos
comentado, Rabía contaba que fueron
sus propios vecinos de Marruecos los
que la acusaron de robar simplemen-
te por haberse comprado ropa y exhi-
birla ante ellos. También contaba có-
mo sus cuñadas, también marroquíes,
no le ayudaban cuando tuvo el niño y
cómo llegaron a no hablar ni a preo-
cuparse de su hermano cuando em-
pezó a vivir con ella. Por lo tanto, nos encontramos an-
te dos trayectorias: la del inmigrante
que está en España y desea volver por
diferentes motivos; y la del extranjero
que quiere llegar a España invirtiendo
todo su dinero y arriesgando su pro-
pia vida. Todo esto al final se vuelve contra
los propios marroquíes, ya que cuan-
do Rabía y su marido volvían a Lara-
che de vacaciones, la gente del lugar
se alegraba de verlos tan bien, con
dinero, con un coche, con piso, con
ropas nuevas, con muchos objetos
que en Marruecos nunca verían. Esta ... además ellos creen que aquí es
una vida fácil, pero no es así, es una Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 233 Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Instinto de mejora. Diferencias entre dos movimientos migratorios... Juan José Muñoz Juan José Muñoz cruzaron el Estrecho para llegar a Es-
paña tienen serias dificultades para en-
contrar trabajo, ganar dinero, conocer
la otra cultura, adaptarse o integrarse. 7. Puntos en común.
Conclusiones La comparación entre ambos mo-
vimientos se antoja sencilla y fácil de
explicar. Se trata de dos procesos si-
milares, con el mismo origen y similar
desarrollo, pero desigual consolida-
ción y resultado. Si ambos nacen ba-
jo las dificultades de la precariedad
económica y los hábitos de un espa-
cio subdesarrollado, basado en la au-
tosuficiencia, ambos también logran
salir de este ámbito gracias a la emi-
gración y a la búsqueda de mejores
condiciones de vida en otro lugar. Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 Bibliografía Del Campo, S. y Navarro, M. (1987), Nuevo aná-
lisis de la población española, Madrid, Ariel. Haddon, A. C. (1912), The wondering of peoples. Camdbridge. Kasdan, L. (1970), “Introduction” in Migration and
Anthropology. Proceedings of the 1970 Anual
Spring Meeting of the American Etnological
Society, Washington. Ambos difieren en una cuestión
muy importante: el resultado de la mi-
gración. Si todas aquellas personas,
como Mari, que dejaron su pueblo pa-
ra llegar a una gran ciudad tuvieron fa-
cilidades para trabajar, adaptarse, co-
nocer gente y ganar dinero suficiente
para tener una vida digna y autosufi-
ciente, otras personas como Rabía que Mangalam, J. J. (1968), Human migration: a gui-
de to migration literature in enghish (1955-
1962), Lexington. Park, R. E. (1950), “Human migrations an the mar-
ginal man” in Race and Culture, Gleonce. Pérez Díaz, V. (1974), Pueblos y clases sociales
en el campo español, Madrid, Alianza. Ravenstein, E. G. (1885), “The laws of migration”
in Journal of Royal Stat. Society. Tamames, R. (1983), Estructura económica de Es-
paña, vol. 1, Madrid, Alianza Universidad. Cuadernos de Trabajo Social
Vol. 15 (2002): 217-234 234
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https://openalex.org/W4226312452
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https://zenodo.org/records/5947287/files/PolicyCLOUD_D5.5%20Cross-Sector%20Policy%20Lifecycle%20Management.%20Software%20Prototype%202_v1.0.pdf
|
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D5.5 CROSS-SECTOR POLICY LIFECYCLE MANAGEMENT: SOFTWARE PROTOTYPE 2
|
Zenodo (CERN European Organization for Nuclear Research)
| 2,021
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|
CLOUD FOR DATA-DRIVEN POLICY MANAGEMENT
Project Number: 870675
Start Date of Project: 01/01/2020
Duration: 36 months
D5.5 CROSS-SECTOR POLICY LIFECYCLE
MANAGEMENT: SOFTWARE PROTOTYPE 2
Dissemination Level
PU
Due Date of Deliverable
31/10/2021, Month 22
Actual Submission Date
02/11/2021
Work Package
WP5
Cross-sector
Policy
Lifecycle
Management
Task
T5.2, T5.3, T5.4, T5.5 & T5.6
Type
Demonstrator
Approval Status
Version
V0.7
Number of Pages
p.1 – p.42
Abstract: This document provides a description of the software components of the Policy
Management Framework Layer, and Policy Development Toolkit Layer.
The information in this document reflects only the author’s views and the European Community is not liable for any use that may be made
of the information contained therein. The information in this document is provided “as is” without guarantee or warranty of any kind,
express or implied, including but not limited to the fitness of the information for a particular purpose. The user thereof uses the information
at his/ her sole risk and liability. This deliverable is licensed under a Creative Commons Attribution 4.0 International License. CLOUD FOR DATA-DRIVEN POLICY MANAGEMENT
Project Number: 870675
Start Date of Project: 01/01/2020
Duration: 36 months
D5.5 CROSS-SECTOR POLICY LIFECYCLE
MANAGEMENT: SOFTWARE PROTOTYPE 2
Dissemination Level
PU
Due Date of Deliverable
31/10/2021, Month 22
Actual Submission Date
02/11/2021
Work Package
WP5
Cross-sector
Policy
Lifecycle
Management
Task
T5.2, T5.3, T5.4, T5.5 & T5.6
Type
Demonstrator
Approval Status
Version
V0.7
Number of Pages
p.1 – p.42
Abstract: This document provides a description of the software components of the Policy
Management Framework Layer, and Policy Development Toolkit Layer. The information in this document reflects only the author’s views and the European Community is not liable for any use that may be made
of the information contained therein. The information in this document is provided “as is” without guarantee or warranty of any kind,
express or implied, including but not limited to the fitness of the information for a particular purpose. The user thereof uses the information
at his/ her sole risk and liability. This deliverable is licensed under a Creative Commons Attribution 4.0 International License. MANAGEMENT: SOFTWARE PROTOTYPE 2 Dissemination Level
PU
Due Date of Deliverable
31/10/2021, Month 22
Actual Submission Date
02/11/2021
Work Package
WP5
Cross-sector
Policy
Lifecycle
Management
Task
T5.2, T5.3, T5.4, T5.5 & T5.6
Type
Demonstrator
Approval Status
Version
V0.7
Number of Pages
p.1 – p.42 Abstract: This document provides a description of the software components of the Policy
Management Framework Layer, and Policy Development Toolkit Layer. The information in this document reflects only the author’s views and the European Community is not liable for any use that may be made
of the information contained therein. The information in this document is provided “as is” without guarantee or warranty of any kind,
express or implied, including but not limited to the fitness of the information for a particular purpose. The user thereof uses the information
at his/ her sole risk and liability. This deliverable is licensed under a Creative Commons Attribution 4.0 International License. Abstract: This document provides a description of the software components of the Policy
Management Framework Layer, and Policy Development Toolkit Layer. The information in this document reflects only the author’s views and the European Community is not liable for any use that may be made
of the information contained therein. The information in this document is provided “as is” without guarantee or warranty of any kind,
express or implied, including but not limited to the fitness of the information for a particular purpose. The user thereof uses the information
at his/ her sole risk and liability. This deliverable is licensed under a Creative Commons Attribution 4.0 International License. he information in this document reflects only the author’s views and the European Community is not liable for any use that may be
f the information contained therein. The information in this document is provided “as is” without guarantee or warranty of an
xpress or implied, including but not limited to the fitness of the information for a particular purpose. The user thereof uses the infor
t his/ her sole risk and liability. This deliverable is licensed under a Creative Commons Attribution 4.0 International License. The information in this document reflects only the author’s views and the European Community is not liable for any use that may be made
of the information contained therein. MANAGEMENT: SOFTWARE PROTOTYPE 2 The information in this document is provided “as is” without guarantee or warranty of any kind,
express or implied, including but not limited to the fitness of the information for a particular purpose. The user thereof uses the information
at his/ her sole risk and liability. This deliverable is licensed under a Creative Commons Attribution 4.0 International License. PolicyCloud has received funding from the European Union’s Horizon 2020 research and
innovation programme under grant agreement No 870675. D5.5 – v1.0 Versioning and Contribution History Version
Date
Reason
Author
0.1
20/09/2021
Updated ToC and assignments
Armend Duzha
0.2
27/09/2021
Contributions from ICCS
Kostas Moutselos
0.3
04/10/2021
Contributions from LXS
Luis Miguel Garcia
Jesús Manuel Gallego
0.4
11/10/2021
Contributions from ATOS
Ana Luiza Pontual
Miquel Mila Prat
0.5
26/10/2021
Consolidated draft ready for internal review
Armend Duzha
0.6
27/10/2021
Internal Review
Giannis Ledakis (UBI) /
Yosef Moatti (IBM)
0.7
28/10/2021
Addressing review comments
Armend Duzha
0.8
02/11/2021
Quality Check
Argyro Mavrogiorgou
1.0
02/11/2021
Final version ready for submission
Submitted version
Armend Duzha Author List Author List Author List
Organisation
Name
MAG
Armend Duzha
UPRC
Ilias Maglogiannis
ICCS
Kostas Moutselos
LXS
Luis Miguel Garcia
Jesús Manuel Gallego
OKS
Chris Maragkos
Stelios Mpetas
ATOS
Ana Luiza Pontual
Miquel Mila Prat 2 www.policycloud.eu 2 D5.5 – v1.0 www.policycloud.eu Abbreviations and Acronyms Abbreviation/Acronym
Definition
API
Application Programming Interface
AT
Analytical Tool
DSS
Decision Support System
EC
European Commission
KPI
Key Performance Indicator
PDT
Policy Development Toolkit
PM
Policy Model
PME
Policy Modelling Editor
PP
Public Policy
SOA
Service Oriented Architecture
UI
User Interface
UX
User eXperience
UML
Unified Modelling Language
WP
Work Package www.policycloud.eu www.policycloud.eu 3 D5.5 – v1.0 Contents www.policycloud.eu
4
Contents
Versioning and Contribution History .................................................................................................................... 2
Author List ................................................................................................................................................................ 2
Abbreviations and Acronyms ................................................................................................................................ 3
Executive Summary ................................................................................................................................................ 7
1
Introduction ...................................................................................................................................................... 8
1.1
Purpose and Scope ................................................................................................................................ 8
1.2
Summary of changes ............................................................................................................................. 8
1.3
Structure of the Deliverable .................................................................................................................. 8
2
Prototype overview ......................................................................................................................................... 9
2.1
Main components of the prototype ................................................................................................... 10
2.1.1
Policy Modelling Editor .................................................................................................................... 10
2.1.2
Policy Development Toolkit ............................................................................................................ 10
2.1.3
Data Visualization ............................................................................................................................. 11
2.1.4
PDT Backend ..................................................................................................................................... 14
2.2
Interfaces ............................................................................................................................................... 16
2.2.1
Policy Modelling Editor .................................................................................................................... 16
2.2.2
Policy Development Toolkit ............................................................................................................ 20
2.2.3
Data Visualization ............................................................................................................................. 28
2.2.4
PDT Backend ..................................................................................................................................... 30
2.3
Baseline technologies and tools ......................................................................................................... 37
2.3.1
Policy Modelling Editor .................................................................................................................... 37
2.3.2
Policy Development Toolkit ............................................................................................................ 37
2.3.3
Data Visualization ............................................................................................................................. 37
2.3.4
PDT Backend ..................................................................................................................................... 37
3
Source Code ................................................................................................................................................... 38
3.1
Availability.............................................................................................................................................. 38
3.1.1
Policy Modelling Editor .................................................................................................................... 38
3.1.2
Policy Development Toolkit ............................................................................................................ 38
3.1.3
Data Visualization ............................................................................................................................. 38
3.1.4
PDT Backend ..................................................................................................................................... 38 D5.5 – v1.0 D5.5 – v1.0
3.2
Deployment ........................................................................................................................................... 39
3.2.1
Policy Modelling Editor .................................................................................................................... 39
3.2.2
Policy Development Toolkit ............................................................................................................ 39
3.2.3
Data Visualization ............................................................................................................................. 39
3.2.4
PDT Backend ..................................................................................................................................... 39
4
Conclusions .................................................................................................................................................... 41
References ............................................................................................................................................................. 42 3.2
Deployment ........................................................................................................................................... 39
3.2.1
Policy Modelling Editor .................................................................................................................... 39
3.2.2
Policy Development Toolkit ............................................................................................................ 39
3.2.3
Data Visualization ............................................................................................................................. 39
3.2.4
PDT Backend ..................................................................................................................................... 39
4
Conclusions .................................................................................................................................................... 41
References ............................................................................................................................................................. 42 List of Figures Figure 1 - PolicyCloud System Architecture ......................................................................................................... 9
Figure 2 - Example of the Heatmap for Maggioli use case (Event Locator) ................................................... 11
Figure 3 – Example of Heatmap for Maggioli use Case (“Hot” zones) ............................................................ 11
Figure 4 - Example of the line and gauge charts for the sarga use case ....................................................... 12
Figure 5 - Example of the Heatmap for The SOFIA use case ........................................................................... 12
Figure 6 - Example of the BAR CHART for The LONDON use case ................................................................. 13
Figure 7 - Example of BAR CHART for The SOFIA use case .............................................................................. 13
Figure 8 - Example of another BAR CHART for The SOFIA use case ............................................................... 14
Figure 9 - Polciy Model Editor Home Page ......................................................................................................... 16
Figure 10 - Existing KPIs selection ....................................................................................................................... 17
Figure 11 - Add New KPI ....................................................................................................................................... 17
Figure 12 - Add New Actor ................................................................................................................................... 18
Figure 13 - Analytical Tool selection ................................................................................................................... 18
Figure 14 - Set parameter value .......................................................................................................................... 19
Figure 15 - Review and Submit ............................................................................................................................ 19
Figure 16 - Policy Model Submitted .................................................................................................................... 20
Figure 17 – Policy selection-evaluation wizard .................................................................................................. 20
Figure 18 - Policy selection and policy properties ............................................................................................ 21
Figure 19 - KPI selecion......................................................................................................................................... 22
Figure 20 – KPI properties .................................................................................................................................... 22
Figure 21 – Evaluation – Analytics tool selection .............................................................................................. 23
Figure 22 – Analytics tool technical details ........................................................................................................ 23
Figure 23 – Analytics tool parameters & submission ...................................................................................... 24
Figure 24 – Analytics tool parameters details ................................................................................................... 25
Figure 25 – Policy overview / saving ................................................................................................................... 25
Figure 26 – PDT components menu ................................................................................................................... 26
Figure 27 - ONLINE help ....................................................................................................................................... 26 5 5 D5.5 – v1.0 Figure 28 - User Data protection information notice ....................................................................................... 27
Figure 29 - User data protection text ................................................................................................................. 27
Figure 30 - User profile properties / logout ....................................................................................................... 28
Figure 31 - Example for heat map chart data .................................................................................................... 29
Figure 32 – Example for Sarga Gauge Chart Data ............................................................................................ 29
Figure 33 – Example for London Bar Chart data............................................................................................... 29
Figure 34 – Example for One Bar Data for Sofia ............................................................................................... 30
Figure 35 - Web Methods related with Actors ................................................................................................... 32
Figure 36 - WEB METHODS RELATED WITH Analytical Tools ........................................................................... 33
Figure 37 - WEB METHODS RELATED WITH Data Models ................................................................................ 33
Figure 38 - WEB METHODS RELATED WITH Domains ...................................................................................... 33
Figure 39 - WEB METHODS RELATED WITH Goals ............................................................................................ www.policycloud.eu List of Figures 34
Figure 40 - WEB METHODS RELATED WITH Job Status..................................................................................... 34
Figure 41 - WEB METHODS RELATED WITH JOBs .............................................................................................. 35
Figure 42 - WEB METHODS RELATED WITH KPIS .............................................................................................. 35
Figure 43 - WEB METHODS RELATED WITH Policies ......................................................................................... 36 www.policycloud.eu www.policycloud.eu 6 6 D5.5 – v1.0 www.policycloud.eu 1.1 Purpose and Scope This document is the second iteration of “Cross-Sector Policy Lifecycle Management: Software prototype”,
and covers tasks T5.2, T5.3, T5.4, T5.5 and T5.6. Its main purpose is to provide a description of the
prototype implementation of the Policy Modelling Editor (PME) and the Policy Development Toolkit (PDT)
to support policy makers in the modelling, creation, update, and evaluation of the policies. This document is consistent with deliverable D5.4 “Cross-Sector Policy Lifecycle Management: Design and
Open Specification 2” [1], delivered in August 2021, which provided the overall architecture and design
specifications of the PME and PDT. This report will be further updated during the last phase of the project duration (at month M34, October
2022) and will include further advancements and contributions from other tasks. Executive Summary This document is an incremental update of deliverable D5.3 Cross-sector Policy Lifecycle Management:
Software Prototype 1, delivered in November 2020, and describes in detail the software components of
the Policy Management Framework Layer and Policy Development Toolkit Layer, which are the frontend
parts of the PolicyCLOUD platform and allow policy makers to create, update and evaluate policies. For
each component, a description of its APIs, specification of the main functionalities, and description of the
source code are provided. 7 www.policycloud.eu www.policycloud.eu 7 D5.5 – v1.0 1.2 Summary of changes The main changes in this deliverable regard the update of Section 2.1 and 2.2. In this second version of
the demonstrator, all components have been integrated and are running on the testbed infrastructure. Furthermore, integration work has been performed for the Maggioli and Sarga use cases, which are now
integrated with the Policy Development Toolkit layer. In addition, we have updated and revised the whole document (including figures) to reflect the current
work done at M22 (October 2021). 1.3 Structure of the Deliverable 1.3 Structure of the Deliverable The rest of the document is structured as follows: The rest of the document is structured as follows: •
Section 2 introduces the main components and sub-systems, their interfaces and technologies
used for their development. •
Section 2 introduces the main components and sub-systems, their interfaces and technologies
used for their development. •
Section 3 describes where the source code is made available. •
Finally, Section 4 provides the conclusion of the document. 8 8 www.policycloud.eu D5.5 – v1.0 2 Prototype overview In this section we provide the description of the prototype implementation of the Policy Modelling
Framework Layer and Policy Development Toolkit Layer, as illustrated in Figure 1. FIGURE 1 - POLICYCLOUD SYSTEM ARCHITECTURE FIGURE 1 - POLICYCLOUD SYSTEM ARCHITECTURE 9 www.policycloud.eu www.policycloud.eu 9 D5.5 – v1.0 2.1.1 Policy Modelling Editor The Policy Model Editor (PME) is the core component that supports and guides the end-user to effectively
create a Policy Model safely. More specifically, the PME is a Single Page Application that relies on the PDT
backend REST API to fetch or store related entities of the Policy Model. 2.1 Main components of the prototype 2.1.1 Policy Modelling Editor 2.1.2 Policy Development Toolkit As described in the deliverable D2.2 “Conceptual Model and Reference Architecture”[2], PDT interacts
with components from Layer 3 - Policies Management Framework (T5.1, T5.2), Analytical Tools (WP4) and
the PDT Backend. PDT provides the integrated prototype of the following components as described in the deliverable D5.4
“Cross-sector Policy Lifecycle Management: Design and Open Specification 2” [1]: •
Policy Model Viewer •
Policy Evaluations / Analytics Integration with the Backend has been implemented, utilising Docker images. This way, PDT can retrieve
default policy models in the structured JSON format which represent the Policy Models (PMs). So, the
functionalities included in this prototype are: Integration with the Backend has been implemented, utilising Docker images. This way, PDT can retrieve
default policy models in the structured JSON format which represent the Policy Models (PMs). So, the
functionalities included in this prototype are: •
The policy maker can obtain a list of available policies and select one for further ex •
Once a policy is selected, the properties of the policy are shown, along with the available
the policy, •
Once a KPI is selected, the relative KPI properties are also shown to the user, •
The Analytical Tools (ATs) which can compute the selected KPI are presented. The policy maker
can see the parameters related with each AT and invoke the respective AT. Finally, the policy
maker can see a short overview of the current selected PMs. The aforementioned functionalities are implemented as a policy wizard component, comprised by three
steps. The relative UI components are described in Section 2.2.2. The relative UI components are described in Section 2.2.2. www.policycloud.eu 10 D5.5 – v1.0 2.1.3 Data Visualization The Data Visualization component has been successfully integrated with the PDT. This second prototype
includes all graphs defined in the first prototype and now can be seen through the PDT. They are: •
A Heatmap as requested by the “Participatory Policies Against Radicalization” Use Case of
Maggioli. This chart uses data extracted from the Data Analytics of WP4. •
A Heatmap as requested by the “Participatory Policies Against Radicalization” Use Case of
l
h
h
d
d f
h
l
f FIGURE 2 - EXAMPLE OF THE HEATMAP FOR MAGGIOLI USE CASE (EVENT LOCATOR) FIGURE 2 - EXAMPLE OF THE HEATMAP FOR MAGGIOLI USE CASE (EVENT LOCATOR) This Heatmap is intended to show, at a glance, all incidents that occurred in a chosen region. Currently,
there are two possible regions: “North America” and “Eastern Europe”; but the list will be enlarged in final
version. Whenever a region is selected, the heatmap shows all incidents that took place in it. It can be
filtered by location and by year, and it is possible to see the incident details, when clicking on it. Users
can change regions as desired. The heat map can also show “hot” zones (zones with a bigger number of
incidents) using colours, instead of numbers: FIGURE 3 – EXAMPLE OF HEATMAP FOR MAGGIOLI USE CASE (“HOT” ZONES) FIGURE 3 – EXAMPLE OF HEATMAP FOR MAGGIOLI USE CASE (“HOT” ZONES) www.policycloud.eu 11 D5.5 – v1.0 •
A Line and Gauge charts as requested by the “Intelligent Policies for Denomination of Origin” Use
Case of Sarga, that plots the evaluation of the Sentimental Analysis considering some Keywords. •
A Line and Gauge charts as requested by the “Intelligent Policies for Denomination of Origin” Use
Case of Sarga, that plots the evaluation of the Sentimental Analysis considering some Keywords. •
A Line and Gauge charts as requested by the Intelligent Policies for Denomination of Origin Use
Case of Sarga, that plots the evaluation of the Sentimental Analysis considering some Keywords. FIGURE 4 - EXAMPLE OF THE LINE AND GAUGE CHARTS FOR THE SARGA USE CASE FIGURE 4 - EXAMPLE OF THE LINE AND GAUGE CHARTS FOR THE SARGA USE CASE Considering a set of keywords, the Sentiment Analysis Line chart shows the sentiments for these
keywords during a given period. The Accumulated Sentiment Gauge chart shows the average of these
sentiments, during the same period. 2.1.3 Data Visualization In order to achieve the internal milestone of having at least one chart by Use Case scenario, the
visualization component has been incremented and has integrated other charts. They are: In order to achieve the internal milestone of having at least one chart by Use Case scenario, the
visualization component has been incremented and has integrated other charts. They are: •
A heatmap, requested by the “Facilitating urban policy making and monitoring through
crowdsourcing data analysis” Use Case of Sofia. In this case, an adaptation of previous Heatmap
has been done, following Sofia Use Case’s requirements. In this way, the adapted Heatmap is
focused on Sofia city and can filter by Sofia’s districts, by year, and, also, differently from Maggioli
Use Case, by months and days. FIGURE 5 - EXAMPLE OF THE HEATMAP FOR THE SOFIA USE CASE FIGURE 5 - EXAMPLE OF THE HEATMAP FOR THE SOFIA USE CASE www.policycloud.eu 12 D5.5 – v1.0 •
A Bar Chart, required both for Sofia and London Use Cases. In the case of London, it is intended
to show unemployment data by gender: •
A Bar Chart, required both for Sofia and London Use Cases. In the case of London, it is intended
to show unemployment data by gender: •
A Bar Chart, required both for Sofia and London Use Cases. In the case of London, it is intended
to show unemployment data by gender: FIGURE 6 - EXAMPLE OF THE BAR CHART FOR THE LONDON USE CASE FIGURE 6 - EXAMPLE OF THE BAR CHART FOR THE LONDON USE CASE FIGURE 6 - EXAMPLE OF THE BAR CHART FOR THE LONDON USE CASE And for Sofia, the chart represents the number of incidents per month and per district (the 24 districts
of Sofia): FIGURE 7 - EXAMPLE OF BAR CHART FOR THE SOFIA USE CASE FIGURE 7 - EXAMPLE OF BAR CHART FOR THE SOFIA USE CASE FIGURE 7 - EXAMPLE OF BAR CHART FOR THE SOFIA USE CASE Both are the same Bar charts, with different data, but that can be seen in two different formats: cluster
and stacked. Both are the same Bar charts, with different data, but that can be seen in two different formats: cluster
and stacked. 2.1.3 Data Visualization 13 www.policycloud.eu D5.5 – v1.0
Also, for Sofia Use Case, there’s another Bar Chart, where the data that can be visualized is the number
of incidents by subcategories per districts: D5.5 – v1.0 Also, for Sofia Use Case, there’s another Bar Chart, where the data that can be visualized is the number
of incidents by subcategories per districts: Also, for Sofia Use Case, there’s another Bar Chart, where the data that can be visualized is the number
of incidents by subcategories per districts: FIGURE 8 - EXAMPLE OF ANOTHER BAR CHART FOR THE SOFIA USE CASE FIGURE 8 - EXAMPLE OF ANOTHER BAR CHART FOR THE SOFIA USE CASE 2.1.4 PDT Backend Those tools can be retrieved either by name, by the domain they are related to or
according to the datasets that they are compatible with www.policycloud.eu 14 D5.5 – v1.0 •
Retrieve information about the available datasets, stored in the central data repository of
PolicyCLOUD and managed by the Data Acquisition and Analytic layer
•
Add, remove, modify and retrieve domains that can be relevant to the entities of the policies
•
Add, remove, modify and retrieve goals of a KPI, and associate those tools with stakeholders
•
Check the status of a job, which has been submitted as the result of the invocation of an analytical
tool
•
Get information about a job, submitted by the invocation of an analytical tool
•
Get the result of an analysis, related to its job
•
Add, remove, modify and retrieve KPIs, based on their name, their corresponding domain or their
relevant actors
•
Add or remove actors, goals, domains and data models from a KPI
•
Add, remove, modify and retrieve policies, according to their names, the user that stored the
policy or their corresponding domain they belong to. •
Add or remove KPIs from policies
•
Store combined types of results of analytical tools, thus giving the opportunity to the end-user to
switch between different views of the same results
•
Correlate policies with the end-users that have defined them
•
Retrieve policies depending on the user that has issued the request
•
Correlate jobs with the users that initially submitted a request to execute an analysis
•
Retrieve the results of the jobs owned by a given end-users
•
Cancel the execution of an analysis if it exceeds the defined maximum response time allowed
•
Provides a warning when the result of an analysis exceeds the maximum number of characters
that can be efficiently processed by the visualization tools at a later phase Regarding the interaction with the Data Acquisition and Analytic layer, the corresponding APIs have been
defined in deliverable D4.4. The Backend of the PDT is integrated with the latter, in order to implement
the higher-level functionality, as described above. The backend of the PDT consists of a single process that runs on a servlet container. It relies on a
relational datastore to store the relational model of the policy, its attributes, related entities, and relations
between them. 2.1.4 PDT Backend The PDT backend provides support to the whole PDT, and can be considered as the layer between the
user interfaces that the policy maker can use to have access the tool, the policy modelling editor, and the
Data Acquisition and Analytics layer of the PolicyCLOUD platform, provided by the components of WP4
and WP3. It exposes its functionality via REST APIs and web sockets, to allow for the integration with the
aforementioned components, while it also makes use of the REST API provided by the Data Acquisition
and Analytics layer. As a result, it serves four main purposes: •
Allows storing, modifying and retrieving policies, along with their relevant corresponding entities
(i.e., KPIs, stakeholders, etc.) (i.e., KPIs, stakeholders, etc.) •
Hold policy meta-information that can be used by the policy modelling editor, in order for the
latter to propose new policies to the domain experts •
Invoke an analytical tool to perform an analysis through the Data Acquisition and Analytics layer,
given the input parameters defined by the KPIs of the involved policy •
Allow for the retrieval of the results of the analytical tools, which can be later fed to the
visualization component of the tool. This involves again the integration of the backend with the
Data Acquisition and Analytics layer More precisely, regarding the functionality implemented at this phase of the project for the PDT and the
Policy Model Editor, the backend provides the ability to: More precisely, regarding the functionality implemented at this phase of the project for the PDT and the
Policy Model Editor, the backend provides the ability to: •
Store, and retrieve actors, based on their name or the domain they are related to •
Retrieve information about the Analytical Tools that are available in the Data Acquisition and
Analytic layer. 2.1.4 PDT Backend It should be highlighted that the datastore which is being used by this component is not
the central data repository of PolicyCLOUD, as it is used for storing the raw data coming from the data
providers, along with the results of the analytical tools. The purpose of the data storage element of this
component is to store the policies and the metadata information that are related with those, along with
the results of the analytical tools, while at the same time it aims at providing a standard way for
communicating with the backend. www.policycloud.eu 15 D5.5 – v1.0 2.2.1 Policy Modelling Editor Upon the end-user’s entrance into the PME, the end-user has to both indicate the domain of its Policy
Model and to provide a short description. Furthermore, the end-user has to provide to the PME the
existing KPIs that might fit in the Policy Model under consideration. Figure 9 illustrates the Home Page interface of the PME. It fetches from the PDT backend KPIs that belong
in the same domain with the Policy Model; and then prompted by the component to continue in the next
page to select existing KPIs. FIGURE 9 - POLCIY MODEL EDITOR HOME PAGE
Figure 10 shows the Existing KPIs selection interface, in which the end-user has the capability to add or 2.2 Interfaces 2.2.1 Policy Modelling Editor 2.2.1 Policy Modelling Editor FIGURE 9 - POLCIY MODEL EDITOR HOME PAGE Figure 10 shows the Existing KPIs selection interface, in which the end-user has the capability to add or
to select existing Actors that are included in the Policy Model domain. www.policycloud.eu 16 www.policycloud.eu D5.5 – v1.0 D5.5 – v1.0 D5.5 – v1.0 FIGURE 10 - EXISTING KPIS SELECTION FIGURE 10 - EXISTING KPIS SELECTION In the Add new KPI interface (see Figure 11) the end-user has the capability to create a new KPI; while
he/she can enter the relevant info of the KPI, describe its Goals, and integrate the additional
Stakeholders. FIGURE 11 - ADD NEW KPI FIGURE 11 - ADD NEW KPI
It should be noted that the user has the following capabilities: (i) to provide description of the Data Model
and add or select Actors, (ii) to select the relevant Analytical Tools from a specific list (drop-down menu)
(Figure 12), (iii) to provide the parameter values that comes for the Analytical Tools (Figure 13), (iv) to
review and submit the Policy Model (Figure 14). FIGURE 11 - ADD NEW KPI It should be noted that the user has the following capabilities: (i) to provide description of the Data Model
and add or select Actors, (ii) to select the relevant Analytical Tools from a specific list (drop-down menu)
(Figure 12), (iii) to provide the parameter values that comes for the Analytical Tools (Figure 13), (iv) to
review and submit the Policy Model (Figure 14). It should be noted that the user has the following capabilities: (i) to provide description of the Data Model
and add or select Actors, (ii) to select the relevant Analytical Tools from a specific list (drop-down menu)
(Figure 12), (iii) to provide the parameter values that comes for the Analytical Tools (Figure 13), (iv) to
review and submit the Policy Model (Figure 14). 17 www.policycloud.eu www.policycloud.eu D5.5 – v1.0 FIGURE 12 - ADD NEW ACTOR
FIGURE 13 - ANALYTICAL TOOL SELECTION FIGURE 12 - ADD NEW ACTOR FIGURE 13 - ANALYTICAL TOOL SELECTION FIGURE 13 - ANALYTICAL TOOL SELECTION www.policycloud.eu 18 D5.5 – v1.0 FIGURE 14 - SET PARAMETER VALUE
Finally, if the process was successful the user is redirected to the last page that promotes the user to visit
the PDT and evaluate the submitted Policy Model or compose a new one. FIGURE 15 - REVIEW AND SUBMIT FIGURE 14 - SET PARAMETER VALUE FIGURE 14 - SET PARAMETER VALUE FIGURE 14 - SET PARAMETER VALUE Finally, if the process was successful the user is redirected to the last page that promotes the user to visit
the PDT and evaluate the submitted Policy Model or compose a new one. www policycloud eu
19
FIGURE 15 - REVIEW AND SUBMIT www.policycloud.eu 19 D5.5 – v1.0 2.2.2 Policy Development Toolkit 2.2.2 Policy Development Toolkit The following figures present the UI implementations of the PDT functionalities, as listed in Section 2.1.2. The web pages are served locally communicating with the dockerised backend API. Figure 17 shows the landing web page of PDT (after the pending Authentication procedure). The page
shows the three-step policy wizard, with the first step as the default location. Here the user selects a
policy from a list of available policy models. FIGURE 17 – POLICY SELECTION-EVALUATION WIZARD www.policycloud.eu 20 D5.5 – v1.0 Figure 18 shows the properties of the policy model selected by the user. After the selection of a policy,
the related KPIs are displayed, and the PM can select one KPI for evaluation. Figure 18 shows the properties of the policy model selected by the user. After the selection of a policy,
the related KPIs are displayed, and the PM can select one KPI for evaluation. FIGURE 18 - POLICY SELECTION AND POLICY PROPERTIES FIGURE 18 - POLICY SELECTION AND POLICY PROPERTIES Figure 18 displays the step for the KPI selection and Figure 19 shows the properties of the selected KPI,
e.g., formula, data, domain, goal, actors and stakeholders. 21 www.policycloud.eu www.policycloud.eu D5.5 – v1.0
FIGURE 19 - KPI SELECION
FIGURE 20 – KPI PROPERTIES D5.5 – v1.0
FIGURE 19 - KPI SELECION D5.5 – v1.0 D5.5 – v1.0 FIGURE 19 - KPI SELECION 22 www.policycloud.eu www.policycloud.eu D5.5 – v1.0 Figure 21 shows the third step of the wizard, which is the evaluation of the selected KPI, by using the
proper Analytics Tool. Figure 21 shows the third step of the wizard, which is the evaluation of the selected KPI, by using the
proper Analytics Tool. FIGURE 21 – EVALUATION – ANALYTICS TOOL SELECTION FIGURE 21 – EVALUATION – ANALYTICS TOOL SELECTION FIGURE 21 – EVALUATION – ANALYTICS TOOL SELECTION Figure 22 shows technical details of the Analytics Tool in an expandable card, as to not distract the PM. Figure 22 shows technical details of the Analytics Tool in an expandable card, as to not dis FIGURE 22 – ANALYTICS TOOL TECHNICAL DETAILS FIGURE 22 – ANALYTICS TOOL TECHNICAL DETAILS 23 www.policycloud.eu D5.5 – v1.0 Figure 23 shows the parameter list that the selected Analytics Tool is expecting for invocation. A
parameter’s values overview is also presented, which includes the default values for the calculation of
the KPI. . FIGURE 23 – ANALYTICS TOOL PARAMETERS & SUBMISSION FIGURE 23 – ANALYTICS TOOL PARAMETERS & SUBMISSION 24 www.policycloud.eu www.policycloud.eu D5.5 – v1.0 Figure 24 shows the details for each parameter of the Analytical tool, also in the form of an expandable
card. FIGURE 24 – ANALYTICS TOOL PARAMETERS DETAILS e 25 shows the fourth step of the wizard, presenting a short overview of the selected policy FIGURE 25 – POLICY OVERVIEW / SAVING FIGURE 25 – POLICY OVERVIEW / SAVING 25 www.policycloud.eu D5.5 – v1.0 Figure 26 shows the UI Menu for selecting different PDT components. Figure 26 shows the UI Menu for selecting different PDT components. Figure 26 shows the UI Menu for selecting different PDT components. FIGURE 26 – PDT COMPONENTS MENU
For each step of the wizard help menu is available. Figure 27 shows the online help for the 1st step of the
wizard. Figure 26 shows the UI Menu for selecting different PDT components. FIGURE 26 – PDT COMPONENTS MENU
For each step of the wizard help menu is available. Figure 27 shows the online help for the 1st step of the
wizard FIGURE 26 – PDT COMPONENTS MENU For each step of the wizard help menu is available. Figure 27 shows the online help for the 1st step of the
wizard. For each step of the wizard help menu is available. Figure 27 shows the online help for the 1st step of the
wizard. FIGURE 27 - ONLINE HELP FIGURE 27 - ONLINE HELP www.policycloud.eu 26 D5.5 – v1.0 At the footnote of each PDT page, the link for the End Users Data Protection Information has been added,
as shown in Figure 28 and Figure 29. FIGURE 28 - USER DATA PROTECTION INFORMATION NOTICE At the footnote of each PDT page, the link for the End Users Data Protection Information has been added,
as shown in Figure 28 and Figure 29. FIGURE 28
USER DATA PROTECTION INFORMATION NOTICE FIGURE 28 - USER DATA PROTECTION INFORMATION NOTICE FIGURE 28 USER DATA PROTECTION INFORMATION NOTICE
FIGURE 29 - USER DATA PROTECTION TEXT FIGURE 29 - USER DATA PROTECTION TEXT 27 www.policycloud.eu D5.5 – v1.0 A user-friendly guide / FAQ on how PolicyCLOUD users should address each parameter required from
them for the registration (in particular, those specified in order to ensure legal/ethical compliance) is
being developed as part of the implementation of the WP4 Legal/Ethical Checklist. After the authentication of the User by providing his/her credentials to the Authentication server, the
User Profile is available to the PDT page. Figure 30 shows the User Profile, as well as the Logout action
button. FIGURE 30 - USER PROFILE PROPERTIES / LOGOUT FIGURE 30 - USER PROFILE PROPERTIES / LOGOUT www.policycloud.eu 2.2.3 Data Visualization From the initial page, users will select the different options offered by the tool, and among them, the
graphics to visualize the results of the selected analytics. When a HeatMap is selected to visualize the analytics results, depending on the Use Case, it will receive
the data in one or another format. These data format has been changed to respect the initial prototype
of the Visualization Tool, but has been also agreed with WP4, and contain data for both, Maggioli and
Sofia Use Cases. The json is like: www.policycloud.eu 28 www.policycloud.eu D5.5 – v1.0 FIGURE 31 - EXAMPLE FOR HEAT MAP CHART DATA FIGURE 31 - EXAMPLE FOR HEAT MAP CHART DATA FIGURE 31 - EXAMPLE FOR HEAT MAP CHART DATA Where data field is the data returned by the Analytics Tool, and the other fields are added by the PDT
Backend, which is the one that does the communication between Analytics and the Visualization. The input data used for Sarga Use Case has been provided by the Sentimental Analysis component from
WP4, and the agreed format is: Backend, which is the one that does the communication between Analytics and the Visualization. The input data used for Sarga Use Case has been provided by the Sentimental Analysis component from
WP4, and the agreed format is: FIGURE 32 – EXAMPLE FOR SARGA GAUGE CHART DATA In the case of the Bar chart for London Use Case, the data received has the format: FIGURE 33 – EXAMPLE FOR LONDON BAR CHART DATA 29 www.policycloud.eu D5.5 – v1.0 In the case of Sofia’s Bar charts, the data received has the format: In the case of Sofia’s Bar charts, the data received has the format: @Path("/starts")
@ApiOperation(value="Returns the list of domains that starts with the given input",
notes="Returns the list of domains that starts with the given input",
responseContainer = "List", response = DomainDTOImpl.class)
@ApiResponses(value= {
@ApiResponse(code=500, message="Exception's message")
})
@Produces(MediaType.APPLICATION JSON) @Path("/starts") 2.2.4 PDT Backend As described in the previous subsection, the backend of the PDT exposes its interfaces via a REST API. In
order to provide an always up-to-date documentation of the definition of the interface to the developers
of other components, we automated this process by making use of the swagger [3] open source
documentation UI. Swagger allows the developer to annotate her source code and provides tools and
plugins that auto-generate the documentation during compilation time. It generates JSON files with all
the relevant information. It also provides a web application that can be used as the UI tool for the
consumer of the REST services to see their documentation. This web application makes use of those
auto-generated JSON files to build the user interface. Moreover, it provides to the REST service consumer
the ability to test the interface online, by invoking them in real-time and check the results. The following code snippet indicates how the developer of a REST service can annotate the latter via the
swagger framework. @Path("domains")
@Api(basePath="http://localhost:54735/pdt", value = "/domains", description = "REST Service that
provides functionalities for domains")
public class DomainResource { @Path("domains")
@Api(basePath="http://localhost:54735/pdt", value = "/domains", description = "REST Service that
provides functionalities for domains")
public class DomainResource { The class DomainResource implements the web methods related with the functionalities for the
domains, and it has been annotated to use the REST path “/domains”. As it can be noted, the developer
can additionally annotate the class with the @Api that informs the swagger that this REST resource should
be documented. Moreover, at the level of web methods, the following code snippet shows how the developer can
document the details of each invocation 30 www.policycloud.eu D5.5 – v1.0 @SuppressWarnings("UseSpecificCatch")
public Response getDomainsStartingWith(
@ApiParam(value = "the start with parameter", required = true)
@QueryParam("name") String name) { @SuppressWarnings("UseSpecificCatch")
public Response getDomainsStartingWith(
@ApiParam(value = "the sta
@QueryParam("name") String This web method that returns all domains whose name starts with a given value. The swagger annotation
indicates to the framework that this should be an operation that will return a List of serialized objects of
the DomainDTOImpl class. Besides the default HTTP code 200 that indicates a successful invocation, this
web method can also return an HTTP error code 500 and provides the parameter that is required. 2.2.4 PDT Backend Swagger, uses Java Reflection to retrieve additional information like the type of the invocation (GET, POST,
PUT etc.) the name and type of the input parameters, the media type used in the body of the request
or/and the response of the HTTP call etc. Finally, the DTOs that will be transferred via the REST calls, can also be annotated, as the following code
snippet indicates. Finally, the DTOs that will be transferred via the REST calls, can also be annotated, as the following code
snippet indicates. @JsonIgnoreProperties(ignoreUnknown = true) @JsonIgnoreProperties(ignoreUnknown = true)
public class DomainDTOImpl implements DomainDTO, Serializable {
private static final long serialVersionUID = 1L;
@ApiModelProperty(value="domain id", required=true)
private Long id;
@ApiModelProperty(value="domain name", required=true)
private String name;
@JsonCreator
public DomainDTOImpl(
@JsonProperty("id") Long id,
@JsonProperty("name") String name) {
this.id = id;
this.name = name;
} public class DomainDTOImpl implements DomainDTO, Serializable { private static final long serialVersionUID = 1L; private static final long serialVersionUID = 1L; The aforementioned DomainDTOImpl used by the web method explained before, has two attributes, the
id and the name of the domain. The ApiModelProperty can be used by swagger to provide this kind of
information to the documentation. After having annotated all our web services, web methods and DTOs, the swagger plugin for maven was
used to auto-generate the stub information in JSON files. The following code snippet was used to do this
work. <plugin>
<groupId>com.github.kongchen</groupId>
<artifactId>swagger-maven-plugin</artifactId>
<version>2.3.4</version>
<configuration>
<apiSources>
<apiSource>
<locations>eu.policycloud.rest.resources</locations>
<apiVersion>${project.version}</apiVersion>
<basePath>/resources</basePath> <groupId>com.github.kongchen</groupId> www.policycloud.eu 31 D5.5 – v1.0 <swaggerDirectory>${project.basedir}/src/main/webapp/</swaggerDirectory>
<outputTemplate>${project.basedir}/src/main/resources/markdown.mustache</outputTemplate>
<mustacheFileRoot>${project.basedir}/src/main/resources/</mustacheFileRoot>
<outputPath>${project.basedir}/src/main/webapp/document.html</outputPath>
</apiSource>
</apiSources>
</configuration>
<executions>
<execution>
<phase>compile</phase>
<goals>
<goal>generate</goal>
</goals>
</execution>
</executions>
</plugin> <swaggerDirectory>${project.basedir}/src/main/webapp/</swaggerDirectory>
<outputTemplate>${project.basedir}/src/main/resources/markdown.mustache</outputTemplate>
<mustacheFileRoot>${project.basedir}/src/main/resources/</mustacheFileRoot>
<outputPath>${project.basedir}/src/main/webapp/document.html</outputPath>
</apiSource>
</apiSources>
</configuration>
<executions>
<execution>
<phase>compile</phase>
<goals>
<goal>generate</goal>
</goals>
</execution>
</executions>
</plugin> It creates the JSON files during compile phase, and adds them to corresponding directory where the
swagger app is expecting to retrieve this information, while it also modifies the markdown mustache
template used by the web app. After setting up everything, the online documentation is now available. The following screenshots show
this documentation per REST Web Service implemented. FIGURE 35 - WEB METHODS RELATED WITH ACTORS FIGURE 35 - WEB METHODS RELATED WITH ACTORS 32 www.policycloud.eu D5.5 – v1.0 FIGURE 36 - WEB METHODS RELATED WITH ANALYTICAL TOOLS FIGURE 36 - WEB METHODS RELATED WITH ANALYTICAL TOOLS FIGURE 36 - WEB METHODS RELATED WITH ANALYTICAL TOOLS
FIGURE 37 - WEB METHODS RELATED WITH DATA MODELS FIGURE 36 - WEB METHODS RELATED WITH ANALYTICAL TOOLS
FIGURE 37 - WEB METHODS RELATED WITH DATA MODELS FIGURE 37 - WEB METHODS RELATED WITH DATA MODELS FIGURE 38 - WEB METHODS RELATED WITH DOMAINS FIGURE 38 - WEB METHODS RELATED WITH DOMAINS FIGURE 38 - WEB METHODS RELATED WITH DOMAINS 33 www.policycloud.eu D5.5 – v1.0
FIGURE 39 - WEB METHODS RELATED WITH GOALS
FIGURE 40 - WEB METHODS RELATED WITH JOB STATUS D5.5 – v1.0 www.policycloud.eu FIGURE 39 - WEB METHODS RELATED WITH GOALS FIGURE 39 - WEB METHODS RELATED WITH GOALS FIGURE 39 - WEB METHODS RELATED WITH GOALS
FIGURE 40 - WEB METHODS RELATED WITH JOB STATUS FIGURE 39 - WEB METHODS RELATED WITH GOALS
FIGURE 40 - WEB METHODS RELATED WITH JOB STATUS FIGURE 40 - WEB METHODS RELATED WITH JOB STATUS 34 www.policycloud.eu www.policycloud.eu D5.5 – v1.0
FIGURE 41 - WEB METHODS RELATED WITH JOBS
FIGURE 42 - WEB METHODS RELATED WITH KPIS D5.5 – v1.0 FIGURE 41 - WEB METHODS RELATED WITH JOBS FIGURE 41 - WEB METHODS RELATED WITH JOBS FIGURE 41 - WEB METHODS RELATED WITH JOBS FIGURE 41 - WEB METHODS RELATED WITH JOBS
FIGURE 42 - WEB METHODS RELATED WITH KPIS FIGURE 41 - WEB METHODS RELATED WITH JOBS
FIGURE 42 - WEB METHODS RELATED WITH KPIS FIGURE 42 - WEB METHODS RELATED WITH KPIS 35 www.policycloud.eu D5.5 – v1.0 www.policycloud.eu FIGURE 43 - WEB METHODS RELATED WITH POLICIES FIGURE 43 - WEB METHODS RELATED WITH POLICIES FIGURE 43 - WEB METHODS RELATED WITH POLICIES Finally, the Backend allows for asynchronous communication with the PDT via web sockets. This is crucial
when other components perform updates that the Backend should be aware of, so that the PDT can
modify the UI in order to reflect those updates, without having to continuously invoke the Backend to
ask for potential pending updates. This helps to improve the overall UX of the end-user. For instance,
when an analytical tool finishes its executions and produces some results, the Data Acquisition and
Analytic API notifies the Backend, which will be triggered in order to send a message via this web socket
to the PDT. By doing this, the PDT can be in position to update the status of a pending job, without
periodically ask this status from the backend. The DTOs that are exchanged have the following format: {
"id": 3242,
"notification": "UPDATED",
"job": {
"id": 3242,
"name": "job name",
"description": "the description of the job",
"status": "FINISHED",
"dateCreated": "2021-10-01T09:23:43" {
"id": 3242,
"notification": "UPDATED",
"job": {
"id": 3242,
"name": "job name",
"description": "the description of the job",
"status": "FINISHED",
"dateCreated": "2021-10-01T09:23:43"
. . . . }
}
In this DTO, the id is the unique identifier of the job its status has been changed and the notification can
accept one of the following values: ADDED, UPDATED or DELETED. Finally, the job contains all information
about the corresponding job, like its id, name, description, status, etc. "description": "the description of the job" }
In this DTO, the id is the unique identifier of the job its status has been changed and the notification can
accept one of the following values: ADDED, UPDATED or DELETED. Finally, the job contains all information
about the corresponding job, like its id, name, description, status, etc. 36 D5.5 – v1.0 2.3.3 Data Visualization As the Data Visualization is part of the PDT, it has been developed using the same base technology,
AngularJS v10 [4]. The main JavaScript libraries used for this first prototype are: As the Data Visualization is part of the PDT, it has been developed using the same base technology,
AngularJS v10 [4]. The main JavaScript libraries used for this first prototype are: •
Leaflet [7] is used to plot the heatmap •
Amcharts [8] is used to plot the line and the gauge charts Although the component is integrated into the PDT, a standalone version can also be used, and the
detailed invocation instructions are detailed into the readme file that accompanies the source code of
the component. Still, the service can be started either with a NodeJS server or using a Docker container
(see Section 3.1.3). Although the component is integrated into the PDT, a standalone version can also be used, and the
detailed invocation instructions are detailed into the readme file that accompanies the source code of
the component. Still, the service can be started either with a NodeJS server or using a Docker container
(see Section 3.1.3). To visualize it as part of the PDT, this component has to be started and all previous steps must be taken. 2.3.2 Policy Development Toolkit As described in the deliverable D5.2 “Cross-sector Policy Lifecycle Management: Design and Open
Specification 1” Section 4.2, PDT is built as a Single Page Application developed using the open-source
Angular framework [4][3]. For UI components and controls we use the Angular Material library [5]. The PDT is using Docker to encapsulate the web serving functionality along with NGINX [6] web server. The PDT is using Docker to encapsulate the web serving functionality along with NGINX [6] web server. 2.3.1 Policy Modelling Editor As described in the deliverable D5.4 “Cross-sector Policy Lifecycle Management: Design and Open
Specification 2”[1], the PME is developed using the open-source Angular framework [4]. For UI
components and controls we use the Angular Material library [5]. 2.3 Baseline technologies and tools 2.3.1 Policy Modelling Editor 3.1.1 Policy Modelling Editor 3.1.1 Policy Modelling Editor The source code of the prototype of the Policy Modelling Editor component is available at: 3.1.2 Policy Development Toolkit
The source code of the prototype of the Policy Development Toolkit component is available at: 3.1.3 Data Visualization
The source code of the Data Visualization component is available separately at: •
Project GitLab repository: https://registry.grid.ece.ntua.gr/ana.pontual/visualization
Or through the PDT at: •
Project GitLab repository: https://registry.grid.ece.ntua.gr/kostas/pdt •
Project GitLab repository: https://registry.grid.ece.ntua.gr/kostas/pdt The Backend of the PDT is currently released under open source license, and its source code has been
uploaded and is publicly available at: •
Project GitLab repository: https://registry.grid.ece.ntua.gr/pavlos_LXS/pdt-backend
It also provides the tools to compile the source code and also build a docker image that can deploy and
run this component. As already mentioned, the PDT Backend requires the use of a relational data store to persistently store
its policies, policy related meta-information and results of the analytical tools. We relied on the datastore
provided by LXS, which can be found at: As already mentioned, the PDT Backend requires the use of a relational data store to persistently store
its policies, policy related meta-information and results of the analytical tools. We relied on the datastore
provided by LXS, which can be found at: •
Project GitLab repository: https://registry.grid.ece.ntua.gr/pavlos_LXS/policy-store 2.3.4 PDT Backend The backend of PDT has been implemented using Java SDK 8. It does not make use of any framework like
Spring, but it relies on the standard JDK. For the implementation of the REST services, it makes use of the
javax servlet API 3.1.0 while it also makes use of an embedded tomcat, as the servlet container that the
web services are deployed. We rely on the Apache Embedded Tomcat 8 [9]. By doing this, there is no
need to maintain an additional standalone container like Tomcat, Glassfish or similar, rather than the
java virtual machine starts the embedded one inside its process. Moreover, Maven 3 was used as the tool
for the automation of the build process. For the persistent storage of this component, we relied on the
LXS relational datastore. However, this is not mandatory and therefore, the Backend component of the
PDT can switch to other relational datastore, with minimal changes in the source code. www.policycloud.eu 37 D5.5 – v1.0 www.policycloud.eu •
Project GitLab repository: https://registry.grid.ece.ntua.gr/pavlos_LXS/policy-store As the datastore is published with LXS IPR licence, its source code cannot be available. However, the
GitLab repositories for the datastore contains all binaries and scripting files necessary for packaging the
distribution and creating a docker container that the datastore can start within. As the datastore is published with LXS IPR licence, its source code cannot be available. However, the
GitLab repositories for the datastore contains all binaries and scripting files necessary for packaging the
distribution and creating a docker container that the datastore can start within. 38 D5.5 – v1.0 3.2 Deployment 3.2.1 Policy Modelling Editor
The Policy Modelling Editor will follow the Cloud provider setups and Kubernetes staging. 3.2.1 Policy Modelling Editor
The Policy Modelling Editor will follow the Cloud provider setups and Kubernetes staging. 3.2.2 Policy Development Toolkit
The Policy Development Toolkit will follow the Cloud provider setups and Kubernetes staging. 3.2.4 PDT Backend git clone http://snf-877903.vm.okeanos.grnet.gr/pavlos/pdt-backend.git And later compile the source code. Assuming maven 3 and Java 8 are pre-installed, it needs to execute
the following: And later compile the source code. Assuming maven 3 and Java 8 are pre-installed, it needs to execute
the following: mvn clean install The component can be executed via a java JRE tools, but we also provide a Dockerfile to fully containerize
the deployment and integrate it with the LXS datastore. Firstly, they would need to locally build a docker
image. Assuming that docker has been already installed in the host machine, they should execute the
following: docker build –t pdt-backend . This will create a docker image that can be found in the host machine’s catalogue. To execute the
component, a docker-compose file has also been provided in the GitLab, which makes use of the LXS
datastore. The latter must have been already available in the host machine’s docker catalogue. To start
everything, the administrator should execute the following: docker-compose up docker-compose up The docker-compose file exposes the 54735 port to the host machine, where the servlet container listens
to. Therefore, the administrators can now open their favorite browser and put the following URL, which
will open the swagger documentation: www.policycloud.eu www.policycloud.eu 39 D5.5 – v1.0 An alternative option for starting the PDT backend is to manually start firstly the LXS relational datastore,
and then the PDT backend. With the assumption that the docker images for both the relational datastore
and the PDT backend has been already created and available, then the user needs to first start the
datastore by executing the following: docker run -d -p 2181:2181 -p 1529:1529 --name datastore --env KVPEXTERNALIP='datastore!9800'
policy-store www.policycloud.eu Once the datastore is running, then the user can start the PDT backend with: docker run -d -p 54735:54735 --name pdt_backend --env DATASTORE_HOST='172.17.0.2' --env
swagger_path=/tmp/pdt-backend Once the backend has been started, then the administration can still open a browser and check that
everything is working by invoking the swagger documentation, as shown before at: http://localhost:54735 http://localhost:54735 www.policycloud.eu www.policycloud.eu 40 D5.5 – v1.0 www.policycloud.eu 4 Conclusions This deliverable provided a description of the software components of the Policy Management
Framework Layer and Policy Development Toolkit Layer, which consists of the frontend part of the
PolicyCLOUD platform and allow policy makers to create, update and evaluate policies. For each
component, a description of its APIs, specification of the main functionalities, and description of the
source code is provided. www.policycloud.eu 41 www.policycloud.eu 41 D5.5 – v1.0 www.policycloud.eu References www.policycloud.eu
42
[1]
PolicyCLOUD. D5.4 “Cross-sector Policy Lifecycle Management Design and Open Specification 2”. Armend Duzha et al 2021. [2]
PolicyCLOUD. D2.2 “Conceptual Model and Reference Architecture”, Panayiotis Michael et al 2021. [3]
Swagger [Online], Available at: https://swagger.io/tools/swagger-ui/
[4]
Angular [Online], Available at: https://angular.io
[5]
Angular Material [Online], Available at: https://material.angular.io/
[6]
NGINX [Online], Available at: https://www.nginx.com/
[7]
Leaflet [Online], Available at: https://leafletjs.com/
[8]
Amcharts [Online], Available at: https://www.amcharts.com/
[9]
Apache Tomcat [Online], Available at: http://tomcat.apache.org/ [1]
PolicyCLOUD. D5.4 “Cross-sector Policy Lifecycle Management Design and Open Specification 2”. Armend Duzha et al 2021. [2]
PolicyCLOUD. D2.2 “Conceptual Model and Reference Architecture”, Panayiotis Michael et al 2021. [3]
Swagger [Online], Available at: https://swagger.io/tools/swagger-ui/
[4]
Angular [Online], Available at: https://angular.io
[5]
Angular Material [Online], Available at: https://material.angular.io/
[6]
NGINX [Online], Available at: https://www.nginx.com/
[7]
Leaflet [Online], Available at: https://leafletjs.com/
[8]
Amcharts [Online], Available at: https://www.amcharts.com/
[9]
Apache Tomcat [Online], Available at: http://tomcat.apache.org/ [2]
PolicyCLOUD. D2.2 “Conceptual Model and Reference Architecture”, Panayiotis Michael et al 2021. [3]
Swagger [Online], Available at: https://swagger.io/tools/swagger-ui/
[4]
Angular [Online], Available at: https://angular.io
[5]
Angular Material [Online], Available at: https://material.angular.io/
[6]
NGINX [Online], Available at: https://www.nginx.com/
[7]
Leaflet [Online], Available at: https://leafletjs.com/
[8]
Amcharts [Online], Available at: https://www.amcharts.com/
[9]
Apache Tomcat [Online], Available at: http://tomcat.apache.org/ www.policycloud.eu www.policycloud.eu 42
|
https://openalex.org/W4386125940
|
https://www.frontiersin.org/articles/10.3389/fpls.2023.1235686/pdf
|
English
| null |
Copper boosts the biostimulant activity of a vegetal-derived protein hydrolysate in basil: morpho-physiological and metabolomics insights
|
Frontiers in plant science
| 2,023
|
cc-by
| 11,898
|
TYPE Original Research
PUBLISHED 24 August 2023
DOI 10.3389/fpls.2023.1235686 COPYRIGHT
© 2023 Rouphael, Carillo, Ciriello,
Formisano, El-Nakhel, Ganugi, Fiorini,
Miras Moreno, Zhang, Cardarelli, Lucini and
Colla. This is an open-access article
distributed under the terms of the Creative
Commons Attribution License (CC BY). The
use, distribution or reproduction in other
forums is permitted, provided the original
author(s) and the copyright owner(s) are
credited and that the original publication in
this journal is cited, in accordance with
accepted academic practice. No use,
distribution or reproduction is permitted
which does not comply with these terms. OPEN ACCESS OPEN ACCESS
EDITED BY
Pushp Sheel Shukla,
Dalhousie University, Canada
REVIEWED BY
Sławomir Kocira,
University of Life Sciences of Lublin, Poland
Arup Ghosh,
Council of Scientific and Industrial
Research (CSIR), India
Sumera Javad,
Lahore College for Women University,
Pakistan
*CORRESPONDENCE
Giuseppe Colla
giucolla@unitus.it
Petronia Carillo
petronia.carillo@unicampania.it
RECEIVED 06 June 2023
ACCEPTED 31 July 2023
PUBLISHED 24 August 2023
CITATION
Rouphael Y, Carillo P, Ciriello M,
Formisano L, El-Nakhel C, Ganugi P,
Fiorini A, Miras Moreno B, Zhang L,
Cardarelli M, Lucini L and Colla G (2023)
Copper boosts the biostimulant activity of
a vegetal-derived protein hydrolysate in
basil: morpho-physiological and
metabolomics insights. Front. Plant Sci. 14:1235686. doi: 10.3389/fpls.2023.1235686 EDITED BY
Pushp Sheel Shukla,
Dalhousie University, Canada
REVIEWED BY
Sławomir Kocira,
University of Life Sciences of Lublin, Poland
Arup Ghosh,
Council of Scientific and Industrial
Research (CSIR), India
Sumera Javad,
Lahore College for Women University,
Pakistan Youssef Rouphael 1, Petronia Carillo 2*, Michele Ciriello 1,
Luigi Formisano 1, Christophe El-Nakhel 1, Paola Ganugi 3,4,
Andrea Fiorini 4, Begoña Miras Moreno 3, Leilei Zhang 3,
Mariateresa Cardarelli 5, Luigi Lucini 4,6 and Giuseppe Colla 5* 1Department of Agricultural Sciences, University of Naples Federico II, Portici, Italy, 2Department of
Environmental, Biological and Pharmaceutical Sciences and Technologies, University of Campania
“Luigi Vanvitelli”, Caserta, Italy, 3Department for Sustainable Food Process, Università Cattolica del
Sacro Cuore, Piacenza, Italy, 4Department of Sustainable Crop Production, Università Cattolica del
Sacro Cuore, Piacenza, Italy, 5Department of Agriculture and Forest Sciences, University of Tuscia,
Viterbo, Italy, 6CRAST Research Centre, Università Cattolica del Sacro Cuore, Piacenza, Italy In addition to be used as a plant protection agent, copper (Cu) is also an essential
micronutrient for plant growth and development. The bioavailability of Cu in
agricultural systems can be limited due to its specific physical–chemical
characteristics, leading to imbalances in plant production. To address this
issue, an experimental trial was conducted on Genovese basil (Ocimum
basilicum L.) in protected conditions to comparatively evaluate the effects of a
vegetable protein hydrolysate (VPH), free Cu and Cu complexed with peptides
and amino acids of vegetal origin (Cu and Cu-VPH, respectively), and a
combination of VPH and Cu-VPH (VPH+Cu-VPH). The study showed that the
combined application of VPH+Cu-VPH led to a significant average increase of
16.3% in fresh yield compared to the untreated Control and Cu treatment. This
finding was supported by an improved photosynthetic performance in ACO2
(+29%) and Fv/Fm (+7%). frontiersin.org Frontiers in Plant Science 1 Introduction Hatamian, 2019). However, in recent years, the unsustainability of
the massive use of synthetic chelating agents in agriculture has
emerged because the main artificial chelating agents used are
persistent in natural systems because of their low biodegradability
(Wu et al., 2004). In addition to this disadvantage, these compounds
can alter the natural evolution of metals in groundwater and soil,
making toxic metals bioavailable and compromising the quality of
agricultural products and food safety (Zuluaga et al., 2023). A balance of nutrients, especially between macro- and
micronutrients, is essential to ensure plant health and the
sustainability of agricultural productions (Souri et al., 2017). Despite being required in smaller amounts than primary
nutrients, micronutrients play a crucial role in plant metabolism
and physiology. Accordingly, plants must efficiently control their
mobilization, absorption, distribution, and storage (Hänsch and
Mendel, 2009). Among micronutrients, Cu, a redox-active
transition metal, plays a fundamental role in various physiological
and biochemical processes (Kaewchangwat et al., 2017). At optimal
levels (between 5 and 30 mg kg–1), this microelement participates in
primary metabolic processes (mitochondrial respiration, electron
transport chain, and photosynthesis), lignin biosynthesis, hormone
signaling, and response to oxidative stress (Mir et al., 2021). However, since the 1880s, the role of Cu has been relegated to an
essential active ingredient in fungicidal and bacteriostatic
formulations (Lopez-Lima et al., 2021). Its broad-spectrum
fungicide action has made Cu essential, especially for organic
agriculture where the management of fungal disease depends
almost exclusively on its use (La Torre et al., 2018). The necessary shift towards sustainable agricultural systems has
highlighted the potential of innovative technical solutions, such as
plant-based biostimulants. These latter can increase nutrient use
efficiency and improve crop productivity and quality (Rouphael and
Colla, 2020). The ability to produce biostimulants from plant waste
has made vegetal protein hydrolysate (VPH) one of the most
promising products (Xu and Geelen, 2018). Their biostimulant
activity is attributable to a mixture of bioactive compounds such as
amino acids, peptides, carbohydrates, minerals, phytohormones,
phenols, and other organic compounds (Ciriello et al., 2022a). As
Colla et al. (2015) stated, protein hydrolysates (PHs) can improve
plant nutrition through three modalities: (i) promoting the growth
of the root system, (ii) stimulating absorption processes, and (iii)
increasing the availability of nutrients in the soil by increasing soil
cation exchange and thus inorganic minerals’ solubility in water
(Zuluaga et al., 2023). KEYWORDS vegetal protein hydrolysates, sustainable agriculture, primary metabolism, oxidative
stress, metabolomics, photosynthetic performance, ionomics Frontiers in Plant Science OPEN ACCESS Furthermore, mineral analysis using ICP OES
demonstrated that Cu and Cu-VPH treatments determined, on average, a
15.1-, 16.9-, and 1.9-fold increase in Cu in plant tissues compared to control,
VPH, and VPH+Cu-VPH treatments, respectively. However, the VPH+Cu-VPH
treatment induced the highest contents of the other analyzed ions, except for P. In particular, Mg, Mn, Ca, and Fe, which take part in the constitution of
chlorophylls, water splitting system, and photosynthetic electron transport
chain, increased by 23%, 21%, 25%, and 32% compared to respective controls. Indeed, this improved the photosynthetic efficiency and the carboxylation
capacity of the plants, and consequently, the physiological and productive
performance of Genovese basil, compared to all other treatments and control. Consistently, the untargeted metabolomics also pointed out a distinctive
modulation of phytochemical signatures as a function of the treatment. An
accumulation of alkaloids, terpenoids, and phenylpropanoids was observed
following Cu treatment, suggesting an oxidative imbalance upon metal COPYRIGHT
© 2023 Rouphael, Carillo, Ciriello,
Formisano, El-Nakhel, Ganugi, Fiorini,
Miras Moreno, Zhang, Cardarelli, Lucini and
Colla. This is an open-access article
distributed under the terms of the Creative
Commons Attribution License (CC BY). The
use, distribution or reproduction in other
forums is permitted, provided the original
author(s) and the copyright owner(s) are
credited and that the original publication in
this journal is cited, in accordance with
accepted academic practice. No use,
distribution or reproduction is permitted
which does not comply with these terms. Frontiers in Plant Science 01 frontiersin.org Rouphael et al. 10.3389/fpls.2023.1235686 exposure. In contrast, a mitigation of oxidative stress was highlighted in Cu-VPH
and VPH+Cu-VPH, where the treatments reduced stress-related metabolites. Overall, these results highlight an interaction between Cu and VPH, hence paving
the way towards the combined use of Cu and biostimulants to optimize
agronomic interventions. Rouphael et al. 2.2 Determination of CIELab colorimetric
and SPAD indexes Color coordinates (L, a*, and b*) were recorded using a Minolta
Chroma Meter CR-300 handheld colorimeter (Minolta Camera Co. Ltd., Tokyo, Japan). Ten measurements were taken on the surface of
fully expanded young leaves of eight plants per experimental unit
on the day before harvest (36 DAT). The same leaves were used to
determine the SPAD index using the SPAD-502 optical device
instrument (Minolta Corp. Ltd., Osaka, Japan). The experimental trial was conducted on the experimental farm
“Torre Lama” (lat. 43°31’N, long. 14°58’E; alt. 60 m above sea level)
of the University of Naples “Federico II” - Department of
Agriculture (DIA). The test was conducted in an unheated
greenhouse with a galvanized iron structure and a glass roof. The
structure has an aisle width of 10 m and a length of 30 m; the ridge
height is 4.5 m, and the height at the eaves is 3.0 m. The plant
material used for the trial was the Ocimum basilicum L. Genovese
cultivar Mammolo (L’Ortolano, Cesena, Italy), grown in 1.2-L plots
filled with agricultural soil at a density of 20 pt m−2, with 25 and
20 cm inter- and intra-rows, respectively. The main physical and
chemical soil characteristics at the experimental site were clay loam
texture (46% sand, 24.2% silt, and 29.8% clay), electrical
conductivity (EC): 0.16 dS m−1, pH: 7.7, total nitrogen (N):
0.11%, and organic matter: 1.21% (w/w). The Olsen phosphorus
and exchangeable potassium were 88 and 980 mg kg−1, respectively. The seeds were transplanted on 12 June 2019, in the two- to the
three-leaf stage and irrigated using a micro-irrigation system with
16-mm inner diameter drip wings and a 10-cm pitch with self-
compensating drippers of 2 L h–1 flow rate. The experimental trial
involved randomized blocks with five treatments replicated three
times. The experimental unit consisted of 15 plants. Seven days after
transplanting (DAT), the crop was subjected to the following
treatments: copper sulfate pentahydrate CuSO4-5H2O (0.23 g L–1;
henceforth Cu), vegetal-protein hydrolysate at the manufacturer’s
recommended dose (4 ml L–1 of Trainer®; henceforth VPH; Hello
Nature, Rivoli Veronese, Italy), Cu complexed with peptides and
amino acids of vegetal origin (Scudo® - Hello Nature, Rivoli
Veronese, Italy—at the dose of 0.5 ml L–1; henceforth Cu-VPH),
Cu complexed with peptides and amino acids of vegetal origin
(Scudo®) enriched with vegetal-protein hydrolysate Trainer®
(0.5 ml L–1 and 3.7 ml L–1, respectively; henceforth VPH+Cu-
VPH), and untreated control. 1 Introduction Combining biostimulants and innovative
fertilizers such as chelates may represent a step forward in solving
nutritional problems in agricultural systems. With their ability to
act as ligand via non-covalent bonds, peptides may function as
biochelators for specific microelements (Zuluaga et al., 2023). The
use of biochelates makes micronutrients more available than
standard fertilization practices (Souri et al., 2017). Moreover, the
enhanced biodegradability of metal biochelates, compared to
synthetic counterparts, provides a more favorable environmental
impact. In the specific case of Cu, PHs might also provide additional
benefits in terms of mitigation of metal-related stress in plants. Despite their potential, no in-depth studies have been conducted on
the single and combined effects of biostimulants and the
corresponding biochelates. The definition of synergism between
Cu and PHs may offer novel solutions in the agronomic
management of both plant nutrition and disease management. For this reason, our research aimed to compare the applications
of copper sulfate pentahydrate, a VPH, Cu complexed with a On the other hand, excessive use of Cu in agriculture and
industry has significantly increased its concentration in cultivable
soils. High levels of Cu inhibit plant development and growth due to
irreversible morphological, physiological, and biochemical
alterations (Mir et al., 2021). In fact, a complex and articulated
system of utility, toxicity, absorption, and transport in plants has led
to a dual outlook for Cu, as a harmful or an essential element
(Shabbir et al., 2020). Predominantly present as a cupric ion (Cu2+),
its bioavailability depends not only on the absolute concentration
but also on the physicochemical characteristics of the soil. Organic
and inorganic colloids immobilize Cu, making it unavailable to
plants, especially in alkaline soils that represent approximately 30%
of cultivable soils worldwide (Kumar et al., 2021; Wairich et al.,
2022). In these cases, the foliar application of Cu is preferable to the
soil application as it improves its own uptake and use efficiency,
thus promoting a more sustainable agriculture. Although
introduced to improve the nutritional status of animal organisms,
synthetic chelating agents (EDTA and EDDHA) have become
essential for the balanced nutrition of crop systems (Souri and 02 frontiersin.org Rouphael et al. 10.3389/fpls.2023.1235686 10.3389/fpls.2023.1235686 At 37 DAT, eight plants per replicate were sampled to
determine biometric parameters. Specifically, leaves and stems
were separated to fresh weight (fw; g plant–1). The fresh yield was
expressed as kg m–2. 2.2 Determination of CIELab colorimetric
and SPAD indexes VPH was produced through
enzymatic hydrolysis of legume biomass as reported by Colantoni
et al. (2017) while the Cu complexed with peptides and amino acids
(Cu-VPH) was obtained by chemical reaction of copper sulfate with
peptides and amino acids contained in the VPH. Copper
concentration in all Cu treatments (Cu, Cu-VPH, and VPH-Cu-
VPH) was 58.5 mg L−1. In both VPH and VPH-Cu-VPH
treatments, nitrogen as free amino acids and soluble peptides was
0.24 g L−1 while Cu-VPH treatment provided 0.02 g L−1 of N as
amino acids and soluble peptides. Treatments were made by foliar
application, always in the early morning, using separators to
prevent drift contamination. By weekly applications, four
treatments were made throughout the crop cycle. 2.3 Determination of leaf gas exchange
and chlorophyll fluorescence At 37 DAT, between 10:30 a.m. and 1:30 p.m., in order to check
the physiological status of plants at harvest, gas exchange and
chlorophyll fluorescence measurements were made. Measurements were performed on fully expanded young basil
leaves, avoiding the midrib. The maximum quantum efficiency of
photosystem II (PSII) Fv/Fm was assessed on the leaves of nine
plants per experimental unit after their adaptation to darkness (for
at least 10 min) using leaf clips specific for the portable fluorometer
Fv/Fm Meter (Opti-Sciences Inc., Hudson, USA). On the same
leaves, the assimilation rate (ACO2) and transpiration rate (E) were
determined using a portable gas exchange analyzer (LCA 4; ADC
BioScientific Ltd., Hoddesdon, United Kingdom). The ratio
between ACO2 and E was used to determine the instantaneous
water use efficiency (WUEi). For leaf gas exchange measurements,
all parameters [CO2 concentration, photosynthetically active
radiation (PAR), and relative humidity (RH)] were set to ambient
values while airflow was set to 400 ml s–1. Frontiers in Plant Science 1 Introduction The sampled plant material was then placed in
a ventilated oven (for 72 h at 65°C) to assess the total dry weight
(dw) of leaves and stems and thus the dry matter (%). vegetable protein hydrolysate (Cu-VPH), and a mixture of VPH
and Cu-VPH on the yield and biochemical traits of Genovese basil
grown in a glass greenhouse. frontiersin.org 2.6 Statistics All data were analyzed with IBM SPSS Statistics software
version 26.0 (SPSS Inc., Chicago, IL, USA) and are presented as
mean ± standard error, n = 3. All mean effects were subjected to
one-way ANOVA analysis. Statistical significance was determined
with Tukey’s HSD test at the p = 0.05 level. Macro- and trace element analysis of the digested samples was
performed by ICP OES. The plasma source was provided by
99.999% purity argon (SOL, Marcianise, Italy). The instrument
was optimized daily for the maximum signal and tuned with a
specific solution for the wavelength range studied. For non-alkali
elements (Fe, Mn, Cu, Zn, and P), the calibration curve was
constructed in the 1.0–100 mg L–1 range. For alkaline elements
(K, Ca, and Mg), the calibration curve was constructed from 2 mg
L–1 to 1,000 mg L–1. Macro (K, Ca, Mg, and P)- and trace elements
(Cu, Fe, Mn, and Zn) were expressed as mg g–1 fw and µg g–1 fw,
respectively, based on the original dw of each basil sample. Mass Profiler Professional 15.1 software (Agilent Technologies)
was used to elaborate metabolomic data. Compound abundance
was Log2 transformed and normalized at the 75th percentile, then
baselined against the median. Thereafter, the unsupervised
hierarchical cluster analysis (Ward linkage method and Euclidean
distance) was carried out to get an overview of phytochemical
patterns and investigate relatedness among treatments. Then, the
raw dataset was imported into SIMCA 13 (Umetrics, Malmo,
Sweden), pareto scaled, and subjected to the supervised
orthogonal projection to latent structures discriminant analysis
(OPLS-DA). The OPLS-DA model was cross- validated (CV-
ANOVA) and R2Y and Q2Y parameters (degree of correlation
and prediction ability, respectively) were recorded. Moreover,
overfitting was excluded through a permutation test (n = 100),
outliers were inspected through Hotelling’s T2, and discriminant
compounds selected by variable importance in projection (VIP) in
the predictive model were noted. Finally, Volcano plot analysis (p-
value < 0.05, Benjamini correction; fold change ≥2) was used to
identify differential compounds, and the PlantCyc pathway tools
software [http://www.plantcyc.org/; Caspi et al. (2014)] was used for
biochemical interpretations. 3 Results Untargeted metabolomic analysis was performed using a 1290
ultrahigh-pressure liquid chromatographic system coupled to a
G6550 electrospray quadrupole-time-of-flight mass spectrometry
(UHPLC-ESI/QTOF-MS) from Agilent Technologies (Santa Clara,
CA, USA). Reverse-phase chromatographic separation used an
Agilent 120 PFP column (100 mm × 2.1 i.d., 1.9 mm particle size)
and a mobile phase linear gradient of water and acetonitrile (6-94%
acetonitrile in 33 min). Both mobile phases were acidified with 0.1%
(v/v) formic acid. The mass spectrometer operated in full scan mode
(range 100–1200 m/z), positive polarity (ESI+), and source
parameters were previously optimized (Mastinu et al., 2021). 2.4 Determination of macro- and
trace elements The determination of macro- and trace elements was carried
out as detailed by Ciriello et al. (2023). All standards and reagents
used to determine macro- and trace elements by inductively
coupled plasma optical emission spectrometric analysis (ICP
OES) were purchased from Sigma-Aldrich (Milan, Italy). Calibration standards were prepared from the multi-element
standard solution ICP Calibration mix EH61 PrimAg (Romil,
Cambridge, UK). For accuracy determination, Standard Reference
Material (SRM) 1570a (NIST) was prepared for trace elements in
spinach (Spinacia oleracea L.) leaves, which was analyzed in
triplicate in the same manner as the samples. All glassware and 03 frontiersin.org Rouphael et al. 10.3389/fpls.2023.1235686 10.3389/fpls.2023.1235686 instruments were decontaminated by soaking them in a sulfuric
acid solution overnight and then rinsed several times, first with
deionized water and then with ultrapure water (Merck Millipore,
Darmstadt, Germany). Compound annotation was achieved with MussHunter
Profinder 10.0 software (Agilent Technologies) by applying the
“find-by-formula” algorithm, based on the combination of
monoisotopic accurate mass (tolerance of 5 ppm) and isotopic
spacing and ratio, following mass and retention time alignment. The PlantCyc 9.6 [http://www.plantcyc.org/; Hawkins et al. (2021)]
database was used for annotation, and post-annotation filtering
retained the compounds present in three replications within at least
one treatment. According to COSMOS Metabolomics Standard
Initiative, Level 2 of annotation, corresponding to putatively
annotated compounds, was adopted (Salek et al., 2015). The basil leaves were washed with ultrapure water and placed in a
forced-ventilated oven at 65°C until they reached a constant weight and
were finely ground. Before the analysis of the ICP OES (Spectroblue,
Spectro Ametek, Berwyn, PA, USA), 0.5 g of the dried plant sample was
weighed and subjected to laboratory analysis. Five grams of the dried
plant sample, weighed in PTFE (polytetrafluoroethylene) vessels, was
treated by microwave-assisted digestion (MLS-1200, Microwave
Laboratory Systems, Milestone, Shelton, CT, USA) in a mixture of
9 ml of nitric acid (70%, for trace analysis), 3 ml of hydrochloric acid
(30%, for trace analysis), and 0.5 ml of H2O2. The heating program
consisted of switching from 20°C to 180°C in 15 min and then staying
at 180°C for an additional 10 min, with a power setting of 800 W. The
fully digested samples, after cooling, were transferred to a volumetric
flask and the volume was increased to 50 ml with ultrapure water
(Volpe et al., 2015). 2.5 UHPLC-ESI/QTOF-MS
untargeted metabolomics Basil leaves were collected, ground with liquid nitrogen using
pestle and mortar, and successively extracted and analyzed
according to Rouphael et al. (2020). Briefly, a 1-g aliquot per
sample was extracted in 15 ml of an aqueous solution with 80%
(v/v) methanol acidified with 0.1% formic acid using an Ultra-
Turrax (Polytron PT, City, Switzerland). Thereafter, the extracts
were centrifugated at 8,000 × g for 10 min and 1 ml of each resulting
supernatant was transferred into a glass vial by filtering through a
0.22-mm cellulose syringe filter for analysis. Frontiers in Plant Science 3.1 Yield and yield parameters With the exception of dry matter, the yield and yield
components presented in Table 1 showed significant differences
among the treatments (Control, Cu, Cu-VPH, VPH+Cu-VPH, and
VPH). Specifically, when compared to the Control group, the VPH
+Cu-VPH treatment exhibited notable increases in leaf fresh
weight, total fresh weight, stem dry weight, total dry weight, and
fresh yield, with improvements of 16.80%, 18.62%, 29.50%, 22.70%,
and 18.48%, respectively. 04 frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. TABLE 1 Leaf, stem, and total fresh weight; leaf, stem, and total dry weight; fresh yield; and dry matter of basil under untreated control, copper
sulfate (Cu), copper complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-
VPH) treatments. Treatments
Leaf fresh
weight
Stem fresh
weight
Total fresh
weight
Leaf dry
weight
Stem dry
weight
Total dry
weight
Fresh
yield
Dry
matter
g plant–1
kg m–2
%
Control
62.56 ± 0.23 c
29.39 ± 0.42 b
91.95 ± 0.53 c
6.97 ± 0.07 b
3.39 ± 0.04 d
10.35 ± 0.08 c
1.84 ± 0.01 c
11.26 ± 0.03
Cu
64.22 ± 0.80 c
31.25 ± 0.93 b
95.47 ± 1.67 c
6.94 ± 0.06 b
3.87 ± 0.07 bc
10.80 ± 0.12 bc
1.91 ± 0.03 c
11.32 ± 0.12
Cu-VPH
66.38 ± 0.52 b
34.47 ± 0.73 a
100.85 ± 1.25 b
7.42 ± 0.14 b
3.71 ± 0.08 c
11.14 ± 0.19 bc
2.02 ± 0.03 b
11.04 ± 0.06
VPH+Cu-
VPH
73.07 ± 0.14 a
36.00 ± 0.43 a
109.07 ± 0.57 a
8.32 ± 0.26 a
4.39 ± 0.07 a
12.70 ± 0.32 a
2.18 ± 0.01 a
11.64 ± 0.24
VPH
67.52 ± 0.27 b
34.69 ± 0.28 a
102.21 ± 0.37 b
7.67 ± 0.19 ab
4.00 ± 0.02 b
11.68 ± 0.19 b
2.04 ± 0.01 b
11.42 ± 0.15
Significance
***
***
***
***
***
***
***
ns
ns and *** denote nonsignificant or significant at p ≤0.001, respectively. Different letters within each column indicate significant differences according to Tukey HSD test (p = 0.05). All data are
expressed as mean ± standard error, n = 3. 3.3 Protein, carbohydrate, and polyphenols
concentration Relative to stem fresh weight, the highest values were obtained
from the Cu-VPH, VPH+Cu-VPH, and VPH treatments, with no
significant differences. Compared with the Cu-VPH treatment, the
combined VPH+Cu-VPH treatment significantly increased leaf dry
weight (+12.13%). Cu-VPH and VPH biostimulant application significantly affected
soluble protein content. The protein content was significantly
decreased by applying Cu-VPH and VPH, which was 25.6% and
31.6% lower than in untreated basil plants, respectively (Table 3). None
of the treatments significantly affected starch (avg. 144.8 µmol g–1 dw)
and sucrose (2.3 µmol g–1 dw) concentrations. (Table 3). However, the
application of VPH+Cu-VPH averaged over other treatments
positively affected the glucose concentrations in basil leaves (an
increase of 76.2% compared to control), even if also Cu and Cu-
VPH determined an increase of this soluble sugar of approximately
40% (Table 3). Fructose increased 24.4% and 30.8% compared to
control when treated with Cu-VPH and VPH+Cu-VPH, respectively
(Table 4). Finally, Cu-VPH and VPH significantly decreased the
polyphenols content, with no significant differences among the three
treatments (Table 3). Frontiers in Plant Science *** significant at p ≤0.001. Different letters within each column indicate significant differences according to Tukey HSD test (p = 0.05). All data are expressed as mean ± standard error, n = 3. Frontiers in Plant Science
frontiersin.org
05 3.2 Physiological parameters All physiological parameters, shown in Table 2, were
significantly affected by the treatments under investigation. The highest values of net CO2 assimilation rate (ACO2; 14.38
mmol CO2 m–2 s–1) and fluorescence (0.81) were obtained in
plants of the VPH+Cu-VPH treatment. In comparison with the
Control, the Cu-VPH and VPH+Cu-VPH treatments increased
transpiration (E) by 49.10% (on average). In contrast, the Cu-
VPH treatment reduced WUEi (–29.91%) compared with
the Control. TABLE 2 Net CO2 assimilation rate (ACO2), transpiration (E), water use efficiency (WUE), and chlorophyll fluorescence (Fv/Fm) of basil under untreated
control, copper sulfate (Cu), copper complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched
with VPH (VPH+Cu-VPH) treatments. Treatments
ACO2
E
WUEi
Fluorescence Fv/Fm
mmol CO2 m−2 s−1
mol H2O m−2 s−1
mmol CO2 mol−1 H2O
Control
11.15 ± 0.11 b
1.66 ± 0.04 c
6.72 ± 0.12 a
0.76 ± 0.00 d
Cu
11.96 ± 0.13 b
2.18 ± 0.18 b
5.57 ± 0.45 bc
0.77 ± 0.00 c
Cu-VPH
12.19 ± 0.13 b
2.59 ± 0.04 a
4.71 ± 0.10 c
0.78 ± 0.00 b
VPH+Cu-VPH
14.38 ± 0.45 a
2.36 ± 0.03 ab
6.09 ± 0.12 ab
0.81 ± 0.00 a
VPH
12.14 ± 0.19 b
2.00 ± 0.02 bc
6.07 ± 0.15 ab
0.77 ± 0.00 c
Significance
***
***
***
*** 05 frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. TABLE 3 Soluble proteins, glucose, fructose, sucrose, starch, and polyphenols concentration of basil under untreated control, copper sulfate (Cu),
copper complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH)
treatments. 3.2 Physiological parameters Treatments
Soluble proteins
Glucose
Fructose
Sucrose
Starch
Polyphenols
mg g–1 fw
µmol g–1 dw
mg GAE g–1 dw
Control
9.11 ± 0.59 a
36.96 ± 1.41 c
22.90 ± 2.87 b
2.65 ± 0.50
146.27 ± 10.43
20.21 ± 0.22 a
Cu
7.85 ± 0.34 ab
51.65 ± 2.33 b
23.17 ± 1.12 b
2.76 ± 0.30
152.57 ± 24.54
18.92 ± 0.91 a
Cu-VPH
6.78 ± 0.47 b
51.23 ± 0.27 b
28.50 ± 0.89 a
2.10 ± 0.31
148.91 ± 25.37
16.40 ± 0.48 b
VPH+Cu-VPH
7.26 ± 0.33 ab
65.12 ± 3.75 a
29.96 ± 0.50 a
2.37 ± 0.28
160.45 ± 3.04
15.79 ± 0.42 b
VPH
6.23 ± 0.14 b
44.88 ± 0.89 bc
27.40 ± 1.58 ab
1.68 ± 0.20
115.66 ± 19.12
15.38 ± 0.36 b
Significance
**
***
*
ns
ns
***
ns, *, **, and *** denote nonsignificant or significant at p ≤0.05, 0.01, and 0.001, respectively. Different letters within each column indicate significant differences according to Tukey HSD test (p =
0.05). All data are expressed as mean ± standard error, n = 3. ns, *, **, and *** denote nonsignificant or significant at p ≤0.05, 0.01, and 0.001, respectively. Different letters within each column indicate significant differences according to Tukey HSD test (p =
0.05). All data are expressed as mean ± standard error, n = 3. 3.6 Principal component analysis To obtain an in-depth overview of growth parameters, mineral
profile, and nutritional and functional quality traits of the studied
basil leaves in response to the diverse treatments, a principal
component analysis (PCA) was carried out for all the analyzed
parameters discussed above. The first three principal components
(PCs) were related with eigenvalues higher than 1 and explained
92.6% of the total variance, with PC1, PC2, and PC3 accounting for
54.2%, 25.1%, and 13.3%, respectively (data not shown). The increase in metabolism reshaping (activation of more
metabolic pathways) contributed to the clear separation of PC1
with VPH+Cu-VPH in the positive side of PC1. In contrast, the
distribution on PC2 was more linked to the efficient use of water
with the control and Cu-VPH in the positive and negative side of
PC2, respectively (Figure 1). PC1 was positively correlated to amino
acids [in particular phenylalanine, asparagine, valine, tryptophan,
branched-chain amino acids (BCAAs), tyrosine, threonine, and
serine], dry matter, leaf fresh weight, stem dry weight, leaf dry
weight, total dry weight, and ACO2 (Figure 1). PC1 negatively
correlated with P, polyphenols, and L* and a* parameters. PC2
was positively correlated with lysine and WUEi, and negatively
correlated with E (Figure 1). In addition, basil plants treated with
VPH+Cu-VPH positioned on the positive side of PC1 close to the x-
axis between the lower and upper right quadrant produced leaves of
premium quality with higher fresh yield and concentrations of free
amino acids and, in particular, essential amino acids. Control and
Cu-treated plants in the upper left quadrant were instead
characterized by the lowest fresh yield but the highest
polyphenols content. Finally, Cu-VPH showed the lowest amino
acid content, lower soluble proteins, and polyphenols than the
control, but a high mineral concentration (K, Mg, Cu, and
Fe) (Figure 1). 3.4 Free amino acid concentration reduced Zn by 21.93% compared to the VPH+Cu-VPH
treatment (Table 5). The free amino acid profile of basil plants as a function of the
different treatments is reported in Table 4. Irrespective of the
treatments, glutamate and alanine were quantitatively the major
amino acids, representing, on average, approximately 31.0% and
9.9% of total free amino acids in basil plants, respectively. Essential
amino acids (the sum of isoleucine, histidine, leucine, lysine,
methionine, threonine, and valine) accounted, on average, for
15.5% of total amino acids. Interestingly, the VPH and VPH+Cu-
VPH determined a significant increase of all amino acids,
particularly glutamate, glutamine, aspartate, and asparagine,
which are involved in nitrogen assimilation and transport
(Table 4). In contrast, the application of Cu-VPH alone
determined the opposite effect, with a strong decrease of total
amino acid content and of GABA (–61.6%), MEA (–67.5%),
phenylalanine (–35.3%), glutamate (–42.9%), and alanine (–
22.8%); even the latter two were not significant (Table 4). In this
condition, the only amino acids that did not decrease compared to
the respective controls were the amides asparagine and glutamine,
proline, and essential amino acids (Table 4). Frontiers in Plant Science 3.5 Leaf mineral composition Among the analyzed minerals, Mn and K were by far the most
abundant micro- and macroelements. Regardless of the treatments,
K values ranged from 52.1 to 59.7 mg g–1 dw, while Mn values
ranged from 56.5 to 74.2 µg g–1 dw (Table 5). None of the
treatments affected the concentration of P (averaging at 2.4 mg g–
1 dw). However, the Cu, Cu-VPH, and VPH+Cu-VPH treatments
significantly increased the Cu concentration by 15.3-, 15.0-, and 7.8-
fold, respectively, compared to the control (Table 5). The Cu-VPH
treatment also significantly increased K (+14.3%), Mg (+18.0%),
and Fe (+29.2%), while the VPH+Cu-VPH treatment significantly
increased Ca (+24.7%) and Mn (+20.8%) compared to their
respective control treatments. The VPH treatment significantly Frontiers in Plant Science 06 frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. TABLE 4 Aminogram, essential amino acids, branched-chain amino acid (BCAAs), and total amino acids of basil under untreated control, copper
sulfate (Cu), copper complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-
VPH) treatments. 3.5 Leaf mineral composition Treatments
Control
Cu
Cu-VPH
VPH+ Cu-VPH
VPH
Significance
Ala
3.64 ± 0.34 b
5.41 ± 0.10 b
2.81 ± 0.39 b
9.68 ± 1.63 a
5.93 ± 0.73 ab
***
Arg
0.908 ± 0.13 b
1.771 ± 0.08 a
1.436 ± 0.11 ab
0.959 ± 0.13 b
1.757 ± 0.22 a
**
Asn
1.142 ± 0.226 b
0.970 ± 0.087 b
1.061 ± 0.087 b
2.634 ± 0.486 a
1.627 ± 0.204 ab
**
Asp
0.513 ± 0.023 b
0.374 ± 0.041 b
0.410 ± 0.047 b
0.957 ± 0.137 a
1.018 ± 0.106 a
***
GABA
1.825 ± 0.367 ab
2.666 ± 0.339 a
0.700 ± 0.126 b
2.816 ± 0.439 a
2.206 ± 0.374 ab
**
Gln
2.028 ± 0.273 ab
1.356 ± 0.099 b
1.209 ± 0.234 b
3.619 ± 0.594 a
3.805 ± 0.532 a
**
Glu
18.49 ± 1.17 ab
16.43 ± 1.85 ab
10.55 ± 0.68 b
20.15 ± 2.78 a
20.49 ± 2.06 a
*
Gly
0.511 ± 0.068 abc
0.443 ± 0.024 bc
0.275 ± 0.044 c
0.714 ± 0.055 a
0.685 ± 0.067 ab
***
His
0.040 ± 0.006 b
0.165 ± 0.018 b
0.136 ± 0.014 b
0.369 ± 0.052 a
0.357 ± 0.040 a
***
Ile
0.781 ± 0.099 ab
0.896 ± 0.048 ab
0.543 ± 0.036 c
1.155 ± 0.090 a
1.006 ± 0.125 a
**
Leu
1.454 ± 0.130 ab
1.465 ± 0.082 ab
1.088 ± 0.096 b
1.850 ± 0.094 a
1.641 ± 0.186 ab
*
Lys
2.454 ± 0.068
2.411 ± 0.097
2.176 ± 0.221
2.318 ± 0.146
2.464 ± 0.057
ns
MEA
0.919 ± 0.156 a
0.773 ± 0.052 ab
0.299 ± 0.034 b
1.207 ± 0.183 a
1.149 ± 0.163 a
**
Met
0.028 ± 0.003 ab
0.033 ± 0.006 a
0.015 ± 0.002 b
0.031 ± 0.003 ab
0.031 ± 0.004 ab
*
Orn
1.108 ± 0.098
1.217 ± 0.125
0.904 ± 0.070
1.313 ± 0.099
1.214 ± 0.127
ns
Phe
0.201 ± 0.028 bc
0.238 ± 0.004 bc
0.130 ± 0.013 c
0.442 ± 0.044 a
0.322 ± 0.066 ab
**
Pro
2.584 ± 0.158
2.051 ± 0.055
2.261 ± 0.215
2.332 ± 0.107
2.067 ± 0.304
ns
Ser
0.611 ± 0.048 bc
0.517 ± 0.049 bc
0.394 ± 0.048 c
0.897 ± 0.037 a
0.634 ± 0.055 b
***
Thr
0.382 ± 0.054 ab
0.250 ± 0.021 b
0.166 ± 0.028 b
0.632 ± 0.095 a
0.465 ± 0.092 ab
**
Trp
0.403 ± 0.083 ab
0.495 ± 0.002 ab
0.251 ± 0.026 b
0.752 ± 0.116 a
0.530 ± 0.101 ab
*
Tyr
0.354 ± 0.063 ab
0.338 ± 0.009 ab
0.218 ± 0.018 b
0.518 ± 0.057 a
0.407 ± 0.076 ab
*
Val
0.850 ± 0.174 ab
1.044 ± 0.057 ab
0.743 ± 0.038 b
1.471 ± 0.128 a
1.231 ± 0.204 ab
*
Essential AA
7.50 ± 0.74 bc
8.77 ± 0.28 abc
6.68 ± 0.54 c
9.98 ± 0.65 ab
10.14 ± 0.44 a
**
BCAAs
3.08 ± 0.40 ab
3.40 ± 0.19 ab
2.37 ± 0.17 b
4.48 ± 0.29 a
3.88 ± 0.51 ab
*
Total AA
51.81 ± 4.34 bc
53.49 ± 2.12 ab
36.83 ± 2.72 c
67.94 ± 4.29 a
68.05 ± 3.02 a
***
ns, *, **, and *** denote nonsignificant or significant at p ≤0.05, 0.01, and 0.001, respectively. Frontiers in Plant Science 3.5 Leaf mineral composition Different letters within each row indicate significant differences according to Tukey HSD test (p =
0 05) All d t
d
± t
d
d
3 All d t
d
l
–1 d ns, *, **, and *** denote nonsignificant or significant at p ≤0.05, 0.01, and 0.001, respectively. Different letters within each row indicate significant differences according to Tukey HSD test (p =
0.05). All data are expressed as mean ± standard error, n = 3. All data were expressed as µmol g–1 dw. 3.7 UHPLC-ESI/QTOF-MS metabolomic
profile of basil 3.7 UHPLC-ESI/QTOF-MS metabolomic
profile of basil frontiersin.org 3.7 UHPLC-ESI/QTOF-MS metabolomic
profile of basil Cu-VPH- and VPH-treated samples—was created, indicating a
more comparable metabolic profile in the basil plant after the
two treatments. UHPLC-ESI/QTOF-MS untargeted metabolomics was carried
out to understand the effect of the different treatments on the basil
metabolome. Overall, more than 3,200 leaf compounds were
putatively annotated and provided as Supplementary Material,
together with individual abundances and composite mass spectra
(Supplementary File 1). Successively, the supervised OPLS-DA multivariate analysis
allowed us to better outline the differences between treatments
and to highlight discriminant (VIP) compounds underlying these
differences. The OPLS-DA score plot, provided in Figure 3, revealed
distinct phytochemical profiles in basil leaves after treatments. In
particular, the first latent vector discriminated VPH+Cu-VPH from
the control, with the treated replicates being considerably separated
from the untreated ones. This result indicated the more pronounced
effect of VPH+Cu-VPH on metabolic reprogramming of basil
leaves, compared to the other treatments. Differently, the second
latent vector pointed out a differential profile of VPH from Cu-VPH
and control. The OPLS model was validated through CV-ANOVA Based on these metabolomic signatures, unsupervised
hierarchical clustering was first performed to describe similarities/
dissimilarities among the treatments. As shown in Figure 2, two
main clusters were generated by the fold-change-based heatmap,
one represented by Cu and VPH+Cu-VPH, and another by control,
Cu-VPH, and VPH. Within this last one, a subcluster—containing 07 frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. 3.7 UHPLC-ESI/QTOF-MS metabolomic
profile of basil Treatments
P
K
Ca
Mg
Cu
Fe
Mn
Zn
mg g–1 dw
mg g–1 dw
Control
2.61 ± 0.11
52.10 ± 1.05 b
16.54 ± 0.62 b
3.20 ± 0.06 c
7.78 ± 0.65 c
34.06 ± 1.18 b
61.37 ± 0.54 bc
19.19 ± 0.59 ab
Cu
2.41 ± 0.12
53.02 ± 1.29 b
16.76 ± 0.48 b
3.30 ± 0.12 c
118.66 ± 6.81 a
33.41 ± 1.58 b
66.10 ± 2.23 b
18.40 ± 0.55 b
Cu-VPH
2.42 ± 0.14
59.59 ± 1.56 a
18.65 ± 0.58 ab
3.78 ± 0.05 ab
116.87 ± 8.78 a
44.01 ± 1.98 a
65.80 ± 1.99 b
18.76 ± 0.74 ab
VPH+Cu-VPH
2.36 ± 0.01
57.73 ± 1.39 ab
20.63 ± 0.98 a
3.94 ± 0.13 a
60.66 ± 2.53 b
44.83 ± 2.76 a
74.17 ± 0.87 a
21.56 ± 0.77 a
VPH
2.32 ± 0.07
59.72 ± 0.93 a
17.17 ± 0.16 b
3.50 ± 0.08 bc
6.98 ± 0.64 c
40.32 ± 1.20 ab
56.52 ± 0.66 c
16.83 ± 0.50 b
Significance
ns
**
**
***
***
**
***
**
ns, **, and *** denote nonsignificant or significant at p ≤0.01 and 0.001, respectively. Different letters within each column indicate significant differences according to Tukey HSD test (p = 0.05). All data are expressed as mean ± standard error, n = 3. (p-value = 1.87E-15), and adequate accuracy parameters (R2Y =
0.99 and Q2Y = 0.88) were obtained. secondary metabolites, terpene and nitrogen-containing
compounds were markedly modulated, followed by
phenylpropanoid derivates. Notably, Cu elicited the highest
accumulation of metabolites of each of these classes, reporting
above all higher concentration of alkaloids [(S)-laudanosine, (S)-
N-methylcoclaurine, 16-hydroxytabersonine, and galanthamine]
and terpenoids, including carotenoids (lactucaxanthin, b-
citraurin, and 9′-cis-neoxanthin), diterpenoids, and
sesquiterpenoids (desoxyhemigossypol and 2-dehydrolubimin). Interestingly, increased levels of terpenoid phytoalexins (e.g.,
momilactone A and 2-dehydrolubimin) were noticed in Cu-
treated samples. Nevertheless, an up-accumulation for both these
two classes was observed with the other treatments, except for VPH,
which exhibited a slight decrease of alkaloid and glucosinolate
compounds (2-benzoyloxy-3-butenylglucosinolate and 2-
sinapoyloxy-3-butenylglucosinolate). The list of VIP compounds, which mostly discriminated the
treatments (VIP > 1.5), is provided in Supplementary File 1. Overall,
isoprenoids were the most represented class of metabolites, mainly
including carotenoids, sesquiterpenes, and gibberellins. Frontiers in Plant Science 3.7 UHPLC-ESI/QTOF-MS metabolomic
profile of basil Finally,
Volcano plot analysis (p-value < 0.05; fold change > 2) was
carried out to select those compounds involved in basil plants’
response to the treatments. A total of 251 differential metabolites
were identified and provided in Table S3 together with their
function class. Therein, compounds related to secondary
metabolite biosynthesis were the most represented in all
treatments, followed by fatty acid and lipid biosynthesis-related
metabolites. The graphical interpretation obtained with the
PlantCyc pathway tool analysis allowed us to get insight into the
metabolic pathways mostly affected in basil leaves (Figure 4). Notably, secondary metabolism was strongly modulated,
showing an overall upregulation after each treatment. Among Differently, improved contents of phenylpropanoids were
highlighted only after VPH and Cu treatments. However, the FIGURE 1
Principal component loading plot and scores of principal component analysis (PCA) of growth parameters, mineral elements, SPAD index, starch,
soluble carbohydrates, polyphenols, and total amino acids in basil leaves untreated (Control) or foliarly treated with copper sulfate (Cu), copper
complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH). FIGURE 1
Principal component loading plot and scores of principal component analysis (PCA) of growth parameters, mineral elements, SPAD index, starch,
soluble carbohydrates, polyphenols, and total amino acids in basil leaves untreated (Control) or foliarly treated with copper sulfate (Cu), copper
complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH). 08 Frontiers in Plant Science frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. FIGURE 2
Fold-change heatmap obtained from the unsupervised hierarchical cluster analysis (Euclidean distance; linkage rule: Ward) on UHPLC-ESI/QTOF-MS
data of basil leaf chemical profiles under untreated control, copper sulfate (Cu), copper complexed with amino acids and peptides (Cu-VPH), vegetal
protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH) treatments. FIGURE 2
Fold-change heatmap obtained from the unsupervised hierarchical cluster analysis (Euclidean distance; linkage rule: Ward) on UHPLC-ESI/QTOF-MS
data of basil leaf chemical profiles under untreated control, copper sulfate (Cu), copper complexed with amino acids and peptides (Cu-VPH), vegetal
protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH) treatments. total fresh and dry weight that increased the basil fresh yield. Probably, it was due to its higher carboxylation activity (ACO2)
and maximum potential quantum efficiency of Photosystem II (Fv/
Fm) compared to all other treatments. 3.7 UHPLC-ESI/QTOF-MS metabolomic
profile of basil It may be supposed that the
VPH+Cu-VPH treatment improved the Cu use efficiency, as well as
the Ca and Mn ones. In particular, it enhanced Cu and Fe
integration as cofactors in thylakoid electron transporters like
plastocyanin and Fe-proteins (e.g., cytochromes, Fe-S clusters,
and ferredoxin), and that of Ca and Mn in the water splitting
system of the photosynthetic electron transport chain, which
became capable of accommodating faster and higher rates of
electron flow under high light (Printz et al., 2016). Also,
photochemical quenching saturated at higher irradiance in these
Cu-sufficient plants, thus increasing the effective quantum yield of
photosynthesis compared to other treatments, particularly to strongest modulation was shown with Cu, highlighting particularly
enhanced concentrations of (2E,4Z,6E)-5-formyl-2-hydroxy-8-
oxoocta-2,4,6-trienoate, (+)-sesaminol, apigeninidin, and (+)-6a-
hydroxymaackiain. In contrast, Cu-VPH and VPH showed a
differential pattern, showing a general downregulation of
phenylpropanoid compounds. Following both treatments, decreased
levels of 8-C-glucosyl chrysin, rutin, (+)-dihydrokaempferol, and
diphyllin were observed. 4 Discussion Accordingly, in
control, the highest levels of polyphenols were found, since they
are a powerful antioxidant particularly efficient in scavenging ROS
and counteracting oxidative stress damage (Caldwell et al., 1983). Nevertheless, considering that treatments with higher levels of
phenols are also less productive, it is probable that both the
control plants and the Cu-treated plants have bolstered secondary
metabolic pathways at the expense of primary ones (Prinsloo and
Nogemane, 2018; Ciriello et al., 2022b). Phenolic compounds have
an antioxidant activity higher than vitamins since they can prevent
ROS formation and improve antioxidant cell response due to their
ability to chelate metal ions (Faraloni et al., 2021). The problem of
excess light correlated to high polyphenols content was not present
in the other plant treatments, which showed higher transpiration
and carboxylation activity, but significantly lower polyphenols,
except for Cu treatment, even if at a lower extent, but without
negative consequences on plant fresh weight. Interestingly, VPH
+Cu-VPH-, VPH-, and Cu-VPH-treated plants also showed a high
SPAD index accompanied by the deepest coloration resulting from
more intense yellowness (b* values; Supplementary Table S1) and
the highest content of glucose and free amino acids. In particular, in
VPH+Cu-VPH plants, the increase in the amounts of leaf
glutamate, aspartate, and amides (glutamine and asparagine) evidenced not only a higher N use efficiency but also (i) a higher
energy availability, probably deriving by the more efficient
photosynthesis, and by the capacity to store nitrogen in glutamine
and asparagine instead of soluble proteins, with a much lower
energy expense (Diab and Limami, 2016), and (ii) the possibility to
use these two amides as long-distance nitrogen transport molecules,
given their high N/C ratio, to supply organic nitrogen to roots to be
used as amine donor for the synthesis of all protein amino acids
boosting root growth (Van Oosten et al., 2019). In contrast, the effect of Cu-VPH alone deserves particular
attention. It determined the highest transpiration and the lowest
WUEi and free amino acid levels, maintaining a high SPAD index,
ACO2, Fv/Fm, mineral nutrients, and fresh yield. Noteworthy,
among amino acids, GABA and MEA strongly decreased. It
seems that GABA, which has a well-known antioxidant activity
against ROS and is useful to stabilize and protect membranes and
macromolecules, was not needed under Cu-VPH treatment
(Carillo, 2018). 4 Discussion Remarkable changes in morpho-physiological and biochemical
traits were observed under VPH+Cu-VPH treatments compared to
other ones. VPH+Cu-VPH treatment showed the highest leaf and FIGURE 3
Orthogonal Projections to Latent Structures Discriminant Analysis (OPLS-DA) score plot for basil leaf under untreated control, copper sulfate (Cu),
copper complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH)
treatments. FIGURE 3
Orthogonal Projections to Latent Structures Discriminant Analysis (OPLS-DA) score plot for basil leaf under untreated control, copper sulfate (Cu),
copper complexed with amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH)
treatments. 09 frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. B
A
FIGURE 4
Leaf basil metabolic processes (A) and the relative details of secondary metabolism (B) as affected by copper sulfate (Cu), copper complexed with
amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH) treatments compared to
untreated control. The compounds resulting from Volcano plot analysis (p value < 0.05; fold change > 0.01) were loaded into the PlantCyc Pathway
Tool (https://www.plantcyc.org/). The x-axis represents each set of metabolic subcategories, while the y-axis corresponds to the cumulative log fold
change (FC). The large dots represent the average (mean) of all FCs for the different metabolites in the class, while the small dots represent the
individual log FC. A A B FIGURE 4
Leaf basil metabolic processes (A) and the relative details of secondary metabolism (B) as affected by copper sulfate (Cu), copper complexed with
amino acids and peptides (Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH) treatments compared to
untreated control. The compounds resulting from Volcano plot analysis (p value < 0.05; fold change > 0.01) were loaded into the PlantCyc Pathway
Tool (https://www.plantcyc.org/). The x-axis represents each set of metabolic subcategories, while the y-axis corresponds to the cumulative log fold
change (FC). The large dots represent the average (mean) of all FCs for the different metabolites in the class, while the small dots represent the
individual log FC. control (Kim and Price, 2017). Instead, in the other treatments,
particularly in control, the absorbed light energy might exceed the
photosynthetic capacity, and over-reduce the photosynthetic
electron transport chain, thus leading to the generation of reactive
oxygen species (ROS) (Sperdouli et al., 2021). Frontiers in Plant Science frontiersin.org 4 Discussion VPH+Cu-VPH treatment increased, even if not significantly
compared to the VPH one, the tryptophan content of plants, and
determined an inhibition of stem elongation and an increase of
compactness of plants, with a more equilibrated leaf enlargement
(especially that of older leaves) and consequent increase of yield
compared to all other differently treated plants (Figure 5). This is puzzling since Colla et al. (2014) demonstrated that VPH
(Trainer®) was able to induce coleoptile elongation in detached
corn (Zea mays L.) seedlings due to the high tryptophan content
equal to 3 g kg–1 that exerted an auxin-like activity (Rouphael et al.,
2018). However, tryptophan, which is the main precursor of indole-
3-acetic acid (IAA), not only can modulate shoot length, but also, in
synergy with the peptides contained in VPH, can certainly
contribute via auxin- and/or gibberellin-like activity, as shown in
in vitro bioassays by Ertani et al. (2009) and Colla et al. (2014), to
modulate/improve root architecture (Fukaki and Tasaka, 2009). This reshaping of root architecture has been correlated with an
enhancement of N uptake and translocation known as “nutrient
acquisition response” and can also be associated with the higher
root assimilation of Ca, Mg, Mn, and Fe, and certainly also Cu, and
their translocation to shoots of basil plants, as previously seen in
Rouphael et al. (2017) and Carillo et al. (2019). This higher
availability of N and other minerals can be responsible for the
better leaf nutritional status and yield of VPH+Cu-VPH treatment. In particular, the higher availability of amino acids and Mg can
contribute to the higher photochemical efficiency and activity of
photosystem II (PSII), possibly due to the increase of
photosynthetic proteins, particularly Ribulose-1,5-bisphosphate
carboxylase/oxygenase (RuBisCO), and their Mg-dependent
activation, thus increasing CO2 assimilation (ACO2) and
translocation of recent photosynthates via the phloem to sink
tissues (Carillo et al., 2019). Therefore, the interaction between Consistently, untargeted metabolomics unraveled a distinctive
reprogramming of basil leaf metabolome in response to the different
treatments, generally displaying a remarkable modulation of
secondary metabolite concentrations. The exposure of basil plant to Cu treatment induced significant
changes in its alkaloid, terpenoid, and phenylpropanoid profiles,
showing the most marked accumulation compared to the other
treatments. This result can be interpreted as plant protective effect
against physiological processes’ damage and oxidative stress, due to
the excess presence of Cu. 4 Discussion This is also confirmed by the low levels of
monoethanolamine (MEA), a serine derivative important for
synthesizing or regenerating phospholipids (Carillo et al., 2020). This suggests that under Cu-VPH treatment, the higher Cu
concentration, probably not yet toxic as in Cu treatment, can
modulate plant metabolism in a way that photo-assimilates are
promptly diverted to roots. In roots, N is mainly used to boost root
growth and soil exploration, instead of remaining in leaves and
being invested in proteins, particularly RUBISCO proteins that
make up a large fraction of N in leaves. The higher amounts of water 10 frontiersin.org 10.3389/fpls.2023.1235686 Rouphael et al. VPH and Cu-VPH exerted a positive effect on the electron transport
chain, CO2 carboxylation, and translocation of newly synthetized
soluble sugars to young leaves, decreasing the photooxidative
damage, the consequent ROS production, and the need to
synthesize polyphenols, while increasing N use efficiency, amino
acid synthesis, and growth. This phenomenon was phenotypically
exteriorized by the change in plant shape and higher crop yield. Moreover, when Cu is supplied by foliar spraying, high contents of
free cytosolic Cu+ in leaves may cause a massive generation of ROS
and oxidative stress or activate the programmed cell death (PCD). To prevent this copper-dependent oxidative burst, Cu+ must be
chelated by intracellular cysteine-rich metallothioneins (MTs) or
specific chaperones whose synthesis has a high energy cost, thus
jeopardizing plant growth (Printz et al., 2016). In contrast, VPH
peptides, particularly amino acids and small peptides that can be
absorbed by plants (Colla et al., 2015), can complex not only Cu+,
but also Cu2+, maintaining its redox capability, and facilitate Cu
transport and allocation to Cu proteins, as also recently seen in
medical studies on Cu-based chemotherapeutics (Peña et al., 2022). and mineral nutrients acquired exert a positive effect on plant
metabolism, positively affecting processes such as chloroplast and
mitochondrial electron transports, ROS detoxification, and redox
status. However, since the high availability of water, Cu-VPH plants
less strictly control transpiration, but without being able to further
increase carboxylation activity; consequently, the lower ACO2-to-
transpiration (E) ratio determines lower WUEi values compared to
that of VPH+Cu-VPH (Liu et al., 2018). 5 Conclusions The widespread awareness of the impact of anthropogenic
activities on the environment calls for careful and critical
reflection on the agricultural sector’s future. The unconditional
use of chemical fertilizers and pesticides is no longer sustainable,
requiring growers to use innovative techniques that are increasingly
eco-friendly. Cu, despite being a broad-spectrum fungicide and
nutritionally essential element for plants, is a heavy metal that may
accumulate and enter trophic chains. For this reason, its use with
new formulations may be a necessary sustainable means of reducing
dosages and increasing crop uptake and use efficiency. Regarding
the mineral profile, the plants treated with Cu-VPH had the same
copper concentration as those treated with inorganic copper. Still, at
the same time, the use of Cu-VPH significantly improved the
concentrations of K, Mg, and Fe, whereas VPH+Cu-VPH
treatments, which showed half of leaf Cu concentration compared
to Cu and Cu-VPH treatments, underwent a better Ca and Mn
uptake capacity, and Cu and N use efficiency. It is likely that Cu
integration as a cofactor in transporters of the photosynthetic
electron transport chain, and that of Mn in the water splitting
system, improved the maximum potential quantum efficiency of
Photosystem II (Fv/Fm) and the electron transport flow under high
light. At the same time, the carboxylation capacity of the plants and
the synthesis and translocation of amides increased, thus enhancing
the physiological and productive performance of Genovese basil and
its fresh production compared to the other treatments and the
untreated control. Finally, untargeted metabolomics pointed out
increased concentrations of stress-related compounds (mainly
alkaloids, terpenoids, and phenylpropanoids) following Cu supply,
differently from VPH-Cu, VPH, and VPH+Cu-VPH treatments,
which showed low levels of these compounds. This suggested the
VPH ability to alleviate the negative effect of oxidative stress due to
plant exposure to substantial amount of Cu. Therefore, in light of
the promising results obtained in this study and with a view to
enhancing agricultural sustainability, producers should promote the
use of complex micronutrients combined with biostimulants. Likewise, plant production of terpenoids with antioxidant
activities to cope and overcome abiotic stress is increasingly
described in the scientific literature (Toffolatti et al., 2021). When
exposed to the above threshold concentration of trace metals, plants
can modify metabolic processes by synthesizing them (Anjitha
et al., 2021). Kunwar et al. 5 Conclusions (2015) accounted significant changes
in basil terpenoid composition by the addition of Cu, showing
improved concentrations of linalool, chavicol, and methyl
cinnamate when compared to the control (Kunwar et al., 2015). Interestingly, Hojati et al. (2017) investigated the response of
Tanacetum parthenium L. to copper chloride dihydrate
(CuCl2·2H2O) by associating stress-related responses with
changes in terpenoids: low Cu (5 mM) supply increased the
proportion of sesquiterpenes, both hydrocarbons and oxygenated,
while elevated Cu levels reduced the total content of isoprenoids, by
depressing the uptake of divalent cations, which are essential
cofactors of enzymes involved in the biosynthesis of
these metabolites. In contrast, VPH-, Cu-VPH-, and VPH+Cu-VPH-treated
plants displayed lower contents of alkaloids and terpenoids. Overall, we infer that the lower presence of these compounds
following VPH supply is related to healthier plants with reduced
levels of ROS. These results are in accordance with the recent
findings of Sorrentino et al. (2021) where Trainer® VPH treatment
on Arabidopsis thaliana showed a clear downregulation of
terpenoid compounds (Sorrentino et al., 2021). Likewise, Lucini
et al. (2020) pointed out a general decrease of defense compounds,
including alkaloids and terpenoids following the application of the
same VPH on Solanum lycopersicum L. (Lucini et al., 2020). Moreover, it is interesting to note that phenylpropanoid
metabolites were markedly accumulated only in Cu-treated
samples, since VPH+Cu-VHP induced just a weak increase and
noteworthy downregulations were observed with VPH and Cu-
VPH treatments. It is proven that the phenylpropanoid biosynthetic
pathway is triggered under heavy metal stress, leading to the
accumulation of phenolic compounds with ROS scavenging
capacity (Park et al., 2022). Mira et al. (2002) extensively
described flavonoid ability to exert pro-oxidant effects under Cu
metal stress through the metal chelating process. This clearly
explains our finding that the levels of phenylpropanoid and
flavonoid compounds were found to be increased by Cu excess. 4 Discussion Several studies on many other plant
species (Brassica napus L., Brassica juncea L., Cucumis sativus L.,
Oryza sativa L., and Coriandrum sativum L.) showed that plant
exposure to substantial amount of Cu may result in increased ROS
production—hydrogen peroxide (H2O2), superoxide anion radicals
(O2−), and hydroxyl radicals (OH−)—causing damage to cellular
constituents (Shaw and Hossain, 2013; Sunita and Shekhawat, 2016;
Nair and Chung, 2017; Mosa et al., 2018; Alquraidi et al., 2019). ROS produced may cause lipid peroxidation, stimulating the
production of highly active signaling compounds competent of
triggering the production of secondary metabolites in the plant
(Anjitha et al., 2021). FIGURE 5
Illustrative picture of basil plants untreated (Control) or foliarly treated with copper sulfate (Cu), copper complexed with amino acids and peptides
(Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH). FIGURE 5
Illustrative picture of basil plants untreated (Control) or foliarly treated with copper sulfate (Cu), copper complexed with amino acids and peptides
(Cu-VPH), vegetal protein hydrolysate (VPH), and Cu-VPH enriched with VPH (VPH+Cu-VPH). 11 11 Frontiers in Plant Science Frontiers in Plant Science frontiersin.org Rouphael et al. 10.3389/fpls.2023.1235686 Rouphael et al. In this regard, it is well known that alkaloids represent key
components of the plant defense system, being released by the plant
to face stress conditions. In particular, the antioxidant activity of
alkaloids includes the upregulation of ROS scavenging, the
inhibition of ROS-producing enzymes, and the chelation of
metals (Yoon et al., 2006). Lala (2020) reported a significant
increase in the total content of alkaloid metabolites following Cu-
based treatment in Bacopa monnieri L., accomplished by an
increase in ROS markers. Moreover, enhanced production of
alkaloids was found in O. basilicum L. and Catharanthus roseus
L., respectively, following copper sulfate (CuSO4) and copper oxide
(CuO) treatments (Pan et al., 2015; Nazir et al., 2021). Our results are in accordance with previous reports in which CuO
nanoparticles enhanced total flavonoid content in Stevia
rebaudiana, Brassica nigra L., and O. basilicum L. (Zafar et al.,
2017; Javed et al., 2018; Nazir et al., 2021). Moreover, our study is
coherent with the previous work of Sorrentino et al. (2021), which
ascribed to the same VPH a significant downregulation of
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10.1002/jpln.200800174 Caspi, R., Altman, T., Billington, R., Dreher, K., Foerster, H., Fulcher, C. A., et al. (2014). Acknowledgments GC and YR: conceptualization and project administration. YR,
PC, MC, LF, CE-N, PG, AF, BMM, MTC, LL, and GC:
methodology, validation, formal analysis, investigation, and
writing—original draft preparation. YR, PC, MC, LF, CE-N, PG,
AF, BMM, MTC, LL, and GC: software. YR, PC, MC, LF, CE-N, PG,
AF, BMM, MTC, LL, and GC: resources. YR, PC, MC, LF, CE-N,
PG, AF, BMM, MTC, LL, and GC: writing—review and editing. YR,
PC, LL, and GC: visualization. YR, PC, LL, and GC: supervision. YR,
PC and GC: funding acquisition. All authors contributed to the
article and approved the submitted version. The authors are grateful to Giuseppe Iandiorio for his technical
assistance in the field trial. This manuscript reflects only the
authors’ views and opinions; neither the European Union nor the
European Commission can be considered responsible for them. Conflict of interest The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could be
construed as a potential conflict of interest. Supplementary material The Supplementary Material for this article can be found online
at: https://www.frontiersin.org/articles/10.3389/fpls.2023.1235686/
full#supplementary-material Funding All claims expressed in this article are solely those of the authors
and do not necessarily represent those of their affiliated organizations,
or those of the publisher, the editors and the reviewers. Any product
that may be evaluated in this article, or claim that may be made by its
manufacturer, is not guaranteed or endorsed by the publisher. This study was carried out within the Agritech National
Research Center and received funding from the European Union
Next-Generation EU (PIANO NAZIONALE DIRIPRESA E
RESILIENZA (PNRR) – MISSIONE 4 COMPONENTE 2,
INVESTIMENTO 1.4 – D.D. 1032 17/06/2022, CN00000022). This research work was partially funded by Università degli Studi
della Campania Luigi Vanvitelli in the frame of project
'VAnviteLli pEr la RicErca (VALERE)'. This research work
was partially funded by MUR in the frame of the project
‘Digitali, Intelligenti, Verdi e Sostenibili (D.I.Ver.So)’ managed
by DAFNE of Tuscia University. Data availability statement The original contributions presented in the study are included
in the article/Supplementary Material. Further inquiries can be
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https://zenodo.org/records/1585306/files/article.pdf
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English
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PERNICIOUS ANEMIA AND ITS RELATION TO GASTRIC DIGESTION, BASED ON TWENTY-FIVE CASES.
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Journal of the American Medical Association
| 1,904
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public-domain
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PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D. BUFFALO, N. Y. The group of twenty-five cases reported in this paper
represents about one-half of the private and hospital
cases which have come under my observation during the
past ten years. Unfortunately, the records of a number
of cases are so faulty as to render them unavailable for
careful analysis. In presenting these reports the aim is
made to consider the disease from each of three stand¬
points : First, the general condition of the patient ; sec¬
ond, the gastric digestion ; third, the blood. y
j
Third, in a few cases where the anemia was marked,
there was not found
a complete achylia gastrica, but
merely hypochlorhydria. Since Austin Flint's original
observation that there is an absence of gastric secretion
in pernicious anemia, this relation has been constantly
recognized; but it is apparent that the failure of gastric
juice is the result of depression in the blood. It is not
a cause of pernicious anemia, but, on the other hand, in
these cases pernicious anemia is the
cause of achylia
gastrica. In most cases of achylia gastrica the motor
activity of the stomach is exaggerated,
as a result of
which the stomach usually empties itself more quickly
than normal. This rule holds true in those cases which
result from pernicious anemia, but, according to my ob¬
servation, not so often as in other cases of achylia gas¬
trica. This is probably true for the reason that with
the failure in the blood there comes about a failure in
the strength of the muscular coat of the stomach, and in
some cases there is the development of
a real gastric
atony and sometimes gastritis. Perhaps for this reason
some patients complain of food stagnation and anorexia
—conditions which are rarely observed in achylia gas¬
trica from other causes. In some of the cases reported
in this paper diarrhea was an occasional and troublesome
symptom. It appears to me that it occurs less often in
this group than in the same number of cases of achylia
gastrica occurring without pernicious anemia. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. When
present it is usually associated with excessive gastric
motility and the disturbance of intestinal peristalsis by
the untimely emptying of the stomach onward ; also, the
exaggerated motor impulse which empties the stomach
is apparently carried over to the intestine, which influ¬
ences result in the appearance of frequent liquid stools
showing undigested food substances. ,
gast c digestion ;
,
The stomach examinations,
so far
as relates to the
chemistry, were made mostly by Dr. Allen Jones; the
blood examinations by Dr. A. E. Woehnert. Nearly all
of these cases were referred to me for diagnosis, and in·
many it became impossible to follow systematically the
subsequent state of the digestion. As will be seen in the
history of individual cases, the patient almost uniformly
complained
of
dyspnea, weakness and digestive dis¬
turbance, sometimes relating to the stomach, at other
times to the intestines. All of them presented the pic¬
ture of marked pallor, associated with
a more or less
marked lemon-colored tint of the skin, differing from
jaundice, and yet in some instances strongly suggesting
it. In most the loss of weight was not great. In almost
all cases the tongue was strikingly pale, usually showing
a loss of epithelium, and therefore not coated, or but
slightly
so. A proportion of the cases suffered from
marked gastric distress; others complained of no stom¬
ach trouble. In some there was anorexia, but in most
there was a fairly good appetite, and in a few instances
the appetite was increased. In all the gastric digestion
was greatly depressed, and in the majority there was
complete achylia gastrica, no appreciable digestion tak¬
ing place in the stomach. A number of times I have
seen moderate improvement in digestion at intervals
corresponding with improvement in the state of the
blood, but this does not apply to the secretion of gastric
juice. In those cases of pernicious anemia in which the
secretion of hydrochloric acid has disappeared, I have
not seen it return, even when gastric motility has be¬
come apparently normal. The cases in which complete
achylia gastrica was present did not differ, so far as the
stomach examinations are concerned, from the ordinary
type of achylia gastrica. From a careful history of these
cases I am convinced that the achylia gastrica is not an
etiologic factor in the development of pernicious anemia. *Read at the Fifty-fifth Annual Session of the American Med-
ical Association, in the Section
on Practice of Medicine, and
ap-
proved for publication by the Executive Committee :
Drs.
J.
M.
Anders, Frank Jones and W. S. Thayer. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User on 05/25/2015 I have observed, not in the gastric digestion. Possibly,
if all cases had been studied with this in view for a pro¬
longed period, some instances of improved gastric secre¬
tion, corresponding with betterment in the blood, might
have been noted. However, in a few cases in which the
condition of the stomach has been ascertained at regular
intervals during several years, no change for the better
was found, although at times the blood has made re¬
markable improvement. Dr. Frank . Wynn, Indianapolis—Answering Dr. Bergey's
question, so far as organic changes are concerned, there were
none observable. I mentioned in the paper symptomatic evi¬
dences of failure in nutrition in the animals, loss of appetite
and of playfulness. The two groups were near at hand and
could be compared with each other. The .experimental rabbits
were not frolicking about
as the other animals
were, and
most of all, perhaps, the physical evidence of failure of nutri¬
tion was most clearly manifest in the loss of luster of the hair. p
Second, in a large number of cases of achylia gastrica
which I have studied, many of which have been under
observation' during periods of several years, there has
not been found in a single instance a change in the blood
that was suggestive of pernicious anemia. In several
cases of achylia gastrica, associated with diarrhea, there
sometimes develops a severe secondary anemia, but the
characteristics of pernicious anemia have been found
wanting. In this respect my experience agrees with
that of Riegel, Einhorn, and most others who have care¬
fully studied the two subjects. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. While I hoped for a
return of the combined chlorids during these periods of im¬
provement in the patient, I have been
so far disappointed. Starch digestion is moderately well performed when the pa¬
tient is doing well, but even this is greatly diminished when
the blood count is very low. y
Following this is a brief statement of each of the re¬
ported cases. A report of the study of the stomach ex¬
aminations is inserted, and, as will be seen, they repre¬
sent practically what is found in ordinary cases of gas¬
tric anacidity. I very much regret that the records of
some of the stomach examinations have been lost, and
the paper is more incomplete in this connection than I
anticipated ; nevertheless, the uniformity of the results
is
so evident that I feel the
same conclusion may be
drawn from these reports that would follow
a much
longer series. o ge
Most of the patients had an incorrect idea as to the
nature of their disease, some complaining of the liver,
others of cancer of the stomach, heart disease, locomotor
ataxia, and so on. g
Most of the patients had an incorrect idea as to the
nature of their disease, some complaining of the liver,
others of cancer of the stomach, heart disease, locomotor
ataxia, and so on. Case 1.—G. K., German, aged 47, carpenter, married, fam¬
ily history negative; personal history negative until the begin¬
ning of the present illness, two years ago. Patient is weak,
short of breath, has tinnitus aurium, numbness of the
ex¬
tremities,
slow reflexes, gaseous pulse. First sound of the
heart at apex is weak, with slight systolic bruit; aortic sound
exaggerated; arteries sclerosed moderately; capillary circula¬
tion poor; skin pale, lemon tint; appetite fair, afraid to eat
because of subsequent distress; tongue pale, shiny and tender;
spleen not palpable. Treatment, Donovan's solution, etc. Im¬
provement wonderful on two occasions. Died after an illness
of four years. y
Blood.—Red cells, 1,021,000; large number of megalocytes,
few megaloblasts and normoblasts; hemoglobin, 26 per cent.;
leucocytes, 3,200; polymorphonuclear neutrophiles, 60 percent. y
p y
p
p
Case 6.—A. M., aged 38, laborer. Mother died at 45 of un¬
known trouble, one brother from hemoptysis, one sister from
hemoptysis, one of Bright's disease, and the third from sudden
failure of the heart. The patient's previous history has been
good. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. Complained that for some months he had been weak, and
of late had shortness of breath; distress after eating, although
he had a good appetite; lost somewhat in flesh, exact amount
unknown. His pulse was weak and gaseous; arteries very com¬
pressible; heart rather enlarged;
apex diffuse; hemic bruit
heard over preeordium; also bruits de Diable; râles at bases of
both lungs; tongue pale, epithelium largely absent;
bowels
constipated; liver large, one inch below ribs; spleen and lymph
glands normal. Patient's skin was dry, scaly, with bran-like
exfoliations and characteristic lemon-colored tint. A letter
from the patient March 28, 1903, reports that he had been
at work since May last without the loss of a day and appears
to be in fair health. y
Blood.—Red cells, 980,000 ; hemoglobin, 25 per cent. ; megalo-
blasts, megalocytes, microcytes, poikilocyt.es marked;
hemo¬
globin index high; leucocytes, 3,300. Case 2.—S. J. M., American, aged 45, physician. Family
history:
Mother probably died of tuberculosis; in other
re¬
spects negative. Patient always well
until present illness. Had great mental strain for
a year, followed by weakness,
dyspnea, and later by digestive disturbances. Pulse frequent,
gaseous; heart normal in size, systolic bruit at apex, bruits de
Diable. Appetite fair, tongue pale, bowels loose, often diar¬
rhea ; skin lemon tint and pale ; occasionally slight cough ;
lungs normal ; spleen not palpable. Three periods of marked
improvement. Died two years and
a half after beginning of
illness. Blood.—Red cells, 1,199,000; hemoglobin, 23 per cent.; many
microcytes, poikilocytes
and megalocytes;
normoblasts
and
megaloblasts. Leucocytes, 8,000; 50 per cent, polymorphonu¬
clear neutrophiles. Blood.—Red cells, 1,130,600; hemoglobin, 25 per cent.;
ma-
crocytes, microcytes and poikilocytes were numerous; megalo-
blasts and normoblasts; leucocytes, 2,933; polymorphonuclear
cells, 84 per cent. p
Case 7.—S. F., patient of Dr. Roos, Wellsville, N. Y., aged
60, merchant; negative family history; suffered from typhoid
fever 25 years ago, subsequently well until 12 years ago, when
he had a decline and gave up business. He improved again and
was fairly well. Five years ago had what was apparently an
epileptic seizure;
since then has had
several
such attacks. Two months ago began to suffer greatly with his stomach,
nausea, distress especially after eating, occasionally followed by
vomiting. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. There have been
four distinct relapses, followed by striking improvement in
general health, still at no time has there been any evidence of
gastric juice. The gastric motility has varied markedly with
the condition of the blood. When the anemia is at its worst
the stomach is found to empty itself almost immediately after
eating, but when the blood is better the stomach retains the
food for a longer period, and with this return of a more natural
gastric motility the general nutrition improves. What rela¬
tions exist between these facts I am unable to say, but it seems
probable that the intestine is better prepared to carry on the
digestion when
the
stomach discharges
itself
more slowly,
and, therefore, when the intestinal peristalsis, especially in the
upper part of the tract, is less disturbed. While I hoped for a
return of the combined chlorids during these periods of im¬
provement in the patient, I have been
so far disappointed. Starch digestion is moderately well performed when the pa¬
tient is doing well, but even this is greatly diminished when
the blood count is very low. p
p
Case 5.—-Mrs. W. D. R., American, aged 45, patient of Dr. Benninghoff of Bradford, Pa., suffers from weakness, anorexia,
dyspnea. A remarkable case, which I have studied for five
years, and which is still under observation ; has shown great
oscillations in the blood counts—from a few over one million
at the lowest to over four millions at the best, the hemoglobin
ranging from 26 per cent, to 92 per cent. There have been
four distinct relapses, followed by striking improvement in
general health, still at no time has there been any evidence of
gastric juice. The gastric motility has varied markedly with
the condition of the blood. When the anemia is at its worst
the stomach is found to empty itself almost immediately after
eating, but when the blood is better the stomach retains the
food for a longer period, and with this return of a more natural
gastric motility the general nutrition improves. What rela¬
tions exist between these facts I am unable to say, but it seems
probable that the intestine is better prepared to carry on the
digestion when
the
stomach discharges
itself
more slowly,
and, therefore, when the intestinal peristalsis, especially in the
upper part of the tract, is less disturbed. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. My reasons for reaching this view are : First when in g
g
I am somewhat surprised, in reviewing these cases, to
find
so little evidence of marked cardiac disease. In
those
engaged in heavy manual work the heart
was
usually slightly dilated, and sometimes there was a rela¬
tive mitral insufficiency, hemic bruits were often heard,
and sometimes bruits de Diable, but the latter I have not
found so commonly present as in chlorosis. There has
been found no constant change in the liver, spleen or
lymph glands. Often the liver has been found slightly
enlarged, sometimes markedly so ; the spleen sometimes
increased in bulk, but rarely very large. No other strik¬
ing features have been found present with sufficient fre-
quencv to warrant notice, with the exception of symp¬
toms of spinal cord disease. Since Billings called atten- et o og c
p
p
My reasons for reaching this view are : First, when in
a given case the blood shows marked improvement for
several weeks together, there usually appears evidences
of improvement in the general nutrition, but, so far as number of microcytes, poikilocytes and megalocytes,
no
nor-
moblasts;
an occasional megaloblast; leucocytes, 3,800; poly¬
morphonuclear cells, 68 per cent. number of microcytes, poikilocytes and megalocytes,
no
nor-
moblasts;
an occasional megaloblast; leucocytes, 3,800; poly¬
morphonuclear cells, 68 per cent. tion to its frequency I have found it in the majority of
cases. The personal history of the patient is remarkable
in the fact of the absence of previous severe illness. Most of the patients insist that they had never before
been seriously ill. The family histories were negative,
so far as relate to pernicious anemia, and I do not know
of an instance in which it has occurred in the second
generation. The results of study of the urine have not
been very notable in these cases. p
p
Case 5.—-Mrs. W. D. R., American, aged 45, patient of Dr. Benninghoff of Bradford, Pa., suffers from weakness, anorexia,
dyspnea. A remarkable case, which I have studied for five
years, and which is still under observation ; has shown great
oscillations in the blood counts—from a few over one million
at the lowest to over four millions at the best, the hemoglobin
ranging from 26 per cent, to 92 per cent. PERNICIOUS
ANEMIA
AND
ITS
RELATION
TO GASTRIC DIGESTION, BASED
ON
TWENTY-FIVE CASES.*
CHARLES G. STOCKTON, M.D.
BUFFALO, N. Y. He now complains of great weakness; has a lemon
tinted skin, suffers from nausea ; the pulse is very weak ;
ar¬
teries not deeply sclerosed; heart dilated, with musical
sys¬
tolic bruit at apex; tongue pale;
bowels constipated; liver
large palpable below ribs; spleen and lymph glands negative. Former weight 150, now 138 pounds. A letter, March, 1903,
reports that the patient was still living and in fair health. p
Case 3.—A. S., German, aged 40, engineer. Good family
history. The present is the patient's first illness ; began
a
year ago, gradually losing strength ; is now very weak, short of
breath; digestion disturbed; pulse gaseous, weak; heart nega¬
tive ;
anorexia ; tongue pale ;
bowels constipated ;
skin very
pale, lemon tint;
liver and spleen negative;
one period of
marked improvement. Died after two years' illness. p
y
Blood.—Red
cells,
1,366,000;
hemoglobin,
35
per
cent.;
macrocytes in great number ; microcytes, poikilocytes marked ;
leucocytes, 2,400 ; polymorphonuclear cells, 54 per cent. y
p y
p
p
Case 4.—J. M.. American, farmer, aged 70. Family history
good. Present illness began eight months ago. He was pale,
weak,
had dyspnea;
pulse weak,
somewhat
gaseous;
heart
sounds weak, arteries sclerosed; appetite good; tongue pink;
digestion bad; bowels loose: skin pale, sallow, sodden; liver
and spleen negative, reflexes active. Made
no improvement. Died a year after the beginning of the illness. p
p
g
Blood.—Red cells,
1,058,000 ;
hemoglobin,
45
per
cent. ;
megalocytes, microcytes, normoblasts; leucocytes, 8,400; poly¬
morphonuclear neutrophiles, 66 per cent. p
p
,
Case 8.—Captain S., patient of Dr. C. W. Howe, lumber
dealer, aged 70; served three years in Civil War; has suffered
from rheumatic arthritis ; for five years has had much disturb¬
ance with stomach, much
worse for past two years; for
a y
g
g
Blood.—Red cells, 1,248,000; hemoglobin, 30 per cent.; large y
g
g
Blood.—Red cells, 1,248,000; hemoglobin, 30 per cent.; large aded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User on 05/25/2015 Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o The pulse was weak and frequent; heart not indicative; blood
vessels normal; liver and spleen negative; tongue almost col¬
orless; appetite poor; bowels irregular. She died in Febru¬
ary, five months after being discharged from the hospital after
an illness of about three years. g
Blood.—Red
cells,
1,250,000";
hemoglobin,
40
per
cent.;
gigantoblasts, normoblasts, microcytes; poikilocytes
marked;
leucocytes,
4,000;
polymorphonuclear neutrophiles,
70
per
cent. Case 11.—B. P. B., aged 42, caterer, married;
claims to
have been well until four weeks ago, then began to suffer from
progressive weakness and shortness of breath on exertion. No
emaciation; marked lemon-tinted skin; tongue pale; appetite
poor; bowels loose; pulse gaseous, frequent; apex of heart at
the nipple line; systolic bruit at apex and heard in axilla;
liver and spleen normal. Case 11.—B. P. B., aged 42, caterer, married;
claims to
have been well until four weeks ago, then began to suffer from
progressive weakness and shortness of breath on exertion. No
emaciation; marked lemon-tinted skin; tongue pale; appetite
poor; bowels loose; pulse gaseous, frequent; apex of heart at
the nipple line; systolic bruit at apex and heard in axilla;
liver and spleen normal. Case 11.—B. P. B., aged 42, caterer, married;
claims to
have been well until four weeks ago, then began to suffer from
progressive weakness and shortness of breath on exertion. No
emaciation; marked lemon-tinted skin; tongue pale; appetite
poor; bowels loose; pulse gaseous, frequent; apex of heart at
the nipple line; systolic bruit at apex and heard in axilla;
liver and spleen normal. Blood.—Red cells, 949,333 ; hemoglobin, 35 per cent. ; mega¬
locytes, poikilocytes marked, normoblasts
and megaloblasts;
leucocytes, 3,240; polymorphonuclear
neutrophiles,
63
per
cent. Blood.—Red cells, 1,118,000; hemoglobin, 25 per
ent. ;
mi¬
crocytes, macrocytes, poikilocytes present; leucocytes, 6,000;
polymorphonuclear neutrophiles, 75 per cent. p
Blood.—Red cells, 949,333 ; hemoglobin, 35 per cent. ; mega¬
locytes, poikilocytes marked, normoblasts
and megaloblasts;
leucocytes, 3,240; polymorphonuclear
neutrophiles,
63
per
cent. p
Blood.—Red cells, 949,333 ; hemoglobin, 35 per cent. ; mega¬
locytes, poikilocytes marked, normoblasts
and megaloblasts;
leucocytes, 3,240; polymorphonuclear
neutrophiles,
63
per
cent. p y
p
p
Case 16.—B. W., German, aged 42, indoors, machinist; fam¬
ily history negative; personal history good; worked in an ice¬
house for
15 years; lost slightly in weight; present illness
began two months ago with weakness, dyspnea
on exercise,
biliousness
and slight
edema of the
ankles. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o He died in
1902, two and a half years after beginning of illness. y
g
g
Blood.—Red
cells,
1,100,000;
hemoglobin,
40
per
cent.;
megalocytes,
numerous;
megaloblasts, 1;
leucocytes,
4,463;
polymorphonuclear neutrophiles, 46 per cent. y
g
g
Blood.—Red
cells,
1,100,000;
hemoglobin,
40
per
cent.;
megalocytes,
numerous;
megaloblasts, 1;
leucocytes,
4,463;
polymorphonuclear neutrophiles, 46 per cent. p y
p
p
,
p
Case 9.—C. J. M., aged 33, farmer; family history negative;
patient had no previous illness; complained of weakness, dysp¬
nea, faintness; pulse weak, gaseous; has hemic bruit over the
heart; arteries very compressible; tongue is denuded of epi¬
thelium, pale; appetite capricious; bowels irregular with
oc¬
casional diarrhea; skin lemon tint; liver and spleen negative. Dr. Shugert, of Tidioute, Pa., his physician, writes me that
three months after the patient's visit to
me he had
so far
improved as tó resume his work, and that he has continued,
with slight interruptions, to work ever since that time, and
seems to be in very good condition. His work is out-of-doors,
quite laborious, and he walks several miles a day. His color,
June, 1904, is good, and the man is at work. g
,
Blood.—Red cells, 2,118,400; hemoglobin, 51 per cent. Large
number of microcytes, poikilocytes and megalocytes ; one mega-
loblast;
leucocytes,
12,334;
polymorphonuclear neutrophiles,
55 per cent. p
Case 10.—H. B., English, aged 46, plumber, married; family
history good; never sick until present trouble; complained of
his stomach first one year ago; had chills with fever several
days in
succession eight
months later. Present condition,
weak, cachectic, losing flesi,, dyspnea; pulse gaseous;
heart
of normal size; pulmonary second sound exaggerated;
appe¬
tite poor; tongue pale; bowels irregular (constipation or diar¬
rhea) ; skin pale, sallow, with lemon tint; spleen just palp¬
able; mentality good; sleep undisturbed. Had several peri¬
ods of improvement,
some of them quite remarkable. Treat¬
ment, Donovan's solution, etc.; died three years after begin¬
ning of illness. y
p
y
Blood.—Red cells, 1,700,000; hemoglobin,
33 1/3 per cent.;
megaloblasts,
normoblasts,
macrocytes,
microcytes,
poikilo¬
cytes;
some leucocytosis ; polymorphonuclear neutrophiles, 38
per cent. Case 15.—Mrs. I. J. S., patient of Dr. C. A. Ellis, Sherman,
N. Y., aged 34. Patient had been ill for two and a half years,
sometimes showing betterment, followed by periods of depres¬
sion. Her phief symptoms
were
weakness
and
dyspnea. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o The-e
was
a
marked lemon-colored tint to the skin; tongue very smooth,
with absence of papillae; muscles flabby; pulse weak, small;
lungs normal ; heart of normal size, systolic bruit at apex ;
right heart slightly dilated, slight bruit de Diable; liver con¬
siderably enlarged; spleen palpable. Case 12.—J. K., telegraph operator, aged 33, single; present
illness began within six months with weakness and dyspnea;
marked pallor, with lemon-colored tint to skin; tongue pale;
appetite fair;
bowels very loose, with yeasty, dark colored
stools; patient appears well nourished; pulse soft, about 100;
temperature,
101
degrees;
apex of heart diffused;
systolic
bruit heard over the base of the heart, with a loud venous hum
of the vessels of the neck; liver a trifle full; spleen normal;
reflexes somewhat exaggerated. y
g
;
p
p p
Blood.—Red cells, 1,560,000; hemoglobin, 30 per cent.; a few
normoblasts; leucocytes, 2,600. y
,
Case 17.—J. L., aged 49, laborer; family history negative;
patient has handled nitroglycerin for
25 years;
no illness
save that two years ago he froze his hands and feet while
shoveling snow; has not been well since; complains of dyspnea
on exercise; great weakness; numbness of extremities; lost 30
pounds in weight; has poor appetite; no cough or hemorrhage;
heart of normal size, apical systolic bruit, no bruit de Diable;
liver slightly enlarged, two inches below ribs; spleen not pal¬
pable; skin sallow and pigmented; tongue smooth and pale;
bowels constipated. y
Case 17.—J. L., aged 49, laborer; family history negative;
patient has handled nitroglycerin for
25 years;
no illness
save that two years ago he froze his hands and feet while
shoveling snow; has not been well since; complains of dyspnea
on exercise; great weakness; numbness of extremities; lost 30
pounds in weight; has poor appetite; no cough or hemorrhage;
heart of normal size, apical systolic bruit, no bruit de Diable;
liver slightly enlarged, two inches below ribs; spleen not pal¬
pable; skin sallow and pigmented; tongue smooth and pale;
bowels constipated. gg
Blood.—Red cells, 1,127,000; hemoglobin, 31 per cent;
ma-
crocytes, microcytes
and poikilocytes marked;
normoblasts
present; leucocytes, 3,900; polymorphonuclear neutrophiles, 75
per cent. Case 13.—-J. D., aged 40, laborer; duration of illness not
known; complained of extreme weakness and digestive dis¬
turbance; some dyspnea; has marked lemon tint to skin. Case 13.—-J. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o Case 14.—S. 0., Freedom, . ., aged 46, excellent family
history; patient had pneumonia when he
was 21 years old,
otherwise perfectly well until
a year ago, when his health
gradually declined;
he complained of weakness, shortness of
breath, disturbance of the stomach; pain above the pubes, re¬
lieved by the action of the bowels. His pulse was extremely
weak, arteries thickened, the heart slightly enlarged, systolic
bruit at the apex, the pulmonary second sound exaggerated. His appetite was good, at times craving; tongue smooth, de¬
nuded of epithelium, pale; bowels very constipated; skin pale,
almost lemon tint; liver large, extending 1% inches below the
ribs; spleen and lymph glands negative; patient weighed 155
pounds, now 137. Patient was given the bichlorid of mercury
for two months and made marked improvement; he had com¬
plete achylia gastrica. Two months later he was given potas¬
sium iodid;
he suffered greatly from proctitis. After six
months patient was not
so well. During the following year
he again made improvement under the influence of bichlorid
of mercury, nux vomica and iron, and on the 9th of January,
1895, he was discharged as cured. During the following sum¬
mer, 1895, he returned with all his former symptoms. This
time the blood showed the characteristics of pernicious anemia. An incomplete examination made two years before showed 35
per cent, hemoglobin, the red corpuscles very large and de¬
formed. This
case possesses special interest for the
reason
that after passing through my hands he was treated by
an¬
other physician, who claimed to have found the ova of anky-
lostoma, and furthermore he reported that the patient made
great improvement under treatment by thymol. The stools
were examined by Dr. Woehnert, who was unable to find the
ova of ankylostoma, and the blood
showed
no eosinophilia. The case may be put down as not being one of ankylostomiasis. The patient died in 1900, eight years after the beginning of
the attack, after at least three periods of distinct improve¬
ment. Not under my observation for the past three years. year has vomited at times excessively;
none for past three
months;
suffered from exaggerated salivation;
suffers most
from eating solid food; has weakness and dyspnea; is pale;
has arteriosclerosis ; pulse of fair tension ; heart slightly large ;
aortic second sound exaggerated; tongue coated and flabby;
bowels constipated;
liver and spleen negative. p
Blood.—Red cells, 1,150,000; hemoglobin, 50 per cent. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o He was very weak, but finally made great improvement and
went to work. A year ago he had a second severe illness, with
great weakness, lemon-colored tint and bleeding from hemor¬
rhoids; marked anasarca lasting two or three months. This
again disappeared and the man was able to return to work. During each of these attacks the liver was large. The present
attack occurred in September last; became yellow, had chills
and fever, great weakness; became nervous with dyspnea on
exercise; bleeding from the hemorrhoids and moderate
ana¬
sarca. Pulse was weak and gaseous; temperature from 98 to
102; respirations, 24 to 26. Heart was moderately dilated,
systolic bruit at the apex and over pulmonary areas. Lungs
were normal ; arteries slightly sclerosed ; had paresthesia in
both legs, some weakness in gait; appetite was poor, with dis¬
tress after eating; tongue was smooth and shiny; liver was
very large, as also was the spleen; bowels were loose; absence
of ova of parasites. Case 22.—W. O'D., aged 48, lumberman ; family history neg
ative; personal history, had been well previous until two years
ago, when he began to suffer from weakness, with vomiting,
dizziness, dyspnea
on exercise, edema of the extremities and
numbness of the legs and feet. Had
a marked lemon-colored
tint; pale tongue; slightly enlarged liver; bowels loose. His
heart was slightly dilated, pulse weak, but not gaseous. He
had had one period of improvement. The urine showed
a few
hyaline casts, a trace of albumin; otherwise negative; stomach
contents showed complete achylia gastrica. p
y
g
Blood.—Red cells, 755,300; hemoglobin, 25 per cent; there
were many macrocytes, microcytes, normoblasts and megalo¬
blasts; leucocytes, 5,200; the white cells showed 73 per cent,
of the polymorphonuclear form. p y
p
Case 23.—Mrs. T. H. R., aged 41, farmer's wife; patient of
Dr. Rich, Kennedy, N. Y. Family history good ; personal his¬
tory good;
never been sick until present illness. Trouble be¬
gan one year ago, suffering from weakness, dyspnea on exer¬
cise, some dizziness, but especially from paresthesia and
some
anesthesia of the extremities ; some difficulty in walking. Ap¬
petite
was good; pulse very weak, not gaseous. SUMMARY. Of the 24 cases here reported, the youngest was 32. the oldest 70 ; average, 45 years + ; there were 20 males
and 4 females; 13 had outdoor occupation and 11 in¬
door ; the longest duration of illness was in one 5 years,
the shortest 6 months, and fatal; 12 are known to be
dead, 5 living, the others unknown; all had dyspnea on
exercise; all suffered from faintness and weakness; 16
had gaseous pulse, 8 did not; 16 had heart slightly en¬
larged, 8 did not; 16 had systolic apical bruit, in 8 it
was absent; 6 had bruit de Diable, in 18 it was absent; 8
had other bruit, 16 had no other; 22 had
a distinct
lemon-colored tint of the skin, in 2 it was absent ; 15 had
the tongue strikingly denuded of epithelium, 9 did not ;
10 had good appetite, 7 capricious, and 7 anorexia; 18
complained of symptoms of stomach trouble, 6 did not ;
10 had constipation, 8 diarrhea, and 6 irregularity of
the bowels; in 11 the liver was enlarged, in 13 it was not
enlarged, small in none ; 1 had four periods of improve¬
ment, 3 two periods, 8 one period, and 8 no period of
improvement. After my attention was called to the sub¬
ject by Billings' paper, I found signs of spinal cord in¬
volvement in 6 out of 9 cases. Nine cases showed slight
dilatation of the stomach; in 13 the gastric digestion
was absent, in 8 it was very low, in the 3 others it was
fairly good. Fnye showed evidences of gastric catarrh ; p
Blood.—Red cells, 1,363,000; hemoglobin, 38 per cent.;
ma¬
crocytes,
microcytes,
poikilocytes
numerous;
no
nucleated
cells; there were 8,000 leucocytes, of which 62 per cent, were
of the polymorphous form. After the first month considerable
improvement
was shown. The patient is
now undergoing
a
period of improvement. p
p
Case 21.—P. J. K., aged 35, cabinet-maker; family history
negative; personal history, had been well previous to present
illness, which began about
18 months
ago with the usual
symptoms of weakness, dyspnea on exercise, indigestion,
occa¬
sional vomiting with constipation. The heart was of normal
size; weak systolic apical bruit, but no bruit de Diable. The
stomach contents showed complete achylia gastrica. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o McN., aged 46, liquor dealer; family his¬
tory negative; personal history good until the beginning of the
present illness last summer, at which time he began to lo>e
strength; became sallow; suffered from shortness of breath on
exercise; had vertigo and diarrhea. His pulse was very soft;
heart of normal size, with apical systolic bruit and systolic
bruit at the aortic orifice and bruit de Diable; blood vessels
were in good condition; lungs sound; appetite fair; moderate
complaint of stomach; skin has the characteristic lemon tint. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o Heart was
moderately dilated, no bruit; had had nose bleed once; little
complaint of the stomach; had complete achylia gastrica; food
present unchanged three hours after meal; tongue pale, but
not denuded; liver large, two inches below ribs; bowels fre¬
quently loose;
skin pale, but absence of lemon tint;
urine
scant, urea low. No period of improvement. p
Blood.—Red cells, 640,000; hemoglobin, 30 per cent.;
a few
megaloblasts
and normoblasts, with
many large
and small
cells; leucocytes, 8,200. Patient made
no improvement and
died April 1. Blood.—Red cells, 1,964,000; hemoglobin, 40 per cent.;
ma¬
crocytes, microcytes, poikilocytes abundant, with a few megalo¬
blasts; lymphocytes showed 57 per cent, of the polymorphous
form. p
Case 19.—Mrs. C. H., aged 39; father died of cancer of the
stomach; in other respects family history good; personal his¬
tory good
save that three months ago suddenly lost strength
and color ; had dyspnea
on exercise, dizziness, bleeding from
the nose; tinnitus; moderate anasarca, and some tremor and
weakness in her extremities;
pulse
gaseous;
temperature
slightly raised; heart slightly enlarged; systolic apical bruit;
bruit de Diable;
blood vessels good;
râles at base of both
lungs; slight cough; mucous expectoration; stomach irritable;
vomiting in the morning, and distress after eating; tongue
was pale;
had had diarrhea for two months;
skin distinct
lemon tint;
some edema; regular but pale menstruation. Pa¬
tient had steadily failed since first attack, with no improve¬
ment;
now in critical condition. Case 24.—B. J. H., aged 39, male, railroad engineer; family
history negative; personal history, health uniformly good until
a year ago, when he began to grow weak and suffer from his
stomach. These symptoms have been progressive until the
present time. Former weight, 225 pounds, now 184. He suf¬
fers from dyspnea on exercise, weakness, dizziness; the tongue
is denuded and dry; appetite poor, gastric distress, occasional
vomiting and constipation. He
came supposing that he suf¬
fered from some disease of the stomach. His heart is dilated,
relative mitral insufficiency
and moderate bruit
de Diable. Stomach shows achylia gastrica. Urine is negative. History
of past luetic infection 20 years before. On Donovan's solu¬
tion he made improvement during the first six months, but his
trouble became worse and he died April 1, 1904. Blood.—Red cells, 1,735,000; hemoglobin not taken; many
normoblasts, megaloblasts. There were 5,000 leucocytes and
70 per cent, of polymorphonuclear cells. p y
p
Case 20.—G J. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User o D., aged 40, laborer; duration of illness not
known; complained of extreme weakness and digestive dis¬
turbance; some dyspnea; has marked lemon tint to skin. y p
Blood.—Red cells, 1,352,000; hemoglobin, 25 per cent; poi¬
kilocytes,
macrocytes,
megaloblasts,
numerous;
leucocytes,
4,000; polymorphonuclear neutrophiles, not given. y p
Blood.—Red cells, 1,352,000; hemoglobin, 25 per cent; poi¬
kilocytes,
macrocytes,
megaloblasts,
numerous;
leucocytes,
4,000; polymorphonuclear neutrophiles, not given. p
Blood.—Red cells, 1,150,000; hemoglobin, 50 per cent. p
Blood.—Red cells, 1,150,000; hemoglobin, 50 per cent. ded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User on 05/25/2015 Case 18.—F. D., aged 47, farmer; patient of Dr. Tompkins
of Randolph, N. Y. Father died of diabetes; in other respects
family history is good. Patient had for years experienced
severe bilious attacks with vomiting and sallowness,
accom¬
panied by chills and fever. Four years ago, after severe influ¬
enza, he had one of these attacks with jaundice, accompanied
by general anasarca, at which time the scrotum was drained. He was very weak, but finally made great improvement and
went to work. A year ago he had a second severe illness, with
great weakness, lemon-colored tint and bleeding from hemor¬
rhoids; marked anasarca lasting two or three months. This
again disappeared and the man was able to return to work. During each of these attacks the liver was large. The present
attack occurred in September last; became yellow, had chills
and fever, great weakness; became nervous with dyspnea on
exercise; bleeding from the hemorrhoids and moderate
ana¬
sarca. Pulse was weak and gaseous; temperature from 98 to
102; respirations, 24 to 26. Heart was moderately dilated,
systolic bruit at the apex and over pulmonary areas. Lungs
were normal ; arteries slightly sclerosed ; had paresthesia in
both legs, some weakness in gait; appetite was poor, with dis¬
tress after eating; tongue was smooth and shiny; liver was
very large, as also was the spleen; bowels were loose; absence
of ova of parasites. Case 18.—F. D., aged 47, farmer; patient of Dr. Tompkins
of Randolph, N. Y. Father died of diabetes; in other respects
family history is good. Patient had for years experienced
severe bilious attacks with vomiting and sallowness,
accom¬
panied by chills and fever. Four years ago, after severe influ¬
enza, he had one of these attacks with jaundice, accompanied
by general anasarca, at which time the scrotum was drained. DISCUSSION. Dr. George Dock, Ann Arbor, Mich.—I agree with the re¬
marks made concerning the condition of the stomach and its
relation or lack of relation to the disease. I have had a great
deal of experience with hydrochloric acid. I have treated
these cases with hydrochloric acid, fresh air and diet. Many
of them have done as well as those with Fowler's solution and
as others treated without any medicine at all. I have seen
many patients with advanced pernicious anemia get well tem¬
porarily under fresh air and the same sort of diet as given to
tuberculous patients. These patients may take 40, 50
or 60
minims of hydrochloric acid, diluted, after
meals. It has
been my experience that if they could not take the acid well
they did not do well. The effect of the hydrochloric acid does
not seem to have to do so much with stomach as with intes¬
tinal digestion. The condition of the intestine predominates
in nutrition. In all cases the condition of the intestinal diges¬
tion should be determined by an examination of the stools. Dr. George Dock, Ann Arbor, Mich.—I agree with the re¬
marks made concerning the condition of the stomach and its
relation or lack of relation to the disease. I have had a great
deal of experience with hydrochloric acid. I have treated
these cases with hydrochloric acid, fresh air and diet. Many
of them have done as well as those with Fowler's solution and
as others treated without any medicine at all. I have seen
many patients with advanced pernicious anemia get well tem¬
porarily under fresh air and the same sort of diet as given to
tuberculous patients. These patients may take 40, 50
or 60
minims of hydrochloric acid, diluted, after
meals. It has
been my experience that if they could not take the acid well
they did not do well. The effect of the hydrochloric acid does
not seem to have to do so much with stomach as with intes¬
tinal digestion. The condition of the intestine predominates
in nutrition. In all cases the condition of the intestinal diges¬
tion should be determined by an examination of the stools. Dr. R. C. Cabot, Boston—Of 150 eases of pernicious anemia
seen by me, five-sixths showed the absence of hydrochloric acid
and yet digestion went
on perfectly well. DISCUSSION. One is almost in¬
clined to say that hydrochloric acid is a luxury and not a ne¬
cessity. In
cases with poor digestion the trouble
seems due
rather to motor insufficiency of the stomach, and not to the
absence of hydrochloric acid. It has been my experience that
there has often been an absence of hydrochloric acid even dur¬
ing periods of improvement. There is also
a curious lack of
cardiac symptoms in
some cases,
even when we know that
advanced fatty degeneration is present. y
g
p
Dr. Max Einhorn, New York—While
some time ago the
opinion prevailed that pernicious anemia was due to atrophy
of the stomach, recently this view had to be changed. This
point has been brought up by Dr. Stockton and myself. -There
have been met many cases of pernicious anemia in which the
gastric digestion is not impaired. Again, most cases of achylia
gastrica do not show a condition of pernicious anemia. Per¬
nicious anemia may be complicated with achylia gastrica and
atrophy of the stomach. But the latter condition is not the
cause, but rather the result of pernicious anemia. y
Dr. Charles G. Stockton—I can not wonder, considering
the strange capriciousness shown in this disease, at the con¬
clusion reached by Dr. Cabot that hydrochloric acid is a lux¬
ury and not a necessity. However, I believe, perhaps for the
first time, that I must take different ground from Dr. Cabot,
as I regard hydrochloric acid as more than a luxury, though
perhaps not a necessity. The fact brought out by Dr. Einhorn
that in pernicious anemia we so often find serious atrophy of. the gastric mucosa seems to me to answer the point raised by
Dr. Osier, who suggests that in some cases the gastric atony
may bear a causal relation to the pernicious anemia. I would
like to ask Dr. Osier if he does not think that in view of the
serious degenerations of other organs in pernicious anemia, it
might easily account for the gastric atrophy, such, for in¬
stance, as he described
some years ago in a case reported in
conjunction with Dr. Henry; that this atrophy is more likely
to be the result of the pernicious anemia than the
cause of
that disease. I am sorry to hear Dr. DISCUSSION. Osier make the state¬
ment, for it leads me to feel that I must be wrong in my view;
nevertheless, my belief is strongly to the contrary. It seems
to me that we should draw a distinct line between ordinary
achylia gastrica with atrophy of the gastric mucosa and those
cases of gastric atrophy which appear in the course of per¬
nicious anemia. Attention has been called by Dr. Dock to the
fact that these cases improve under hydrochloric acid, and I
wish to say that I have universally given that remedy where
patients could take it. In some instances improvement has fol¬
lowed, but it is difficult to say what actually does good in this
disease. Formerly patients apparently did well under arsenic,
and now we wonder why they sometimes do well without it. One patient whom I have had under observation for three years
wrote me of his condition a month ago. He is still at work and
under no treatment. However, there
are
cases that improve
under hydrochloric acid, and this I can not deny. In some of
my cases examinations of the stools have been made with nega¬
tive results as regard evidences of parasites. I am sorry that
more new knowledge has not been added by our consideration
of this disease. ,
p
De. Allen Jones, Buffalo—I studied the gastric contents
in these cases that Dr. Stockton has reported and I feel like
confirming the view that pernicious anemia is not caused by
achylia gastrica so far as we can tell. For a number of years
after the idea was exploited that pernicious anemia was due
to failure of the secretory functions of the stomach, it was
found that achylia gastrica
appeared
without
pernicious
anemia more often than with it, and yet I felt like hesitating
before stating positively that this condition of the stomach
was not the cause of pernicious anemia. It is very gratifying
to hear Dr. Stockton and others with larger experience than
mine make this statement as a clinical fact. The point I wish
particularly to emphasize is that in gastric work achylia gas¬
trica without pernicious anemia is
more often encountered
than with it. Db. William
Osler,
Baltimore—My
experience
agrees
largely with Dr. Stockton's and Dr. Cabot's. It is wonderful
to see, in the great majority of cases, when a diagnosis has
been made, how rapidly these cases improve. SUMMARY. He died
in the spring of 1903 after an illness of little over two years,
with one period of improvement. p
p
Blood.—Red cells, 950,000; hemoglobin, 35 per cent.; many
megalocytes
and poikilocytes;
there
were four
normoblasts
and 10 macroblasts in one field. There were 2,500 leucocytes,
of which only 15 per cent, belonged to the polymorphonuclear
type. Downloaded From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User on 05/25/201 nicious anemia and not with
a secondary anemia due to un-
einariasis or other intestinal infection. Little has been said
regarding treatment. It has long been thought that arsenic
did good because of its antiseptic effect, and those who have
laid especial stress on the frequent presence of pyorrhea alveo-
laris have thought that here the antiseptic effect was especially
prominent, yet I have recently seen one patient improve won¬
derfully, so that he has resumed his practice, under the hypo¬
dermic use of one-half grain of cacodylate of sodium daily. He
had utterly failed to improve under the long-continued use of
Fowler's solution. If the arsenic did any good here it
cer¬
tainly was not through its antiseptic effect in the intestinal
canal as when given by mouth. Pyorrhea was present in this
case. in the others it was absent. Six showed no acidity of the
gastrici contents, 1 (the highest) showed a total acidity
of 46, one 26, and the average about 10. Four showed
the presence of combined chlorids; in the others it was
absent. One showed free hydrochloric acid, .05 per cent.,
one .06 per cent., and one .022 per cent. ; in all the others
it was absent. Six showed lactic acid present; in the
others it was absent. Four showed other organic acids;
the others, none. The acid salts were very low, 18 being
the highest, 5 having none whatever; 7 showed biuret
reaction ; six showed rennet present, in the others it was
absent. The digestion of starch was poor in all save
two cases. DISCUSSION. There are
cer¬
tain cases in which from the outset the stomach symptoms so
dominate the whole scene as to suggest that the profound and
increasing anemia is due to the gastric condition. Such cases,
though, are extremely
rare. In
a
case reported
some years
ago the entire mucous membrane of the stomach was cuticular,
similar to that found in the esophageal end of the stomach in a
horse. I should like to ask Dr. Stockton if he has made any
observations on the condition of the mouth, if oral sepsis had
anything to do with it. I do not think it has in the majority
of
cases. We have had a large number of cases this spring,
almost an epidemic of pernicious anemia, but in not one was
the oral condition extreme. We are still "at sea"
as to the
exact pathology of pernicious anemia. p
gy
p
Dr. J. N. Hall, Denver—Much has been said regarding the
absence of hydrochloric acid. I have, in such cases, seen fail¬
ure to improve when not administered and yet improvement
was striking when hydrochloric acid was given in 25-minim
doses. The absence of any
excess of eosinophiles should be
noted, as it generally indicates that we are dealing with per- What to eat and what to drink will always be decided by
national custom and individual preference, so far as the public
is concerned, but both may be influenced in the right direction
by the guidance of skilled medical opinion.-—Brit. Med. Jour. d From: http://jama.jamanetwork.com/ by a Penn State Milton S Hershey Med Ctr User on 05/25/2015
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https://openalex.org/W1978643214
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https://zenodo.org/records/2410530/files/article.pdf
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German
| null |
Ueber den inneren Widerstand des Westonelementes
|
Annalen der Physik
| 1,900
|
public-domain
| 1,928
|
848 848 11. Ueber dem i m e r e m Wiclerstamd des Weston-
e Iernerbtes; vom Ipn ax K I e m eln d i E. 1) I. KlemenEi6, Wied. Ann. 66. p.917. 1898.
2) E. H o s e , Zeitschr. f. Elektrochemie Nr. 35. 1900. 2) E. H o s e , Zeitschr. f. Elektrochemie Nr. 35. 1900. 1) I. KlemenEi6, Wied. Ann. 66. p.917. 1898. 2) I. KlemenbiZ, Ber. d. naturw.-med. Yereinee in Innsbruck,
23. Jahrg. 1896-1897. 11. Ueber dem i m e r e m Wiclerstamd des Weston-
e Iernerbtes; vom Ipn ax K I e m eln d i E. Vor einiger Zeit habe ich in einer Notiz') auf die prak-
tische Bedeutnng des inneren Widerstandes der Normalelemente
hingewiesen. Im allgenieinen wird durch einen kleineii inneren
Widerstand die praktische Brauchbarkeit des Elementes wesent-
lich erhiiht. Es sind schon von verschiedenen Seiten Versuche gemacht
worden, den Elementen eine Form zu geben, bei welcher dieser
Widerstand thatskchlich sehr klein ausfallt. Insbesondere ware
in dieser Richtung die Form zu erwahnen, welche vor nicht
langer Zeit Hr. Emil Bose2) beschrieben hat. g
In vielen Fallen muss man jedoch bei der Construction
auf andere Factoren , z. B. die Versandfahigkeit Riicksicht
nehmen und kann daher nicht zugleich auch einen kleinen
inneren Widerstand erzielen. Ein grosser Teil der Experimentatoren wird die An-
schaffung fertiger Elemente vorziehen. Die Fabriken , welche
solche Elemente lieferii, werden dabei natiirlich ' immer die
Versandfahigkeit im Auge behalten miissen, es wird daher auch
der innere Widerstand ziemlich betrachtlich ausfallen. Dieser
Uebelstand ist jedoch nicht so gross, wenn nur der Wider-
stand wenigstens niit der Zeit constant bleibt, ein Umstand,
der bei den bisher meistenteils gebrauchten Clarkelementen nicht
zutrifft. In neuerer Zeit liefert die Firma ,,The European Weston
Electrical Instrument Co." in Berlin ein Normalelement, welches
von der Firma kurz als Westonelement bezeichnet wird. Es
ist gebaut wie das Clarkelement, nur hat es Cadmium statt
des Zinkes und eine bei 4 concentrirte Cadmiumsulfatlosung
ohne iiberschiissige Sulfatkrystalle. 8 49 Annalen cler Physlk. IV. Polge. 2. 1) W. J iiger, Centralblatt fur Accumulatoren uud Elementenkunde
1900.
. ..
._ . . lnnerer Widerstand des Westoneleme7ites. lnnerer Widerstand des Westoneleme7ites. Dieses Element zeichnet sich durch viele wertvolle Eigen-
schaften dern Clarkelemente und auch dem Cadmiumelemente
Init stets conce~~t~-irter
Sulfatlosung gegenuber aus. Ks ver-
dient riach Hrn. W. Jager’) vor allen ancleren Elementen
den Vorzug
Einige Angaben fiber dieses Element, insbe-
sondere ub er seinen inneren Widerstand durften daher allgemein
interessiren. Jn erster Linie waren meine Messungen auf die Unter-
suchung cles inlieren W-iderstandes gerichtet. Der innere Widerstand wurde nach einer Methode be-
stimmt, welche ich vor einiger Zeit beschrieben hsbe.2) Die
Methode ist besoriders gut durchfiihrbar, wenri man zwei gleiche
Normalelemente zur Verfiigung hat. Sie besteht in Folgendem: Kin Galvanometer init feindrahtiger Rolle vom Wider-
stande Q sei durch ein Element voii der elektromotorischen
Kraft 4 und durch den Widerstaaid W geschlossen. Vor dem
Galvanometer befinde sich ein Nebenschluss vom Widerstande w,
wobei w gegen
und W sehr klein ist. Dann ist bekanntlich
die Stromstirke im Galvanometerzweige j gegeben durch . Ew
j = ?-w-
= & a , wo R der Reductionsfactor und cc der Ausschlag des Galvano-
meters ist. wo R der Reductionsfactor und cc der Ausschlag des Galvano-
meters ist. Ich lege nun statt des Widerstandes 10 ein Normalelement
mit dem Widerstande T und der elektromotorischen Kraft 71
in den Nebenschluss. Um die Wirkung dieser elektromotori-
schen Kraft auf das Galvanometer teilweise zu compensiren,
lege ich ein zweites Normalelement mit der elektromotorischen
Kraft e und passender Richtung derselben in den Galvano-
meterzweig. g
Die Stromstirke im Qalvanometerkreise ist danii wobei der Widerstand des compeilsirenden Elementes gegen 0
vernachlassigt ist. Durch Commutiren von B bekommt man wobei der Widerstand des compeilsirenden Elementes gegen 0
vernachlassigt ist. Durch Commutiren von B bekommt man 1) W. J iiger, Centralblatt fur Accumulatoren uud Elementenkunde
1900. . .. - ._
-. - . 54 54 850 I. KlemenEi8. leicht den der elektromotorischen Kraft B und dem Wider-
stande T entsprechenden Ausschlag. Bas /?, a und w h
~
sich der Widerstand T berechnen. Bei den vorliegenden Mes-
sungen bewegte sich der Widerstand W zwischen 40000 und
100000 Q, p zwischen 30000 und 70000 9, und T zwischen
100 und 300 Q. Nur im Falle eines Clark'schen Nementes
war T an 9OO.Q. Es wurden untersucht: Zwei Westonelemente, geliefert von ,,The European Weston
Electrical Instrument Co." in Berlin. Sie werden hier be-
zeichnet rnit /Pi6 und
W,6 ist am 17. October 18A8
und IV,, am 29. lnnerer Widerstand des Westoneleme7ites. October 1898 von Berlin angekommen. ,,
g
Zwei Westonelemente, H-Form, rnit uberschussigen Cad-
mium - Sulfatkrystallen , verfertigt im hiesigen physikalischen
Institute am 8. ,Juli 1898 nach den Vorschriften der Physi-
kalisch-Technisclien Reichsanstalt und bezeiclinet mit W, und WI,. Zwei Clarkelemente. Das eine, H-Form, bezeichnet mit 11,
ebenfalls verfertigt am 25. Mai 1898 im hiesigen physikalischcn
Institute nach den Normen der genaniiten Anstalt; das zweite,
bezeiclinet mit Nr. 160, coristruirt von Fiiss, spater (Sommer
1898) reconstruirt von H a r t m a n n und Braun. Da der innere Widerstand der Elemente von der Tem-
peratur rtbhiingt, musste bei jeder Restimmung auch die Tem-
peratur gemessen werden. Sie wurde im allgemeinen bei den
Westonelementen an einem neben den Ellementen gelegenen
Thermometer abgelesen, was natiirlicli lteine sehr genaue An-
gabe gestattet,; diese diirften daher wohl nur eine Genauig-
keit auf etwa 0,5-1,0"
haben. In Tab. I sind die zu verschiedenen Zeiten geniachten
Messungsresultate verzeichnet. Unter 1' sind die Temperaturen 1. XI. 1898
16. XII. 1895
22. 11. 1899
13. XI. 1899
7. VI. 1900
162
__
161
~
159
158
153
150
147
177
I
169
159
I ,
152
7'
- .. .. -_ - -
-
16,5
17,O
19'0
19,5
16,4 85 1 Innerer Widerstand des Westonelementes. Fur 77, und W,, giebt Tab. II: die Resultate. Fur 77, und W,, giebt Tab. II: die Resultate. Fur 77, und W,, giebt Tab. II: die Resultate. Zum Vergleiche folgeri in Tab. 111 noch die Widerstands-
bestimmungen mit den Clarltelementen ,,H"
und Nr. 160. 11 befand sich in Paraffin61 und die Temperatur wurde an
einem im Bade befindlichen Thermometer abgelesen. Nr. 160
ist von der Fabrik aus mit einem eingesenkten Thermometer
versehen. m
1
1 1
1 T T m
1
1 1
1 T T
. ' Anschliessend moclite ich einige Beobachtungen mitteilen,
welche niit
gcniltclit wurden, um die Aenderung des innereii
Widerstandes mit der Temperntur zu bestimmen. Zu diesem
Zwecke wurdc IV,, in ein Bad von Paraffinol getaucht, welches
auf verschiedene Temveraturen gebracht werden ltonnte. g l!!s
wurde die Temperatur jedcsmal nur langsam erhijht
oder erniedrigt und erst dann beobachtet, wenn die Temperatur
2-3
Stunden constant auf gleicher Hohe war. * 54 * 54 * 852 I. Klem en EiE. Aus den angefuhrten Uaten ergiebt sich ein Temperatur-
coefficient e fur den iiineren Widerstand gleich 0,025. Der Widerstand bei einer gewissen niedrigen Temperatur
wird durch die Temperaturerhohung nicht geandert, im Gegen-
sake Zuni Clark'schen Elemente und vermutlich auch zum
Westonelemente lriit uberschussigen Cadmium-Sulf~tltrystalleii. DAY Clnrltelement 11 hatte z. B. eineri Widerstand von 281 9
bei 18,7 O, nnch einer TemperaturerhGhung auf ca. '28" liatte
es 261 S! bei 1(3,5(' und nach ciner abermaligen ErhGhung
auf 29O tien Widerstand von 243 fi, bei 16,9O. Der Wider-
stand fallt riucli der Erwkmung und es ist ja bekannt, dass
in der Physikalisch- Technischen Reiclisanstnlt der mit der
Zeit zunehmende innere Widerstand der Clarkelemente anf
diese Weise wieder auf einen niederen Wert gebrxcht wird. Der Widerstand bei einer gewissen niedrigen Temperatur
wird durch die Temperaturerhohung nicht geandert, im Gegen-
sake Zuni Clark'schen Elemente und vermutlich auch zum
Westonelemente lriit uberschussigen Cadmium-Sulf~tltrystalleii. DAY Clnrltelement 11 hatte z. B. eineri Widerstand von 281 9
bei 18,7 O, nnch einer TemperaturerhGhung auf ca. '28" liatte
es 261 S! bei 1(3,5(' und nach ciner abermaligen ErhGhung
auf 29O tien Widerstand von 243 fi, bei 16,9O. Der Wider-
stand fallt riucli der Erwkmung und es ist ja bekannt, dass
in der Physikalisch- Technischen Reiclisanstnlt der mit der
Zeit zunehmende innere Widerstand der Clarkelemente anf
diese Weise wieder auf einen niederen Wert gebrxcht wird. Die Teniperaturerhiihung wurde mit dem Clarkelemente 11
zwischen dem 22. Pebruar und 14. November vorgenommen. Ohne diese waren wahrscheinlich die Werte fur H vom
14. Kovember 1899 und 28. Xai 1900 in der Tab. I11 noch
hoher ausgefalleii. Aus den angefuhrten Tabellen ergiebt sich
also, dass der Widerstarid der Westonelemente, wie sie von
der Firrna ,,The Kuropean Weston Electrical Instrument Co."
geliefert werden, zwar im Verlaufe von 1,s ,Jahren ein wenig
gestiegen ist, dass aber dieser Anstieg weitaus nicht jene Hohe
erreicht, die bei den Elementen mit uberschussigen Cadmium-
Sulfatkryatallen vorkommt. Auch die Clarkelemente zeigeii
rnit der Zeit eine sehr starke Zunahme des inneren Wider-
standes; iiberdies ist der absolute Wert desselben bei den in
den Handel gebrachten Clurk'schen Elementeii vie1 grosser
als bei den aus der gemarinten E'abrik stnmmenden Weston-
elenienten. Die Teniperaturerhiihung wurde mit dem Clarkelemente 11
zwischen dem 22. Pebruar und 14. November vorgenommen. Ohne diese waren wahrscheinlich die Werte fur H vom
14. Kovember 1899 und 28. Xai 1900 in der Tab. I11 noch
hoher ausgefalleii. Aus den angefuhrten Tabellen ergiebt sich
also, dass der Widerstarid der Westonelemente, wie sie von
der Firrna ,,The Kuropean Weston Electrical Instrument Co."
geliefert werden, zwar im Verlaufe von 1,s ,Jahren ein wenig
gestiegen ist, dass aber dieser Anstieg weitaus nicht jene Hohe
erreicht, die bei den Elementen mit uberschussigen Cadmium-
Sulfatkryatallen vorkommt. Auch die Clarkelemente zeigeii
rnit der Zeit eine sehr starke Zunahme des inneren Wider-
standes; iiberdies ist der absolute Wert desselben bei den in
den Handel gebrachten Clurk'schen Elementeii vie1 grosser
als bei den aus der gemarinten E'abrik stnmmenden Weston-
elenienten. Die Westonelernerite der ,,The Europeaii Weston Electrical
Instrument Co." siiid also nicht nur n i t Rucksicht auf die
Unabhangigkeit der elektromotorischen Kraft von der Tem-
peratur, sondern auch vori dem fur die praktische Bedeutung
allerdings nur secundar wichtigen Staridpunlrte der Constanx
und des niederen Wcrtes des inneren Widerstandes alleii
anderen Normalelementen iiberlegen. Schliesslich mochte ich noch einige Daten uber die elektro-
motorischen Kriifte der uritersuchten Elemente mitteilen. Hier Innerer Nrider.stand des Westonelementes. 853 kommen nur die Verldtnisse und nicht die absoluten Werte
in Betracht. Das Verliiiltnis der elektromotorischen Krafte
von iV, und 7YIl wurde am 17. (Eingegangen 9. Juli 1900.) October 1898 auf ca. 0,002 Proc. gleich gefunden. An diesem Tage war hei der Zimmertemperatur. Ilics stimmt mit. den Angnben der Physilralisch- Technischen
Reichsaiistalt, wonach die Differenz zwischen der elelrtro-
motorischen Kraft fur die beiden Sorten der Elemente etwa
0,04 Proc. bei 20" betragen soll. Bei 4') solleii die elelrtro-
motorischen Krafte gleich sein. g
Am 7 . Juni 1900 wurde gefunden
1 = 1,00001, 1
'VII
WI = 1,00001, 1
'VII
WI Die Xlemente behielten dalier durch mehr als 1,5 Jalire ein
constantes Vcrh~ltnis der elektromotorischen Kriifte. Die Xlemente behielten dalier durch mehr als 1,5 Jalire ein
constantes Vcrh~ltnis der elektromotorischen Kriifte. AUS den vorliegenden Daten kann man aber auch er-
sehen, dass der Verschluss hei den Westonelementen ein guter
ist, sodass Iteine Verduiistung der Llisung eintritt, was auf
die elektromotorische Krzlft, nicht ohne EinHuss ware. Zwar
sind die Verhaltnisse dcr elel~tromotorischen Kriifte gegen CY,,
irn Verlaufe von 1,5 Jahren etwas gefallen, doch diirfte dies
in dcr Verschiedenheit der Beobachtungsteinperaturen liegen.')
I n n s bruck, Physikal. Inst. d. Univ., Juli 1900. 1) Wiihrcmd sich bci den Westonelementen mit eincr bci 4 O ge-
sattigten Lijsung die elrlrtromotorischc Kraft mit der Temperatur gar
nicht Bndert, ist bei den Elemcnten mit iiberschhsigen Krystallen ein
lrleinrr Temperaturcoefficient vorhanden. (Eingegangen 9. Juli 1900.)
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https://zenodo.org/records/7465069/files/KlGrBr_XXIII-Plenary-7-13.pdf
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Russian
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Стратегия и структура инклюзивного управления предприятием
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Zenodo (CERN European Organization for Nuclear Research)
| 2,022
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cc-by
| 2,477
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Г.Б. Клейнер Г.Б. Клейнер Пленарное заседание Пленарное заседание ПЛЕНАРНОЕ ЗАСЕДАНИЕ ПЛЕНАРНОЕ ЗАСЕДАНИЕ DOI: 10.34706/978-5-8211-0809-8-7-13 СТРАТЕГИЯ И СТРУКТУРА
ИНКЛЮЗИВНОГО УПРАВЛЕНИЯ ПРЕДПРИЯТИЕМ Клейнер Георгий Борисович, член-корреспондент РАН, руководитель научного направле-
ния «Мезоэкономика, микроэкономика, корпоративная экономика» ЦЭМИ РАН, зав. кафед-
рой «Системный анализ в экономике» Финансового университета при Правительстве РФ,
зав. Кафедрой институциональной экономики Государственного университета управления,
george.kleiner@inbox.ru Ключевые слова: инклюзивное управление, расщепление власти, социальное лидерство,
экономика замкнутого цикла. Отечественная экономика вступает в сложный и ответственный период сво-
его развития. Политика санкций, ограничений и давления, которую применяют мно-
гие страны – традиционные партнеры России в области экономики – существенно
затрудняет функционирование и развитие экономики России. Проблемы продолже-
ния деятельности предприятий в сферах производства, науки, спорта и других сфе-
рах заставляют искать новые способы управления экономическими агентами, позво-
ляющие не только выдержать условия стресса, в которых оказались российские
предприятия, но и создать предпосылки для дальнейшего развития в средне- и дол-
госрочной перспективе. 7
По данным Национального агентства финансовых исследований (НАФИ) на
март 2022 г., доля предприятий, считающих свое положение неудовлетворительным,
выросла с 22% в январе – начале февраля до 40% в марте. Доля предприятий, рас-
сматривающих свое состояние как устойчивое, снизилась с 36 до 25%. При этом 51%
предприятий дают негативные прогнозы на ближайшие три месяца, 22% считают,
что ситуация останется без изменений, 27% ожидают улучшений (Оперативный со-
циально-экономический мониторинг…, 2022). По прогнозам Центра стратегических
разработок, экономике предстоит лишиться около 2 млн рабочих мест (Мануйлова,
2022). Меняется, таким образом, и состав самостоятельных экономических агентов
(предприятий) и обстановка внутри предприятия. Это приводит к необходимости ак-
тивизировать выявление новых резервов производства, недостаточно эффективно
использовавшихся ранее. Основным направлением такого поиска является опреде-
ление возможностей перехода к инклюзивному управлению предприятием, т.е. управлению, ориентированному на вовлечение в хозяйственную деятельность всех 7 XXIII Всероссийский симпозиум «Стратегическое планирование и развитие предприятий» материальных и нематериальных ресурсов, работников и др. участников деятельно-
сти предприятия. При этом в хозяйственную деятельность включается и координа-
ционно-управленческая деятельность, так что тенденция демократизации управле-
ния рассматривается как одна из составляющих инклюзивного управления. В настоящей статье излагаются результаты поиска новой структуры управ-
ления предприятием, которая в максимальной степени способствовала бы активиза-
ции деятельности работников предприятий по преодолению последствий изоляции
экономики России в пространстве международного разделения труда. Задача со-
стоит в том, чтобы максимально реализовать потенциал человеческих ресурсов на
предприятиях, приблизиться к инклюзивному управлению, интегрирующему все по-
зитивные факторы деятельности предприятия. Пути решения этой задачи видятся в
оптимизации сочетания формальной структуры управления и неформальных инсти-
тутов социального лидерства на предприятии. СТРАТЕГИЯ И СТРУКТУРА
ИНКЛЮЗИВНОГО УПРАВЛЕНИЯ ПРЕДПРИЯТИЕМ Отличительной особенностью функционирования экономики России в крат-
косрочной перспективе является ее частичная изоляция, работа в режиме «эконо-
мики замкнутого цикла» (Валько, 2020). В таком режиме реализация продукции и
пополнение израсходованных ресурсов осуществляются главным образом на терри-
тории самого государства. Понятия «страна-производитель» и «страна-потреби-
тель» данного вида продукции практически смыкаются. 8
Здесь возникает вопрос общего характера: может ли какая-либо страна или
группа стран существовать в условиях автаркии? Общий положительный ответ дает
сама жизнь: такая экономическая система, как мировая экономика, с самого начала и
до сегодняшнего дня функционирует в условиях автаркии, поскольку не ведет обмена
товарами и услугами ни с одной инопланетой цивилизацией. Если мировая экономика
в течение тысяч лет находится в изоляции, то и Россия как самая большая страна в
мире, видимо, тоже может существовать в режиме частичной изоляции достаточно
длительное время. Главным фактором, обеспечивающим возможность функциониро-
вания в таком режиме, является разнообразие условий экономической деятельности
на территории страны (что аналогично разнообразию экономик стран, входящих в ми-
ровое экономическое сообщество). Такое разнообразие должно учитывать админи-
стративно-территориальное деление страны и распространяться на институциональ-
ные, отраслевые природно-географические и ментальные особенности регионального
развития. Инклюзивное управление страновой экономикой в значительной мере
должно концентрироваться на управлении разнообразием, т.е. поиске и применении
управленческих средств, обеспечивающих оптимальный (не слишком высокий, но и
не слишком низкий) уровень разнообразия. Представляется, что определение опти-
мальных границ (в более общем случае – оптимальное распределение фрагментов
уплотнения и разрежения) условий экономической деятельности является одной из 8 Пленарное заседание актуальных задач современной экономической науки. Проблема разнообразия отно-
сится и к усилению экономической специализации регионов, и к определению раз-
мерностной структуры предприятий и организаций, и в целом к организации разме-
щения и развития производительных сил в страновом пространстве и времени. Здесь
должны взаимодействовать такие отрасли экономической науки, как пространствен-
ная экономика, экономическая динамика и экономика разнообразия. В настоящее время страна должна готовиться к режиму частичной изоляции,
т.е. к экономике замкнутого цикла. Следует подчеркнуть, что Россия обладает
огромными запасами природных ресурсов, включая уникальные природные ре-
сурсы; гигантским запасом человеческого ресурса, включая интеллектуальный, ко-
гнитивный и абсорбционный потенциал; значительным опытом развития за «желез-
ным занавесом». Основными ресурсами производства являются, как известно, труд (включая
предпринимательский и организаторский потенциалы) и капитал (включая физиче-
ский, финансовый, социальный и др. виды). В условиях продолжения изоляционист-
ких тенденций следует ожидать сокращения доступа к физическому капиталу, к со-
временным технологиям и техническим устройствам. СТРАТЕГИЯ И СТРУКТУРА
ИНКЛЮЗИВНОГО УПРАВЛЕНИЯ ПРЕДПРИЯТИЕМ Единственным способом про-
должения производственной деятельности для большинства российских предприя-
тий становится расширение и активизация процессов замещения: во-первых, им-
портозамещения; во-вторых, замещения трудом капитала. При этом процесс соб-
ственного производства импортируемых комплектующих и устройств требует не-
мало времени, в то время как процесс замещения недостающих элементов капитала
за счет активизации трудовой деятельности и модернизации процессов управления
может быть выполнен в достаточно сжатые сроки. Инклюзивное управление пред-
полагает, в том числе, и решение последней задачи. По сути дела, речь идет об инвестициях в эластичность замещения между
физическим капиталом и человеческим трудом. Задача управления эластичностью
замещения не является новой для экономической науки. В 1970–1980 гг., в период
расцвета теории производственных функций, были заложены основы построения
производственных функций с переменной эластичностью замещения факторов
(Клейнер, 1986; Revankar, 1971), см. также (Хацкевич, Проневич, 2017) В настоящее
время внимание к моделям производства с управляемой эластичностью замещения
должно быть усилено; эластичность замещения должна войти в состав предметов
инклюзивного управления. 9
Более конкретно речь идет о замещении физического капитала интеллекту-
альным трудом. Необходимо усилить внимание к уникальным специалистам
(«умельцам»), способным развивать производство в условиях сокращения доступа к
импорту сырья, материалов, оборудования; западное управление талантами XXIII Всероссийский симпозиум «Стратегическое планирование и развитие предприятий» превратить в российское управление «умельцами». Управление «умельцами» при-
звано организовать внутри предприятия непрерывные цепочки добавленной ценно-
сти, в настоящее время терпящие разрывы из-за слома традиционных, внешних для
предприятия, цепочек поставок. В определенном смысле проблемы логистического
обеспечения предприятий должны быть сняты или, по крайней мере, ослаблены за
счет интериоризации – «точечного» управления внутрифирменной логистикой. Та-
ких «умельцев» немного, но в сегодняшних условиях к ним надо относиться более
внимательно. Следует более бережно относиться к таким работникам, обеспечивать
для них определенные стимулы привилегии и в целом более бережно относиться к
человеческим ресурсам предприятий и организаций. Если мы сможем активизиро-
вать креативный потенциал работников, мы сможем преодолеть ограничения, свя-
занные с физическим недостатком капитала, технологий, сырья и т.д. Как же должно быть организовано инклюзивное управление предприятием в
этих условиях? Чтобы обеспечить охват всех аспектов деятельности предприятия,
обратимся к универсальной структуре предприятия, представленной четырьмя базо-
выми секторами (подсистемами) предприятия: объектным, процессным, проектным
и средовым (Клейнер, 2011). Объектный сектор составляют организационные под-
разделения; процессный – совокупность бизнес-процессов, протекающих на пред-
приятии; проектный – краткосрочные локализованные, значимые для предприятия,
мероприятия различного характера; средовой – различные виды таких внутрифир-
менных сред, как институциональная, информационная, социальная и т.п. Эти че-
тыре сектора предприятия нуждаются во внутрисекторном управлении и межсектор-
ной координации. СТРАТЕГИЯ И СТРУКТУРА
ИНКЛЮЗИВНОГО УПРАВЛЕНИЯ ПРЕДПРИЯТИЕМ Для целенаправленного эффективного управления предприятием
необходимо, чтобы указанные секторы были представлены в административной
структуре управления и имели бы административных руководителей. Этого, однако,
недостаточно для инклюзивного управления. Необходимо привлечение сил социаль-
ного регулирования внутрифирменной среды. Предлагается двухкомпонентная
структура инклюзивного управления, основанная на сочетании структур и процес-
сов административного управления, с одной стороны, и социально-лидерского
управления, с другой. В формальной структуре управления выделяются администра-
тивные лидеры каждого из 4-х секторов: объектный лидер, процессный лидер, про-
ектный лидер и средовой лидер. В обязанности объектного лидера входит обеспече-
ние развития предприятия в пространственно-временном аспекте и обеспечение без-
опасности функционирования предприятия. Процессный лидер осуществляет кон-
троль за протеканием внутрифирменных процессов обмена товарно-материаль-
ными, информационными и трудовыми ресурсами между различными подразделе-
ниями предприятия. Сфера действия проектного лидера – технологические и иные 10 Пленарное заседание нововведения и мероприятия. Должностные обязанности средового лидера связаны
с созданием и поддержанием в работоспособном состоянии внутренней инфраструк-
туры предприятия, обеспечением благоприятного социального климата и повыше-
нием привлекательности предприятия для новых работников и контрагентов. нововведения и мероприятия. Должностные обязанности средового лидера связаны
с созданием и поддержанием в работоспособном состоянии внутренней инфраструк-
туры предприятия, обеспечением благоприятного социального климата и повыше-
нием привлекательности предприятия для новых работников и контрагентов. Параллельно со структурой административного управления на предприятии,
переходящем к инклюзивному управлению, создается структура неформального ли-
дерства, также включающая четырех лидеров, осуществляющих свою деятельность
в рамках социальной среды предприятия. В структуру социального лидерства вхо-
дят: инспиративный лидер, обладающий способностями влиять на поведение работ-
ников с помощью личного примера или убеждения; культурный лидер, обладающий
знаниями в сфере технологии и организации производства; интеллектуальный ли-
дер, владеющий информацией о состоянии и динамике рынка в зоне хозяйствования
предприятия и определяющий состав товаров и услуг, предлагаемый предприятием;
духовный лидер, способный определить положение предприятия в перспективной
внешней среде (Клейнер, 2022). Данная структура социального лидерства коррелирует со структурой функ-
ционалистского лидерства, предложенной И. Адизесом (Адизес, 2007) (табл. 1) Системное лидерство, социальное лидерство и целевые функции лидеров
Системные
лидеры
Социальные
лидеры
Функции по
И. Адизесу
Характеристика деятельности организа-
ции, обеспечиваемая данной функцией
Объектный
Инспиративный Е-функция
Результативность в долгосрочной перспек-
тиве
Процессный
Культурный
А-функция
Эффективность в краткосрочной перспек-
тиве
Проектный
Интеллектуаль-
ный
Р-функция
Результативность в краткосрочной перспек-
тиве
Средовой
Духовный
I-функция
Эффективность в долгосрочной перспек-
тиве Системное лидерство, социальное лидерство и целевые функции ли В целом социальная сфера предприятия может рассматриваться как своего
рода полевая структура, индуцированная влиянием соответствующих социальных
лидеров: сфера инспиративного капитала; сфера культурного капитала; сфера интел-
лектуального капитала; сфера духовного капитала. СТРАТЕГИЯ И СТРУКТУРА
ИНКЛЮЗИВНОГО УПРАВЛЕНИЯ ПРЕДПРИЯТИЕМ При этом система управления утра-
чивает свойство иерархической расслоенности и консолидируется. 2. Системное инклюзивное управление предприятием предполагает опору на
сочетание формальной (административной) структуры управления и неформальной
структуры социального лидерства. Такой подход обеспечит максимальную эффек-
тивность использования уникальных специалистов, чей интеллект способен решить
возникающие в условиях изоляции задачи. 3. Принципы бережливого производства должны быть распространены на
бережное отношение к работникам предприятия. В особенности это должно отно-
ситься к тем специалистам, чья квалификация позволит преодолеть разрывы во внут-
рифирменных технологических и внешнефирменных рыночных цепочках поставок
полупродуктов, сырья и комплектующих. 4. Органической частью инклюзивного управления на предприятиях должны
стать процессы демократизации управления, создания на каждом предприятии атмо-
сферы солидарности и взаимного уважения. Особое внимание требуется уделить
процессам расширения в стране популяции бирюзовых, а впоследствии и перламут-
ровых предприятий (Клейнер, 2020). СТРАТЕГИЯ И СТРУКТУРА
ИНКЛЮЗИВНОГО УПРАВЛЕНИЯ ПРЕДПРИЯТИЕМ Соответственно, полевая струк-
тура предприятия может быть представлена как совокупность четырех полей: поля
влияния силы власти; поля влияния культуры; поля влияния интеллекта; поля влия-
ния силы духа. Инклюзивное управление интегрирует интересы, возможности и им-
перативы участников хозяйственной деятельности, обусловленные их принадлежно-
стью к соответствующим внутрифирменным силовым полям. 11 XXIII Всероссийский симпозиум «Стратегическое планирование и развитие предприятий» Социальные лидеры принимают на себя неформальную ответственность за
содержание ответов на следующие вопросы: «что производить и кому продавать?»
(стратегия) – интеллектуальный лидер; «кто должен производить?» (организация
производства) – инспиративный лидер; «как производить?» (технология производ-
ства) – культурный лидер; «зачем производить?» (высшие цели и смыслы производ-
ства) – духовный лидер. Мы видим, что номенклатура производства, состав участ-
ников производства, технология и высшие цели деятельности находятся под идей-
ным руководством неформальных социальных лидеров. Описанное выше распределение функций управления между социальными
лидерами можно назвать «расщеплением власти». Такое расщепление, однако, не
ослабляет власть, а, наоборот, способствует ее укреплению, поскольку опирается на
структуру социальных сил предприятия. В том или ином виде подобное расщепле-
ние распространяется не только на микроуровень (предприятие и его подразделе-
ния), но и на мезоуровень (региональное и отраслевое управление), и на макроуро-
вень (управление социально-экономическим развитием страны). Можно найти эле-
менты подобного расщепления, скажем, в структуре управления судном и в других
управленческих структурах армии. Учитывая, что данная статья отражает содержание пленарного доклада, сде-
ланного 12 апреля 2022 г. в День космонавтики на XXIII Всероссийском симпозиуме
«Стратегическое планирование и развитие предприятий», в качестве примера при-
ведем распределение социальных лидерских ролей в команде С.П. Королева по со-
зданию первого искусственного спутника Земли (реконструкция выполнена на ос-
нове книги воспоминаний С. Хабарова (2021), работавшего вместе с С.П. Короле-
вым, а также ряда других источников): инспиративный лидер – В.П. Мишин, первый
заместитель и впоследствии наследник дела С.П. Королева; духовный лидер –
Л.А. Вознесенский, ответственный за испытания техники; культурный лидер –
С.О. Охапкин, заместитель по конструированию; интеллектуальный лидер –
К.Д. Бушуев, начальник проектно-исследовательского отдела. Формальные институты управления (Chief Executive Officer (CEO), Chief
Technology Officer (CTO), Chief Operating Officer (COO), Chief Financial Officer
(CFO)) должны быть дополнены неформальными институтами лидерства (Leading
Executive Officer (LEO), Leading Spiritual Officer (LSO), Leading Cultural Officer
(LCO), Leading Intelligence Officer (LIO)). Изложенное выше позволяет сделать следующие выводы. Изложенное выше позволяет сделать следующие выводы. 1. Переориентация управления предприятиями на принципы инклюзивного
управления, согласно которым внутрифирменное социальное пространство стано-
вится более однородным, является тем более необходимой, чем более 12 Пленарное заседание изолированной становится экономика страны. СПИСОК ИСПОЛЬЗОВАННЫХ ИСТОЧНИКОВ Адизес И. (2007). Идеальный руководитель: почему им нельзя стать и что из этого следует. М.: Альпина
Бизнес Букс. 262 с. Адизес И. (2007). Идеальный руководитель: почему им нельзя стать и что из этого следует. М.: Альпина
Бизнес Букс. 262 с. Валько Д.В. (2020). Циркулярная экономика: основные бизнес-модели и экономические возможности //
Журнал экономической теории. Т. 17. № 1. С. 156–163. DOI: 10.31063/2073-6517/2020.17-1.12 Валько Д.В. (2020). Циркулярная экономика: основные бизнес-модели и экономические возможности //
Журнал экономической теории. Т. 17. № 1. С. 156–163. DOI: 10.31063/2073-6517/2020.17-1.12 Клейнер Г.Б. (1986). Производственные функции: теория, методы, применение. М.: Финансы и стати-
стика. 238 с. Клейнер Г.Б. (2011). Новая теория экономических систем и ее приложения // Вестник Российской акаде-
мии наук. Т. 81. № 9. С. 794–808. Клейнер Г.Б. (2020). Спиральная динамика, системные циклы и новые организационные модели: перла-
мутровые предприятия // Российский журнал менеджмента. Т. 18. № 4. С. 471–496. DOI:
10.21638/spbu18.2020.401 Клейнер Г.Б. (2022). Социальное лидерство, расщепление власти и инклюзивное управление организа-
цией // Вопросы экономики. № 4. С. 26–44. DOI: 10.32609/0042-8736-2022-4-26-44. Клейнер Г.Б. (2022). Социальное лидерство, расщепление власти и инклюзивное управление организа-
цией // Вопросы экономики. № 4. С. 26–44. DOI: 10.32609/0042-8736-2022-4-26-44. Мануйлова А. (2022). Безработица примеряется к росту // Коммерсант. 2022. 29 марта. URL: р
(
)
р
р
у р
р
цией // Вопросы экономики. № 4. С. 26–44. DOI: 10.32609/0042-8736-2022-4-26-44. Мануйлова А. (2022). Безработица примеряется к росту // Коммерсант. 2022. 29 марта. URL:
https://www.kommersant.ru/doc/5281886 (дата обращения: 11.07.2022). Мануйлова А. (2022). Безработица примеряется к росту // Коммерсант. 2022. 29 марта. URL:
https://www.kommersant.ru/doc/5281886 (дата обращения: 11.07.2022). Оперативный социально-экономический мониторинг. Результаты первой волны мониторинга среди
предпринимателей. (2022). Март
2022
г. НАФИ
//
Сайт
НАФИ. URL:
https://nafi.ru/projects/predprinimatelstvo/pervaya-volna-operativnogo-sotsialno-ekonomicheskogo-
monitoringa-sredi-predprinimateley/ (дата обращения: 11.07.2022). Хабаров С. (2021). Сюжет в центре. М.: Litres. Хацкевич Г.А., Проневич А.Ф. (2017). Квазиоднородные производственные функции единичной эла-
стичности замещения факторов по Хиксу // Экономика. Моделирование. Прогнозирование. № 11. С. 135–140. Revankar N.S. (1971). A class of variable elasticity of substitution production functions. Econometrica, Vol. 39,
No. 1, pp. 1–71. Revankar N.S. (1971). A class of variable elasticity of substitution production functions. Econometrica, Vol. 39,
No. 1, pp. 1–71. 13
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The bird, its form and function,
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a THE BIRD THE BIRD THE BIRD esperornis, — a wingless, toothed, diving bird, about five feet in length, which inhabited
the great seas during the Cretaceous period, some four millions of years ago. Hesperornis, — a wingless, toothed, diving bird, about five feet in length, which inhabited
the great seas during the Cretaceous period, some four millions of years ago. ROBERT DRDMMOND, PRINTER, NEW YORK American j]5ature Series
Group II. The Functions of Nature
THE BIRD
ITS FORM AND FUNCTION
BY
C. WILLIAM BEEBE
Curator of Ornithology of the New York Zoological Park and Life Member of the
New York Zoological Society ; Member of the American Ornithologists'
Union and Fellow of the New York Academy of Sciences
Author of " Two Bird- Lovers in Mexico"
WITH OVER THREE HUNDRED AND SEVENTY ILLUSTRATIONS
CHIEFLY PHOTOGRAPHED FROM LIFE
BY THE AUTHOR
NEW YORK
HENRY HOLT AND COMPANY
1906 Copyright, 1906
BT
HENRY HOLT AND COMPANY
Published September, 1906 DEDICATED
IN GRATITUDE AND ESTEEM
TO
professor "fccnrg ffairfielo ©sborn
BY HIS FORMER PUPIL
THE AUTHOR PKEFACE WE find to-day some thirteen or fourteen thousand
different forms, or species, of birds upon the earth. For
many years ornithologists have laboured to name, and to
arrange in some rational order, these multitudinous forms
of bird life. Some such arrangement is, of course, a neces-
sity— without a handle we should indeed be handicapped
in studying a bird; but let us not forget that classification
is but a means to an end. Far too many students of birds follow some such mode
of procedure as this: When a new bird is found, it is shot,
labelled, preserved in a collection and forgotten; or, if
studying the bird with a glass, all effort is centred in
finding some characteristic by which it can be named,
and, succeeding in this, search is at once made for still
another species, whose name can in turn be added to a list. Observing the habits, the courtship and nest-building,
and memorizing the song, is a third phase of bird-study —
the best of all three methods; but few indeed have ever
given a moment's thought to the bird itself. given a moment's thought to the bird itself. I have lectured to an audience of teachers, every one
of whom was able to identify fifty birds or more, but not
one among them knew the significance of the scales on Preface Vlll a bird's foot. It is to bridge this gap that this book is
intended — an untechnical study of the bird in the abstract. This, it seems to me, is the logical phase of bird life,
which, with an earnest nature-lover, should follow the
handbook of identification — the study of the physical
life of the bird itself preceding the consequent phase
of the mental life, with its ever-varying outward ex-
pression. pression. Far from considering this treatment exhaustive, one
must remember that any chapter subject could easily
be elaborated into one or more volumes. I have intended
the book more as an invitation than aught else: for each
to observe for himself the marvellously fascinating drama
of evolution; to pass on from the nature stories of ideal-
ized composite animals and birds to the consideration of
the evolution of all life; to the tales of time and truth
which have been patiently gleaned by the life-long labours
of thousands of students. PKEFACE Whenever possible I have illustrated a fact with a
photograph from a preparation or from a living bird,
believing that, where verbal exposition fails, pictorial
interest will often fix a fact in the memory. First of all
we must consider a few of the more important and sig-
nificant of the bird-forms of past ages; because no one
who is interested in living birds from any standpoint
should be entirely ignorant of a few facts concerning the
ancestors of these creatures: Otherwise it is as if one,
entirely ignoring the rest of the plant, studied certain
leaves and flowers, knowing not whether they came from
tree or vine. Pre NEW YORK ZOOLOGICAL PARK, May, 1906. Pre ix In my treatment of the various phases of the bird's
physical life I have been considerably influenced by the
many questions which I have heard asked by visitors to
the New York Zoological Park. The short list of books
in the Appendix will indicate the sources whence much
more detailed information may be obtained by those who
desire it. Some two dozen of the illustrations are from outside
sources, and for permission to use these I am indebted
to Dr. William T. Hornaday, the American Museum
of Natural History, Prof. A. Smith-Woodward, Prof. R. S. Lull, A. E. Brown, Esq., Mr. R. H. Beebe, Mr. T. H. Jackson, Mr. Harold Whealton, and Mr. E. H. Baynes;
and for the use of specimens to Dr. F. A. Lucas, Dr. Robert Ridgway, and Dr. Jonathan Dwight, Jr. Unless
otherwise indicated, the illustrations were taken by the
author. The work of Mr. Walter King Stone in the paint-
ing for the frontispiece and a number of text cuts is
gratefully acknowledged; and for the skilful printing of
many of the photographs my thanks are due to Mr. E. R. Sanborn. To my wife, for constant and valuable help, criticism,
and suggestion in all departments of the book, I render
my sincere appreciation. To take a few dead facts and clothe them with the
living interest which will make them memorable and full
of meaning to any lover of birds, and at the same time
to keep them acceptable in tenor and truth to the most
critical scientist — this has been my aim. Preface Preface x A few chapters of this volume have already appeared
in print in "Outing," "Bird-Lore," and the "New York
Evening Post." C. W. B. CONTENTS CHAPTER PAOB
1. ANCESTORS 1
II. FEATHERS 19
III. THE FRAMEWORK OF THE BIRD 62
IV. THE SKULL 103
V. ORGANS OF NUTRITION 116
VI. THE FOOD OF BIRDS 142
VII. THE BREATH OF A BIRD 1 65
VIII. MUSCLES AND NERVES 188
IX. THE SENSES 203
X. BEAKS AND BILLS 223
XI. HEADS AND NECKS 252
XII. THE BODY OF A BIRD 285
XIII. WINGS 319
XIV. FEET AND LEGS 353
XV. TAILS 398
XVI. THE EGGS OF BIRDS 427
XVII. THE BIRD IN THE EGG 462
APPENDIX — Brief List of Useful Books 483
INDEX 485
xi
39966 39966 ANCESTORS I ITH the exception of Astronomy, the science
which most powerfully dominates our imagina-
tion is Palaeontology, or the study of the life of
bygone ages. Of all things in Nature, the stars symbolize
absolute immensity, their distances stretching out beyond
our utmost calculation. So the revelations of Palaeon-
tology take us far beyond the sciences of life on the earth
to-day, and open vistas of time reaching back more than
five-hundred-fold the duration of the sway of mankind. Fossil bones — philosophically more precious than any
jewels which Mother Earth has yielded — are the only
certain clews to the restoration of the life of past ages,
millions of years before the first being awakened into
human consciousness from the sleep of the animal mind. Until recently, Palaeontology has been popularly con-
sidered one of the dryest and most uninteresting of the
'ologies, but now that the fossil collections in our museums
are being arranged so logically and so interestingly, the
most casual lover of Nature can read as he runs some of The Bird 2 the "poems hidden in the bones." As Professor Huxley
once said, "Palaeontology is simply the biology of the
past, and a fossil animal differs only in this regard from
a stuffed one, that the one has been dead longer than
the other, for ages instead of for days." the other, for ages instead of for days."
A great many more fossil mammals and reptiles have
been discovered than birds, and the reason may perhaps
be conjectured. The bones and bodies of birds were in
former times as now very light, and if death occurred on
the water, the body would float and probably be de-
voured by some aquatic reptile. Then, again, when some
cataclysm of nature or change of climate obliterated
whole herds and even races of terrestrial creatures, the
birds would escape by flight, and when death eventually
came, they would be stricken, not in flocks, but singly
and in widely scattered places as to-day. and in widely scattered places as to-day. For perhaps a million years in the past, birds have
changed scarcely at all, — the bones of this period belong-
ing to the species or at least genera of living birds. ANCESTORS But
in the period known as the Cretaceous, when the gigantic
Dinosaurs flourished and those flying reptile-dragons —
the Pterodactyls— flapped through the air, a few remains
of birds have been found. Some of these are so com-
plete that almost perfect skeletons have been set up,
enabling us vividly to imagine how the bird looked when
swimming through the waters of our globe, or flying
through the air, perhaps four millions of years ago. through the air, perhaps four millions of years ago. The most remarkable peculiarity of these birds was
the possession of teeth. Two of the most well-known
examples are called Ichthyornis and Hesperornis. The Ancestors 3 bones of these birds were discovered by Professor Marsh
imbedded in the rocks of western Kansas, and they are
now preserved in the museum of Yale University. Pro-
fessor Marsh tells us that Hesperornis, the Bird of the IG. 1. — Restored skeleton of Ichthyornis (after Marsh). 1/2 natural size. FIG. 1. — Restored skeleton of Ichthyornis (after Marsh). 1/2 natural size. West, "was a typical aquatic bird, and in habit was
doubtless very similar to the loon, although, flight being
impossible, its life was probably passed entirely upon
the water, except when visiting the shore for the purpose
of breeding. The nearest land at that time was the sue- The Bird 4 cession of low islands which marked the position of the
present Rocky Mountains. In the shallow tropical sea,
extending from this land five hundred miles or more to
the eastward, and to unknown limits north and south,
there was the greatest abundance and variety of fishes,
and these doubtless constituted the main food of the
present species. Hesperornis, as we have seen, was an
admirable diver; while the long neck, with its capabilities
of rapid flexure, and the long slender jaws armed with
sharp recurved teeth, formed together a perfect instru-
ment for the capture and retention of the most agile fish. The lower jaws were united in front only by cartilage,
as in serpents, and had on each side a joint which admitted
of some motion, so the power of swallowing was doubt-
less equal to almost any emergency." less equal to almost any emergency."
Hesperornis had numerous teeth set in grooves like
those of serpents and crocodiles, but in Ichthyornis ('Fish-
bird/ so called because its vertebra? ANCESTORS are biconcave like
those of a fish) the teeth were in separate sockets as in
alligators. The latter bird was not large, being about
the size of a pigeon, and it had well-developed wings. the size of a pigeon, and it had well-developed wings. It is interesting to compare Hesperornis with the
group of penguins, both being highly specialized, although
in ways so different, for an almost wholly aquatic life. Hesperornis swam by strong strokes of its great webbed,
or lobed, toes, its wings dangling uselessly for genera-
tion after generation, until all trace, save a vestigial
humerus, of their bony support disappeared. Penguins,
however, make but little use of their feet in swimming,
only occasionally aiding the tail in steering; but they Ancestors Ancestors literally fly through the water by means of their flipper-
like wings. The large size of the leg and toe bones of Hesperornis
shows that great speed was attainable in the water, FIG. 2. — Lower jaw of Ichthyornis (after Marsh). 4/5 natural sizf. FIG. 2. — Lower jaw of Ichthyornis (after Marsh). 4/5 natural sizf. FIG. 3.— Lower jaw of Alligator. 1/6 natural size. The teeth are set in distinct
sockets both in the extinct bird and in the living reptile. FIG. 3.— Lower jaw of Alligator. 1/6 natural size. The teeth are set in distinct
sockets both in the extinct bird and in the living reptile. while only a single bone remained to show where the
wings of its ancestors were situated. It is doubtful if it
could stand erect upon land, being in this respect more
helpless even than a grebe. Its nest, if it made one,
must have been at the very edge of the shore, from which The Bird 6 it could wriggle or push itself with its powerful toes into
the water. The thought of the untold generations of
birds which must have preceded this toothed, wingless,
feathered being, makes the mind falter at the vast stretches
of time during which evolution has been unceasingly at
work. When we examine the skull of Hesperornis we get a
clew to the reason why this great creature, nearly as large
as a man, succumbed when some slight change in its
environment called for new adjustments in its habits of
life. ANCESTORS Its brain was comparatively smaller than that of
any existing bird; and this absence of brain power im-
plied a total lack of that ingenuity, so prominent in the
crow, which, when man alters the face of the land, changes
its habits, and with increasing wit holds its own against
guns and traps. guns and traps. When Hesperornis passed, it was succeeded by birds
much smaller in size but of greater wit — loons and grebes
—which hold their own even to the present day. —which hold their own even to the present day. When in the depth of the winter, a full hundred miles
from the nearest land, one sees a loon in the path of the
steamer, listens to its weird, maniacal laughter, and sees it
slowly sink downward through the green waters, it truly
seems a hint of the bird-life of long-past ages. seems a hint of the bird-life of long-past ages. We must now pass back, as nearly as can be estimated,
over two millions of years, through the ages when the
Iguanodonts and Megalosaurs lived, long before the first
serpents had evolved and about the time when the first
timid forerunners of the mammals made their appear-
ance,— tiny insect-eating creatures which were fated to Ance 7 remain so long subordinate to the masterful giant reptiles. This was about the middle of the Jurassic period, and in
deposits of this epoch have been found remains of the
very first birds of which we know anything. Two specimens have been discovered and named Archce-
opteryx (ancient-winged-creature). From these two little
stone slabs, one in the British Museum and the other at
Berlin, we know that these birds were about the size of
a crow. Instead of the broad, fan-shaped tail of modern
birds, the tail of the ArcJiceopteryx was a long, jointed
affair like that of a lizard, and was fringed with large
feathers — a pair growing from each of the twenty joints. The wings were not large, and instead of the fingers being
concealed by feathers, there were three entirely free
digits, each armed with a. claw, in front of each wing. The
skin-covered jaws were furnished with teeth, but the feet
and legs were much like those of an ordinary crow. ANCESTORS Taken all in all, this was a most wonderful discovery,
linking birds and reptiles together, and proving beyond
all dispute the fact of their common origin. Perhaps the
most surprising fact was the remarkable development of
the plumage of the wings and tail, showing that perfect
feathers were in existence at least six millions of years
ago. ago. In the rocks deposited in very ancient epochs are
found many footprints which were supposed to be those
of huge birds, but it is more probable that they were
made by certain three-toed reptiles which, like birds,
walked or hopped on two feet. Indeed Nature seems to
have made several abortive attempts to produce bird- The Bird 8 like creatures before she struck the right adjustments. Pterodactyls failed to become birds because they depended
on a broad web of skin, like the wing of a bat, thus miss-
ing the all-necessary feather-ideal; Dinosaurs began at
the wrong end, learning to stand on their hind feet and
to hop, but never the delights of flight. These offshoots
sooner or later were forced to the wall, but Archceop-
teryx seems to have been very near the true line of
descent. But after all, what a meagre record we have of the un-
told myriads of generations of birds which have succeeded
each other through ages past! It is to be hoped that
many more fossils may be discovered, for the hints given
us in the anatomy of birds, and the glimpses of past his-
tory which flash out from the development of the chick
within the egg, — all this evidence is becoming ever more
and more clouded and illegible. Having learned that birds are descended from a rep-
tile-like ancestor, it is interesting to search among living
reptiles for the one which most resembles birds, and we
have no choice but to select the alligator — cold-blooded,
scaly, bound to the earth though he is. A second near
relation is to be found in the group of long-extinct Dino-
saurs. A complete record of past ages would show the
ancestral stems of alligators, Dinosaurs, and birds grad-
ually approaching each other until somewhere, at some
time, they were united in a common stock. * There are one or two reasons for regarding Archceopteryx as merely
the tip of a parallel branch, but one sprouting close to the base of the avian
tree. ANCESTORS But we
must guard against the notion that birds are descended
from any group of living reptiles; which is as fallacious
an idea as that we Americans trace our direct descent from FIG. 4. — Anhceopteryx preserved in the British Museum. The pelvic girdle, le
and tail show best in this specimen. 1/5 natural size. FIG. 4. — Anhceopteryx preserved in the British Museum. The pelvic girdle, leg
and tail show best in this specimen. 1/5 natural size. The Bird i the Chinese, or that mankind is descended from the chim-
panzee or gorilla. For the purpose of making more clear and interesting
the ways in which birds have become especially adapted
to their surroundings and needs, we may consider Archce-
opteryx as resembling closely the typical original bird-
type from which all others have at least indirectly evolved ;
and thus having obtained a definitely fixed starting-point,
we may consider how some of the more representative
birds of the present day came to acquire their widely
differing structure and characteristics.* differing structure and characteristics.*
The tree of evolution of reptiles may be compared to
a growth where several great trunks spring from the
ground close together, towering up separately but equally
high; the topmost twigs of which are represented by the
living species of serpents, turtles, lizards, and crocodiles
respectively. A very different arboreal structure is pre-
sented in the genealogical tree of the Class of birds. Here,
from a short trunk, we have many radiating branches,
widely spreading and with thickly massed twigs, confu-
sedly intermingled ; so slight are the divergences between
adjoining groups and so equally do almost all share be-
tween them various reptilian characteristics. It is not necessary to concern ourselves now with the
processes of evolution, especially as scientists are still in
doubt as to the exact methods. Let us read our Darwin,
and hope for another, philosophically as great, to com- FIG. 5. — Archoeopteryx preserved in the Berlin Museum. The skull, vertebrae,
forelimbs and flight-feathers are remarkably distinct. 1/3 natural size. FIG. 5. — Archoeopteryx preserved in the Berlin Museum. The skull, vertebrae,
forelimbs and flight-feathers are remarkably distinct. 1/3 natural size. ize. The Bird 1 plete the work, meanwhile adding our own mite of truth-
ful observation to swell the whole, and help prepare the
way for this other. ANCESTORS For even Darwin's theory of evolu-
tion was but the consummation of theories of former
years and centuries, — beginning with Thales and Anaxi-
mander, in the days of early Grecian civilization: in-
deed Aristotle, coming but two hundred years later, is
the only name in the history of zoology worthy of a
place with that of Darwin. From the fragmentary evidence afforded by Archceop-
teryx we may conclude that this Bird of Old had a short,
blunt, skinny bill of moderate size, furnished with teeth
which would enable the owner to feed upon Jurassic
berries and fruit, or more probably a carnivorous diet of
lizards and insects. Its wings were weak, hinting that
it was a flutterer rather than a true flyer, perhaps only
scaling like a flying squirrel from the summit of one
tree to the base of the next. Even this would give it
an immense advantage over its terrestrial and arboreal
non-flying enemies. The three free fingers on each wing
would allow it to climb easily, to pry into crevices for
insects, or to draw a berry-laden branch close to its
bill. Doubtless it frequently walked or ran on all fours, the
more probably from its weak-loined condition, — the bones
of the thigh-girdle not being fused together as in modern
birds. Its tail has already been mentioned — a long
double-feathered appendage, composed of a score of
little vertebrae jointed together, — as we will later see
the true forerunner of the modern fan-like tails. Its FIG. 6. — Restoration of Archceopteryx (adapted from Smit). Notice the teeth,
three fingers, and lizard-like tail. 13 FIG. 6. — Restoration of Archceopteryx (adapted from Smit). Notice the teeth,
three fingers, and lizard-like tail. The Bird H feet and legs were little different from those of perching
birds of to-day, with strong toes well adapted to cling
to a branch. Finally, from a cast of the brain, which
fortunately was found with one of the fossils, we know
that, although small, it was that of a true quick-witted
bird. As yet science has no more to tell us. bird. As yet science has no more to tell us. ANCESTORS Our fancy may add an archaic attempt at song — a
lizard's croak touched with the first harmony, which
was to echo through all the ages to follow; we may also
imagine, if we will, leathery eggs deposited in a rotten
knot-hole of a Jurassic conifer. In both islands of New Zealand well-preserved remains
of giant birds have been discovered, to which has been
given the name of moas. One species must have reached
a height of ten or eleven feet, which would make it
tower above the largest living ostrich. They were, in
fact, not unrelated to these latter birds and, like them,
were flightless (in some cases absolutely wingless), and
they had great massive feet and legs. Native legends
among the Maoris hint that these birds were in existence
during the last few centuries before the coming of the
white men. In South America also, giant birds lived in ages past. One, the Phororhacos, stood seven to twelve feet in
height, with a head and beak like that of a gigantic eagle. Unlike all eagles, however, this bird could not fly and
doubtless ran down its prey, as a chicken runs down a
grasshopper. grasshopper. It is an interesting fact that in South America there
lives to-day a bird known as the Seriema, which is prob- Ance 1 5 ably at least an indirect descendant of the Phororhacos. The Seriema defies exact classification, sharing characters
of cranes, bustards, and eagles. Its beak and inner
claw are like those of a bird of prey, while in form of
body, and in the other claws of the toes, and in the legs
it is crane-like. One of these birds which I have ob-
served for years in captivity is as gentle and as fearless
as a bird can be. It will chase insects and field-mice
outdoors in the Zoological Park, and will occasionally
stalk solemnly into my office and, coming close to my
desk, watch me closely. It has most beautiful gray-
blue eyes, with long eyelashes (Fig. 199), and if the
birds of past ages were as comely and as lovable as this
interesting species, I regret that only their fossil bones
are left to us. As the Seriema runs down and kills a
mouse, so the giant Phororhacos, doubtless, overtook
and slew creatures as large as a deer. Its skull (Fig. ANCESTORS 7)
is drawn to the same scale as that of the living Seriema
(Fig. 8). The evolution which has gone on since these epochs
of old, bringing into being the wonderfully varied forms
of penguin, ostrich, albatross, peacock, and humming-
bird, may be summed up in two words which it is well
to know and remember, — Adaptive Radiation. This is
the spreading out or radiating of bird-forms descended
from the ancient stem, into all parts of the earth, each
coming into contact with a particular environment, to
adjust itself to which, its various organs and parts exer-
cise different functions, until the friction of the "struggle
for existence" has moulded each to its particular niche. The Bird i6 If its lines lie in happy places, its race is established,
and it pursues and flees, it fights and plays, it sings with
joy or pants with fear, and Evolution marks another
success in its inexorable movement onward and upward,
— a new species is born! — a new species is born! Earth has few secrets from the birds. With wings
and legs there is hardly a spot to which they cannot and
indeed have not penetrated. Some find food and con-
tentment in the desolate wastes of the far North; others
spend almost all of their life on or above the sea far from FIG. 7. — Skull of Phororhacos, drawn to scale with Fig. 8. 1/6 natural size. IG. 7. — Skull of Phororhacos, drawn to scale with Fig. 8. 1/6 natural size. land; thousands revel in the luxuriance of reeking trop-
ical jungles; a lesser number are as perfectly suited to
the blazing dust of the desert; and there are birds which
burrow deep into the very earth itself. Day and night;
heat and cold; water, earth, and air, have all been con-
quered by the thirteen or fourteen thousand species of
birds which share the earth with us at the present day. These brethren of ours, whose clans have so bravely
conquered the dangers of millions of years, and at last
have gained a foremost rank in the scale of living crea- Ance 1 7 tures, now find themselves face to face with the culmi-
nating effort of Nature, — Mankind. They cannot escape 8. — Seriema, a living descendant of Phororachos , with characters
Bustards, and Eagles. 1/6 natural size. FEATHERS ANY definitions of the Class of birds have been
given, but all fall short in some particular, or
are weak in having exceptions. FEATHERED is
the one word which always holds true. All birds have
feathers, and nowhere else in the world are similar struc-
tures found. A feather, like an egg, is perfect in its
adaptation to the bird's requirements, and also, like the
egg, its structure is rather complicated. A ANCESTORS m us, though the least among them laughs to FIG. 8. — Seriema, a living descendant of Phororachos , with characters of Cranes,
Bustards, and Eagles. 1/6 natural size. from us, though the least among them laughs to scorn
our efforts at following through the air. Yet all must from us, though the least among them laughs to scorn
our efforts at following through the air. Yet all must The Bird 1 8 return sooner or later to earth for rest and food, and thus
all are at our mercy. Let us beware of needlessly destroying even one of
the lives — so sublimely crowning the ages upon ages of
evolving; and let us put forth all our efforts to save a
threatened species from extinction; to give hearty aid
to the last few individuals pitifully struggling to avoid
absolute annihilation. The beauty and genius of a work of art may be recon-
ceived, though its first material expression be destroyed;
a vanished harmony may yet again inspire the composer;
but when the last individual of a race of living beings
breathes no more, another heaven and another earth
must pass before such a one can be again. Structure and Development First let us look at the skin itself in which the feathers
grow. To skin a bird is an easy matter, for the skin,
or integument as it is called, is very slightly attached
to the muscles underneath. The skin of a dove is almost
like tissue-paper, and tears so easily that it is a marvel
how the hundreds of feathers find a sufficiently strong
attachment. Thin as is this skin, it is made up of three
separate layers, but in order to make our feather-study
enjoyable by not overburdening it with too many details,
we will consider only the two more important layers of
the skin — a deeper one, the dermis, and an outer, more
horny covering, the epidermis. The Bird 20 A list of all the structures of animals which are prod-
ucts of the outer layer alone would be a long and sur-
prising one, and we would be very ready to grant the
importance of skin. Such an enumeration would include
all claws and talons, nails and teeth, the rattles of a
snake, spurs, hairs, the scales of fishes and reptiles, \\
FIG. 9. — Tarpon-scale, shark-tooth, and peacock-feather; showing diversity of
structure derived from the skin alone. 1/2 natural size. FIG. 9. — Tarpon-scale, shark-tooth, and peacock-feather; showing diversity of
structure derived from the skin alone. 1/2 natural size. spines, whalebone, beaks and feathers. Even the horn
of a rhinocerus is only a solid mass of agglutinated
hairs, while as the antithesis to this may be mentioned
all down and feathers : the tiniest fluff from a humming-
bird to the great pinion of a condor. spines, whalebone, beaks and feathers. Even the horn
of a rhinocerus is only a solid mass of agglutinated
hairs, while as the antithesis to this may be mentioned
all down and feathers : the tiniest fluff from a humming-
bird to the great pinion of a condor. If we examine a newly hatched dove or sparrow,
the little, ugly, sprawling creature, at first glance, seems Feathers 21 to be entirely naked; but a closer inspection shows
scanty tufts of down scattered irregularly over the body. This, like the set of milk-teeth in mammals, is useful
only for a time, and is later pushed out by the second
or true plumage. Even more numerous than the down- FIG. 10. — Brown Pelican nestlings, showing feather papillae on body and wings. About 1/4 natural size. FIG. 10. — Brown Pelican nestlings, showing feather papillae on body and wings. About 1/4 natural size. tufts are little pimples or dots, many hundreds of which
cover certain parts of the skin. Each of these will event-
ually give rise to a perfect feather — quill, vane, barbs,
and all. tufts are little pimples or dots, many hundreds of which
cover certain parts of the skin. Each of these will event-
ually give rise to a perfect feather — quill, vane, barbs,
and all. The under layer of skin, or dermis, is very thin in
birds, much more so than in reptiles and other animals. The Bird The Bird 22 The first intimation of the appearance of a feather, or
of down, is shown by a thickened group, or pimple, of
cells in this under layer of skin, which grows and presses
upward toward the outer layer — the epidermis. This is
exactly the way in which the scales of fishes and reptiles
begin to form; and if, at this stage, the tiny projection
should flatten out, the shining scale of a carp, the armor FIG. 11. — Sprouting feathers of a 12-day embryo chick. Magnified 25 diameters. FIG. 11. — Sprouting feathers of a 12-day embryo chick. Magnified 25 diameters. of an alligator, or the cobble-scale of an iguana lizard
might result. Indeed, in the feathers of a penguin we
find transition stages of flat, almost unsplit feather-
scales; while on the legs and feet of birds are reptile-like
scales. The evolution of scales, hair, and feathers is a most
interesting problem, most of the details of which are
beyond the scope of this work. Suffice it to say that Feat 23 in sharks, which are among the most primitive forms of
fishes, the skin is covered with tiny denticles or spines,
which consist of enamel and dentine, and which rest on
small bony plates. This form of scale is the most ancient
known, and the hint of teeth which the description con-
veys is not misleading; for we find that in some of these
voracious fishes the spines in the skin become enlarged
near the edge of the mouth, merging imperceptibly into
the rows of cruel teeth which, to a certain extent, are
homologous with the teeth of all higher animals. In
other fishes the denticles become flattened scales, and
many of these fish have teeth of corresponding plate-
like form. So it is interesting to know that the scales of
fishes and reptiles, the feathers of birds, and the teeth
of animals have all evolved from skin structures which
at an early stage of growth bear considerable resemblance
to each other. But, in our young bird, the slender finger of cells
which reaches upward, and whose base at the same time
sinks deeply into the dermis, does not broaden out, but
splits longitudinally into a number of folds, which grad-
ually dry apart and harden into the slender, silky fila-
ments which we know collectively as down. The Bird At the base of, and in fact attached to, the little pro-
jection which gives rise to the nestling down is a small
circular body of cells, which grows but little while the
down plumage is serving its use; but when the bird is
ready for a coat of true feathers this lower cellular mass
begins to grow upward into a second finger, or column,
of cells, pushing the base of the down feather out of its The Bird 2 socket. This growth continuing, the down is lifted clear
of the skin, being supported on the new structure, and FIG. 12. — Early stages in the development of a down feather, showing close
resemblance to scale of fish or reptile. FIG. 12. — Early stages in the development of a down feather, showing close
resemblance to scale of fish or reptile. FIG. 12a. — Later stages of Fig. 12, showing the first splitting up of the
feather pulp. FIG. 12a. — Later stages of Fig. 12, showing the first splitting up of the
feather pulp. soon brushed off and lost. Thus, little by little, in
shreds and tatters, the baby plumage is shed and replaced soon brushed off and lost. Thus, little by little, in
shreds and tatters, the baby plumage is shed and replaced Feat 25 by true feathers, which overlap, protecting the body from
heat and cold, dust and rain. heat and cold, dust and rain. FIG. 126. — Last stages in the formation of a down feather, showing the plumes
well above the surface of the skin, as in a newly hatched chick. All greatly
enlarged. — Last stages in the formation of a down feather, showing the
above the surface of the skin, as in a newly hatched chick. All
ged. the time of the first moult, this successi
s can be observed in almost any young bird, FIG. 126. — Last stages in the formation of a down feather, showing the plumes
well above the surface of the skin, as in a newly hatched chick. All greatly
enlarged. At the time of the first moult, this succession of
feathers can be observed in almost any young bird, being
more noticeable in large species, which have very thick The Bird 2 or lengthened down, as gulls and ducks. The Bird A Red-winged
Blackbird, or for that matter almost any passerine nest-
ling, looks very odd when it rises up in the nest, gaping for
food; the long gray streamers of down waving like an
aureole around its head. In some water-birds this nest-
ling down retains its usefulness for nearly two months. FIG. 13. — Feather from the head of a young Bobolink, with down still attached
to its tip. Twice natural size. FIG. 13. — Feather from the head of a young Bobolink, with down still attached
to its tip. Twice natural size. The feathers which replace the down are, when they
first appear above the skin, rolled tightly and bound up
in the thin tissue of the horny sheaths, so that they
resemble a bundle of withes wrapped together in a cloth. In many young birds the feathers remain in this condi-
tion until they are nearly full grown, and a young cuckoo Feat 27 or kingfisher is a curious-looking object, most of the
bird's body seeming to be tiled with small, bluish sticks. FIG. 14. — Tip of feather from the crown of a young Song Sparrow, showing
connection with down. Magnified 25 diameters. FIG. 14. — Tip of feather from the crown of a young Song Sparrow, showing
connection with down. Magnified 25 diameters. FIG. 15. — Duck Hawk moulting into juvenal plumage, with the natal down coming
away in shreds and tatters. 1/4 natural size. FIG. 15. — Duck Hawk moulting into juvenal plumage, with the natal down coming
away in shreds and tatters. 1/4 natural size. When the folds of the developing feathers are sufficiently
dry, they burst their sheaths and rapidly spread out. The Bird 2 The appearance of a young kingfisher or heron may be
completely changed within a few hours time, so quickly
and simultaneously does the first suit of feathers unroll. The condition of young birds when hatched varies
greatly in birds of different groups. Nestlings are, in
many ways, like human babies, and there are as many
differences in the one class as there are in the other, FIG. 16. — Growth of an Ostrich feather from sheath to plume. 1/5 natural size. FIG. 16. — Growth of an Ostrich feather from sheath to plume. 1/5 natural size. between those from different countries, only Nature does
for the little birds what parents do for the babies. The Bird We see American babies wrapped in furs and blankets,
wheeled in carriages, and rocked to sleep; while a tiny
savage is strapped tightly to its mother's back, and as
soon as possible allowed to run where it pleases, find its
own toys and -develop its little muscles, gaining a degree
of health and strength which many a civilized child would
envy. So with birds, the highest — such as crows and
thrushes — are hatched almost naked and must be warmed Feat 29 and cuddled and fed for many weeks, before they learn
to take care of themselves; while birds lower in the
scale — as our quail — are born covered thickly with
down and with wings nearly feathered, and in a few days
can fly and find their own food. So a bird naked at birth is very helpless, one covered
with down is more capable of taking care of itself, while FIG. 17. — Nestling Kingfisher with feathers still in their sheaths. 2/3 natural size. FIG. 17. — Nestling Kingfisher with feathers still in their sheaths. 2/3 natural size. the few which are completely feathered when hatched
may be said to have no chickhood except in the egg. In the Crested Screamer (Fig. 264) the down-like
character of the body-feathers of the adult birds may be
a hint of the plumage of very ancient types of birds such
as Archceopteryx. Now we are ready to begin our study of the perfect
feather itself, and we will, for once, have to disregard
our rule of starting with the simpler form — the scale of The Bird 3°
a reptile — and working up to a feather; for, if we except
the down, there seems to be no connecting link left. FIG. 18. — Young Brown Pelicans; hatched naked and helpless (altricial). 1/4 natural size. FIG. 18. — Young Brown Pelicans; hatched naked and helpless (altricial). 1/4 natural size. FIG. 19. — Young Red Jungle Fowl one day old; hatched covered with down and
able within a few hours to help itself (precocial). Almost natural size. FIG. 19. — Young Red Jungle Fowl one day old; hatched covered with down and
able within a few hours to help itself (precocial). Almost natural size. The Bird Although that old, old fossil bird Archceopteryx
still retained reptile-like teeth, fingers and tail, it had Feathers feathers which were apparently as perfect as any we may
examine to-day. When some form of scale had once
changed so that it was of use in flight, the hollow elastic
vane took first place at once, and all intermediate stages,
which perhaps had been acquired merely for warmth,
went to the wall. A creature could have flight if pro-
vided with perfect feathers, or it could retain its scales
and find existence possible along the old reptilian planes
of life, but no awkward scale-flutterer could long be
tolerated. All through the evidences of evolution we
find instances like this, — a change for the better beginning
slowly, through many channels, then the one best suited
forging ahead with inconceivable swiftness, and crushing
out all other less adapted structures. Hence the rarity
of "missing links." of "missing links."
Feathers are certainly among the most beautiful
objects in Nature; and when we learn a little about their
structure, they will be still more interesting. No matter
how closely we may examine them, with hand-lens or
microscope, their beauty and perfection of structure only
increase. If we study a feather, say from the wing of a
pigeon, we see that its whole structure is subservient to
two characteristics — lightness and strength. What won-
derful elasticity it has! We can bend the tip so that it
touches the base and it will spring back into shape with-
out breaking. If we look closely, we will see that each feather is
composite — feathers within feathers. The quill gives off
two rows of what are called barbs which together form
the vane of the feather; each of these barbs has two The Bird 3 rows of barbules, and these give rise to a series of curved
hooks, known as barbicels, which work into opposite
series of grooves, so tightly that air cannot force its way
through the feather. When the wings are pressed down-
ward, the phenomenon flight is made possible by the
accumulated resistance which the flight-feathers offer to
the air. At the lower end of our pigeon's feather, bar-
bicels are present only near the quill. Therefore the FIG. 20. — Two interlocked barbs from the vane of a Condor's wing-feather, show-
ing barbules and barbicels. Magnified 25 diameters. FIG. 20. The Bird — Two interlocked barbs from the vane of a Condor's wing-feather, show-
ing barbules and barbicels. Magnified 25 diameters. tips of the barbs are loose and fluffy, unconnected and
useless for flight. This is the condition in all down and
in the feathers of the ostrich and cassowary. We might
naturally think that feathers stiffened by so many close
rows of interlocking barbicels would be useful in many
ways beside flight. But fluffy feathers are evidently just
as efficient in keeping warmth in and rain out as the
other kind; so Nature, economical to the most micro-
scopic degree, has lessened the number of, or has never
provided, barbules and barbicels wherever a feather is
not needed for flight or steering. tips of the barbs are loose and fluffy, unconnected and
useless for flight. This is the condition in all down and
in the feathers of the ostrich and cassowary. We might
naturally think that feathers stiffened by so many close
rows of interlocking barbicels would be useful in many
ways beside flight. But fluffy feathers are evidently just
as efficient in keeping warmth in and rain out as the
other kind; so Nature, economical to the most micro-
scopic degree, has lessened the number of, or has never
provided, barbules and barbicels wherever a feather is
not needed for flight or steering. Feathers Feathers 33 The two lines of barbs which grow out on each side
of the quill are very elastic and so intimately hooked to
each other that they will bend some distance before sepa-
rating. If we ever tried to force our way through a FIG. 21. — Model showing interlocking barbules and barbicels of feather,
greatly enlarged. FIG. 21. — Model showing interlocking barbules and barbicels of feather,
greatly enlarged. bramble of sweet-brier or blackberry- vines, we can more
readily appreciate how these barbs and the interlocking
barbules clutch each other. The thorns in the bramble
catch our clothes and, when we move, the elasticity of the
long stems tends to make them hold the tighter. We notice that one line of barbs — that along the
inner curve of the quill — is much longer than that on the
outer curve and we might think the air would force this The Bird 3 upward and escape beyond the edge. So it would, if it
were not for the arrangement of the feathers on the wing,
which overlap like the tiles on a roof, each vane over-
lying and holding down the long barbs of the feather
in front, while, above and below, other shorter feathers
help to bind the whole tightly, thus enabling the bird at
every stroke to whip a wingful of air downward and
backward. A feather and its parts, like all the rest of the bird, is
composed of cells — empty and hollow ones in this in-
stance, as we can easily see for ourselves by placing a
barb from a pigeon's feather in a drop of water and
looking at it under a low-power magnify ing-lens. The
network of horny cells is very plain. It is a simple matter to say that a feather consists of
quill, barb, barbules, etc., but to appreciate the wonder-
ful complexity of this structure let us make a little cal-
culation. Suppose we have a wing-feather from a com-
mon pigeon with a vane about six inches long. If we
have patience enough to count the barbs on one side of
the quill, we will find there are about six hundred of
them. So the vane of the entire feather has twelve
hundred of these little side featherlets. FIG. 22. — Feathers illustrating conditions where barbicels are unnecessary and
are hence reduced or entirely lost, causing downiness. 3/5 natural size.
(a) Primary of Pigeon — an important flight-feather; hence possessing a stiff
vane. (6) Under wing-covert of a Great Blue Heron; downy portion was over-
lapped by the adjoining feather, (c) Wing-covert of Owl; the downy edge makes
possible the all-important noiseless flight of this bird. (d) Feather of Ostrich;
the power of flight being lost, the feathers are downy throughout the entire vane.
35 Feathers One of these,
from a narrow part of the vane, will show under the micro-
scope about two hundred and seventy-five pairs of bar-
bules, which multiplied by the number of barbs on that
side amounts to three hundred and thirty thousand. Making a very low estimate of the whole vane, we have
nine hundred and ninety thousand separate barbules on
this one feather, and when we think of the innumerable . I l*L'
FIG. 22. — Feathers illustrating conditions where barbicels are unnecessary and
are hence reduced or entirely lost, causing downiness. 3/5 natural size. (a) Primary of Pigeon — an important flight-feather; hence possessing a stiff
vane. (6) Under wing-covert of a Great Blue Heron; downy portion was over-
lapped by the adjoining feather, (c) Wing-covert of Owl; the downy edge makes tire
35 tire
35 The Bird finer booklets, and then the number of feathers on the
pigeon's body, we can echo the exclamation of Solomon:
"The way of an eagle in the air" is "too wonderful for me!"
Another beautiful adaptation to flight is seen in our FIG. 23. — Feathers of Condor and Emeu. The aftershaft in the former is reduced
to a downy filament at the base of the vane; in the latter it equals the feather
itself in size. FIG. 23. — Feathers of Condor and Emeu. The aftershaft in the former is reduced
to a downy filament at the base of the vane; in the latter it equals the feather
itself in size. feather. The upper part of the wing must of course be
perfectly level, with no projections to catch the air and
retard motion. So, on the upper side of the feather, we
notice that the lines of barbs spring out flush with the
flattened quill-top, while below, the shaft projects promi-
nently from the vane. The obliquely forward direction
in which the barbs grow, the change in shape of the feather. The upper part of the wing must of course be
perfectly level, with no projections to catch the air and
retard motion. So, on the upper side of the feather, we
notice that the lines of barbs spring out flush with the
flattened quill-top, while below, the shaft projects promi-
nently from the vane. Feathers The obliquely forward direction
in which the barbs grow, the change in shape of the Feat 37 quill — round where the body or body-feathers conceal
it, square where it supports the vane, — and many other
niceties which we can each detect for ourselves, show
how exquisitely exact is the adaptation of a feather to
its uses. FIG. 24. — Powder-down patch on the breast of a live Great White Heron. 2/3 natural size. FIG. 24. — Powder-down patch on the breast of a live Great White Heron. 2/3 natural size. Growing from the under side of the quill, at the be-
ginning of the vane, is a tiny feather known as the after-
shaft. In an ordinary down-feather of a young bird this
is of considerable size, but it is either small or entirely
absent in an ordinary feather. It reaches its greatest
development in the emeu and the cassowary, where it is The Bird 3 8 as long and as perfect as the main feather. The origin
and use of this feather-double is not known. and use of this feather-double is not known. Parrots, herons, and some other birds have a most
convenient arrangement — a kind of automatic clothes-
cleaner and valet combined. Concealed by the long body-
plumage are several dense patches of down-feathers which
grow quite rapildy, but instead of constantly increasing
in length, the tips break up into a fine, white, greasy
powder. This works its way through the entire plumage,
and is doubtless of use in keeping the feathers in good
condition and the body dry. Most of the birds possess-
ing this convenience are comparatively free from lice, so
this natural dressing may be as unpleasant to these ver-
min as camphor-balls are to clothes-moths. min as camphor-balls are to clothes-moths. The forms and textures of feathers are innumerable,
and the uses to which they are put, more than we would
ever imagine, but these will be spoken of under the chap-
ters treating of the different parts of the body where they
are found. Arrangement In examining a nestling we will notice that the feather-
dots are not scattered at random over the surface of the
skin, but grow in lines and tracts, whose limits are very
sharply defined. In an adult bird, say an English Spar-
row, this is even more noticeable. If we part the feathers
on the centre of the breast, a broad, bare area is seen,
with only a thin scattering of soft downy feathers. Under
the wings are other naked spaces, and several more are
on other parts of the body. The most ancient birds were Feat 39 probably covered uniformly with scale-feathers, but as
these increased in length there was less need for an un-
broken covering, the feathers of one portion overlapping
and protecting the surrounding parts, and besides, for
ease in active motions, bare patches of skin were required. It has been found that the arrangement of the feathers
on a bird's body varies in different groups, and, such
variation being rather characteristic of these larger divi- FIG. 25. — Nestling Crow, showing feathered and infeathered portions of the body
(pterylae and apteria). 1/2 natural size. FIG. 25. — Nestling Crow, showing feathered and infeathered portions of the body
(pterylae and apteria). 1/2 natural size. sions, pterylosis — as it is called — is of some importance
in classification. Penguins only, of existing birds, have
feathers growing uniformly on all parts of the body. In
the ostrich, which has given up flight and taken to run-
ning, the body feathers have grown over almost all the
bare spaces which existed in its flying ancestors. There
are two marked exceptions due to the present habits of
these birds. Like the camel, when resting, these giant
birds lean upon their breasts. This portion of the body sions, pterylosis — as it is called — is of some importance
in classification. Penguins only, of existing birds, have
feathers growing uniformly on all parts of the body. In
the ostrich, which has given up flight and taken to run-
ning, the body feathers have grown over almost all the
bare spaces which existed in its flying ancestors. There
are two marked exceptions due to the present habits of
these birds. Like the camel, when resting, these giant
birds lean upon their breasts. Arrangement This portion of the body The Bird 4 is provided with a thick, callous pad, which, by constant
use, is thus kept bare of feathers. In addition, the under
sides of the degenerate wings are also free of plumage,
owing no doubt to the continual close application of
these organs to the sides of the body. The other bare
areas are almost obliterated, but the legs are bare, thus
allowing perfect freedom in action. Some birds, such as vultures and cassowaries, have lost
all feathers on the head and neck, or other portions of
the body, from various causes, as for cleanliness, or, in
some cases, probably for ornament. This will be spoken
of more in detail in a later chapter. Moult The waste of internal tissues and organs in animals
is repaired by means of the blood which brings them
fresh material and carries away worn-out cells, as it
traverses arteries and veins. Entire parts, as the tails of
tadpoles, may even be absorbed; but, in general, skin
structures when old and worn out are cast off and renewed
from the lower, or derm, layer. This takes place in various
ways. The skin, even to the covering of the eyeballs,
may come off entire, as is the case among snakes, or por-
tions peel off and tear away, as in lizards. Warm-blooded
animals also shed, or cast, their outside covering; mam-
mals shedding their coats of hair, and birds their feathers. In the latter class this process is called moulting. The nestling down and the feather which replaces it
can hardly be considered as separate structures, as the Feat 41 same channel perforates both and the nutriment pith
which supplies the down traverses the hollow quill of
the succeeding feather. A bird's swaddling-clothes and
his first full dress are cut from the same piece. But when
these perfect feathers reach full size, the aperture at the
base closes, all blood-supply is cut off, and the feather at
the commencement of its usefulness becomes a dead
thing. There is no vital connection between the feathers
of all the following moults. Each is separate, the papilla
or feather-cells reawakening to new activity every time
the process occurs. So when a bird's wing is clipped, no
pain is felt, any more than when a person's hair is cut. Such feathers are of course not renewed until the succeed-
ing moult. If a feather in a living bird be pulled out,
it will be replaced immediately by another, and this will
be repeated as often as the feather is removed. In cassowaries, each moult is advertised by dangling
streamers of the old plumage still attached to the tips
of the incoming feathers, but this connection is not a
living one, the adult feathers being as lifeless as those of
other birds. As powerful savages often exhibit very
childlike traits, so these great birds are absurdly marked
with what, in other species, are sure signs of recent chick-
hood. The changing of plumage of the Brown Pelican is well
shown by the illustrations. The naked young (Fig. 18)
become covered with papillae (Fig. Moult 10) which soon burst
into a coating of the softest white down (Fig. 36) ; this
in turn gives place to the juvenile plumage of gray, the
features of the wings and shoulders appearing first (Fig. The Bird 4 37). This is also the winter plumage of the adult birds,
both sexes moulting alike into the rich-hued breeding
plumage (Fig. 38) of yellow, chocolate, and silver-gray. plumage (Fig. 38) of yellow, chocolate, and silver-gray. The feathers of the entire bird are moulted or fall out
naturally at least once a year, and in some cases twice
or even three times. If we were asked at what season the FIG. 26. — Flight-feathers of Chimney Swift clogged with soot, showing necessity
for moulting. FIG. 26. — Flight-feathers of Chimney Swift clogged with soot, showing necessity
for moulting. principal annual moult would be most likely to occur,
the fall of the year would suggest itself, and such is the
case, for a number of good reasons. First, the hardest work which birds have to do, hatch-
ing and caring for their young, has, at this season of the
year, just been accomplished, and has doubtless told
heavily on their plumage. Breast-feathers are worn thin,
tails are badly frayed, and wing-pinions are broken and
ragged. Two alternatives confront birds at this period. Those species which are to take their migratory flight Feat 43 over hundreds of miles of land and water must have
perfect wings and rudders to carry them safely, against
contrary winds and sudden accidents. Others which are
contented with the food found near their homes, and
elect (by the laws of their kind) to remain, must be pre-
pared to withstand the blasts of winter. Their plumage
must be abundant and thick to keep out the cold and
snow, and to enable them to bury their tender eyes and
feet in its warm mass. Otherwise the tiny round fluffs
huddled close to the trunks in the evergreens would drop
stiffened to the ground during some long winter night. So a renewal of plumage in the fall is most necessary to
the life of birds. A baby robin, secure from most enemies in his nest,
with parents to supply his every want, acquires his wing-
quills only when his nestling down is shed. Moult He is care-
fully watched and tended during his first flights, and
takes such good care of these flight-feathers that they
serve to carry him to his winter home far to the south-
ward. But a brood of a dozen or more little Bob-whites
whose wing-feathers sprout with the most marvellous
rapidity, from the moment the birds tumble out of their
white shells, would fare ill indeed if they had to trust to
these nursery quills all the first winter, with hungry
foxes sniffing for their scent, and more-to-be-dreaded owls
shadowing their trembling covey. Nature has come to
their aid, and when they have fairly worn out their wings
in the first awkward attempts at flight, new feathers
come in, and this succession of quills keeps them in fine
flying condition until full grown. Indeed so solicitous is The Bird 4 Mother Nature about the ground-nesters that she puts
strength and vigor into the coverts, or upper feathers on
the little wings; so that these shoot forth with an energy
far beyond what is usual, for a time lending their aid in
flight, although they are not true primaries. Later they FIG, 27. — Iridescent feather from the breast of a Rufous Humming-bird, showing
wearing off of the tips of the barbs, caused perhaps by rubbing against the
petals of flowers. Magnified 25 diameters. FIG, 27. — Iridescent feather from the breast of a Rufous Humming-bird, showing
wearing off of the tips of the barbs, caused perhaps by rubbing against the
petals of flowers. Magnified 25 diameters. are far outgrown by the flight primaries, and then func-
tion only as protectors of these more important feathers. The extreme in this precocious development of chicks
is found in those strange Australian birds, the mound-
builders, which are left from the first to shift for them-
selves; even the duties of incubation being shirked by
the parents. This necessitates a perfect ability on the Feat 45 part of the young birds to take care of themselves as
soon as hatched. They pass the entire first moult within
the egg itself, and are covered with perfect feathers and
fully developed flight-quills when they emerge from the
shell. Moult A wild duckling, although provided with a thick
waterproof coat of down, has, like the robin, to wait a
long time for his flight-feathers; but his aquatic habits
and powers of diving make the dangers to which he is
exposed far less than is the case with the young Bob-
white. The causes of wear and disablement to feathers would
make a long list if we but knew them all. As one instance
take the wings of a Chimney Swift after she has reared
her brood in the depths of some blackened chimney, or
even a lightning-struck hollow tree. Her primaries are
so matted and clogged with balls of soot that she would
often find the migratory flight difficult indeed, were the
feathers not replaced by new ones. When birds return from the South, and when a hint
of spring warns winter residents to cease their roving,
they prepare to develop all the advantages which may
in any way aid them in securing a mate. Some indus-
triously practise dance-steps, others flight-evolutions, a
larger number rehearse their songs under their breath,
while still others passively await the development of
plumes, gorgets, spots and splashes of colour which, if
the feathers come out large and brilliant, may stand them
in as good stead in their wooing as any song or antic. Thus we find a class of birds which have a partial or com-
plete moult in the spring. These feathers may last all The Bird 4 summer, or may drop out as soon as begins the hard
work of building the nest or feeding the young, with which
labor they might interfere. To return for a moment to the fall moult. If a spar-
row or lark should shed all of its large wing-feathers
simultaneously, it would have slight hope of ever living
long enough for new ones to grow out again. If such
defenceless birds were compelled to hop helplessly along FIG. 28. — Wings of English Sparrow, showing two feathers of each wing being
moulted simultaneously. FIG. 28. — Wings of English Sparrow, showing two feathers of each wing being
moulted simultaneously. the ground, weasels and cats would be able to catch
hundreds of them without effort. This is avoided in all
land birds by the moulting of only a pair of primaries,
as the large flight-feathers are called, at a time, one from
each wing. Moult This process usually starts with the pair
farthest from the front of the wing, and the second pair
does not fall out until the first pair of new feathers is
nearly of full size. Thus all danger of a crippled flight
is avoided. the ground, weasels and cats would be able to catch
hundreds of them without effort. This is avoided in all
land birds by the moulting of only a pair of primaries,
as the large flight-feathers are called, at a time, one from
each wing. This process usually starts with the pair
farthest from the front of the wing, and the second pair
does not fall out until the first pair of new feathers is
nearly of full size. Thus all danger of a crippled flight
is avoided. One of the most interesting phases of Nature is the
way she provides for exceptions to what we are pleased
to call her laws. Some birds, unlike those mentioned Feathers above, shed every primary in their wings at once, so that
their angular stump-feathered wings are perfectly useless
for flight. In this class are many water birds — ducks,
geese, flamingoes, snake-birds and others. Just before this
wholesale moulting occurs, a flock of wild ducks will FIG. 29. — Wing of adult Mallard Duck, with the new set of flight-feathers
just appearing. FIG. 29. — Wing of adult Mallard Duck, with the new set of flight-feathers
just appearing. make their way, by an unfailing instinct, to some large
body of water where they can swim and dive in safety
and, if need be, never come within reach of enemies on the
shore until the new feathers are strong enough to bear
them up. Associated with this temporary disablement is another
provision for the safety of certain birds of this class. Our
common Mallard Duck, for example, is sometimes com- The Bird 4 pelled to undergo the fall moult in a rather small body
of water, where danger menaces on all sides. Although
when flightless he swims low among the thick water-reeds,
yet his brilliant colours — iridescent green and white —
would too frequently mark him out. Moult So the invisible
cloak of his brooding mate is dropped over him for a
while — his colours vanish, and by a partial moult thus
sandwiched in, the hues of his plumage change to an
inconspicuous mottling of brown, hardly distinguishable
from the female. Then when the splitting of his quill-
sheaths hints of coming power to take care of himself
again, the dusky mantle is lifted, and, triumphantly
treading water, he stands upright and shakes his glisten-
ing wings, daring his enemies to catch him if they can. This has been happily termed the " eclipse" plumage. In
certain portions of the Old World where foxes are scarce
and the ducks have been persistently pursued by men
in boats, the knowing birds have changed their habits
and, when their wing-quills fall, they make their home
in deep woods, finding greater safety there than on ponds
or lakes. A somewhat similar condition occurs in the Black
Grouse of Europe, which loses the conspicuous black
feathers of the head and neck during the helpless period
caused by the moult of its tail-feathers. caused by the moult of its tail-feathers. This additional moult brings us to the consideration
of the birds which have no less than three changes of
plumage, and here we find the cause intimately connected
with the colour of the birds' surroundings. Ptarmigans,
which are species of grouse living in the far North, moult Feathers 49 FIG. 30. — Eclipse plumage of Mallard Duck. Male in full breeding p
(the brilliant green of the head and neck is lost in the photograp FIG. 30. — Eclipse plumage of Mallard Duck. Male in full breeding plumage
(the brilliant green of the head and neck is lost in the photograph). FIG. 31. — Male in eclipse plumage during moult of wing-feathers. FIG. 32.— Female Mallard. FIG. 32.— Female Mallard. The Bird 5°
after the breeding season into a special gray or dark
plumage, harmonizing well with the autumnal shades of
the grass and lichened rocks. In the late fall a second
plumage of immaculate white is assumed, affording these
birds great protection on the snowy wastes where they FIG. 33. — Willow Ptarmigan in early spring, with brown feathers beginning to
replace the white. 1/4 natural size. FIG. 33. — Willow Ptarmigan in early spring, with brown feathers beginning to
replace the white. 1/4 natural size. live. Moult In spring a third suit is donned — brown and parti-
coloured like the environment, which late in the year is
still covered with patches of snow here and there. This
too is the nuptial plumage, and lasts until the gray garb
completes the cycle of the year's changes. The wing-
feathers are white all the year, but when the wings are live. In spring a third suit is donned — brown and parti-
coloured like the environment, which late in the year is
still covered with patches of snow here and there. This
too is the nuptial plumage, and lasts until the gray garb
completes the cycle of the year's changes. The wing-
feathers are white all the year, but when the wings are Feat 5 1 closed they telescope so neatly beneath the feathers of
the shoulder that they are not noticeable while the bird
is in either the autumnal or vernal plumage. As the feathers on the flipper-like wings of a penguin
resemble the scales of reptiles in appearance, so this FIG. 33a. — Ptarmigan in the fall, showing the gray autumnal plumage (which
has replaced the brown of summer on the upper parts of the body and wings),
gradually giving place to the white of the corning winter feathers. Wild
birds in Alaska. (Harold Whealton, photographer.) FIG. 33a. — Ptarmigan in the fall, showing the gray autumnal plumage (which
has replaced the brown of summer on the upper parts of the body and wings),
gradually giving place to the white of the corning winter feathers. Wild
birds in Alaska. (Harold Whealton, photographer.) homology is carried out in the method of shedding them. Unlike the dropping out of feathers one by one, as in
other birds, these come off in flakes, like the skin of a
lizard. The feathers of the back loosen, shrivel up, and
fade to a brownish hue before they peel away. homology is carried out in the method of shedding them. Unlike the dropping out of feathers one by one, as in
other birds, these come off in flakes, like the skin of a
lizard. The feathers of the back loosen, shrivel up, and
fade to a brownish hue before they peel away. The Bird The Bird 5 We have seen how birds, by moulting their feathers,
change the colour of their plumage; in some cases several
times each year. There is, however, still another way in
which the appearance of new colour is brought about. Not by increase of pigment, for the feather when once
full grown is dead; but by the mere breaking or fraying FIG. 34. — The three moults of the Ptarmigan, shown in three individuals. (Courtesy of American Museum.) FIG. 34. — The three moults of the Ptarmigan, shown in three individuals. (Courtesy of American Museum.) of the edges of each feather. It is thus that the Snow-
flake brushes off the rusty trimmings of his winter's suit
and returns to his home in the far North, dressed in spick-
and-span black and white. A much more familiar exam-
ple is to be seen at our very doorstep. The cock English
Sparrow in midwinter is even more sombrely clad than
usual; but as spring approaches, although he can attain
to no elaborate song or flowing plume, yet even this of the edges of each feather. It is thus that the Snow-
flake brushes off the rusty trimmings of his winter's suit
and returns to his home in the far North, dressed in spick-
and-span black and white. A much more familiar exam-
ple is to be seen at our very doorstep. The cock English
Sparrow in midwinter is even more sombrely clad than
usual; but as spring approaches, although he can attain
to no elaborate song or flowing plume, yet even this Feathers 53
commoner feels the call of love for beauty, and day by
day the dusty brown tips of his throat-feathers wear
away one by one, and leave exposed the clear black
centres; and behold, the vulgar frequenter of our streets FIG. 35. — Two male English Sparrows, showing the difference in colour caused by
wear of the feather-tips between October and April. FIG. 35. — Two male English Sparrows, showing the difference in colour caused by
wear of the feather-tips between October and April. and alleys, flaunts a jet cravat before the eyes of his lady-
love! and alleys, flaunts a jet cravat before the eyes of his lady-
love! Colour. The very interesting uses which the colours of birds
serve, the part they take in courtship, in evading danger,
or in enabling birds to find each other, are many. These
uses have been much written about, but of the nature
and formation of colour less is known. Few of us have The Bird 5 probably ever given a thought to the colours themselves
Why is that feather blue? Why — because it is blue! Why is that feather blue? Why — because it is blue! There are two principal ways in which colours are
produced in feathers: first, when a real colour-pigment
is present, and again when the structure of the feather is
more or less like miniature prisms in shape, breaking up FIG. 36. — Young Brown Pelicans in the downy plumage. 1/6 natural size. FIG. 36. — Young Brown Pelicans in the downy plumage. 1/6 natural size. the rays— rainbow-like — into the iridescence of the spec-
trum. In the case of almost all the beauties of Nature,
the more closely we examine them, the more beautiful
they become. But this is not true of the iridescent
colours of birds such as hummingbirds, unless we con-
sider the structure. The colour itself disappears under
the microscope, and only gray or black tints are seen. the rays— rainbow-like — into the iridescence of the spec-
trum. In the case of almost all the beauties of Nature,
the more closely we examine them, the more beautiful
they become. But this is not true of the iridescent
colours of birds such as hummingbirds, unless we con-
sider the structure. The colour itself disappears under
the microscope, and only gray or black tints are seen. The black, red, brown, and yellow colours of feathers Feat 55 are almost always due to pigment or colouring-matter in
the shaft or vane. If we take a black feather and hold
it to the light, it will still look black; if we pound it with
a hammer, it will not change. a hammer, it will not change. Green is never found as a pigment except in the
feathers of a small family of birds called plantain-eaters
or turacous, which inhabit West Africa. Colour. For some time
it was thought that the natives dyed the birds artificially,
as when these birds were kept captive, the magnificent
scarlet patch on the wing would gradually fade and
become a dull gray. It is a fact that this colouring-
matter washes out when the feather is washed in alkaline
water. Even ordinary water will be slightly tinged if the
feather is soaked in it. The pigment contains about ten
per cent of copper, and this can be extracted chemically
in the form of a metallic powder. The plumage of almost
all brightly coloured birds will fade in the course of years,
if the feathers are left exposed to direct sunlight; but, like
photographic plates, the hues of some birds are more sen-
sitive than others to the light. The delicate reds and
yellows on the lower parts of Mexican Trogons are par-
ticularly evanescent, and the rose-pink of the African
Fairy Warbler disappears a short time after death. We might speak of a third class of colours, which are
due to both pigment and structure. For instance, no
blue pigment is known to exist in the feathers of birds,
but blue feathers contain a brown or yellowish pigment
which is encased in the horny coating of the feather. Between this outer sheath and the underlying pigment
is a layer of many-sided cones or small projections which The Bird 5 have numerous little ridges extending down the sides,
and in some way, by reflection, these change the yellow
or black to blue. If we take a parrot's feather and
pound the blue portion, that colour will disappear and
the vane will become black. It is surprising to see how the colours of many beau-
tiful feathers will vanish when we hold them between
our eye and the light. When we look at feathers under
the microscope, and see their horny rays, we forget, for
a time, the delicacy and fluffiness which the bird's plumage
as a whole exhibits, and we are constantly reminded of
the scales of reptiles. And in colour we have another
similarity between the two: lizards have both pigment
and prisms, and the scales of large snakes glow like opals
when the sunlight falls on them. Colour. White never exists as a pigment in the feathers of
birds, but is always due to innumerable air-spaces in the
substance of the feather, by which the rays of light are
reflected and deflected until, as in snow or foam, all
colour is lost and white results. In any one Order of birds there may often be found
a series of species with colour patterns grading into each
other and connecting two extremes, perhaps very diverse
in appearance. But it is seldom that we can examine
such a series at once, and, except in a large collection of
birds' skins in a museum, these wonderful life-chains, or
twig-tips of the tree of evolution seldom appeal to us
very forcibly. But in a feather it is different. We may
find on one bird a most delicately graduated series, show-
ing every step in the process by which simple unicoloured i
05 m
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8PS
SoO i
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SoO m m The Bird 5 or spotted feathers assume most intricate and complex
colour masses and patterns. Darwin illustrates this very plainly in the case of the
Argus Pheasant, and pays a fitting tribute to the evolu-
tion of the marvellous colour patterns among birds. "The
ocelli on the wing-feathers of the Argus Pheasant are
shaded in so wonderful a manner as to resemble balls
lying loose within sockets. That these ornaments should
have been formed through the selection of many succes-
sive variations, not one of which was originally intended
to produce the ball-and-socket effect, seems as incredible
as that one of Raphael's Madonnas should have been
formed by the selection of chance daubs of paint made by
a long succession of young artists, not one of whom in-
tended at first to draw the human figure. In order to
discover how the ocelli have been developed we cannot
look to a long line of progenitors, nor to many closely
allied forms, for such do not now exist. Colour. But fortunately
the several feathers on the wing suffice to give us a clue
to the problem, and they prove to demonstration that a
graduation is at least possible from a mere spot to a
finished ball-and-socket ocellus." finished ball-and-socket ocellus."
Two feathers from the wing of a Vulturine Guinea-
fowl have been chosen to illustrate a more simple but
no less beautiful colour evolution. On the less exposed
side of one of the feathers are three or four series of irregu-
lar white spots which tend in places to form transverse
bands. On the opposite side of the shaft near the tip
these spots are still distinct, but as our glance passes
gradually toward the base of the feather, the spots con- FIG. 38. — Adult Brown Pelicans in full breeding plumage. 1/8 natural size. FIG. 38. — Adult Brown Pelicans in full breeding plumage. 1/8 natural size. ral The Bird 6o verge more and more, until two distinct longitudinal
lines are formed, with traces of a third near the quill. A smaller feather from the same wing is marked with
spots which are nearly circular and which show faint
traces of encircling bands of white pointing toward a
still more elaborate system of decoration. FIG. 39. — Evolution of a colour pattern upon two feathers of a ulturme Gun.ea-
fowl; a stripe breaking up into dots, these forming cross-vars, and on the
second feather a regular series of dots encircled with white. FIG. 39. — Evolution of a colour pattern upon two feathers of a ulturme Gun.ea-
fowl; a stripe breaking up into dots, these forming cross-vars, and on the
second feather a regular series of dots encircled with white. FIG. 39. — Evolution of a colour pattern upon two feathers of a ulturme Gun.ea-
fowl; a stripe breaking up into dots, these forming cross-vars, and on the
second feather a regular series of dots encircled with white. It is interesting to conjecture in which direction the
decoration of feathers is proceeding. In the case of the
guinea-fowl, are the spots converging into lines or are
the lines the more ancient, and for some reason grad-
ually splitting up into smaller divisions? This is hard to
decipher, and if we look at the rest of the guinea-fowl's
body, the matter becomes only the more complicated. Colour. Feat 6 1 For, higher up on the wings, and on the shoulders, we
find that the fine specks which were barely noticeable on
the tips of some of the wing-feathers, are in the ascend-
ant, and absorb or replace the white spots over the whole
feather. The faint trace of the third line near the shaft
of which I spoke, has suddenly assumed an unexpected
importance and has spread out into a broad central band. The young or the female might give us a clew; for in
rnany birds the coloration of these shows a more ancient
arrangement of colour pattern than the feathers of the
male. The Indian Wood Ibis — what an imbecile it looks to
our eyes when we observe it in a zoological garden; what
a fishy smell it generally diffuses, how unpleasant are its
feeding habits, and what a dull black and white colora-
tion it has! Surely here is a bird with nothing which
could possibly appeal to our aesthetic sense. But we are
mistaken. Some of the innermost feathers of its wings,
seldom visible, except when the bird partly spreads them,
are of the most beautiful rose hue, shading at the tip
into a deeper pink. Seldom, even in Nature, will we
find tints comparable to the delicacy and bloom of these
hidden feathers. We have gone into these details only to show the
possibilities of a little feather-study. Even our common
Plymouth Rock chickens and hundreds of other birds
will show us unthought-of beauties, and in the fields or
in a zoological park we have only to use our eyes more
carefully to realize how ^auch we usually pass by un-
noticed. FIG. 40. — X-ray photograph of the front view of a homing Pigeon, showing the
bones clearly through the surrounding flesh. Observe the light, spongy
character of the skull and the bones of the limbs, the latter appearing almost
hollow. The crop filled with corn is visible spread out at the base of the
neck, and low down in the body, near the right thigh, the grit and pebbles
within the gizzard are very distinct. About the tarsus of the right leg is
seen the metal tag which was used for the identification of the living bird.
(Photographed by Dr. Henry G. Piffard.) THE FRAMEWORK OF THE BIRD H HEN we look at a living bird, we see only feathers,
horn, and skin, and we sometimes forget that
hidden beneath all these are many bones, — the
framework of the body. If we wish to alter the style of
architecture of a house, we need only to change the ex-
terior, columns, arches and windows, while the stone
foundation and brick walls may remain as they are. So
in fashioning new forms of life, Nature has often altered
the covering, and even the muscles and organs, of ani-
mals to such an extent that we would have little clew
as to the relations of these creatures, were it not for the
underlying bones, which are so deeply seated that they
react less slowly to changes in the outside life. If a fish,
a lizard, a bird, a whale, and a man should be presented
to us for classification, we might well hesitate until we
had seen their bones, when there would flash upon us
the same moulded type running through all. The study of the skeleton, or Osteology, is like all
other 'ologies; it can be made as dry as the bones them-
selves; or the very opposite, by leaving the minor details
and less important particulars to text-books, choosing
only the most significant facts. One may smile at the at FIG. 40. — X-ray photograph of the front view of a homing Pigeon, showing the
bones clearly through the surrounding flesh. Observe the light, spongy The Bird 6 thought of bestowing an encomium on a jaw-bone, and
yet the history of the lower part of a sparrow's beak
opens a vista so far-reaching that the mind of man falters
at the thought; it shows the last roll of an evolving
which, could we follow it back, would merge the man, the
whale, the bird, the lizard, the fish, into one. Let us look at some of the bones of a sparrow or
dove or chicken. One way to do this is to place a dead
bird in a box pierced with numerous holes, leave it near
an ant-hill, and wait for the industrious insects to do
their work. Another way is to clean as much flesh as
possible from the skeleton and deposit the bones in a
pail of water. In a few days they can be washed white
and clean. THE FRAMEWORK OF THE BIRD Perhaps the easiest way of all is to save what
bones you can of a boiled chicken. These are of large
size and will show us all we wish to know. The framework of a bird consists of a long jointed
string of bones called vertebrae, with the brain-box or
skull at one end and a blunt tail at the other. Near
the middle, the outcurving ribs extend around the organs
of the body, and, with the breast-bone, form an encircling
protective sheath. Two short series of bones project in
front of the ribs — the bones of the wings, — and two more
behind the ribs — those of the legs and feet; while at the
point of attachment of each of these four limbs there
radiates a trio of bones. The back-bone is the fundamental and oldest part of
the skeleton, and though we cannot follow its evolution
directly backward through the long ages, yet there is
sufficient gradation among living creatures to give us FIG. 41. — Common Fowl, showing relation of the bony framework or skeleton
to the contour of the body. Notice large eye, long and mobile neck, the
knee wholly within the body plumage, and the well-developed keel hinting
of ancestors with strong powers of flight. 1/4 natural size. 6? FIG. 41. — Common Fowl, showing relation of the bony framework or skeleton
to the contour of the body. Notice large eye, long and mobile neck, the
knee wholly within the body plumage, and the well-developed keel hinting
of ancestors with strong powers of flight. 1/4 natural size. The Bird 6 hints of the way it originated. In the lowest of fish-like
creatures — the Amphioxus, a tiny animal, an inch or two
in length, living in the sand along our shores — there is
a thread-like cord of a gelatinous substance (not carti-
lage, however) extending down the back, known as the
notochord. He looks like some kind of worm, but this
little gristle is his badge of nobility and lilts him clear FIG. 42. — Amphioxus, one of the lowest vertebrates, with a mere thread of
gristle foreshadowing the back-bone of higher animals. This creature bur-
rows in the sand along the Atlantic coast. FIG. 42. — Amphioxus, one of the lowest vertebrates, with a mere thread of
gristle foreshadowing the back-bone of higher animals. THE FRAMEWORK OF THE BIRD This creature bur-
rows in the sand along the Atlantic coast. of corals, snails, insects, and worms, into the realm of
back-boned animals. This notochord lies underneath a
thin white line which is all the spinal chord he has, and,
at the front end of this, a tiny dot of pigment stands
for brain, eye, and ear. Indeed Amphioxus has neither
skull, brain, nor limbs. The history of the back-bone, like human history, is
not altogether a majestic upward evolution; it has its
tragedies and set-backs, its hopes and failures. In the
waters along our Northern seashores are creatures, some
sponge- or lichen-like, others with strange bulb-like bodies * The actual evolution of birds was of course not through fish, tadpoles,
and reptiles as we know them, but by some line of creatures unknown to us
forever, and resembling some of these other living Classes at least in the pos-
session of gills, scales, etc. The Framework of the Bird growing on the end of long stalks. We call them almost
plants. But they hold a secret from the crabs and snails
which crawl about, and when the fishes brush against IG. 43. — A colony of living Boltenia, photographed by the author in the Bay
of Fundy. The Boltenia is one of Nature's failures to make a vertebrate. The larva is active and has a notochord; the adult is degenerate and fixe
on a stem. Found in five fathoms and deeper off rocky coasts north of Cap
Cod. FIG. 43. — A colony of living Boltenia, photographed by the author in the Bay
of Fundy. The Boltenia is one of Nature's failures to make a vertebrate. The larva is active and has a notochord; the adult is degenerate and fixed
on a stem. Found in five fathoms and deeper off rocky coasts north of Cape
Cod. them — if their poor dull senses only knew it — they
might claim a blood-brotherhood. When they were
young, for a little while, a gelatinous notochord was
theirs also, but this, with all the hopes that such a be- 6 The Bird ginning brings, of fish, of bird, of man even, soon melted
away and there they nod and sway in the watery cur-
rents, never to know of the opportunity Nature has
snatched from them — why, who can tell? snatched from them — why, who can tell? In adult sharks, the back-bone has become jointed
and flexible, and a crude kind of skull is present, but
still more important is the presence of four fins which
correspond to the four legs of lizards and to the wings
and legs of birds. A curious basket-like skeleton pro-
tects the delicate gills, and it is probable that this existed FIG. 44. — Back-bone of Dogfish, with simple cartilaginous vertebrae. FIG. 44. — Back-bone of Dogfish, with simple cartilaginous vertebrae. long before the limbs appeared. All of this is composed
of gristly cartilage. In the higher fishes, bone replaces
the cartilage, and when the lowly tadpole — fish-like at
first, swimming about by means of the fin around his
tail — pushes forth his legs and climbs upon the land, our
skeleton is well on its way birdwards.* Reptiles of old
took to trees; their back-bones grew less flexible, so that
they might safely sail through the air; feathers replaced The Framework of the Bird The Framework of the Bird scales, two ringers of each hand were lost, and one from
each foot; teeth disappeared; a beak of horn proved
best; intelligence increased and the forehead rose high,
and behold, — a bird! Can we then despise even an Eng-
lish Sparrow? All these things we have learned from a comparison
with creatures other than birds, and we may, without
trouble, take one more glimpse into the dim past. Let
us go to the hencoop, where for three days the patient
biddy has been sitting on her precious eggs. We will FIG. 45. — Neck vertebrae of an Ostrich, highly complex and bony in structure. FIG. 45. — Neck vertebrae of an Ostrich, highly complex and bony in structure. rob her of one — she will not miss it — while from it we
may learn many wonderful things. Rest the warm egg
in a dish of sand, carefully picking away the shell from
the upper part. A glance at the tiny embryo lying on
the yolk within will show a double series of tiny squares
extending down the long diameter of the body. These
are the first hints of the spinal column, and if we could
follow its further development we would see something
of great interest. The squares are now divided up like
beads, just as are the bones of our bird's vertebrae; but
in reality this first segmentation is a false one. It is sim- The Bird 7°
ply a copy of the primitive flakes or joints of the tiny
muscle-beginnings, and is comparable to the joints or
rings in the body of a beetle, butterfly, or earthworm. In a short time all the squares will fuse together, and not
until later will they separate again into divisions which
will ultimately form the real bones of the spinal column. Every little chick, before it hatches, goes through the
same strange changes, — living reminders of the evolution
which has gone on in past ages of the earth. It is inter- FIG. 46. — Muscle-plates, or false vertebrae, of third-day embryo chick. Magnified 25 diameters. FIG. 46. — Muscle-plates, or false vertebrae, of third-day embryo chick. Magnified 25 diameters. esting to note that the vertebrae of the embryo chick
pass through a stage when they are biconcave, — a condi-
tion found both in Amphioxus and Archseopteryx. The Framework of the Bird This digression upon the back-bone history may seem
out of place, but in reality such a bird's-eye survey of
the past, imperfect as it is, will add a new interest to
our handful of chicken-bones. The Framework of the Bi * This bone is formed chiefly of two intercentra, which are small bones,
very characteristic of reptiles (chevron-bones of the tail) and are not uncom-
mon among the lower Orders of birds.
t In Hornbills the atlas and axis are fused together. The Framework of the Bi 71 Let us suppose that we have strung a wire through
the hollow centre of the back-bone of our chicken, to
which the ribs are still attached, and that we have be-
sides the skull and the bones of one wing and one leg. Compare them with those in the illustrations and we will
see if they can tell us aught of interest. The bones of the neck are all separate, and slide back
and forth on the wire, like beads on a string. How unlike FIG. 47. — Front and rear views of seventeenth and eighteenth cervical vertebrae
of Ostrich, showing complicated saddles and sliding surfaces, giving great
freedom of motion. FIG. 47. — Front and rear views of seventeenth and eighteenth cervical vertebrae
of Ostrich, showing complicated saddles and sliding surfaces, giving great
freedom of motion. the long smooth ribs are these vertebrae, bristling with
spines and projections! How is it that a bird can be
comfortable with a string of such irregular-looking ob-
jects run through its body? But fit two of these bones
together and see how beautifully they saddle end to end,
every convexity or projecting knob exactly adjusted to
a corresponding concave portion of the neighbouring bone. These saddles are characteristic of birds alone. Every
one of the sixteen bones of the neck is different from its the long smooth ribs are these vertebrae, bristling with
spines and projections! How is it that a bird can be
comfortable with a string of such irregular-looking ob-
jects run through its body? But fit two of these bones
together and see how beautifully they saddle end to end,
every convexity or projecting knob exactly adjusted to
a corresponding concave portion of the neighbouring bone. These saddles are characteristic of birds alone. Every
one of the sixteen bones of the neck is different from its The Bird 7 fellows and exactly suited to the requirements of its
position, but the first two following just behind the skull
are so radically unlike the others that we know at once
that they must serve some particular purpose. The first
is little more than a simple ring * of bone, and is called
the atlas, after the mythological giant who held up the
heavens upon his shoulders; named very aptly too, for FIG. 48. — Atlas and axis of Jabiru, separated FIG. 49. — Atlas and axis of
Jabiru, joined. The Framework of the Bi FIG. 48. — Atlas and axis of Jabiru, separated FIG. 49. — Atlas and axis of
Jabiru, joined. this tiny collar of bone supports the skull itself. The
next vertebra is ring-like too, but has a curious knob in
front, which projects forward through the atlas and forms
a pivot on which the head turns, hence its name, — the
axis.f Let us compare the neck-bones with those of a reptile
and a man. Although, as a whole, the bones of the The Framework of the Bi 73 skeleton of a bird are more or less soldered together,
yet the neck is far more flexible than in either of the
other examples. Indeed the neck of a bird has greater
freedom of motion than that of a snake. A lizard can
turn his head only a little way around, and we ourselves
can look only across our shoulder, but with a bird it is FIG. 50. — American Egret, showing curves into which the neck naturally falls
when the bird is at rest. When striking at a fish the vertebra? straighten
out. FIG. 50. — American Egret, showing curves into which the neck naturally falls
when the bird is at rest. When striking at a fish the vertebra? straighten
out. very different. Watch a heron or, better still, a fla-
mingo and see its neck describe figures of eight as he
arranges the feathers on its back. Few people would
ever imagine that there are exactly twice as many neck-
bones in a sparrow as in a giraffe, but such is the case,
there being fourteen in the former and seven in the latter. In the neck of a swan there are twenty-three of these
bones. The Bird 74 The remaining vertebrae, those of the upper and lower
back, are very different from those of the neck. The
flexible neck enables the bird to reach all parts of its
plumage with its beak, and to pick up food from the
ground or from twigs overhead, but the all-important
function of flight must be provided for by means of a FIG. 51. — White-throated Sparrow, three inches tall, with fourteen neck vertebrae. (Compare with Fig. 52.) FIG. 51. — White-throated Sparrow, three inches tall, with fourteen neck vertebrae. (Compare with Fig. 52.) rigid body-frame. In reptiles and in the embryos of
birds only two pelvic vertebrae are fused together, but
in adult birds many dorsal and caudal vertebrae (as many
as 23 in some cases) are fused into a single bone. Thus
the rib-bearing tipper back vertebrae are partially fused
together, and below them those of the lower back have
merged until it is difficult to realize that this portion of The Framework of the Bird The Framework of the Bird The Framework of the Bird 77 hence the separate bones which unite it to the vertebrae
of the lower back. The evolution of the tail will be
treated of in another chapter. The Framework of the Bird 75 the skeleton was not originally one bone. Passing on for
a moment to the bones of the tail, we find a number of
separate pieces, ending in a curious-shaped bone, called
the ploughshare. This is at the tip of the tail, or " pope's FIG. 52. — Giraffe, reaching with tongue for leaves perhaps eighteen feet above the
ground, with but 7 neck bones. FIG. 52. — Giraffe, reaching with tongue for leaves perhaps eighteen feet above the
ground, with but 7 neck bones. nose," of the chicken and really consists of many verte-
brae fused together. It is necessary for this to be large
and strong: for it supports all the feathers of the tall. But to be of efficient aid in steering, the tail, like the
rudder of a ship, must have freedom of motion, and nose," of the chicken and really consists of many verte-
brae fused together. It is necessary for this to be large
and strong: for it supports all the feathers of the tall. But to be of efficient aid in steering, the tail, like the
rudder of a ship, must have freedom of motion, and FIG. 53. — Pelvic vertebrae of young Alligator. FIG. 53. — Pelvic vertebrae of young Alligator. Fia. 54. — Pelvic vertebrae of American Flamingo. Fia. 54. — Pelvic vertebrae of American Flamingo. FIG. 55. — Pelvic vertebrae of Bald Eagle. In the reptile, where there is no need
for rigidity, only two typical pelvic vertebrae are joined together; in the birds
many dorsal and caudal vertebrae are joined with these to make a rigid frame
for flight and for bipedal locomotion. 76 FIG. 55. — Pelvic vertebrae of Bald Eagle. In the reptile, where there is no need
for rigidity, only two typical pelvic vertebrae are joined together; in the birds
many dorsal and caudal vertebrae are joined with these to make a rigid frame
for flight and for bipedal locomotion. 76 frame
76 The Framework of the Bird Ribs :AV7vv
The ribs are the long, narrow, double-headed bones
which curve out from the vertebrae of the upper back
and, uniting with the breast-bone, form a barred pro-
tection for the heart, lungs,
liver, and other organs. These
are the ribs proper, but there
are other smaller ones, called
false or floating ribs, which
reach only part of the way
around the body. Look at
the largest ribs of the chicken
and an added provision for
making this box of bone more
solid will be seen. From near
the centre of the upper part
of each rib a small bony projection laps across the rib
next behind and thus forms a kind of lattice-work,
movable but of great strength. A further interest
attaches to these cross-rib pieces when we learn that every
bird except the Screamer possesses them, while else-
where they are found only in crocodile-like reptiles
and in the Hatteria Lizard of New Zealand. FIG. 56. — Ribs of Hatteria Lizard,
with uncinate processes. :AV7vv
FIG. 56. — Ribs of Hatteria Lizard,
with uncinate processes. FIG. 56. — Ribs of Hatteria Lizard,
with uncinate processes. The similarity of the ribs — slanting one after another The Bird 7 around toward the breast-bone — hints of something
which perhaps has never occurred to us. We spoke of
the worm-like appearance of the lowly Amphioxus —
the sand-fish with the shadow of a back-bone. When we
think of a worm we think of a creature very much alike
from head to tail, one in which a section across the neck
is not very unlike one across the centre of the body or
near the tail; indeed that is exactly what the word Am-
phioxus means, — like head, like tail. This repetition of
segments or similar parts is a sign of low degree in the
scale of life, as it harks back to the time when the very
highest form of life was worm-like. highest form of life was worm-like. The flesh of a salmon or of a trout shows such a con-
dition very well, the body consisting of flake after flake
of flesh. Now in birds and the higher animals this divi-
sion into successive segments is hardly noticeable, and
almost every inch of a man or bird, from head to toe,
seems very distinct and individual. Ribs But ribs bring back
the old ancestral condition very vividly, and when a
peacock, strutting proudly before us, resplendent from
beak to tail, picks up and swallows an unfortunate angle-
worm, we may remember that, no matter what geological
eras or inexplicable physical gulfs separate the two, the
bird carries within his body indelible imprints which
insolubly link his past with that of the lowly creature
of the dust. As in various other cases throughout nature, when the
many ribs of the bird's ancestors began to be reduced in
number, some attained to other uses beside that of arch-
ing around the whole body and protecting the heart, the The Framework of the Bi 79 lungs, and other organs. Look at the two neck-bones of
the ostrich in Fig. 47, where in addition to the central
aperture, through which the spinal nerve-cord passes, two
other openings will be seen, one on each side. Through
these the vertebral arteries carry their burden of pure
blood. The outer wall of this bony canal, extending up
the whole length of the neck, is formed principally by
what is left of the ribs which were once long and free,
like their fellows farther down the back. Though re-
duced to a tiny fragmentary arc of bone, yet they still
perform a protecting function. perform a protecting function. In Archceopteryx (Fig. 5) there existed well-developed
abdominal ribs, exactly like those found in crocodiles
and other reptiles. In no living bird, however, are these
found. Breast-bone The lower portions of the true ribs of our chicken
are separate pieces of bone, slanting in a forward direc-
tion and attached by a movable joint to the upper parts. These end close together along the sides of the large
breast-bone, or sternum as it is called. In fact the origin
of the sternum can be traced to the fused ends of these
ribs, and in the sternum of an immature, ostrich (Fig. 58) the line of juncture between the two lateral halves is
still distinct or even open. To the edge of this bone,
nearest the head, two column-like shoulder-bones are
attached, and in some birds the wish-bone is also joined
to it (Fig. 103). The sternum is one of the largest single bones in The Bird 8 the body of the chicken, and is very different from our
own breast-bone, which is long and narrow. The pos-
terior edge of the sternum is of many shapes, varying in
birds of different species. Deep channels may extend
into each side, leaving long slender splinters or spines FIG. 57. — Ribs and sternum of Flamingo; notice what a complex box of bone
is formed by the vertebrae, scapula, ribs with their uncinate processes, and
the sternum; notice large keel for the attachment of flight-muscles. FIG. 57. — Ribs and sternum of Flamingo; notice what a complex box of bone
is formed by the vertebrae, scapula, ribs with their uncinate processes, and
the sternum; notice large keel for the attachment of flight-muscles. of projecting bone, or this channel may be partly closed,
forming a round hole quite through the bone. The
sternum and its various processes are of considerable
value in classification, the same configuration being found
throughout allied groups, in consequence, doubtless, of
the slight chance of modification resulting directly from
any specialized habits in the life of the bird. The portion The Framework of the Bi 81 of this bone which is most characteristic of birds is the
central ridge or keel which projects straight out from
the surface of the sternum. This is of the utmost im-
portance in giving firm anchorage for the great flight-
muscles of the breast. FIG. 58. — Ribs and sternum of Ostrich; notice absence of keel correlated with
loss of power of flight. The suture through the centre of the sternum reveals
its paired origin. FIG. 58. Breast-bone — Ribs and sternum of Ostrich; notice absence of keel correlated with
loss of power of flight. The suture through the centre of the sternum reveals
its paired origin. The keel is of many shapes, but when well developed
is generally very high at the upper end of the breast-
bone and becomes lower as it slopes gradually backward. In birds which have lost the use of their wings the keel
has disappeared completely, the sternum being flat, as
in ourselves. This is its condition in the ostrich and
cassowary, and it is the character which has given a The Bird 8 name to two great divisions of birds : Rati'tce (those with
flat breast-bones, raft-like), including the ostrich, rhea,
emeu, cassowary, and apteryx; and Carina'tos (birds
with keeled breast-bones, boat-like), including all other
living birds, whether flyers, as the thrushes, storks, and
gulls, or swimmers like the penguins. But this differ-
ence in breast-bones is far from being as profound as
other differences existing between certain birds which
are alike in having keels to their sternums. The pres-
ence or absence of a keel is not of great taxonomic im-
portance. portance. The size of the keel is a pretty sure criterion of the
flying powers of a bird, that is, judging not the actual
duration of flight, but the actual muscular power and
amount of energy used in flying (Fig. 59). The alba-
tross, and other birds which, trusting to the air-currents
to bear them upward, flap seldom and soar much, have
comparatively smaller keels than do those birds which
flap their wings more frequently. Thus the pigeon has
a very good-sized keel; while in the humming-bird this
bone is enormous, compared to its spread of wings. Dr. Frederick A. Lucas has expressed this very graphically
in a diagram, where it is supposed that the albatross,
pigeon, and humming-bird have an equal spread of wings. On comparison, the keel of the first is seen to occupy but
a small fraction of the surface of the same bone in a
humming-bird. To account for this we must realize
that the wings of the humming-bird execute from six
hundred to a thousand strokes a minute; while the alba-
tross may soar for miles with wings held outstretched The Framework of the Bi 83 and all but motionless. Breast-bone It is said that, comparatively,
the muscular energy is greater and the wing-bones more
powerful in a hummingbird than in any other animal. powerful in a hummingbird than in any other animal. Nature has a puzzling way of achieving similar results
in a very similar manner in creatures wholly unrelated. We have a good example of this in bats and birds, both FIG. 59. — Comparison of the size of the keel of the Albatross, Pigeon, and Humming-
bird, supposing all to have an equal spread of wing. (Courtesy of Dr. F. A. Lucas.) FIG. 59. — Comparison of the size of the keel of the Albatross, Pigeon, and Humming-
bird, supposing all to have an equal spread of wing. (Courtesy of Dr. F. A. Lucas.) of which have independently learned to propel them-
selves through the air by means of their front limbs. If we take the breast-bone of a common bat and that
of a small bird and place them together, few persons
unacquainted with the bones of the two types could tell
which was that of the bat, — different as that little crea- The Bird 8 ture is in external appearance from a bird. Their keels
and sternums are very much alike. This is called par-
allelism, and sometimes it gives a great deal of trouble
to naturalists when they are trying to find the right
relationships between living animals. Shoulder-girdle It will be remembered that mention was made of the
trios of bones which radiate near the juncture with the
body, of each wing and each leg, — girdles or arches they
are called. The pectoral, or shoulder, girdle meets the
upper arm-bone of each wing at the shoulder- joint. If
we run our hand along the back of our shoulders, we
will feel a prominent bone, called the shoulder-blade, and
in almost the same place in our chicken we notice a very
long and thin bone. This is the scapula, and is one of
the pectoral-girdle trio, the other two being known as
coracoid and clavicle. The coracoid is a short, but stout, column of bone
joined to the shoulder-blade and extending down and
backward to the breast-bone. This coracoid bone is
especially developed in birds as compared with other
creatures. When their forefathers began to scale through
the air, thus putting a great strain on the muscles of
the breast, Nature seized on these coracoid bones, giving
them such strength and thickness that they have become
the pivots upon which, at each swift vibration through
the air, turn the marvellous wings of a modern bird. In
reptiles, this bone is divided into two weak, thin plates The Framework of the Bi 85 which would hardly afford strength for a single wing-
flutter. Since mammals in their high evolution have
found no use for this bone, it has become reduced to a
small projection on the shoulder-blade. small projection on the shoulder-blade. The clavicle we will recognize instantly, when we
give it another name — the wish-bone or merry-thought. FIG 60. — Pectoral girdle of bird (scapulas, coracoids, and clavicles); compared
with the scapula and coracoid of a young Leopard, the latter bone in the
Leopard being reduced to a tiny process. FIG 60. — Pectoral girdle of bird (scapulas, coracoids, and clavicles); compared
with the scapula and coracoid of a young Leopard, the latter bone in the
Leopard being reduced to a tiny process. In birds the wish-bone is generally V-shaped, the two
clavicles usually meeting and fusing at their tips. Through
this V-shaped opening in the neck, the resophagus and
the windpipe pass from the throat into the body cavity. We too have wish-bones, although they are not placed
exactly as are those of a chicken. Shoulder-girdle We call them collar-
bones, but by whatever name we know them they are
of importance, both in ourselves and in birds, in serving
to brace out the shoulders. In creatures which, unlike The Bird 8 mankind and most birds, have less varied movements of
the fore limbs, the clavicles have fallen into disuse, as
in the lion and the horse. But in climbing, burrowing,
and flying animals, such as the squirrel, mole, and bat,
these bones have been of active use and are well devel-
oped. But to keep its wish-bone a bird must continue
to fly: for Nature is opposed to useless parts. So, in
the flightless cassowary and ostrich, the wish-bone is
very small or altogether absent. Parrots are almost
alone in appearing to suffer no inconvenience in flight
by the lack of clavicles, — these being greatly reduced in
some species. In that anomalous bird the Hoatzin, the clavicles
are fused not only at their base, but the tips are ossified
firmly to a projecting spine of bone from the upper part
of the breast-bone. In glancing back over the lower back-boned animals
we realize that a shoulder-girdle of bones is of no use
without a limb. Therefore we find the first hint of the
shoulder-girdle in sharks, in which we also find the first
limbs, or fins. In these fishes it is nothing but a single
bar of soft cartilage. In the girdle supporting the pec-
toral fin of such a fish as the trout or other bony fish,
we find the adumbration of some of our bird's bones. When we remember how very wing-like is the movement
of a fin in the water, we will not be surprised to learn
that the girdle is almost all epiclavicle; these bones
being the forerunners of clavicles, and giving place, in
the higher forms, to the real wish-bones which steadily
increase in size and importance. We would hardly The Framework of the Bi The Framework of the Bi 87 recognize in these primitive types the wish-bone of our
Christmas turkey. In terrestrial quadrupeds and birds we usually find
the front limbs near the front part of the body and the
hind limbs much farther back, but it is interesting to FIG. 61.— Girdle of a bony fish. FIG. 61.— Girdle of a bony fish. FIG. 61.— Girdle of a bony fish. notice that in the fish, Fig. 61, all four limbs or fins are
very far forward, almost or quite in the head region
itself. This is a result of the function of balance which
these structures almost wholly perform, the fin of the
tail furnishing the locomotive power. It is very interesting to notice how many bones have
kept to their respective places in the evolution of animals, The Bird 8 no matter how much change has occurred in their
shape and size. Take, for instance, the shoulder-blades. When a tiger crouches they are very conspicuous, and
whether we take a frog, a turtle, a lizard, an armadillo,
a mouse, or a horse, we may always be sure of finding a
scapula in the region where we have observed it in the
bird. This is an important fact, and one which makes
the identification of many bones an easy matter. * They were separate also in Archcenpteryx. Thigh-girdle The shoulder-girdle which we have just examined
was not joined to the back-bone, but only saddled on
the ribs, the scapula extending backward, just clearing
them. What kept it in place in the chicken's skeleton
was the fact that it was strongly attached to the sternum,
and this in turn joined to the back-bone by means of
the ribs. But the pelvic arch or thigh-girdle is very
different. If the entire framework of the bird is to be
supported on two legs, the point of attachment of these
limbs must be solidly fixed to the back-bone of the body. limbs must be solidly fixed to the back-bone
Although there are as many bones supporting the leg
or thigh as there are bracing the shoulder, we would
never know this from examining our chicken. As in
other places in a bird's skeleton, the bones — six in this
case — have fused together in one solid piece, and only
in very young birds are they separate.* in very young birds are they separate.*
The names of the bones composing the pelvic girdle,
or arch, are the ilium, ischium, and pubis. The easiest The Framework of the Bi 89 way to locate these is first to find the socket in which
fits the head of the thigh-bone. This is the deep cup-
shaped depression on each side, and all three bones join
in making the socket. The ilium lies along the back
and forms a sort of roof over the portion of the back-
bone in this region. If we look at the under side of this
bone, we may see the fused vertebrae more distinctly —
fourteen or fifteen of them. The two deep depressions
in which the kidneys of the bird were located are also FIG. 62. — Pelvic girdle of a bird. FIG. 62. — Pelvic girdle of a bird. now visible. As the coracoid is the great pivot of the
wing, so the ilium helps most to bear the strain of hop-
ping and running. In the frog, which progresses by
hops or great leaps, the ilium is also largely developed;
indeed we can see it through the skin, thus giving the
broken-back appearance to that creature. now visible. As the coracoid is the great pivot of the
wing, so the ilium helps most to bear the strain of hop-
ping and running. Thigh-girdle In the frog, which progresses by
hops or great leaps, the ilium is also largely developed;
indeed we can see it through the skin, thus giving the
broken-back appearance to that creature. broken-back appearance to that creature. Each side of the thigh-bone box is formed by the ischium,
which is closely fused with the ilium except in most of
the ostrich-like birds, the tinamous, and in reptiles,
where these bones are free throughout their entire length. We can readily make out the pubis as a slender bar of The Bird
-JL 9°
bone extending backward from the thigh-socket, sepa-
rated from the ischium
(except at the extreme
end) by a long open slit. In other animals these
bones are as different in om the thigh-socket, sepa-
rated from the ischium
(except at the extreme
end) by a long open slit. In other animals these
bones are as different in
shape as can be imagined,
but, almost without excep-
tion, the position of each
is relatively the same. The
extreme extension, forward
and back, of the ilium
above the back-bone, thus
joining with more vertebrae,
is directly connected with
two-legged erect locomo-
tion. A parallel condition
is found in some Dinosaurs
— those extinct giant rep-
tiles -- certain of which
walked more or less on
their hind legs. PuA
FIG. 63 — Pelvic arch of a Dinosaur FIG. 63 — Pelvic arch of a Dinosaur FIG. 63 — Pelvic arch of a Dinosaur is found in some Dinosaurs
— those extinct giant rep-
tiles -- certain of which
walked more or less on
their hind legs. Another fundamental
resemblance is found be-
tween the thigh-girdles of
-Js. Dinosaurs and other rep-
tiles and that of a bird in
the egg. As is shown in
Figs. 63-65, the pubis slants
slightly forward in both reptile and embryo bird ; but in
FIG. 64. — Pelvic arch of an embryo bird,
to show similarity of the two as
contrasted with Fig. 65. Pur
FIG. 65. — Pelvic arch of an adult bird. (The three figures by courtesy of
Prof. H. F. Osborn.) -Js. FIG. 64. — Pelvic arch of an embryo bird,
to show similarity of the two as
contrasted with Fig. 65. Pur
FIG. 65. — Pelvic arch of an adult bird. (The three figures by courtesy of
Prof. H. F. Osborn.) FIG. 64. The Framework of the Bird the adult bird the shifting backward of this bone until it
is parallel with the ischium is wholly an avian feature. FIG. 66. — Bullfrog. The bend in the back shows the great development of the
ilium for bipedal locomotion in the sense of leaping ability. FIG. 66. — Bullfrog. The bend in the back shows the great development of the
ilium for bipedal locomotion in the sense of leaping ability. Thigh-girdle — Pelvic arch of an embryo bird,
to show similarity of the two as
contrasted with Fig. 65. FIG. 64. — Pelvic arch of an embryo bird,
to show similarity of the two as
contrasted with Fig. 65. Another fundamental
resemblance is found be-
tween the thigh-girdles of
Dinosaurs and other rep-
tiles and that of a bird in
the egg. As is shown in
Figs. 63-65, the pubis slants
and embryo bird ; but in Another fundamental
resemblance is found be-
tween the thigh-girdles of
-Js. Dinosaurs and other rep-
tiles and that of a bird in
the egg. As is shown in
Figs. 63-65, the pubis slants
slightly forward in both reptile and embryo bird ; but in
Pur
FIG. 65. — Pelvic arch of an adult bird. (The three figures by courtesy of
Prof. H. F. Osborn.) Pur
FIG. 65. — Pelvic arch of an adult bird. (The three figures by courtesy of
Prof. H. F. Osborn.) Pur
FIG. 65. — Pelvic arch of an adult bird. (The three figures by courtesy of
Prof. H. F. Osborn.) The Framework of the Bird Wings We will now consider the framework of the fore limb,
or wing, of a bird, and a glance at the illustration show-
ing the arm of a man and the wing of a bird will at once
make plain the relation between the two. Here we again
find a great help in the fact that many of the bones keep
to their respective places in frogs, lizards, birds, and man. We know but little of the direct change from a fin to
a hand or foot, although there are some fishes living at
the present day with large finger-like bones in their pec- The Bird 9 toral fins. Even among fossil forms there have as yet
been found no " missing links" in this respect. But how-
ever it came about, it is certain that when the fish-
amphibians of olden time, venturing into shallow water,
felt more or less solid mud under them, and tried to
move about upon it, their fins must have become pressed
downward, and before they could safely push themselves
about on dry land or lift their bodies clear of the ground,
the stiff fin-rays must have become split up into a few,
thick, bony rays or toes. We know that these were
originally five in number on all four limbs, and when-
ever, among living creatures, we find a lesser number, the
reduction has been brought about by some subsequent
change in the life of the animal. As yet, however, we
know of no direct transitions from fins to feet. The requirements of flight demanded a fin-like stiff-
ness in the wings of birds, and therefore many of the
smaller bones of lizards, counterparts of which we find
in our own wrists and hands, are in the bird fused together. in our own wrists and hands, are in the bird fused together. The upper arm-bone, or humerus, corresponds exactly
to our bone of that name, and when we feel the two long
bones of our forearm and look for them in the bird, we
find both very plainly represented, the large one with
notches, where the great wing-feathers are fastened, being
called the ulna, and the smaller, straighter bone the
radius. In our wrist there are eight little bones which
are joined to each other so delicately that we can move
and turn our hand in every direction. * Some morphologists homologize the fingers of a bird's wing with the
second, third, and fourth digits of a pentadactyl hand. The question is still
a mooted one.
f In the embryos of some birds, traces of a fourth finger have been found. Wings But when a bird's
wing is extended, if the wrist was at all flexible, the pres-
sure of air on the great wing-feathers would turn the The Framework of the Bi The Framework of the Bi 93 f the Bird 93
FIG. 67. — Wing of Pigeon, leathered. FIG. 68. — Wing of Pigeon, bare, compared with Fig. 69. FIG. 67. — Wing of Pigeon, leathered. FIG. 67. — Wing of Pigeon, leathered. FIG. 68. — Wing of Pigeon, bare, compared with Fig. 69. FIG. 68. — Wing of Pigeon, bare, compared with Fig. 69. FIG. 69. — Arm and three fingers of a man. FIG. 69. — Arm and three fingers of a man. The Bird 9 wing-tip around and make flight impossible. So but
two of these small bones are free in our chicken's wrist,
although in the small chick several more (six in all) are
separate. separate. If we double back our fourth and fifth fingers arid
imagine that they have disappeared, extend our other
three fingers and then suppose that all our wrist-bones,
save two, have fused with the three long bones leading
to the base of our thumb, index and middle fingers,* we
will have an idea of the condition of our chicken's wing,
and indeed there is very little difference between this and
the wings of all other birds, f We have two separate
bones in our thumb, and three in each of the next two
fingers, and the bird has the same number, except in
its third finger, in which there is but one. The principal
value of this comparison is to show us that the bird,
even in its most characteristic and specialized organ, —
the wing, is not physically so unlike ourselves as we
might at first glance suppose. When a bird folds its
wing against its body, the joints are bent sharply, and
the Z, formed by the elbow and the wrist, almost closes
up. We can place our arm and hand in much the same
position. position. If we move our arms slowly up and down, little by
little greatly increasing the speed, we will realize how
much greater strength and rigidity the whirring wings FIG. 70. — Skeleton of wing of Condor, compared with Fig. 71. FIG. 70. — Skeleton of wing of Condor, compared with Fig. 71. FIG. 71. Wings — Skeleton of a man's arm; notice close correspondence of bones in the
two. (The extended thumb of the human hand is not silhouetted against
the background, and hence not very distinct.)
95 FIG. 71. — Skeleton of a man's arm; notice close correspondence of bones in the
two. (The extended thumb of the human hand is not silhouetted against
the background, and hence not very distinct.)
95 95 95 The Bird 9 of a hummingbird or a grouse require than do the slowly
flapping pinions of a gull or an albatross. When we
compare the relative shortness of the upper arm-bone,
or humerus, in the former groups with the long wing-
bones of the sea-birds, we again realize what exquisite
adaptations exist everywhere in Nature. The proportionate length of the various parts of the
fore limb of a bird forms an interesting coiollary to its
habits of life. For example, the hand in penguins and
in hummingbirds is very long indeed; while in the os-
trich the humerus is considerably longer than the fore-
arm and hand combined. (Compare Figs. 269 and 272.) Before we leave the wing-bones, it will be well worth
our time to consider for a moment how limbs first origi- ....., . . ...... . -^mmiim^ • •
^•^•iiVV;---- ' *
FIG. 72. — Diagram showing the origin of paired fins (limbs) from a continuous
fin-fold. (After Wiederscheim.) -^mmiim^ • •
^•^•iiVV;---- ' *
72. — Diagram showing the origin of paired fins (limbs) from a continuous
fin-fold. (After Wiederscheim.) nated. In the lowest of fishes, such as our friend the
Amphioxus and in lampreys, limbs are altogether absent,
but in embryos of sharks we get a hint of what the first
^reat fish-like forms were like. Along the lower part of
each side of the body there runs a continuous fin, so that nated. In the lowest of fishes, such as our friend the
Amphioxus and in lampreys, limbs are altogether absent,
but in embryos of sharks we get a hint of what the first
^reat fish-like forms were like. Along the lower part of
each side of the body there runs a continuous fin, so that The Framework of the Bird 97 7 97 The Framework of the Bi 97 the front view of a section would be something like this,
the dorsal fin being above and the lateral
fins on each side. * To the lower end of this are fused, in the bird, the bones which corre-
spond to our heel-bone and the small astragalus. The Bird 9 Wings Now owing to certain laws
of mechanics, whenever such a creature as this
moved about in the water, the stress of bal-
ancing would be thrown most heavily on two points in
these side fins, and gradually at these two nodes the fin
became more strongly developed; while between these
points it degenerated and finally disappeared. So in
modern fishes we find the quartet of limbs alone left of
this continuous fin or fold of skin. Look at a little embryo in the egg, taking one which
has been incubated for six or seven days, and see the
curious paddle or fin-like wings and feet — simply four
rounded flaps projecting from the body — as unlike the
limbs of the chick when it emerges from the egg as can
be imagined (see Fig. 367). The ridge or fin of skin in
the early, soft-backboned creatures could have been of
no use whatever, except in balancing. In fact if we
watch a trout carefully, we will see that it is the tail-fin
which does almost all the propelling, the front- and hind-
limb fins simply acting as guides and balances. So in this instance (as indeed in almost every organ
in ourselves as well as in birds) we learn that the original
function was entirely unlike that which the part now
serves. The idea of miraculous change, which is sup-
posed to be an exclusive prerogative of fairy-tales, is a
common phenomenon of evolution, and the shadows of
these miracles of the past are forever coming and going,
over the growth of the tiny bird hidden in the egg. The Bird * These correspond in ourselves to the bones of the upper foot and the
second row of carpals, this joint of the leg coming, therefore, really between
the bones of the ankle, instead of, as in ourselves, between the ankle and the
lower part of the leg. Legs The leg of our chicken, as we have seen, is attached
to the great bone of the thigh-girdle. Being used for
locomotion on land, the foot is not very different from
that of a lizard, but there seems something very strange
about the leg. Can it be possible that a chicken's knees
bend backward? If so, it must be different from all
other two-legged or four-legged creatures. Much of a
bird's leg is concealed beneath its feathers, and when we
see the bones as far up as the thigh-joint, we understand
our mistake at once, and see that a bird has knees which
bend in the same way as our own, that is, forward in an
opposite direction from the elbow. The knees of a bird
are usually concealed within the skin of the body, as in
the short-legged ducks, and are never visible outside
the plumage. Hence the wide-spread mistake concern-
ing them. For this reason the femur, or thigh-bone, is,
in birds, relatively very short, even in the long-shanked
herons and flamingos, the extra length of limb resulting
from the elongation of the next two lower joints. The thigh-bone, or femur, alone forms the upper leg,
or " second joint," and two bones, as in the forearm,
the next portion below. One of these, the tibia, is much
the larger and is the " drumstick" of the chicken.* When
we cut the dark meat from this portion, our knife some-
times slits off a splinter, which is the second bone of this
joint, the fibula. The Framework of the Bird The Framework of the Bird 99 In the chicken, we next come to a single long bone
called the tarsus, which is, in life, covered with scales. , in life, covered with scales. FIG. 74. — Human leg and foot. FIG. 73. — Skeleton of an Ostrich
leg and foot. FIG. 74. — Human leg and foot. FIG. 73. — Skeleton of an Ostrich
leg and foot. In the embryo it is composed of several separate bones.*
The simplest interpretation we can give to the foot of The Bird TOO the chicken, in comparing it with our own, is to imagine
that our small toe has completely disappeared; the great
toe (corresponding to the thumb of the forelimb) is
turned backward, and the heel is lifted high from the FIG. 75. — Living Ostrich, showing entire leg; notice the knee almost
within the body. FIG. 75. — Living Ostrich, showing entire leg; notice the knee almost
within the body. ground, the several bones of the upper foot being greatly
lengthened and fused into one. So we, like bears and
raccoons, walk with our whole foot, from toe to heel, flat
upon the ground, while a bird, like a cat or a horse, walks
on its toes alone. FIG. 76. — Heron, standing naturally upon its eight toes. FIG. 76. — Heron, standing naturally upon its eight toes. FIG. 77. — Jaguar, showing progression upon toes alone (digitigrade). (Sanborn, photo. Courtesy of N. Y. Zoological Society.)
101 FIG. 77. — Jaguar, showing progression upon toes alone (digitigrade). (Sanborn, photo. Courtesy of N. Y. Zoological Society.)
101 FIG. 78. — Wood Ib:s, resting temporarily upon its whole foot. (Sanborn, photo. Courtesy of N. Y. Zoological Society.) FIG. 78. — Wood Ib:s, resting temporarily upon its whole foot. (Sanborn, photo. Courtesy of N. Y. Zoological Society.) FIG. 79. — Bear, walking upon the whole foot (plantigrade). Compare with
Wood Ibis. 102 FIG. 79. — Bear, walking upon the whole foot (plantigrade). Compare with
Wood Ibis. 102 102 102 THE SKULL BIRD'S skull has been called a "poem in bone —
its architecture is the frozen music of morphol-
ogy; in its mutely eloquent lines may be traced
the rhythmic rhymes of the myriad amcebiform animals
which constructed the noble edifice when they sang
together." We should all "be able to whistle some bars
of the cranial song — the ptery go-palatine bar at least." BIRD'S skull has been called a "poem in bone —
its architecture is the frozen music of morphol-
ogy; in its mutely eloquent lines may be traced
the rhythmic rhymes of the myriad amcebiform animals
which constructed the noble edifice when they sang
together." We should all "be able to whistle some bars
of the cranial song — the ptery go-palatine bar at least."
We perhaps know that there are twenty-eight bones
in our own head, and if we attempt to dissect the skull of
a fish we will find many more, but at first glance the
skull of our chicken seems to be composed of but one
solid bone. Indeed, if we except the lower jaw and a
few others, such as the two little bones which unite it
to the skull, the entire cranium is soldered together, and
the lines of junction obliterated. In young birds these
seams are more or less visible, although the soldering
process begins very early. of the cranial song — the ptery go-palatine bar at least."
We perhaps know that there are twenty-eight bones
in our own head, and if we attempt to dissect the skull of
a fish we will find many more, but at first glance the
skull of our chicken seems to be composed of but one
solid bone. Indeed, if we except the lower jaw and a
few others, such as the two little bones which unite it
to the skull, the entire cranium is soldered together, and
the lines of junction obliterated. In young birds these
seams are more or less visible, although the soldering
process begins very early. The origin of the skull is wrapped in obscurity, and
neither the student of fossil bones, nor of those beneath
the skin of living creatures, nor yet the diligent watcher
of the mysterious panorama of life in the egg, can tell
us very much, although many theories have been sug-
103 en s The Bird 1 gested. THE SKULL The poet Goethe thought the skull was merely
a continuation of the neck-bones, very much expanded
and changed, and although the division of the skull into
three roughly outlined rings is possible, yet we have no
direct proof of the truth of this theory. direct proof of the truth of this theory. Fortunately, in the skulls of most animals, the bones
are separate, and by keeping in mind the constancy of
their position, the puzzle of the skull of a chicken begins
to clear up. Just as the first back-bone was a gelatinous or gristly FIG. 80. — Cranium of Dogfish, cartilaginous, generalized in structure. FIG. 80. — Cranium of Dogfish, cartilaginous, generalized in structure. one, so the old type of skull was entirely gristly or car-
tilaginous. Sometimes on the seashore near the huts of
the fishermen, we may pick up a strange-looking object —
translucent and looking as if it were made of hard white
rubber. Clinging to it is perhaps a long string of delicate
beads of the same substance. This is the skull and back-
bone of a dogfish or shark, and although the skull is
very unlike the chicken's cranium, yet many of the parts
in the latter are faintly foreshadowed in the cartilage
skull washed up by the waves. Through all the long ages of geological epochs, myri- The Sk 105 ads of creatures were changing in form and structure,
some growing too bulky and helpless and vanishing,
others developing powers of running, burrowing, flying
and leaping. But it is a very remarkable and wonder-
ful thing and very fortunate for us poor mortals, striving
after knowledge of the past, that in each general class
of creatures, certain ones should have found a niche
where they were removed from the fierce struggle for FIG. 81. — Skull of young Alligator. Bones massive and solid, adapting their
owner to an active aquatic life but to sluggish terrestrial movements; eye-
cavities and brain-case very small, the jaws (organs of prehension) composing
by far the major part of the head. FIG. 81. — Skull of young Alligator. Bones massive and solid, adapting their
owner to an active aquatic life but to sluggish terrestrial movements; eye-
cavities and brain-case very small, the jaws (organs of prehension) composing
by far the major part of the head. THE SKULL existence, and where for year after year, century upon
century, they and their descendants changed but little. We might mention Amphioxus and sharks among fishes,
Necturus among amphibians, Sphenodon among reptiles,
and the duck-billed mammal and others among hair-
covered creatures. These may be meaningless names, but
if one will read about them and then examine their skins
and skeletons in our museums, many a glance will be
given into the ages of the past, compared to which the existence, and where for year after year, century upon
century, they and their descendants changed but little. We might mention Amphioxus and sharks among fishes,
Necturus among amphibians, Sphenodon among reptiles,
and the duck-billed mammal and others among hair-
covered creatures. These may be meaningless names, but
if one will read about them and then examine their skins
and skeletons in our museums, many a glance will be
given into the ages of the past, compared to which the The Bird 1 few thousand years during which man has reigned seems
but a day. When we study the early structure of some creature,
say a bird, we find that before it emerges from the egg
the skull is soft and cartilaginous, open and quite differ-
ent in shape from what it will be eventually, and it is
most startling to find a living creature — a shark — with FIG. 82.— Skull of Bald Eagle. Bones light and spongy, fitting for a very active
aerial life; orbit very large and brain-case capacious, showing great advance
beyond reptilian condition. FIG. 82.— Skull of Bald Eagle. Bones light and spongy, fitting for a very active
aerial life; orbit very large and brain-case capacious, showing great advance
beyond reptilian condition. a skull which never gets beyond this condition. It is
as if the curtain of eternity had been, for a moment, drawn
aside for us, and a glimpse given into the past — a past
so remote and clouded that our keenest searches seem
to reveal but dim, skeletal forms of weird shapes, which
yet we know must have blended and imperceptibly
merged, through millions of years, into the present life
of the earth. a skull which never gets beyond this condition. THE SKULL It is
as if the curtain of eternity had been, for a moment, drawn
aside for us, and a glimpse given into the past — a past
so remote and clouded that our keenest searches seem
to reveal but dim, skeletal forms of weird shapes, which
yet we know must have blended and imperceptibly
merged, through millions of years, into the present life
of the earth. Looking at the chicken's skull as a whole, we notice
a number of uses which the various parts serve. The The Skull large rounded portion taking up most of the skull proper
is, of course, the box of bone which protects the brain. On each side, a large cavity shows where the eyes are
placed, and if we compare this skull with that of a cat
or dog or with that of a human being, we will see what
great importance eyes must be to a bird; the cavities
for them are so much larger than in other animals. Back 3r«5u nCd.se 3r«5u nCd.se
Oriit
ifa.Occ. Tin*
FIG. 83. — Skull of Fowl, showing orbit, brain-case, ear, lower jaw, premaxillary
(Pmx.), maxillary (Mx.), vomer (Vo.), lacrymal (Lc.), jugal (Ju.), palatine
(Pal.), pterygoid (.Ptg), quadrate (Qd.), and supra (Sup. occ.), ex (Ex. occ.), and basioccipital bones. (After Parker.) FIG. 83. — Skull of Fowl, showing orbit, brain-case, ear, lower jaw, premaxillary
(Pmx.), maxillary (Mx.), vomer (Vo.), lacrymal (Lc.), jugal (Ju.), palatine
(Pal.), pterygoid (.Ptg), quadrate (Qd.), and supra (Sup. occ.), ex (Ex. occ.), and basioccipital bones. (After Parker.) of each eye-case we see an irregular opening, the portal
of the ear; and in front of the brain two apertures in the
beak open toward the organ of the nose. The prominent
beak and wide-spreading lower jaw are chiefly concerned
in the procuring of food. We need not bother with the
names of all the bones, but there are some too interest-
ing and with too strange a history to be passed by. Let us glance at the back of the skull for a moment. Here we find a large round opening through which the The Bird io8 spinal chord passes into the brain, and below it is a
small knob, which in the living bird fitted into the first
vertebra of the neck. THE SKULL It is a very tiny projection of
bone, but fraught with significance: for if we look at
the skulls of a frog, a mouse, a cat, a horse or a man, we
will see that the head hinges upon two bony projections,
but in all birds and reptiles there is but one, — a very FIG. 84. — Rear views of bird (Hornbill) and mammal (Yaguarondi) skulls. Notice
single facet (occipital condyle) in the bird and two in the mammal, connecting
skull with the neck-bones. FIG. 84. — Rear views of bird (Hornbill) and mammal (Yaguarondi) skulls. Notice
single facet (occipital condyle) in the bird and two in the mammal, connecting
skull with the neck-bones. plain hint of the relationship of these two Classes, so dif-
ferent in external appearance. The head thus pivoting
upon a single point, the bird can turn its head much
farther around than if there were two points of attach-
ment. Before we leave this great opening, as the scien-
tists call it when they speak of it as the foramen magnum,
it may be worth while to mention the remarkably con-
stant position of the bones around it. Whether these
are all separate, or solidly fused into one, we may always plain hint of the relationship of these two Classes, so dif-
ferent in external appearance. The head thus pivoting
upon a single point, the bird can turn its head much
farther around than if there were two points of attach-
ment. Before we leave this great opening, as the scien-
tists call it when they speak of it as the foramen magnum,
it may be worth while to mention the remarkably con-
stant position of the bones around it. Whether these
are all separate, or solidly fused into one, we may always The Sk 109 know them by their position relative to the brain open-
ing; the upper edge of the hole is always formed by the
supraoccipital, the lower by the basioccipital, the two
sides by the exoccipitals. sides by the exoccipitals. Although many bones of the skull, such as the supra-
occipital, keep their names, whether found in salmon,
frog, lizard, bird, or man, others identical in position
have had new names given them. THE SKULL For instance, a small
bone directly in front of the eye is known as the lacrymal,
from its close relation to the tear-duct, but in fishes the
bone is called the preorbital, as a suggestion of fish-tears
would be rather absurd. We may find the dried ear-drum, or tympanum,
stretched tight across the entrance of the ear-cavities,
and if we break this, or even look carefully through the
transparent membrane, a long thin bone may be seen
beneath, extending backwards from the under surface of
the drum. This is the columella, or little column of
bone, and will have an interest for us later on. If we examine the way in which the upper and lower
mandibles or jaws are joined to the skull, we will find a
very ingenious arrangement; one very different from
that in ourselves. If the beak of a bird is to serve as
hand, lips, and mouth, it must be as free and movable as
possible, and instead of the upper jaw being fixed im-
movably to the skull, and the lower jaw swinging up
and down from it, we find that the upper jaw is attached
very loosely, while each side of the lower mandible hinges
upon a loose irregular-shaped bone, known as the quad-
rate. A long slender bone connects the quadrate with The Bird I 10 the upper mandible, which bone we may call the jugal. Indeed when we come to look closely at the quadrates
we find that they are very important, and in addition to
supporting the bar of bone from the upper jaw, and
pivoting the lower jaw, they bear another pan1 of bones FIG. 85. — Columella in ear of Snowy Owl (magnified 2 diameters). FIG. 85. — Columella in ear of Snowy Owl (magnified 2 diameters). extending inward from them, beneath the skull, to the
broad thin palate or roof of the mouth. In a chicken the individual movement of the upper
jaw is not very great, but in some birds, such as parrots,
it is much more noticeable. With a sharp knife we can
entirely detach the upper and lower jaws of most birds,
without cutting through a bone, the connection consist-
ing only of exceedingly tough tendons. THE SKULL When we found
our shark's skull we perhaps wondered what had become The Sk 1 1 1 of the jaws with the many rows of teeth, and we may
now guess that they were attached so loosely to the skull
that the action of the water washed them away with
the flesh. This was the case, and in all fishes we find
both jaws as separate bones. Among reptiles we find
the quadrate bone free only in snakes, an admirable
adaptation which enables them to swallow their prey
entire. The bones forming the palate, or roof of the mouth,
are of the greatest value in classification. No matter
how specialized the habits or the food of a bird may be,
the palate appears to be the last portion of its structure
to respond to any recent outside influences. Thus while
the absence or presence of a keel to the sternum is a
character of little value in separating the ostriches and
their allies from all other birds, yet the radical differ-
ence shown by the palate bones in the two groups is
reliable evidence of their early divergence from each
other. These taxonomic characters may be found in
any good book on systematic ornithology and need
not detain us here. Although we have the skull and both jaws of our
chicken, yet some very important and interesting bones
are lacking, and to find them we must find the tongue
of the bird. For a bird's tongue, as well as that of other
creatures, is not all flesh or horn, but underneath there
is a jointed framework of bone, which is called the hyoid. We may compare its shape to that of an arrow, with a
central head and four barbs, two very short and blunt
and two long jointed ones. The Bird 112 It may seem to us that the mandibles, the jugal, the
palate, the quadrates, the ear-bones, and the hyoid are
an unmeaning jumble of irregular bones, apparently
bearing no relation to one another,
and with absolutely no interest
outside the fact that each is very
well suited to its particular use. If scientists had studied only the
bones of adult animals, we might
have groped in vain for any an-
swer to the question of how these
bones came to be what they are. But the science of embryology, or
egg-life, has unfolded wonderful
things, and, as we shall soon see,
nothing more marvellous than the
strange story of these bones. The eel-like lampreys which It may seem to us that the mandibles, the jugal, the
palate, the quadrates, the ear-bones, and the hyoid are
an unmeaning jumble of irregular bones, apparently
bearing no relation to one another,
and with absolutely no interest that the mandibles, the jugal, the
the ear-bones, and the hyoid are
of irregular bones, apparently
bearing no relation to one another,
and with absolutely no interest
outside the fact that each is very
well suited to its particular use. If scientists had studied only the
bones of adult animals, we might
have groped in vain for any an-
swer to the question of how these
bones came to be what they are. But the science of embryology, or
egg-life, has unfolded wonderful
things, and, as we shall soon see,
nothing more marvellous than the
strange story of these bones. an unmeaning jumble
FiG.86.-Gill-basket of Lamprey. The eel-like lampreys which
crowd up our shallow brooks in
April to spawn are curious crea-
tures, and not the least remark-
able thing about them is the fact
that they have no jaws, although
they have an elaborate cartilagi-
FiG.86.-Gill-basket of Lamprey. noug net_WOrk protecting and
supporting the gills. We perhaps thought that every
vertebrate animal in the world had jaws of some kind, and
perhaps even lampreys had them long ago, before their
habit of sucking did away with any need for chewing. The Bird But the reason I have spoken of the lamprey is because The Sk 1 1 3 it brings vividly to mind the image of an animal which
must have once existed — a fish-like creature with no
jaws, but with a gristly mass which held up and pro-
tected the delicate blood-fringes, or gills, by means of
which all true fishes breathe. Our shark is a very convenient starting-point, and
before going further we should mention the technical name FIG. 87.— Gill-bars of Shark. FIG. 87.— Gill-bars of Shark. of this group — Elasmobranchs, or strap-gilled fishes. of this group — Elasmobranchs, or strap-gilled fishes. There are usually five of these gills, and within each
strap or fold of skin is a jointed arch of gristle. of this group — Elasmobranchs, or strap-gilled fishes. There are usually five of these gills, and within each
strap or fold of skin is a jointed arch of gristle. All this may be very true, say you, but what bearing
has it on the skull of the chicken? We have seen that in sharks the number of gills has
been greatly reduced, and a pair of very loosely attached
jaws has been acquired, — and the truth gradually dawns The Bird i upon us: the jaw of a shark is nothing but a greatly
changed gill-arch, which has doubled up, bent forward
and hinged to the skull. The skin has grown over the
edge, and the bony scales in the skin, standing up on
end, have become teeth. And now to our bird. In the embryo chick four
gill-arches are at first distinguishable, but these soon
begin to alter their position, to fade away, or to change
in some way, and in our bony skull we may trace them
as follows (see Fig. 89). The upper half of the first gill- FIG. 88. — Lower mandible, tongue, and hyoid bones of Bald Eagle. FIG. 88. — Lower mandible, tongue, and hyoid bones of Bald Eagle. arch forms the bones of the upper jaw, palate, jugal, and
quadrate, and the lower jaw completes the entire arch. The central part of the second gill fades into nothing,
but the top is present as the columella-bone of the ear,
while the base is transformed into the head and two
blunt barbs of the arrow-like bone of the tongue. The Bird The
two long barbs of this bone correspond to the third gill The.S 115 and, from their rod-like jointed character, they look
very much like the real gill-arches of a fish. The fourth
arch vanishes. Such is the almost incredible alchemy which Nature
has wrought from a plastic rod of gristle, — transforming
it into beak, tongue, and ears. Few of us, when watch- FIG. 89. — Ultimate distribution of the four embryonic gill-arches in the skull
of the adult bird. The dotted portions are not developed. (Adapted from
Newton.) Compare with Figs. 83 and 88. ing the gently waving gills of a fish, have realized how
much we indirectly owe to them. A noted German
anatomist — Karl Gegenbaur — believes that we owe even
our hands and arms (by way of the pectoral fins of fishes)
to portions of the gill framework, but this theory is not
generally accepted. ing the gently waving gills of a fish, have realized how
much we indirectly owe to them. A noted German
anatomist — Karl Gegenbaur — believes that we owe even
our hands and arms (by way of the pectoral fins of fishes)
to portions of the gill framework, but this theory is not
generally accepted. ORGANS OF NUTRITION N N other pages we shall consider some of the things
upon which birds feed, and shall see how surely
the methods used in the search and capture of
this food mould the bird's structure, modifying its form
from beak to toe ; and now is it not possible to find some-
thing of interest in the food after the bird swallows it? Indeed even before the swallowing takes place, if we
watch carefully we may notice something which we did
not before know. In the first place the bill of a bird is, of course, a
primary factor, not only in procuring food, but often in
killing and preparing and also holding it while it is being
made ready to swallow. Less confusion will result, how-
ever, if we leave the consideration of the beaks and bills
to a later chapter. After the bill (which corresponds to our mouth and
lips) come the glands of the mouth and here we again
enter the portals of physiology, — for some unknown
reason dreaded by many of us, and systematically shunned,
as dry and ultra-scientific. On the contrary there are
interesting facts awaiting us in all its branches. After
a brief consideration of the more important, we shall we sh Organs of Nutri 117 surely return to outdoor study of the daily life of these
creatures, with more balanced interest, and a "little
knowledge" which, instead of being a " dangerous thing,"
will, in this instance, add many fold to our appreciation
of the external results of these functions, whose work-
ings are ever concealed from the light of day. ings are ever concealed from the light of day. The digestive apparatus of a bird, or indeed of any
creature with a back-bone, is in reality a tube or canal,
which begins at the mouth and extends through the
body. Certain portions are contracted or expanded, and
specialized to store up, moisten, grind, dissolve, digest, or
absorb the food substances which pass through. The Salivary Glands The mucous membrane, or lining, of the entire diges-
tive canal is very delicate and requires to be kept con-
stantly moist. The lining of the mouth and throat, being
so exposed to contact with the outer air, requires some
special provision to lubricate it. This is accomplished
by certain glands, some beneath the tongue, others situ-
ated in the upper portion of the mouth. These are not
found in fishes, nor in other creatures which live alto-
gether in the water; but in reptiles several groups are
distinguishable. In birds they vary greatly, some having
scarcely a trace, while others have large well-developed
glands. Salivary is the common name given to certain
of these, and we will let that name represent all. will let that name represent all. In ourselves, saliva is an important aid in digestion. Besides moistening the food and softening all hard por-
tions, it exerts active chemical effects, as, for example, The Bird u8 changing starch to sugar and in many other ways making
ready the food, that the important changes which take
place in the stomach may begin at once. In birds, how-
ever, the saliva has but little chemical effect on the food,
its principal use being to moisten the substances before
they are swallowed. It is not often that Nature, when she has produced FIG. 90 — Nest of Chimney Swift; twigs glued together with saliva. FIG. 90 — Nest of Chimney Swift; twigs glued together with saliva. an organ or special tissue by the elaborate synthesis of
evolution, confines its use to any one function. If birds
were provided with salivary glands intended only for the
purpose mentioned above, they soon found other uses for
them. In a woodpecker we will find very large salivary
glands on each side of the mouth. These secrete a sticky
liquid which covers the long, many-barbed tongue and is
an efficient aid in picking out insects from their holes in
the bark and wood of trees. an organ or special tissue by the elaborate synthesis of
evolution, confines its use to any one function. If birds
were provided with salivary glands intended only for the
purpose mentioned above, they soon found other uses for
them. In a woodpecker we will find very large salivary
glands on each side of the mouth. The Salivary Glands These secrete a sticky
liquid which covers the long, many-barbed tongue and is
an efficient aid in picking out insects from their holes in
the bark and wood of trees. Organs of Nutri Organs of Nutri 119 Many birds carry in their beaks the grasses and
twigs with which they construct their nests, and if a sticky
fluid helped them to get their food, why would it not
also soften the twigs and make them easy to bend? Not
only this, but certain birds, such as our Chimney Swifts,
are provided with saliva in such quantities, and of such
tenacious consistency, that the entire nest — a mosaic of FIG. 91. — Nest of Esculent Swiftlet (edible bird's nest) composed entirely of saliva. FIG. 91. — Nest of Esculent Swiftlet (edible bird's nest) composed entirely of saliva. small twigs, each about an inch in length — is set in saliva
cement, and fastened to the vertical side of a chimney
or charred tree- trunk by the same means. Even this is
only a step, or link, in the direction of the extreme use-
fulness of saliva, for the little East Indian birds known
as Swiftlets (one of which ornithologists call Collocalia
fuciphaga) make their nests entirely of saliva or mucus. The second part of the scientific name, which means an
eater of seaweed, refers to the idea formerly held, that small twigs, each about an inch in length — is set in saliva
cement, and fastened to the vertical side of a chimney
or charred tree- trunk by the same means. Even this is
only a step, or link, in the direction of the extreme use-
fulness of saliva, for the little East Indian birds known
as Swiftlets (one of which ornithologists call Collocalia
fuciphaga) make their nests entirely of saliva or mucus. The second part of the scientific name, which means an
eater of seaweed, refers to the idea formerly held, that The Bird I the nests consisted entirely of half-digested seaweed, but
it has been proved that this is not the case. The homes
of these birds are the " edible birds'-nests " held in such
esteem by Chinese gourmands. The Tongue If we had a long series of birds' tongues before us,
we would be surprised at the great variety of shapes and
sizes. Observing the good-sized tongues of sparrows and
other small birds, we would turn to a pelican expecting
to see an enormous affair to correspond with the huge bill
of that bird. On the contrary we will find a tiny incon-
spicuous flap not larger round than a toothpick. This
reminds us of the condition of the tongue in some fishes,
where it is a very simple structure indeed. The king-
fisher also has a small tongue, and the same is true of
many other fish-eating birds, such as pelicans and most
of those which feed on large insects. The reason is ob-
vious. The food, which is swallowed whole, is of such
large size that a tongue of even moderate proportions
would be only in the way. The tongues of many birds are provided with oblique
series of teeth, either soft and fleshy or horny in struc-
ture, which point backward toward the gullet and must
be of great help to the bird in guiding and swallowing its
food. These teeth are especially abundant around the
glottis, or opening to the windpipe, guarding it from
chafing or from the chance of food choking it up. In ducks and geese we find tongues large and of com-
plicated appearance. The edges are often toothed or Organs of Nutri I 2 1 fringed to correspond with the serrated or otherwise
indented edges of the mandibles. One which is before me
as I write is very elaborate. It is that of a wild Mallard
Duck. At the tip is a thin, distinct flap or lamella, horny
and with smooth edges. Behind it the tongue enlarges
abruptly into a thick oblong mass, deeply grooved down FIG. 92. — Bill of Brown Pelican, showing extreme reduction of tongue in a bird
which swallows whole fish. FIG. 92. — Bill of Brown Pelican, showing extreme reduction of tongue in a bird
which swallows whole fish. the center. The edges of the anterior half are fringed
with a double line of horny hairs, while in the posterior
portion the upper line is replaced with tooth-like struc-
tures. The upper surface is smooth in front, but farther
back two central folds arise and curve over laterally,
forming tube-like grooves. The Tongue Still more posteriorly, fleshy
recurved teeth are visible, singly, in groups, or in regular
lines. The tongue of our common barnyard duck is the center. The edges of the anterior half are fringed
with a double line of horny hairs, while in the posterior
portion the upper line is replaced with tooth-like struc-
tures. The upper surface is smooth in front, but farther
back two central folds arise and curve over laterally,
forming tube-like grooves. Still more posteriorly, fleshy
recurved teeth are visible, singly, in groups, or in regular
lines. The tongue of our common barnyard duck is The Bird 1 similar to this and is well worth examining. The use of
such a complicated organ in a bird of so simple feeding
habits as the duck is hard to explain. We will hardly find two tongues that are alike, and
even the tips differ, and show as wide a range of varia- FIGS. 93 and 94. — Top and side views of the tongue of a Mallard Duck, showing
complicated structure in the tongue of a bird which sifts its food from the mud. FIGS. 93 and 94. — Top and side views of the tongue of a Mallard Duck, showing
complicated structure in the tongue of a bird which sifts its food from the mud. tion as the remaining portions. In many birds, such as
owls, larks, and swifts, the tip is bifid, or double-pointed,
bringing to mind the forked tongues of snakes and cer-
tain lizards. In woodpeckers the tongue is round and
exceedingly long, and can usually be thrown out some
distance beyond the tip of the bill. Our common Flicker, or Golden-winged Woodpecker, Organs of Nutrition possesses a tongue of remarkable length, even for a wood-
pecker, and while feeding, the bird will often shoot it FIG. 95. — Head of Flicker, showing tongue slightly protruding FIG. 95. — Head of Flicker, showing tongue slightly protruding out two or three inches beyond the tip of the beak. Easily and without a hitch it disappears again, appa- FIG. 96. — Skull of Flicker, showing rear branches of the hyoid bone, curving up
over the skull and down into the right nostril. The front of the tongue is
visible beyond the tip of the beak. FIG. 96. The Tongue — Skull of Flicker, showing rear branches of the hyoid bone, curving up
over the skull and down into the right nostril. The front of the tongue is
visible beyond the tip of the beak. rently down the very throat of the bird. If we carefully
remove the skin from the skull of a dead Flicker, the
magic will become plain. When we spoke of the skull rently down the very throat of the bird. If we carefully
remove the skin from the skull of a dead Flicker, the
magic will become plain. When we spoke of the skull The Bird I 2 of a bird, mention was made of the two long bones which
branched out from the rear of the tongue and which are
all that remain of the third ancestral gill-arch. In the
Flicker, the slender, white tongue divides into these two
branches just in front of the glottis and from here they
extend backward, passing one on each side of the wind-
pipe, and on upward, following the curve of the skull,
then forward, lying together upon the forehead. Not
even here do they end, however, but actually reach some
distance into one nostril! So when this bird stretches
out its tongue, the tips of the rear branches leave the
opening of the nose and shoot around over the surface
of the skull until they have gone as far as possible. No
wonder the poor ants have but little chance when a
Flicker visits their hill and sets the marvellous mechanism
of his tongue rapidly to work. And no wonder the
enthusiasm of an ornithologist never fails, when he thinks
of the scores of similarly interesting structures still await-
ing investigation. ing investigation. The tip of the tongue in the sap-sucking woodpeckers
is beset with numerous hairs forming a brush-like instru-
ment, but spines take the place of hairs in the species
which feed exclusively on insects. It is known that the
exact proportion of insects in the diet of any particular
kind of woodpecker is reflected in the more or less per-
fect adaptation of the minute structure of its tongue to
that end. In the sapsuckers, too, the tongue is comparatively
short, doubtless because the sap flows readily from the
holes which these birds bore. The Tongue Hence they require no Organs of Nutrition such extension of tongue as the deep burrows of the
ants necessitate in the case of the Flicker. Thus the tongue of a bird seems a very unstable
character, acted upon quickly and radically by any
change in the diet of the species. The entire tip of the
tongue is frequently frayed out into a kind of brush,
remarkably developed in
the parrot-like lories. Yet
this curious structure is
probably only an elonga-
tion of the papillae, hom-
ologous with those which
make the tongue of a cat
or lion so rough. Cocka-
toos, although first cousins
to the lories, have very
different tongues, thick and
fleshy with club-shaped
tips. Thus the tongue of a bird seems a very unstable
character, acted upon quickly and radically by any
change in the diet of the species. The entire tip of the
tongue is frequently frayed out into a kind of brush,
remarkably developed in rd seems a very unstable
ly and radically by any
ies. The entire tip of the
out into a kind of brush,
remarkably developed in
the parrot-like lories. Yet
this curious structure is
probably only an elonga-
tion of the papillae, hom-
ologous with those which
make the tongue of a cat
or lion so rough. Cocka-
toos, although first cousins
to the lories, have very
different tongues, thick and
fleshy with club-shaped
tips. FIG. 97. — Thick fleshy tongue of Cockatoo, tips. In our common gold-
finch, the sides of the
tongue curl inward, form-
ing an admirable seed-
scoop, while the same or-
distinctly cleft into sev-
been likened to a "four-
s little owner impales the
which it so industriously
he great particoloured bill
curious tongue — a long tips. In our common gold-
finch, the sides of the
tongue curl inward, form-
ing an admirable seed-
scoop, while the same or-
gan in the chickadee, being distinctly cleft into sev-
eral prongs at the tip, has been likened to a "four-
tined pitchfork" on which its little owner impales the
myriad grubs and insects for which it so industriously
searches twigs and leaves. The great particoloured bill
of a toucan conceals a very curious tongue — a long
FIG. 97. — Thick fleshy tongue of Cockatoo, FIG. 97. The Tongue — Thick fleshy tongue of Cockatoo, scoop, while the same or-
gan in the chickadee, being distinctly cleft into sev-
eral prongs at the tip, has been likened to a "four-
tined pitchfork" on which its little owner impales the
myriad grubs and insects for which it so industriously
searches twigs and leaves. The great particoloured bill
of a toucan conceals a very curious tongue — a long The Bird 126 thin affair, narrow throughout its whole length and so
thickly bordered with a deep, delicate fringe that it
bears a decided resemblance to a feather. FIG 98. — Feather-like tongue of Toco Toucan. FIG 98. — Feather-like tongue of Toco Toucan. The tongue of a flamingo is thick and fleshy, filling
the entire cavity of the lower mandible and in shape
reflecting its crookedness. The upper edges of the man- • •»
FIG. 99. — Tongue of Flamingo within lower mandible (natural position), FIG. 99. — Tongue of Flamingo within lower mandible (natural position), dible approach each other so closely that they perma-
nently inclose the tongue, motion being only possible in Organs of Nutri 1 27 a forward or backward direction. Along the sides of the
tongue are two series of fleshy teeth, in shape exactly
like the poison-fangs of a rattlesnake. like the poison-fangs of a rattlesnake. Mention must be made of the unique and greatly
specialized tongues of Hummingbirds and Honey Creepers. The outer edges of this organ are curled into two tubes, FIG. 100. — Side view of Flamingo's tongue, forcibly lifted above mandible, shopp-
ing fleshy recurved teeth. FIG. 100. — Side view of Flamingo's tongue, forcibly lifted above mandible, shopp-
ing fleshy recurved teeth. which are more or less split and frayed near the tip,
forming delicate brushes — efficient instruments either to
suck up nectar or to flick out insects from the heart of
corollas. The Crop From the back of the throat to the stomach extends
a tube, the gullet or oesophagus, through which the food .3
a .3
a .3
a Organs of Nutri 129 descends after it leaves the mouth. In some birds this
is a simple tube of the same diameter throughout, always
moist from the secretion of (mucous) glands which are
found abundant in its walls, but serving merely as a
passage for the food on its way to the stomach. passage for the food on its way to the stomach. In another class of birds an enlarged chamber is
present, called the crop. This serves a somewhat simi-
lar purpose as the external pouch of the pelican; that
is, it acts as a receptacle for food. No especial digestive
glands are found here, and the only agents acting on FIG. 102.— Brush, or tube-like tongue of Honey Creeper. Twice natural size. FIG. 102.— Brush, or tube-like tongue of Honey Creeper. Twice natural size. the food are water, the secretions of the salivary glands,
and the heat of the bird's body. The crop exists only
superficially in some birds, the dilation being hardly
noticeable. From these we find a succession of more distinctly
marked permanent crops, until in grain-eating birds this
organ is very prominent. If we examine an English
Sparrow after it has made a hearty meal in the chicken-
yard, we will find the crop filled with grains of wheat,
some cracked in pieces by the bill, others entire. The Bird 1 When we sought the extreme in the provision of saliva
in birds, we had to refer to a swift, living in caves in
islands of the Malay Archipelago, but to find the highest
degree of development of crops is a much easier matter. In the ordinary pigeon the crop is of very great size
and divided into two lobes. The capacity of the crop
in some birds of this class is astonishing. As many as
sixty-three acorns have been found in the crop of the
English Wood-pigeon. If we look at the crop of a pigeon
before its young leave the nest, we will discover a func-
tion of this organ which would otherwise never be sus-
pected. We know that herons and some other birds
feed their young on fish half -digested by themselves. This process is known as regurgitation. The Crop If we have ever
seen a pigeon with the beak of its young half down its
throat, pumping something into the offspring's mouth,
we have probably thought that a similar habit was being
shown, — half -digested grain taking the place of the
heron's fish. But such is not the case. At the time
of the breeding season, the folds of membrane in the
crops of both parent pigeons thicken and secrete or
peel off in curdy cheesy masses — "pigeon's milk" some
call it — and this forms the food of the young birds So
in pigeons the crop not only receives food, but at times
provides it. Now for a glance at some of the oddities in the struc-
ture of crops. The Hoatzin — a strange bird of Brazilian
swamps — which harks back to its reptilian ancestors in
many ways, has a very curious crop. There are strong
muscles in its walls, the use of which, it is said, is to Organs of Nutri 131 squeeze out the juice of the thick leaves of the Arum
arborescens which forms its food. Thus it has a gizzard-
like function, and has become so important in the life-
economy of the bird that it has developed out of all
proportion, and occupies so much space that the keel
on the breast-bone has had to give way in part to make
room for it, and even the arms of the " wish-bone" have
been bent outward. In this remarkable bird the pro-
ventriculus and gizzard are reduced, their functions being
usurped by the crop. usurped by the crop. The facility with which most birds are able to eject
the contents of their crops serves several useful pur-
poses, besides the feeding of the young of herons, cor-
morants, and others. When vultures have gorged them-
selves to repletion on the flesh of any animal, they usually
retire to some near-by retreat and sleep until digestion
has taken place. But if they are suddenly approached
or alarmed, they will instantly eject all they have swal-
lowed and, thus lightened, take safety in flight. Pelicans
and Wood Ibises also have the habit of ' unswallowing '
their fishy meals when frightened. The Crop Petrels and many
fish-eating sea-birds appear to have a supply of oil always
in readiness, which they shoot from the mouth to a con-
siderable distance, surving as an efficient means of self-
defence when taken in the hand after being wounded. In birds of prey generally, but especially in owls,
another use for this habit is found. Owls always swallow
their smaller prey entire, sometimes crushing the skulls
of mice and plucking out the longer wing and tail-feathers
of birds. Although their crops are not nearly so mus- The Bird 132 cular as that of the Hoatzin, yet there must be powerful
movements of the walls, for the mice and birds are de-
nuded of hair and feathers and even the bones are in
some way removed from the body, and all are ejected
in a neat oval ball. If we find some hollow tree where an owl has its FIG. 103. — Keel of Hoatzin, repressed by crop. FIG. 103. — Keel of Hoatzin, repressed by crop. regular sleeping-place, many of these pellets may be
found on the ground below, showing the results of each
night's hunting. The skulls in them are often in such
perfect condition that the species of rodents may be
identified. Besides allowing the bird conveniently to get
rid of the indigestible portions of its food, this habit
seems to be necessary to the health of the bird. In Organs of Nutrition captivity, owls and hawks are never so healthy and active
when fed on fleshy meat alone, as when a dead mouse or
sparrow, rat or pigeon is given occasionally. In dissecting
specimens which have had nothing but a flesh diet for a
year or more, I have found the throat and gullet in a very
bad condition, as if the lack of some scouring process, FIG. 104. — Food-pellets ejected by Great Horned Owl, containing
remains of rodents. FIG. 104. — Food-pellets ejected by Great Horned Owl, containing
remains of rodents. such as is afforded by the pasage up and down of the
indigestible hair and feathers, had actually resulted in
the death of the bird. I have seen owls try to eat the
straw on the floors of the cages, when not provided with
food in the condition in which they find it when at lib-
erty. The Crop In cormorants and birds of similar voracious fish-eating
habits, the entire gullet serves as a receptacle for food,
while the fish first swallowed are undergoing the process The Bird '34 of digestion lower down. Here, as in many other in-
stances, we have a condition very similar to that in some
reptiles — crocodiles in particular. These ravenous scaly
creatures have such powerful organs of digestion that even
bones are dissolved, but the stomach is comparatively
small, and when a crocodile makes a large meal, it is at
first stored away in the wide gullet. The Stomach and Gizzard In the present chapter we might easily be led into
details which would strand
us in the midst of dry
technicalities, but we will
try to avoid all this and
choose only the interesting
facts. FIQ. 105. — Caracara, showing crop dis-
tended with food The chief organ of di-
gestion, in birds as in other
animals, is of course the
stomach. In many fish-
eating birds this organ is
merely a simple, more or
less enlarged chamber, rather
crop-like except that it con-
tains numerous digestive
lands. FIQ. 105. — Caracara, showing crop dis-
tended with food FIQ. 105. — Caracara, showing crop dis-
tended with food
glands. The typical bird-stom-
ach, however, is compound,
or formed of two more or less distinct parts. The first Organs of Nutrition portion — known as the proventriculus — is the smaller, and
contains very active digestive glands, sometimes ar-
ranged in patches, but more usually forming a band. If
the lower part of the oesophagus of an English Sparrow
is removed, slit open and washed, these glands can be
easily seen, being more of a rose tint than the paler tissue
of the portion nearer the mouth. The walls are thicker
in this glandular area. This can be seen to better ad- portion — known as the proventriculus — is the smaller, and
contains very active digestive glands, sometimes ar-
ranged in patches, but more usually forming a band. If
the lower part of the oesophagus of an English Sparrow
is removed, slit open and washed, these glands can be
easily seen, being more of a rose tint than the paler tissue
of the portion nearer the mouth. The walls are thicker
in this glandular area. This can be seen to better ad-
vantage in a young chicken,
where the glands take the
form of conical protuber-
ances which dot the entire
surface. Nature has pro-
duced curious modifications rtion — known as the proventriculus — is the smaller, and
ntains very active digestive glands, sometimes ar-
ranged in patches, but more usually forming a band. If
e lower part of the oesophagus of an English Sparrow
removed, slit open and washed, these glands can be
ily seen, being more of a rose tint than the paler tissue
the portion nearer the mouth. The walls are thicker
this glandular area. The Stomach and Gizzard This can be seen to better ad-
vantage in a young chicken,
where the glands take the
form of conical protuber-
ances which dot the entire
surface. Nature has pro-
duced curious modifications
of this typical fore-stomach,
as in snake-birds, whicn have
the glands of this portion en-
closed in a sac, in shape not
unlike a small crop. Here
the food is softened and acted
106.— Glands of the stomach of a upon chemically by the secre-
young chicken. ,. ,,
tions from the walls. portion — known as the proventriculus — is the smaller, and
contains very active digestive glands, sometimes ar-
ranged in patches, but more usually forming a band. If
the lower part of the oesophagus of an English Sparrow
is removed, slit open and washed, these glands can be
easily seen, being more of a rose tint than the paler tissue
of the portion nearer the mouth. The walls are thicker
in this glandular area. This can be seen to better ad-
vantage in a young chicken,
where the glands take the
form of conical protuber-
ances which dot the entire
surface. Nature has pro-
duced curious modifications
of this typical fore-stomach,
as in snake-birds, whicn have
the glands of this portion en-
closed in a sac, in shape not
unlike a small crop. Here
the food is softened and acted
FIG. 106.— Glands of the stomach of a upon chemically by the secre-
young chicken. ,. ,,
tions from the walls. is removed, slit open and washed, these glands can be
easily seen, being more of a rose tint than the paler tissue
of the portion nearer the mouth. The walls are thicker
in this glandular area. This can be seen to better ad-
vantage in a young chicken,
where the glands take the
form of conical protuber-
ances which dot the entire
surface. Nature has pro-
duced curious modifications
of this typical fore-stomach,
as in snake-birds, whicn have
the glands of this portion en-
closed in a sac, in shape not
unlike a small crop. Here
the food is softened and acted
FIG. 106.— Glands of the stomach of a upon chemically by the secre-
young chicken. ,. ,,
tions from the walls. The Stomach and Gizzard The second division of the stomach is the gizzard,
an organ made to perform most powerful compressing
motions, thus crushing and macerating the food, so that
when passed on into the intestine, every particle of nour-
ishment may be extracted from it. When we think of
beauty of colouring in birds, it is their plumage which
at once presents itself to the mind, and yet a gizzard
has a real beauty both of shape and hue. This organ, FIG. 106.— Glands of the stomach of a
young chicken FIG. 106.— Glands of the stomach of a
young chicken The second division of the stomach is the gizzard,
an organ made to perform most powerful compressing
motions, thus crushing and macerating the food, so that
when passed on into the intestine, every particle of nour-
ishment may be extracted from it. When we think of
beauty of colouring in birds, it is their plumage which
at once presents itself to the mind, and yet a gizzard
has a real beauty both of shape and hue. This organ, The Bird i 3 in a chicken, is in shape like a double convex lens. The
cavity in the centre is lined with a tough yellow membrane,
sometimes almost as hard as bone. Two great tendons
spread over the outer surface on each side, and although
in life forever buried in the absolute darkness of the
bird's body, yet when brought into sunlight they shine
with an iridescence like the beam from a spectrum. It is hardly possible for the gizzard to grind up food
in the sense of having much lateral motion, like the move-
ment of the jaws in chewing, but it shuts together again
and again with great force. Gravel and sharp stones are
swallowed by many birds, and are of great importance
in helping to grind the food. The number and size of
these stones are sometimes almost beyond belief. I
have known a cassowary to swallow over a quart of rubble
in one day, and have given a quartz pebble twice as large
as a hen's egg to one of these birds and watched it slip
down the bird's throat as easily as a cube of carrot. This
particular bird preferred smooth white quartz pebbles,
and would search through a whole heap, picking out stones
of this character. The Stomach and Gizzard The same preference was exhibited
by the gigantic extinct birds of New Zealand called moas. Mr. Frederick Chapman, writing of a portion of New
Zealand where the skeletons of moas were found in great
abundance, says: "When we came upon the ground
disturbed by the wind (the soil being shifting sand)
we soon found a number of distinct groups of gigantic
gizzard-stones. It was impossible to mistake them. In
several cases they lay with a few fragments of the heavier
bones. In all cases they were in distinct groups; even Organs of Nutri 137 where they had become scattered, each group covered
only a few square yards of ground, and in that space
lay thickly strewn. . . . The peculiar feature of the
stones was that they were almost all opaque, white quartz
pebbles. In one place I found a small group of small
pebbles of different colour, more like the few brown water-
worn pebbles which may be picked up hereabouts. These
lay with a set of bones much smaller than the very large
bones I found with most of the clusters of pebbles. "I did not gather these brown pebbles, as I thought
it uncertain whether they were gizzard-stones or not,
though it is possible that the species to which the smaller
stones belonged was not so careful in selecting white
stones. "A glance at the pebbles lying around in the sur-
rounding country showed that the quartz-pebbles were
not collected here. . . . Mr. Murdock and I collected
three sets of pebbles, and these I can pronounce com-
plete, or nearly so. It is beyond question, too, that each
set belongs to a distinct bird. No. 1 weighs 3 Ib. 9 oz.;
No. 2 weighs 4 Ib.; while No. 3 weighs no less than 5 Ib. 7 oz. ! This giant set contains individual stones weighing
over 2 oz.; indeed, I have picked out eight stones weigh-
ing almost exactly 1 pound." ing almost exactly 1 pound."
The gizzard of a bird is reflective of its diet, and is
very quickly affected by any change in the food. For
example, a captive gull when fed exclusively on fish
has but little muscular power in the gizzard, but a diet
of grain will produce a change in that organ, giving it
grinding power sufficient to crush the kernels of corn. 1 The Bird That this is something more than an abnormal con-
dition brought about by artificial means is proved by
the fact that in the Orkney Islands the wild gulls feed
in winter, spring, and summer on fish, and at this time
are gizzardless; but in the fall they change to a diet of
corn and develop a very respectable gizzard. So we
see that this organ, apparently so independent in func-
tion and individual in appearance in many birds, is in
reality only a physiological change from the stomach
proper. proper. The history of the development of this organ may
be traced in various living species, from the soft mem-
branous sac of a fish-eating bird to the knot of tendons
which forms the gizzard of certain Fruit-pigeons. These
birds feed on nutmegs and other very hard, almost stony
nuts, and to enable the bird to crush these, the lining of
the gizzard is covered with several score of conical pro-
jections, horny in consistence. These are probably the
nearest approach to " hen's teeth "we are likely to find. nearest approach to " hen's teeth "we are likely to find. What a boon to a business man who indulges in a
daily " fireman's lunch/' if his masticatory function
could be an internal and unconscious one, as in a bird! A crocodile, which has so much in common with a
bird, is provided with a gizzard, which, like that of a
chicken, is round, muscular, and has two great side ten-
dons, and no less than five pounds of grinding-stones
have been found in one of these reptiles. Many interesting adaptations are found in the stom-
achs of birds, made necessary by special requirements
in the diet. As an instance of this, the snake-bird has Organs of Nutri I 39 a dense mat of hair at one end of the stomach, the free
ends of which point outward, brush-like, and prevent
the accidental entrance of any small fish-bones which
otherwise might get into the small intestine. The giz-
zard of a cuckoo, when opened, often gives the impression
of a similar coating of hair lining the entire organ, but
these are in reality only the hairs of caterpillars upon FIG. 107. — Cluster of matted hairs in the stomach of a Snake-bird. FIG. 107. — Cluster of matted hairs in the stomach of a Snake-bird. The Bird which these birds feed, which have become detached
and have lodged in the folds of the gizzard lining. When considering the crops of birds we noticed the
curious way in which a pigeon feeds its young, by re-
gurgitating a cheesy substance which forms in its crop,
and we will now speak of something still more remark-
able. The strange nesting habits of the hornbills are
foreign to this volume, but we cannot leave the subject
of gizzards without touching on the manner in which
the male birds of this group probably feed their impris- The Bird 1 oned mate and young. I say probably, because no one
has seen them do this, but as in captivity the operation
occurs repeatedly during the breeding season, there can
be but little doubt concerning its evident significance. After walling up his mate and her egg in some hollow
tree, the male hornbill takes upon himself the labour of
supplying her with food throughout the period of incu-
bation and the subsequent rearing of the young bird. Instead of bringing food piecemeal, — nut by nut, grape
by grape, — the lining of the entire gizzard peels off at
certain frequent intervals, appearing, when ejected at
the mouth, like a small bag or purse, the puckered open-
ing (heightening the simile) serving to retain securely
the contents of the gizzard, — a dozen or score of grapes
or other fruit. This, the male bird, in his native land,
doubtless takes in his beak to the tiny opening of the
walled-up nest and delivers into the bill of his mate. How admirable a spouse this, who not only seeks and
provides sufficient food for his temporarily helpless
family, but bears it to them wrapped in a packet torn
from his very body — if not a "pound of flesh," at least
enough to make a lunch-basket! The Intestines Beyond the gizzard is the intestinal canal, which
varies greatly in length in different birds. The ostrich
has forty-six feet of this digestive tube, while the nectar
and tiny insects snatched by a hummingbird in its flight
are digested in a delicate hair-like duct but two inches in
length. Although comparatively of such great length, the Organs of Nutri 141 way in which this part of the digestive tract is coiled and
twisted in the body cavity of the bird allows it to take
up the least possible amount of room. up the least possible amount of room. The function of this long tube is to absorb the nutri-
ment from the food after this has been moistened by
the salivary glands, crushed by the gizzard, and acted
on by the stomach acids, and secretions from the liver
and pancreas. The digestible parts are then taken up
by the blood through the walls of the intestine. In
many of the lower types of birds, such as the cassowary,
ostrich, and screamer, the arrangement of this long
digestive tract is very simple, much like the condition
to be found in alligators. THE FOOD OF BIRDS HE organs and physiological functions of a bird,
as of animals generally, are so interrelated and
intimately dependent on each other that it is a
rather difficult matter to consider any single one by
itself without being led into another's province. For
example: we have for the subject of this chapter the
food of birds, and unless we are very careful, we shall
overstep the bounds of our theme. To limit our subject
clearly we will consider only adult birds. H clearly we will consider only adult birds. We have all seen the pestiferous sparrows picking
up grain in the chicken-yard; we have admired the skill
which the red-breasted robin exhibits in spying and
extracting earthworms on our lawns; our memory re-
calls the osprey dropping upon his fish, and the wood-
pecker chiselling to the wood-borer; but did we ever stop
a while and attempt a "bird's-eye view" of all the classes
of substances which birds find good as food? The ways in which this food is sought and caught,
killed and prepared are wonderfully varied, and some
idea of the remarkable variety of substances laid under
contribution as food by birds of different orders may
be had from a brief review of the principal divisions
142 ivisi
142 The Food of Bi 143 into which these substances are classified, and the part
they take in supplying birds with food. As with all animals, certain mineral salts are very
necessary to a bird's existence, such as the substances
from which the calcium phosphate for the bones, and
the calcium carbonate for the shells of the eggs, are de- FIG. 108. — Finch, a bird with heavy, thick bill adapted for crushing seed. FIG. 108. — Finch, a bird with heavy, thick bill adapted for crushing seed rived. The gravel and pebbles swallowed by birds in
the course of their daily feeding should hardly be men-
tioned in this connection, as this is only done for the
mechanical assistance, derived from the hard surfaces,
in triturating the food. Vegetable-feeders form a large group among birds, and
they alone would offer an interesting field for study, as The Bird 1 there is such specialization for feeding on particular
varieties or portions of plants. We find fruit- and grain-
eaters, besides those which feed almost entirely on buds,
leaves, berries and nuts, nectar, sap, and even pollen. THE FOOD OF BIRDS Lichens form a considerable item in the bill of fare of
ptarmigans, the Arctic grouse. We have even dedicated
certain plants to birds which show a decided partiality
for them, — duckweed and partridge-berry. FIG. 109. — Vireo, an insect-hunter, with a delicate, hooked bill. FIG. 109. — Vireo, an insect-hunter, with a delicate, hooked bill. There is no doubt that a great many plants benefit
from the cross-fertilization of their flowers by humming-
birds carrying the pollen from blossom to blossom. Of
one of the sugar-birds of South Africa it is said: "When
sucking up the nectar of one of the larger protea-blossoms,
the bird perches on the edge of the flower, plunges its
long bill and the greater part of its head downwards
among the petals, and retains it in this position until The Food of Birds satisfied. As a result the narrow, shaft-like feathers of
the forehead frequently become saturated and stained
with juice and dusted over with pollen, and it is probable
that this bird plays an important part in the cross-fer-
tilization of several species of protea." tilization of several species of protea."
Desmids and diatoms, those one-celled microscopic
organisms which are almost on the border-line between FOOD OF VARIOUS BIRDS. FIG. 110. — Sea-urchin. FIG. 110. — Sea-urchin. plants and animals, I have found in large numbers in
the digestive tracts of ducks and other birds which are
accustomed to find their food by sifting the mud at the
edges of ponds and lakes. plants and animals, I have found in large numbers in
the digestive tracts of ducks and other birds which are
accustomed to find their food by sifting the mud at the
edges of ponds and lakes. Sponges, at least in a decayng state, are devoured by
crows, as I can testify from observation after dredging
expeditions in the Bay of Fundy. The Bird The Bird 146 We should scarcely think that those watery creatures
sea-anemones, hydroids, and jelly-fish (some of the latter FIG. 111.— Caterpillar. FIG. 111.— Caterpillar. consisting of over 95 per cent water) could afford much
nourishment to any animal, and when crows and gulls
are seen tearing large stranded jellies into pieces, it is consisting of over 95 per cent water) could afford much
nourishment to any animal, and when crows and gulls
are seen tearing large stranded jellies into pieces, it is FIG. 112.— Cocoon. FIG. 112.— Cocoon. FIG. 112.— Cocoon. probably only for the sake of the semi-parasitic shrimps
which make their home in the interior canals of the The Food of Bi 147 masses of animate gelatine. But the fresh-water hydra,
belonging to the same division as the hydroids, is eaten
in myriads by ducks and geese. These and many other
birds are remarkably fond of duckweed, which they
devour with such evident pleasure that they must enjoy
it as much as cats do catnip, or canaries hempseed. As
the under surface of these small water plants is the fa- FIG. 113.— Butterfly with wing torn by bird. FIG. 113.— Butterfly with wing torn by bird. vourite home of the hydra, they necessarily form a por-
tion of the food of these water birds. Roundworms, flatworms, and leeches are devoured
by many aquatic birds, while earthworms form a staple
article of diet with such different species as thrushes,
woodcocks, and cranes. A favourite morsel of the curi-
ous apteryx of New Zealand is a gigantic species of worm,
twelve to twenty inches in length, which is highly phospho-
rescent. The apteryx seeks its food by night, and when The Bird 1 devouring one of these worms, the whole bird is lighted
up, and after its meal the bird's bill is illumined by the
mucus which adheres to it. mucus which adheres to it. Starfish and sea-urchins are sought out by crows,
ravens and gulls, and perhaps other birds. They break
into them by main force, or
else carry them to a height
and drop them on the rocks. I have even seen a Bald
Eagle, when fish and Fish-
hawks were scarce, deliber-
ately break into and devour
a green-spined urchin. If, as is said, immense bow-
head whales subsist entirely
FIG. 114.-Snail. The Bird shrimps,
then it is not surprising that many thousands of shore-
birds are well nourished by the myriads of shrimps and
prawns, large and small, which every tide leaves exposed. Starfish and sea-urchins are sought out by crows,
ravens and gulls, and perhaps other birds. They break
into them by main force, or
else carry them to a height
and drop them on the rocks. I have even seen a Bald
Eagle, when fish and Fish-
hawks were scarce, deliber-
ately break into and devour
a green-spined urchin. FIG. 114.-Snail. a green-spined urchin. If, as is said, immense bow-
head whales subsist entirely
FIG. 114.-Snail. shrimps,
then it is not surprising that many thousands of shore-
birds are well nourished by the myriads of shrimps and
prawns, large and small, which every tide leaves exposed. FIG. 114.-Snail. It is a mere truism to say that insects form the sole
food of scores of species of birds, and enter into the diet
of many hundreds. It has been said that without birds,
within a space of ten years, the earth would not be habit-
able for man, owing to the unrestricted increase of nox-
ious insects. There is doubtless not a single group of in-
sects which does not suffer from the appetite of one or
more species of bird. The eggs and larvae are dug and
pried out of their burrows in the wood by woodpeckers
and creepers; those underground are scratched and
clawed up to view by quail, partridges, and many spar- The Food of Birds rows; warblers and vireos scan every twig and leaf;
flycatchers, like the cat family, lie in watch and spring
after their prey, only in the air instead of on the ground,
feeding more particularly on low-flying insects; while
swifts, swallows, and martins glean their harvest from the
diurnal hosts of high-flying winged creatures. Many FIG. 115.— Crab. FIG. 115.— Crab. times when we think hummingbirds are taking dainty
sips of nectar from the flowers, they are in reality pick-
ing minute spiders and flies from the deep cups of the co-
rollas. The Bird When night falls, the insects which have chosen
that time as the safer to carry on their business of life
are pounced upon by nocturnal feathered beings — the
cavernous mouths of the whippoorwills engulf them as
they rise from their hiding-places, and the bristles of times when we think hummingbirds are taking dainty
sips of nectar from the flowers, they are in reality pick-
ing minute spiders and flies from the deep cups of the co-
rollas. When night falls, the insects which have chosen
that time as the safer to carry on their business of life
are pounced upon by nocturnal feathered beings — the
cavernous mouths of the whippoorwills engulf them as
they rise from their hiding-places, and the bristles of The Bird 150 night-hawks brush them into rapacious maws, if per-
chance they have succeeded in reaching the upper air. chance they have succeeded in reaching the upper air. In tropical forests, where insects are everywhere
abundant, the birds seemed to have realized the fact that
to each is apportioned certain phases of insect life, and
that by hunting in large flocks, instead of competition
resulting between birds of different species, they play
into each other's hands (or rather beaks). It is of such
a flock that Hudson writes: "The larger creepers ex- FIG. 116.— Squid. FIG. 116.— Squid. plore the trunks of big trees, others run over the branches
and cling to the lesser twigs, so that every tree in their
route, from its roots to the topmost foliage, is thoroughly
examined, and every spider and caterpillar taken, while
the winged insects, driven from their lurking-places, are
seized where they settle, or caught flying by the tyrant-
birds.^ birds.^
The Wattled Starlings or Locust-birds of South Africa
live in flocks of thousands, and so dependent are they
on locusts as food, that their habitat and place of nest- The Food of Birds '5* ing is influenced by the presence or absence of
sects. "When pursuing a flight of mature loc
starlings perform various extraordinary and
aerial evolutions with the object of intercepting
rounding a portion of the swarm, and in doing
movements closely resemble those of anothe
destroying starling, the beautiful rose-colour
of eastern Europe and Asia. Individually the t
are very different; collectively and under simi
tions their actions are quite
similar. The Bird Starting in a dense
1 ball-like ' mass, they suddenly
open out into a fan-shaped
formation, then assume a
semicircular arrangement, and
finally end by forming a
hollow cylinder in which a
portion of the locusts are
enclosed; as the imprisoned
insects are destroyed, the
starlings gradually fill up the
hollow of the cylinder until they again assume t
formation and proceed to follow the remainin
The ground below the flock is covered with the
of the birds and the snipped-off legs and wings
At other times the starlings station themselve
tops of bushes and trees, from which they da
flying insects like flycatchers. "In Cape Colony the Locust-birds usually
oc
FIG. 117.— Ratt ing is influenced by the presence or absence of these in
sects. "When pursuing a flight of mature locusts these
starlings perform various extraordinary and beautiful
aerial evolutions with the object of intercepting and sur-
rounding a portion of the swarm, and in doing this their
movements closely resemble those of another locust-
destroying starling, the beautiful rose-coloured Pastor
of eastern Europe and Asia. Individually the two species
are very different; collectively and under similar condi-
tions their actions are quite
similar. Starting in a dense
1 ball-like ' mass, they suddenly
open out into a fan-shaped
formation, then assume a
semicircular arrangement, and
finally end by forming a
hollow cylinder in which a
portion of the locusts are
enclosed; as the imprisoned
insects are destroyed, the
starlings gradually fill up the
hollow of the cylinder until they again assume their 'ball'
formation and proceed to follow the remaining locusts. The ground below the flock is covered with the droppings
of the birds and the snipped-off legs and wings of locusts. At other times the starlings station themselves on the
tops of bushes and trees, from which they dart on the
flying insects like flycatchers. FIG. 117.— Rattlesnake. FIG. 117.— Rattlesnake. FIG. 117.— Rattlesnake. flying insects like flycatchers. "In Cape Colony the Locust-birds usually breed in
very large colonies, in localities in which the locusts have The Bird 1 deposited their eggs. For hundreds of yards every thorny
bush is packed full of cup-shaped nests, even the spaces
between the nests being often filled up with sticks or
rubbish, through which narrow passages are left for the
ingress and egress of the birds. The Bird Many starlings that can
find no room in the bushes build on the ground, or under FIG. 118. — Brown Pelicans diving for fish. (Sanborn, photographer. Courtesy
N. Y. Zoological Society.) FIG. 118. — Brown Pelicans diving for fish. (Sanborn, photographer. Courtesy
N. Y. Zoological Society.) stones, or in holes, and these unfortunates, together with
their eggs or young, ultimately become the victims of
the smaller carnivorous mammals or of snakes. It fre-
quently happens also that either the young locusts are
hatched in insufficient numbers or that they migrate before
the young starlings are fledged. In either case large
numbers of birds perish of hunger, the majority of the stones, or in holes, and these unfortunates, together with
their eggs or young, ultimately become the victims of
the smaller carnivorous mammals or of snakes. It fre-
quently happens also that either the young locusts are
hatched in insufficient numbers or that they migrate before
the young starlings are fledged. In either case large
numbers of birds perish of hunger, the majority of the The Food of Bi 153 old birds and the more advanced young following the
locusts." locusts."
Slugs and snails are eaten by thrushes and many other
small birds with avidity, and the name " Snail-hawk"
has been given on account of one bird's fondness for these
mollusks. The Oyster-catcher feeds on clams and oysters
and derives its name from the facility with which it in-
serts its bill and pries open the shells. The Courlan,
a near relative of the rails, feeds on clams and mussels
and has a most ingenious method of obtaining its food. In shallow water it feels about with its feet for these
mollusks, and when they are found the bird inserts its
bill between the valves with a sudden quick stroke, and,
thus suspended, the heavy shell and its occupant are
carried to the shore, where the shell is forced open and
the animal eaten. Crows treat shell-fish in the same
way that they do sea-urchins and crabs, carrying them
aloft and, after dropping them, descending to feed
on the nutritious flesh exposed by the shattered
shells. Squids, the "head-footed" leaders of the division of
mollusks, are eaten by penguins at least, and so numerous
and at times so conspicuous a feature of marine life are
they that probably many other birds also feed upon
them. The Bird Even deep-water snails and crabs are not safe,
as the sturdy sea-ducks will sometimes dive to a depth
of one hundred and fifty feet to feed upon them. Fish count many enemies among birds, which have
numerous ways of obtaining their victims from ocean or
lake. Some of these are so ingenious that they well The Bird 1 deserve notice. In their variety they rival the methods
of man himself, and we find many analogies between the
two. Penguins earn their food with perhaps the hardest
work, as they follow the fast-swimming fish of the open
ocean in their own icy element and capture them not-
withstanding their speed and quick turns. withstanding their speed and quick turns. We must not forget the slim, evil-looking snake-
birds of the tropical swamps, which also dart through
the water, but impale their victims on their needle-pointed
beaks, suggesting the fish-spears of mankind. Cormo-
rants and sheldrakes also dive after the fish on which
they feed. they feed. Next in the list of strenuous seekers after fish we
must mention the osprey, which hovers on slowly vibrat-
ing wings, treading the air, as it were, over some favourite
spot, until a finny back shows itself near the surface,
when, giving itself to gravitation, the bird drops like a
plummet. It seizes its prey in its talons, while our com-
mon kingfisher, after watching patiently from some
branch overhanging the water, uses its bill to capture
the fish. Terns dive for their fish, gulls usually snatch
them from the surface, and skuas and jaegers get theirs
at second hand, stealing fish from the more skilful fishers
of the sea. When schools of mullet leap in frantic fear
from the water to escape the attacks of porpoises, or
when the dolphins force the flying-fish above the surface,
the merciless Frigate-bird has but to pick and choose. Certain cormorants are the analogues of man's gill-nets,
a flock of these birds surrounding a school of fish in a
half-circle and driving them ashore or into shallow water. The Bird i56 Herons are the 'still-fishers' of the bird world, and stand
in the shallows, silent and motionless as the reeds around
them, with their lance-like beaks in rest and their necks IG. 120.— Great Blue Heron, a still hunter. (Sanborn, photographer. Cou
N. Y. Zoological Society.) FIG. 120.— Great Blue Heron, a still hunter. (Sanborn, photographer. Courtesy
N. The Bird Y. Zoological Society.) at a hair-trigger poise. So we see that few kinds of fish,
from the lowly lamprey to the jewelled brook trout,
escape the sharp eyes of birds, and even when decayed at a hair-trigger poise. So we see that few kinds of fish,
from the lowly lamprey to the jewelled brook trout,
escape the sharp eyes of birds, and even when decayed The Food of Bi 157 masses of fish are thrown ashore, feathered scavengers
are always alert. masses of fish are thrown ashore, feathered scavengers
are always alert. Frogs always suggest storks to our minds, the rela-
tion being of course solely a gastronomic one, and indeed
most of the near relatives of the frog pay their tithe to
birds in a similar way. V. r
FIG. 121. — Wild Mouse, the most frequent victim of birds. FIG. 121. — Wild Mouse, the most frequent victim of birds. Turtles, lizards, and snakes enter largely into the food
of certain birds, some of which, such as the Secretary-
bird and our native Road-runner, are adepts in the cap-
ture and killing of members of the latter division of
reptiles. Certain sea-eagles subsist chiefly upon sea-
snakes. 1 The Bird The most unpleasant items in the bill of fare of the
bird kingdom are birds themselves, although few, if any,
hawks or owls feed exclusively on members of their
own Class. The most systematic cannibal among birds
is the Peregrine Falcon or Duck Hawk, and, where birds
are abundant, this fastidious gourmand merely eats the
flesh of the head and neck and the eyes of each victim,
leaving the remainder of the body untouched. Occa-
sionally, as among other animals, a bird of strictly vege-
tarian habits will attack another bird, even one of its
own kind, and kill and eat it in the most matter-of-fact
way. Owls are the terror of many birds, from the tiny Elf-
Owl which sometimes finds a sparrow too great a match
for him, to the great Strenuous Owl of Australia, which
snatches full-grown Lyre-birds from their perches. But
these birds of the night are ever ready to vary their
diet; as we read of certain owls in India feeding chiefly
on fish and crabs which they snatch from the water. In
that same country, too, bats form a large part of the
Barn Owl's diet. Barn Owl's diet. The Bird Every class of living
beings appears, at certain phases
of its existence, to check or come
into intimate contact with other
unrelated groups, radically affect-
ing the most isolated, in ways
too subtle for our observation. A little green flycatcher snatch-
ing a tiny gnat from its hiding-
place beneath a leaf seems
a trivial incident, and yet ^— FI
the effects of accumulated ^^^
events no more important than this a
world, so delicate is the balance of Natu surprising in its numbers and
extent. Every class of living
beings appears, at certain phases
of its existence, to check or come
into intimate contact with other
unrelated groups, radically affect-
ing the most isolated, in ways
too subtle for our observation. A little green flycatcher snatch-
ing a tiny gnat from its hiding-
place beneath a leaf seems
a trivial incident, and yet ^— FIG 122._Red.tailed Hawk
the effects of accumulated ^^^ (thunteTatcher) an aetive
events no more important than this are felt around the
world, so delicate is the balance of Nature. surprising in its numbers and
extent. Every class of living
beings appears, at certain phases
of its existence, to check or come
into intimate contact with other
unrelated groups, radically affect-
ing the most isolated, in ways
too subtle for our observation. A little green flycatcher snatch-
ing a tiny gnat from its hiding-
place beneath a leaf seems
a trivial incident, and yet ^— FIG 122._Red.tailed Hawk
the effects of accumulated ^^^ (thunteTatcher) an aetive
events no more important than this are felt around the
world, so delicate is the balance of Nature. FIG 122._Red.tailed Hawk
(thunteTatcher) an aetive
his are felt around the
f Nature. FIG 122._Red.tailed Hawk
(thunteTatcher) an aetive
are felt around the The Bird The eggs of birds are delicacies which many feathered
robbers, such as jays and crows, can never resist. There
are two birds, however, one a raven and one a hawk,
which well deserve the eggs which they steal, — so inge-
nious is their method of obtaining them. In South Africa,
on an ostrich-farm, when a female bird has left her nest
for a few minutes, a black form will often appear and
hasten toward the great white eggs. Hovering over
them the raven will let fall a stone into their midst, The Food of Bi 159 instantly swooping down and regaling himself on the
yolk pouring out through the crack in the shell. His
beak being too weak to break the shell, he has learned
to adopt this effective method. A similar remarkable
habit is related of the Black-breasted Buzzard of Aus-
tralia, but in this case it is an emeu which is the victim. After breaking a hole in the thick shell, this bird inserts
its foot and carries the egg to its nest. Perhaps every Order of the higher warm-blooded
animals may be included in our list, from the sloth which
mutely resigns itself to the terrible grip of a Harpy
Eagle to the human child which is powerless before the
attack of some bird of prey frenzied with hunger. In
certain districts eagles and hawks have been shot
smelling strongly of skunk, but whether that fearless
animal really figured in their diet remains to be proved ! If any entire group of mammals is to be excepted from
the birds' bill of fare, it is only that of the whales, although
indeed, when one of these leviathans dies from any cause,
his blubber and oil furnish food for sea-birds of many
kinds. The small gnawers of wood, the rodents, suffer most
heavily, and untold thousands are devoured by hawks
and owls, while cranes, shrikes, and ducks make away
with their share. This brief and very imperfect review of the vast
variety of substances eaten by birds is at least instruc-
tive in revealing vividly the complex interrelations of
all organic life on the earth. A counter-list of animate
creatures which cause the death of birds would be as The Bird i6o surprising in its numbers and
extent. Oddities of Birds' Diet To give any adequate idea of the vagaries of the diet
of birds would require a volume by itself, but certain changes
in feeding habits, due to
some increased pressure in
the struggle for existence,
are too interesting to be To give any adequate idea of the vagaries of the diet
of birds would require a volume by itself, but certain changes
in feeding habits, due to
some increased pressure in
the struggle for existence,
are too interesting to be
passed by unnoticed. They
show us how plastic and
adaptive birds as a whole
are, — how, often, instead
FIG. 123.-Red Squirrel (the watched), of giving Up and becoming
food of hawks and owls. . . (R. H. Beebe, photographer.) extinct, a Certain race Will FIG. 123.-Red Squirrel (the watched),
food of hawks and owls. (R. H. Beebe, photographer.) FIG. 123.-Red Squirrel (the watched),
food of hawks and owls. (R. H. Beebe, photographer.) The Food of Birds 161 instantly accept changed conditions and flourish under
the new regime. the new regime. As the range of diet of the whole Class of birds is so
vast, doubtless the food of the individual species varies
more than we should ever suppose, but many instances
are recorded of birds regularly feeding on food for whose
capture they seem very ill adapt-
ed. Insects form the staple food
of all flycatchers and tyrant-
birds, but the Sulphur Tyrant and
several others readily devour
snakes. They dash down at one
of these reptiles, catch it up in
their beak, and, flying back to
a branch or stone, hammer the
snake flail-like, until its life is
battered out. Certain small king-
fishers living in New Zealand
have deserted the habits of their group, and subsist on the
remarkable diet of " flies, young birds, and cherries"! FIG. 124.— Texas Kingfisher
fishing for insects. As the range of diet of the whole Class of birds is so
vast, doubtless the food of the individual species varies
more than we should ever suppose, but many instances
are recorded of birds regularly feeding on food for whose
capture they seem very ill adapt-
ed. Insects form the staple food
of all flycatchers and tyrant-
birds, but the Sulphur Tyrant and
several others readily devour
snakes. They dash down at one FIG. 124.— Texas Kingfisher
fishing for insects. FIG. 124.— Texas Kingfisher
fishing for insects. Oddities of Birds' Diet remarkable diet of " flies, young birds, and cherries"! The change in habits of the Kea Parrot is only too
well known, especially to the sheep-raisers in New Zea-
land, the home of these birds. Originally exclusive
fruit-eaters, they have lately become so fond of the fat
from the backs of living sheep that they have developed
into ravenous birds of prey, vivisecting their victims and
rejecting all but the choicest morsels. Gulls have long
been known to enjoy an insect diet, and on the pampas
in the vicinity of Buenos Ayres the people look and pray
for flocks of gulls as the only relief from the hordes of The Bird 1 6 grasshoppers which occasionally devastate that region. In the antipodes we find a gull with crepuscular habits,
whose entire food consists of night-flying moths. whose entire food consists of night-flying moths. Birds in captivity may sometimes be induced to eat
food which they would never touch when in a state of
freedom, but there are three species of birds the variety
of whose natural diet will challenge that of any living
creature. The first is a Burrowing Owl. This bird will
not disdain vegetable food, and in its underground dining-
chambers have been found remains of ducklings, spar-
rows, mice, and many other small birds and rodents ;
snakes and frogs, besides spiders, beetles, and apparently
all small forms of life which these little birds are able to
catch and kill. But leaving even the Burrowing Owl
far behind in this respect is the Chimango Carrion-hawk
of southern South America. Hudson tells us that noth-
ing comes amiss to these birds. The vulture habit is per-
haps strongest, and all offal and decaying meat is pounced
upon with eagerness. All wounded and sickly creatures
are closely watched until they die, or, if the opportunity
offers, are despatched at once. When a large extent of
grass is burned, bountiful repasts are ready for these
birds in the shape of roasted snakes and small mammals. Eggs and young birds are especial dainties for the Chi-
mango, and young sheep are often attacked, bringing to
mind the Kea Parrot. The bird is, at times, a vegetable-
feeder, and in fact it would be hard to find any organic
object near its home, the edibility of which it has not
tested. whose entire food consists of night-flying moths. Oddities of Birds' Diet Birds in captivity may sometimes be induced to eat
food which they would never touch when in a state of
freedom, but there are three species of birds the variety
of whose natural diet will challenge that of any living
creature. The first is a Burrowing Owl. This bird will
not disdain vegetable food, and in its underground dining-
chambers have been found remains of ducklings, spar-
rows, mice, and many other small birds and rodents ;
snakes and frogs, besides spiders, beetles, and apparently
all small forms of life which these little birds are able to
catch and kill. But leaving even the Burrowing Owl
far behind in this respect is the Chimango Carrion-hawk
of southern South America. Hudson tells us that noth-
ing comes amiss to these birds. The vulture habit is per-
haps strongest, and all offal and decaying meat is pounced
upon with eagerness. All wounded and sickly creatures
are closely watched until they die, or, if the opportunity
offers, are despatched at once. When a large extent of
grass is burned, bountiful repasts are ready for these
birds in the shape of roasted snakes and small mammals. Eggs and young birds are especial dainties for the Chi-
mango, and young sheep are often attacked, bringing to
mind the Kea Parrot. The bird is, at times, a vegetable-
feeder, and in fact it would be hard to find any organic
object near its home, the edibility of which it has not
tested. The Red-winged Starlings of South Africa during The Food of Birds the greater part of the year feed upon larvae and insects,
but grapes, figs, and other soft fruits are eagerly devoured. They catch locusts and flying ants and occasionally devour
the young of small birds. When their travels take them
near the seashore they search the seaweed for snails
and shrimps, and one of the greatest delicacies is the FIG. 125.— Moth and Hummingbird. Both half natural size. FIG. 125.— Moth and Hummingbird. Both half natural size. fruit of the syringa-tree, "on which they sometimes gorge
themselves until they are no longer capable of flight, . . . affected by some narcotic property of the berry itself." affected by some narcotic property of the berry itself."
This state of semi-intoxication is by no means rare
among fruit-eating birds, when over-ripe or fermented
fruit is abundant. Oddities of Birds' Diet 1 The Bird The great extent to which all the external organs
and parts of birds are adapted to facilitate the obtain-
ing of food is evident in every species; but in humming-
birds this adaptation is especially apparent, because we
can compare these feathered mites with other creatures
far beneath them structurally, but with feeding habits
and general environment so similar that such a com-
parison is fraught with interest. These other creatures
to which I refer are hummingbird moths. Again and
again collectors have shot the moths, mistaking them
for hummingbirds, as the manner of flight is the same
in both, and the way in which each species poises before
a flower, probing it with proboscis or bill, is identical. Of the way this wonderful resemblance is carried out
even in details of the body Bates writes: "It is certainly
very curious, and strikes me even when both are in the
hand. Holding them sideways, the shape of the head
and position of the eyes in the moth are seen to be nearly
the same as in the bird, the extended proboscis represent-
ing the long beak. At the tip of the moth's body there
is a brush of long hair-scales resembling feathers, which
being expanded look very much like a bird's tail." being expanded look very much like a bird's tail."
It seems very improbable that this resemblance can
be attributed to mimicry, as neither has many danger-
ous enemies, their marvellous powers of flight being an
all-sufficient protection. So we are left to conclude that
it is solely to similarity in method of seeking their food
that the likeness is due. THE BREATH OF A BIRD HINK of a mite of a hummingbird shooting
southward mile after mile; his singing wings
beginning their throbbing in the cool damp air
of an Alaskan fall, whirring through the dry heat of des-
erts and around the wind-eddied spurs of mountain-ranges,
until they hum in the warm atmosphere of Mexico or
Brazil, where tiny insects are never lacking throughout
the winter! How exquisite an adjustment must exist
in his organs; how mankind's engines of locomotion are
put to shame! The only comparison of which we can
think is with an insect, —a sphinx-moth or a beetle, whose
wings of gauze lift and carry their owners so easily, so
steadily. It will be interesting to keep this similarity
in mind, superficial though it is. H Birds require, comparatively, a vastly greater strength
and "wind" in traversing such a thin, unsupporting
medium as air than animals need for terrestrial locomo-
tion. Even more wonderful than mere flight is the per-
formance of a bird when it springs from the ground, and
goes circling upward higher and higher on rapidly beating
wings, all the while pouring forth a continuous series of
musical notes, the strength of the utterance of which
165 whi The Bird i66 is attested by their distinctness in our ears after the bird
has passed beyond the range of vision. A human singer
is compelled to put forth all his energy in his vocal ef-
forts, and if, while singing, he should start on a run even
on level ground, he would become exhausted at once. The apparatus which gives to a duck the "wind" to out-
strip an express train, and to a Mockingbird notes which
hold us spellbound as by a motif of grand opera, is most
interesting, and as easy to understand in its general scheme
as it is effective in operation. The Trachea, or Windpipe Look into the beak of a sparrow or
back of the tongue, on the floor of
the mouth, a narrow slit is visible
—the glottis, or opening of the
windpipe. In the gaping yellow
mouth of a nestling robin this
may be seen to excellent advan-
tage, and watched as it widens and
narrows with each breath. But
give the young bird a mouthful
of food, and this air-passage closes
instantly and remains so until all
danger of an intruding substance
is past. No matter how suddenly
you may eject a stream of water
from a medicine-dropper into the
bird's mouth, reflex action wi
anticipate the danger of choking an Look into the beak of a sparrow or pigeon and directly
back of the tongue, on the floor of
the mouth, a narrow slit is visible
—the glottis, or opening of the
windpipe. In the gaping yellow
mouth of a nestling robin this
may be seen to excellent advan-
tage, and watched as it widens and
narrows with each breath. But
give the young bird a mouthful
of food, and this air-passage closes
instantly and remains so until all
danger of an intruding substance
is past. No matter how suddenly
you may eject a stream of water
from a medicine-dropper into the
FIG. 126— Open glottis of
bird's mouth, reflex action will a Pelican,
anticipate the danger of choking and close the aperture. FIG. 126— Open glottis of
ll a Pelican, FIG. 126— Open glottis of
ll a Pelican, The Breath of a Bird The swollen rim of this opening suffices to close it, and
there is no elaborate trap-door arrangement as in mam-
mals, only a few backwardly directed fleshy points. Birds have no trace of an "Adam's apple." The vocal
chords and other adjuncts to the voice of mammals
are entirely absent in birds, not a single note or song
being produced in the upper
throat. ,, FIG. 127. — Windpipe and oesophagus of
bird compared; the former always dis-
tended; the latter soft and collapsed. Passing down the neck
from this orifice is the wind-
pipe, which follows the
course of the oesophagus, or
food canal, passes to one
side of the crop and be-
tween the two branches of
the wish-bone, and finally
divides into two equal parts
called bronchi, which carry
the air directly to the lungs. ,, FIG. 127. The Trachea, or Windpipe — Windpipe and oesophagus of
bird compared; the former always dis-
tended; the latter soft and collapsed. FIG. 127. — Windpipe and oesophagus of
Comparison Of the tWO bird compared; the former always dis-
tended; the latter soft and collapsed. tubes which traverse the
throat and neck of birds shows them to be very different
in appearance and structure, and consideration of their
respective functions gives us the key to this dissimilarity. The only occasion for the oesophagus to open is to permit
the passage of food, and thus a limp, fleshy canal answers
all requirements. The windpipe, on the contrary, must
always be wide open, and not only this, but it must be kept
open no matter what the pressure upon it. In addition,
it must be flexible, yielding to every motion of the neck, The Bird 1 and elastic, in order to stretch and contract as the bird
reaches out or draws back its head. We find a most ingenious arrangement fulfilling all
these requirements. A series of bony rings is imbedded
in the wall of the trachea, beginning with that portion FIG. 128. FIG. 129. FIG. 128. — Windpipe of Flamingo, extended and contracted, showing delicate
mechanism of supporting rings. FIG. 129. — Syrinx-drum of Mallard Drake; the windpipe above; the bronchi
below leading to the lungs. . FIG. 129. FIG. 128 FIG. 129. FIG. 1 FIG. 128. — Windpipe of Flamingo, extended and contracted, showing delicate
mechanism of supporting rings. FIG. 129. — Syrinx-drum of Mallard Drake; the windpipe above; the bronchi
below leading to the lungs. immediately back of the glottis, and extending through-
out its entire length. The membrane which connects these
rings is so elastic that a section of trachea can be drawn
out until it is twice as long as when contracted. When
in the latter condition (I have in my hand an inch of
the windpipe of a flamingo, but the general structure immediately back of the glottis, and extending through-
out its entire length. The membrane which connects these
rings is so elastic that a section of trachea can be drawn
out until it is twice as long as when contracted. The Trachea, or Windpipe When
in the latter condition (I have in my hand an inch of
the windpipe of a flamingo, but the general structure The Breath of a Bi 169 is common to all birds) the trachea appears to be com-
posed of alternating half-rings, but when elongated
these are seen to be complete, the illusion being pro-
duced by the overlapping of half of each ring by an equal
part of the rings on each side. When the trachea is ex-
tended, the only hint of this clever device is a small notch
on the sides of every ring. The illustration demonstrates
the working better than any description. Nature is ingenious but not perfect, as is seen even
in the inch of Flamingo's trachea which I have utilized
for illustration and description. Two of the rings do not
"jibe" on one of their sides, and overlap the wrong way,
but the loss in motion is infinitesimal, the defect being
hardly noticeable even when the rings are bent into a
semicircle. In a very young English Sparrow there are about
fifty rings around the trachea, appearing to be of a more
cartilaginous nature than those of the flamingo. This
latter long-necked bird has no less than four hundred and
fifty rings. In some members of the Class of amphibians (frogs,
toads, and newts) the trachea is supported by small ir-
regular pieces of cartilage, tending in the higher forms
toward ringed areas. Among reptiles an intermediate
condition exists, complete rings being present, but of
cartilage instead of bone. The wonderful music of birds is produced in a rela-
tively small area, known as the syrinx. This organ is
situated at the point where the trachea divides into the
two bronchi. The latter arise as if by a splitting of the The Bird 170 windpipe, and the effect is heightened by the rings which
extend as far as the lungs, which are half-rings or semi-
circles, the inner halves being replaced by membrane. This organ is peculiarly characteristic of birds, there
being not a trace of it in any reptile. being not a trace of it in any reptile. The Bird The Bird 7 The Trachea, or Windpipe But though the syrinx alone is concerned in the pro-
duction of sound, this may be modified, made resonant,
or given a reverberating quality by a special structure
or by windings of the trachea before it reaches the syrinx,
and which are perfect analogies of human musical instru-
ments. Many species of ducks have an enlarged box of
bone, a kind of drum, on the lower portion of the trachea,
sometimes of one shape, sometimes of another, serving,
doubtless, to give power to the bird's voice. Cranes and
swans have veritable French horns in their breast-bones. The windpipe enters between the arms of the clavicles
or wish-bone, and describes an S or even a more intricate
figure before passing out and dividing into the two bronchial
tubes. When a Trumpeter Swan stretches out its neck
and utters a musical clang, most maligned by comparing
it to a whoop, we should remember the cause of its mellow-
ness. In the majestic Whooping Crane of our Western
States, which in a few years will have vanished from the
earth, the windings of the trachea reach their maximum. The entire windpipe of this bird is four feet in length, and
of this, one-half is coiled within the sternum, or breast-
bone, giving remarkable volume and resonancy to the
voice. The Breath of a Bird 171 FIG. 130.— Breast-bone of Sandhill Crane. FIG. 130.— Breast-bone of Sandhill Crane. FIG. 131. — Breast-bone of Whooping Crane, showing convolutions of trachea
within the keel. FIG. 131. — Breast-bone of Whooping Crane, showing convolutions of trachea
within the keel. The Bird The Syrinx This organ is peculiar to birds and, as stated before,
is alone concerned in the production of the voice, although
the tongue in parrots may be of some aid in distinctness
of articulation. But this is not true of any other Order
of birds, and the operation of splitting the tongue of a
magpie or crow to "make it talk" is as unnecessary as
it is inhumanly cruel. The syrinx is singularly uniform among birds, and
this seems the more remarkable when we consider the
great variety of vocal sounds which are produced. The
position and the structure of this organ vary within nar-
row limits, but in general it is composed of several modi-
fied rings of the lower trachea or upper bronchial tubes. The membranes which cover the inner half of each bron-
chial tube unite at their juncture with the windpipe
and extend some little way into it as a thin median fold
of tissue, supported by a bony framework. The tense-
ness or looseness of this membrane is governed by special
muscles, of which there are from one to seven pairs. It
is by the action of these muscles that the varying tones
of croak, scream, warble, or trill are produced, the air
from the lungs rushing out through the bronchial tubes
and past the varying aperture controlled by the syringeal
membrane. We may dissect out every muscle and study trachea,
syrinx, and bronchi with all the apparatus and instru-
ments afforded by modern science, and yet the mystery The Breath of a Bi 173 of song is not solved. The marvel of the Canyon Wren's
melody becomes but the more wonderful; the voice of
the Seriema, carrying over a mile, and the never-to-be-
forgotten evening song of the Solitaire only impress us
with the failure of the scalpel and microscope to explain
more than superficially the varied expressions of life. Lungs and Air-sacs At the beginning of this chapter a bird was compared
to an insect, and the reason will now be apparent. The
body of an insect is aerated by means of an intricate sys-
tem of tubes ramifying throughout the body, which in
many instances are connected with air-sacs. The com-
parison with a bird is not to its lungs, which are small
and compact, but to a series of nine air-sacs, distributed
through much of the body, — four pairs, and two which
have coalesced into one. When a bird is dissected, the thin membranous walls
of these air-cavities are collapsed and rather difficult
to make out, being very similar in appearance to other
connective tissues of the body. But if we insert a small
blowpipe into the trachea of a dead bird, tie it tightly
about with a piece of string and blow into it, all the air-
sacs will become distended and bladder-like and can
easily be made out. It is remarkable how closely these
sacs fit around the viscera and muscles, occupying every
crevice and filling the whole body of the bird with air,
thus reducing its specific gravity, and making it a crea-
ture literally "of the air." There is sometimes a layer of The Bird 1 air between the muscles and the skin, and when we
handle a bird thus aerated the skin crackles under our
touch. The lungs and air-sacs send off tiny membranous
tubes which enter the bones of the limbs and skull and
sometimes even the small bones of the wings and toes,
which are hollow and thus filled with air. It seems in-
credible, but nevertheless it is true that the connection
between the lungs and the upper arm-bone of a bird is
so substantial that a bird which has had its wing broken
with shot is able to breathe through the splintered end
of this hollow bone when its windpipe is completely
choked with blood. We may compare the body of a bird to a submarine
boat with many water-tight compartments, and as such
a vessel is made buoyant by admitting air to these bulk-
heads, so a swimming bird may float high out of water
by inflating its sacs and filling its bone-cavities with air. Conversely, when we see a grebe slowly and mysteriously
submerge its body, we conclude that it has but emptied
its lung auxiliaries. Lungs and Air-sacs We now come to the most important part of the re-
spiratory system, where the blood and the air come into
closest contact and exchange gases, the oxygen of the
air vitalizing the entire body. If we follow the two
bronchial tubes after they leave the syrinx, we shall find
that each enters a lung, and passes through it, giving
off a number of side branches which open into the vari-
ous air-sacs. The lungs are not elastic and, instead of
lying freely in .the body, are flattened against the back- The Breath of a Bird bone and ribs, and when carefully removed show fur-
rows made by these latter bones. There is still much
to be learned of the manner of a bird's breathing, but
it is probable that there is a sidewise or dorsal expan-
sion of the ribs, rather than of that portion corresponding
to our chest. In a bird the latter region is chiefly an im-
mense flight-muscle, which could hardly yield to the
action of breathing while carrying on the tremendous
work of keeping the wings in motion, and when a FIG. 132. — Cross-section of wing bone of Ostrich and Black Swan. In life the
bone of the Ostrich is filled with marrow; that of the Swan with air. FIG. 132. — Cross-section of wing bone of Ostrich and Black Swan. In life the
bone of the Ostrich is filled with marrow; that of the Swan with air. bird squats on a branch with its breast pressed close
to the perch, " chest expansion" must be all but
impossible. We cannot help being surprised at first when we see
how small are the lungs of a bird in comparison with
the size of its body. The first thought that occurs to us is that the air-
sacs in birds, and the hollow cavities of the bones, must
function chiefly as aids to flight, and we should expect
to find as best flyers those birds in which the air-cavities The Bird 1 7 are most numerous, but there are many exceptions. The
bones of storks and vultures (birds of great powers of
flight) are extremely pneumatic, while the bones of the
flightless ostriches are filled with marrow, and in the
aquatic penguins even this is reduced to a thread, the
bones being almost wholly osseous tissue. Lungs and Air-sacs A swan,
although a heavy bird, flies remarkably well when once on
the wing, and is highly aerated, but, on the other hand,
terns and swifts — past-masters both in aerial evolutions
— have solid bones! Now an athlete who is trained in running has always
a very large lung capacity. Two persons of equal health
and strength, one of whom has run many races or who
has the power of keeping up a dog-trot for hour after hour,
while the other has led a more sedentary life, may show
a remarkable difference in the amount of air which they
can draw into their lungs — perhaps one hundred or one
hundred and fifty cubic inches more in the case of the
runner. The average person uses only about one sev-
enth of his lung capacity in ordinary breathing, the rest
of the air remaining at the bottom of the lung, being
termed "residual." As this is vitiated by its stay in the
lung, it does harm rather than good by its presence. When
great exertion is required, as in running, the person who
can admit the largest amount of fresh air to his lungs in
each breath has command of an equally great power
of action. As we have seen, the lungs of a bird are small and
non-elastic, but this is more than compensated by the
continuous passage of fresh air, passing not only into The Breath of a Bi 177 but entirely through the lungs into the air-sacs, giving,
therefore, the very best chance for oxygenation to take
place in every portion of the lungs. When we compare
the estimated number of breaths which birds and men
take in a minute — thirteen to sixteen in the latter, twenty
to sixty in birds — we realize better how birds can per-
form such wonderful feats of song and flight. Birds, having no sweat-glands in the skin, and the
action of the capillaries being impeded by the feathers,
would have no way of regulating the temperature of the
body, much as this is necessary in flight, if it were not
that the great quantity of air exhaled with each breath
relieves the body of any excess of heat. Lungs and Air-sacs However directly or indirectly the air-sacs are con-
cerned with flight, a bird which sings uninterruptedly
as it flies upward must be immeasurably aided by the
great quantity of air at its command. And again, when
a Prairie Hen inflates the orange-hued air-sacs on both
sides of its neck, there is only one explanation as to
their use, at least at the time of courtship, namely, an
added decoration, and as an aid in the "booming" —
factors both of which, for aught we know, may help to
soften the hearts of the coquettish females. Looking down the scale of life we find an animal among
the reptiles with a lung which at once suggests that of
a bird. The lungs of a chameleon are spongy and com-
pact in front, but farther back they are hollow, and
give off a dozen or more finger-like tubes or lobes, thus
foreshadowing, at least in appearance, the air-sacs
of birds. i78
The The Bird i78 i78
The Bird
We have learned that the chick
through a stage when it possess
veloped gills. This proves that in t
the ancestors o
aquatic and fish-
lungs? Fishes ha
in their aquatic l
be useless. Neve
see, the lungs of
mammals are leg
tures of the sea. We have learned that the chick in the egg passes
through a stage when it possesses several well-de-
veloped gills. This proves that in the dim, distant past
the ancestors of birds were once
aquatic and fish-like. But how about
lungs? Fishes have none, and indeed
in their aquatic life such organs would
be useless. Nevertheless, as we shall
see, the lungs of reptiles, birds, and
mammals are legacies from the crea-
tures of the sea. Many fishes have within their
bodies a thin-walled sac, known as
the swim-bladder. This is filled with
gas, and as the fish ascends to the
surface, or dives to where the pres-
sure of the water is very great, the
amount of gas varies; so that the
specific gravity of the fish changes
with that of the water. This swim-
bladder is generally connected with
the throat by a delicate tube; and
FIG. 133. — Lung of Chame- , , i_ xi
Icon, foreshadowing con- in these tWO Structures W6 have the
dition in bird. , , /• ,LI i • i » i i
homologues 01 the birds lungs and
trachea. Proof of this is to be found in the growth of the
lungs in all young chicks. A tiny bud appears upon the
primitive oesophagus, just behind the little gill-clefts, and We have learned that the chick in the egg passes
through a stage when it possesses several well-de-
veloped gills. This proves that in the dim, distant past
the ancestors of birds were once
aquatic and fish-like. But how about
lungs? Fishes have none, and indeed
in their aquatic life such organs would
be useless. Nevertheless, as we shall
see, the lungs of reptiles, birds, and
mammals are legacies from the crea-
tures of the sea. FIG. 133. — Lung of Chame
Icon, foreshadowing con-
dition in bird. Many fishes have within their
bodies a thin-walled sac, known as
the swim-bladder. The Bird This is filled with
gas, and as the fish ascends to the
surface, or dives to where the pres-
sure of the water is very great, the
amount of gas varies; so that the
specific gravity of the fish changes
with that of the water. This swim-
bladder is generally connected with
the throat by a delicate tube; and
e- , , i_ xi
in these tWO Structures W6 have the
, , /• ,LI i • i » i i
homologues 01 the birds lungs and
his is to be found in the growth of the
chicks. A tiny bud appears upon the
, just behind the little gill-clefts, and
l it is larger than the food-canal itself. es into two equal parts which become
r canals — the beginnings of the lungs. Many fishes have within their
bodies a thin-walled sac, known as
the swim-bladder. This is filled with
gas, and as the fish ascends to the
surface, or dives to where the pres-
sure of the water is very great, the
amount of gas varies; so that the
specific gravity of the fish changes
with that of the water. This swim-
bladder is generally connected with
the throat by a delicate tube; and
FIG. 133. — Lung of Chame- , , i_ xi
Icon, foreshadowing con- in these tWO Structures W6 have the
dition in bird. , , /• ,LI i • i » i i
homologues 01 the birds lungs and
trachea. Proof of this is to be found in the growth of the
lungs in all young chicks. A tiny bud appears upon the
primitive oesophagus, just behind the little gill-clefts, and
increases in size until it is larger than the food-canal itself. It then in turn divides into two equal parts which become
diminutive flaps, or canals — the beginnings of the lungs. FIG. 133. — Lung of Chame
Icon, foreshadowing con-
dition in bird. FIG. 133. — Lung of Chame
Icon, foreshadowing con-
dition in bird. The Breath of a Bi The Breath of a Bi 179 A simple experiment will show what fishes have a
canal, or duct, leading from the throat to the swim-blad-
der and what have not. If a goldfish and a perch or
sunfish be placed in a bowl of water and the air exhausted,
the two latter will be forced to the surface, while the gold-
fish will soon eject a few bubbles of air, or gas, from its
mouth and stay at the bottom. Thus we can see the ad- FIG. 134. — Diagram of growth of lungs. X, the lower part of the primitive diges-
tive tract, divides into two parts, XX, the lungs. 4. — Diagram of growth of lungs. X, the lower part of the primitive diges-
tive tract, divides into two parts, XX, the lungs. vantage of such a canal in enabling the fish to regulate
the amount of gas in the bladder. When the fish-like creatures of old took to living on
land, the change from swim-bladder and gas to lung
and air was a remarkable example of change of function
of an organ, and the more we learn of the lungs of living
creatures the more marvellous does this transformation The Bird i seem to us. In changing, Nature seems to have tried
numberless experiments, only a few of which have sur-
vived. For example, we know that fish breathe by a
sort of swallowing, the water being taken in at the mouth
and poured out through the gill-clefts. So in frogs and
salamanders we find that, although they possess lungs,
yet they still employ a swallowing process to get the air
down their throats. This is the reason why a frog will
suffocate if its mouth is held open. There are certain
salamanders which are wholly without lungs, their moist
skin being so vascular that the blood is purified through
it. But strange to say, these amphibians still swallow and
swallow, as did their ancestors, although no air passes
down their throats, and indeed there is no place to which
it could go! As we have seen elsewhere, birds exhale
air largely by the action of certain abdominal muscles. The Breath of a Bi Watch a goldfish rise to the top of the water and eject or
gulp down a bubble of air, and observe the rapid breath-
ing of a bird, and you have the two extremes before you
—the swim-bladder of ages ago and the wonderful lungs
of a bird of to-day. The Heart and the Life-blood Perhaps the most wonderful organ in a bird's body
is its heart. In the very lowest of back-boned animals
the heart is merely a long tube, in fact a simple artery
or vein, which contracts at certain intervals and so pro-
pels in a forward direction the fluid which it contains. A fish may almost be said to have its heart in its head,
so far forward in its body is it placed; nevertheless, as The Breath of a Bi 181 is the case in all the warm-blooded creatures above it,
the heart is nearer the under side of the body — the breast
— than near the back. And herein lies an important
difference between the two great divisions of the Ani-
mal Kingdom, vertebrates and invertebrates, — the former
always having the heart near the breast, while in the back-
boneless organisms it is near the back. boneless organisms it is near the back. The heart of a fish is fairly concentrated and muscu-
lar, but the blood which passes through it is but an im-
pure and sluggish stream. In reptiles both pure and
impure blood is found in the heart, but they mingle, and
thus half destroy the purifying action of the lungs. This
explains why these animals are cold-blooded, and also
accounts for their usual lethargic disposition and low
mental plane of life. In crocodiles we find a significant condition. There
are four chambers in the heart, as in mammals and in
birds, but this avails nothing; for, leading from the heart
are two arteries instead of one, and where these cross
each other there is a tiny aperture — a small opening in
the partition which allows the impure blood to leak into
the stream of pure, red blood, and so a crocodile is only
a crocodile, although evolution has lifted his heart al-
most to a level with birds and the warm-blooded ani-
mals. If this tiny hole could become closed, and the
two streams of blood be kept separate, the eyes of the
crocodile would brighten, his activity increase many fold,
and in fact his entire plane of life would be changed. I have thus briefly reviewed the heart in the lower
vertebrates in order to give a more vivid idea of this organ The Bird 1 8 in birds. The Heart and the Life-blood Here we find an organ remarkably large in
proportion to the size of the bird's body — a conical knot
of muscle, the power of which is almost beyond belief. The heart of a bird is said to beat a " hundred and twenty
times a minute when the bird is at rest. The first flap
of the wings doubles the pulsations, and when the bird
is frightened or exhausted the number of beats are too
many to be counted." many to be counted."
There are four separate chambers, known as right and
left ventricles and auricles, and the partition which di-
vides the heart in the middle is blood-tight so that not a
particle of "bad" blood can get through and vitiate the
life-giving stream which has just come from the lungs. life-giving stream which has just come from the lungs. A Bluebird is perched on a twig near its nest mur-
muring its sweet warble; a Wood Pewee, half hidden
in the shadows of some dense, moist forest, speaks to
us in its sad dreamy phrase; how calmly, how quietly
they sit! It seems impossible to believe that every drop
of blood in their bodies is rushing back and forth with
inconceivable rapidity — from heart to head, from body
to wings and legs, and back again ! Let us take the blood as it is just leaving the heart
in the breast in one of these little feathered beings, and
trace its course through the body and back again to
the starting-point. The left ventricle opens into the
aorta, the greatest artery, or blood-tube leading from
the heart, in the body. The clean oxygen-food-bearing
stream rushes through this channel, which we may com-
pare to the trunk of a tree, and is carried into branch
arteries, dividing finer and finer, just as the trunk of FIG. 135. — Circulatory system of Pigeon (injected), showing blood-vessels rami-
fying from the heart to every part of the body. 183 FIG. 135. — Circulatory system of Pigeon (injected), showing blood-vessels rami-
fying from the heart to every part of the body. The Bird 1 8 the tree merges into limbs, and these into branches, twigs,
stems, and at last into the delicate foliage. The Heart and the Life-blood This last
we may liken to the capillaries or hair-tubes in which
the blood does its real work of supplying nourishment
directly to the tissues, and where it receives the waste
matters, carrying them away in its current. When we have followed the divisions of a tree out
to the foliage, we may find that they touch and interlace
with the foliage of another tree, and this is very much
like what occurs in the course of the blood. The capil-
laries run together and form larger vessels, these in turn
coalesce, and soon the blood — dark now and filled with
the waste matters of the body-cells — is flowing through
only two large veins (veins always lead toward the heart). These enter the right auricle, which opens into the right
ventricle. From here the blood rushes to the lungs to be
purified and back again to the left auricle and ventricle,
and its cycle is complete. If we look at a drop of bird's blood (or that of any kind
of warm-blooded creature) under the microscope, we shall
see thousands upon thousands of oval discs, or corpuscles,
like tiny platters floating in a fluid. These flow about
under the cover-glass through little channels, mechanic-
ally and very slowly of course, and giving but a faint
idea of the way they must tumble and rush after each
other through the veins and arteries of the bird. Scat-
tered among these oval bodies will occasionally be seen
others of indefinite shape and white in colour. As we
watch one of these tiny cells, the thought suddenly comes
over us, — what are birds indeed but collections of untold The Breath of a Bird millions of one-celled animals! For here before us we
have what is almost exactly like the little flowing drops
of jelly called Amcebse which we may find in quiet ponds
and watch as they move about in search of food ; flowing
around a bit of nutriment, digesting it and flowing away
from the waste matter which is left. This is just what the FIG. 136. — Blood-corpuscles of bird. FIG. 136. — Blood-corpuscles of bird. white corpuscles do; they flow around the food which is
absorbed by the walls of the digestive canal, and in fact
act like tiny independent animals, parts though they are
of the great whole. The Heart and the Life-blood The oval corpuscles carry and dis-
tribute the oxygen, and here we have in a sentence the
inner 'living' of a bird: the food-canal bringing in food
and preparing it; the windpipe and lungs admitting white corpuscles do; they flow around the food which is
absorbed by the walls of the digestive canal, and in fact
act like tiny independent animals, parts though they are
of the great whole. The oval corpuscles carry and dis-
tribute the oxygen, and here we have in a sentence the
inner 'living' of a bird: the food-canal bringing in food
and preparing it; the windpipe and lungs admitting The Bird i86 oxygen; and the blood taking up and transporting both
to every part of the body. The normal temperature of our body is about 98£°;
if it rose to 106°, we should soon succumb to the burning
fever, while the little bird before us is healthy and com-
fortable with a temperature of 110° to 112°! FIG 137. — Amoeba, greatly magnified. (Courtesy of Dr. G. L. Calkins.) FIG 137. — Amoeba, greatly magnified. (Courtesy of Dr. G. L. Calkins.) The next time you see a wee chickadee, calling con-
tentedly and happily while the air makes you shiver
from head to foot, think of the hard-shelled frozen in-
sects passing down his throat, the icy air entering lungs
and air-sacs, and ponder a moment on the wondrous little
laboratory concealed in his mite of a body; which his
wings bear up with so little effort, which his tiny legs sup- The Breath of a Bi 187 port, now hopping along a branch, now suspended from
some wormy twig. Can we do aught but silently marvel at this alchemy? A little bundle of muscle and blood, which in this freez-
ing weather can transmute frozen beetles and zero air
into a happy, cheery little Black-capped Chickadee, as he
names himself, whose bravery shames us, whose trust-
fulness warms our hearts ! And the next time you raise your gun to needlessly
take a feathered life, think of the marvellous little en-
gine which your lead will stifle forever; lower your weap-
on and look into the clear bright eyes of the bird whose
body equals yours in physical perfection, and whose tiny
brain can generate a sympathy, a love for its mate, which
in sincerity and unselfishness suffers little when compared
with human affection. The Heart and the Life-blood FIG. 138. — Chickadee in the snow. FIG. 138. — Chickadee in the snow. I IRDS exhibit probably a greater degree of activity
than any other class of animals. Some seem
never to be still, and, whether soaring, fluttering,
running, hopping, climbing, dancing, or swimming, every
motion is the result of the action of one or more muscles. I The entire flesh of a bird is divided up into layers
or bundles of distinct muscles, each having its function, —
raising, lowering, or in some way moving feathers, eye-
lids, legs, wings, tail, and other portions of the body. The
number and intricacy of these muscles can be imagined
when it is stated that in a goose there are more than twelve
thousand muscles or parts of muscles immediately be-
neath the skin, which serve to raise or otherwise move
the feathers. In a penguin the muscles immediately beneath the
skin are unusually well developed, and for an excellent
reason. By means of them the water "may be readily
expelled from the interstices of the plumage so soon as
the bird quits the water. Were it otherwise, in the low
temperature of the Antarctic region, which the majority
of these birds inhabit, their plumage would soon be frozen
1 88 e fro
1 88 Muscles and Ner 189 into an icy mass, the high temperature of the bird being
of itself insufficient to obviate this, although assisted
by the great development of the subcutaneous fatty
layer, which far exceeds in thickness that of the corre-
sponding structure in the member of any other group
of birds, and recalls to mind the fatty deposit of 'blub-
ber7 of the seals and cetaceans." ber7 of the seals and cetaceans."
When we looked at the blood of a bird, we saw the
tiny white corpuscles, which in life flow and move in
every direction, constantly changing their outline; and
now if we take a piece of a bird's muscle or flesh and
examine it carefully, after "teasing" it out into shreds
with a needle, we shall see another kind of cell-animal. These are long and generally pointed, each a single cell
with a tiny spot or nucleus in it, differing from the white-
blood animals in being able to stretch out and contract
in only one direction. When we will our arm to close
together, bringing our hand close to our shoulder, a thick
colony or bunch of these muscle-animals shortens, be-
comes stouter, and bulges up under the skin on our upper
arm. In our own body the bones of the spinal column are
movable, and we can bend in almost every direction,
and so we are provided with many important back-muscles. But if we have ever carved a chicken, we shall remem-
ber that the ribs and shoulder-bones are close to the sur-
face, and but poor pickings are to be had from them. The breast and chest, on the contrary, are hidden in a
thick mass of muscles, most of which are concerned with
moving the wings in flight. The immense pectoral or The Bird I 90 breast muscle, which makes possible the all-important
downward sweep of the wings, weighs one-fifth as much
as the entire bird, bones and all. This arrangement of
a great weight of muscle hung below the point of attach- FIG. 139. — Wing and breast of Pigeon, showing immense pectoral muscles, and
tendons of wing used in flight. FIG. 139. — Wing and breast of Pigeon, showing immense pectoral muscles, and
tendons of wing used in flight. ment of the wings is, for mechanical reasons, the only
one possible in a bird of flight; since any excess of weight
above the wings would instantly overbalance the bird. If we remove the skin from the upper arm of a bird, Muscles and Nerv 191 we shall see a tangle of bundles of red flesh — the muscles
which unite to make the arm of a bird such an exquisite
flying-machine. Where a muscle narrows and is fastened
to a bone, its fibres merge into a thin, tough white cord — FIG. 140. — Model of bird's foot, showing perching tendons; toes extended. FIG. 140. — Model of bird's foot, showing perching tendons; toes extended. a tendon. This is not elastic like the main portion of
the muscle, but is much more tough. In the slender legs and feet of birds there is little more
than bone, tendon, and skin. The tendons which clasp
and unclasp the toes are very interesting, and if we will The Bird 1 bend the tarsus back and forth in the leg of a dead chicken,
the workings of these strands of tissue may be traced
beneath the scales. Reference to the photograph, where
catgut replaces these tendons, will make their workings
still more plain. FIG. 141. — Same as Fig. 140; toes contracted. FIG. 141. — Same as Fig. 140; toes contracted. Many birds cannot flex the leg without drawing the
toes up, and we may be sure that these birds are safe
when they perch; the closer they sit to the branch the
tighter becomes their grip. But this safety mechanism
is not found in all perching birds by any means (Fig. 143). A strange thing about muscles is that there are fine Muscles and Nerves wavy cross-lines, or striations, on those which are moved
voluntarily, such as the muscles of the neck or wing;
but those which are moved involuntarily, as the gizzard,
are smooth and without the cross-lines. Amazon Parrot in sleeping position, hanging by its toes;
the wonderful strength of the tendons. ng birds new muscles have appeared, or o FIG. 142, — Amazon Parrot in sleeping position, hanging by its toes; illustrating
the wonderful strength of the tendons. FIG. 142, — Amazon Parrot in sleeping position, hanging by its toes; illustrating
the wonderful strength of the tendons. Among birds new muscles have appeared, or old ones
have split up or so changed in position that it is all but
impossible to compare them, muscle for muscle, with other The Bird 1 animals. There are so many resemblances between birds
and reptiles that we naturally turn to the latter for com-
parison, but even here we find a great unlikeness. We
learned, when we reflected on the number of ribs of a
bird, that the repetition of so many similar structures
was merely the last remaining vestige of ancestral body
segments, which reach their extreme development (in
number and similarity) among the worms; but in regard
to muscles birds show little or nothing of this. In liz-
ards we may count dozens upon dozens of bands of muscles
succeeding one another, all more or less alike, from head
to tail, but it is only in the neck of a bird that we shall
find anything like this. In order to give to muscles a firm anchorage, they
must of course be attached to the bones. At these
points of attachment deep furrows or cavities are often
found in the surface of the bones, and in still other ways
we are reminded, even in fossil bones, of the flesh and
muscle which once moved them. These muscle impres-
sions are often a valuable source of identification in
naming the bones of creatures which, many thousands
of years ago, disappeared from the earth. And indeed
so great variety exists in the muscles of living birds that
many of them, those of the upper arm for example, are
of considerable value in classification. Nerves The last great system of internal organs which we
shall consider, and perhaps the most mysterious of all,
is that of the nerves. We have learned that the back- The Bird 1 bone supports the entire body and gives a point of at-
tachment for the limbs, but long before limbs were found
among animals on the earth, in fact long before bone
existed, a sheath of cartilage surrounded and supported
the primitive spinal cord of creatures which lived long
ago in earlier epochs of the earth's history. So we may
say this protection to the nerve-trunk is the most im-
portant, as it was the original, function of our vertebrae. When "brainy" creatures appeared, that is, when the
front end of the nerve-cord became enlarged, it needed
some special protection, so a box — the skull, — first of
cartilage, then of bone, was evolved. One more fact which may hark back to old, old times,
and then we shall leave the past as perhaps trespassing
too much on the province of the chick while he is yet
within the egg. Birds (and all the higher classes of ani-
mals) have what we may call two separate systems of
nerves, although in some ways they are insolubly con-
nected with each other. The brain and spinal cord send
numerous branches which subdivide into countless nerve-
lets, permeating every portion of the body, as we can
easily prove by the feeling, on pricking our skin anywhere
with a needle. This is the principal nervous system of
back-boned animals, and it is by this that birds, and all
creatures with well-developed nerves, see, hear, taste,
smell, and by which they send messages to the muscles
when they desire to move them. Below the vertebral
column is another lesser system which sends nerves to
the digestive tract and other organs, the movements and
functions of which are not under control of the will, and . 144. — Nervous system of Pigeon, show
leading to wings FIG. 144. — Nervous system of Pigeon, showing large eyes and brain, and nerves
leading to wings and legs. The Bird 1 this is the sympathetic or reflex system. It is a very
wonderful thing, this not having to think about the heart
beating or the lungs expanding. We can understand how a muscle (such as the heart)
can pump the blood through the body, but we know little
or nothing of the action of nerves. An eagle soars high
above the clouds; a rabbit is discovered crouching in
a field far below; the eye of the eagle telegraphs this
discovery to the brain; a message is sent along the spinal
cord, switches off to the wings, repeats to the muscles,
which half close and set the great pinions firmly; the
eye is the pilot, never leaving the mark; a triple message
now goes out, to the wings to hold back, to the legs to
reach forward, to the talons to open and clutch! All
is done without a break or hesitation, so quickly that
one's eye can hardly register the act, and all by means
of impulses sent through the finest of white, hair chan-
nels, consisting of a substance so unstable that it tears
and falls apart, like wet tissue-paper, when we examine
it. And if the sending and receiving of impulses seems
wonderful to us, what can we say of the brain, the master
of all, where instinct, mind, soul, — no matter what we
call it, — directs the whole life? It is here that fact upon
fact, experience upon experience, is stored from the mo-
ment the bird breaks its shell throughout its whole life-
time, and it is from the brain that the benefit derived
from this perception of experience, failures and successes,
causes and effects, is intelligently brought into play and
made to redound to the bettering of the subsequent
life. Muscles and Ner Muscles and Ner 199 When we carefully remove the upper part of a bird's
skull, we find that the brain occupies the whole interior, FIG. 145. — Comparison of skulls of Heron and Hawk, showing unlikeness caused
by difference in manner of procuring food. FIG. 145. — Comparison of skulls of Heron and Hawk, showing unlikeness caused
by difference in manner of procuring food. the shell or box of bone which protects it being very thin,
although strong. It would be very interesting if we could
compare the short and thick bullet-shaped skull and brain the shell or box of bone which protects it being very thin,
although strong. It would be very interesting if we could
compare the short and thick bullet-shaped skull and brain The Bird 2 of a rapacious hawk with the thin-templed head of a
timid heron and say, "phrenologically," in the first we
have the bump of combativeness well developed, analo-
gous to a prize-fighter; in the second case, timidity is
prominent! But unfortunately, characteristics such as
these are compound, and made up of many simple fac-
tors, the synthesis of which is not confined to any par-
ticular "bump." ticular "bump."
At the first sight of the bird's brain we are struck
with the very great size of the two larger masses of brain-
matter — cerebral hemispheres these are called. It is in
these that the higher faculties reside, and when these are
destroyed, all knowledge, all power of voluntary move-
ment passes from the bird. These great brain-halves
are much larger than in the brain of a reptile, in fact the
cerebral hemispheres, set deep in the great buttressed
skull of a full-grown crocodile, are no larger than those
of the duck which he snaps up. Not only this, but in
the days of the Archceopteryx (which had a typical bird-
brain), the monster Dinosaur, Triceratops, 25 feet long,
had, in its 6 feet of skull, a brain proportionately only
one tenth as large as that of a modern crocodile! When
compared with a mammal there is seen to be a conspicu-
ous difference, since the outer surface is perfectly smooth
in birds, but is wound about in convolutions in the higher
four-footed animals. Muscles and Ner This latter condition is said to indi-
cate a greater degree of intelligence, but when we look
at the brain of a young musk-ox or walrus and find convo-
lutions as deep as those of a five-year-old child, and when
we compare the wonderfully varied life of birds, and Muscles and Nerves 201 realize what resource and intelligence they frequently
display in adapting themselves to new untried con-
ditions, a smooth brain does not seem such an inferior
organ as is often inferred by writers on the subject. I
would willingly match a crow against a walrus any day,
in a test of intelligent behaviour! Between the hemispheres is a small projection which FIG. 146. — Vertical section through skull of bird, showing great size of brain. FIG. 146. — Vertical section through skull of bird, showing great size of brain. is called the pineal body. It is very tiny, and we know
little of its function at present, but its history is one of
the most interesting chapters in the evolution of the
bird, which we shall leave to the chapter on the senses. The other most conspicuous part of the brain is the
cerebellum, or "little brain," a section of which shows
a most remarkable tree-like appearance. This has been
called the arbor vitce — the tree of life. It is in this portion of the brain that a few tiny drops The Bird 2 of blood are found when a bird dies of fright, which oc-
curs more often than in any other class of animals. Sports-
men have fired at a bird, missed it completely, and yet
have seen it drop dead as suddenly as if it had received
the full charge. In captivity, herons succumb more
frequently to fright apoplexy than other birds. When
we assume the care of any creature, bird or beast, we
should treat it as a timid child, and the person who moves
quietly but unhesitatingly will win the confidence of
wild creatures much sooner than when he alarms them'
by sudden motions, or arouses their suspicions by jerky
half-hearted approaches. half-hearted approaches. There are twenty-four nerves given off in pairs from
the brain, which pass out through minute holes in the
skull, and energize eye, ear, tongue, and other organs. THE SENSES E E have seen that the brain is the storehouse of
facts and experiences, but whence come these
and how do they gain admittance to that soft
gray matter which is one of the wonders of the world? There are five channels (and sometimes there seems the
shadow of a metaphysical sixth) which are cognizant of
and receptive to environmental influences. These are
the nostrils, eyes, ears and tongue, and the tactile nerves
of the surface of the body; or in other words the bird is
in direct connection with his surroundings on land or
water or in the air, by means of the senses of smelling,
seeing, hearing, tasting, and feeling. Muscles and Ner Each of these has an individual name, and as they are
homologous with similar nerves in ourselves, the same
name is retained, such as the olfactory, or that leading
to the nostril; and the pathetic, the function of which
is to control the obliquely raising eye-muscle, producing
a pathetic expression, although it must be confessed that
the effect of this in the immobile face of a bird is not
especially affecting. Back of the cerebellum is a thickening of the spinal
cord, and after again narrowing it enters the bones of the
neck and back, as the true spinal cord. At the base of
the neck and near the thigh- joints this cord increases
in size, large nerves being given off at these places to the
wings and legs. It terminates in a fine white thread. The Senses In all animals the mucous membrane which lines the
nasal cavity is very delicate and filled with nervelets. These nervelets unite and form a single nerve on each
side which passes to the brain and transmits the impres-
sions derived from the odours in the air. The thin bones
within the nostril, which, in dogs and deer, curl and re-
curl in delicate lines and
scrolls and thus expose
such a large surface to
the odour-bearing air, are
but poorly represented in
birds. The simple curve
of the bone in the nos-
trils of birds is very simi-
lar in structure to that
found in reptiles. The question whether
vultures perceive their
prey by sight or smell has
been decided in favour of the former sense. Lacking the
ability readily to distinguish delicate odours, we find
among birds none of the glands which are so common
among hairy-coated creatures: the oil-gland is the only
one on the body, and this is practically odourless. But
slight as is the scent which diffuses from birds, it is
FIG. 149. — Turbinal scrolls of dog. In a
bird these bones are far more simple. In all animals the mucous membrane
nasal cavity is very delicate and fille
These nervelets unite and form a singl
side which passes to the brain and trans
sions derived from the odours in the air. within the nostril, which, in dogs and d
curl in delicate lines and
scrolls and thus expose
such a large surface to
the odour-bearing air, are
but poorly represented in
birds. The simple curve
of the bone in the nos-
trils of birds is very simi-
lar in structure to that
found in reptiles. In all animals the mucous membrane which lines the
nasal cavity is very delicate and filled with nervelets. These nervelets unite and form a single nerve on each
side which passes to the brain and transmits the impres-
sions derived from the odours in the air. The thin bones
within the nostril, which, in dogs and deer, curl and re-
curl in delicate lines and
scrolls and thus expose
such a large surface to
the odour-bearing air, are
but poorly represented in
birds. The simple curve
of the bone in the nos-
trils of birds is very simi-
lar in structure to that
found in reptiles. The Sense of Smell The sense of smell is dependent upon the diffusion in
the air of minute particles of objects, and naturally is
effective at very short distances compared to the senses
of sight and hearing, which require only vibrations in the
atmosphere. When we remember that the nostrils of
birds are usually encased in horn and that there is no
exposed moist surface, as in the nose of a dog, we shall
see how it is that this sense is but little developed among
feathered creatures. 203 The Bird 204 FIG. 147. — Nostrils of bird encased in horn. FIG. 147. — Nostrils of bird encased in horn. FIG. 148.— Nostrils of deer encased in moist flesh. FIG. 148.— Nostrils of deer encased in moist flesh. The Senses The Senses The question whether In all animals the muco
nasal cavity is very deli
These nervelets unite and
side which passes to the b
sions derived from the odou
within the nostril, which,
curl in delicate lines and
scrolls and thus expose
such a large surface to
the odour-bearing air, are
but poorly represented in
birds. The simple curve
of the bone in the nos-
trils of birds is very simi-
lar in structure to that
found in reptiles. FIG. 149. — Turbinal scrolls of dog. In a
bird these bones are far more simple. The question whether
vultures perceive their
prey by sight or smell has
been decided in favour of the former sense. Lacking the
ability readily to distinguish delicate odours, we find
among birds none of the glands which are so common
among hairy-coated creatures: the oil-gland is the only
one on the body, and this is practically odourless. But
slight as is the scent which diffuses from birds, it is
sufficient to enable a dog, with his wonderful keenness
of smell, to detect a crouching bird some distance
away. FIG. 149. — Turbinal scrolls of dog. In a
bird these bones are far more simple. FIG. 149. — Turbinal scrolls of dog. In a
bird these bones are far more simple. The woodcock of our inland swamps and marshes,
and the apteryx of New Zealand, probably have the The Bird 2o6 sense of smell most acutely developed, although in both
cases it is the delicate nerves of touch in the bill which
are most helpful in detecting the presence of the earth-
worms which constitute the food of these birds. To whatever degree the nostrils of land birds aid
their owners in procuring food, it is certain that those FIG. 150. — Head of Apteryx, showing tactile hair-like feathers, nostrils at tip
of beak, and small eyes. FIG. 150. — Head of Apteryx, showing tactile hair-like feathers, nostrils at tip
of beak, and small eyes. species which feed entirely on fish, which they swallow
whole, have little use for nostrils, except for breathing. Thus Nature, ever on the watch to economize, has re-
duced these organs, in such birds as pelicans and cor-
morants, and, at least in the adults, the nostrils are com-
pletely filled up with bone and horn. The Sen 207 The Sense of Sight The Sense of Sight Birds, so wonderful and interesting in all their structure
and life, have that most treasured of all the senses —
sight — so highly developed that there is nothing with
which we can compare it among living creatures. With
our great telescopes we can see to a greater distance than
any bird; with the high-power lenses of our microscopes
we can distinguish infinitely smaller objects than any
feathered creature is capable of perceiving, but where
else on the earth is there an organ of vision which in a
fraction of time can change itself from telescope to micro-
scope; where is the eye that, seeing with wonderful clear-
ness in the atmosphere, suddenly adapts itself to the re-
fraction of water, or (less slowly, although no less surely)
to the darkness of night? Next to our powers of reasoning, we value sight above
all things, and fortunate indeed should we be could we
but exchange our imperfect vision for sight like that of
an eagle! Little need of spectacles or binoculars has he,
for the perfection of his eye enables him to become near-
sighted or far-sighted at will. sighted or far-sighted at will. "The eye," says Professor Coues, "is an exquisitely
perfect optical instrument, like an automatic camera
which adjusts its own focus, photographs a picture upon
its sensitized retinal plate, and telegraphs the molecu-
lar movements of the nervous sheet to the optic 'twins'
of the brain, where the result is translated from the phys-
ical terms of motion in matter to the mental terms of
consciousness. But no part of the nervous tract, from The Bird 2 the surface of the retina to the optic centre, sees or knows
anything about it, being simply the apparatus through
which the bird looks, sees, and knows. In this Class of
vertebrates the optic organs, both cerebral and ocular,
are of great size, power, and effect; their vision far tran-
scends that of man, unaided by artificial instruments, in
scope and delicacy. The faculty of accommodation, that
is of adjusting the focus of vision, is developed to a marvel-
lous degree; rapid, almost instantaneous changes of the
visual angle being required for distinct perception of
objects that must rush into the focal field with the
velocity at least of the bird's flight. Observe an eagle
soaring aloft until he seems to us but a speck in the blue
sky expanse. The Sense of Sight He is far-sighted, and, scouring the earth
below, descries an object much smaller than himself,
which would be invisible to us at that distance. He
prepares to pounce upon his quarry; in the moment re-
quired for the deadly plunge he becomes at once near-
sighted, seizes his victim with unerring aim, and sees well
how to complete the bloody work begun. A humming-
bird darts so quickly that our eyes cannot follow him,
yet he instantaneously settles as lightly as a feather upon
a tiny twig. How far off it was when first perceived
we do not know; but in the intervening fraction of a
second the twig has rushed into the focus of distinct
vision, from many yards away. A woodcock tears
through the thickest cover as if it were clear space, avoid-
ing every obstacle. The only things to the accurate per- The Sen 209 thousands of birds are annually hurled against these
objects to their destruction." objects to their destruction."
A bird's eye is very large in proportion to the size of
its head, and is correspondingly perfect and delicate in its
workings. It rests in a deep cavity hollowed out of the
skull, and is protected by soft cushions of fat and controlled
by bands and pulleys of muscle which control its motions. Looking closely at the eye of a live bird, we at once
remark its brightness — that alertness of expression which
so truly reflects the virile life of these creatures. The
eye, more than any other part of a living organism, is
an index to the relative power of its intelligence — more
surely than all the other facial features taken together. The eyes of a sloth are expressionless black spots, and
even those of an orang-utan are bleary and watery. But a crow or magpie, or any other bird you may choose,
though with horny, shapeless lips, nose, and mouth, looks
at us through eyes so expressive, so human, that no won-
der man's love has gone out to feathered creatures through-
out all his life on the earth. A dog is a four-legged, hairy
animal with the eyes of a bird. The eye of a bird appears perfectly round, and is
composed of a central area of black, encircled by a ring,
sometimes hardly distinguishable from the inner divi-
sion, or again it may be highly coloured. The Sense of Sight The circular
centre or pupil is always of a uniform black, and no won-
der, for "it is not a thing — it is the hole in a thing." As
when we look through the lens of a camera, only the
blackened inside of the bellows is reflected to us, so in
the eye of a bird, the delicate living lens, itself invisible, The Bird 2 reflects the black pigmented tissue at the back of the eye-
ball. The image passes through this lens and is thrown
upon the curtain of jet, and here the brain nerves find it
and know it — how, we cannot even guess. and know it — how, we cannot even guess. If the eye-camera of the bird has no long bellows to
focus out and back, it has something infinitely better —
the coloured ring or iris which surrounds the pupil. We FIG. 151.— Head of living Sloth. FIG. 151.— Head of living Sloth. are all familiar with the way the oval iris of a cat nar-
rows to a slit in bright sunlight and broadens at night
to let in all the light possible. Look closely at the eye
of an owl or parrot, even in broad daylight, and the
circle of the iris will be seen to contract and enlarge at the
will of the bird. We have always been inclined to pity
the poor "blind" owl during the daytime, but the truth
is that, because of this power of adjustment, almost The Senses 21 I all owls can see very well, even in sunshine, although of
course their eyes are especially adapted for use in the dim
light of the evening and of the stars. In Nova Scotia I have
noticed Barred Owls flying about and feeding at noonday. FIG. 152.— Duck Hawk. (Courtesy of N. Y. Zoological Society.)
Compare the alert expression with the sloth in Fig. 151. FIG. 152.— Duck Hawk. (Courtesy of N. Y. Zoological Society.)
Compare the alert expression with the sloth in Fig. 151. Birds have well-developed lachrymal glands, although
it is seldom that they actually shed tears. Still I have
seen a flamingo in a flying-cage weeping copiously from
terror, anticipating all sorts of torture from a harmless The Bird 2 condor which was playfully galloping around the fright-
ened bird. The Sense of Sight Millions of years ago, in the geological period of time
known as the Jurassic, there existed gigantic sea-lizards,
which we call Ichthyosauri. All we know of them we
have learned by study of their fossil bones which, through
the ages, have been preserved in rocks. One notable FIG. 153. — Brown Thrasher with eyes wide open. FIG. 153. — Brown Thrasher with eyes wide open. thing about them was the great size of their eyes — meas-
uring as much as twelve and fourteen inches across. These orbits were surrounded by a series of bony plates,
and in certain birds of to-day we find a similar circle of
small overlapping bones. To make the simile between a camera and an eye hold
good, we must show that the latter is provided with a The Senses FIG. 154.— Brown Thrasher with nictitating membrane drawn. FIG. 154.— Brown Thrasher with nictitating membrane drawn. FIG. 155. — Same with eyelids closed. FIG. 155. — Same with eyelids closed. The Bird 2 shutter, and in fact our bird has not one, but three —
eyelids we call them. So "between winks" all day our
bird is taking snapshots, inconceivably more perfect and
continuous than any cinematograph ever produced. We
have but two eyelids, and every time we wink these
shoot toward each other, moisten the surface of the eye-
ball, clear it of dust, and are back in their places so quickly
that we are not aware of any interruption of our vision. The upper lid has most to do with covering the eye. In
almost all birds this condition is unusual and the lower
lid comes far upward over the eyeball. Perhaps the most
notable exception to this is among the Great Horned Owls,
where the action of the two lids is like that of our own. When birds are sleepy these lids close, but usually
in winking, the third eyelid, or nictitating membrane, alone
is drawn across the eye. This lid is a delicate, semi-
transparent sheet of tissue, which, when not in use, lies
snugly packed away in folds at the inner corner of the
eye, held back out of sight by its own elasticity. It is
drawn across the front of the eye by a slender thread of
tendon which is suspended, pulley-like, from a muscle
which keeps it from pushing against the optic nerve. The Sense of Sight When you see an owl in the daytime with eyes dull
and glazed, this third eyelid is drawn partly across them,
diluting the strong glare of light and yet enabling the
bird to distinguish much that is going on. When an eagle
turns his head upward and looks full at the sun, it is not
" unwinkingly, " but with the help of this eyelid shield. " unwinkingly, " but with the help of this eyelid shield. It is interesting to know that this membranous lid
is found in many other creatures, from sharks to monkeys, The Senses although usually much less perfectly developed than it
is in birds. Alligators, however, have it fully functional. In the inner corner of our own eyes we may detect a trace
of it, useless to us, but showing that far back in dimly
imaginable geological epochs our forebears had need of a
third eyelid. FIG. 156. — Vestige of nictitating membrane in a human eye. FIG. 156. — Vestige of nictitating membrane in a human eye. The Sense of Hearing "The Gauls," says Livy, " having discovered that the
rock Carmentalis was accessible, one night when it was
pretty clear, sent a man to examine the way, without
his arms which were afterward handed to him. Others
followed, lifting and assisting each other, according to
the difficulties which they encountered in the ascent, till
they reached the summit. They proceeded with so much
silence that neither the sentinels nor even the dogs, ani-
mals usually so vigilant as to be aroused by the slightest
noise, took any alarm. They did not, however, escape
the notice of the geese, which, being sacred to Juno,
had been fed by the Romans notwithstanding the famine
caused by the siege. This saved the capitol; for, by The Bird 2 their cackling and beating their wings, they roused Mar-
cus Manlius, a brave soldier and formerly consul, who,
snatching up his arms and giving the alarm, flew to the
ramparts, set upon the Gauls, and by precipitating one
of them over the rocks terrified the rest so much that
they threw down their arms." So also Pliny, ^Elian, and
Columella vaunt the hearing of Geese. But leaving leg-
endary lore, it is certain that birds would not have the
power of producing the most varied as well as the sweet-
est sounds in all Nature, had they not been provided
with powers of hearing, correspondingly acute and dis-
criminating. The organ of hearing is complicated and there are
many points about it which are still mysteries to scientists. The flap of skin to which we give the name of ear is
entirely absent in birds, and indeed in ourselves is a very
unimportant part of the auditory apparatus, serving
only as a collector of sound-waves. The opening of the
inner ear on each side ol the head, in birds, is usually
protected by a cover of feathers which are bristle-like,
partly denuded of barbicels, doubtless to avoid any muf-
fling of sound-waves. In owls this opening is of very
large size and protected by a movable flap of skin which
may serve to aid in focussing the sounds from below —
a very useful function to an owl at night, silently wing-
ing its way over field and meadow in search of mice and
other terrestrial prey. The Sense of Hearing A rather singular fact is that in
many owls the two ear-openings are unlike, one being
larger and of a different shape from the other, and this
asymmetry extends even to the form of the skull itself. The Senses The Senses Two membranes are stretched across the ear-tube,
and between these a tiny bone, the columella, is sus-
pended, taking the place of the chain of three bones in
the ear of a mammal. When sound-waves strike against
the outermost membrane, or drum of the ear, vibrations
are transmitted by the little bony suspension bridge to
the inner membrane, and this in turn troubles the fluid FIG. 157. — External ear of Barred Owl. FIG. 157. — External ear of Barred Owl. which fills the inner ear. The hair-like endings of the
nerve of hearing are affected by the vibrations of the
fluid and thus is hearing accomplished. Rather say,
thus the disposition of the physical components of the
ear may be explained; but how anything more than the
monotone of a sea-shell's cavity is translated to the brain,
no one can say. The fluid contained in three semicircular canals, situ- The Bird 2i 8 ated in the inner ear — which occupy the three planes of
space, — exercises a most important function, that of equi-
libration. They have been compared to the glass tube
filled with water and a shifting bubble of air, by centring
which a surveyor knows his instrument is perfectly level. which a surveyor knows his instrument is perfectly level. If these canals be injured or cut, the bird loses all
control of his actions ; if a certain one of the three canals
suffers, the bird moves its head rapidly sideways and
spins around in a circle; if another of the trio is by an
accident severed, the motion of the head is back and
forth, and the bird is compelled to execute forward som-
ersaults; when the third of these canals is cut the bird
continually falls backward. In reptiles and mammals
the same thing occurs, so the wisdom of Nature in pro-
tecting these delicate organs by a sheafh of hard bone
is very apparent. The Sense of Taste and Touch "The hands of birds being hidden in the feathers
which envelop the whole body, — their feet and their lips
and usually much, if not all, of their tongue, being
sheathed in horn, — these two faculties would appear to
be enjoyed in but small degree." be enjoyed in but small degree."
The sense of taste is probably the least developed
of all. The nerves which find their way through the
pores of the bill and tongue are more properly those of
touch than of taste, and this seems the more credible
when we consider the food of many birds, which is swal-
lowed entire, besides being so hard that nerves of taste
would be useless. Parrots and ducks, with their fleshy The Sen 219 tongues and ample membranes of the mouth, doubtless
possess this sense to a considerable degree, while in birds
which are exclusively fish-eaters we may expect to find
taste least developed, the character of their food pre-
cluding all need for this faculty. But from no bird is taste entirely absent, as we may
easily see by presenting some nauseous insect, which
will be instantly rejected with very evident signs of dis-
gust, the bird wiping its bill on a branch and shaking
its head violently. The sense of feeling, although much deadened by the
feathery and horny character of a bird's integument, is
most active at the tip of the tongue and the beak. At
the base of the feathers, especially those of the wings and
tail, tactile nerves are found, so that even a touch on the
tips of the feathers awakens a response in the nervous
system. system. The delicacy of the tactile touch is remarkable in those
long-billed birds which seek their food in the muddy
bottom of shallow water, detecting by means of their
sensitive bills the presence of worms and snails, — aided
little or not at all by eyesight. In the woodcock and
apteryx this dependence on the senses of touch and smell
has even wrought a change in the position and character
of the eyes. The upper mandible of the woodcock is
probably unique in being so sensitive and mobile that the
distal third can be curved some distance upward, the base
of the two mandibles remaining close together. The Sense of Taste and Touch This is
an admirable provision by which, when the bird has driven
its beak deep down into the moist soil, it may feel about I
O
I
g
2 I
O
I
g
2 The Senses 221 and seize the earthworm for which it is seeking. The
eyes have become unusually large in consequence of its
nocturnal habits and in addition are placed far back
upon its head, permitting a clear lookout for danger,
above and even behind, while probing with its head held
close to the ground. FIG. 160 FIG. 159. FIG. 160
FIG. 159. — Photograph of living Woodcock with bill closed. FIG. 160 — Same with bill open, showing mobility of upper mandible. The bird
is thus enabled to feel about and seize the earthworms deep down in the mud. FIG. 159. FIG. 159. FIG. 159. FIG. 160 FIG. 159. — Photograph of living Woodcock with bill closed. FIG. 160 — Same with bill open, showing mobility of upper mandible. The bird
is thus enabled to feel about and seize the earthworms deep down in the mud. FIG. 159. — Photograph of living Woodcock with bill closed. FIG. 160 — Same with bill open, showing mobility of upper mandible. The bird
is thus enabled to feel about and seize the earthworms deep down in the mud. Thus ends our brief survey of the five senses; that of
smell taking note of minute particles of matter diffused
in the air; sight and hearing depending on vibrations of
the atmosphere; the sense of taste detecting matter which
is dissolved in water, and that of feeling making the bird
cognizant of the qualities of bodies by actual contact. o>
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£ BEAKS AND BILLS I IE a man's hands and arms tightly behind his
back, stand him on his feet, and tell him that
he must hereafter find and prepare his food,
build his house, defend himself from his enemies and
perform all the business of life in such a position, and
what a pitiable object he would present! Yet this is not
unlike what birds have to do. As we have seen, almost
every form of vegetable and animal life is used as food
by one or another of the species. Birds have most in-
tricately built homes, and their methods of defence are
to be numbered by the score; the care of their delicate
plumage alone would seem to necessitate many and varied
instruments: yet all this is made possible, and chiefly
executed, by one small portion of the bird — its bill or
beak. If one will spend an afternoon at a zoological park,
or with any good collection of live birds, watching the
ways in which the bills of the various species are used,
one will not boast of his own accomplishments, when
it is realized how much more, comparatively, the bird is
able to achieve with the aid of two projecting pieces of
horn. 223 The Bird 224 More than a single volume could be filled with in-
teresting facts about the bills of birds and the uses to
which they are put, — hardly any two species using their
beaks in a similar manner. The general way in which
the vast subject of the adaptation of the bird to its needs
and to its surroundings is treated in this volume will,
it is hoped, be a stimulus to the reader to observe for
himself, — to discover the thousand and one facts to FIG. 162. — Beak of Snapping-turtle. (Courtesy of N. Y. Zoological Society.) FIG. 162. — Beak of Snapping-turtle. (Courtesy of N. Y. Zoological Society.) which Nature has not yet given us the key. Our lan-
guage is too often lacking in phrases expressing delicate
shades of meaning, and thus we are compelled to identify
structures among the creatures which rank below us
with portions of our own anatomy corresponding only in
relative position or a general vague likeness of function. We are accustomed to speak of the mouth of a starfish,
the arms of a sea-anemone, the foot of a snail: in these
respective cases, structures specialized for receiving food, Beaks and Bi 225 reaching about, or for progression being understood. But
no one would think of alluding to a bird's lips or nose ; both
are included in the terms beak, or bill, and nostrils. The finding and securing of food being the most im-
portant problem which birds have to solve for themselves,
it is for these purposes, and especially the last mentioned,
that we find bills most adapted. This is so universally FIG. 163. — Bill ot American Raven. FIG. 163. — Bill ot American Raven. the case that we may often judge accurately of the kind
of food of a certain bird from a glance at its beak. As is the case with so many other avian structures,
the horny, toothless beak or bill is duplicated elsewhere
in Nature only in a group of reptiles, the turtles and tor-
toises, whose mandibles furnish a splendid example of
parallel evolution. In certain of those long-extinct Dinosaurs, such as The Bird 2 Triceratops, an interesting transitional condition is found. The front of the mouth was beak-like and horny, while
farther back were the masticatory teeth. The Bird Starting with the generalized beak of the Archce-
opteryx, which, we remember, was furnished with teeth,
we are almost at a loss in which direction to turn,
so many and so varied are the beaks of modern birds. No trace of teeth, however, is to be found in the adults
of any of them. The bill of a crow or raven and, to a
lesser extent, that of his near relatives, the jay and the
blackbird, is perhaps in shape most like that of the 'bird
of old,' and is suited to the many purposes which the
varied life of these intelligent birds requires. The crow or the raven is an excellent example of a
modern bird with a remarkably generalized diet, in striking
contrast to those birds whose bills show them to be fitted
for feeding only on some strictly defined food. With
his strong, ample beak the crow can dig up recently
planted corn, or crack the hard shells of acorns; he en-
joys stealing the eggs and the young birds of thrushes,
orioles, sparrows, warblers, and quail, and I have seen
a crow chase, capture, and carry off a half-dozen wild Mal-
lard ducklings in one morning ! These birds are, in ad-
dition, able to capture insects of all kinds, besides pick-
ing berries, and ducking their heads under water in quest
of the shrimps which live in tide-pools. In short, their
bill serves them well in procuring many kinds of food1,
from earth, water, or tree; as well as in carrying great
quantities of sticks, which they use in the construction
of their nests. These birds are so skilful with their Beaks and Bills FIG. 164.— Beak of Gannet. FIG. 164.— Beak of Gannet. FIG. 165. — Beak of Cormorant. Birds closely related, but procuring food in different ways. FIG. 165. — Beak of Cormorant. Birds closely related, but procuring food in different ways. The Bird 2 beaks that a new trick is learned in a very short time. In captivity a crow, when it thinks no one is watching,
will often take a morsel of food, thrust it beneath a piece
of sod, and cover it up with grass, almost with one
motion of the beak. The Bird Functional or adaptive radiation is beautifully il-
lustrated by the beak of a gannet, cormorant, snake-
bird, and pelican — birds which are closely related to
one another structurally, also having in common a fish
diet, swallowing their prey whole. The gannet's beak
is thick and very strong, and along the inner edge is a
series of fine serrations pointing backward. The bird
dives, from a great height, into the water and seizes the
fish in a grip of steel. The upper mandible of the cor-
morant is furnished with a large, sharp hook, with which
the bird gaffs its prey, pursuing it under water. The
snake-bird, or darter, has a bill like a needle, with which
it spears the fish, impaling it through and through; while
the pelican, because of its great pouch least vicious of
all in its methods, simply engulfs the fish, the water in
which it is swimming and all, then straining out the liquid,
tosses the unfortunate into the air and swallows it
head first. The under mandibles of this bird are long
and pliable and so arranged that they can bend far apart,
thus making of the great bag of skin beneath the bill
and throat an admirable fish-trap. and throat an admirable fish-trap. This is one of the many instances where several closely
related species, with needs so similar that there is danger
of fatal competition, are able to exist in great numbers
and to avoid all undue struggle for existence by having Beaks and Bills Beaks and Bills
FIG. 166.— Beak of Snake-bird. FIG. 167.— Beak of Pelican. Birds related to each other and to Figs. 164 and 165, but with different feeding habits. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. FIG. 166.— Beak of Snake-bird. FIG. 166.— Beak of Snake-bird. FIG. 166.— Beak of Snake-bird. FIG. 167.— Beak of Pelican. Birds related to each other and to Figs. 164 and 165, but with different feeding habits. FIG. 167.— Beak of Pelican. Birds related to each other and to Figs. 164 and 165, but with different feeding habits. The Bird 23° each an individual method — a niche into which it fits
perfectly in the great scheme of Earth's hungry creat-
ures. The snake-bird's prey is in the water of dense FIG. 168. — Brown Pelican catching fish, showing bag-like distension of lower
mandible. (Sanborn: photographer. Courtesy of N. Y. Zoological Society.) FIG. 168. — Brown Pelican catching fish, showing bag-like distension of lower
mandible. (Sanborn: photographer. Courtesy of N. Y. Zoological Society.) swamps and bayous; cormorants and pelicans amicably
share inland lakes and tidal waters; while the haunt of
the gannet is the high seas. swamps and bayous; cormorants and pelicans amicably
share inland lakes and tidal waters; while the haunt of
the gannet is the high seas. Even more closely related to each other are terns Beaks and Bills 231 and Black Skimmers. Except in their bills these birds
are almost identical in structure, but the bill makes a and Black Skimmers. Except in their bills these birds
are almost identical in structure, but the bill makes a FIG. 169.— Bill of Tern. FIG. 169.— Bill of Tern. FIG 170.— Bill of Skimmer
Closely related birds which differ in their feeding habits. FIG 170.— Bill of Skimmer
Closely related birds which differ in their feeding habits. vast difference in the appearance of a bird, as is very
apparent when these two species are seen flying about The Bird 232 •
together on their breeding-grounds, — low, sandy islands
along our coast. The small, delicately pointed beak of
the tern finishes off its neat appearance, and the entire
bird is the personification of grace, as it dashes through
the air, or plunges headlong into the sea, — to rise almost
immediately with a
small fish in its beak. •
the air, or plunges headlong into the sea, — to rise almost
immediately with a
small fish in its beak. The beak of the
skimmer lends a heavy
aspect to the whole
bird. It is long
and high, and
the lower man-
FIG. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. 172.— Two-year-old Skim-
mer, reared in captivity,
showing abnormal growth
of mandible, due to lack
of friction against water. PIG. 171. — Bill of young Skimmer, showing under
mandible already slightly the longer. The beak of the
skimmer lends a heavy
aspect to the whole
bird. It is long
and high, and
the lower man- F
PIG. 171. — Bill of young Skimmer, showing under
mandible already slightly the longer. FIG. 172.— Two-year-old Skim-
mer, reared in captivity,
showing abnormal growth
of mandible, due to lack
of friction against water. PIG. 171. — Bill of young Skimmer, showing under
mandible already slightly the longer. dible extends a full inch beyond the upper. Both are
as thin and as pliable as paper-knives. A unique method
of obtaining food is the secret of this apparent deformity:
the strong wings of the bird enable it to fly very close to dible extends a full inch beyond the upper. Both are
as thin and as pliable as paper-knives. A unique method
of obtaining food is the secret of this apparent deformity:
the strong wings of the bird enable it to fly very close to Beaks and Bills. the surface of the water, so close in fact that the lower
mandible dips below the surface, thus ploughing a zig-
zag furrow and catching up any organisms, shrimps or
fish, which chance to be floating on the water. FIG. 173. — Bill of Merganser, a fish-eating duck. FIG. 173. — Bill of Merganser, a fish-eating duck. FIG. 174. — Bill of Shoveller Duck, a bird which strains its food from the mud. FIG. 174. — Bill of Shoveller Duck, a bird which strains its food from the mud. Among ducks, we find those which feed on fish, and
those which sift their food from the mud at the bottom
of ponds, and these differ radically in respect to their
beaks. The fish-eating merganser has perhaps, of all
living birds, the nearest resemblance to a toothed beak. The Bird 2 The deep serrations, however, are but indentations in
the substance of the strong, narrow bill of the bird. When once in this saw-like grasp, the most slippery fish
is helpless. The beak of the Shoveller Duck shows how
well Nature has provided for its wants. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. The beak is
arched and spatulate, while the sensitive epidermis is pro-
longed at the edges into a series of comb-like teeth,—
analogous to the whalebone in the mouth of a whale. Through this sieve the water is drained out, leaving
entangled the edible worms and insects. If we should elevate our Shoveller Duck, placing him
on long, slender legs and providing him with a corre-
spondingly long neck, he would indeed be in a predica-
ment, since only the tip of his beak could be brought
to bear in feeding. Now a flamingo is really a long-
legged duck, which feeds in much the same way as the
Shoveller, and the difficulty mentioned is overcome in
a most ingenious way. The mandibles are bent down-
ward, almost at right angles, so that, when the head
reaches the ground, not the tip but the whole inverted
bill is in a position to sift out food. To meet the reversed
condition, the lower mandible is deeply arched, instead
of the upper as in the Shoveller Duck. We are able to follow the probable evolution of such
remarkable beaks as those of the flamingo and skimmer
by observing the growth of this organ in any individual
from the time when the bird hatches from the egg until
it is full-grown. In the very young flamingo chick there
is no sign of the subsequent deflection, the mandible
being short, perfectly straight, and rather slender. As the FOUR STAGES IN THE DEVELOPMENT OF THE BILL OF THE
FLAMINGO. FIG. 175. — Young bird in down. FIG. 176. — Young in gray plumage. FIG. 175. — Young bird in down. FIG. 175. — Young bird in down. FIG. 175. — Young bird in down. FIG. 175. — Young bird in down. FIG. 176. — Young in gray plumage. FIG. 176. — Young in gray plumage. FIG. 177. — Young in gray plumage, later stage. FIG. 177. — Young in gray plumage, later stage. FIG. 178. — Adult living bird. 235 FIG. 178. — Adult living bird. FIG. 178. — Adult living bird. FIG. 178. — Adult living bird. The Bird 2 bird at first feeds upon regurgitated food, taking it drop
by drop from the bill of the old bird, it of course has
no need of the curved beak of its parents. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. Later, when
its bill has increased in length and has begun to be marked
by the ultimately sharp angle, the birds begin to sift
from the coral mud the small mollusks of which their
food consists. Until its wings are full-feathered the young skimmer
is compelled to limit its wanderings to the sand-dunes
along the shore near its nest. Thus, although at birth
the lower mandible is a trifle longer than the upper, yet
even when the birds are half-grown the disparity in length
between the two mandibles is but slight. Later, when
the young bird is able to join its parents in their skimming
of the seas, the lower mandible quickly attains its full
development. The friction of the water upon the bill
must be considerable, as in a skimmer which I have had
for years in captivity, the lower mandible grew remark-
ably fast, measuring 6f inches from base to tip when
the bird was eighteen months old. Herons and ibises, through all the years, sought their
food in much the same places as have ducks ; the straight-
billed herons seizing their living prey with a single light-
ning dart, as it swims past them; the spoonbills spatter-
ing in the shallows; and the curved-beaked ibises prob-
ing every crevice along shore. The spoonbills swing
their necks and heads from side to side, as they walk
slowly through the water, gleaning their food with the
motion of a mower wielding his scythe. Two of the
herons are interesting enough to hold our attention for Beaks and Bills a moment. The common Black-crowned Night Heron
is abundant throughout most of North America, and he
fishes in legitimate heron fashion; but his near relative,
the Boat-billed Heron, is a more tropical species. In
voice, appearance, and structure there is little to choose
between the two birds, — except that the latter has a broad,
scoop-like beak, — a pelican's fish-trap in miniature, which
seems to answer every requirement of this strange-look-
ing bird. From the
muddiness of the water
in the tropical swamps
from which I have
flushed these birds, it
seems probable that much
of their food may be
lesser fry than fish. FIG. 179.— Bill of Great Blue Heron. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. Pebbles and shells,
which shelter so many
toothsome morsels along
the shallows of our sea-
shore, offer sumptuous
feasts to birds furnished with beaks adapted to prying
and probing, and we find all sorts of sizes and shapes. A collection of bills of various wading-birds would look
like a complete set- of surgical tools ! There is the stilt,
whose bill is almost straight; the ibis, with mandibles
curved downward to probe the crevices between the
pebbles on which he stands; the avocet has a pair of
recurved pliers, which search out the worm or snail in
the deepest fissures ahead of him. At the slightest touch
FIG. 179.— Bill of Great Blue Heron. FIG. 179.— Bill of Great Blue Heron. The Bird 238 of such a beak, the oysters and other large bivalves
close with a snap, defying these birds to penetrate their
living armour. Indeed, more than once a gull or wader
has rashly pecked at the sweet flesh, when the two tight-
fitting doors have suddenly closed, pinning the bird help- FIG. 180. — Boat-billed Heron. Figs. 179 and 180 represent birds with slightly different feeding habits. FIG. 180. — Boat-billed Heron. Figs. 179 and 180 represent birds with slightly different feeding habits. less and holding it captive despite its struggles, until
the rising tide has ended its life. But along comes a bird, well named Oyster-catcher,
and woe to the mollusks now. It allows them to close
tightly upon its bill, the mandibles of which are thin
like blades, many years antedating man's oyster-knives. The mollusk is wrenched free by the sturdy bird, car-
ried from the water still gripping the bird's bill, and is Beaks and Bills FIG. 182.— White Ibis, showing
curved bill. FIG. 181. — Spoonbill, with spatulate
mandibles
FIG. 182.— White Ibis, showing
curved bill. FIG. 181. — Spoonbill, with spatulate
mandibles FIG. 182.— White Ibis, showing
curved bill. FIG. 181. — Spoonbill, with spatulate
mandibles FIG. 183.— Bill of Avocet, recurved for probing. FIG. 183.— Bill of Avocet, recurved for probing. The Bird 240 then pried open and eaten. The bill of this bird shows
the wear and tear of forcing apart the shells, and it is
sometimes slightly bent to one side. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. The short-billed
gulls are denied the power of opening these oysters and
mussels, but they sometimes get an unlawful feast by fol-
lowing up and robbing the Oyster-catchers of the shells
which the latter have opened. The bill of the Shell Ibis of India may be likened to
an ordinary lemon-squeezer, having a cavity in which FIG. 184. — Bill of Oyster-catcher; used for prying open the shells of mollusks. FIG. 184. — Bill of Oyster-catcher; used for prying open the shells of mollusks. the half -lemon rests before it is compressed. When
the mandibles of this bird are closely opposed the central
portion of the beak gapes slightly. In this cavity the
bird firmly holds the shells of the land-snails upon which
it feeds, until it can bring the pressure of both mandibles
to bear and so crush the shell of the mollusk. the half -lemon rests before it is compressed. When
the mandibles of this bird are closely opposed the central
portion of the beak gapes slightly. In this cavity the
bird firmly holds the shells of the land-snails upon which
it feeds, until it can bring the pressure of both mandibles
to bear and so crush the shell of the mollusk. The asymmetry of the bill — as seen in the Oyster-
catcher — is not accidental, but constant, in the Crook-
billed Plover of New Zealand. In this bird the bill is Beaks and Bi 241 permanently bent to the right, a beautiful adaptation
to help the bird in its search for insects, which, in the
dry country that it inhabits, are found almost entirely
under stones. As a rule, beaks are rather immovable throughout
their length, but in the woodcock, and to a less extent FIG. 185. — Bill of Crook-billed Plover, for probing under stones. FIG. 185. — Bill of Crook-billed Plover, for probing under stones. in the Dowitcher Snipe, the extremity of the upper man-
dible can be raised some distance (Figs. 159, 160). This
extreme sensitiveness is especially necessary, as the eyes
of the woodcock are placed very far back on the top
of its head, and are of little or no use in seeking food. What an interesting study the various beaks of land
birds would offer, were we able to devote to them the The Bird 242 space which they deserve! FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. They defy classification and
refuse to be arranged in any linear sequence. The ma-
jority of those birds which have their beaks armed with
a strong hook feed upon living prey, — from the great
mandible of the Golden Eagle to the tiny vireo, which
snaps up the dancing gnats. sna
up the da
gn
The owls and the parrots, which, by the way, are
much more closely related than most of our classifications
would indicate, have bills very much alike, and afford FIG. 186. — Bill of Golden Eagle, hooked for tearing prey. FIG. 186. — Bill of Golden Eagle, hooked for tearing prey. a striking example of two large related groups of birds
whose diet has become radically unlike, although even
in this case "blood will tell" and the Kea Parrot slips
back into carnivorous habits with ease. Owls tear their prey apart with their beaks, or swallow
it entire, but parrots gnaw and gnaw upon their nuts
and seeds, reducing their food to powder. This grind-
ing and rasping is aided by several file-like ridges which
many parrots have within their beaks. The hinging
of the upper mandible with the skull is more evident in a Beaks and Bi 243 parrot than in any other bird. This arrangement allows
much freedom of motion. parrot than in any other bird. This arrangement allows
much freedom of motion. It is not clearly known what use the immense beaks
of toucans may serve, although there seems little excuse
for this ignorance in those who know the birds in their
native haunts. The delicate, spongy texture renders the FIG. 187. — Toucan, showing enormous bill used perhaps for reaching fruit on
the tips of branches. FIG. 187. — Toucan, showing enormous bill used perhaps for reaching fruit on
the tips of branches. clumsy-looking appendages exceedingly light, and they
are usually banded or marked with brilliant hues, — blue,
yellow, red, brown, green, or black. But light as the
beaks are in these birds, in the unrelated but similarly
monstrous-beaked hornbills the weight must be con-
siderable, for the first two vertebrae of the neck in these The Bird 244 latter birds are fused together, to yield a firmer support
for the muscles of the neck. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. Chimney Swifts and hummingbirds both feed upon
insects and are rather closely related to each other, but
here again the most decided difference is to be found in
their bills. The broad, flattened mandibles of the swifts FIG. 188. — Bill of Toucan; cut open to show its light, spongy structure. FIG. 188. — Bill of Toucan; cut open to show its light, spongy structure. open wide, as the birds dash through the air, engulfing
gnats and flies with wonderful skill; while the humming-
birds, as we all know, probe the deepest calyxes. Could
two bills more unlike be imagined? In very young hum-
mingbirds the bill is short and broad, very like the swift
type, and later its long and slender shape is acquired Beaks and Bi 245 very rapidly, as we can see in Figs. 190 and 191. There
are many resemblances between hummingbirds and in-
sects, due entirely to the similarity in their feeding habits. Certain flowers are especially adapted in structure to
attract certain bees or moths, which in return unconsciously
cross-fertilize the blossoms; and certain of the various
bills of hummingbirds reflect the exact contour of the
corollas in which the birds seek their food. Among
hummingbirds the various shapes of bills of other groups
are reproduced again. Humming through the air about
us in the tropics speed miniature avocets, ibises, stilts,
mergansers, and we realize, as never before, the never-
ending devices of Nature, providing for the needs of all,
from the greatest to the least; endless patterns paral-
leling each other, but never identical. Indeed, in the
great family of South American birds known as Wood-
hewers the diversity in shape, size, and direction of bills
is so great that it seems as if not a niche, or crack, or hollow
in the bark of any tree in the forest where these birds
abound would afford a safe retreat to an insect! It remains to mention the woodpecker's bill, which
is used chisel-like, for excavating his home as well as
in boring for grubs. With his beak the nuthatch ham-
mers his acorns, and the tailor-bird sews his nest. The
thick conical beaks of all sparrows and finches are for
cracking seeds; while the weaker, more slender beaks
of warblers, thrushes, and wrens reflect a diet of insects. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. Among the finches is a group of several species which,
by a thrust of the bill, have at their command a new
source of food, one which there are none to dispute with The Bird 246 FIG. 189. — Bills of adult Hummingbird and Chimney Swift, showing great dis-
similarity in form, due to different methods of procuring food. FIG. 189. — Bills of adult Hummingbird and Chimney Swift, showing great dis-
similarity in form, due to different methods of procuring food. FIG. 190. — Bills of young Rufous Hummingbirds, showing swift-like character. (Photograph by Finley & Bohlman.) FIG. 190. — Bills of young Rufous Hummingbirds, showing swift-like character. (Photograph by Finley & Bohlman.) 191. — Slightly older Hummingbirds, with bills half as long as the adults. (Photograph by Finley & Bohlman.)
247 FIG. 191. — Slightly older Hummingbirds, with bills half as long as the adults. (Photograph by Finley & Bohlman.) The Bird 2 them. Both mandibles of the crossbill are curved into
sharp hooks which cross one another, either to the right
or left, thus forming a unique pair of pliers, with which
the bird pries out the seeds shut tight behind the over-
lapping scales of pine-cones. FIG. 192.— Two extreme types of Hummingbirds' bills adapted for insertion
in flowers with shallow and with deep calyxes. FIG. 192.— Two extreme types of Hummingbirds' bills adapted for insertion
in flowers with shallow and with deep calyxes. The beak of a bird is always growing, and in captivity,
from lack of proper use, the mandibles sometimes grow
to a great length, and, if not trimmed, will often inter-
fere with the bird's feeding. fere with the bird's feeding. Perhaps the most remarkably adapted beaks in the
world are those of the male and female Huia birds — Beaks and Bi 249 natives of New Zealand — in which not only is the bill
of the species designed for a special method of procuring
food, but the bills of the two sexes are very different in
form and use, and complement each other's methods. Concerning the peculiar use of the bill in the Huia birds, FIG. 193. — Bill of Purple Finch and Crossbill compared; the latter specialized
for extracting seeds from pine-cones. FIG. 193. — Bill of Purple Finch and Crossbill compared; the latter specialized
for extracting seeds from pine-cones. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. Professor Newton writes: "Its favourite food is the grub
of a timber-boring beetle, and the male bird with his short
stout bill attacks the more decayed portions of the wood,
and chisels out his prey, while the female with her long
slender bill probes the holes in the sounder part, the hard-
ness of which resists his weapon; or when he, having
removed the decayed portion, is unable to reach the grub, The Bird 25° the female comes to his aid and accomplishes what he
has failed to do." has failed to do."
The bill of a bird, besides serving in so many other
ways, is invaluable in preening the plumage, arranging
disordered feathers, drying them, and, most important
of all, in pressing out the oil from the gland on the lower
back, and with it carefully dressing all the feathers, giv- FIG. 194. — Bill of male and female Huia Birds, showing difference of the bill
in the two sexes. FIG. 194. — Bill of male and female Huia Birds, showing difference of the bill
in the two sexes. ing to them that brightness and gloss and also the water-
proof quality — so surely a sign of perfect health in a
bird. When, after the bath of a caged bird, you see the
drops roll from its feathers, literally like " water off a
duck's back," then the good health of the bird is certain. duck's back," then the good health of the bird is certain. The all-important use of the bill as a needle, shuttle,
pick and shovel, auger, or trowel in nest-building does not
concern us here, nor does its function in expressing emo-
tion, or in taking the place of the voice or of the foot. Beaks and Bi 251 All this is expressive rather of the mental than the phys-
ical life of the bird. All this is expressive rather of the mental than the phys-
ical life of the bird. Within a period of five minutes I have observed the
following uses of the beak of a parrot perching in my
study. FIG. 166.— Beak of Snake-bird.
FIG. 167.— Beak of Pelican.
Birds related to each other and to Figs. 164 and 165, but with different feeding habits. With its mandibles it picked up a sunflower
seed and comminuted it; it then hooked the upper man-
dible into a wire and swung itself along; gnawed at a nest-
ing-hole it had begun to excavate; nibbled gently at
my finger, showing affection; bit fiercely in anger and
fear at a dead snake which I presented; preened several
feathers of one wing, smoothing out all the dislodged
barbs; rattled its beak along the wires to make a sound
to attract my attention; and finally seized its water-pan
and turned it over in pure playfulness! HEADS AND NECKS H HE head of a bird is indeed a wonderful object,
when we consider its comparatively small size
and yet realize that it contains the brain, as
well as being the seat of the five senses. It also sup-
ports that most important organ the beak, which, as we
have seen, takes the place of hands and tools in the life
of its owner. Of expression, with the exception of that caused by
raising its feathers, the bird has but little; although
fear, that emotion which must needs be expressed all too
often in the life of these timid and comparatively de-
fenceless creatures, is made apparent by the dilating
eyes and the open, panting beak. The only exception
which comes to mind is the Crowned Crane, Fig. 206,
the suffusing of whose bare, white cheeks indicates the
changing emotions. Perhaps the best index is to be
found in the crest, which we will find to be developed to
a very remarkable degree. a very remarkable degree. The eyes of most birds are placed at the sides of the
head, in such a position that the bird cannot bring both
to bear simultaneously upon the same object, but is com-
pelled to turn its head and look sideways. As birds
252 s bi Heads and Necks spend so much of their time in the air, or in trees, where
danger may threaten from all sides, above or below, this
arrangement is most useful to them, giving them com-
mand of almost their whole surroundings, whereas, with-
out turning the head, we can see only ahead of us. In
much the same relative position, the two ears are placed, FIG. 195. FIG. 196. FIG. 195. — Head of Dove, with eyes at side
FIG. 196.— Head of Owl, with eyes in front Showing difference in position of
eyes in a pursued and a pursuer in Life's race FIG. 196. FIG. 195. FIG. 195. FIG. 196. FIG. 195. — Head of Dove, with eyes at side
FIG. 196.— Head of Owl, with eyes in front Showing difference in position of
eyes in a pursued and a pursuer in Life's race FIG. 195. — Head of Dove, with eyes at side
FIG. HEADS AND NECKS 196.— Head of Owl, with eyes in front Showing difference in position of
eyes in a pursued and a pursuer in Life's race and the absence of a directive outer ear renders the bird
susceptible to sounds coming from every direction. Owls, for very obvious reasons, are interesting excep-
tions to the above statements. Living most of their
active life at night, playing always the role of pursuers,
these raptorial birds have few enemies to fear; and their
subsistence depends upon the keenness of their senses
when focussed in one direction — downward. When its
strong, soft-feathered pinions carry a mousing owl over The Bird 2 field and stubble, the head, like the nose of a hound, is
held low, and, that not a rustle nor a motion of the little
field-mice may be lost, the ear-openings are turned down-
ward and the eyes look full upon the ground. Look a
Barn Owl in the face and you will see the entire cir-
cumference of both eyes, but a dove — one of the pursued
in life's race — shows in the front view only the profile
of the eyeballs. The same story is told in the eyes of
the fox and the rabbit — examples of Nature's parallels,
which are never repetitions. It is interesting to compare the eyes of owls with those
of mammals in general. With the exception of man,
and of some of the monkeys, we find that when the eyes
show but slight divergence the animal is invariably a
lover of the dusk, or is wholly nocturnal. We know that
when we are asleep, or are under the effects of ether, our
eyes tend to roll upward and outward, and now we realize
that the cause of this is the old ancestral pulling outward,
toward monocular vision, as in the fish or rabbit or dove. Our distant ancestors, far from having books or work
which focussed their attention directly in front, had
most vital need of looking out for dangers in all direc-
tions. Another adaption found in the eyes of almost all noc-
turnal birds is the great size of the orbit, fully one half
of the skull being hollowed out to receive the eyeballs. HEADS AND NECKS No degeneration of the eyes, as a result of nocturnal habits,
is recorded among birds, such as exists in moles and bats,
except in the case of the apteryx, the diminutive New
Zealand representative of the ostrich-like birds. The Heads and Necks Heads and Necks FIG. 197.— Skull of Owl. FIG. 197.— Skull of Owl. FIG. 198.— Skull of Apteryx. Showing opposite effects of nocturnal habits on the size of the eyes. FIG. 198.— Skull of Apteryx. Showing opposite effects of nocturnal habits on the size of the eyes. FIG. 198.— Skull of Apteryx. FIG. 198.— Skull of Apteryx. Showing opposite effects of nocturnal habits on the size of the eyes. Showing opposite effects of nocturnal habits on the size of the eyes. The Bird 2 small eyes of this bird become dazzled by strong light,
its food being detected by the senses of touch and smell. its food being detected by the senses of touch and smell. The eyes of the woodcock show an interesting adapta-
tion to its habits. The bird feeds at night in marshes,
probing the mud for worms and, being in frequent danger
of attack from owls or other foes, it has need of constant
vigilance. So we find that its eyes, which are large and
lustrous, are placed far back on its head and also up near
the top of the skull. Useless in guiding the bird in its
search for food, they have become altered in size and posi-
tion and so best fulfil their function of aiding their owner
to all but look through the back of its head. Even the iris of a bird's eye may share in the won-
derful colour scheme of its feathers, although the most
common hue is a hazel-brown. And in birds of two
related species or races, there is sometimes a marked dif-
ference in the colour of the iris; such, for example, as be-
tween the Red-eyed and White-eyed Vireos, or the simi-
larly named Towhees. Puffins have blue irides, pigeons
pink ones, while young Bald Eagles have brown eyes
which, in the adult, turn yellow. The eyes of Barred
Owls seem to be a deep, lustrous black, but they are really
dark brown; while the great yellow eyes of Snowy and
Horned Owls are the most brilliant bits of colour about
these birds. In cormorants the irides are a glittering
emerald-green. emerald-green. HEADS AND NECKS It might be thought that "making eyes" was con-
fined to the more frivolous of our own race, but certain
it is that, whether or not it plays a part in charming the
females, the irides of the males of a number of species Heads and Necks of birds change, at the season of courtship, from a dull
hue to some bright tint, either red, green, or yellow. Although eyelashes, as we understand them, are
merely stiffened hairs which have been inherited from
hairy-coated ancestors, yet among birds we sometimes
find lashes similar in appearance and function, but struc-
turally derived from feathers. The ostrich has well- FIG. 199. — Head of Seriema, showing eyelashes. FIG. 199. — Head of Seriema, showing eyelashes. developed eyelashes, which must be of value in helping
to exclude the dust of the desert; but why such birds
as hornbills and the Seriema should possess them we
cannot say. A savage thrusts feathers into his hair, warriors of
old bedecked their helmets with flowing plumes, the
opera hat of milady is by way of wonderful and strange
creations; but withal feathers are really beautiful only The Bird 258 where they by rights belong — upon a bird. Among
birds we find a more remarkable development of crests
than in any other class of animals. Indeed nearly all
birds have the power of slightly raising the feathers on
the head. FIG. 200.— Crest of Java Peacock. FIG. 200.— Crest of Java Peacock. Most, if not all, plumes and crests are probably orna-
mental, and, since many are more highly developed in
the male sex and at breeding-time, we must conclude
that they are of value in attracting and holding the at-
tention of the females during the period of courtship. What a list of these crests we may compile in a walk
through a zoological park! The photographs show the
grace and delicacy of these feathers, to which words can Heads and Nec 259 add nothing. Note the slender shafts which rise from
the head of the Indian Peacock, each one tipped with a
dainty feather tuft; and the variation in the crest of its
splendid cousin from Java. In one of the Birds of Para-
dise, six long, fan-tipped shafts extend backward from
the head, much longer than, but similar to, the crest of
the Indian Peacock. The California Partridge has a tiny, FIG. 201. HEADS AND NECKS — California Partridges, showing difference in crest of male and
female birds. FIG. 201. — California Partridges, showing difference in crest of male and
female birds. club-shaped crest which points in a forward direction,
and, when the bird is excited, the feathers which com-
pose it spread out, breaking into a fan. The ornament
of the Plumed Partridge is a long, sweeping plume. The
crest of the curassow is most peculiar, being composed
of curly, recurved feathers, resembling in texture and
appearance jet-black or parti-coloured shavings. club-shaped crest which points in a forward direction,
and, when the bird is excited, the feathers which com-
pose it spread out, breaking into a fan. The ornament
of the Plumed Partridge is a long, sweeping plume. The
crest of the curassow is most peculiar, being composed
of curly, recurved feathers, resembling in texture and
appearance jet-black or parti-coloured shavings. appearance jet-black or parti-coloured shavings. The nuptial plumes of the Night Heron hang far down
upon its shoulders, and the soft barbs are curved inward, The Bird 2 forming a slender tube. The glory of the Great Crowned
Pigeon is a maze of lavender lacework, — one of the most
beautiful of all crests; while the most graceful, perhaps,
is the mist of filmy whiteness which, at the slightest breath
of air, floats about the neck of the Snowy Egret, like
the mantilla of a senorita. Cockatoos are decorated
with a profusion of beautiful crests, each characteristic. These are under the complete control of the birds, and
take an important part in expressing changing moods
and emotions. The crests may lie so flat as to be ordi-
narily invisible, when, in a flash, the whole head is sur-
mounted by an auricle of colour or whiteness. An ex-
cited Leadbeater Cockatoo is a wonderful sight. Before
the crest is raised, all that is visible is a single, rather
elongated white feather, but a wealth of colour is hid-
den, which flares out, showing a band of scarlet close to
the head, next a streak of bright yellow, then a second
band of red, and finally the white tips of the crest feathers. The nod or jerk of the head in spreading wide the crest
reminds one of the sudden flick with which a fan is thrown
open. open. HEADS AND NECKS Concealed crests bring to mind the Kingbird and the
Ruby-crowned Kinglet, both of which derive their names
from their crowns of ruby. It is said that the former
bird is aided in its search for food by the bright spot
of colour which, flower-like when exposed, attracts in-
sects. This, however, should be confirmed before being
accepted as a fact; although in a tropical flycatcher,
which has a beautiful red and purple transverse crest, the
evidence of this novel use seems fairly well corroborated. FIG. 202.— Crest of Banded Curassow (female). FIG. 202.— Crest of Banded Curassow (female). l" :>-"
e
II
FIG. 203. — Crest of Victoria Crowned Pigeon. 261 FIG. 203. — Crest of Victoria Crowned Pigeon. 261 FIG. 203. — Crest of Victoria Crowned Pigeon. 261 FIG. 204. — Harpy Eagle. (Courtesy of Dr. Frank Baker.)
262 FIG. 204. — Harpy Eagle. (Courtesy of Dr. Frank Baker.)
26 FIG. 204. — Harpy Eagle. (Courtesy of Dr. Frank Baker.) Heads and Necks Heads and Necks The Laughing Thrush of the Himalaya Mountains
has every feather upon its head lengthened and perma-
nently erect, forming a soft, spreading halo. IG. 205. — Hooded Merganser. (From a photograph provided by the America
Museum of Natural History.) FIG. 205. — Hooded Merganser. (From a photograph provided by the American
Museum of Natural History.) In almost every Family of birds we find certain spe-
cies with long, well-developed crests. Among the ducks,
the Hooded Merganser has a compressed, semicircular
halo of delicate feathers, while the Mandarin Duck has
a broad, many-coloured, erectile crown, which is con- The Bird 2 spicuous even in contrast with the gorgeous ornamenta-
tion of the body of this feathered harlequin. Of birds
of prey, the Harpy Eagle has perhaps the most imposing
crown of feathers. Sometimes the crest is sharply set off from the rest
of the bird's plumage, as in the scarlet-plumed wood-
peckers, whose crests give them the appearance of having
long hair, which is gracefully brushed straight backward
and upward. and upward. We must not discuss the subject of crests without men-
tion of two birds of extraordinary appearance, the Crowned
Crane and the Umbrella-bird. The former illustrates
admirably what strange and unfeather-like forms, feathers
may assume in the course of evolution. The illustration
shows better than words can describe the dense, velvety
cap of plush-like feathers, and the glorious crown of a
myriad radiating points — a decoration unrivalled, even
among birds, in exquisite colour and delicacy. In addi-
tion to this, the cheeks are entirely bare of feathers, and
the lower half suffused with blood, which shows through
the skin, — an ever-changing blush of deep pink. the skin, — an ever-changing blush of deep pink. The decorations of the Umbrella-bird are as beautiful
as they are bizarre; while if shorn of its crest and streamers,
this bird would resemble a small crow in appearance. The crest really bears a resemblance to the article which
has given the bird its name— a high, arching mass of
feathers, overshadowing the entire head and beak, con-
tinually spreading and partly closing again, as the bird's
emotions change. From the neck of the bird dangles a
streamer of black feathers, as long as the bird's entire Heads and Necks Heads and Necks
FIG. 206 —Crowned Crane. FIG 207. — Demoiselle Crane. FIG. 206 —Crowned Crane. FIG. 206 —Crowned Crane. FIG. Heads and Necks 206 —Crowned Crane. FIG. 206 —Crowned Crane. FIG 207. — Demoiselle Crane. FIG 207. — Demoiselle Crane. The Bird 266 body and which, when it flies, blows back between its
feet. The filament of feathers looks for all the world
as if a strip of the bird's plumage had caught on a thorn
and torn loose. The core of the streamer is a very slender FIG. 208.— Umbrella-bird. FIG. 208.— Umbrella-bird. FIG. 208.— Umbrella-bird. ribbon of skin which hangs from the neck. Would that
we could state the causes and the manner of the devel-
opment of these curious structures which our fancy likens
to an umbrella and a feathery handle! ribbon of skin which hangs from the neck. Would that
we could state the causes and the manner of the devel-
opment of these curious structures which our fancy likens
to an umbrella and a feathery handle! One or two small tufts of feathers may spring from Heads and Nec 267 some part of the head of a bird, such as the feather " horns "
of owls, motmots, and larks. In Screech Owls these
prominent "ears" certainly play a useful part in breaking
up the outline of the bird, rendering it very difficult of
detection when it is perched upon some jagged limb or
stub. Or again, tufts or pencils of feathers may arise
from near the ear, or over the eye; as shown by the Dem-
oiselle Crane (Fig. 207), some of the Puffins, and the Man-
churian Pheasants (Fig. 209). The Great Bustard has long
tufts of chin-feathers which, like wide-spreading whiskers,
spread to each side, and the Bearded Vulture has a simi-
lar goatee of stiff, black bristles. lar goatee of stiff, black bristles. Of the wonderful crests, frills, ruffs, breastplates, and
cloaks of hummingbirds there is no room to speak, and
indeed no words or pictures can aught but parody them. The eye alone can record their marvels, in the collection
of a museum, or, better still, in the living birds, as the
little creatures hover over, their favourite flowers, or
vibrate before us, fanning the air in our very faces with
their invisible wings. Brief mention should be made of two Birds of Para-
dise, those beautiful creatures inhabiting a region where
the eye of man seldom sees them. The Six-shafted Bird of Paradise is found only in
New Guinea. Heads and Necks "The plumage appears at first sight black,
but it glows in certain lights with bronze and deep pur-
ple. The throat and breast are scaled with broad, flat
feathers of an intense golden hue, changing to green and
blue tints and certain lights. On the back of the head
is a broad recurved band of feathers, whose brilliancv FIG. 209.— Head of Eared Pheasant. FIG. 209.— Head of Eared Pheasant. FIG. 210.— Head of Great Horned Owl. 268 FIG. 210.— Head of Great Horned Owl. 268 Heads and Necks is indescribable, resembling the sheen of emeralds and to-
paz, rather than any organic substance. Over the fore,
head is a patch of pure white feathers, which shine like
satin; and from the sides of the head spring the six won-
derful feathers." IG. 211. — Six-shafted Bird of Paradise (a mounted bird). (From a phot
graph provided by the American Museum of Natural History.) FIG. 211. — Six-shafted Bird of Paradise (a mounted bird). (From a photo-
graph provided by the American Museum of Natural History.) Head decorations reach the acme of strangeness in
the King of Saxony's Bird of Paradise. The bird itself
is sombre-hued and small, about the size of our robin,
with nothing unusual about its appearance, except for
the two streamers springing from opposite sides of the The Bird 270 head. They are twice (or more) the length of the body,
and, far from being feather-like, they are best described
as a series of thirty or forty tiny flags of blue enamel,
each separate, each hanging pendent from the main
shaft (Fig. 212). It would seem as if Nature herself
could go no farther in unusual decoration than this. FIG. 212. — King of Saxony Bird of Paradise. (From a photograph provided
by the American Museum of Natural History.) FIG. 212. — King of Saxony Bird of Paradise. (From a photograph provided
by the American Museum of Natural History.) In the Double-crested Pigeon of Australia the core
or fleshy covering of the beak is completely feathered;
while some of the birds known as plantain-eaters are
feathered to the very tip of the short beak with plumes
of delicate green, tipped with white. Heads and Necks The extreme of
feathering is shown by the Cock-of-the-Rock, in which Heads and Necks 271 the whole beak, in fact every part of the head except the
eyes, is buried in a maze of soft, orange plumes. the whole beak, in fact every part of the head except the
eyes, is buried in a maze of soft, orange plumes. eyes, is buried in a maze of soft, orange plumes. As the antithesis to this condition, we find many
birds which have the head partly or entirely bare of
feathers, such as the vultures and some of the waders. In the former group this lack of feathers is doubtless FIG. 213. — Head of male Condor FIG. 213. — Head of male Condor of value in enabling the birds to avoid soiling their plu-
mage, when engaged in their scavenger work. The great
Condor of South America has, just below this naked area,
a necklace of the whitest of fluffy down, and in addition
the male has a large wattle of skin upon the front of the
head. The Caracara of Mexico is partly vulturine in its
habits, and the feathers have disappeared from part of
its face. Wherever the skin of the head and neck is even of value in enabling the birds to avoid soiling their plu-
mage, when engaged in their scavenger work. The great
Condor of South America has, just below this naked area,
a necklace of the whitest of fluffy down, and in addition
the male has a large wattle of skin upon the front of the
head. The Caracara of Mexico is partly vulturine in its
habits, and the feathers have disappeared from part of
its face. Wherever the skin of the head and neck is even The Bird 272 FIG. 214. — Caracara. partly vulturine in habits. FIG. 214. — Caracara. partly vulturine in habits. FIG. 215.— Young King Vulture. FIG. 215.— Young King Vulture. Heads and Nec 273 partly bare, ornamentation often takes the form of many-
shaped and often highly coloured wattles, such as we see
highly developed in a King Vulture. The most common example of this is seen in a domestic
rooster or a turkey, but in many other birds these wat-
tles of skin are very brilliant in hue. Indeed the casso-
waries are resplendent in their gorgeous hues of blue, yel-
low, red, and many other intermediate shades. Heads and Necks Turkeys
too, at the breeding season, develop bright colours. The Yucatan Wild Turkey, which has thus far resisted
all attempts at domestication, has the bright blue naked
skin of the head, dotted with tubercles of the most bril-
liant orange, while a long tube-like wattle, also tipped
with orange, dangles down over the beak. The wattles,
or caruncles, of the Bell Bird are interesting as being con-
nected with the windpipe in such a way that they become
inflated with air when the bird utters its wonderful note. In the White Ibis the face only is bare, in the spoon-
bill the head and face, and the whole head and neck in
the Marabou Stork; the effect of this condition in the
latter bird being heightened by the enormous pouch
which hangs suspended from the neck. The same is true
of the Adjutant. A close inspection of the neck of one of these storks
will show that, while ordinary feathers are absent, there
is a scanty covering, here and there, of what looks like soft,
curling ringlets of chestnut hair. The resemblance is
absolutely perfect, and no naturalist in the world, if shown
one of these locks, would say that it came from a bird
and not from one of the hair-covered mammals ! FIG. 216. — Head of domestic cock. Extreme development of comb. FIG. 216. — Head of domestic cock. Extreme development of comb. FIG. 217. — Head of domestic cock. Extreme development of crest. 27 FIG. 217. — Head of domestic cock. Extreme development of crest. Heads and Necks Heads and Necks We must pass by all the strange ornaments of horn
upon the heads of birds, such as the scarlet plate of the
gallinules, the immense recurved casques of the hornbills,
the use of which is as yet unknown, unless it be purety
ornamental. But the impressive helmets of the cassowaries, FIG. 218.— Head of Wild Turkey. FIG. 218.— Head of Wild Turkey. Fig. 228, demand especial notice in this volume, as being very
useful adaptations to life in a dense forest. These great
running birds are the only members of their Sub-class
which inhabit thickly forested regions, and in speeding
with great leaps and bounds through the undergrowth, the
tall, smooth helmet of horn protects the head of the bird The Bird 2/6 FIG. 219.— Head of Adjutant. FIG. 219.— Head of Adjutant. FIG. 219.— Head of Adjutant. FIG. 220. — Ringlet of hair from the neck of an Adjutant. FIG. 220. — Ringlet of hair from the neck of an Adjutant. Heads and Nec 277 and shunts off the hanging lianas and vines which would
otherwise impede its progress. In our hasty paragraphs we have seen to what a re-
markable extent the ornamentation of the heads of birds
is carried, and as many extraordinary examples could
be given of decoration of the neck. The Loon has a
speckled black and white throat with a long colour band-
age of black feathers wound about its neck; the cervical
ruffs of our Ruffed Grouse are like wings in miniature;
in the Golden and Amherst Pheasants this form of orna-
ment is extended into a circular ruff of black and gold
and black and white respectively; while in the Superb
Bird of Paradise a shoulder-cape flares back, large enough
to cover almost the entire body of the bird, giving it the
appearance of being clothed in two distinct sets of plu-
mage! The nuptial attire of the Ruff, a species of sand-
piper, is as greatly developed, except that it forms a
double cloak over the breast. This cloak or shield of feathers in the Ruff plays a
vital pa*rt in the life of the bird. We must first notice
the remarkable variation in the pattern of this cloak of
battle, — for such it really is. Heads and Necks If we could see fifty Ruffs
standing side by side, some would be seen to have ruffs
of pure white, others of gray, black, orange, buff, or chest-
nut, while the waving ear-plumes are also independent
in colour, varying from white to purple, green, or blue. Then there is a type of Ruff with barred cloaks, another
with spotted patterns, and so on in almost endless
variation. This condition of affairs is wholly unlike
the uniform pattern of colouring of other wild birds. The Bird 278 FIG 221. — Lady Amherst Pheasant. FIG 221. — Lady Amherst Pheasant. FIG. 222.— Casque of Hornbill. FIG. 222.— Casque of Hornbill. Heads and Necks We can only compare these little Joseph-coated birds
with the unnatural sports among domestic poultry and
pigeons. pigeons. But whatever their colour, these Fighting Snipe find
their ruffs of service in their encounters at the breeding
season. Four male birds which I observed in captivity FIG. 223.— Ruff with battle-cloak partly moulted. FIG. 223.— Ruff with battle-cloak partly moulted. were adorned respectively, — plain gray, dotted gray,
chestnut barred with black, and a rich golden rufous. Though no females were present, yet their fighting instinct
often cropped out and a pair of them would dart and side-
step about each other, bills held low and far advanced,
ruff spread out from the breast and trailing low, hiding
almost the whole body. Now and then one of the fencers
would make a vicious dash, sending his bill through the were adorned respectively, — plain gray, dotted gray,
chestnut barred with black, and a rich golden rufous. Though no females were present, yet their fighting instinct
often cropped out and a pair of them would dart and side-
step about each other, bills held low and far advanced,
ruff spread out from the breast and trailing low, hiding
almost the whole body. Now and then one of the fencers
would make a vicious dash, sending his bill through the The Bird 280 feather shield of his opponent. But the force of the
blow would spend itself on the inch of space between the
shield and the feathers of the bird's breast. When, in
his native haunts, the Ruff has conquered his rival, his
triumphant dances before the female are most elaborate. While these facts are not exactly pertinent to the physical FIG. 224. — Breast ornament of a Wild Turkeycock. FIG. Heads and Necks 224. — Breast ornament of a Wild Turkeycock. life of the bird, yet I mention them to show to what prac-
tical, as well as aesthetic, uses the development of some
portion of the bird's plumage may be devoted. portion of the bird's plumage may be devoted. What a contrast to the cloak of the Ruff is the pectoral
decoration of the Wild Turkeycock: a great tuft of
coarse, black hair-like feathers, like the tail of a horse Heads and Necks 281 in miniature, growing almost a foot in length from the
centre of the breast ! The length of the neck of birds is often correlated
with that of the legs, — a long-legged bird of necessity re-
quiring a long neck to permit its bill to reach the ground. Geese and swans are an exception, and in their case we FIG. 225. — Flamingoes Correlation of long neck with long legs. FIG. 225. — Flamingoes Correlation of long neck with long legs. find that the long, mobile neck is of great use in making
up for the awkwardness of their waddle when on land,
and in allowing them to reach beneath them while floating
in shallow water, thus feeding along the bottom. Herons are uniformly so light of body that they would
have difficulty in steadying themselves in the air, were
it not that, when in flight, their necks become compressed
to an incredible thinness, thus acting as does the cut- The Bird 282 water of a ship's prow. The perpetual crook in the necks
of these birds is significant of their method of fishing — a
patient watch until the prey comes within striking distance-
In the snake-bird this crook, or Z-shape, has, by the it
FIG. 226. — Swan. Correlation of long neck and short legs due to feeding habits. (Sanborn, photographer.) FIG. 226. — Swan. Correlation of long neck and short legs due to feeding habits. (Sanborn, photographer.) adaptation of three of the neck-bones, become a veritable
trigger, by the springing of which the bird literally spears
the fish. Heads and Necks If the mention in this chapter of a few examples of
crests and other decorations has seemed in the least to Heads and Necks verge upon the monotony of a mere catalogue, my plea
is that they have been cited with the intention of empha-
sizing the fact of the remarkable degree which decoration,
pure and simple, plays in courtship. Viewed from such
a standpoint, these facts and comparisons become im-
portant data in the observation of the courtship of birds, FIG. 227. — Snake-bird, showing crook in neck. FIG. 227. — Snake-bird, showing crook in neck. which in its turn is one of the most important and interest-
ing corollaries of the psychology of these beings. Whether
female birds have highly developed aesthetic feelings, or
whether the songs and dances and colour masses act more
along the line of the passes of a hypnotist, is yet to be
ascertained. It is also hoped that a realization of the more immedi- The Bird 2 ately practical uses of such structures as the cassowary's
horny helmet, the feather shield of the Ruff, perhaps the
crest of the kingbird, and many others as yet unknown,
will impel amateur observers to further efforts in the
investigation of the life-habits of birds. THE BODY OF A BIRD N
, N experimenting with balloons and flying-ma-
chines, weight is a question of prime import-
, ance, and among birds there seem to be certain
limits to the bulk of the body, beyond which flight is
impossible. The tiny hummingbirds, with bodies weigh-
ing less than some insects, have remarkable powers of
flight, and throughout all the groups of larger birds we
find certain species with exceptional flight ability, until
in the birds of widest extent of wing, such as the condor
and the albatross, flight seems to reach the acme of perfec-
tion. But the flying birds of actual heaviest bulk are
perhaps the Wild Turkey, the Great Bustard, and the
Trumpeter Swan, the two latter reaching weights of thirty-
two and twenty-five pounds respectively. Even the
gigantic Pterodactyls, those flying reptiles of olden time,
some of which had heads a yard long, and an expanse of
eighteen feet or more of bat-like wings, are estimated
to have weighed but twenty pounds or thereabouts. 285 But when the necessity for flight ceases, a bird may
begin to assume larger . proportions and greater weight
without detriment; just as a mammal which adopts a life
in the dense medium of water may attain a much more
285 The Bird 2 gigantic size than one which has to support its body in
the thinner atmosphere: a whale is to a horse as an
ostrich is to a dove. The ostrich is the largest of all living birds, a full-
grown male being able to reach to a height of nine feet
and weighing as much as three hundred pounds; but
even these figures were exceeded by its extinct relative
of Madagascar, the moa, whose height is variously esti-
mated at from ten to eleven feet, and whose massive leg-
bones show that its weight must have been much greater
than that of the ostrich. There is a great difference in the relative condition
of the body in various birds. Herons, even when fish
are abundant, with opportunities of feeding from morn-
ing to night, are thin to emaciation. Truly they belong
to the "lean kine." A fat heron would be an anomaly. On the other hand, the flesh of many sea-birds seems as
constantly encased in thick, oily layers of fat. THE BODY OF A BIRD Petrels
are used by the inhabitants of some islands as candles,
simply by threading the body of the dead bird with a
wick, the excess of fat burning steadily until the whole is
consumed. Penguins are well protected against the icy
waters of their Antarctic home by a layer of fat under
the skin, so thick in proportion to their size as to remind
one of the blubber of whales. If we were writing of the bodies of the fur-bearers
instead of birds, we would have much to say concerning
the various kinds of scent-glands and secreted odours;
but in birds the only gland is that above the tail, which
furnishes the oil with which the bird preens its plumage, The Body of a Bi 287 thus both cleansing it and rendering it water-proof. That
birds, and especially those which, like quail, are found
in flocks, possess odours is borne witness to by the ability
of dogs to point successfully the hidden game; but that
this is of much use in enabling the birds to find one another
is doubtful, both from the fact of the slight development
of the sense of smell, and because of the loud call-notes
which are so characteristic of these birds. One exception,
however, may be noted, that of the apteryx, which is
said to have a strong and persistent odour, with corre-
spondingly well-developed nostrils. spondingly well-developed nostrils. Again, among fur-covered animals we find usually
a poor development of the sense of sight and but few
of them exhibit bright colours, while, as we have seen,
birds excel in the power of seeing, and, correlated with
this, possess an unparalleled array of colours upon the
body. There are many ways in which the body or its feathers
are adapted to aid the bird in some special way. For
example, the Puff-back Shrike of Africa has a habit of
suddenly puffing out and erecting a patch of long, loose,
white feathers on its back, giving the appearance of a
large powder-puff, an act so startling and unexpected
being well calculated to make any attacking hawk or
other bird hesitate. The general texture of the body feathers is usually
an accurate index to the bird's power of flight. THE BODY OF A BIRD Although
the feathers of the breast and back are never as compact
or as stiff as those of the wings and tail, yet in birds of
good flight their barbs are quite firmly connected. In a 288 The Bird small African bird, called from its habits the Rock-jumper,
the wings are so small that the power of flight is almost
nil, and we find an interesting corollary in the plumage, FIG. 228. — Cassowary, showing the loose plumage of a flightless bird. (Sanborn,
photographer. Courtesy of N. Y. Zoological Society.) FIG. 228. — Cassowary, showing the loose plumage of a flightless bird. (Sanborn,
photographer. Courtesy of N. Y. Zoological Society.) which is so loose and fluffy that it blows about in the
least wind. In the ostrich and rhea this down-like char-
acter is still more noticeable and extends even to the
feathers of the wings and tail. The extreme is to be found which is so loose and fluffy that it blows about in the
least wind. In the ostrich and rhea this down-like char-
acter is still more noticeable and extends even to the
feathers of the wings and tail. The extreme is to be found The Body of a Bi 289 in the apteryx and emeu (Fig. 23). Compare a feather
of the latter with one of a condor and the difference is
remarkable. So unfeatherlike is the emeu's plume and
so loose are its barbs that it brings to mind the much-
divided leaflets of an Acacia. The plumage of the snake-bird is inexplicable. This
bird is so emphatically aquatic that we would expect
a dense, compact covering of the body; but in reality
it more nearly resembles hair or fur, soaking through so
quickly and thoroughly that, after immersion for some
time, the bird becomes waterlogged and has to hang
itself out to dry by seeking some sunlit perch, opening
wide its wings and waving them to and fro. The feathers of the penguin are small, flat, and rigid,
approaching in these respects the scales of fishes — an
interesting reacquinng of characters consequent upon
an all but wholly aquatic life. It is interesting to com-
pare the colouring of such a bird as the Scaled Partridge
with a fish like the Carp, the dark margins of the feathers
and scales bringing about a remarkable resemblance. THE BODY OF A BIRD Taking up the subject of colour in general, we realize,
after even a superficial glance at a collection of birds,
that in gorgeousness of hue and diversity of shade and pat-
tern, they are to be compared only with insects. In a
former chapter we have briefly considered the chemical
and optical causes of colour in feathers; but the causes
due to environment (using that word in its widest sense)
cover a vastly greater field and one as yet comparatively
unexplored. Advancement of actual knowledge of any subject in The Bird 290 science depends upon two things: first, the accumula-
tion of facts; and secondly, a philosophical spirit capable
of generalizing and bringing order out of the chaos of of generalizing and bringing order out of the chaos of
FIG. 229.— Breast of Scaled Partridge. FIG. 230. — Carp, a fish with distinctly marked scales. (Keller, photographei.)
these myriad observations. A knowledge of museum
facts is of but slight use in such a subject as the one under
consideration, which requires more adequate knowledge FIG. 229.— Breast of Scaled Partridge. FIG. 229.— Breast of Scaled Partridge. FIG. 229.— Breast of Scaled Partridge. FIG. 230. — Carp, a fish with distinctly marked scales. (Keller, photographei.) FIG. 230. — Carp, a fish with distinctly marked scales. (Keller, photographei.) these myriad observations. A knowledge of museum
facts is of but slight use in such a subject as the one under
consideration, which requires more adequate knowledge these myriad observations. A knowledge of museum
facts is of but slight use in such a subject as the one under
consideration, which requires more adequate knowledge The Body of a Bird 291 than we now possess of the life-habits and the psychology
of birds. This is especially true of the great number of
cases which we can explain only by calling them orna-
mental and decorative. Hence we find one ornithologist FIG. 231. — White-throated Sparrows. The light-coloured bird is in normal plu-
mage; the dark bird was subjected to moisture-laden air through two moults. FIG. 231. — White-throated Sparrows. The light-coloured bird is in normal plu-
mage; the dark bird was subjected to moisture-laden air through two moults. explaining a certain colour as due to one cause, while
another scientist gives an entirely different interpretation
of the same fact. THE BODY OF A BIRD From personal observation among the birds of the The Bird 292 New York Zoological Park, I have had opportunity to
record many cases of the effect of food upon colour. An
experiment very commonly known is that of feeding
canaries on red pepper, thus causing their plumage, after FIG. 232. — Variation due to climate, etc., in races of North American Song Sparrows
(From a photograph provided by the American Museum of Natural History.) FIG. 232. — Variation due to climate, etc., in races of North American Song Sparrows
(From a photograph provided by the American Museum of Natural History.) successive moults, to become of an intense orange colour. This is the more remarkable since the actual red pigment,
or capsicin, of red pepper is not the direct cause of the
canaries' changed hue, but a fatty substance known as
triolein, which is a constituent of the pepper. The Body of a Bird The Body of a Bird It is generally thought that the fact that, in captivity,
Purple Finches and orioles frequently moult into yellow-
ish hues, instead of their rightful tints, is due to some
change in food. Indeed in many species the bright colours
are wholly lacking after a year or two in captivity. But
I have transferred a male Purple Finch, which had for
several years moulted yellow, from a dark cage to one
which was exposed to
bright sunlight, and in
one moult the bird as-
sumed his original and
normal colour. FIG. 233.— Effect of environment on Bob-
white, shown by specimens from Min-
nesota, Florida, and Cuba. (From a
photograph provided by the American
Museum of Natural History.) A more probable ex-
ample of the effect of
food upon colour is seen
in our American Flamin-
goes. In captivity these
birds fade out moult by
moult, until they become
almost white, like the
European species. By
mixing with their food
a quantity of some
strong but harmless dye,
I have had them either retain their original colour for
years, or at least the fading process has been appreciably
lessened. FIG. 233.— Effect of environment on Bob-
white, shown by specimens from Min-
nesota, Florida, and Cuba. (From a
photograph provided by the American
Museum of Natural History.) FIG. 233.— Effect of environment on Bob-
white, shown by specimens from Min-
nesota, Florida, and Cuba. (From a
photograph provided by the American
Museum of Natural History.) FIG. 233.— Effect of environment on Bob-
white, shown by specimens from Min-
nesota, Florida, and Cuba. (From a
photograph provided by the American
Museum of Natural History.) The effect of climate upon colour is even more readily
proved, and may be noticed in wild birds as well as in
those in captivity. In regions which have a very dry The Bird 294 climate, the birds, and in fact all of the animals, are of
a much lighter hue than those living in an atmosphere
of great humidity, where moisture does not readily evapo- FIG. 234. — Male Scarlet Tanagers, showing moult from the scarlet summer dress,
(a), through the parti— coloured garb (6), into the green winter plumage (c). FIG. 234. — Male Scarlet Tanagers, showing moult from the scarlet summer dress,
(a), through the parti— coloured garb (6), into the green winter plumage (c). rate. The Body of a Bird In such a place birds tend to be very dark-coloured. In the case of captive birds, I have seen White-throated
Sparrows and Wood Thrushes become almost like black- rate. In such a place birds tend to be very dark-coloured. In the case of captive birds, I have seen White-throated
Sparrows and Wood Thrushes become almost like black- The Body of a Bird birds in colour when confined in a bird-house where the air
was constantly moist. Correlated with the effect upon
colour is often a difference in size, and in many instances
among birds the more northerly individuals are larger,
those inhabiting warmer regions being less in stature. Among wild birds, the Quail, or Bob-white, shows an
almost unbroken series from the northern, light-coloured
variety, ten inches in length, to the Cuban bird, very FIG. 235. — Siberian Black Lark, male bird in the spring. FIG. 235. — Siberian Black Lark, male bird in the spring. much darker in shade and measuring only eight inches
from beak to tip of tail. The race of Bob-whites seems
very susceptible to climatic influence; as in Mexico there
are nearly a dozen different geographical races, each in-
habiting a distinct portion of the country. Many other
wide-spread groups of birds, such as the Song Sparrows,
vary in a similar manner. It is strange what a marked
effect this greater or less amount of moisture has upon
birds, even in very limited districts. A South Ameri-
can pipit, the individuals of which spend their lives on
very circumscribed plots of earth, exhibits two colour much darker in shade and measuring only eight inches
from beak to tip of tail. The race of Bob-whites seems
very susceptible to climatic influence; as in Mexico there
are nearly a dozen different geographical races, each in-
habiting a distinct portion of the country. Many other
wide-spread groups of birds, such as the Song Sparrows,
vary in a similar manner. It is strange what a marked
effect this greater or less amount of moisture has upon
birds, even in very limited districts. A South Ameri-
can pipit, the individuals of which spend their lives on
very circumscribed plots of earth, exhibits two colour The Bird 2 forms entirely different, and thought to be due solely
to the amount of moisture in the ground on which it lives. The Body of a Bird Very dark-coloured and very pale individuals live within
a few hundred yards of each other, in dry and swampy
situations respectively, each, it is said, keeping entirely
to its own little beat. We are all fairiliar with the changes of colour due to FIG. 236. — Nighthawk perching lengthwise on a fallen branch. FIG. 236. — Nighthawk perching lengthwise on a fallen branch. age, as, for instance, in the young Rose-breasted Gros-
beaks, which are very different from the male parent, and
the young Bald Eagles, which lack the white colour of the
feathers of head and tail. Certain wild pigeons show
marked differences in colour patterns between the young
birds and the adults, and very good evidence of the gradual
evolution which must have preceded these changes is The Body of a Bi 297 to be found by plucking out a few of the feathers of the
young bird. Those which replace the ones pulled out
will show intermediate stages, which have long since been
dropped from the sequence of patterns, as observed in
the regular moults of the birds. Another important phenomenon is the seasonal moult,
which was spoken of in the chapter treating of feathers-
In the fall of the year the brilliant Scarlet Tanager assumes
the olive-green dress of the female, and the Indigo Bunt-
ing and the Bobolink likewise don the dull garb of their
mates. There is another very interesting cause of change in
colour, namely, the wearing off of the brittle tips of the
feather-vane. An excellent example of this is seen in the
Snowflakes, which come south in the depth of severe
winters, flying in small flocks about our fields, like an
animated flurry of the actual crystals. When we see
the birds at this time they are brownish and brownish
white. In the spring, in their northern home, they change
to a clear-cut black and white, not by shedding the entire
plumage, but merely by the breaking off of the brown
feather-tips. By a similar process the Bobolink changes
from the buffy female dress to his rich black-and-white
spring suit, and, as we saw in Chapter II, Fig. 35, the
English Sparrow gains his cravat of jet. English Sparrow gains his cravat of jet. The Body of a Bird Another excellent example is found in the Black Larks
of Siberia, the males of which, in winter, are of an almost
uniform sandy colour, like a Skylark, but by the wearing
off of the buff tips of the feathers, the birds become jet-black
in the summer — a most remarkable and radical change. The Body of a Bi The Body of a Bi 299 The relation of a bird's colours to its haunts and its
habits of life is a subject of intense interest. This is,
of course, not in the same category as the subjects of the
foregoing paragraphs, but indeed includes them all. The
most common class of colours is known as protective. These are such that the bird resembles its environment
or surroundings and is thus given a better chance of escap-
ing the observation of its enemies. It is evident that,
in a study of this nature, observation of the bird in its
natural haunts is of far greater value than any other
method. We find that the majority of sparrows, sandpipers,
and quail are gray or brown, like the grasses, sedges, and
leaves among which they live; while the birds which
spend their lives higher up among the branches of trees
are greenish, or at least more brightly coloured. are greenish, or at least more brightly coloured. Many birds which are protectively coloured are dark
above and white or whitish beneath. The significance
of this pattern of coloration has been beautifully demon-
strated by an American artist, Mr. Abbott Thayer. His
experiment, which is as follows, may be repeated by any
one: Take two wooden decoy ducks, and place them
against a sand-bank. Colour one the exact tint of the
sand, or even coat it with that substance. Repeat this
with the upper parts of the second decoy, making its
back darker than the surrounding sand, but grade the
under part of this one to pure white below. At a little
distance away, decoy number one will still be distinctly
seen; while number two will absolutely disappear, merg-
ing perfectly into its background. The reason for this is The Bird 00 300
that the conspicuous white of the under surface of the
second bird is, when normally lighted up by the sun,
neutralized by the shadow of the bird, and the darker
upper parts are softened and toned down by the strong
direct light; while if the entire bird be unshaded, although FIG. 238.— Sooty Tern on her nest. FIG. 238.— Sooty Tern on her nest. coloured like the environment, the dark shadow beneath
will reveal it clearly. The Body of a Bi Whether or not birds really appreciate the value of
the protective colour of their plumage, it is certain that
a quail or ptarmigan will remain crouching on a brown
bit of turf, until all hope of evading danger is gone; while
birds which are very evidently not protectively coloured
are invariably more wary and difficult of approach. When FIG. 239. — Seven young Flickers clinging to a tree. (R. H. Beebe, photographer.) FIG. 239. — Seven young Flickers clinging to a tree. (R. H. Beebe, photographer.) The Bird 302
a ptarmigan, while yet in the brown garb of summer,
is exposed against a hillside of snow, it becomes very
wary. It is interesting, in this connection, to observe how a
Nighthawk carries out its colour resemblance to a knot
or a rough piece of bark, by perchiyg, not crosswise, but
lengthwise, along a branch or fallen tree-trunk. lengthwise, along a branch or fallen tree-trunk. A volume might easily be written of the various ways
in which protective coloration works out among birds,
but there is so great a difference of opinion, and indeed
so many exceptions to every theory which may be ad-
vanced, that it is better, for the most part, to go to Nature
without a priori theories, and putting ourselves as nearly
as possible in the position of the creatures themselves,
to hope for better ability to see with their eyes. And
it is right along this line that we most need fresh data
and experiments, namely, the actual ability of birds
and insects to distinguish shades, colours, forms, and
motion, — whether efficient in certain ways or not. We
know that many men cannot distinguish a scarlet ball
lying upon green grass; that is, they are partly colour-
blind. If this were the case with certain hawks, a male
Scarlet Tanager would be forever safe from them among
the green foliage. An important fact, which for years had been appar-
ent to me, but unexpressed until Mr. Abbott Thayer put it
into words, is that colours which we would ordinarily term
conspicuous are often exactly the opposite when found in the
plumage of a bird. The Body of a Bi Writing of the Motmot in my volume
"Two Bird-lovers in Mexico," I say: "I have often The Body of a Bird 3°3 wondered, when I saw mounted specimens in museums,
with what special immunity from danger these birds
were blessed, their beautiful colouring would seem to be
such a startling advertisement of the bird's whereabouts. But in reality the very diversity in hue is their protec-
tion, and they merge per-
fectly into their environ-
ment of green foliage and
bright sunlight." FIG. 240. — Brown Creeper circling up the
trunk of a spruce. bright sunlight."
Indeed absolute uniform-
ity of coloration instantly
reveals the outline of the
bird entire, and renders it
very conspicuous. Birds
which have but few ene-
mies are often thus mono-
crome in hue. But look
at the photographs and
see how a broken colora-
tion baffles the eye. If
the Sooty Tern, Fig. 238, were totally black, it would
be conspicuous even against a patch of dark-coloured
mottled shingle. But the transverse lines of white across
the back totally destroy the symmetry of form, while
the white wing-edges fairly force the eye to call them,
not part of a bird sheltering her eggs, but only two among
a myriad irregular edges of coral rock! FIG. 240. — Brown Creeper circling up the
trunk of a spruce. FIG. 240. — Brown Creeper circling up the
trunk of a spruce. Observe closely the seven young Flickers clinging to
their natal stump. As the warriors of Jason sprang forth
from the ground fully armed, so the very bark, mottled The Bird 3°4 with spots of lichen and sunlight, seems to have gendered
these baby birds. Yet they were hatched in a dark hole
from the whitest of white eggs. Is this and a thousand
of other resemblances to be termed accidental? Then
is all Nature one great accident! When the Flicker flies
with swift wing-beat from tree to tree, then the white FIG. 241. — Laughing Gull on nest. FIG. 241. — Laughing Gull on nest. rump blazes forth. At such moment no protection is
needed; but in these young Flickers upon the tree-trunk,
how exquisitely do their spots deceive the eye! The Body of a Bi They
are, we say, perhaps sunlight splashes, — nothing more-
Yet others which, like the Brown Creeper, haunt the
tree-trunks of the forest, seem veritably to be but stray
bits of roughened bark creeping here and there. tt tt tt The Bird 3 Let us glance at one more bird upon her nest, — a Laugh-
ing Gull. At a distance we see a shapeless blotch of white
sand among the reeds, that is all. We walk over a hundred
other similar patches; but when near enough, we at
last are able to distinguish the dark head and wing-tips,
all but invisible among the shadows, and even through
the centre of the head we can see two spots of light be-
yond,— or no, it is the little subtle ring of white about
the eye! Two majestic Black-necked Swans may swim closely
along in full view near the opposite bank of a pond, and
yet be totally unrecognizable; showing to the eye as
bodiless necks or neckless bodies, according to the chang-
ing conditions of light and shade around them. ing conditions of light and shade around them. We see a troop of ostriches rushing past. Surely
nothing could hide birds such as these! Again we see
one of these birds prone upon the ground, and a mighty
creature towering eight feet or more above the earth,
becomes naught but a dark ant-hill, which the photo-
graph picks out clearly, but which in the desert, dotted
with ant-hills, would seldom be noticed even by the hungri-
est of lions. Of course, like most other theories, this of protective
coloration can easily be carried too far, but there are
hundreds of instances where it seems to answer every
requirement of the case. Few fields offer such opportuni-
ties for original work of the most delightful character. As one example out of untold numbers, what explanation
can we give of the Blood-breasted Pigeon or Bleeding-
heart Pigeon, which, as its name denotes, has a splash The Body of a Bird 3°7 FIG. 243. — Group of Ostriches on the run. (Cawston, photographer.) FIG. 243. — Group of Ostriches on the run. (Cawston, photographer.) FIG. 244. — Ostrich as it hides from an enemy. FIG. 244. — Ostrich as it hides from an enemy. The Bird 8 3o8
of blood-like scarlet in the centre of its breast? The Body of a Bi The re-
markable and inexplicable resemblance is heightened
by the stiffened vanes of the centre feathers, causing them
to appear bedraggled and clotted, as if by an actual wound ! The photograph does but little justice to the bird's real
appearance. appearance. Another class of colours, while still protective, is so
for a purpose very different from those cases which we FIG. 245. — Bleeding-heart Pigeon. FIG. 245. — Bleeding-heart Pigeon. have been considering. The colours which we are now
to mention have been aptly called aggressive colours, as,
by their means, a bird of prey is enabled to approach
its victim more easily. So, throughout the entire animal
world we find two phases of phenomena constantly pres-
ent: on the one hand the pursued ones, striving to escape
by all means in their power; and on the other hand the
pursuers, ever trying to outwit those upon which they
prey. If a duck acquires great speed of flight, the Duck have been considering. The colours which we are now
to mention have been aptly called aggressive colours, as,
by their means, a bird of prey is enabled to approach
its victim more easily. So, throughout the entire animal
world we find two phases of phenomena constantly pres-
ent: on the one hand the pursued ones, striving to escape
by all means in their power; and on the other hand the
pursuers, ever trying to outwit those upon which they
prey. If a duck acquires great speed of flight, the Duck The Body of a Bi 309 Hawk must learn to fly still faster. If the duck learns
to crouch close to the reeds when his flight-feathers are
moulted and he is helpless, the hawk must develop
ever sharper eyesight. We may puzzle and puzzle over
a characteristic habit or a colour of some bird, finding
no solution, until we discover some special enemy or
other factor in its life which makes all clear. So, among aggressive colours we may mention the
garb of the penguin, which is steel-gray on the back and
silvery white below; not to protect it from danger, but
to enable it the better to approach fish without alarming
them. The Body of a Bi It is curious how fish-like the coloration of these
birds really is, and they are said frequently to lay feet
and tail together and, drawing their flipper-like wings
to their sides, spring clear of the water again and again,
by a single motion of the back muscles, exactly as the
mammalian dolphins leap ahead of a vessel's bow. mammalian dolphins leap ahead of a vessel's bow. Again, while we find the ptarmigan mimicking the
snow in colour, we find the Arctic Fox, the Snowy Owl,
and the Gyrfalcon, all of which are enemies of this bird,
also garbed in white. The ptarmigan may crouch upon
a drift, but it must ever be on the alert, lest from amid
the snowflakes a white death come suddenly upon it. Nature is terribly just in her plan of life's battles. Nature is terribly just in her plan of life's battles. In the same region with these lives the Ivory Gull,
immaculate as the ice-floe over which it flies, and in its
whiteness we can perhaps read two purposes: a better
chance to elude the fierce Gyrfalcon, and a better chance
to float cloud-like unperceived over the unsuspecting fish
which it seeks for food. FIG. 246.— Black footed Penguin. FIG. 246.— Black footed Penguin. FIG. 247. — Pickerel. (Keller, photographer. From life, swimming.)
PREDACIOUS, AQUATIC ANIMALS, SHOWING AGGRESSIVE
COLORATION. FIG. 247. — Pickerel. (Keller, photographer. From life, swimming.)
PREDACIOUS, AQUATIC ANIMALS, SHOWING AGGRESSIVE
COLORATION. FIG. 252.— Young Robin. FIG. 252.— Young Robin. FIG. 253. — Fallow Deer fawn one day old. BOTH SHOWING SPOTS WHICH ARE ABSENT IN THE ADULTS. FIG. 253. — Fallow Deer fawn one day old. BOTH SHOWING SPOTS WHICH ARE ABSENT IN THE ADULTS. The Body of a Bi 3 1 7 thickly spotted. This gives a clue to the coloration of its
ancestors, — birds probably resembling our Wood Thrush,
and lacking the rufous, immaculate breast of the parents. We find a similar condition existing among many deer,
whose young are spotted, entirely unlike the brown coats
of their parents. FIG. 254. — Nestling Turkey Vulture. (T. H. Jackson, photographer.) FIG. 254. — Nestling Turkey Vulture. (T. H. The Body of a Bi Jackson, photographer.) In many cases the colouring of the downy young is
the opposite of the adult, as in the Turkey Vulture, the
nestling being clad in down of purest white, and ultimately
moulting into the blackish plumage of the parent birds. It would be out of place in this volume to speak further
of the wonderful colours which the Class of birds, as a The Bird 3 1 8 whole, exhibits, or of the beautiful plumes which, as
in the case of the Snowy Egret, are assumed only during
the season of courtship. The great majority are now
explained either as decorations to charm the female, or
as mere by-products of the vitality of the bird, according
as to whether one believes in a greater or less degree of
aBsthetic appreciation among birds. When we consider
the nervous, high-strung natures of birds and realize with
what ease they are thrown into what seems a kind of
trance, it seems unnecessary to credit them with too great
an appreciation of pure beauty. The repetition of many
similar bright spots, as, for example, the eyes of a pea-
cock's train, may well serve to attract and hold the atten-
tion of the female; while the antics and sounds which
many birds bring into play in courtship may appeal in
some more directly psychic way than we know. That
birds do have a certain appreciation of beauty and har-
mony there can be little doubt. When we remember the
jarring discords and clashing tints in which a human
savage takes delight under the name of music and beauty,
we should be very willing to admit some degree of
appreciation to the demure Impeyan Pheasant hen which
chooses among her suitors, clad each in hues such as
artist could never imitate; or the fair Hermit Thrush,
which selects a singer from the incomparable choir of her
serenaders. I believe that future field study and experi-
ments with caged birds will reveal much that we do not
suspect in regard to the causes of coloration. * Extremes in regard to the number of primaries are the three flightless
groups, penguins with approximately 36, ostriches with 16, and cassowaries
with perhaps but 2 feathers which can be called primaries. WINGS E EFORE the front limbs of any creature had be-
come adapted to flight through the air, they
served to assist the hind legs in locomotion on
the ground, and, ages before this, a many-rayed mem-
brane stretched across the primitive fin, aided its owner
in cleaving a way through the water. So, like a palimp-
sest, if we look beneath the outer covering of feathers,
we see, in the wing of the modern bird, the three fingers
hinting of widely different ancestral habits. hinting of widely different ancestral habits. The general structure and appearance of the bills, the
feet, and wings of various birds is the result of a function
characteristic of each. The bills are used to procure
food, the feet to walk or perch, and the wings to propel
the bird through the air. But, as we have seen in the
case of the bill, these organs are put to many other uses
besides the one for which they were primarily adapted. This is only what we should expect when we consider the
relative high position which avian intelligence holds, and
the remarkable extremes of environment with which
these structures — bill, feet, and wings — are brought into
close touch. The photograph of the young heron's wing shows sho The Bird 320 320
the two principal divisions into which the flight-feathers
are divided: the primary feathers, or those growing on
the fingers and wrist-bones, and the secondaries which
sprout from the bone of the forearm. The several feathers FIG. 255. — Young Green Heron, showing various divisions of wing-feathers. FIG. 255. — Young Green Heron, showing various divisions of wing-feathers. supported by the thumb are also very distinctly shown. When a wing is greatly elongated it is the secondary
feathers which are increased in number, the two extremes
being represented by the hummingbird and the albatross, Wings 32 Wi 321 each of which has ten primaries, but the one has six and
the other forty secondaries (Figs. 259, 260).* the other forty secondaries (Figs. 259, 260).*
Let us observe the wings of living birds in the woods
and fields or in a zoological park and see what of interest
we can discover. We have all noticed how well adapted FIG. 256. — Great White Heron stretching its wing. (E. R. Sanborn,
photographer.) FIG. 256. — Great White Heron stretching its wing. (E. R. WINGS Sanborn,
photographer.) to its owner's many uses is the foot of a parrot — how
hand-like it is, — and now if we again watch one of these
birds we will see that, as we should expect from its being to its owner's many uses is the foot of a parrot — how
hand-like it is, — and now if we again watch one of these
birds we will see that, as we should expect from its being The Bird 2 322
so much like a human hand, it is not a good walking foot. When a parrot is in great haste to reach some object
on the ground without flying, it waddles awkwardly, " toe-
ing in" and frequently tripping up. When this happens,
out fly the wings, and, as if reverting to some clouded
memory of the habits of its pre-Jurassic forefathers, it
walks on all fours. A young Canada Goose, when climb-
ing about its nest, or a Fish Hawk in the downy nestling FIG. 257. — Nestling Catbird, supporting itself, lizard-like, on all four limbs. 257. — Nestling Catbird, supporting itself, lizard-like, on all four limbs. plumage, does the same thing, and young birds of many
species, when too young to stand, push themselves along
the ground with feet and wings; a young grebe doubtless
being the most accomplished in this motion. In certain
adult birds, such as the swan, Osprey, Turkey Vulture,
and the various ostrich-like birds, there are perfect claws
at the tips of one or more of the skin-bound wing-fingers. These are true relics of a lizard-handed ancestry. plumage, does the same thing, and young birds of many
species, when too young to stand, push themselves along
the ground with feet and wings; a young grebe doubtless
being the most accomplished in this motion. In certain
adult birds, such as the swan, Osprey, Turkey Vulture,
and the various ostrich-like birds, there are perfect claws
at the tips of one or more of the skin-bound wing-fingers. These are true relics of a lizard-handed ancestry. These are true relics of a lizard-handed ancestry. Before going on to find the more curious uses to which Wings 323 wings are put, we will look at certain birds whose flight
can teach us something interesting. WINGS If a pheasant in
captivity becomes suddenly alarmed, or its spacious aviary
tempts it to rise from the ground, we hear a great whirr, —
broad, round-curved wings buzz in a half -circle of haze
around the bird and it is off like a shot to the farther FIG. 258. — Young Green Heron, reaching out with its wing toward a branch which
it hooked with the sprouting feathers, and steadied itself for a new foothold. FIG. 258. — Young Green Heron, reaching out with its wing toward a branch which
it hooked with the sprouting feathers, and steadied itself for a new foothold. end of the runway. It may go right through the sash
and pane of glass — such is the impetus gained in this mad
rush. Fortunate it is for these birds, and for their cousins,
tne grouse and quail, that they can thus spring up and
escape from foxes and other enemies to whom their scent
so often betrays them. If the pheasant were at liberty, end of the runway. It may go right through the sash
and pane of glass — such is the impetus gained in this mad
rush. Fortunate it is for these birds, and for their cousins,
tne grouse and quail, that they can thus spring up and
escape from foxes and other enemies to whom their scent
so often betrays them. If the pheasant were at liberty, The Bird 3 we should see that this burst of speed would end in
a long, slowly descending sail, and with wings held mo-
tionless the bird would sink into the nearest cover. It is
most interesting and exciting to walk through a field
of tall grass where many pheasants are feeding, and see
them shoot up to the right and to the left; a hen with
her brood waiting until one's foot is almost upon her
before booming away. In a zoological park we may observe another extreme
of bird flight by watching a condor take wing. He waits
until a breeze is blowing and then, facing the direction
from which it comes, he runs with all his might, flapping
awkwardly until sufficient headway is gained, when strong
downward strokes carry him to the perch he has selected. WINGS We may, at first thought, pity him, but if we could see
him soaring for hours high among the cloud-peaks of his
native Andes, we should instead pity the low-flying pheas-
ant. These two examples — the pheasant and the condor —
show what differences may be found in flying birds, and
as we examine the wings of other species, we find that
each is perfectly adapted to the wants of its owner. A
wing is a most delicately adjusted organ; its feathers
being just strong enough to lift the body of its owner
into the air, and, like evenly balanced scales, the least
excess or lack of use is quickly met by a reaction. Com-
pare the Black Skimmer of the seas, which is only eighteen
inches in length, but whose long wings expand four feet,
with a stubby-winged quail or grouse. with a stubby-winged quail or grouse. There are some species of flycatchers with wonderful Wings 325 FIG. 259. — Whig of Hummingbird with 16 flight-feathers. 1/2 natural size. FIG. 259. — Whig of Hummingbird with 16 flight-feathers. 1/2 natural size. FIG. 260.— Wing of Albatross with 50 flight-feathers. 1/28 natural size. FIG. 260.— Wing of Albatross with 50 flight-feathers. 1/28 natural size. The Bird 326 326
powers of flight. When perched on a branch, they can
evade the shot from a shot-gun. It is said that one will
sometimes "chase another for three or four minutes,
doubling, turning, twisting, and shooting, now brushing
the grass, now rising to a height of at least two or three FIG. 261. — Condor about to take flight. FIG. 261. — Condor about to take flight. hundred feet, and all the movements so rapid that the
eye can scarcely follow them; and at the end of it would
go back to his own chosen weed-stalk, apparently without
a feather ruffled." a feather ruffled."
Any attempt to explain the mechanics of the way of a
bird in the air would at most be imperfect in the present Wi 327 state of our knowledge. Suffice it to say that if we will
think of a bird flying through the air when we ourselves
are swimming in the water, we can realize the achievement
more vividly than from any amount of descriptions and
diagrams. The under surface of a bird's wing is concave; and FIG. 262. WINGS — Wing of living Golden Pheasant; rounded and curved for short,
sudden flight. FIG. 262. — Wing of living Golden Pheasant; rounded and curved for short,
sudden flight. while the front edge is rather straight and firm, the hinder
rim of the feathers is soft and yielding; thus a downward
stroke both raises the bird or holds it sustained at the
height already reached and urges it in a forward direction. Similarly we push our hollowed palms backward and pro-
pel ourselves through the denser medium of water. The manner of flight varies greatly in different birds The Bird 328 328
and is often so characteristic that when too far off to dis-
tinguish the colour of its plumage, or for its notes to
reach our ears, the bird may be recognized by the undu-
lations or the directness of its flight. No one who has
ever visited the tropics can have failed to admire the Fro. 263. — Wing of living Herring-gull; long and narrow for slow,
continuous flight. Fro. 263. — Wing of living Herring-gull; long and narrow for slow,
continuous flight. soaring vultures, — spots of black swinging across the
heavens or swooping low in grand arcs over the palms. Gulls and their kindred fly steadily with continuous wing-
beats, which, however, are much less rapid than in the
flight of a duck or a parrot. Many sparrows have an
abrupt jerking motion, hitching themselves over trees soaring vultures, — spots of black swinging across the
heavens or swooping low in grand arcs over the palms. Gulls and their kindred fly steadily with continuous wing-
beats, which, however, are much less rapid than in the
flight of a duck or a parrot. Many sparrows have an
abrupt jerking motion, hitching themselves over trees Wi 329- and bushes; while goldfinches and woodpeckers swing
past in long undulations, a loop and a catch, a loop and
a catch, — with wings wide extended, then quickly closed. Hummingbirds have a remarkably insect-like flight; the
rapid reflex whirr of the wings holding them perfectly
still, poised in mid-air. still, poised in mid-air. When ornithologists think that they have formed a
correct theory of flight and that, given such and such
conditions, certain results must follow, such a bird as the
Crested Screamer soars into their mental atmosphere and
upsets every calculation. WINGS Such a bulky and short-winged
bird, by all good "rules" of flight, should confine itself
to short laboured efforts, barely skimming the low bushes
of its South American haunts! But it refuses to be thus
limited. Of this species it is said : " The Screamer is a very
heavy bird, and rises from the ground laboriously, the
wings, as in the case of the swan, making a loud noise. Nevertheless it loves soaring, and will rise in an immense
spiral until it wholly disappears from sight in the zenith,
even in the brightest weather; and considering its great
bulk and dark colour, the height it ultimately attains
must be very great. On sunny windless days, especially
in winter and spring, they often spend hours at a time
in these sublime aerial exercises, slowly floating round
and round in vast circles, and singing at intervals. How
so heavy and comparatively short-winged a bird can sus-
tain itself for such long periods in the thin upper air to
which it rises has not yet been explained." which it rises has not yet been explained."
I find in my journal the following account of a flight
of vultures which we saw in a desolate alkali desert in The Bird 33° 33°
western Mexico: "One of the most wonderful exhibitions
of bird-flight came to us to-day as we left the alkali plain
and rode among the mesquite scrub. A confused mass
of black appeared in the air which, as we advanced, re-
solved itself into hundreds of individual black specks. FIG. 264. — Crested Screamers. FIG. 264. — Crested Screamers. The atmosphere was so deceptive that what at first seemed
to be a vast cloud of gnats close at hand, was soon seen
to be a multitude of birds, and when a quarter of a mile
away we knew them to be vultures. Three burros lay
dead upon the plain. This we knew yesterday, and here
were the scavengers. Never had we seen Vultures more Wings 33
numerous or in more orderly array. A careful scrutiny
through our glasses showed many scores of Black and
Turkey Vultures walking about and feeding upon the
carcasses of the animals, and from this point there ex-
tended upward into the air a vast inverted cone of birds,
all circling in the same direction. WINGS From where we sat
upon our horses there see ed not one out of place, the FKJ. 265. — Turkey Vulture soaring. FKJ. 265. — Turkey Vulture soaring. outline of the cone was as smooth and distinct as though
the birds were limited in their flight to that particular
area. It was a rare sight, the sun lighting up every bird
on the farther side and shadowing black as night those
nearest us. Through one's partly closed eyes the whole
mass appeared composed of a myriad slowly revolving
wheels, intersecting, crossing each others' orbits, but never
breaking their circular outline. The thousands of soaring outline of the cone was as smooth and distinct as though
the birds were limited in their flight to that particular
area. It was a rare sight, the sun lighting up every bird
on the farther side and shadowing black as night those
nearest us. Through one's partly closed eyes the whole
mass appeared composed of a myriad slowly revolving
wheels, intersecting, crossing each others' orbits, but never
breaking their circular outline. The thousands of soaring The Bird 3 forms held us spellbound for minutes before we rode
closer. Now a change took place, as gradual but as sure
as the shifting clouds of a sunset. Until this moment
there was a tendency to concentrate at the base of the
cone, that portion becoming more and more black until it
seemed a solid mass of rapidly revolving forms. But,
at our nearer approach, this concentration ceased, and
there was perfect equilibrium for a time; then, as we
rode up a gentle slope into clearer view, a wonderful
ascent began. WINGS Slowly the oblique spirals swing upward;
the gigantic cone, still perfect in shape, lifts clear of
the ground and drifts away, the summit rises in a
curve which, little by little, frays out into ragged lines,
all drifting in the same direction, and before our very eyes
the thousands of birds merge into a shapeless undulating
cloud which rises and rises, spreading out more and more
until the eye can no longer distinguish the birds which
from vultures dwindle to motes, floating and lost among
the clouds." the clouds."
Concerning the greatest extent of wing which any
bird possesses, there are records of a Wandering Albatross
which measured fourteen feet from tip to tip, but the
condor of South America exceeds this, certain individuals
having an expanse of fifteen feet. Having considered the finest flyers among the birds,
we may now begin to go down the scale and see what
has happened when certain species have deliberately dis-
carded the wonderful power of flight with which Nature
has provided them and for which human inventors are
so earnestly striving. But always we must remember Wi 333 that this restriction and disuse have been to subserve some
good and useful purpose, — food perhaps being more easily
obtained, or enemies avoided by terrestrial or aquatic
locomotion. Functional radiation, working always for
the good of the race, once gave to all birds the power of
traversing the globe, passing high over sea and land; but
later this was withdrawn, until in some cases their wings
have become a mockery. The wings of the Owl Parrot
of New Zealand are of full size, but the muscles are so en-
cased in fat that they are useless for flight. These par-
rots feed on ground-mosses, and being nocturnal and
tnerefore having few enemies, their only use for wings
is occasionally to sail gently to earth, like a Flying Squir-
rel, from the trees in the hollows of which they some-
times roost. For this purpose their flabby muscles are
perfectly suited. perfectly suited. The Spotted Tinamou of South America is one of a
number of birds which have not quite lost the power of
flight, but in which, as in the first attempts of a young
bird, almost no control is possessed over the direction
or height of their flight. WINGS In fact, the condition is much
the same as that of a man in an ordinary balloon, who
is at the mercy of the wind and the sustaining power of
the gas. Hudson gives the following interesting account
of this bird: "It is an exceedingly rare thing to see this
bird rise except when compelled. I believe the power
of flight is used chiefly, if not exclusively, as a means of
escape from danger. The bird rises up when almost trod-
den upon, rushing into the air with a noise and violence
that fill one with astonishment. It continues to rise The Bird 3 at a decreasing angle for fifty or sixty yards, then gradu-
ally nears the earth, till, when it has got to a distance of
two or three hundred yards, the violent action of the wing
ceases, and the bird glides along close to the earth for
some distance, and either drops down or renews its flight. I suppose many birds fly in much the same way; only FIG. 266. — South American Tinamou. FIG. 266. — South American Tinamou. this tinamou starts forward with such amazing energy
that, until this is expended and the moment of gliding
comes, the flight is just as ungovernable to the bird as
the motion of a brakeless engine, rushing along at full
speed, would be to the driver. The bird knows the danger
to which this peculiar character of its flight exposes it Wi 335 so well that it is careful to fly only to that side where
it sees a clear course It is sometimes, however, compelled
to take wing suddenly, without considering the obstacles
in its path; it also often miscalculates the height of an
obstacle, so that for tinamous to meet with accidents
when flying is very common. In the course of a short
ride of two miles, during which several birds sprang up
before me, I have seen three of these tinamous dash
themselves to death against a fence close to the path,
the height of which they had evidently misjudged. I
have also seen a bird fly blindly against the wall of a
house, killing itself instantly. A brother of mine told
me of a very curious thing he once witnessed. He was
galloping over the pampas, with a very violent wind blow-
ing in his face, when a tinamou started up before his
horse. WINGS The bird flew up in the air vertically, and, beat-
ing its wings violently, and with a swiftness far exceeding
that of its ordinary flight, continued to ascend until it
reached a vast height, then came down again, whirling
round and round, striking the earth a very few yards
from the spot where it rose, and crushing itself to a pulp
with the tremendous force of the fall. It is very easy to
guess the cause of such an accident: while the tinamou
struggled blindly to go forward, the violent wind, catch-
ing the under surface of the wings, forced it upward, until
the bird, becoming hopelessly confused, fell back to earth. I have often seen a swallow, gull, or hawk, soaring about
in a high wind, suddenly turn the under surface of its
wings to the wind and instantly shoot straight up, appar-
ently without an effort, to a vast height, then recover The Bird The Bird 3 itself and start off in a fresh direction. The tinamou,
when launched on the atmosphere, is at the mercy of itself and start off in a fresh direction. The tinamou,
when launched on the atmosphere, is at the mercy of FIG. 267.— Feathers of Ostrich and Condor. FIG. 267.— Feathers of Ostrich and Condor. chance; nevertheless, had this incident been related to
me by a stranger, I should not have recorded it." me by a stranger, I should not have recorded it."
So in this bird we have a most rare and suggestive Wi 337 instance of a condition where an important organ is actu-
ally in process of losing its primary function, and in so
doing becomes a source of danger to the bird. doing becomes a source of danger to the bird. In the waters of the sea near the Falkland Islands is
a duck known as the Steamer or Side-wheel Duck. The
young birds of this species are good flyers and whistle
through the air on strong pinions. But maturity, instead
of bringing, as in most birds, a fully perfected power of
flight, takes from them what they have, and after the first
moult they are helpless to rise above the great waves
of their haunts. However, this duck finds another use
for its wings, and the stiffness which forbids their being
used in the air makes of them bladed paddles which are
all the better for their lack of flying power, and with wings
and feet these birds make remarkable speed through the
water — "twelve or fifteen miles an hour" — and they are
thus able to live out their lives in safety. Thus the study
of the flight of these birds carries us a step farther than
the tinamou, with the all-important difference that, in this
case, loss of the primary function is compensated by a
direct adaptation of the wing to tlie new conditions of life. In the ostriches and their near allies the extreme reduc-
tion of wings is to be found, and yet in the true ostriches
and rheas the great expanse of soft feathers is a consid-
erable help to the birds when running at full speed, acting
as a sail or aeroplane to assist in the onward motion. The Bird But the contrast between a loose, open-work feather from
the wing of one of these birds and a compact, firmly vaned
plume from a condor's wing is very striking. The casso-
wary has from four to six flight-feathers, but, far from The Bird 3 being of any use in supporting his great frame, they are
so vestigial that they look exactly like black slate-pencils
projecting in a row from the little fleshy flap which con-
tains the evidence of his full-winged ancestors tains the evidence of his full-winged ancestors
A full-grown ostrich was once imported to this country
from Abyssinia. When the native keepers learned that the
bird was to be sent away, they surreptitiously plucked the FIG. 268. — Wing of Cassowary, showing degenerate flight-feathers. FIG. 268. — Wing of Cassowary, showing degenerate flight-feathers. poor creature, until but few feathers were left on its body. The bird was tame, and, by keeping its attention busy
with a basket of carrots, I inserted a piece of white card-
board beneath one of its skinny, denuded wings and se-
cured an excellent photograph (Fig. 269). This clearly
shows the black, curved claws on the first two fingers. In this same bird I noticed that occasionally the crooked
forearm would be raised, the claw at the end of the wing poor creature, until but few feathers were left on its body. The bird was tame, and, by keeping its attention busy
with a basket of carrots, I inserted a piece of white card-
board beneath one of its skinny, denuded wings and se-
cured an excellent photograph (Fig. 269). This clearly
shows the black, curved claws on the first two fingers. In this same bird I noticed that occasionally the crooked
forearm would be raised, the claw at the end of the wing Wi 339 drawn up, and the ostrich would scratch its body or head
with this interesting finger relic ! When the plume feathers
of the wing are full grown, the foot or leg is thus used, FIG. 269. — Wing of Ostrich, showing reptile-like claws. FIG. 269. — Wing of Ostrich, showing reptile-like claws. the head or neck being rubbed against its roughened
scales. The Bird The Great Auk — a sea-bird which has become extinct
within the last sixty years — was without the power of
flight, and its living allies, the Razor-billed Auks and
Murrelets, have very small wings and are rather weak The Bird 3 flyers. The latter, in fact, use their wings, the feathers
of which have very stiff and long quills, as much in diving
under water as in flying in the air, and, strangely enough,
they are said to swim breast upward, propelling themselves
by means of both wings and feet. Grebes, too, are very
weak of wing, and these birds cannot rise from level ground,
no matter how much of a fluttering run is taken, and even
in the water much splashing and headway are needed. in the water much splashing and headway are needed. Perhaps the most wonderful birds in the world are
penguins, and the strangest part of these strange birds
is the wing. There is no doubt that they are descended
from birds which possessed the power of flight; but the
penguins have discarded this gift and have returned to
a life in the sea, whence in long ages past their forebears
had crawled out upon land. As in the ostriches, the
relics of flight-feathers have increased greatly in number,
but have become small and scaly, and the wings have
virtually become flippers or fins. Instead of a given num-
ber of feathers, divided into well-marked series, the pad-
dles of a penguin are covered thickly with small feather-
scales, and the rigidity of the wings, together with the
rotary movement at the shoulder- joint, make the propeller
of a ship an apt simile. The colour of the feather-scales
on the upper side of the wmg is dark, like the back of
the bird, but those on the under side have run rampant,
the white and black being mixed irregularly, not corre-
sponding even in the two wings of an individual bird. The outline of the wing is exactly like that of a shark's
fin, the flatness and breadth including even the bones,
while (also like a fin) all of the bending quality of a wing The Bird 2 342
is lost, — all the flexibility of wrist and elbow. With
these propellers the penguins fly through the water, with
almost the identical motion of a bird in the air. The Bird Though
it is usually asserted that the wings move alternately,
this was never the case with a pair of Black-footed Pen-
guins which I carefully observed. As regards the speed
of swimming, I found that one of these birds, though in
bad health at the time and so weak that it could take
but a few steps on land, was able to progress under water FIG. 271. — Penguin swimming with its wings. FIG. 271. — Penguin swimming with its wings. considerably faster than a man could walk an equal dis-
tance on land. The greatest speed was about seven miles
an hour; but I have no doubt that when in full health
this rate can be far surpassed. Surely no fairy-tale can match the marvellous evo-
lution of a penguin's wing : fin becoming hand, hand
evolving into wing, and wing reacting to the environ-
ment of long ago and again taking on all the outward
characteristics of a fin! Wings 343 FIG. 272. — Wing of Black-footed Penguin, top view. FIG. 273.— Wing of Black-footed Penguin, side view. FIG. 272. — Wing of Black-footed Penguin, top view. FIG. 272. — Wing of Black-footed Penguin, top view. FIG. 273.— Wing of Black-footed Penguin, side view. FIG. 27-4. — Wing of Black-footed Penguin, under surface of wing. ALL FROM THE LIVING BIRD. FIG. 272. — Wing of Black-footed Penguin, top view. FIG. 272. — Wing of Black-footed Penguin, top view. FIG. 273.— Wing of Black-footed Penguin, side view. FIG. 273.— Wing of Black-footed Penguin, side view. FIG. 27-4. — Wing of Black-footed Penguin, under surface of wing. ALL FROM THE LIVING BIRD. FIG. 27-4. — Wing of Black-footed Penguin, under surface of wing. ALL FROM THE LIVING BIRD. The Bird The Bird 3 We have seen how wings guard their owners from the
risk of sudden surprises from enemies, and now let us
observe how, in a sleeping bird, the tender nostrils and
eyes are protected against cold and other dangers. Birds
do not put their heads under their wings, but behind them,
often using the shoulder-feathers as cover. It is inter- FIG. 275. — Green Heron with head behind wing. FIG. 275. — Green Heron with head behind wing. esting to see how many birds, from all quarters of the
earth, have this same habit. The pelican, however, de-
parts from this custom and snuggles his tremendous bill
between the feathers in the centre of his back, and flops
both wings up so as completely to cover it. Even the
cassowary vainly tries to tuck his bill behind his absurd
wing. His smaller wing-coverts are mere soft, loose hair- esting to see how many birds, from all quarters of the
earth, have this same habit. The pelican, however, de-
parts from this custom and snuggles his tremendous bill
between the feathers in the centre of his back, and flops
both wings up so as completely to cover it. Even the
cassowary vainly tries to tuck his bill behind his absurd
wing. His smaller wing-coverts are mere soft, loose hair- Wi 345 like shafts, while the larger quills, as mentioned before,
are reduced to four or six horny sticks. If we watch an owl flying about its cage at night, or
if, in the woods, an owl passes near, his shadow in the
moonlight is all that warns us of his presence. The feathers
of an owl's wing are soft and downy, and the bird moves
as lightly as a falling leaf. Little warning, except by FIG. 276.— Trumpeter Swan asleep. FIG. 276.— Trumpeter Swan asleep. sight, the mice and birds have of its deadly presence. Few birds have a flight as noiseless as that of owls, and
in some species the motion of the wrings makes, as we
noticed in the pheasant, a very audible sound. When a
widgeon rises from the water, the whistling of its quills,
so dear to the ears of the sportsman, is quite shrill. A
dove claps its wings together above its back while gain-
ing impetus for flight. The characteristic sound from
which a hummingbird takes its name is well known. The Bird 3 When wild geese and swans nest in captivity, their
wings are put to most excellent use as weapons of de-
fence, and of course this use must come into play fre-
quently when nesting in their native haunts. I have
seen a man knocked breathless by a Canada gander who
thought his nest in danger. When preparing for attack,
the bird approaches hissing, with head stretched low along
the ground, and suddenly, without warning, launches FIG 277.— Trumpeter Swan preparing to attack an intruder with its wings. FIG 277.— Trumpeter Swan preparing to attack an intruder with its wings. itself straight at one's breast and, clinging with bill and
claws, beats a tattoo with the hard bend of its wings. One is not likely to forget such a drubbing for a long
time. The wings of certain birds are armed with weapons
of offence, such as the Spur-winged Goose, Jacana, Plover,
and Screamer. The Spur-winged Goose is a really danger-
ous antagonist and can strike incredibly strong blows,
bringing the sharp spur to bear with telling effect. These itself straight at one's breast and, clinging with bill and
claws, beats a tattoo with the hard bend of its wings. One is not likely to forget such a drubbing for a long
time. The wings of certain birds are armed with weapons
of offence, such as the Spur-winged Goose, Jacana, Plover,
and Screamer. The Spur-winged Goose is a really danger-
ous antagonist and can strike incredibly strong blows,
bringing the sharp spur to bear with telling effect. These Wings 347 spurs are not claws, but correspond in structure to the
ordinary spurs on the legs of a rooster. The great heavy-headed and heavy-bodied hornbills
fly with great effort, and it is said upon good authority
that when passing low overhead they make a noise like
a steam-engine. Although not strictly within the prov- FIG. 278. — Spur-winged Goose. FIG. 278. — Spur-winged Goose. ince of this volume, mention should be made of the inten-
tional use of the wings as instruments of sound, — to at-
tract the females, as in our Ruffed Grouse and other birds. A little Bush Warbler of Africa has indeed never been heard
to utter a note, seeming to depend upon an occasional
whirr of wings, in lieu even of the usual call-note or chirp. The Bird ince of this volume, mention should be made of the inten-
tional use of the wings as instruments of sound, — to at-
tract the females, as in our Ruffed Grouse and other birds. A little Bush Warbler of Africa has indeed never been heard
to utter a note, seeming to depend upon an occasional
whirr of wings, in lieu even of the usual call-note or chirp. 3 The Bird In the woodcock we find the vane of the three outer
primaries of the wing remarkably narrowed and stiffened;
probably a direct adaptation for the production of the
high, whistling sound which plays so important a part
in its aerial courtship performance. FIG. 279. — Wing ornaments of Twelve-wired Bird of Paradise. FIG. 279. — Wing ornaments of Twelve-wired Bird of Paradise. As upon all other parts of the bird's body, we find
beautiful decorations upon the wings — inexplicable unless
we are willing to credit the females with appreciation of,
or at least a reaction to, these beauties. Otherwise we
know not the uses of the brilliant wing-mirrors of ducks,
or the scarlet wax-like tips of the Cedar-bird's feathers, Wings 349 349
or the bizarre decoration of the Twelve-wired Bird of
Paradise. A strange appendage is found in the wing of the West
African Goatsucker. Conspicuous enough when the bird
is flying, it is wonderfully protected when the bird rests, FIG. 280.— Wing of Woodcock. FIG. 280.— Wing of Woodcock. as is its wont, upon the ground among tall, feathery-
topped grasses. From each wing a single long feather
extends in an upward direction, almost bare of barbs
for most of its length, but tipped with a mottled, loose-
vaned tuft which corresponds very perfectly with the
flower-heads of the grasses among which it lives. As The Bird 3 this decoration, so protective and yet so beautiful, is
assumed only during the breeding season, its use is doubt-
less to aid in attracting the attention of the females. Herons and other birds make still another use of their
wings and the long, tough flight-feathers: as shields for
parrying the blows of a rival, or to catch the poison of
a snake when it strikes and thus give an opportunity
to seize and despatch the reptile. Two Snowy Egrets
will sometimes fence with each other in play, and use
beak and wing as a soldier would use sword and shield. beak and wing as a soldier would use sword and shield. I once saw the wing of a bird used in an entirely original
manner — a use peculiar, doubtless, to this individual. The Bird Several spoonbills suffered severely from the frozen
ground upon which they were forced to stand, and no
method of relief was found, except by one of their num-
ber, who every night stretched one wing beneath him,
drew up one foot deep into his plumage, and with the
other stood upon the tips of the primaries. Much might be written concerning the swiftness of
birds' flight, but so much of exaggeration has entered
into estimates of this kind that it would be difficult to
select facts and figures of indisputable verity. However,
it may be asserted as at least within the actual facts that
ducks can attain a speed of ninety miles an hour. An
apparently well-authenticated record of a swallow's flight
at Antwerp is as follows: A gentleman arranged a flight
of homing pigeons from Compiegne to Antwerp, — a dis-
tance of one hundred and forty-eight miles, — and with the
pigeons he liberated a swallow captured on her nest under
the eaves of his house in Antwerp. The swallow, which 03
I
a
O
r
0, 03
I
a
O
r
0, O
r
0, The Bird The Bird 3 was marked for identification, covered the distance in
one hour and eight minutes, or at the extraordinary speed
of about two miles and three hundred yards per minute. The first pigeon to arrive took four hours and a quarter
to make the journey. \
FIG. 282. — Terns in flight. (Photograph provided by the American Museum
of Natural History.) FIG. 282. — Terns in flight. (Photograph provided by the American Museum
of Natural History.) FEET AND LEGS ARRIED far and wide by the power of flight,
no two species of birds have exactly similar
environments. When the wings cease their la-
bour and are folded close to the sides, the bird must depend
upon its feet to carry it to its food and to keep it out of
danger, whether its footing be in a tree-top or on a cliff;
in shallow water or on the deep; in mud, sand, or snow. Thus we realize the need for many varied adaptations
in the way of feet and legs. A Although birds are descended from five-toed ancestors,
yet no living wild bird, and none of those which we know
only as fossils, has more than four toes on each foot. The
disposition of these toes — four, three, or two, as the
case may be — is always in accordance with the habits of
the bird. The most common type of avian foot is that in which
the arrangement is of three toes in front, with the fourth,
corresponding to our great toe, pointing backward. This
was the arrangement in our first bird, the Archceopteryx,
and for perching birds, as well as for many others with
very different habits, it has stood the test of six millions 353 The Bird 354 of years, or thereabouts, since the days of its venerable
prototype. prototype. This is the kind of generalized organ which, we should
think, would be able to cope with changes in the bird's
surroundings more successfully than any other; but that
this theory fails when put to the test is proved by the
variety of specialized toes and legs which we may observe FIG. 283.— Foot of Alligator. FIG. 283.— Foot of Alligator. among the birds on the earth to-day. Indeed, in the
variety of uses which they subserve, the feet and legs
of birds are second only to the bills. A classification of birds, generally accepted for many
years, was based on the uses of the feet, or mode of loco-
motion. In this scheme birds were divided into runners,
scratchers, climbers, swimmers, perchers, etc. Although Feet and Legs 355 these, as exact divisions, have long since been abandoned,
yet it is worthy of note that even in the most modern
classifications many of these groups hold good in the
main, although based on other and more fundamental FIG. 284.— Foot of Brown Pelican. FIG. FEET AND LEGS 284.— Foot of Brown Pelican. characters. Examples of these are the ostrich-like birds,
or runners; the fowl-like birds, or scratchers; and the
Passeres, or perching birds. But there is no doubt that
several unrelated groups have independently acquired the The Bird 3 specialized type of foot which is adapted to climbing or
to swimming, so that any classification based on such
similarity of locomotion is obviously false. similarity of locomotion is obviously false. From the tiny limbs of a hummingbird to the gigantic
shanks of an ostrich, the legs of birds, with a very few
exceptions, are covered with scales, most emphatic re-
minders of the reptilian ancestry of both these extreme
forms of feathered life. The real foot of a bird, as the
term is used in speaking of other animals, extends to the
backward-bending joint, or heel. Part of the lower leg FIG. 285.— Foot of Raven. FIG. 285.— Foot of Raven. is thus concealed by the feathers and skin, while the upper
leg, or thigh, is generally wholly within the body, as we
saw in the chapter treating of the framework. As before, we must call on the crow, in many respects
standing very near the top of the scale of bird life, yet
which has found it good to hold to the typical bird's foot. And indeed it serves him well, for with it he can walk on
snow or ice; w&de in shallow water; perch in trees; scratch
or claw the ground and hold down a crab's carapace,
while he extracts the edible portion. Not only this, but
he can hop like a sparrow or walk like a lark at will. Feet and Le 357 We have hardly to leave the group of birds to which
the crow belongs to find dozens of interesting and unex-
pected adaptations of the feet to unusual habits. For
example, the Rhinoceros-birds of Africa attach them-
selves to some of the larger mammals, such as buffalos,
rhinoceri, or antelopes, and spend much of their time
in freeing these animals from troublesome ticks and other
parasites. The power which these birds possess in their
feet and legs is remarkable. FEET AND LEGS Millais says of them: "The
prehensile power of the claws is, as I found by experience,
so great that when a dead bird which had grown stiff
was thrown on to the back or sides of an ox, so that the
feet touched the animal's hide, the claws held fast at
once and could not be withdrawn. It is most interesting
to note the way in which a party of these birds will move
about on the body of a horse or ox, searching every part of
him as they run or hop over it in the most lively fashion. At the risk of being accused of telling a traveller's yarn,
I must state the fact that they can hop backward quite
as well as forward, and they often make long drops down-
ward from the shoulders to the foreleg, or down the side
of the animal whose coat they are engaged upon. It
is quite immaterial to them how or in what direction
they move." they move."
No hard and fast laws can be laid down, but it is gener-
ally the rule that birds which are especially at home in
the trees usually hop with both feet simultaneously
when on the ground. Ground nesters and feeders, such
as the Meadow Lark, Bob-white, and Vesper Sparrow,
usually walk or run. The Bird 3 The great Order of perching birds (Passeres) shows to
what varied uses the typical foot can be put. All birds
of this Order have three toes in front and one behind,
and there is scarcely a place on the globe to which these
birds have not adapted themselves; and recently too,
as would seem probable from the similarity of the foot-
type running through all. type running through all. This very foot holds much of interest too, if we con-
sider it from another point of view. Many apes and
monkeys, and we ourselves, still have the five fingers and
toes which we suppose was the number originally devel-
oped upon the limbs of the vertebrate prototype; while
horses and deer — animals much lower in the scale of
life — have had the five original digits reduced to one
or two. FEET AND LEGS So among birds the ostriches and some other
low forms have become extremely specialized in the same
respect, possessing but two or three toes, while those
birds which in mental and physical attributes excel all
others of their Class are still more reptilian, and thus
more primitive — more Archseopteryx-like — in possessing a
larger number of digits — four. Thus when we speak
of an animal as high or low in the scale of life, we must
carefully distinguish between mere specialization and
actual upward progress, mentally or physically, toward
some ideal goal. The branch of a tree, which stretches
horizontally farthest from the parent trunk, is not likely
to be the one which reaches upward high enough to catch
the first rays of the morning sun. The majority of the Passeres are arboreal and the
strength of the tiny tendons which run down the leg and 359 through each toe is sufficient to clasp and unclasp a thou-
sand times a day, and to hold and balance the bird on
whatever bending twigs or wind-blown foliage it chances
to alight. In this matter of perching the hind toe plays
an important part, so much so that when the necessity
for grasping ceases, this digit begins to wax flabby and
weak and often becomes reduced in size. FIG. 286.— Nuthatch on tree,
clinging upside down. FIG. 287. — Nuthatch clinging to a gloved
hand. (Bowdish, photographer.) FIG. 287. — Nuthatch clinging to a gloved
hand. (Bowdish, photographer.) FIG. 286.— Nuthatch on tree,
clinging upside down. The creepers, Fig. 240, are passerine woodpeckers
in habit and forever wind their spiral paths about the
tree-trunks. But the nuthatch is the marvel of the
whole Class of birds in this climbing ability. With no
support whatever from the tail, and without special
adaptation of toes, it defies all laws of gravitation and
creeps up and down or around the vertical trunks, as if
on a level surface. Never a misstep, never a slip, but The Bird 36o 36o
each foothold as secure as if its feet were vacuum-
cupped. In the swallows the feet are very small, having fallen
into disuse with the great increase of the power of flight. FEET AND LEGS Orioles and weaver-birds make occasional use of their
feet to hold a strand of grass or string which they are
weaving with their beaks into their elaborate nests, and
certain flycatchers pounce upon and hold their insect FIG. 288. — Swallow, showing small size of feet. FIG. 288. — Swallow, showing small size of feet. prey as an owl grips a bird, or a jay clings to a nut; but
with the exception of a few such cases, the feet of perching
birds serve principally the function of locomotion. As variation in habitat or haunt depends so much upon
the power of locomotion, it will not be out of place to
mention here, in rather more detail than usual, a splendid
example of adaptive radiation which we can all verify for
ourselves. There is no more wonderful fact in Nature than the way Feet and Le 361 in which birds have inherited the earth. When we realize
the immense advantage which the power of flight gives
to them, we do not marvel at this remarkable distribu-
tion, but the more we think about it the more wonders
appear. The utmost efforts which man has made to
reach the North Pole have shown flocks of birds winging
their way still farther to the North, heedless of the ter-
rible cold. In the heat of deserts and the . sweltering
jungles of the tropics, birds find congenial haunts and
abundant food. Thousands of miles out at sea, on the
highest mountains, and even in dark underground tun-
nels; the whole day — twilight, midnight, and dawn, —
all have been conquered by these tireless, energetic feath-
ered ones. When we see a large collection of birds, we can appre-
ciate how they are adapted to such varying conditions
of temperature, of moisture, of light, and of altitude. Their bodies, wings, legs, feet, and tails — in fact every
organ and member is of all sizes and shapes, and shows to
what condition of life the individual is suited. FEET AND LEGS But when
we come to know birds better, and we realize that there
are wheels within wheels, that behind these very evident
divisions into Families and Orders there are lesser groups,
among the members of which the competition is no less
keen, we look for and find gentler gradations and adapta-
tions which, in their way, are more to be wondered at
than the larger, more radical differences; for these birds
have changed their habits and haunts without waiting
for Nature to adjust their wings or their feet. They have
taken the initiative as it were, and, like a man of letters The Bird 3 who is suddenly forced to work at some arduous manual
labour, they have entered on new ways of life — ways to
which their structure seems but ill adapted, and yet,
by the very daring of their efforts, they have won success. by the very daring of their efforts, they have won success. The great-grandfathers, many times removed, of the
modern Families of birds lived lives which were much
broader and more generalized than those of their descend-
ants of to-day, and it is this variety, this seeking of new
opportunities and overcoming of new difficulties by the
feathered sons, which makes the study of birds so fascinat-
ing a pursuit. Let us follow the diverging paths of the later gen-
erations of some of our own birds. Take the wood-
warblers of our own country. The only way we can
imagine what the earlier ancestors of the warblers were
like is to make a composite of the whole Family. All
its members are tiny, delicate birds which feed on the
smallest insects, their bills are slender and pointed, and
their feet and toes like the finest wire. Yet, far from
waiting for Nature to alter these delicate organs, they
have struck out boldly for themselves and, to avoid a
fatal competition with one another, have varied their
methods of hunting and the limits of their preserves so
successfully that a dozen may live in close proximity
and yet never poach on each other's domains. and yet never poach on each other's domains. FEET AND LEGS Our well-known little Maryland or Northern Yellow-
throat has chosen the low bushes of a marsh as his sphere
in life, and, although he has hidden his face behind a black
mask, yet he is a true warbler, and the blood of his fathers
forces him up now and then into some exposed position, Feet and Le 363 where he bursts into a joyous bubbling and warbling,
calling to his brethren of the tree-tops that, though his
haunts are changed, his heart is true to the clan. His
cousin, the Worm-eating Warbler, is tending in his direc-
tion, living in low bushes and in his habits drifting ever
marshward, where there may not be sufficient competition
to prevent his eventually sharing it with his more original
kinsman. The Yellow Palm Warblers, although more
conventional in their ordinary tree-top haunts, have de-
parted from ancient customs in their feeding habits. They dine on the ground, then fly back to the trees; ob-
serving, like some humans, the traditions of their family
in the spirit, if not in the letter. The brilliant Redstart clings even more closely to the
ancestral ideas of high trees, and cares little what kinds
he may find himself in; but he has a failing for water,
and if he may not descend, as have his two cousins men-
tioned above, yet he overlooks them and often swings
low through the air toward them. For in his feeding
habits he is one of the most radical of warblers. Has
he not seen the little green flycatchers in the woods, sit-
ting so lazily upon some favourite perch, and with an occa-
sional swoop snapping up an unfortunate insect? Why,
indeed, search all day for the tiny mouthfuls? Why not
wait for them to appear? So Redstart attempts fly-
catching and with perfect success. But the active blood which surges through his veins
will not allow him to assume the patient waiting tactics
of the genuine flycatchers. He may imitate their meth-
ods of actual capture, bagging his game on the wing, but The Bird 3 he is still ever on the move, from twig to twig, from tree
to tree. FEET AND LEGS Nevertheless, he has gained an advantage which
ensures to his race a long life; for in a tree whose foliage
and twiglets are being scanned with the microscopic glances
of his relatives, he gets more than the others by watch-
ing for the many insects which are alarmed at the dis-
turbance of the tiny hunters, and which flutter out in
the bright sunshine only to flutter straight down his
throat. Mother Nature has seen his efforts in the new
field with satisfaction, and has given to him a little re-
ward; for from either side of his mouth several stiff bristles
project, and many times, when he has misjudged the dis-
tance or the dodging powers of his prey, these little hairs
shunt the gnat or fly into his mouth. The Myrtle Warbler is an expert catcher of flies, and
has in addition another string to his bow, which bids fair
to place him at the head of the list of new departures
in warblers. He has learned that bay berries are not only
an occasional welcome variety to the everlasting diet of
insects, but that a warbler can comfortably live upon
them when the cold has benumbed the little winged and
crawling creatures. So, instead of migrating south at
the first hint of winter, these hardy little Myrtle Warblers
sometimes remain with us throughout the whole season of
cold and snow. A most daring departure from old-established prin-
ciples of the warbler clans is that of the Water Thrushes. Ages ago, perhaps, we may imagine that some member
of this group, while drinking at a stream or pond, watched
the little bobbing sandpipers as they scurried past along Feet and Le 365 the brim, now wading in a short distance, then leaping
to a soft rim of clay, everywhere finding the most delicious
morsels abundant. A strange fascination took hold of the
tree-haunting warbler, and although perhaps you and
I would have said he was a very silly bird and that such
a thing as a warbler turning into a sandpiper was utterly
absurd, yet the little fellow and his descendants persisted. Sandpipers and sandpipers only they wished to be, and
Nature has given them their wish. Study the Water Thrushes of to-day. Their whole
life is spent along some stream or pond, searching for
worms and snails in true sandpiper fashion. FEET AND LEGS Not only
this, but even the dipping gait of the pipers has been
copied, and though we cannot give a reason for this char-
acteristic, yet the warblers have learned it by heart,
and many an amateur bird-lover do they confuse! But
the heart of the old clan instinct can never be entirely
eliminated, and even if a warbler should attempt to hum
away his life on the wing like a hummingbird, or to run
with the speed of the wind through dry deserts like an
ostrich, yet, like the Water Thrushes, he would occasion-
ally drift back to the old tree-tops and there sing of the
happiness which is within his heart. A 'strange whim of evolution in one member of the
warbler tribe results in his mimicking the sandpiper as far as
terrestrial locomotion, a walking gait, and the peculiar tilt-
ing habit go, but the fondness for water did not accompany
these changes, and so we find the Oven-bird content with
the deep woods where he builds his home upon the ground. He often returns for a time to the trees, but, like a college The Bird 3 boy whose whole ideas of life have been changed by ab-
sence from his rural home, the Oven-bird carries aloft
with him the mincing gait of the littoral sandpipers, walk-
ing sedately along the limbs among his agile, hopping,
creeping cousins. creeping cousins. Of the conventional aristocracy of the warblers there
would be much to say had we the space. The Black-
and-white Creeping Warbler has been transformed into a
woodpecker, as far as mode of progression goes; and
lucky for him too, for he never fails to find cocoons
and small edible things among the cracks and crevices
of the bark, no matter how vainly the others may be
searching the overworked twigs and leaves. And Nature
has helped him, too. She has dipped him in a bath of
the essence of these very same crevices and cracks, and
out he has come, covered with the semblance of the rough
surface and the long, dark shadows which may shield
and hide him from many enemies. Of the typical tree-loving species, the Pine Warbler
haunts the growths which have given him his name; the
Black-throated Green also loves the evergreens, and the
beautiful Magnolia delights in thick forests of spruces. FEET AND LEGS Thus we have taken a brief survey of the recent branch-
ing of the warbler's genealogical tree. Each has found
a niche in which to live, and the food and safety which
permit him to rear a nestful of young each year. So far
so good, but we must not forget to give a thought to the
untold thousands and tens of thousands of generations
which have failed in their attempts. Nature has removed
all traces from view and in the general advancement of Feet and Le 367 the race as a whole they are forgotten, but it is well for
us to think of them occasionally: their birth, the chance
which came, which seemed so full of promise, which they
so eagerly accepted and which betrayed them; the myriad
little dead forms which gave up their lives in ages past,
and upon whose bodies and whose efforts the birds of
to-day have risen to their present high place in the scale
of the creatures of the world. We might have used this same illustration, or many
others like it, in connection with almost any other portion
of the bird's body. Although, indeed, it pertains more
strictly to the mental characters, and so is in a way out-
side the province of this volume, yet its application to
physical adaptations is so evident that its omission would
leave incomplete a most interesting phase of the possi-
bilities of the adaptation of bird structure. Although among perching birds the bill is the important
organ for procuring food, yet such birds as the Chewink,
the White-throated Sparrow, and the jays, in search of
small insects use their feet to scratch away dead leaves
and rubbish, kicking backward with both feet at once. There are many curious things about toes to which
we have not yet found the key. Who can tell why the
Horned Lark, Pipit, and some other birds have such
elongated claws on their rear toes? Perhaps the fact
that these birds live almost entirely on the ground may
have something to do with this peculiarity. Any one who
has kept a cage full of small birds will soon have learned
the fact that the claws of birds are continually growing. FEET AND LEGS In a remarkably short time their claws become long and The Bird 8 368
curved, and in a neglected aviary I have seen birds which
were prisoners on their perch, unable to untwist their
claws from it. When wild, birds wear down these struc-
tures by constant rubbing, and if given plenty of rough
bark and wood in their cages, their claws will remain of
usual length. or that portion of the foot which,
is, in almost all birds, covered with
horny scales, yet these vary con-
siderably in different groups. la
many the scales are small, six-
sided or oblong, as in plovers. In some of the higher song-birds
these scales have become joined
together until, as in our robin,
the front of the leg is covered
with a long "boot" of horn. The cause of this coalescence yet
remains to be discovered. Although the tarsus, or that portion of the foot which,
we usually call the leg, is, in almost all birds, covered with
horny scales, yet these vary con-
siderably in different groups. la
many the scales are small, six- Although the tarsus, or that portion of the foot which,
we usually call the leg, is, in almost all birds, covered with
horny scales, yet these vary con-
siderably in different groups. la
many the scales are small, six-
sided or oblong, as in plovers. In some of the higher song-birds
these scales have become joined
together until, as in our robin,
the front of the leg is covered
with a long "boot" of horn. The cause of this coalescence yet
remains to be discovered. FIG. 289. — Foot of American FIG. 289. — Foot of American
Pipit. FIG. 289. — Foot of American
Pipit. A considerable degeneration of
the legs and feet is found among goatsuckers, humming-
birds, and chimney swifts; but, small as are the feet of
the latter birds, they make frequent use of them to break
off the short twigs which are used in the construction
of their nests. One may take a young swift and place it
against the vertical surface of an ordinary brick (Fig. 326),
and the bird will hold fast without slipping a fraction,
of an inch. The slender nails fasten in the slightest
irregularity of the surface and hold the bird safely. FIG. 289. — Foot of American
Pipit. FEET AND LEGS Some species of swifts have all four toes pointing Feet and Legs 369 forward, forming a four-tined grapple by which they
hang themselves up in their hollow nesting-trees. Whip-
poorwills and some other birds have a curious comb,
or pecten, along the edge of the middle claw, which is
perhaps of use in cleaning the long bristles about their
mouths, or in arranging their very delicate, soft plumage. Kingfishers and several related groups of birds make so
little use of their feet, except in motionless perching, that FIG. 290. — Comb on toe of Chuck-wilPs-widow. FIG. 290. — Comb on toe of Chuck-wilPs-widow. not only are the toes small and weak, but two of the
front ones have grown together for over half their length. Perhaps the most interesting condition of toe struc-
ture is found among the woodpeckers, parrots, cuckoos,
and owls. In these groups we find a similar plan of gen-
eral arrangement: two toes in front and two behind. With few exceptions it is the great, or first, toe and the
fourth, or outer, toe which are reversed. This arrange-
ment of toes is known as yoke-toed, or zygodactyl. The Bird 0 We have seen that in perching birds the arrangement
is three toes in front and one behind; and now turning
to the woodpeckers we are struck with the excellent
toe arrangement of these climbing birds,— their claws
spreading so that they point almost to the four points
of the compass, thus forming an admirable grapple or
vise, which makes a vertical position as safe for a wood-
pecker as a horizontal one for a percher. pecker as a horizontal one for a percher. Woodpeckers, the world over, have feet and toes
which are remarkably alike; but in Canada and the most
northern parts of our own country, and in certain por-
tions of the Old World, there are several woodpeckers
which are unique among the birds of this Order in pos-
sessing but three toes. For some unknown reason their
first, or great, toe, which in all other woodpeckers points
backward, has disappeared, leaving but a vestigial trace
beneath the skin, while the outer toe is reversed to take
its place. We may see one of these hardy three-toed fel-
lows sliding and hitching up a pine-tree, pounding and
hammering vigorously, the loss of an entire toe evidently not
handicapping him in the least. In such fashion does Nature
occasionally upset our hard-worked-out theories, leaving
us confused and baffled before her inexplicable surprises. us confused and baffled before her inexplicable surprises. Is it not rather disconcerting to find that this same
arrangement of two toes in front and two behind also
holds good for the other Orders of birds mentioned above,
the parrots, cuckoos, and owls, — their toes all arranged
in pairs, fore-and-aft? This is an excellent example of
what is called parallelism, or the independent develop-
ment of similar structures. Feet and Le 371 Parrots use their feet for more different purposes than
do any other birds : they are the monkeys of the feathered IG. 291. — Cockatoo perching with one foot and holding food with the other FIG. 291. — Cockatoo perching with one foot and holding food with the other. world. They climb wires or branches one step after the
other, their beaks taking the place of a third foot in this
style of locomotion. They pick up food, such as a banana The Bird 3 or a nut, and, holding it in the foot while eating, turn
it from side to side as we revolve an apple in our hand. With their claws they preen their plumage, and push each
other aside when too closely crowded. In fact the functions
of the feet and toes of parrots approach nearer to those
of a human hand than the limb of any other Order of birds. PIG. 292. — Foot of Cuckoo, perching, and with toes outstretched. PIG. 292. — Foot of Cuckoo, perching, and with toes outstretched. Cuckoos are perching birds, and when we see the skil-
ful way in which they creep through a dense thicket, never
missing their hold, we wonder why all perching birds do
not have this arrangement of two toes in front and two
behind. So completely are our theories set at naught
that we should hardly be surprised to see a bird with
one toe in front and three behind cheerfully hopping
from branch to branch! In the deserts of the south- Feet and Le 373 western part of the United States much of the vegetation
consists of prickly cacti and thorny mesquite, most un-
pleasant to perch upon, and here we find the Road-runner,
a kind of ground cuckoo, who has the fore-and-aft toe
arrangement of his arboreal relations, but whose terres-
trial life has developed remarkable powers of running
and leaping. One of these birds can outstrip a horse
for a hundred yards or more and, almost without effort,
can leap upward ten or twelve feet, to all appearances
unaided by its wings. The owls can move their outer toes backward or for-
ward at will, thus being able to assume the arrangement
of toes both of a crow and of a parrot. However the
yoke, or two-and-two, plan is the one most commonly seen
among these birds. Feet and Le With such an automatic vise-trap
ready to descend silently and with deadly swiftness upon
him, the little mouse in the grass has indeed need to be
ever on the alert. The talons of owls are curved and
under the control of tendons of great strength. Their
chief use is to capture living prey and then to hold it firmly
while it is torn to pieces by the beak. The deserts and plains where the Road-runner dwells
are also the home of the Burrowing Owl, Fig. 351, which
finds in its sharp little talons admirable picks and shovels,
certainly a novel use for yoked toes. The feet and toes
of birds are, in zero weather, their most vulnerable points
(except their eyes), and they are most liable to be frozen. In the black wastes of the frozen boreal regions, the Arctic
Owl is able to defy the intense cold, by means of a furry
covering of hair-like feathers, which extends to the very The Bird 374 claws, and even the soles of the feet are thickly covered ,
so that the skin of the bird is never in contact with the
snow and ice on which it roosts. The osprey, or fish-hawk, can, like the owls, reverse
its outer toe, but all typical hawks and eagles have the FIG. 293. — Owl gripping a piece of meat. FIG. 293. — Owl gripping a piece of meat. perching-bird arrangement. The talons of the osprey
are immensely strong, and the scales on the soles of its
feet and toes are hardened and roughened to such a degree
that they are almost spike-like. A more efficient fish-
trap cannot be imagined. The Golden Eagle has a splendid
foot, with great curved talons, which, when they have perching-bird arrangement. The talons of the osprey
are immensely strong, and the scales on the soles of its
feet and toes are hardened and roughened to such a degree
that they are almost spike-like. A more efficient fish-
trap cannot be imagined. The Golden Eagle has a splendid
foot, with great curved talons, which, when they have Feet and Le 375 once clasped an object, never let go. It required two
men and two pairs of the thickest buckskin gloves to
obtain Fig. 295, and even then the foot could be held
still for only a moment. As the photograph shows, the FIG. 294.— Foot of Snowy Owl. Feet and Le FIG. 294.— Foot of Snowy Owl. leg is feathered all the way down to the toes in this eagle,
for some unexplained reason, while in almost all its rela-
tives, as in the Bald Eagle, the legs are covered with
scales. The feet and toes of the Harpy Eagle, Fig. 204,
are probably the most terrible of their kind in the world : The Bird 76 376
certainly they are the strongest. When once they have
closed on an object, and remain clutched, nothing short
of severing the bird's leg will avail to loosen the fearful FIG. 295.— Foot of Golden Eagle. FIG. 295.— Foot of Golden Eagle. FIG. 295.— Foot of Golden Eagle. grip. Besides capturing their food, birds of prey carry
the sticks for their nests in their talons. grip. Besides capturing their food, birds of prey carry
the sticks for their nests in their talons. When, instead of killing its prey, a species of bird
feeds upon carrion, the change in its habits is reflected Feet and Legs 377 clearly in the appearance of its feet. Compare the feet
of a vulture (Fig. 296) with those of one of the true birds
of prey (Fig. 295). The muscles are weaker and the claws
are shorter, more blunt, and, as a result, the toes have lost
their clasping power, while the hind toe is higher and so FIG. 296. — Feet of Vulture. (E. R. Sanborn, photographer.) FIG. 296. — Feet of Vulture. (E. R. Sanborn, photographer.) small that it is of no use even in perching. Such is the
condition in the condor of South America. When in captivity an eagle is given a piece of meat, it
seizes the food in its talons and flies to some favourite
spot to devour it, but a condor transports its meal in The Bird 3 its beak, then holding it down firmly with one of its feet,
it pulls upward and so tears the meat. it pulls upward and so tears the meat. So exactly correlated are these changes of habit and
of feet that in the Caracara, a Mexican bird of mixed
habits, partly rapacious and partly vulturine, the toes and
claws are correspondingly midway between the two groups
of birds. Feet and Le This bird lacks sufficient grasping power to
enable it to lift its prey from the ground after the manner
of a true Hawk; but it will overcome this difficulty by
carrying up the object in its beak, and then reaching for-
ward with its feet, while in full flight, and taking a careful
grip with its talons. In South Africa is a bird known as the Secretary,
which is really a terrestrial hawk, rarely flying, but spend-
ing most of its time stalking about in search of food. Any
one who has seen an eagle progressing upon the ground
by means of its awkward ga41op, can realize the impos-
sibility of such a short-legged bird preferring terrestrial
life, but the legs of the Secretary are as long as those of
a crane, although in other respects the bird would pass
for a very long-tailed species of hawk; it is really a hawk
on stilts. However, there are reasons for supposing that
the Secretary Bird may be, not a more or less recent off-
shoot from the hawks, but a surviving type of old, old
days when there were no hawks and cranes and herons,
but instead, a few strange birds which combined the
characteristics of all these groups. of all these groups. The skilful way in which the Secretary Bird brings its
feet into play in the capture of serpents, of which it is
very fond, has been described as follows: Feet and Legs Feet and Legs 379 "When the snake strikes, the bird either evades the
blow, by skipping to one side or the other, jumping back-
ward, or springing into the air, or else, as frequently hap-
pens, he simply receives the venomous thrusts of his
antagonist on the broad stiff feathers of the outer half \
FIG. 297. — Secretary Bird. FIG. 297. — Secretary Bird. of the long wing, with which he knocks the reptile down,
following up the fall with a vigorous kick. His extreme
agility enables him in a very short time to baffle and
overcome a snake of four or five feet in length, whereupon
he finally seizes it near the head with his bill, and hold- The Bird o 38o
ing the body down with one foot, proceeds to swallow it. In case a snake proves unusually hard to manage on
the ground the dauntless bird watches his opportunity,
seizes his adversary close to the head, and, flying aloft
to a considerable height, lets it drop on the hard ground,
which is usually sufficient to prepare it for the final
ceremony of swallowing." eremony
FIG. 298. — Feet of Ruffed Grouse, showing snowshoes of horn. FIG. 298. — Feet of Ruffed Grouse, showing snowshoes of horn. Quail, grouse, pheasants, turkeys, and all the fowl-
like birds are scratchers, according to the old classification,
and they well deserve the name; for scratching first with
one foot and then the other among the leaves and soft
dirt for insects is a very pronounced habit of them all. The arrangement of toes is the same as in the perching
birds, but the claws are very different. These birds are Feet and Le 381 true horny-handed sons of the soil : their claws are stubby,
short, and blunt. Sharp edges would soon be dulled by
scratching, and elongated ones would sliver and break. So, with his blunt claws, our chicken and his kind are
well provided for. The most interesting feet among these birds are those
of the grouse. The ruffed drummer of our woods walks
about, in summer, on slender toes over moss and logs, but,
when soft deep snows come, his weight would make it
difficult to keep from being buried at each step. So
Nature provides him with snowshoes. Feet and Legs From each side of
each toe a broad, horny comb-like fringe grows out; not
a web of skin which might soon freeze, but rows of horny
projections, as of a myriad extra claws. This distributes
his weight so that he trots merrily over snow through which
a fox sinks deep and flounders awkwardly at every step. But what of the ptarmigan, that snow-white grouse
of the far North, whose home is amid those frigid barren
regions? This bird is much more of a walker than the
Snowy Owl, and its feet would surely freeze during
the long winters if they were bare of feathers. So we
find indeed that scarcely a claw is visible beyond the
thick feathers which cover legs, toes, and soles. Such
a provision against cold is evident and reasonable enough,
but how are we to account for the feet and toes of the
House Martin of Europe, which are densely feathered to
the very claws? It breeds in Iceland and Lapland, but
only in summer, when it would need no such protection
against cold, and it is also true that it breeds upon the
cliffs of Persia and southern India. The Bird 2 382
If we watch a duck as it settles itself for the night
upon the snow, we will see it squat down, snuggle its
beak deep among the feathers of the back, and finally
draw up each foot from the frozen surface and tuck them
up out of sight. Thus they are protected from freezing
during the long, cold night. The pugnacity of the males of the Order of game-
birds has become proverbial; almost all are "fighting
cocks" and yet their beaks are not fitted for defence or FIG. 299. — Mallard asleep on the snow, with its feet drawn up to avoid freezi FIG. 299. — Mallard asleep on the snow, with its feet drawn up to avoid freezing. offence, nor can they clutch and tear with their claws. But we find spurs developed on the tarsus, or upper
foot, in fowls, turkeys, pheasants, and peacocks, which
are used with remarkable skill in their battles. In
structure these outgrowths are identical with the horns
of antelopes and cows, consisting of a bony projection
over which grows a sheath of horn. The spurs of the
peacock are long and sharp and are occasionally used
with such effect that the results are fatal to each of the
contestants. A diminutive relative of Pavo, the Pea- Feet and Le 383 cock Pheasant of the East Indies, has two, three, or even
four spurs of full size on the legs. The bird photographed
on page 419 had two on the right leg and three on the
left. Yet these birds are not as correspondingly pug-
nacious as we should imagine from their increased arma-
ture. There is a small group of peculiar birds, known as
Sand-grouse, which in many respects stand midway FIG. 300. — Spur of Java Peacock. FIG. 300. — Spur of Java Peacock. between the true grouse and the pigeons. In certain of
these the toes, to their very tips, are encased in the skin
of the foot, the effect being of a mitten with only the
claws free. The reason for this is yet to be found. We now come to the water-loving birds, and we find
that their varying associations with this element have
wrought many interesting changes in their feet and legs. The Bird Those birds which are content to wade along the shallow
margins of ponds and streams require long legs and long The Bird 3 toes, the latter to distribute their weight as they walk
over the soft muddy bottom, the former to lift their bodies
above the surface of the water. Such, broadly speaking,
are the plovers and sandpipers and herons. Let us see
how the feet of these birds reflect theh1 habits. With
the exception of the tribe of plovers, almost all have four
toes. The plovers have but three, and these are slender
and not webbed, for although they usually feed on aquatic
forms of life, yet their food is gleaned from the upper
part of beaches, or from the sand-flats when the tide is
out, and they therefore seldom have occasion to swim. The sandpipers venture into the shallows and are some-
times lifted from their feet by a small inrushing wave. But the majority even of these go through life unwebbed. One, the Semipalmated Sandpiper, shows a beginning of
this in the half-webbed condition of the toes, but the
group of phalaropes are actually sandpipers of the sea. I have seen them in flocks of thousands, resting upon
the surface of the ocean, scores of miles from land. Yet
when ashore they have need to be as active as other mem-
bers of their Order in order to find sufficient food; so, in-
stead of being hampered with a confining web, each toe
has a series of broad scalloped lobes, serving admirably
as water propellers, yet allowing the toes freedom of motion
when the owner is scurrying over the sand. I have observed Great Blue Herons almost hip-deep I have observed Great Blue Herons almost hip-deep
in the breakers along the Florida beaches, yet this is not
a usual haunt for members of this group of birds. They
usually prefer quiet inland waters, where they wade and
watch — ever striving to satisfy their insatiable hunger. Feet and Le 385 So, in the case of herons, webs would be superfluous,
length of limb being their only requirement. length of limb being their only requirement. The Wood Ibises (or more properly Storks), which
are more active searchers after food than the herons,
make use of their toes to stir up the bottom mud of shal- FIG. 301.— Toes of Gallinule outstretched. FIG. 301.— Toes of Gallinule outstretched. The Bird low water, keeping the bill ready to snap up any small
creatures thus disturbed. When one sees a flock of gal-
linules or jacanas feeding quietly in their haunts they
appear to be walking on the water, and we find an in-
teresting connection between the structure of their feet
and toes and certain tropical plants. Such are the The Bird 6 386
great pads of water-lilies, which in places cover miles of
water, over whose trembling surfaces the birds are able
to run or walk. To enable them to do this without sink-
ing! both the toes and claws are remarkably long and
slender, so that in a bird which stands but ten or eleven
inches in height the weight is distributed over an area
of some fifty square inches. This makes it possible for
them to feed in places too deep for wading birds and too FIG. 302.- Gallftmle holding food in its foot. FIG. 302.- Gallftmle holding food in its foot. tangled with aquatic vegetation for swimmers readily to
make their way. This is but another forceful example of
the successful adaptive radiation of birds. Gallinules have found that their long toes can be made
useful in other ways besides locomotion, and we find that
they are well-nigh as skilful as a parrot in grasping and
holding. One of these birds perhaps spies a tuft of
water-soaked reeds. He clasps it firmly, draws it up,
and, holding it in the air near his bill, picks the small
worms and snails from among the stems, finally discarding Feet and Legs it for another footful. forming such an action. are so long and slender,
they can swim quite rapi
their feet with such effo
concert. Their cousins,
the coots, resemble the
phalaropes in having
broad lobes of skin
along each toe, so that,
although they and the
gallinules are often
seen feeding in the same
locality, yet the nata-
tory ability of the coot
allows it to venture
beyond the reserves of
the other species. The
toe-lobes also serve an-
other important func-
tion in permitting the
coots to feed upon soft
mud, thus keeping them
from sinking below the
surface, just as the horn
ports it on the snow. Herons are furnished it for another footful. We cannot imagine a
forming such an action. The Bird Although the toes of
are so long and slender, yet, when the necess
they can swim quite rapidly for a short distanc
their feet with such effort that the whole bod
concert. Their cousins,
the coots, resemble the
phalaropes in having
broad lobes of skin
along each toe, so that,
although they and the
gallinules are often
seen feeding in the same
locality, yet the nata-
tory ability of the coot
allows it to venture
beyond the reserves of
the other species. The
toe-lobes also serve an-
other important func-
tion in permitting the
coots to feed upon soft
FIG. 303.— Foot of
mud, thus keeping them
from sinking below the
surface, just as the horny "snowshoe" of the
ports it on the snow. Herons are furnished with a comb-like edge FIG. 303.— Foot of Coot. FIG. 303.— Foot of Coot. FIG. 303.— Foot of Coot. surface, just as the horny "snowshoe" of the grouse sup-
ports it on the snow. Herons are furnished with a comb-like edge to one of
the claws, similar to that on the claw of the whippoor-
will, but as yet we have no clue to its use. Although
differing so greatly from hawks in their method of feeding, The Bird 388 388
yet, when put upon the defensive, herons resort to much
the same tactics as do the birds of prey. When cornered
or wounded, so that escape by flight is impossible, the
bird throws itself upon its back and, with uplifted claws
and levelled beak, awaits the attack of its assailant. The
talons and mandibles of a hawk offer ten sharp points
which can all do severe damage; but the heron depends
only on the grasping power of its toes to hold fast, while
it strikes savage, spear-like blows with its beak. it strikes savage, spear-like blows with its beak. On the borderland of the fully webbed aquatic birds FIG. 304. — Comb on the toe of Heron. FIG. 304. — Comb on the toe of Heron. we find the flamingo, combining characters of the herons
and ducks. Its haunts are the exposed coral-flats of
tropical keys, where at any time a high tide or a
severe storm may sweep all, old and young, from their
feet. Then it is lucky indeed that the youngsters have
webs between their toes in addition to their long legs. The Bird It is a case of swim or be drowned. In the great Orders of sea-birds, and in the ducks and
their allies, the three front toes are joined together by Feet and Legs 389 389
a web of skin which, when swimming, offers a large area
of resistance to the water when the foot is pushed back-
ward. The chick in the egg has a shadow-membrane
of his fish-like ancestors between his toes, and in these
water-birds the web of skin continues throughout life. In
the terns or sea-swallows, which swim much less than
they fly, the web is excised, or scalloped out deeply, a
return to an almost semipalmated condition. FIG. 305. — Rough-legged Hawk in position of defence. FIG. 305. — Rough-legged Hawk in position of defence. A duck or swan out of sheer laziness will often hold
one foot up out of the water and propel itself with the
other, slightly altering the angle at which the web meets
the water, so as to maintain a perfectly direct course. There is a little-known habit which I have frequently
observed in captive ducks and several times in wild ones,
of swimming thus with one foot when both eyes are shut
and the bird is apparently fast asleep. But, in such a The Bird 9° 39°
case, no attempt is made to proceed in a straight line. In a pool only thirty feet square I have seen a duck
revolving thus for an hour or more at a time, impelled
with slow, rhythmical (and apparently reflex) strokes. We FIG. 306.— Foot of Black-necked Swan. FIG. 306.— Foot of Black-necked Swan. can imagine that such a habit would sometimes be of much
advantage to a wild bird, enabling it to keep away from
enemies on the shore and yet at the same time secure rest. The name Steganopodes is applied to the gannets,
pelicans, snake-birds, tropic-birds, and cormorants, be- Feet and Le 391 cause the toes of these birds are all bound together with
a single web. The hind toe points almost in a forward
direction when the foot is in action, and, to complete
the adaptation for a perfect swimming foot, the outer
toe is the longest, a rare condition among birds. The Bird If one
will watch the snake-birds in a zoological park, as they
swim about their glass tank, the extreme delicacy of the
foot mechanism becomes apparent at once. Not only is the flat side of the leg used as a cutwater,
but the toes curl and uncurl with a slight oblique revolving
motion like the blades of a propeller. When drawn for-
ward through the water they are rolled up into a very
small compass and then instantly spread out as widely
as possible on the return stroke. To the eye it seems
as if the bird was constantly grasping something tangible
in the water and thrusting it behind. This propeller motion may be observed even better
in a captive grebe. If the bird's head is placed in a glass
of water, its feet will move back and forth in the air with
all the motion of swimming. The adaptation for swimming
in these birds is so fundamental and thorough that even
the claws are broadened and flattened until they resemble
finger-nails. On land, grebes are absurdly awkward,
although they can walk upright even up a slight incline. But they are powerless to rise from the ground, even
with the aid of the wind,— needing the greater speed which
a swimming take-off from the water will give them. a swimming take-off from the water will give them. The most aquatic of all birds, the penguins, make
much more use of their wings than of their feet in swim-
ming and diving. The toes are webbed, however, and The Bird 92 392
are doubtless of considerable use when the bird is emerg-
ing from the water, which it generally does with a sudden
spurt of speed and a strong leap which lands it on its feet. In landbirds which have either lost or are losing the
power of flight there is often an interesting correlation
to be observed between the lapsing of this mode of loco- FIG. 307.— Feet of Penguin. FIG. 307.— Feet of Penguin. motion and an increased use and consequent greater de-
velopment of the legs and feet. The Bird Of a South African bird,
about the size of our American Robin, known as the Rock-
jumper, it is said: "These curious birds are only to be
found on the rock-strewn slopes and summits of mountain-
ranges where they are able to hop from rock to rock for a Feet and Le 393 distance without having to cross level or open ground;
... at the slightest alarm they either drop into a crevice
or bound from rock to rock with extraordinary speed, look-
ing more like india-rubber balls than birds, for there is no
perceptible interval between the end of one leap and the
beginning of the next, and the distance they can clear at a
single hop must be seen to be believed. Should they have
to cross a piece of level ground between two rocks which
they cannot clear with a single bound, they run across it with
great speed and usually with outspread wings. So feeble
are their powers of flight that they seldom attempt to
fly, and never when in a hurry or alarmed; at the most
they flutter feebly for a few hundred yards down hill. I have occasionally amused myself by trying to drive
these birds across a piece of open ground, but I have never
succeeded in getting them to quit the shelter of the rocks,
where they easily avoid one by leaping over the stones
or hiding in the crevices. In spite of his loose, fluffy
plumage, which blows about in the slightest breeze and
gives him a rather untidy appearance, the cock is an ex-
ceedingly handsome bird." So we have here an isolated
case of direct relation between two organs, the balance
of power changing from wing to feet and affecting much
of the bird's structure, even the plumage losing its cohe-
siveness. The weak-flying Tinamou have unusually sturdy
legs, and many other instances might be mentioned. any reasons the most interesting of all birds' legs, and many other instances might be mentioned. For many reasons the most interesting of all birds'
feet are those of the ostriches and their allies, and among
them the most extreme examples of this same cause and
effect are to be found. The Bird The Bird 394 When one trains in college for a long-distance race,
one rule to observe is, never touch your heels to the ground ;
run wholly on the ball of the foot. Untold centuries
ago, wise old Nature whispered the very same direction
to those of her children who had most need to run for
their lives in life's great race, and down through the
ages some of them have never broken training. When
an animal acquires
great speed in running
or leaping, there is a
tendency for one toe
to become greatly en-
larged at the expense
of the others, as is
seen in the case of the
horse, the kangaroo,
and the ostrich. In the horse only When one trains in college for a long-distance race,
one rule to observe is, never touch your heels to the ground ;
run wholly on the ball of the foot. Untold centuries
ago, wise old Nature whispered the very same direction
to those of her children who had most need to run for
their lives in life's great race, and down through the
ages some of them have never broken training. When FIG. 308. — Feet and legs of Cassowary. In the horse only
the middle toe is
functional, the second
and fourth having de-
generated into the
small splint-bones at
FIG. 308. — Feet and legs of Cassowary. the side of the leg. The kangaroo progresses upon
the fourth and fifth toes, the second and third being
small and skin-bound. The ostrich has but two toes,
one of which, the third, as in the case of the horse, is
very large and armed with a thick claw, which, hoof-
like, grows close to the toe. This toe supports most of
the bird's weight, while the fourth or outer toe is only FIG. 308. — Feet and legs of Cassowary. small splint-bones at
the side of the leg. The kangaroo progresses upon
the fourth and fifth toes, the second and third being
small and skin-bound. The ostrich has but two toes,
one of which, the third, as in the case of the horse, is
very large and armed with a thick claw, which, hoof-
like, grows close to the toe. This toe supports most of
the bird's weight, while the fourth or outer toe is only Feet and Legs
395
mm
FIG. 309. The Bird — (a) Front and (b) side view of foot of Ostrich. Feet and Legs Feet and Legs 395 FIG. 309. — (a) Front and (b) side view of foot of Ostrich. FIG. 310. — Feet of Donkey. FIG. 311. — Feet of young Kangaroo. FIG. 310. — Feet of Donkey. FIG. 311. — Feet of young Kangaroo. The Bird 9^ 39^
one quarter as large; and indeed it bids fair to disappear
altogether in the course of time, and even now the dimin-
utive nail which is often present is only as large as the
claw of a chicken. claw of a chicken. The power of the ostrich to defend itself by kicking
is proverbial, but the claw on the
large toe is blunt and the ability to
inflict injury lies in the terrible force
of the blow. Its ally, the cassowary,
has three good-sized toes, and on
the innermost one a specially adapt-
ed weapon in the shape of a strong,
pointed, talon-like claw, four inches
in length. The two photographs (Figs. 312
and 313) show how similar the
tracks which the modern cassowary
makes in walking over moist clay,
are to those made by the bipedal
reptilian Dinosaurs millions of years
ago, which have been found in the
Connecticut valley. Thus in our brief review we have
seen how the feet and legs of birds
serve them well in walking, hopping, running, perch-
ing, scratching, climbing, burrowing, swimming, diving,
in addition to the finding of their food, fighting,
FIG. 312.— Tracks of Casso-
wary in soft clay. The power of the ostrich to defend itself by kicking
is proverbial, but the claw on the
large toe is blunt and the ability to
inflict injury lies in the terrible force
of the blow. Its ally, the cassowary,
has three good-sized toes, and on The power of the ostrich to defend itself by kicking
is proverbial, but the claw on the
large toe is blunt and the ability to
inflict injury lies in the terrible force
of the blow. Its ally, the cassowary,
has three good-sized toes, and on
the innermost one a specially adapt-
ed weapon in the shape of a strong,
pointed, talon-like claw, four inches he ostrich to defend itself by kicking
is proverbial, but the claw on the
large toe is blunt and the ability to
inflict injury lies in the terrible force
of the blow. The Bird Its ally, the cassowary,
has three good-sized toes, and on
the innermost one a specially adapt-
ed weapon in the shape of a strong,
pointed, talon-like claw, four inches
in length. FIG. 312.— Tracks of Casso-
wary in soft clay. The two photographs (Figs. 312
and 313) show how similar the
tracks which the modern cassowary
makes in walking over moist clay,
are to those made by the bipedal
reptilian Dinosaurs millions of years
ago, which have been found in the
Connecticut valley. Thus in our brief review we have
seen how the feet and legs of birds
serve them well in walking, hopping, running, perch-
ing, scratching, climbing, burrowing, swimming, diving,
in addition to the finding of their food, fighting,
preening their feathers, and in countless other ways. The story of the bird's foot has not half been told,
FIG. 312.— Tracks of Casso-
wary in soft clay. FIG. 312.— Tracks of Casso-
wary in soft clay. Feet and Legs 397 but enough has been said to arouse our interest in
this member and to put us on the watch for new
facts. FIG. 313.— Fossil Dinosaur tracks, found at Middletown, New York. (Courtesy of Prof. R. S. Lull.) FIG. 313.— Fossil Dinosaur tracks, found at Middletown, New York. (Courtesy of Prof. R. S. Lull.) TAILS E have found that almost every organ of a bird's
body may be compared directly with the corre-
sponding structure in the body of a lizard or
of some reptile, and the tail is no exception: although
a lizard with a fan-shaped group of feathers sprouting
from the root of his tail would certainly be an anomaly;
and even if we substitute scales for the feathers, the result
would be ridiculous and unmeaning. But glance at the
photograph of the tail of our ancient, original-bird ac-
quaintance, the Archseopteryx, Fig. 315, which was taken
expressly for this purpose. E Take twenty feathers and arrange them as in Fig. 314 a,
representing the tail of Archa3opteryx ; then rearrange
them as in 314 6, corresponding to the tail of modern birds,
and the whole matter will be clear. Archaopteryx had
twenty bones in its tail, all separate, long and slender, and
arranged end to end, just as are the bones of a lizard's
tail to-day. But in the case of the bird of olden time
a pair of feathers grew out, one on each side of the tail-
bone, making forty tail-feathers in all. As we have seen,
this bird was rather weak-winged and probably more mo a) Arrangement of 20 feathers, as in Archceopteryx ; (6) t
of a Sparrow in place. FIG. 314. — (a) Arrangement of 20 feathers, as in Archceopteryx ; (6) tail-feathers
of a Sparrow in place. 399 399 The Bird 4 of a flutterer, or sealer, than a true flier, but as time went
on, and birds became more and more expert on the wing,
their wings grew stronger and their tails shorter and more
compact. We can readily see the reason for this, if we
imagine a ship which has been built with a rudder as long
as its whole deck. What an awkward thing such a rudder
would be! The waves would beat against it and great
force would be necessary to turn it and to steer the ship. As long as a bird was content to climb a tree with its
hands and feet, and then scale, like a flying squirrel, to
the base of the next, a lizard-like tail would be all-sufficient. TAILS So conspicuous and so unbirdlike was the long appendage
of the Archseopteryx that Saururce — lizard-tailed — has
been given as the name of the Sub-class which it occu-
pies all to itself. pies all to itself. When we look at the bones of the tail of a modern
bird, we find that many interesting changes have taken
place since the days of the lizard-tailed ancestors. Thus
in the common duck, for example, we find eight free
bones followed by a large upturned bone, which, from
its shape, is known as the ploughshare. It is this terminal
bone which supports all the tail-feathers of modern birds,
and in the duck it represents ten of the lizard-tail bones
all telescoped and fused into one. Some of the feathers
have been lost, as there are but sixteen in this bird's
tail. This loss of tail-feathers is of no value in classifica-
tion, as it may vary within narrow limits. For example,
one species of cormorant has seven pairs of tail-feathers,
while a closely related species has but six. Not only
this, but the variation may be merely sexual, as in the • FIG. 315. — Tail of Archceopteryx in British Museum. • FIG. 315. — Tail of Archceopteryx in British Museum. The Bird 4 peacock, which has ten pairs, while the peahen has one
pak less. pak less. The fusing together of these bones has resulted in the
drawing together of the feathers, so that, instead of the
long, unwieldy, paired affair, they are arranged in fan
shape, although still in pairs, and usually showing a slight
graduation reminiscent of the old-style tail. Some birds
have as few as four pairs of tail-feathers, while others FIG. 316.— Tail-hones of Ostrich. FIG. 316.— Tail-hones of Ostrich. have as many as twelve. In the abnormal domestic breed
of pigeons known as fantails, as many as forty tail-feathers
are sometimes found. The cassowary and the emeu
have none at all, while the ostrich seems to have an in-
definite number; the tails of these two unrelated groups
of birds seeming, like their wing-feathers, to have lost
uniformity from little use. Besides these true tail-feathers
there are others, usually smaller, which grow from above
and below the tail, being known as upper and under tail- have as many as twelve. TAILS In the abnormal domestic breed
of pigeons known as fantails, as many as forty tail-feathers
are sometimes found. The cassowary and the emeu
have none at all, while the ostrich seems to have an in-
definite number; the tails of these two unrelated groups
of birds seeming, like their wing-feathers, to have lost
uniformity from little use. Besides these true tail-feathers
there are others, usually smaller, which grow from above
and below the tail, being known as upper and under tail- Ta 403 coverts. Mention is here made of these because of the
important part they take in certain sham tails which
will soon be described. In the embryos of most birds of true flight the tail-
tip of the back-bone is represented by six or ten separate
pieces, which, before the chick hatches from the egg,
fuse into the ploughshare bone. In the ostrich-like birds FIG. 317. — Tail-bones of Bald Eagle, showing greater fusion and more
specialization than in Fig. 316. FIG. 317. — Tail-bones of Bald Eagle, showing greater fusion and more
specialization than in Fig. 316. these small bones never fuse, but remain separate through-
out life — a reptilian character persistent in these strange
birds (Fig. 3 16). The ploughshare bone is seen splendidly
developed in such a bird of strong flight as the Bald Eagle. Now that we have explained the origin of the tail,
let us consider what part it plays in the lives of the birds
about us. So diverse are the modes of life, and so varied
are the surroundings of this class of creatures, that we The Bird 404 FIG. 318. — Fan-tailed Pigeon, showing extreme development of tail. FIG. 318. — Fan-tailed Pigeon, showing extreme development of tail. Fio. 319. — Emeu, a tailless bird. (Courtesy of N. Y. Zoological Society.) Fio. 319. — Emeu, a tailless bird. (Courtesy of N. Y. Zoological Society.) Tails shall find many unexpected uses to which the tail is put,
and yet those which have been explained are a mere frac-
tion of the problems which still await solution. ch still await solution. TAILS The principal use of the tail-feathers in birds is, of
course, to perform the function of a rudder, and we find
that the arrangement of the bones perfectly carries out
the simile of a tail to the rudder of a ship; namely, a
broad, expanded surface which is closely hinged to the FIG. 320. FIG. 321. Tail of Barn Swallow, closed (320) and spread (321). 320. FIG. 321. Tail of Barn Swallow, closed (320) and spread (321). 320. FIG. 321. Tail of Barn Swallow, closed (320) and spread (321). body by several movable joints. The real tail of a bird
is the small, fleshy protuberance which in our roast
chicken we call the "pope's nose"; but in common par-
lance the word tail has come to be applied to the large
feathers which sprout from this structure. Thus, although
not comparable to the appendages of mammals, the so-
called tail of a bird is superficially more like the correspond-
ing organ of a whale than the tail-fin of a fish, since it is
expanded horizontally instead of vertically. The Bird 06 406
One interesting analogy to the fin of a fish is found
in the tail of the Blue Duck of New Zealand. This bird
lives in swift mountain streams and when swimming carries
its long tail entirely submerged. By vigorous sidewise
flicks of these tail-feathers it can turn around, as if on a
pivot, without being carried down-stream, even when
in the centre of a rapid, swirling current. PIG. 322. — Murre showing tail. (Compare with Fig. 246.) PIG. 322. — Murre showing tail. (Compare with Fig. 246.) It is interesting to compare this use of the tail-feathers
with the function of the tail in the flightless penguins. In the Black-footed species, at least, the tail-feathers are
stiff and short, but the bones of the tail are unusually
elongated and the flesh which covers them is flattened
into a kind of vertical rudder. Strong muscles control Ta 407 this, and by it the extremely quick dives and turns are
made possible. No feather would be stiff or rigid enough
to offer to the water the resistance which these feathered
seals require. Exceptions to the rudder use in flying birds are found
in the murres — sea-birds which share the cliffs of our north-
ern coast with cormorants and gulls. The tail-feathers
of a murre are so short as to be useless for steering pur-
poses, so in flight the bird uses its webbed feet instead,
stretching them out behind, opening, turning, and twist-
ing them in harmony with the wings, with as satisfactory
results as could be desired. The shape of the tip of the tail varies greatly in birds. It may be square or rounded, or cuneate, or indented
in the centre, or swallow-tailed, as we appropriately call
the latter deeply forked condition. These conditions may
be paralleled or duplicated in many different Families of
birds. For example, the forked type is seen in our com-
mon Barn Swallow, in those dainty relatives of the gulls,
the terns — " Swallows of the Sea," — and again in the
Forked-tailed Kite and the Scissor-tailed Flycatcher. By closely watching a swallow as it courses swiftly over
a meadow, or shoots upward, buoying itself against the
breeze, we can appreciate the delicate adjustment of the
muscles which govern the tail-feathers. The Bird Each feather
seems vital with life, now sliding one over the other until
all are in a narrow line, then expanding, with less friction
than ever a fan opened, into a wide-spreading, gently
graduated fork. The quartet of forked-tailed birds men-
tioned above are splendid fliers, but we shall see that skill The Bird 08 408
in flight depends but little upon the shape of the tip,
when we consider certain birds with cuneate tails, or
those in which the central feathers, soft and not rigid,
are elongated, instead of the outer ones. The Undulated Grass Parrakeet shows a condition
almost the opposite of the swallow. The Mexican long-
tailed jays, the magpies, and the tropic-birds are also
all of this type, the latter being especially fine fliers and
capable of remarkable aerial evolutions. Again, some FIG. 323.— Tail of Grass Parrakeet. FIG. 323.— Tail of Grass Parrakeet. of the flycatchers with moderate, rounded tails can exe-
cute most wonderful flight movements, steering in erratic
darts through the air, or darting aside at right angles
while at full speed, this being accomplished principally
by means of the tail. of the flycatchers with moderate, rounded tails can exe-
cute most wonderful flight movements, steering in erratic
darts through the air, or darting aside at right angles
while at full speed, this being accomplished principally
by means of the tail. A tail serves also an important use as a brake. When
a great pelican settles gradually toward the surface of
the water, or a duck momentarily hovers before alighting,
the tail, wide-spread and brought downward, gives effi-
cient aid in retarding the impetus. Tails 409 We notice that birds which have very short tails are
unable to turn quickly and that their flight is very direct,
or even where there is a long tail, if it is principally for
ornament and not well muscled, it is of little use in help-
ing its owner to change the direction of flight. The
partridge-like tinamous of South America are good exam-
ples of the first-mentioned group. Their tails are small
and useless, and when once the bird launches itself into FIG. 324. — Tail of Pelican alighting. FIG. 325.— Tail of Tern in flight. FIG. 324. — Tail of Pelican alighting. FIG. 325.— Tail of Tern in flight. the air, it can keep on only in a straight line and is at
the mercy of every cross-current of air. A more familiar
case, which any one may observe, is a Song Sparrow, or
other small bird, which, from accident or from some irregu-
larity of moult, has lost all or most of its tail-feathers. Instead of rising with the strong, darting flight with
which such a bird is accustomed to make its escape from
our path, its flight under such conditions is weak and
direct, like the trial efforts of a young bird. The Bird 41 o Reserving the mention of partly ornamental tails until
the last, we may now consider the use of this member as
a prop or support to the bird as it clings to or makes its
way up vertical surfaces. Four groups of birds which
are thus distinguished are the woodhewers — a tropical FIG. 326.— Chimney Swift clinging to wall, resting upon tail. FIG. 326.— Chimney Swift clinging to wall, resting upon tail. family, — the creepers, woodpeckers, and swifts. These
birds really sit upon their tails, the feathers of which are
adapted for this special use, while retaining perfectly the
rudder function in flight. The tail-feathers of the Chim-
ney Swift are peculiar in having the ends, for a short dis- Ta 411 tance, free of barbs, the tips being thus composed of a num-
ber of bare spines which are admirably adapted to catch
in the irregularities of hollow trees, or, as now in their
recently adopted homes, in the roughness of chimney-
bricks. I one day caught a Chimney Swift and placed
it against a varnished wall composed of composition
bricks; and, smooth though the surface was, the bird's
tail and toes held it firmly, not slipping even a quarter
of an inch. After photographing it, I watched it for
some minutes and saw the bird shift its position several
times, moving always with a certainty and surety of
grasp most inexplicable. The tails of woodpeckers and creepers are not thus
denuded at the tip, but they are stiffened throughout
and are very elastic (Fig. 240). When a woodpecker
brings up against a comparatively smooth tree-trunk, its
certainty of hold is a perfect bit of magic. Then when
it braces itself and sets to work to hammer a hole into
the wood, or to excavate its nest, how the tail-feathers
bend and spread, buttressing themselves against every
roughness, the elasticity of the feather-tips allowing them
to slip into every crevice! In many birds the tail is a perfect index of the emotions,
doing much to compensate for the lack of facial expression. Especially is this true of the wrens, those feathered bundles
of tireless energy and curiosity, whose tails, upturned so
high that they fairly tilt forward over the back, twitch
and jerk with every passing mood. The Bird Even the genetic
individuality of a species may be hinted at in the way
it carries its tail; quiet, soft-mannered birds holding it The Bird 412 412
low, beneath the wing-tips, while active, nervous species
carry it more or less raised. FIG. 327.— Tail of Chimney Swift. In certain of the flycatchers
the tail, which hangs demurely
downward, reacts with a jerk
to every note of the bird, as if
connected with the bird's vo-
cal apparatus, as in our com-
mon Least Flycatcher at every
" Che-bee'!" " Che-bee'!"
The jerking motion of the
FIG. 327.— Tail of Chimney Swift. tail seems to have become
a regular habit with many birds, and, curiously enough, FIG. 327.— Tail of Chimney Swift. FIG. 328.— Tail of Flicker. FIG. 328.— Tail of Flicker. especially with those which spend their lives chiefly along Ta 4 1 3 the borders of streams. We are all familiar with the tip-
ping of the tail in sandpipers, and, including the Green
Heron, we will see much the same motion in birds which
haunt the stream borders ; even in the Water Thrush the
same habit prevailing, although, as we saw in the pre-
ceding chapter, this bird is closely related to the bright-
coloured warblers of our tree-tops. The wagtails have
received their name from this same habit, of which no
explanation has yet been offered. The Road-runner, a ground cuckoo of the Western
plains, has a tail as long as its entire body, which is as
expressive as the gestures of a Frenchman. When sitting
quietly in the shade of a mesquite-bush in Mexico, I have
seen one of these birds dash into sight and drop, like an
arrow, upon a luckless lizard. At the moment of attack
all ten tail-feathers of the bird were wide-spread and
a-tremor, indicative of the extreme excitement attendant
upon the capture of the reptile. While eating what choice
parts were desired, the tail was folded and lifted out of
the way. Soon the bird spied some motion of mine, and
with the suspicion came the high extended neck, while the
tail turned up and forward, until almost touching the
bird's head. A second motion on my part, and the tail
manoeuvred to a line and trailed limply after the bird,
as it half-flew, half-leaped to a high rock and on out of
sight. sight. The Bird The white under sides of the tails of the wild rabbit
and the white-tailed deer have been explained as warning
signals to others of the family or herd: white guides
which the less experienced members may follow and so The Bird 414 escape from danger. Again, the theory has been advanced
that these white patches merge with the sky when the rab-
bit or deer makes the first high frantic leap to escape an
assailant, the white spots thus tending to confuse the
creature making the attack. We are, however, far from
certain whether any such interpretations can be applied
to those birds, such as the Junco, the Meadowlark, and
the Vesper Sparrow, which have the lateral feathers of escape from danger. Again, the theory has been advanced
that these white patches merge with the sky when the rab-
bit or deer makes the first high frantic leap to escape an
assailant, the white spots thus tending to confuse the
creature making the attack. We are, however, far from
certain whether any such interpretations can be applied
to those birds, such as the Junco, the Meadowlark, and
the Vesper Sparrow, which have the lateral feathers of
the tail white; but in these
cases the first theory seems
at least more probable, as
these birds live in flocks and
in a more or less open en-
vironment, where such a sig-
nal would have the greatest
chance for use. When a
Junco is upon the ground,
its black and gray plumage
renders it very inconspicu-
, FIG. 329.— Tail of Junco. ous> but the instant it takes
to wing, out flashes the white V in its tail. , FIG. 329.— Tail of Junco. FIG. 329.— Tail of Junc We have seen that not a portion of the external parts
of the bird has escaped, in one species or another, being
utilized for ornament; generally, as well as we can tell? as some decoration to attract or charm the female. Tails
bear even more than their share of adornment, which we
cannot pass by without mention, although, as dealing with
the psychological side of bird life, any discussion of this
question is outside the province of this volume. Some-
times it is only some slight addition to the feathers of Ta 415 the tail proper, as the elongated middle feathers of the
male Pintail Duck and the Sharp-tailed Grouse. The Bird male Pintail Duck and the Sharp-tailed Grouse. Turning to a few of the more decorative tails in the
world of birds, we find a small Australian bird, known
as the Emeu-wren, bearing aloft a half-dozen long feathers,
so scantily clothed with barbs as to resemble somewhat
the plumage of the Emeu itself. These skeleton plumes,
for they are little else, while giving a striking appearance FIG. 330.— Tail of Emeu-wren. (Cf. with Fig. 23.) FIG. 330.— Tail of Emeu-wren. (Cf. with Fig. 23.) to the owner, must radically weaken its flight, as regards
steering capacity; since the open-work mesh of the vanes
can offer no resistance to the air. Indeed it is said of
this bird that it is such a poor flier that it is seldom seen
on the wing, but it runs rapidly and is able to leap into
the lower branches of trees. The penalty of danger from
weakened flight which the Emeu-wren must pay for his
caudal decoration is paralleled by certain little whydah-
finches of Africa, the males of which at the breeding
season are decorated with several tail-feathers over five to the owner, must radically weaken its flight, as regards
steering capacity; since the open-work mesh of the vanes
can offer no resistance to the air. Indeed it is said of
this bird that it is such a poor flier that it is seldom seen
on the wing, but it runs rapidly and is able to leap into
the lower branches of trees. The penalty of danger from
weakened flight which the Emeu-wren must pay for his
caudal decoration is paralleled by certain little whydah-
finches of Africa, the males of which at the breeding
season are decorated with several tail-feathers over five The Bird 4i 6 times as long as their tiny bodies. When a heavy dew
falls during the night, drenching the plumage of these
birds, they become helpless and quite unable to fly. At
such times many are killed by their natural enemies;
and such indeed is their helplessness, brought about by FIG. 331. — Male Paradise Whydah-bird showing tail. FIG. 331. — Male Paradise Whydah-bird showing tail. this excess of nuptial dress, that a person can pick them
up in the hand without difficulty. this excess of nuptial dress, that a person can pick them
up in the hand without difficulty. The Bird The beautiful tails of pheasants are in harmony with
the wealth of colour which many of these birds display
upon other parts of the body; the long graceful tail of the
Reeves being especially striking. he folded, roof-shaped tail of the common rooster, Japanese Long-tailed Fowls. (From a photograph provided by the
American Museum of Natural History.)
417 Fio. 332.— Japanese Long-tailed Fowls. (From a photograph provided by the
American Museum of Natural History.)
417 417 The Bird 4i8 4i8
and of his wild ancestors the Jungle-fowl, with the graceful
overarching feathers, is a type of tail found elsewhere FIG. 333.— Roof-like tail of Jungle-fowl. FIG. 333.— Roof-like tail of Jungle-fowl. only in certain pheasants. In the Boat-tailed Grackle
the arrangement is reversed, the apex of the slope being only in certain pheasants. In the Boat-tailed Grackle
the arrangement is reversed, the apex of the slope being FIG. 334. — Decorative tail of Reeves Pheasant. FIG. 334. — Decorative tail of Reeves Pheasant. FIG. 334. — Decorative tail of Reeves Pheasant. beneath instead of above. The possibilities of abnormal
feather growth are well shown in the tails of the Japanese Tails 419 Long-tailed Fowl — a breed of birds in which, by artificial
stimulation, such perhaps as periodical pulling of the
feathers or else retardation of moult, has produced, in the
cocks, tails from twelve to twenty feet in length. This
process dates back, in Corea at least, to A.D. 1000, and
necessitates keeping the birds continually upon high
perches, or else wrapping the
feathers carefully in paper. The arrangement of feathers
in this artificially induced
character is duplicated in
nature in the Paradise Why-
dah-finch mentioned above. Long-tailed Fowl — a breed of birds in which, by artificial
stimulation, such perhaps as periodical pulling of the
feathers or else retardation of moult, has produced, in the
cocks, tails from twelve to twenty feet in length. This
process dates back, in Corea at least, to A.D. 1000, and
necessitates keeping the birds continually upon high
perches, or else wrapping the
feathers carefully in paper. The arrangement of feathers
in this artificially induced
character is duplicated in
nature in the Paradise Why-
dah-finch mentioned above. FIG. 335.— Useful tail of Peacock
Pheasant. dah-finch mentioned above. The Bird In both the male and fe-
male Peacock Pheasant the
tail is quite long and the
feathers are decorated with
beautiful iridescent " eyes."
But in this bird usefulness ex-
ists as a corollary of beauty. When the young chicks are
reared under a bantam hen,
they invariably keep close be-
hind their foster-mother, for
no apparent reason; indeed this position often results in
their death, a kick from the bird's foot generally being
fatal. The reason for this strange instinctive act is at
once clear when we see the chicks with their rightful
mother. They spend much of their time hidden beneath
the shelter of her long, sloping tail, coming out now
FIG. 335.— Useful tail of Peacock
Pheasant. FIG. 335.— Useful tail of Peacock
Pheasant. The Bird 420 420
and then to feed when she calls them, then hurrying
back to their snug shelter. Thus when she walks from
place to place, the tiny feet of the chicks may be seen
scurrying along beneath the beautiful tail-feathers, all FIG. 336.— Tail of Lyre-bird. FIG. 336.— Tail of Lyre-bird. but their legs concealed from view, giving a most re-
markable appearance to the mother bird. markable appearance to the mot
Among ornamental tails assumed for show during the
breeding season, that of the Lyre-bird of Australia is
unequalled. The name is well given, since the outer
tail-feathers carry out the graceful, curving outline of Tails 421
the classic form of a lyre; while twelve of the central
feathers, so scantily barbed that their stems are plainly
visible, hold positions corresponding to the strings of
that ancient instrument. The two elongated middle
feathers cross each other and curve outward, adding still
more to the decorative effect of
this strangely beautiful member. Naturally we find that these
birds are better runners than
fliers. The females lack the or-
namental tail. FIG. 337.— Tail-coverts of Quezal. If we judge from analogy
with the human race, when an
inordinate amount of ostenta-
tious show is noticeable among
birds, we occasionally find that
it is, in a sense, a sham display ;
although the analogy ceases when
we find that such a case among
birds is no less interesting than
where the phenomenon is really
what it appears to be. Upon
seeing a specimen of the beau-
tiful trogon commonly called
j.i rv i , i , FIG. 337.— Tail-coverts of Quezal. The Bird the (Quezal, the involuntary ex-
clamation is, " What a magnificent tail!" And no wonder;
for, while the bird is only about the size of a small dove,
behind it, for three and a half feet, there stream long,
iridescent green plumes, soft as down, brilliant as emeralds. Yet the true tail is a short, squarish affair, completely If we judge from analogy
with the human race, when an
inordinate amount of ostenta-
tious show is noticeable among
birds, we occasionally find that
it is, in a sense, a sham display ;
although the analogy ceases when
we find that such a case among
birds is no less interesting than
where the phenomenon is really
what it appears to be. Upon
seeing a specimen of the beau-
tiful trogon commonly called
j.i rv i , i ,
the (Quezal, the involuntary ex-
clamation is, " What a magnificent
for, while the bird is only about t
behind it, for three and a half
iridescent green plumes, soft as dow
Yet the true tail is a short, squ FIG. 337.— Tail-coverts of Quezal. The Bird The Bird 2 422
hidden by the overhanging train of gorgeous plumes,
which are in reality the upper tail-coverts. which are in reality the upper tail-coverts. The same thing is true of the peacock, whose real tail,
while it has the power of spreading, consists solely of FIG. 338. — Train of Peacock spread. FIG. 338. — Train of Peacock spread. short, dull, brownish feathers, acting as a support to the
glorious train of ocellated plumes which springs from the
lower back. Indeed the tail-feathers of a turkey-cock
are far more beautiful than the real tail of a peacock. This is especially evident when, after a peacock has moulted Ta 423 his long train, he sometimes spreads the real, incon-
spicuous tail. Large and heavy as this decoration of
the peacock is, the birds fly with remarkable ease. In
such places as the New York Zoological Park, after roost- FIG. 339. — Rear view of train of Peacock, showing real tail. FIG. 339. — Rear view of train of Peacock, showing real tail. ing all night in the tallest trees, they sail down in the
early morning, the long train waving gracefully behind — a
sight which, once seen, is never forgotten. The Bird ing all night in the tallest trees, they sail down in the
early morning, the long train waving gracefully behind — a
sight which, once seen, is never forgotten. We must leave unmentioned scores of beautiful and The Bird 424 interesting types of tail-feathers — those of hummingbirds,
birds of paradise, and many others; but there is one which
deserves especial mention. These birds, of which there
are a number of species, are the motmots, abundant in
many parts of Mexico and southward. The tail-feathers
of the Mexican motmot, which are bluish green in colour, FIG. 340. — Tails of Motmot: (a) young male; (6) adult female. FIG. 340. — Tails of Motmot: (a) young male; (6) adult female. have nothing peculiar about them, except the middle pair,
which are two inches longer than the others. Of this
extra length one inch is bare shaft, while at the tip the
barbs are normal, forming a racket-shaped extremity. The
fact which places this slight decoration above all other more
elaborate examples in point of interest is that the birds
themselves voluntarily produce the racket condition. Even
the youngest birds, of both sexes, when the long central have nothing peculiar about them, except the middle pair,
which are two inches longer than the others. Of this
extra length one inch is bare shaft, while at the tip the
barbs are normal, forming a racket-shaped extremity. The
fact which places this slight decoration above all other more
elaborate examples in point of interest is that the birds
themselves voluntarily produce the racket condition. Even
the youngest birds, of both sexes, when the long central Tails 425 tail-feathers have grown beyond the others, instinctively
begin to pick at the vane, soon denuding the shaft so
symmetrically that the rackets are equal in size. The FIG. 341. — Motmot swinging its tail. FIG. 341. — Motmot swinging its tail. photographs show this perfectly. Figure a is the tail
of a young male where the operation of ornamental de-
nudation has just begun; while b shows the condition
in an adult female. The photograph of the entire liv-
ing bird also shows the rackets, as well as the peculiar
pendulum motion of the tail from side to side, although The Bird 4 the motion is more abrupt than is the motion of a pendu-
lum. The Bird The Mexican motmot is brilliantly coloured, yet in a
densely foliaged tree, among the bright spots of sunlight,
it becomes almost invisible. It is the motion of the tail
which most often betrays the bird. In the tail marked a in Fig. 340 it will be noticed that,
where the barbs have not yet been picked off, the unmu-
tilated vane is considerably narrowed — an interesting fact
for the consideration of evolutionists, as it offers strong
circumstantial evidence, but by no means absolute proof,
of a case of the inheritance of acquired characters, a much-
rr ooted question not many years ago among scientists. If
we choose to accept the evidence thus, we may presume
that if this habit is continued through a sufficient number
of generations, the vane will, at the point of continued
denudation, ultimately become naturally bare. But, in any case, it is a fact which must hold the inter-
est of the most superficial bird student that here is a
bird which voluntarily tears away a portion of its plu-
mage. To the best of our present knowledge this is solely
to ornament itself, but the fact that both sexes equally
possess this habit makes such an explanation the merest
theory. The interest which this has for us here is not the
ultimate psychological significance of the habit, but the
fact that there is a bird which thus voluntarily mutilates
its plumage. As in so many other cases, we must
depend on future study of live birds in their natural
haunts to clear up the difficulty. It is this very com-
plexity of Nature's problems which makes a naturalist's
life ever one of enthusiasm and zest. THE EGGS OF BIRDS ERHAPS the most fascinating phase of Nature
is the way in which she cares for her chil-
dren during the early part of their lives. The
story of seeds and eggs has not been half told. Think
of the tiny thistle-fluff which soars away, borne on the
lightest breath of air; of the great cocoanuts in their
husks, so hard that they will turn the edge of a knife;
of the burrs which ever patiently reach out for some
passing creature to carry them to a distant home; of the
cones of the forest, whose seeds may be transported by
birds, or dropped to the ground only to smother in the
shadow of the parent tree. E In that "mother of life" the sea, the wonder of
the first beginnings holds us spellbound. We see the
tiny hydroids, those animal plants, flowering and budding
on their waving stalks, and presently setting free their
"seeds" — jelly-fish, — throbbing with life, drifting away
on the ocean currents. Again observe these jellies scatter-
ing behind them an untold host of eggs, as a rocket marks
its path with a myriad sparks. Think of the salmon
seeking her spawning-grounds in the uppermost reaches
of rivers, or the cod boldly playing for her offspring the
427 The Bird 4 chance in the lottery of life in the open ocean. Of her
nine millions of eggs, will one survive? How strange is the four-tendriled, purse-like cradle
of the baby shark; how delicate the forms and patterns
of butterflies' eggs ! and was there ever a more model
parent than that frog which holds its eggs in its mouth
until the tadpoles grow up? The white leathery eggs of turtles and lizards bring
us to our subject. Leading all in beauty and interest are
the eggs of birds. Precious stones have always exerted
a great fascination over mankind, and in appearance
birds' eggs may be compared with gems; indeed the shell
itself is almost wholly composed of mineral matter. But,
far from being an inanimate crystal, an egg shelters one
of the marvels of the world — an embryo bird. The
gaudy sea-shell cloaks a slimy snail, but from the beautiful
egg of a bird emerges a greater beauty. Reptiles lay white eggs whose shells are not brittle,
but, when broken, curl up like a celluloid film. THE EGGS OF BIRDS Some
of these reptilian eggs are oblong in shape, but most are
spherical and the great majority are deposited in the
ground, or under bark, and are hatched by the heat of the
decaying vegetation or by the direct rays of the sun. Thus we see that there is little need for variation in
shape or colour. Among birds, however, we find very
different conditions. As we know that birds have evolved from reptiles,
we have a right to suppose that the early forms of birds
laid white, leathery eggs, perhaps in hollow trees; but
the power of flight has taken birds entirely out of the FIG. 342.— Comparison of eg
(a) Egg of Hen. (b) Egg of Skate,
Egg of Alligator. FIG. 342.— Comparison of eggs of reptiles and birds. (a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turtl
Egg of Alligator. 429 FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 FIG. 342.— Comparison of eggs of reptiles and birds. (a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turtle. (e) Egg of Alligator. 429 429 The Bird 43°
reptilian horizon, and greatly altered all the conditions
of their life. The history of the egg of a bird, from the
time it is laid until it hatches, has an all-important effect
on its form, colour, and even upon the number of eggs
laid. This is not strange when we consider that every
minute of the bird's life is open to many dangers, and
that the egg stage — that bridging over of generations —
is a most precarious period. That which adds the greatest interest to anything is
the why of it, and a vast collection of eggs, beautiful
though they are, yet, if ignorantly looked at, is worse than
useless. Why one bird lays twenty eggs and another
but two ; why one bird's eggs are white, another's of varied
colours, we will never learn from blown museum speci-
mens. Not until we have the patience and skill to watch
and to find the most deadly enemies which threaten the
nests and eggs of birds, their number and modes of attack,
can we hope for successful solutions to the thousand and
one problems which offer themselves. What we know in
respect to eggs is fragmentary and rests on so slight a
degree of proof that every theory is attacked and re-
attacked in turn. Supposing that the eggs of the early forms bf birds
were round, — that being the most typical form of a single
cell, — we find many variations in shape among the eggs
of living species. Many of the eggs which are laid in
hollow trees still retain the primitive spherical form, per-
haps an advantage in keeping the eggs in a close group
in the centre of the floor of the cavity. So characteristic of the eggs of birds is the pear-shape The Eggs of Bir 4^1 — one end blunt and narrowing to the other — that they
have given to it its name: oval. In the eggs of certain
sea-birds which breed on the narrow ledges of perpendicular
cliffs this oval shape is carried to an extreme, and ap-
parently for an excellent reason, mechanical, but of ines-
timable value to the birds. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 Eggs laid in such positions FIG. 343.— Egg of Murre. FIG. 343.— Egg of Murre. are of course especially exposed to danger from the wind
or from some sudden movement of the birds, which gener-
ally nest very close together. Were it not that the eggs,
on account of their peculiar shape, describe an arc of
very small diameter when they roll, doubtless a far greater
number wrould roll off and be dashed down upon the
rocks below. Among the plovers, sandpipers, and phal-
aropes we again find a peculiarly pronounced pyriform are of course especially exposed to danger from the wind
or from some sudden movement of the birds, which gener-
ally nest very close together. Were it not that the eggs,
on account of their peculiar shape, describe an arc of
very small diameter when they roll, doubtless a far greater
number wrould roll off and be dashed down upon the
rocks below. Among the plovers, sandpipers, and phal-
aropes we again find a peculiarly pronounced pyriform The Bird 432 432
shape of egg, serving in these instances a very apparent
and useful end. These birds almost invariably lay four
eggs, which are of large size in comparison with the birds,
and their shape allows them to be fitted closely together,
each forming one of the four segments, their points all
but meeting in the centre. Thus the little body of the FIG. 344.— Eggs of Killdeer. FIG. 344.— Eggs of Killdeer. parent is large enough to cover them all, which would be
impossible were the eggs arranged at random. The eggs
of grebes are peculiar in having both ends alike. The number of eggs which a bird lays has been found
to bear a definite relation to the amount of danger to
which the species is exposed — a fact which holds good The Eggs of Bir 433 in the young of many, if not all, other Phyla of animals,
and which is one of the most interesting provisions brought
about by the slow but sure working of evolution. We
may instance the few eggs of the voracious and masterful
sharks and the millions of spawn necessary to enable
the halibut and the cod to continue in existence. Mr. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 Ernest Ingersoll has so admirably summed up
the matter of this relation of the number of eggs to the
corresponding danger that I cannot do better than to
follow his argument, quoting his words with a slight
change here and there. This phase of the study of eggs
being so clearly understood, it is well worth a little detail
as an illustration of how interesting all the other problems
will become when we once get on the right road to their
solution. Among the majority of birds the average number of
eggs in a nest is from three to six; we may take five as a
typical average. "Any considerable departure from this
normal number in a species or Family must then be ac-
counted for by some specific or tribal peculiarity in cir-
cumstances. " Beginning with the ostrichlike group at the bottom
of the list, we find ourselves face to face with an inter-
esting state of things, to which the number of eggs is
an index. Ostriches, rheas, and emeus incubate large
clutches — a dozen or more, — those inhabiting the conti-
nents of Africa and South America, however, producing
twice as many eggs annually as their relatives of Australia
and the neighbouring smaller islands. " Immediately following and contrasting with them are The Bird 4 the three groups characterized by the curious elephant-
footed, often gigantic moas, and similar birds of Mada-
gascar, Mauritius, New Zealand, and the Papuan region,
which have become extinct within the historic period,
except the kiwis, to be spoken of later. All of these,
so far as we know, laid only one egg at a time, which^
plainly enough, was sufficient to keep the race going in
the limited space afforded to each species by its island,
but which did not suffice to prevent an almost immediate
extinction of these species as soon as mankind discovered
that the birds and their eggs were serviceable. But Provi-
dence, or Nature, or natural selection, or whatever has
been the ruling influence in determining means and limits
for animal life, seems never to have taken man into ac-
count. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 " Turning now to the sea-birds — penguins, gannets,
murres, puffins, auks, petrels, guillemots, tropic-birds,
and the like, — we find that none of them is in the habit
of laying more than one egg, as all breed on such remote
and inaccessible rocks, often in holes, that harm can
rarely happen to their young, and therefore a very high
percentage comes to maturity. Many of these breed in
companies, and are so unacquainted with danger that
they make no attempt to hide their eggs or to leave the
nest when the place is visited by some wandering natu-
ralist or egging party. "The habit of the King Penguin deserves a note to
itself. This big Antarctic bird guards its one white egg
from harm by carrying it somewhat as a marsupial does
its young, in a pouch formed by a fold of the skin of the The Bird 6 436
body between the thighs. Both sexes are provided with
this contrivance during the breeding season, and relieve
each other of the burden at intervals. " The gull tribe, however, are far more exposed to acci-
dent and to enemies, both in adult life and as to their
eggs and young, than are the penguins, petrels, and others
mentioned above; and here the rule is from two (skuas)
to four (gulls and terns) eggs in a nest. When we come FIG. 346. — Eggs of Ostrich, Cassowary, Hummingbird, and Hen,
showing comparative size. FIG. 346. — Eggs of Ostrich, Cassowary, Hummingbird, and Hen,
showing comparative size. to the shore- and marsh-birds — the plovers, snipe, sand-
pipers, jacanas, all of which nestle on the ground, usually
near the shore of the sea or lakes — we judge them to be
exposed to about the average of dangers, since their nest
complement is from four to six. The northern, tundra-
loving cranes need raise few young, and hatch only two
eggs; but when we come to the water-birds — the rails,
gallinules, ducks, and geese — we find an extensive group to the shore- and marsh-birds — the plovers, snipe, sand-
pipers, jacanas, all of which nestle on the ground, usually
near the shore of the sea or lakes — we judge them to be
exposed to about the average of dangers, since their nest
complement is from four to six. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 The northern, tundra-
loving cranes need raise few young, and hatch only two
eggs; but when we come to the water-birds — the rails,
gallinules, ducks, and geese — we find an extensive group The Eggs of Birds 437 whose nests average a dozen eggs in each set. Explana-
tions are ready for this: the birds themselves are exposed
to unusual peril, from weather as well as from active
enemies, since they mostly emigrate to the extreme North
and nest in the edges of marshes, where the sitting birds, FIG. 347.— Nest of Laughing Gull. FIG. 347.— Nest of Laughing Gull. eggs, and young are all subjected to freezings, floods,
and countless marauders that depend largely upon them
for food during the Arctic summer, so that a heavy annual
recruiting must be made to repair losses. Few birds are
liable to so many misfortunes and mishaps as the water- eggs, and young are all subjected to freezings, floods,
and countless marauders that depend largely upon them
for food during the Arctic summer, so that a heavy annual
recruiting must be made to repair losses. Few birds are
liable to so many misfortunes and mishaps as the water- The Bird 438 438
fowl, except perhaps the big and pugnacious swans, who
can take better care of themselves, and lay only five
eggs or fewer. The long-legged wading birds also, such
as the storks, ibises, herons, and the like, are fairly safe
in the breeding season, because they nest in trees, as
a rule, (Fig. 356,) and consequently we here find only two FIG. 348. — Nest and eggs of California Partridge. FIG. 348. — Nest and eggs of California Partridge. to four young in the annual brood; so with the snake-
birds. "This brings us to the game-birds — the world-wide
tribes of partridges, pheasants, grouse, turkeys, jungle fowls,
peacocks, and the like — which are of large size, run about
on the ground, and are of interest to sportsmen and epi-
cures. With few exceptions, these must put forth a The Bird 4 large complement of eggs (eight to twenty) in order to
bring to maturity enough young to replace the yearly
mortality, for the ground-built homes and huddling chicks
encounter a multitude of dangers to which birds in trees, or
even the small-sized ground-nesters, are not exposed. One
exception here singularly favours the rule. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 The Thibetan
Peacock Pheasant inhabits the heights of the Himalayas,
where it has to contend with only three or four nest-
robbers, instead of the countless foes that infest the lower
jungles; hence its ample breast warms but two eggs. " The doves and pigeons lay only two eggs, and a few
lay but one; but this seems to be due to the fact that
their extraordinary powers of flight render them, as adults,
unusual immunity from capture and famine, rather than
to any special safety pertaining to their method of nidifi-
cation. " Hawks and owls in general have four or five eggs,
and as this is about the average number of the small
birds on which they largely prey, it seems evident that
their chances of life and the difficulty of sustaining it
are, on the whole, no less than are met with by their
victims. The owls, however, vary much among them-
selves in this respect; the Snowy Owls, whose home is in
the snowy north, where a nest in the tundra moss is acces-
sible to every marauder, and the Burrowing Owls, whose
underground homes are constantly robbed, being obliged
to lay twice as many eggs as the remainder of the family
in order to overcome the high percentage of casualties
due to these unfortunate situations. An odd feature in the nidification of some of the The Eggs of Bi 441 Arctic-breeding owls, where the nesting must take place at
an unreasonably early and cold date in order to give the
fledglings time to reach mature strength before the suc-
ceeding winter assails them, is that these birds deposit
their eggs at intervals of a week or ten days. In this
way the mother can envelop in her plumage and keep
thoroughly warm one egg and a callow fledgling at a FIG. 350.— Eggs of Screech Owl. FIG. 350.— Eggs of Screech Owl. time, and is assisted, in respect to the later eggs and
fledglings, by the warmth of the older young in the nest. fledglings, by the warmth of the older young in the nest. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 ' The parrots are a wide-spread and numerous tribe,
and none of the larger species need lay more than two
or three eggs, for they protect them in deep holes in the
earth or in trees, and are able to defend them; but some
of the smaller parrakeets lay as many as twelve eggs,
reflecting the greater dangers with which they have to The Bird 442 442
contend. Toucans are able to get along with a pair of
eggs; while a hornbill, by sealing its mate up in its little
arboreal cavern during nidification, is so adequately
protected that one to three eggs in each family suffice
to keep the race going, since practically every young FIG. 351. — Burrowing Owl at nesting hole. FIG. 351. — Burrowing Owl at nesting hole. bird is brought to maturity. Of the host of smaller and
weaker birds nesting in cavities, two to five eggs are the
usual quota. This brings us to the tribes of little singing
birds with which we started, whose average is about
five; but a few interesting exceptions may be noted. Our whippoorwills and night-hawks, for instance, lay
only two eggs. These are placed on the ground in the The Eggs of Bi 443 woods, surrounded by no nest, and are so precisely the
colour of the dead leaves that nothing but the merest
accident would lead to their discovery by the eye alone. The same is eminently true of the bird itself. None of
the almost uncatchable hummingbirds needs to lay
more than two eggs in order to recruit the ranks of its FIG. 352. — Nest and eggs of the Anna Hummingbird. FIG. 352. — Nest and eggs of the Anna Hummingbird. species to the full quota permitted it in the numerical
adjustment of bird life. " I have gone into this matter somewhat at length,
though by no means exhaustively, because I am not
aware that the matter has ever been exploited, and be-
cause it embodies a general law or principle. Thus we
see that the nest complement of eggs of any bird is in
exact proportion to the average danger to which that " I have gone into this matter somewhat at length,
though by no means exhaustively, because I am not
aware that the matter has ever been exploited, and be-
cause it embodies a general law or principle. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 Thus we
see that the nest complement of eggs of any bird is in
exact proportion to the average danger to which that Bird 444 species is exposed. I believe that this factor is fairly
constant for species or tribes of similar habits, and that
exceptions indicate peculiarities of circumstances which
in many cases we can easily perceive, because I believe
that Nature is strictly economical of energy, allowing
no more eggs to be laid, and consequently young to be
produced, than the conditions justify in each case. Thus
the uniformity of avine population — the balance of bird-
life — is maintained." life — is maintained."
When a bird's nest and eggs are destroyed, she will
often lay another setting, and some birds raise two and
even three broods in a season under normal conditions. If the eggs of a bird are removed as fast as they are laid,
the bird will sometimes continue to lay, one of the most
remarkable instances of this in an uncaged bird being
a Flicker which laid seventy-one eggs during the space
of three-and-seventy days. A tiny African Waxbill in
captivity has been known to rear fifty-four young in the
course of a year, during the same period laying an addi-
tional sixty-seven eggs! The domestic hen has become
a veritable egg-laying machine, thanks to careful breed-
ing in the past, since the wild Red Jungle Fowl from which
all varieties of poultry are descended, lays only one nestful
of seven to twelve eggs once a year. Many birds still hold to the old style of nesting in
hollow trees and such concealed places. Whether they
hunt around until they find a cavity ready-made by the
elements, or whether, like the woodpeckers, they pro-
ceed to excavate a home in a dead branch, or, kingfisher-
like, to tunnel deep into a sand-bank, their eggs are almost The Eggs of Bi 445 invariably white. Many indeed have such glossy, highly
polished shells that, were they laid in exposed situations,
their shining surface would be a sure guide to hungry FIG. 353. — White eggs of Hairy Woodpecker in hollow tree. (Bowdish, photographer.) FIG. 353. — White eggs of Hairy Woodpecker in hollow tree. (Bowdish, photographer.) egg-eaters. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 Among such birds may be mentioned the
owls, woodpeckers and parrots, trogons, motmots, king-
fishers and puffins, besides many others which hide their The Bird 4 eggs in domed nests. On the other hand we find a num-
ber of birds laying spotted eggs in concealed nests, and
white eggs in open places; so that no universal law can
be framed to account for the varied colouring. This is
not surprising when we think of the great difference of
conditions under which each species lives. Take for ex-
ample the two species of marsh wrens which live so happily
among the reeds of the marshes of our Eastern States. Both birds build globular mouse-like nests, both hide their
treasures deep in the interior, but the eggs of the Long-
billed species are dark chocolate-brown, while the Short-
bill's eggs are like pearls. We do not know why this
difference exists, but that need not deter us from accept-
ing the facts to which the majority of eggs seem to point:
that eggs which are concealed, having no need for colour-
ing, are white like those of reptiles. If, as many writers
have suggested, the colours of eggs are only meaningless
by-products, there is no reason why these hues should
not run riot upon each egg or nestful of eggs, as is the
case in one or two interesting isolated cases to be men-
tioned shortly. Perhaps the most marked exceptions to the theory
of the protective coloration of eggs is to be found in
doves and pigeons, which lay white eggs in open nests
(Fig. 349) ; with the exception, curiously enough, of the
Rock Dove, the wild progenitor of our domestic birds,
which places its nest in inaccessible caverns in the face of
cliffs. The almost total extermination of the Passenger
Pigeon has been instanced as an example of a "mistake"
of Nature in allotting to it white eggs; the absurdity of The Eggs of Bir 447 which statement is apparent when we consider that the
havoc was wrought upon the adult birds and by manl Wallace has suggested that the nests of doves are so
loosely and so flimsily built — being in reality mere plat-
forms of sticks — that, looking up at them, the eggs simu-
lated the colour of the sky beyond and so became incon-
spicuous; but unfortunately that argument is so decidedly FIG. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 354. — Nest and eggs of Mallard Duck. FIG. 354. — Nest and eggs of Mallard Duck. suggestive of human presence that it loses much of its
value when we remember that egg-hunters among the
mammals and birds do not stand on the ground to take
observations, but either climb the trees in search of nests
or fly low above the branches. The eggs of ducks and grouse are white or very light-
coloured, and are laid in open nests upon the ground. The mother duck's plumage is the very essence of the
mottled lights and shadows among the reeds, and when The Bird 4 she leaves her eggs she backs carefully away, drawing
over them, at the same time, a coverlet of beautiful down,
the protective colouring of which is ample to shield the
eggs. Ordinarily this coverlet is rolled up at the edge
of the nest. It is to such a habit that the eider-down
hunters owe their supply. A grouse does not pluck the
down from her breast, but in devotion and ability to
remain close upon her eggs she has few equals. It is
rare indeed to find the nest of a grouse unguarded, and
the mother bird will all but wait until your hand is upon
her before leaving her eggs exposed. The many species of hummingbirds lay the whitest
of eggs, but here it is the nest which is protected, — fash-
ioned of dull-hued plant-down, with beams and rafters
of cobweb, covered outside in our Eastern species with
lichens exactly like those which are growing upon the
limb to which the tiny air-castle is attached. The nests
of vireos, also, are much like their surroundings. of vireos, also, are much like their surroundings. Herons and egrets, pelicans, cormorants, storks,
swans and geese, all lay white or whitish eggs in open
nests; but obviously these birds require little protection, all
being able to defend themselves with beak or wing. Some
of them nest, too, in large colonies, adding the advantage
of numbers. The constant need of vigilance in protect-
ing eggs thus exposed is at once evident when mankind
—that disturber of Nature for whose intrusion she seems
never prepared — comes upon the scene. FIG. 342.— Comparison of eggs of reptiles and birds.
(a) Egg of Hen. (b) Egg of Skate, (c) Egg of Snake, (d) Egg of Turt
) Egg of Alligator.
429 If we make our
way into the heart of a Florida rookery of herons, ibises,
or cormorants, many of the birds will be frightened from
their nests and the Fish Crows take instant advantage, The Bird The Bird 45° 45°
swooping down one after another upon the nests and
each impaling an egg upon its beak and flying off with it. They would never dare such open villainy were the herons
undisturbed. FIG. 356. — Colony of Great Blue Herons. Many of the more isolated cases of exposed whi FIG. 356. — Colony of Great Blue Herons. Many of the more isolated cases of exposed white
eggs are to be explained, I think, by the fact that the
habits of birds often change rapidly, while their structural The Eggs of Bi 45 1 adaptation follows more slowly. For example, let us
take the group of owls. The majority of these birds nest
in hollow trees, but even these occasionally make use of
an open hollow or a very shallow one, and individual,
radical departures from the conventional owl-habitation
are doubtless not uncommon. But these exposed eggs
are soon destroyed; for no crow, jay, or squirrel could ever
resist any opportunity to avenge himself for the wrongs
inflicted by his ancestral enemy, the owl. But when,
urged on by that impulse which ever tends to make birds
vary their habits in all directions, some owl, such as the
Short-eared, finds good feeding on marshes and open,
treeless plains, it naturally takes to nesting on the ground,
in nests but partly concealed by the overhanging grasses. in nests but partly concealed by the overhanging grasses. Three things might now happen. If sufficient varia-
tion occurred and the conditions demanded it, natural
selection might bring about a protective colour on the
shells of the eggs; if enemies were few and easily over-
awed, the eggs might remain white; while, on the other
hand, the enterprising race might be wiped out of exist-
ence for no more reason than the colour of the egg-shells. The second result seems to be the good fortune of the
Short-eared Owls. All of these fates have undoubtedly
overtaken birds again and again, and it is by the inter-
action of such condition , combined with an ever-chang-
ing environment, that many phenomena are brought about. ing environment, that many phenomena are brought about. It was by reason of the general similarity in colour
which the eggs of related groups of birds tend to show
to each other that oology, or the science of egg-shells,
was able to initiate an important scientific discovery. The Bird The Bird 452 452
At one time the sandpipers and plovers were classed as
wading birds, and the gulls and terns in an Order placed
at a remote distance in the scheme of classification from
the former birds; no one suspecting that the two groups
were in any way related. The striking resemblance
which their eggs showed, however, suggested an affinity FIG. 357. — (a) Egg of common Tern compared with (6) egg of Black-necked St FIG. 357. — (a) Egg of common Tern compared with (6) egg of Black-necked Stilt which was later perfectly confirmed by anatomists and
embryologists. The few thousands of years during which our race
has risen to inheritance of the earth is all too short
a time, geologically speaking, for us to flatter ourselves
that any of the protective colours of animals were de-
veloped on our account; but in many instances we,
sharing the same five senses of animals, may put our-
selves in their position. Imagining ourselves egg-hunting FIG. 358. — Eggs of Cassowary. FIG. 358. — Eggs of Cassowary. FIG. 359.— Eggs of Ostrich. 453 FIG. 359.— Eggs of Ostrich. Bird 454 animals, let us consider some of the more patent cases
where eggs are coloured for protection — where they
mimic their surroundings so perfectly that only the most
careful search reveals their whereabouts. Ostriches and
Cassowaries are two interesting examples, the former
bird laying its white eggs upon the white sands of the
desert; while the cassowary, in the depths of its jungle
home, incubates a nestful of eggs of the most exquisite
emerald hue, matching perfectly the green moss upon
which they rest. I knew of one of these birds confined
in a small paddock of green grass, whose splendid eggs,
measuring three by six inches, once remained undiscov-
ered for weeks, although laid openly upon the ground. Special search was necessary to find even these great eggs. Special search was necessary to find even these great eggs. If we walk in the woods in June and happen to flush a
night-hawk from the ground, the most careful scrutiny of
the place where the bird rose will often fail to reveal to
our sight what at last our fingers detect — two eggs, their
shells imbued with the colours of the forest floor. The Bird I have
led persons to a spot on a beach of shells and sand,
told them that there were twenty-one good-sized eggs
within a radius of fifteen feet, and seen them utterly baffled. The olive-gray, blotched shell of a tern's egg rests among
dark pebbles, or more often upon a wisp of seaweed, into
whose irregularities the hues of the eggs melt and mingle
perfectly. The Black Skimmer, that most interesting
bird of our coast, lays its eggs upon the bare sand among,
or sometimes in, the large clam-shells which the storms throw
up in windrows. Against man's systematic search their
wonderful assimilative colouring is of course often useless, The Eggs of Birds 455 but sharp as is the eye of passing crow or beach-patrolling
bear, the eggs to them would appear but bits of sand and
shadow. And thus we might go on with many other examples
of protection derived from the pigment on the shells —
protection which in a hundred instances might prove FIG. 360.— Eggs of Night-hawk. FIG. 360.— Eggs of Night-hawk. futile, but which in the great summing up and balancing
of Nature's profit and loss is of inestimable value to the
race. We find an unusual condition in the colouring of the
eggs of sea-birds, — of certain of those species which nest
on inaccessible cliffs. If pigment was developed in the The Bird 56 456
eggs of the ancestors of these birds for the sake of protec-
tion, all need for it is now lacking, and as an apparent
result the various hues seem to have run riot. One may
place a hundred murres' eggs side by side and find no two
alike, while the extremes would never be recognized as
belonging to the same species of bird. FIG. 361. — Nests of Tern and Skimmer. FIG. 361. — Nests of Tern and Skimmer. Another instance of extreme variability in the colour
of eggs and an instance of intensely interesting import is
found in the English Cuckoo, which may be taken as an
example of species which are parasitical, — in the sense
that the females make no nest of their own, but deposit
their eggs in the nests of other birds, the young being thus The Bird 4 hatched and reared by foster-parents. Such an unusual,
almost unique habit has brought about a considerable
modification of the eggs. The Bird Anything which would tend to
deceive the greatest number of intended victims would,
of course, greatly redound to the advantage of parasitical
birds. The remarkable similarity of the English Cuckoo's egg
to those in the nest in which it is laid has been explained
as due to each individual bird being accustomed to lay its
egg in the nest of the same species favoured by its parents
and its more distant ancestors ; its eggs in course of time, by
natural selection, thus coming to resemble the eggs of that
particular species. Other adaptations are the extremely
small size of the egg in comparison with the parent bird,
and also the unusual strength and weight of the shell. This last is doubtless of great value; for, strange as it
may seem, the bird first deposits its egg upon the ground
and then picks it up in its beak and places it in the nest
selected. Thus a strong shell is a very necessary require-
ment. The colours of eggs have been carefully examined with
the spectroscope and are found to consist, chemically, of
seven pigments: a brownish red, two delicate blues, two
clear yellows, a peculiar brown hue, while the seventh is a
rather indefinite shade, known as lichenixanthine — most
interesting of all as being identical with a colour substance
common in plants and especially in lichens and fungi. These substances somewhat resemble those found in the
blood and the bile. They are deposited on the shell while
the egg is passing down the oviduct, and it is to the circular The Eggs of Birds 459 or erratic motion of the egg that the curious scrawls and
blotches upon some eggs are due. The shell is deposited
in successive layers, and from the dim, clouded appearance
of many colours we judge that the pigment is often partly
concealed by the outermost layers of the shell. FIG. 363. — Nest and eggs of Skimmer, showing the remarkable variation in colour
of the eggs in a single nest, heightening their resemblance to pebbles or sea-
shells. FIG. 363. — Nest and eggs of Skimmer, showing the remarkable variation in colour
of the eggs in a single nest, heightening their resemblance to pebbles or sea-
shells. The Bird Occasionally, in the eggs of birds which number only
two in a nest, one egg will be almost white and the other
coated with an abnormal density of pigment. In certain
species of small birds which lay four or five eggs, one egg
always differs remarkably from the rest. Can we not The Bird 4 account for this latter condition on the hypothesis that an
actual change — an increase — is slowly taking place in the
number of eggs of this species, the abnormal shell reflect-
ing the as yet only partial readjustment of the pigment-
gland to meet the extra demand? gland to meet the extra demand? The carbonate of lime, of which the shell is chiefly
composed, varies in its composition, being sometimes so
fine that the surface has a high gloss, the eggs of wood-
peckers being a good example, or again loose and chalky,
as in cormorants. In tinamous the glossiness is carried
to an extreme, their eggs resembling ovals of highly
burnished metal, green and purple in colour. The shells of ducks' eggs are impregnated with an oily
substance, which must be of great use in resisting the
dampness and moisture of their surroundings. The eggs of some entire Families of birds are easily
recognized by the resemblance of the grain of the shell;
while, on the other hand, this microscopic appearance in
the eggs of individual species may differ considerably,
as in the case of the eggs of the Mute and Whooping Swans. The eggs of the North African Ostrich have a surface
smooth as ivory, while the eggs laid by the South African
birds are deeply pitted. The beautiful eggs of the casso-
wary show an extreme condition, the light green surface
of the egg being covered with raised irregularities of a
darker green colour. The thickness of the shells of ostrich eggs is remarkable,
and their strength permits their use as water-bottles — an
invaluable boon to the Arabs of the desert. With the relative size of the egg and the bird which The Eggs of Bi 46 1 lays it we will not here concern ourselves, except to remark
that the largest egg in proportion to the size of the bird
is that of the apteryx. The Bird If we imagine a rather smallish
hen laying an egg 3X5 inches in size, we will get a vivid
idea of this bird's ability, and it lays two at a setting! The smallest of all eggs is that of the hummingbird, while
the largest is the egg of the extinct giant jfipyornis of
Madagascar, the shell of which measures 9X13 inches. In some cases the fossil egg is all that is left to us to hint
of the existence of these great feathered creatures. Many
of these shells have been found buried with some old
native chief, the whole egg placed beside him to furnish
food for the long journey after death. Whether we look at eggs from the standpoint of an
artist's delight in harmonious and delicate colouring, or
from the wonder of their scientific composition, or even
from the point of view of a hungry man sitting down to
breakfast, we must admit that they deserve all the appre-
ciation which their beauty and their utility demand. THE BIRD IN THE EGG ]HE embryology, or life of the bird in the egg, is
the most mysterious and wonderful part of the
entire physical aspect. Many of the lesser de-
tails of growth are very difficult to study without the
use of microscopic sections and wax models; but a little
knowledge of the subject is more interesting and simple
than one would imagine. ] The very best way to begin our study of the life in the
egg will be to go to the nearest pond or marsh, if it is spring-
time, and bring home a pailful of freshly laid frog's eggs
— those queer, gelatinous masses filled with black dots. Place them in a flat, white basin, and into a smaller saucer
near by break a fresh hen's egg, being careful not to injure
the yolk. Separate one of the frog's eggs with a spoon
and put it beside that of the fowl. Now examine them
carefully with a good dissecting-microscope or even with
a hand-lens. We see a large, round, yellow yolk in the case of one
egg, and a tiny speck of black and white in the other, —
both apparently inanimate bits of matter, but which,
merely by the application of heat in the one instance insta The Bird in the Egg 463 and the presence of water in the other, will slowly take
on the semblance of living creatures; the one eventually
to swim forth, live the life of a fish for a time, then to
leap upon the land and croak among the reeds. The
other yolk would have evolved into a downy, yellow chick. We cannot hope to solve the mystery of life, but there
is a fascination in seeing how near its beginnings we can
approach. FIG. 364. — Egg of Hen, opened to show a 3-day embryo in position on the yolk. (Slightly enlarged.) FIG. 364. — Egg of Hen, opened to show a 3-day embryo in position on the yolk. THE BIRD IN THE EGG (Slightly enlarged.) If we have ever watched under the microscope the strange
little creatures which live in the mud at the bottom of
ponds, we will have realized the wonderful possibilities of a
single drop of living matter, — a single cell, — from the
amceba with its ever-changing shape to the swiftly moving
slipper paiamecium and the beautiful animal vases, — the The Bird 4 vorticella, on their queer little corkscrew stems. All these
are made up of but a single cell, and in the beginning all
seeds of plants and all eggs of animals likewise consist of
one cell. If we examine a chicken while it is being dressed for the
table, we can easily find the ovary, a mass of hundreds
of tiny golden spheres, — eggs which would have been
laid during the coming years. So we realize that the most
essential part, in fact the real egg, is only the yolk; all
else being merely protective. The shell protects the yolk
while the chick is developing during incubation, and
although formed of crystals of lime, yet it is so porous
that oxygen can enter and carbonic acid gas escape. The
viscid white, or albumen, is nutritious as well as protective,
while the yolk itself is the real food of the embryo and also
acts as a support to the developing chick. If we look
carefully, we will see two whitish, twisted strands which
extend from the yolk through the white. These two strands
have whitish opaque knots strung along them, and from a
fancied resemblance to hailstones they are called chalazce. These act as pads to protect the yolk from sudden jars,
but they do not act as suspensories. A hen never turns
her eggs, as many people imagine, to warm the different
sides equally, for the germ-dot — the position of the future
embryo (of which we will speak presently) — is always
on the lightest side of the yolk, and whichever way the egg
is turned it always swings uppermost, nearest the heat
from the body of the sitting hen. The turning, however,
may be of advantage in allowing moisture to act upon a
greater surface of shell. The Bird in the Eg The Bird in the Eg 465 Now let us examine closely the egg of the frog. It,
too, has a protective gelatinous outer coating. Before the
egg was laid it was enveloped with several very delicate
membranes, which were sponge-like in their property of
absorbing water, and when deposited in a pond they
immediately swelled up to the present gelatinous con-
sistency. If the egg has been deposited but an hour or
two, it will show a perfectly smooth surface under the
lens, but look at it intermittently for a half-hour, or even
longer, and you will be well repaid. Slowly but surely,
as the shadow of an eclipse darkens the face of the sun,
a tiny furrow ploughs its way over the surface of the dark
end of the egg. It lengthens and deepens and soon divides
the egg into two equal halves. Let us stop a minute and realize what we have seen. It is all but the beginning of life, the first hint of a higher
order of things than those one-celled creatures which we
dredged from the mud, — than the life which, untold ages
ago, was all that the earth boasted. The original cell of
the egg has, before our eyes, divided into two! But
while we have been lost in wonder and awe, — for the lover
of Nature must indeed be stolid if the first sight of such a
happening does not stir his deepest emotions, — the life
has ceased its progress never an instant. A new furrow
appears, crossing the first at right angles, dividing the egg
into quarters; then other furrows dividing it into eighths,
then cross-furrows, and the count is lost; the multitude of
cells repeating themselves hour after hour, day and night,
arranging themselves, each in its right position, obeying
some inscrutable law, until at the end of about 300 hours The Bird 4 the tadpole wriggles his way through the cloudy mass of
gelatine and swims into the water. The first steps of this dividing or cleaving of the original
single cell is similar in all eggs. The deep significance of
the equality of the first two cells may be better appre-
ciated when we know that if one of these be destroyed by
a touch from a red-hot needle, a perfect half tadpole will
develop from the other unharmed twin cell. The Bird in the Eg If we observe
the cleavage of the whiter portion of the frog's egg, we will
notice that the furrows, though ultimately extending all
the way around, yet grow very slowly in that portion. This
is because much of the white part consists of yolk, or true
food-matter, the more active formative material being
confined to the black portion. If we follow this segmentation of the cells for some
time, the egg of the frog will come to look like a diminutive
blackberry — a single layer of cells thickly covering its
entire surface, like the rounded protuberances of the berry. Now a curious thing happens. A tiny nick appears in one
side, which gradually deepens and widens until it extends
deep into the egg, pressing two rows of cells into close
proximity to each other. This will be perfectly clear if
we take a small rubber ball and squeeze it until one hollow
hemisphere is pressed into the other. This stage of em-
bryological life is called the gastrula, and is of the greatest
significance, as we shall soon see. Without further comment at present, let us now leave
the frog's egg and consider that of the fowl. When the
yolk or egg has but just left the ovary a tiny dot is visible
on one side, — the germinal vesicle, which after fertilization FIG. 365. — Stages in the development of frog's egg, from first division into twe
cells up to well-formed larval tadpole. (From original drawings by the author.)
467 FIG. 365. — Stages in the development of frog's egg, from first division into twe
cells up to well-formed larval tadpole. (From original drawings by the author.)
467 The Bird 4 immediately begins to divide into numerous cells, as in
the case of the frog's egg. This goes on until the egg is
laid, and when we break the shell, we see at the uppermost
part of the sphere of yellow yolk a well-defined portion,
in appearance a tiny ring of cloudy, opaque matter enclos-
ing a transparent circle. So now we see the use of begin-
ning our investigation with the frog's egg, that of the fowl
having reached quite an advanced stage before it is laid. having reached quite an advanced stage before it is laid. The Bird in the Eg The ring and circle of the embryonic spot on the yolk
consists of a layer of small, even cells, like cobblestones. These are spread over the top of the yolk, while just be-
neath is a jumbled mass of many larger cells. The opaque
ring is caused by a thicker, denser concentric layer of these
lower cells. When heat is applied, this outer layer begins
to segment rapidly, the new cells spreading down over
the surface of the great ball of yolk; a curving depression
dimples the surface of the little transparent circle, pushing
in deeper and deeper; and behold! we have the very same
condition — the gastrula stage — which we saw in the frog's
egg. To make this stage in the egg of the hen more real,
squeeze the rubber ball into a hemisphere and clap it
upon an orange so that the two layers of rubber fit, cap-
like, upon the fruit. This is all very wonderful, but what special significance
has it? What particular point upon which we may sus-
pend it in our memory, so that it will always return to us
with a thrill of interest and wonder whenever we see an
egg ? Just this. When we first examined the frog's egg,
and when the egg of the chick was still attached to the
ovary, they were comparable to the one-celled creatures The Bird in the Egg 469 living in the mud of the pond, which are the most lowly
organized beings in the world. The gastrula stage — the
double-walled cup, into which the real egg-part of each double-walled cup, into which the real egg-part of each
j&cain
Ear
FlG. 366.— Third-day stage of embryo chick. (See Fig. 364.) Greatly enlarged-
M, Muscle-plates (false vertebrae). j&cain FlG. 366.— Third-day stage of embryo chick. (See Fig. 364.) Greatly enlarged-
M, Muscle-plates (false vertebrae). yolk forms itself, is comparable with the next higher class
of living creatures, the sponges. For the simplest of
these are nothing more than a cup of cells, two layers deep The Bird 4 (these layers being known as the ectoderm and endoderm,
or outer skin and inner skin). The name gastrula, or little
stomach, is certainly most applicable, for an animal of
this kind consists of hardly more than stomach and mouth. The Bird in the Eg But the embryo of the frog's egg does not long remain
in this sponge-like condition; for almost immediately a
third layer, the mesoderm, or middle skin, appears between
the other two. From these three layers of cells all the
parts of the body of the future chick arise, by the continued
dividing of the cells. The details are far too involved to
be followed without going into technicalities. Suffice it to say that in the development of the embryo
chick we have one of the surest proofs of the truth of the
theory of evolution, — of the gradual evolving of each of
the higher groups of animals from some lower, more
generalized form, until all are originally derived from an
organism consisting of a single cell, with its tiny germ-
spot. The dividing of this germ-spot in the dawn of
creation was the beginning of that wonderful unrolling
of life which to-day culminates in birds and the higher
mammals, — even in man himself. mammals, — even in man himself. It would be too much to expect that the growing
embryo chick distinctly reflects in its successive stages of
growth, during a short three weeks, the embryonic states
of all its unnumbered generations of ancestors. The record,
like that of palaeontology, is imperfect. Many important
phases are slurred over or apparently entirely omitted;
in order, evidently, to give freer play to the development
of organs which will be of vital importance in the future
active life of the bird. Now and then, however, a gleam — The Bird in the Eg 471 a spark of life reflected from the far- distant past shines forth
so vividly as to hold us spellbound, almost instantly to
fade out forever, having no part in the actual life of the
chick. Like the finding of the Archseopteryx, these dim
reflections seem to have been preserved by some kind
Providence, especially to aid our groping efforts to find
the truth of ages that are past. Were it not for these
we should never dare to voice such an incredible theory
as the story of evolution would be, were it not supported
by unanswerable proofs. The question which interests
scientists to-day is not whether evolution is true, but how
its processes and changes have been brought about. The Bird in the Eg The difficulty of seizing upon these evanescent bits of
realism of the past will be appreciated when we know
that while, in the case of the hen's egg, three weeks are
required before the chick is ready to break the shell, yet
when incubation has proceeded but eighteen hours, a tiny
rod of cells shows where the notochord will be formed —
that gelatinous foreshadowing of the back-bone. Thus a
character, found first in living organisms as high in the
scale of life as fish and primitive fish-like creatures, makes
its appearance in a few hours, giving but the scantest
opportunity for the passing in review of embryonic features
of the great group of invertebrates, or those animals, like
starfishes, crabs, worms, and insects, which lack a back-
bone. The simplest way to study the growing embryo is to put
a number of eggs in an incubator, or under a hen, and
examine one on each successive day. If the egg is held
firmly, by pressing it down into a box of loose sand, the The Bird 4 upper part of the shell may be carefully picked away with
a pin and the little embryo exposed to view. a pin and the little embryo exposed to view. When thirty-six hours old it measures almost one
quarter of an inch in length and shows many interesting
things. The embryo is set off from the rest of the yolk,
much as one's hand is if placed under a piece of cloth, the
latter then being tucked in beneath the palm in all direc-
tions, until the gathered portion is closely constricted. We are able with a good lens to make out which is the
head and which the tail end of the future chick, the former
being broader and showing the beginning of the two tiny
swellings — the future eyes. Behind these, four faintly
outlined enlargements along the central line show the
anlagen of the various parts of the brain. These take up
about one third of the entire length of the embryo, showing
the importance of the organs of the head. Still farther
back are two rows of little segments strung along the
centre line — the false back-bone, hinting of the worm-like
series of muscles, of which we have already spoken (page
69). A heart is even now hinted at, but is seen better in a
later stage. The Bird in the Eg An interesting thing about it, however, is
that, at this stage, it is really in the head region, vividly
recalling the condition existing in fishes, where it is very
far forward in the body, in fact only just behind the gills. At this period in the chick embryo the heart, instead of
being a complicated organ, divided into four complete
cavities, is very similar to that organ in our old friend
Amphioxus, that lowliest of all fishes, where it is nothing
but a slightly enlarged, contractile blood-vessel. In this The Bird in the Eg 473 latter creature there have been found as many as a hun-
dred and eighty pairs of gill-clefts, such a remarkable
number aerating the blood with but little necessary pro-
pulsion, but when in the higher fishes the number of gills
in many species is reduced to four, we realize at once the
need for a stronger engine to force the blood through the
lessened number, this accounting for the increased com-
plexity of the heart. plexity of the heart. Up to about the twelfth day the tiny foreshadowings of
bones are cartilaginous, like those of the shark, but at this
time real osseous, or bony, tissue begins to be deposited
in spots which spread rapidly. In the various portions of
the skull these bony centres spread until the bones are
separated only by narrow sutures, and in the adult bird
even these are obliterated, unlike the condition in the
skull of a cat or a dog. The bones of the adult bird are so neatly joined together,
and are so mutually dependent, that we might easily
imagine that they were formed in the order of size or
importance, or in a regular series, following their connection
with one another; but this is not true. The ribs, for
example, are formed between the segments of the primitive
sheets of muscle, independently of the back-bone, and
only later become attached to it. There is no trace of the
great keel-bone, or even of the sternum of the adult fowl,
until after the ends of the ribs have met in the middle line
of the body, when they grow together and give rise to the
sternum — a structure not found in fishes. The Bird in the Eg We have
learned that the repetition of similar structures (as the
ribs) is a sign of a low degree of organization, and the truth The Bird 474 of this is emphasized in the development of the embryo,
during which process a number of additional ribs dis-
appear. The abortive ribs of the neck-bones are especially
noticeable during the egg-life of the bird, so that in some
species we can make out traces of as many as fifteen ribs
all told. age 97 a short account was given of the origin of
the wings and feet, in the case
of the chick — from a primi-
tive fin-fold in some general-
ized aquatic ancestor. About
the fourth day of incuba-
tion, sections of our embryo
chick will show a low, round-
ed ridge, extending the whole
length from the neck to the
tail. While we can never be
absolutely certain that perfect
homology exists between the
two, yet it is very significant
that soon after its develop-
ment it dwindles away, leav-
ing four conical, isolated
he beginnings of the limbs of the bird. Within
hree days after the appearance of the limbs, faint
become visible upon the tips of the extremities,
e hints of the bones of fingers and toes, for such
, soon push out beyond the edge, still bound
by their transparent membrane, and for some
ey present the appearance of webbed paws or
arly embryo of Canada
howing fin-like limbs. . page 97 a short account was given of the origin of
the wings and feet, in the case
of the chick — from a primi-
tive fin-fold in some general-
ized aquatic ancestor. About
the fourth day of incuba-
tion, sections of our embryo
chick will show a low, round-
ed ridge, extending the whole
length from the neck to the
tail. While we can never be
absolutely certain that perfect
homology exists between the
two, yet it is very significant
that soon after its develop-
ment it dwindles away, leav-
ing four conical, isolated
the beginnings of the limbs of the bird. * This process is quite pronounced in the case of the Ostrich. The Bird in the Eg Within
three days after the appearance of the limbs, faint
become visible upon the tips of the extremities,
ese hints of the bones of fingers and toes, for such
are, soon push out beyond the edge, still bound
er by their transparent membrane, and for some
they present the appearance of webbed paws or
— Early embryo of Canada
, showing fin-like limbs. ount was given of the origin of
the wings and feet, in the case
of the chick — from a primi-
tive fin-fold in some general-
ized aquatic ancestor. About
the fourth day of incuba-
tion, sections of our embryo
chick will show a low, round-
ed ridge, extending the whole
length from the neck to the
tail. While we can never be
absolutely certain that perfect
homology exists between the
two, yet it is very significant
that soon after its develop-
ment it dwindles away, leav-
ing four conical, isolated
the limbs of the bird. Within
appearance of the limbs, faint
on the tips of the extremities,
es of fingers and toes, for such
beyond the edge, still bound
rent membrane, and for some
ppearance of webbed paws or On page 97 a short account was given of the origin of
the wings and feet, in the case
of the chick — from a primi-
tive fin-fold in some general-
ized aquatic ancestor. About FIG. 367. — Early embryo of Canada
Goose, showing fin-like limbs. FIG. 367. — Early embryo of Canada
Goose, showing fin-like limbs. The Bird in the Eg The Bird in the Eg 475 radiate fins. But as early as the tenth day, except for
the absence of feathers and claws, the limbs are, in appear-
ance, very perfect wings and feet. The most interesting
fact in connection with the limbs is that their develop-
ment begins superficially and works inward, not, as would
be thought, starting at the shoulder and ending at the
digits. Even the deep-seated shoulder- and thigh-girdles of bone
(pp. 85 and 89) are not derived from the axial skeleton. The former, in the long ago, was gradually pushed inward
from the surface by the deep-reaching rays of the fin-like
fore limbs, and it is believed that the pelvic girdle had
its origin in the spliced scales of some fish-like ancestor
of old, which had scales like those of some of the fossil
ganoids. These probably covered over the cartilage girdle
and then sunk in. An example of one out of many reptilian structures
which appear for a time and then vanish, is found in the
procoracoid bone which has apparently much to do with
the development of the typical coracoids, but which is
absent or reduced to a mere process in the adult bird.*
Strangely enough, in the embryo of the common chick
the coracoid and scapula fuse together at an early stage,
being then in a condition comparable only to that found
in the full-grown ostrich. Later this inexplicable fusion
is dissolved and the bones complete their development as
they began, — two wholly independent structures. they began, — two wholly independent structures. Again, in the embryo of a tern, faint vestiges of teeth The Bird 4 have been observed, instantly bringing to mind that some-
what gull-like, toothed bird of old — Icthyornis. what gull-like, toothed bird of old — Icthyornis. The origin and subsequent changes, in the embryo chick,
of the vascular system, including the heart, nerves, and
arteries, are more intricate than the development of any
other system of organs, and for an excellent reason. We
know that the frog's egg hatches as a tadpole, which breathes
by means of gills and lives, for a considerable time, in the
water. The Bird in the Eg We learned in Chapter IV that important parts
of the head and sense-organs of birds are derived from
metamorphosed gills ; so the inference is that all the changes
in the blood-channels, which in the tadpole and frog take
place during several months, are in the embryo chick
gone through with in a period of a few days. The blood in the heart of a fish is sent from the single
ventricle to the gills, and from there it is distributed all
over the body. In the gills it passes through the paired
series of red fringes and is oxygenated by the water. Now
in the chick there are six pairs of these gills, or paired
blood-vessels (although not more than three or four are
found at one time). The chick breathes by means of a
membranous sheet of blood-vessels spread out just beneath
the shell, and even the lungs are not brought into use until
just before the bird hatches. But strange to say, although
there is no water to supply the gill-channels with life-
giving oxygen, yet blood actually flows through them, in
obedience to the long-forgotten ancestral life-habits —
useless these many millions of years. Of all the gill-channels, but three remain in the adult
bird. The great aorta, which springs from the heart and . 368. — Pineal eye in Lizard
477 . 368. — Pineal eye in Lizard
477 . 368. — Pineal eye in Lizard The Bird turns to the right (in ourselves, the left-side gill-channel
forms this aortic arch), is what is left of the fourth pair
of aortic gill-arches, while the two arteries which, in aH
higher animals, leads to the right and left lungs, are the sel>
same channels which in the creatures of olden time encircbd
the sixth pair of gill-bars. F.E. the sixth pair of gill-bars. Although the eye of the bird is far superior to thaf of
a fish in seeing abiity,
yet in actual structure
there is not very nuch
difference, except that
the bird has gained
eyelids, tear^lands,
and a few other struc-
tures. Fishe-, frogs,
lizards, bins, and
mammals, tlrough all
the ages, Iwe depend-
ed on thesJ two eyes
and have found them
FIG. 369.— Pineal eye in Chick (P. E.). The Bird in the Eg all-sufficirfit; but there
are hin# that once,
long ago, the ancestors of all the higher animls had a sense-
organ, probably of sight, situated, like thafof the mythical
Polyphemus, in the centre of the head, In lizards this
vestigial organ is sometimes quite well Developed, having
a nerve which leads up from the centr of the brain to a
kind of translucent, lens-like scale wJch lies among the
other scales of the skin, upon the cerre of the forehead. In the long-extinct Ichthyosaurs this^edian eye was prob- F.E. FIG. 369.— Pineal eye in Chick (P. E.). FIG. 369.— Pineal eye in Chick (P. E.). long ago, the ancestors of all the higher animls had a sense-
organ, probably of sight, situated, like thafof the mythical
Polyphemus, in the centre of the head, In lizards this
vestigial organ is sometimes quite well Developed, having
a nerve which leads up from the centr of the brain to a
kind of translucent, lens-like scale wJch lies among the
other scales of the skin, upon the cerre of the forehead. In the long-extinct Ichthyosaurs this^edian eye was prob- The Bird in the Egg 479 ably functional. In an embryo chick of even the third
day this organ is remarkably prominent; but although
traces of it always remain, yet it fades away to a vestige. Look with a hand-lens at the head of a polywog, and
see the whitish dot between the eyes; or when you touch FIG. 370. — Forty-day embryo Ostrich, showing position in the shell. FIG. 370. — Forty-day embryo Ostrich, showing position in the shell. the "soft spot" on the head of a human baby, let it recall
the strange third eye which is its cause. And so we might continue to tell of the wonder of
embryo life: how up to the sixth day the little being
might be mistaken for the embryo of a reptile or a mammal,
but from this day onward the bird characteristics become
more and more noticeable. On the ninth day feathers The Bird 4 begin to be seen, looking, however, more like tiny cones
than anything else (Fig. 11). The muscles and the cartilage
skeleton are well defined on the fourteenth day, and about
this time the tiny beak with its white egg-tooth is pressed
against the membrane of the air-chamber at the large end
of the egg. The Bird in the Eg Reptiles also show this tiny bit of sharp lime
upon the head, which drops off soon after its function is
completed. Not until almost the last day is the mem-
brane pierced and the first gasp of air breathed into the
little lungs. By an instinctive moving of the head back
and forth the shell is filed through and cracked, and the
chick rolls out into the world, weak and helpless and for
a while absolutely dependent upon warmth and the care
of its mother, before it is fit for its future life. (Figs. 18
and 19.) Thus do all wild birds begin life, passing through
similar phases within the egg; and although we so often
admire a nest full of eggs, yet how seldom do we give
thought to the tiny creatures within, — their hearts even
at that very minute, perhaps, giving their first fluttering
beat! The instant that its eyes have cleared and its shaky
legs have gained strength to support its body, the chick
begins to use its senses and to store up experiences, taking
note of this sound and that taste, learning to fear or to
ignore, to flee or to pursue, to call or to remain silent. And thus does the brain of the chick and of all wild nestlings
begin to act and its psychological life commences, with
intermingled perceptions, instincts, and gleams of intelli-
gence. Here belong the making of nests and journeys, be The Bird 4 courtship and songs, the rearing of young birds, the avoid-
ing of enemies, the selection of food and suitable haunts,
and, lastly, the encountering and overcoming of dangers, —
new and wide-spread, — which are now affecting the environ-
ment of every creature of this world. Of greater impor-
tance than ever before is this adaptation to new con-
ditions; since man and his traps and his guns have come
upon the scene, upsetting all the world-old order of
Nature and slowly, surely, claiming the whole earth for
himself. May the naturalists of to-day realize their opportunity
and do their best to preserve to us and to posterity what
is left to us of wild life ! If not, let us pity the Nature-
lover of two hundred years hence! Appendix 4 EVOLUTION ORIGIN OP SPECIES. Charles Darwin. Appleton & Co., New York. Charles Darwin. Appleton & Co., New York. FROM THE GREEKS TO DARWIN. Henry Fairfield Osborn. Macmillan Co., New York. Henry Fa
ORGANIC EVOLUTION. Henry Fairfield Osborn. Macmillan Co., New York. ORGANIC EVOLUTION. M. M. Metcalf. Macmillan Co., New York. M. M. Metcalf. Macmillan Co., New York. VARIATION IN ANIMALS AND PLANTS. VARIATION IN ANIMALS AND PLANTS. H. M. Vernon. Henry Holt & Co., New York. VARIATION IN ANIMALS AND PLANTS. H. M. Vernon. Henry Holt & Co., New York. BOOKS FOR THE IDENTIFICATION OF NORTH AMERICAN BIRDS
GUIDE TO THE BIRDS OF NEW ENGLAND AND EASTERN NEW YORK. BOOKS FOR THE IDENTIFICATION OF NORTH AMERICAN BIRDS
GUIDE TO THE BIRDS OF NEW ENGLAND AND EASTERN NEW YORK. GUIDE TO THE BIRDS OF NEW ENGLAND AND EASTERN NEW YORK. Ralph Hoffmann. Houghton, Mifflin & Co., Boston. Ralph Hoffmann. Houghton, Mifflin & Co
HANDBOOK OF BIRDS OP EASTERN NORTH AMERICA. HANDBOOK OF BIRDS OP EASTERN NORTH AMERICA. F. M. Chapman. Appleton & Co., New York. HANDBOOK OF BIRDS OP EASTERN NORTH AMERICA. F. M. Chapman. Appleton & Co., New York. F. M. Chapman. Appleton & Co., New York. HANDBOOK OF BIRDS OP THE WESTERN UNITED STATES. Florence M. Bailey. Houghton, Mifflin & Co., Boston. Florence M. Bailey. Houghton, Mifflin &
KEY TO NORTH AMERICAN BIRDS (2 vols.). Elliot Coues. Dana Estes & Co., Boston. KEY TO NORTH AMERICAN BIRDS (2 vols.). Elliot Coues. Dana Estes & Co., Boston. Elliot Coues. Dana Estes & Co., Boston. BIRDS OF NORTH AND MIDDLE AMERICA (3 parts; others to follow). Robert Ridgway. Bulletin of the United States National Museum,
No. 50, Washington, D. C. No. 50, Washington, D. C. HISTORY OF NORTH AMERICAN LAND BIRDS (3 vols.). Baird, Brewer, and Ridgway. Little, Brown & Co., Boston. NORTH AMERICAN LAND BIRDS (3 vols.). Baird, Brewer, and Ridgway. Little, Brown & Co., Boston. 483 483 BIRDS IN GENERAL DICTIONARY OF BIRDS. Alfred Newton. A. & C. Black, London. RIVERSIDE NATURAL HISTORY, Vol. IV. Birds. L. Stejneger. Houghton, Mifflin & Co., New York. Abdominal ribs, 79 Abdominal ribs, 79 Abdominal ribs, 79
Adaptation of feet, 361, 362 Adaptation of feet, 361, 362
Adaptative Radiation, 15-18; in war-
blers, 361-367 blers, 361-367
Adjutant, head of, 273, 276; hair
from neck of, 276 INDEX Figures in heavy-faced type indicate illustrations. B from neck of, 276
^Epyornis, com. size of egg of, 461
Aftershaft, 36 ^Epyornis, com. size of egg of, 461
Aftershaft, 36 ^Epyornis, com. Aftershaft, 36 ^Epyornis, com. size of egg of, 461
Aftershaft, 36 Back-bone, of Amphioxus, 66; evolu-
tion of, 64-70; of shark, 68
Barbs, of Condor's feather, 32
Barbicels, 32, 34
Barbules, 32, 34
Bats, keel of, 83-84; used as food, 158
Beak, see Bill
Beaks and Bills, 223-251
Bear, feet of, 102
Bellbird, wattles of, 273
Bill,
function of, 223, 224, 250, 251
of Avocet, 237, 239; Archaeopteryx, Back-bone, of Amphioxus, 66; evolu-
tion of, 64-70; of shark, 68
Barbs, of Condor's feather, 32
Barbicels, 32, 34
Barbules, 32, 34
Bats, keel of, 83-84; used as food, 158
Beak, see Bill
Beaks and Bills, 223-251
Bear, feet of, 102
Bellbird, wattles of, 273
Bill,
function of, 223, 224, 250, 251
of Avocet, 237, 239; Archaeopteryx, Aggressive coloration, use of, 308;
in Arctic fox, 309; in Cuckoo, 311;
in Gyrfalcon, 309, 312; in Ivory Gull,
309, 311; in Penguin, 309, 310;
in pickerel, 310; in Snowy Owl, 309,
313 313
Air-sacs, cf. B with respiratory system
of insects, 173: extent of, 173;
function of, 174, 177; in Prairie Hen,
177 Albatross, wing of, 320, 321, 325, 332
Albinism, 314 Albatross, wing of, 320, 321, 325, 332
Albinism, 314 function of, 223, 224, 250, 251
of Avocet, 237, 239; Archaeopteryx,
226; Cormorant, 227, 228; Cross-
bill 24, 249, 245; Crow, 226;
Shoveller Duck, 233, 235; Purple
Finch, 249; Golden Eagle, 242;
Flamingo, 128, 234, 235, 236;
Gannet, 227, 228; Boat-billed
Heron, 237, 238; Great Blue Heron,
237; Night Heron, 237; Huiabird,
248, 249, 250; Hummingbirds, 244,
245, 246, 247, 248; Ibis, 237, 239;
Shell Ibis, 240; Merganser, 233,
234; Nuthatch, 245; Owls, 242;
Oyster-catcher, 238, 240; Parrots,
242; Pelican, 228, 229, 230;
Crook-billed Plover, 240, 241;
American Raven, 225; Black Skim-
mer, 231, 232, 236; Snakebird,
228, 229; Dowitcher Snipe, 241;
Spoonbill, 220, 236, 239; Stilt, 237;
Chimney Swift, 244, 246, 245;
Tailor-bird, 245; Tern, 231, 232;
Toucans, 243, 244; Triceratops,
485 Alligator, egg of, 429; foot of, 354;
nictitating membrane of, 215; re-
lation to birds, 9; skull of, 105 Alligator, egg of, 429; foot of, 354;
nictitating membrane of, 215; re-
lation to birds, 9; skull of, 105
Altricial nestling, 30 Alligator, egg of, 429; foot of, 354;
nictitating membrane of, 215; re-
lation to birds, 9; skull of, 105
Altricial nestling, 30
Amoeba, 185, 186, 463
Amphioxus, 66; gill-clefts in, 473;
notochord of, 66; segments of, 78;
trachea of, 169; breathing motions
in, 180
Anaximander, 12
Ancestors of birds, 1-18
Aorta, 476
Apoplexy in birds, 202
Apteryx, ody-feathers of, 289; eyes
in, 254, 255, 256; sense of touch in,
219
Archseopteryx, as parallel branch, 10;
as ancestral type, 10; foot of, 353;
general description of, 7; in Berlin
Museum, n; in British Museum, 8;
probable habits of, 12-13; restora-
tion of, 14; tail of, 398, 399, 400, 401
Aristotle, cf. with Darwin, 12 Amphioxus, 66; gill-clefts in, 473;
notochord of, 66; segments of, 78;
trachea of, 169; breathing motions
in, 180 Apoplexy in birds, 202
Apteryx, ody-feathers of, 289; eyes
in, 254, 255, 256; sense of touch in,
219 Archseopteryx, as parallel branch, 10;
as ancestral type, 10; foot of, 353;
general description of, 7; in Berlin
Museum, n; in British Museum, 8;
probable habits of, 12-13; restora-
tion of, 14; tail of, 398, 399, 400, 401
Aristotle, cf. Apoplexy in birds, 202
Apteryx, ody-feathers of, 289; eyes
in, 254, 255, 256; sense of touch in,
219 Figures in heavy-faced type indicate illustrations. Artery, 182
Arteries, course of vertebral, 79
Atlas of Jabiru, 72
Auk, wing of Great, 339, 340; wing of
razor-billed, 339, 340
Avpcet, bill of, 237, 239
Axis of Jabiru, 72
B MISCELLANEOUS BIRDS OF ESSEX COUNTY, MASSACHUSETTS. C. W. Townsend. Memoir of Nuttall Ornithological Club, No. III. Cambridge, Mass. (A type of a local bird-study )
THE WOODPECKERS. Fannie H. Eckstorm. Houghton, Mifflin & Co., Boston. (A popu-
lar study of a single group of birds.) BIRDS OF ESSEX COUNTY, MASSACHUSETTS. C. W. Townsend. Memoir of Nuttall Ornithological Club, No. III. Cambridge, Mass. (A type of a local bird-study )
THE WOODPECKERS. Fannie H. Eckstorm. Houghton, Mifflin & Co., Boston. (A popu-
lar study of a single group of birds.) BIRDS OF ESSEX COUNTY, MASSACHUSETTS. C. W. Townsend. Memoir of Nuttall Ornithological Club, No. III. Cambridge, Mass. (A type of a local bird-study )
THE WOODPECKERS. Fannie H. Eckstorm. Houghton, Mifflin & Co., Boston. (A popu-
lar study of a single group of birds.) B Coloration, aggressive, see Aggressive
coloration; protective, see Protec-
tive coloration
Columella, of Owl, no, 109
Comb, of Condor, 271; Domestic Cock,
274; Heron, 387, 388; on toe of
Chuck-will's-widow, 369
Condor, feathers of, 36; flight of, 324,
326; wattles of, 271
Condyles of skull, 108
Coot, foot of, 387
Coracoid, function of, 84; in man, 85;
in reptiles, 84
Cord, spinal, 202
Cormorant, bill of, 227, 228; character
of egg-shell of, 460; gullet of, 133;
iris of, 256; method of fishing of, 154;
tail feathers of, 400
Corpuscles, see Blood
Courlan, food of, 153
Cuckoo, aggressive coloration of, 311;
colour of eggs of English, 456, 458;
foot of, 372
Curassow, Banded, crest of, 261
Crab, used as food, 149
Crane,
Crowned, crest of, 264; head of, 252,
265
Demoiselle, crest of, 267, 295 Classification, based on toes, 354, 355
Clavicle, function of, 85; of Hoatzin, 86; Blackbird, down on nestling, 26
Blood, circulation of, 182, 183, 184;
compared with amoeba, 185; red
corpuscles of, 184, 185; white cor-
puscles of, 185 ion,
Clavicle, function of, 85; of Hoatzin, 86;
in mammals, 86
Claws of foot, 368 Claws of foot, 368
three on wing of, Osprey, 322; Ostrich,
338, 339; Swan, 322 puscles of, 185
Bobolink, cause of colour change in
spring, 297 338, 339; Swan, 322
Climate, effect on plumage of, Bob-
white, 292, 295; Song Sparrow, 292,
295; South American Pipit, 295, 296;
Turkey Vulture, 322; White-throated
Sparrow, 291, 294; Wood Thrush, 294 338, 339; Swan, 322
Climate, effect on plumage of, Bob-
white, 292, 295; Song Sparrow, 292,
295; South American Pipit, 295, 296;
Turkey Vulture, 322; White-throated
Sparrow, 291, 294; Wood Thrush, 294 spring, 297
Bob-white, effect of climate on, 293
295; moult of, 43 295; moult of, 43
Body, of Herons, 286; Petrels, 286 Body, of Herons, 286; Petrels, 286
Body-feathers, of Apteryx, 289; Cas-
sowary, 288; Snowy Egret, 305,
322; Emeu, 36, 289; Ostrich 236,
288; Scaled Partridge, 289; Pen-
guin, 289; Rock-jumper, 288;
Snake-bird, 289 Sparrow, 291, 294; Wood Thrush, 294
Cockatoo, feet of, 371; Leadbeater,
crest of, 260
Cocoon, as food, 146 Snake-bird, 289
Boltenia, notochord of, 67, 67 Bones, of embryo chick, 473; hollow-
ness of, 175, 176; of mammal skull,
103; relation to flight, 176; of skull,
107 Breast-bone, 79-84; evolution of, 79 Breast-bone, 79-84; evolution of, 79
Breast ornament of Wild Turkey, 280 Breast ornament of Wild Turkey, 280
Brain, of bird compared with that of
crocodile, 200; with that of Tricera-
tops; 200; with that of Walrus. B Coloration, aggressive, see Aggressive
coloration; protective, see Protec-
tive coloration
Columella, of Owl, no, 109
Comb, of Condor, 271; Domestic Cock,
274; Heron, 387, 388; on toe of
Chuck-will's-widow, 369
Condor, feathers of, 36; flight of, 324,
326; wattles of, 271
Condyles of skull, 108
Coot, foot of, 387
Coracoid, function of, 84; in man, 85; 486
226; Woodcock, 219, 221, 222;
Woodpeckers, 245
Blackbird, down on nestling, 26 Chuck-will's-widow, comb on toe o(, 369
Circulatory system of pigeon, 183
Classification, based on toes, 354, 355 Chuck-will's-widow, comb on toe o(, 369
Circulatory system of pigeon, 183
Classification, based on toes, 354, 355
Clavicle, function of, 85; of Hoatzin, 86;
in mammals, 86
Claws of foot, 368
three on wing of, Osprey, 322; Ostrich,
338, 339; Swan, 322
Climate, effect on plumage of, Bob-
white, 292, 295; Song Sparrow, 292,
295; South American Pipit, 295, 296;
Turkey Vulture, 322; White-throated
Sparrow, 291, 294; Wood Thrush, 294
Cockatoo, feet of, 371; Leadbeater,
crest of, 260
Cocoon, as food, 146
Cod, eggs of, 428
Collar-bone, see Clavicle
Colour,
of birds, 287; of young birds, 316, 317;
blindness, 301; breast of Bleeding
Heart Pigeon, 306, 308
change in moulting of Bald Eagle,
296; Scarlet Tanager, 294, 297;
Siberian Black Lark, 295, 297
of eggs, 430
of feathers, 53-61 ; causes of, 54-56;
patterns of, 56-60
in mammals, 287; phases, double, 314;
relation to haunts, 296-314; use of,
322 . B with Darwin, 12 Archseopteryx, as parallel branch, 10;
as ancestral type, 10; foot of, 353;
general description of, 7; in Berlin
Museum, n; in British Museum, 8;
probable habits of, 12-13; restora-
tion of, 14; tail of, 398, 399, 400, 401
Aristotle, cf. with Darwin, 12 486 Index Chuck-will's-widow, comb on toe o(, 369
Circulatory system of pigeon, 183
Classification, based on toes, 354, 355
Clavicle, function of, 85; of Hoatzin, 86;
in mammals, 86
Claws of foot, 368
three on wing of, Osprey, 322; Ostrich,
338, 339; Swan, 322
Climate, effect on plumage of, Bob-
white, 292, 295; Song Sparrow, 292,
295; South American Pipit, 295, 296;
Turkey Vulture, 322; White-throated
Sparrow, 291, 294; Wood Thrush, 294
Cockatoo, feet of, 371; Leadbeater,
crest of, 260
Cocoon, as food, 146
Cod, eggs of, 428
Collar-bone, see Clavicle
Colour,
of birds, 287; of young birds, 316, 317;
blindness, 301; breast of Bleeding
Heart Pigeon, 306, 308
change in moulting of Bald Eagle,
296; Scarlet Tanager, 294, 297;
Siberian Black Lark, 295, 297
of eggs, 430
of feathers, 53-61 ; causes of, 54-56;
patterns of, 56-60
in mammals, 287; phases, double, 314;
relation to haunts, 296-314; use of,
322 . B 200;
embryo, thirty-six hours old, 472;
great size in birds, 199, 200, 201;
importance of, 198; nerves of, 202;
protection of, 196 protection of, 196
Bulk, birds of largest, 285 Bulk, birds of largest, 285
Bustard, chin-feathers of Great, 267;
weight of. 285 weight of. 285
Butterfly, eggs of, 428; torn by birds, 147 weight of. 285
Butterfly, eggs of, 428; torn by birds, 1 Butterfly, eggs of, 428; torn by birds, 147
Buzzard, Black-breasted, feeding on
Emeu eggs, 159 Butterfly, eggs of, 428; torn by birds, 147
Buzzard, Black-breasted, feeding on
Emeu eggs, 159 Canals, semicircular, 218
Caracara, head of, 271, 272; foot of, 378
Carp, compared with Scaled Partridge,
289, 290
Carrion Hawk, Chimango, food of, 162
Cassowary,
body feathers of, 288
eggs of, 436, 453; colour of, 453, 454;
character of shell of, 460
helmet of, 275, 288; lack of tail of,
402; tracks of, 396; wing of, 321,
337, 338
Catbird, use of wing in young. 322
Caterpillar, used as food by birds, 146
Cells, of blood, see Blood; of feathers,
34; of muscle, see Muscle
Cerebellum, 201
Chewink, foot of, 367
Chicken, stomach glands of, 135 Canals, semicircular, 218 Canals, semicircular, 218 Canals, semicircular, 218
Caracara, head of, 271, 272; foot of, 378 Carp, compared with Scaled Partridge,
289, 290 Carrion Hawk, Chimango, food of, 162
Cassowary,
body feathers of, 288
eggs of, 436, 453; colour of, 453, 454;
character of shell of, 460
helmet of, 275, 288; lack of tail of,
402; tracks of, 396; wing of, 321,
337, 338
Catbird, use of wing in young. 322
Caterpillar, used as food by birds, 146
Cells, of blood, see Blood; of feathers,
34; of muscle, see Muscle
Cerebellum, 201
Chewink, foot of, 367
Chicken, stomach glands of, 135 Index of adult Crested Screamer, 29;
Duck Hawk, 27. of
Duck, colour of eggs of, 447; oddities
of diet of, 160-163; eggs of, 447;
food of, 147; crest of Mandarin,
263; nest of Mallard, 447, 448;
position of feet in, 382, 389, 390;
Side-wheel, see Steamer Duck; bill
of Shoveller, 233, 234; gradual loss
of flight in Steamer, 337; tail-feathers
of, 400; tail of Blue, 406; tail of
Pintail, 415 Crane, trachea of, 170, 171 of adult Crested Screamer, 29;
Duck Hawk, 27. E Eagle, Bald, iris of eye of, 256; foot
of Golden, 374, 376; foot of Harpy,
375; Golden, 242; Harpy, 262, 264;
Hyoid of, 114; nictitating membrane
of, 214; sight of, 208
Ear, canals of , 218; in owls, 216 217;
structure of, 217, 218
Eclipse plumage, 48, 49
Edible birds' nest, see Swiftlet. Eggs,
abnormal number laid by African
Waxbill, 444; domestic Hen, 444;
Flicker, 444 Eagle, Bald, iris of eye of, 256; foot
of Golden, 374, 376; foot of Harpy,
375; Golden, 242; Harpy, 262, 264;
Hyoid of, 114; nictitating membrane
of, 214; sight of, 208 of, 214; sight of, 208
Ear, canals of , 218; in owls, 216 217;
structure of, 217, 218
Eclipse plumage, 48, 49
Edible birds' nest, see Swiftlet. Eggs,
abnormal number laid by African
Waxbill, 444; domestic Hen, 444;
Flicker, 444 264, 266; Woodpe
Crests, use of, 258 Crests, use of, 258
Crocodile, brain of, 200; gizzard of, 138;
gullet of, 134; heart of, 181 of ^Epyornis, 461; alligator, 429;
Anna Hummingbird, 443; Archae-
opteryx, 13; butterflies, 428; Cali-
fornia Partridge, 438; Cassowary,
436, 453 gullet of, 134; heart of, 181
Crop, 127-134; of Caracara, 134;, capa-
city of, in Wood Pigeon, 130; ejection
of food-pellets from, 132, 133; of
English Sparrow, 129; extreme devel-
opment of, in Pigeon, 130; in Hoatzin,
130, 131; oil in Petrel's, 131 436, 453
character of surface of, in Casso-
wary, 460; Cormorant, 460; Duck,
460; North and South African
Ostriches, 460; Tinamou, 460;
Mute and Whooping Swans, 460;
Woodpeckers, 460 130, 131; oil in Petrel's, 131
Crossbill, bill of, 245, 248, 249 Crossbill, bill of, 245, 248, 249
Cross-fertilization by birds, 144, 145 Cross-fertilization by birds, 144, 145
Crow, brain of, compared with that of
Hesperornis, 6; Fish Crow, stealing
eggs of herons, 448, 449; foot of,
356; method of feeding on shell-fish,
153; pterylosis of nestling, 39 Cross-fertilization by birds, 144, 145
Crow, brain of, compared with that of
Hesperornis, 6; Fish Crow, stealing
eggs of herons, 448, 449; foot of, of cod, 428
coloration of eggs, 430; of Casso-
wary, 453, 454; causes of, 458, 459;
Doves, 446; Ducks, 447: English
Cuckoo, 456, 458; Goose, 448;
Grouse, 447; Herons, 448; Long-
billed Marsh Wren, 446; Murres,
456; Nighthawk, 454, 455; Ostrich,
453, 454; Owls, 451; Parrots,
445; Pelicans, 448; Sea-birds,
455; Short-billed Marsh Wren, 446;
Skimmer Black, 454, 455, 456, 459;
Tern, 454, 456. E 457; compared
with stilt, 452; variability of, 456,
457, 459; Woodpecker, 444 356; method of feeding on shell
153; pterylosis of nestling, 39 356; method of feeding on shell
153; pterylosis of nestling, 39 of
Duck, colour of eggs of, 447; oddities
of diet of, 160-163; eggs of, 447;
food of, 147; crest of Mandarin,
263; nest of Mallard, 447, 448;
position of feet in, 382, 389, 390;
Side-wheel, see Steamer Duck; bill
of Shoveller, 233, 234; gradual loss
of flight in Steamer, 337; tail-feathers
of, 400; tail of Blue, 406; tail of
Pintail, 415 Crane, trachea of, 17
Cranium, see Skull Cranium, see Skull
Creeper, Brown, protective coloration
of, 303, 304; food of, 150; tail of,
410, 411 Cranium, see Skull
Creeper, Brown, protective coloration
of, 303, 304; food of, 150; tail of,
410, 411 410, 411
Crest of, California Partridge, 259;
Cock-of-the-Rock, 270; Condor, 271;
Crowned Crane, 264, 265; Crowned
Pigeon, 260, 261; Banded Curassow,
261; Demoiselle Crane, 265, 267;
Domestic Cock, 274; Double-crested
Pigeon, 270; Eared Pheasant, 268;
Harpy Eagle, 262, 264; Hooded
Merganser, 263; Hummingbirds, 267;
India Peacock, 259; Java Peacock,
258; Kingbird, 260; Ruby-crowned
Kinglet, 260; King of Saxony Bird
of Paradise, 269, 270; Laughing
Thrush, 283; Leadbeater Cockatoo,
260; Mandarin Duck, 263; Night
Heron, 259; Plumed Partridge, 259;
Six-shafted Bird of Paradise, 267;
Snowy Egret, 260; Umbrella-bird,
264, 266; Woodpeckers, 264 Falcon, Peregrine, see Duck Hawk. chick, 22; forty-day Ostrich, 479
gastrula stage of, 466, 468; gill-clefts,
476; hatching of, 480; hatching of
egg-tooth of, 480; hatching of
Ostrich Chicks, 481; illustrating
evolution, 470, 471; limbs of, 97;
lung in, 178, 179; method of study-
ing, 462, 471; muscles of, 480;
muscle-plates of, 69, 70; pineal
eye in chick, 478,479; precoracoid,
475; respiration of, 476; ribs, 473,
474; segmentation of Chick, 466,
468; segmentation of frog, 465,
466, 467; sixth-day chick com-
pared with reptile and mammal,
479; tarsus of, 99; teeth in tern,
476; third day in egg, 463 Falcon, Peregrine, see Duck Hawk. Family, habits of Warbler, Black-and-
white Warbler, 366; Black-throated
Green Warbler, 166; Magnolia, 366;
Maryland Yellow-throat, 362; Myrtle
Warbler, 364; Ovenbird, 366, 375;
Pine Warbler, 366; Redstart, 363,
364; Yellow Palm Warbler, 363;
Water Thrushes, 364, 365; Worm-
eating Warbler, 363 eating Warbler, 363
Fat, of Penguins, 286; Petrels, 286
Feather, aftershaft of, 36; barbicels of, Fat, of Penguins, 286; Petrels, 286
Feather, aftershaft of, 36; barbicels of,
32, 34; barbs of, 34; barbules of, 32,
34; cells of, 34; divisions on wing,
320; growth of Ostrich, 28; model
of structure of, 33; pattern on, 58-61;
structure of, 31-38 structure of, 31-38
Feathers, arrangement of, 38-40; in
tail of Archseopteryx, 398, 399
400, 401; in modern bird, 399, 400
of Cassowary, 338; colour of, 53-61;
of Condor, 36, 336; development
of, 17-38; downy condition of. 35;
of embryo bird, 479, 480; of structure of, 31-38
Feathers, arrangement of, 38-40; in
tail of Archseopteryx, 398, 399
400, 401; in modern bird, 399, 400 476; third day in egg, 463
thirty-six hours, 472; brain in, 472;
eyes, 472; false vertebrae, 472; thirty-six hours, 472; brain in, 472;
eyes, 472; false vertebrae, 472;
heart, 472
twelfth-day, 473 thirty-six hours, 472; brain in, 472;
eyes, 472; false vertebrae, 472;
heart, 472 400, 401; in modern bird, 399, 400
of Cassowary, 338; colour of, 53-61;
of Condor, 36, 336; development
of, 17-38; downy condition of. 35;
of embryo bird, 479, 480; of 400, 401; in modern bird, 399, 400
of Cassowary, 338; colour of, 53-61;
of Condor, 36, 336; development
of, 17-38; downy condition of. D Darwin, evolutionary theory of, 12
Decorations, use of, 322
Deer, nostrils of, 204; spots in young,
316, 317, tail of, 413
Dermis, 21
Dinosaur, relation to birds, 9; thigh-
girdle of, 90; tracks of, 396, 397
Distribution of birds, 361
Dogfish, see Shark
Donkey, foot of, 39, 395
Dove, colour of eggs, 446; nest of
Mourning, 439; position of eyes in,
252, 253, 254
Down, development of, 23; from head
of young Bobolink, 26; magnified
from young Song Sparrow, 27;
models of development of, 24, 25; Darwin, evolutionary theory of, 12
Decorations, use of, 322 Darwin, evolutionary the
Decorations, use of, 322 Deer, nostrils of, 204; spots in young,
316, 317, tail of, 413
Dermis, 21 316, 317,
Dermis, 21 Dinosaur, relation to birds, 9; thigh-
girdle of, 90; tracks of, 396, 397
Distribution of birds, 361 girdle of, 90; tracks of, 396, 397
Distribution of birds, 361 girdle of, 90; tracks of
Distribution of birds, 361 Dogfish, see Shark Dogfish, see Shark
Donkey, foot of, 39, 395
Dove, colour of eggs, 457, 459; Woodpecker, 444
comparative size of eggs of Hum-
mingbird, 461 Donkey, foot of, 39, 395
Dove, colour of eggs, 446; nest of
Mourning, 439; position of eyes in,
252, 253, 254 mingbird, 461
of Emeu, devoured by Buzzard, 159;
Fowl, 429, 436 252, 253, 254
Down, development of, 23; from head
of young Bobolink, 26; magnified
from young Song Sparrow, 27;
models of development of, 24, 25; of frog, 428, 465; development of, 465,
466, 467; for embryonic study, 462 466, 467; for embryonic study, 462
of Grebe, 432; Hairy Woodpecker, 445; Index Eggs,
of Environment, relation of birds to, 480. D 482
Epiclavicle of fish, 86 ggs,
of Hen, development of, 466; for
embryonic study, 462; structure of,
464 Epiclavicle of fish, 86
Evolution, of bill of Fla Epiclavicle of fish, 86
Evolution, of bill of Flamingo, 235, 236;
bill of Skimmer, 231, 232, 236;
breast-bone, 79; Class of birds, 15;
colour patterns, 58-61; embryo chick,
470, 471; foot, 353, 358, 361; gill-
arches, 114, 115; gizzard, 138; heart,
180, 181; history of theories of, 12;
lung, 178, 179, 180; Penguin's wing,
341; ribs, 78; shoulder-girdle, 86,
87; skull, 104-106; tail, 398 399,
400, 402, 403; tree (of birds and
reptiles), 10; warblers, 361-367;
wings, 91-97
Eye, of Hummingbird, 436; Jellyfish, 427;
Junco, 434; Killdeer Plover, 431,
432; Mallard Duck, 447; Mourn-
ing Dove, 439; Murre, shape of,
number laid by, average bird, 433; 43i
number laid by, average bird, 433;
Doves, 440; Game-birds, 438;
Gulls, 436; Hawks, 440; Hum-
mingbirds, 443; Jungle Fowl, 444;
Xing Penguin, 435; Moas, 435;
.Nighthawk, 442; Ostrich, 433;
Owl, 440; Parrots, 441; Peacock
Pheasant, 440; Rhea, 433; Sea-
birds, 435; Shore-birds, 436; Wad-
ing birds, 438; Water-birds, 436,
437, 438 Apteryx, 254, 255, 256; compared
with camera, 207, 208; iris of, 210
lashes of Hornbill, 257; Ostrich, 257;
Seriema, 257
lids of, 214, 215; Brown Thrasher, with camera, 207, 208; iris of, 210
lashes of Hornbill, 257; Ostrich, 257;
Seriema, 257
lids of, 214, 215; Brown Thrasher, 437, 438
relation of number to danger,433-444;
relative size compared with bird,
460, 461
of reptiles, 428, salmon, 427; show-
ing relation of orders, 452; skate,
428, 429; snake, 429 lids of, 214, 215; Brown Thrasher,
212, 213; Woodcock, 221, 256
mammals, 209, 254; nocturnal birds, 212, 213; Woodcock, 221, 256
mammals, 209, 254; nocturnal birds,
254
pineal eye in, embrvo chick, 478, 479; 460, 461
of reptiles, 428, salmon, 427; show-
ing relation of orders, 452; skate,
428, 429; snake, 429 pineal eye in, embrvo chick, 478, 479;
child, 479; lizard, 477, 478; polly-
wog, 479
position of, in Dove, 252, 253, 254; 428, 429; snake, 429
Egg- tooth, of embryo chick, 480;
reptiles, 480 wog, 479
position of, in Dove, 252, 253, 254;
Owl, 252, 253, 254 reptiles, 480
Egret, crest of Snowy, 315, 322; neck
curves of, 73; use of wing of, 350
Embryo,
aorta, 476, 478
development of limbs, 474, 475; of
girdles, 475; of three toes, 475
drawing of third-day embryo, 469
feathers of, 479, 480; twelve-day
chick, 22; forty-day Ostrich, 479
gastrula stage of, 466, 468; gill-clefts, structure of, 207-210; thirty-six-
hour embryo, 472
Expression of face of birds, 252 Expression of face of birds, 252 Expression of face of birds, 252 35;
of embryo bird, 479, 480; of twelfth-day, 473
Emeu, body-feathers of, 239; feather
of, 36; lack of tail in, 402, 404
Emeu-wren, tail of, 415 twelfth-day, 473
Emeu, body-feathers of, 239; feather
of, 36; lack of tail in, 402, 404
Emeu-wren, tail of, 415 Index 489 Feathers (continued),
Emeu, 36; of embryo of twelfth-
day, 22; moult of, 40-53
number of, in tail, of Archaeopteryx,
400; of Cormorant, 400; of Duck,
400; of Ostrich, 402; of Peafowl,
402; of Fantail Pigeon, 402 Food, affecting structure, 164; indu-
cing intoxication, 163; of Apteryx,
147; Buzzard, 159; Courlan, 153;
Creepers, 150, Crows, 153; Ducks,
147, 153; Bald Eagle, 148; Gulls,
161; Carrion Hawk, 162; Duck
Hawk, 159; New Zealand King-
fisher, 161; Locust-birds, 150-152;
Osprey, 155; Barn Owl, 158; Bur-
rowing Owl, 162; Elf Owl, 158;
Strenuous Owl, 158; Oyster-catcher,
153; Kea Parrot, 161; Penguins,
153; Ptarmigan, 144; Raven, 158;
Road-runner, 157; Sea-birds, 148;
Sea-eagles, 157; Secretary-bird, 157;
Red-winged Starlings, 162, 163;
Storks, 157; Sugar-birds, 144; Sul-
phur Tyrant, 161 ; whales, 148
Food-pellets of Owl, 132, 133 Food, affecting structure, 164; indu-
cing intoxication, 163; of Apteryx,
147; Buzzard, 159; Courlan, 153;
Creepers, 150, Crows, 153; Ducks,
147, 153; Bald Eagle, 148; Gulls,
161; Carrion Hawk, 162; Duck
Hawk, 159; New Zealand King-
fisher, 161; Locust-birds, 150-152;
Osprey, 155; Barn Owl, 158; Bur-
rowing Owl, 162; Elf Owl, 158;
Strenuous Owl, 158; Oyster-catcher,
153; Kea Parrot, 161; Penguins,
153; Ptarmigan, 144; Raven, 158;
Road-runner, 157; Sea-birds, 148;
Sea-eagles, 157; Secretary-bird, 157;
Red-winged Starlings, 162, 163;
Storks, 157; Sugar-birds, 144; Sul-
phur Tyrant, 161 ; whales, 148
Food-pellets of Owl, 132, 133 day, 22; moult of, 40-53
number of, in tail, of Archaeopteryx,
400; of Cormorant, 400; of Duck, umber of, in tail, of Archaeopteryx,
400; of Cormorant, 400; of Duck, 400; of Cormorant, 400; of Duck,
400; of Ostrich, 402; of Peafowl, 400; of Ostrich, 402; of Pe
402; of Fantail Pigeon, 402 402; of Fantail Pigeon, 402
of Ostrich, 35, 336; papillae of nestling,
21 ; powder-downs, 37, 38; ptery-
losis, 39; sheaths of, 26; sheaths
of young Kingfisher, 29; texture of,
287-289; function of, in Puff-back
Shrike, 287; of wing, 320-352;
worn-out Hummingbird's breast, 44 worn-out Hummingbird's breast, 44
Feather-ears, of owl, 267, 268 worn-out Hummingbird's breast, 44
Feather-ears, of owl, 267, 268 worn-out Hummingbird's br
Feather-ears, of owl, 267, 268 Feather-ears, of owl, 267, 268
Feather-tips, in Bobolink, 297; in
Black Lark, 295, 297; in Snow-
flake, 297; in English Sparrow, 52,
53; wearing off of, 52, 53, 297 Red-winged Starlings, 162, 163;
Storks, 157; Sugar-birds, 144; Sul-
phur Tyrant, 161 ; whales, 148 phur Tyrant, 161 ; whales, 14
Food-pellets of Owl, 132, 133
Foot, claws of, 368; tendons 53; wearing off of, 52, 53, 297
Feeding, method of, in Flamingo, 128;
in Woodcock, 222 Food-pellets of Owl, 132, 133
Foot, claws of, 368; tendons of, 191,
192; of alligator, 354; Archseop-
teryx, 353; bear, 102; of bird cf. with
man, 99, 100; Caracara, 378; Casso-
wary, 394, 396; Chewink, 367; Chuck-
will's- widow, 369; Cockatoo, 371;
Coot, 387; Crow, 356; Cuckoo, 372;
Donkey, 394, 395; Duck, 382, 389,
390; Golden Eagle, 374, 376; Harpy
Eagle, 375; Flam i go, 388; Fly-
catchers, 360; Gallinule 385, 386,
387; Grebe, 391; Ruffed Grouse, 380^
381; Rough-legged Hawk, 389; Heron ,. 387,388; Hummingbirds, 368; Wood
Ibis, 102, 385; Kangaroo, 394, 395;
Horned Lark, 367; European House
Martin, 381: Nuthatch, 359; Oriole,
360; Osprey, 375; Ostrich, 394, 395,
396; Burrowing Owl, 373; Snowy
Owl, 375; Owls, 373, 374; Parrots, 372;
Passeres, 358, 359; Peacock, 383;
Brown Pelican, 355; Penguin, 391,
392; Pheasant, 380; Phalarope, 384;
Pipit, 367, 368; Plover, 384; Ptar-
migan, 381; Quail, 380; Raven, 356;
Rhinoceros-bird, 357; Road-runner,
373; Rock-jumper, 393; Sand-grouse,
383; Semipalmated Sandpiper, 384;
Secretary-bird, 378, 379; Snakebird,
391; Swallow, 360; Black-necked
Swan, 390; Swift, 368, 369; Vulture,
377; Woodpeckers, 370 Feet, adaptation of, 361, 362; classifi-
cation based on, 354, 355; evolu-
tion of, 353, 358, 361 tion of, 353, 358, 361
Femur, 98
Fibula, 98 Femur, 98
Fibula, 98 Femur, 98
Fibula, 98 Fighting, method of, in Ruff, 277, 279,
280 Finch, 143; bill of Purple, 249; colour
change in Purple, 293 change in Purple, 293
Fins, origin of paired, 96
Fish, used as food, 153; hear change in Purple, 293
Fins, origin of paired, 96 Fins, origin of paired, 96
Fish, used as food, 153; heart of, 181;
muscle-flakes of, 78 muscle-flakes of, 78
Fishing, method of, in Cormorants, 154;
Frigate-birds, 154; Herons, 156;
Kingfishers, 154; Ospreys, 154; Peli-
cans, 152; Penguins, 154; Snake-
birds, 154; Terns, 154 birds, 154; Terns, 154
Flamingo, bill of, 128, 234, 235; colour
change in, 293; foot of, 388; neck of,
73, 281; sternum of, 80; tongue of,
126, 127; trachea of, 168, 169 126, 127; trachea of, 168, 169
Flicker, abnormal number of eggs laid
by, 444; hyoid of, 123, 124; protec-
tive colouring in young, 302, 303, 304;
tongue of, 123, 124 tongue of, 123, 124
Flight, cf. Fox, aggressive coloration in Arctic,
309 with swimming, 327; Condor,
324, 326; Steamer Duck, 337; cer-
tain Flycatchers, 324, 326; Gold-
finches, 329; Grebes, 341; Gulls, 328;
Hummingbirds, 329; Owl Parrot, 333;
Pheasant, 323, 324; Crested Screamer,
329; Sparrows, 328; swiftness of, in
Pigeon, 350; swiftness of, in Swallow,
352; of Tinamou, 333-336; of Wood-
peckers, 329; of Vultures, 328, 330,
33i, 332 377; Woodpeckers, 370
Fowl, egg of, 429, 430; Japanese Long-
tailed, 417, 419; number of eggs laid
by Jungle, 444; spurs of, 382; tail of
Jungle, 418 377; Woodpeckers, 370
Fowl, egg of, 429, 430; Japanese Long-
tailed, 417, 419; number of eggs laid
by Jungle, 444; spurs of, 382; tail of
Jungle, 418 Fox, aggressive coloration in Arctic,
309 33i, 332
Flipper, of Penguin, 341, 342, 343
Flycatcher, feet of, 360; flight of, 324, Framework, 62-102 Framework, 62-102 Framework, 62-102
Frigate-bird, method of feeding of, 154
Frog, egg of, 428; fresh egg of, 462; Flipper, of Penguin, 341, 342, 343
Flycatcher, feet of, 360; flight of, 324,
326; tail of Least, 412 Framework, 62-102
Frigate-bird, method of feeding of, 154
Frog, egg of, 428; fresh egg of, 462; Index 490 490
gastrula of egg of, 466; segmentation
of egg ot, 467; used as food, 157 H Hatching of embryo Chick, 480; Os-
trich, 481 onic, 480
Limbs, origin of, 96, 97; evolution of,
96, 97; of embryo chick, 474, 475
Lobed toes, of coots, 387; of phalaropes, Ibis, 239, 237; feet of Wood, 385;
Indian Wood, beautiful feathers of, 61 ;
Wood, resting upon whole foot, 102;
Shell, 240 96, 97; of embryo chick, 474, 475
Lobed toes, of coots, 387; of phalaropes,
384
ocust birds, method of feeding, 150-
152; relation of habits to food, 152
Loon, in winter haunts, 6; neck of, 277
Lory, tongue of 125 Ibis, 239, 237; feet of Wood, 385;
Indian Wood, beautiful feathers of, 61 ;
Wood, resting upon whole foot, 102;
Shell, 240
Ichthyornis, lower jaw, 5; restored
skeleton of, 3; structure of, 4
Icthyosaurus, eye-plates of, 213
Ilium, 89, 90; function of, 89; of bull-
frog, 91
Ingersoll, Ernest, quoted, 433
Insects, use as food of birds, 148-152
Intestines, function of, 141; length in
Hummingbird, 140; length in Ostrich,
140; in alligator, 141
Iris of eye, 210; of Bald Eagle, 256;
Cormorant, 256; Owl, 256; Puffin,
256; Towhee, 256; Vireo, 256
Ischium, 89, 90
Jabiru, atlas and axis of, 72
Jaguar, standing on toes, 101
Jelly-fish, eggs of, 427
Junco, eggs of, 434; nest of, 434; tail
of, 414
K
Kangaroo, feet of, 394, 395
Keel, in Flamingo, 80; Hoatzin, re-
pressed by crop, 132; model com-
paring various, Albatross, Humming-
bird, Pigeon, 83; value in classifica-
tion of, 82
Kingbird, crest of, 260
Kingfisher, method of fishing of, 154;
nest of, 444; New Zealand, food of,
101
Kinglet, Ruby-crowned, crest of, 260
Lacrymal glands, 211
Lamprey, gill-basket of, 112
Lark, foot of horned, 367; tail of
meadow, 414; cause of colour change
in Siberian black, 295, 297 Lory, tongue of 125
Lungs, evolution of, 178, 179, 180;
character of, 174, 175; cf. with those
of chameleon, 177, 178 Lyre-bird, tail of, 420, 421 Lyre-bird, tail of, 420, 421 M Mallard, eclipse plumage of, 48. 49;
tongue of, 122; trachea of, 168;
syrinx drum of, 168, 170
Mammals, used as food, 159
Mankind vs. birds, 18
Maoris, legends of, 13
Martin, feet of European House, 381
Merganser, bill of, 233, 234; crest of
Hooded, 263
Mimicry, of Hawk by Cuckoo, 311, 312; Jabiru, atlas and axis of, 72
Jaguar, standing on toes, 101
Jelly-fish, eggs of, 427
Junco, eggs of, 434; nest of, 434; tail
of, 414
K
Kangaroo, feet of, 394, 395
Keel, in Flamingo, 80; Hoatzin, re-
pressed by crop, 132; model com-
paring various, Albatross, Humming-
bird, Pigeon, 83; value in classifica-
tion of, 82
Kingbird, crest of, 260
Kingfisher, method of fishing of, 154;
nest of, 444; New Zealand, food of,
101
Kinglet, Ruby-crowned, crest of, 260 G with those
of chameleon, 177, 178
Lyre-bird, tail of, 420, 421
M
Mallard, eclipse plumage of, 48. 49;
tongue of, 122; trachea of, 168;
syrinx drum of, 168, 170
Mammals, used as food, 159
Mankind vs. birds, 18
Maoris, legends of, 13
Martin, feet of European House, 381
Merganser, bill of, 233, 234; crest of
Hooded, 263
Mimicry, of Hawk by Cuckoo, 311, 312;
of Hawk by Hawk, 312
Moa, cf. with Ostrich, 13; number of
eggs laid by, 435
Monkeys, toes of, 358
Motmot, protective coloration of, 303;
tail of Mexican, 424, 425, 426
Moult, 40-53; of Black Grouse, 48;
of Bob-white, 43; of Brown Pelican,
41; causes for, 42-50; of body-
feathers of English Sparrow, 52, 53;
of wing-feathers of English Sparrow,
46; of feathers, 41; of feather-tips,
52, 53; of Mallard Duck, 47, 48, 49;
of mound-builders, 44, 45; of Pen-
guin, 51; of Ptarmigan, 48, 50, 51, 52;
of reptiles, 40; of Robin, 43; in Thomas, quoted, 2
Hyoid, 111; of eagle, 114 Life in the egg, 462-479; post-embry-
onic, 480 K Kangaroo, feet of, 394, 395 Motmot, protective coloration of, 303;
tail of Mexican, 424, 425, 426 Kangaroo, feet of, 394, 395
Keel, in Flamingo, 80; Hoatzin, re-
pressed by crop, 132; model com-
paring various, Albatross, Humming-
bird, Pigeon, 83; value in classifica-
tion of, 82 Moult, 40-53; of Black Grouse, 48;
of Bob-white, 43; of Brown Pelican,
41; causes for, 42-50; of body-
feathers of English Sparrow, 52, 53;
of wing-feathers of English Sparrow,
46; of feathers, 41; of feather-tips,
52, 53; of Mallard Duck, 47, 48, 49;
of mound-builders, 44, 45; of Pen-
guin, 51; of Ptarmigan, 48, 50, 51, 52;
of reptiles, 40; of Robin, 43; in
spring, 45; time of, 42-47
Mouse, used as food, 157 Kingbird, crest of, 260
Kingfisher, method of fishing of, 154;
nest of, 444; New Zealand, food of,
101 101
Kinglet, Ruby-crowned, crest of, 260 Kinglet, Ruby-crowned, crest of, 260 Mimicry, of Hawk by Cuckoo, 311, 312;
of Hawk by Hawk, 312 Jelly-fish, eggs of, 427
Junco, eggs of, 434; nest of, 434; tail
of, 414
K of Hawk by Hawk, 312
Moa, cf. with Ostrich, 13; number of
eggs laid by, 435 G G Hawk Duck, food of, 158; head of, 211;
feet of Rough-legged, 389; mimicked
by Cuckoo, 311, 312; number of
eggs laid by, 440; skull of, compared
with that of Heron, 199, 200 Gallinule, foot of, 385, 386, 387 Gallinule, foot of, 385, 386, 387
Game-birds number of eggs laid by, 438 Game-birds number of eggs laid by, 438
Gannet, bill of, 227, 228 Game-birds number of eggs laid by, 438
Gannet, bill of, 227, 228 Heads and necks, 252-284 Gannet, bill of, 227, 228
Gastrula, 466, 468; compared with
sponge, 469, 470 Heads and necks, 252-284
Head of, Adjutant, 276: Apteryx, 254;
256; Barn Owl, 253; Bell-bird, 273,
Caracara, 271, 272; Eared Pheasant,
267, 268; Condor, 271; Crowned
Crane, 252, 265; Domestic Cock, 274;
Dove, 253; Duck Hawk, 211; Java
Peacock, 258; King Vulture, 272,
273; Seriema, 257; Sloth, 210;
Wild Turkey, 273, 275 sponge, 469, 470
Gill-arches, of chick, 114; evolution of,
114, 115; ultimate distribution of , in
chick, 114, 115 chick, 114, 115
Gill-bars of shark, 113 Gill-bars of shark, 113
Gill-basket of lamprey, 112 Gill-basket of lamprey, 112
Gill-clefts in Amphioxus, 473; embryo
chick, 476; fishes, 473 chick, 476; fishes, 473
Giraffe, neck-vertebrae of, compared with
those of bird, 73, 75
Girdles, embryonic development of, 475 Heart, beats of that of bird, 182;
chambers of, 182, 184; of croco-
dile, 181; of fish, 181; of embryo
36 hours old, 472; evolution of, 180,
181; position of, in vertebrates, 181
Helmet of Cassowary, 275, 288; Horn-
bill, 278
Hemispheres, cerebral, 200
Hen, abnormal number of eggs laid by
domestic, 444 Heart, beats of that of bird, 182;
chambers of, 182, 184; of croco-
dile, 181; of fish, 181; of embryo
36 hours old, 472; evolution of, 180,
181; position of, in vertebrates, 181
Helmet of Cassowary, 275, 288; Horn-
bill, 278 Girdles, embryonic development of, 475
Gizzard, 134-140; change in structure
of, 137, 138; function of, 135; of
Fruit Pigeons, 138; of Hornbills, 139,
140 Gizzard, 134-140; change in structure
of, 137, 138; function of, 135; of
Fruit Pigeons, 138; of Hornbills, 139,
140
stones of Cassowary, 136; crocodiles,
138; extinct Moas, 136, 137
of pigeon shown by X-ray, 63
Glottis of nestling Robin, 166; Peli-
can, 1 66
Goat-sucker, West African, wing orna-
ments of, 349
Goldfinch, flight of, 329; tongue of, 125 181; position of, in vertebrates, 181
Helmet of Cassowary, 275, 288; Horn-
bill, 278 140
stones of Cassowary, 136; crocodiles,
138; extinct Moas, 136, 137
of pigeon shown by X-ray, 63
Glottis of nestling Robin, 166; Peli-
can, 1 66
Goat-sucker, West African, wing orna-
ments of, 349
Goldfinch, flight of, 329; tongue of, 125 stones of Cassowary, 136; crocodiles,
138; extinct Moas, 136, 137
of pigeon shown by X-ray, 63 Hemispheres, cerebral, 200
Hen, abnormal number of eggs laid by
domestic, 444 Hen, abnormal number of eggs laid by
domestic, 444 of pigeon shown by X-ray, 63
Glottis of nestling Robin, 166; Peli-
can, 1 66 domestic, 444
Heron, a still hunter, 156; Boat-billed,
237, 238; body of, 386; cause of
downiness in feather of, 35; colony
of Great Blue, 450; colour of eggs
of, 448; comb on toe of, 387, 388;
divisions of feathers in wing of young
of, 320; Great Blue, 237; in sleep,
344; Night, 237; Night, crest of,
259; skull of, compared with that
of Hawk; 199, 200; standing on
toes, 101; tail of Green, 413; use
of wing in young Green, 323; wing
of Great White, 321 can, 1 66
Goat-sucker, West African, wing orna-
ments of, 349 ments of, 349
Goldfinch, flight of, 329; tongue of, 125
Goose, Canada, embryo of, 474; colour Goldfinch, flight of, 329; tongue of, 125
Goose, Canada, embryo of, 474; colour
of eggs of, 448; muscle of skin in, 188;
sense of hearing in, 216; Spur-wing,
wing of, 346, 347; use of wing in
young Canada, 322
Crackle, tail of Boat-tailed, 418
Grebe, eggs of, 432; flight of, 341; foot
of, 391 Goldfinch, flight of, 329; tongue of, 125
Goose, Canada, embryo of, 474; colour
of eggs of, 448; muscle of skin in, 188;
sense of hearing in, 216; Spur-wing,
wing of, 346, 347; use of wing in
young Canada, 322 young Canada, 322
Crackle, tail of Boat-tailed, 418 Crackle, tail of Boat-tailed, 418
Grebe, eggs of, 432; flight of, 341; foot
of, 391 of, 391
Grouse, Black, moult of, 48; colour of
eggs of, 447; Ruffed, ruff of, 277;
Ruffed, use of wing of, 547; toes of
Ruffed, 380, 381 Hesperornis, habits and structure, 3-6;
restoration of, Frontispiece Hesperornis, habits and structure, 3-6;
restoration of, Frontispiece Hoatzin, clavicles of, 86; crop of, 130,
131; keel repressed by crop, 132 Hoatzin, clavicles of, 86; crop of, 130,
131; keel repressed by crop, 132 Ruffed, 380, 381
Guinea Fowl, Vulturine, evolution of
colour pattern on wings of, 58, 59,
61 131; keel repressed by crop, 132
Honey Creeper, tongue of, 127, 129 131; keel repressed by crop, 132
Honey Creeper, tongue of, 127, 129 Honey Creeper, tongue of, 127, 129
Hornbill, casque of, 275, 278; eye-
lashes of, 257; feeding mate from
its gizzard, 137, 138 Honey Creeper, tongue of, 127, 129
Hornbill, casque of, 275, 278; eye-
lashes of, 257; feeding mate from
its gizzard, 137, 138
Horse, toes of, 358 Gullet, of Cormorant, 133; crocodile, 134
Gull, change in gizzard of, 137, 138; Gull, change in gizzard of, 137, 138;
flight of, 328; food of, 161; Her-
ring, wing of, 328 its gizzard, 137, 1
Horse, toes of, 358 Hudson, quoted, 329, 333
Huia Bird, bill of, 248, 24 ring, wing of, 328
Ivory, aggressive coloration in, 309,
311; protective coloration in 309, Humidity, effect on plumage of, Bob
White, 293, 295; Song Sparrow, 292,
295; White-throated Sparrow, 291,
294; Wood Thrush, 294 3"
Laughing, nest of, 437; protective
coloration of, 309, 312; number of
eggs laid by, 436 295; White-throated S
294; Wood Thrush, 294
Hummingbird, Anna, nest eggs laid by, 436
Gyrfalcon, aggressive coloration in, 309, 294; Wood Thrush, 294
Hummingbird, Anna, nest of, 443; eggs
of, 443; bills of, 244, 245, 246, 247, Index 49 248; crests of, 267; eggs of, 436, 461;
feet of, 368; flight of, 329; flight of,
compared with that of insect, 165;
length of intestines in, 140; nest of
Ruby-throated, 449; number of eggs
laid by, 443; tongue of, 127; wing
of, 320, 321, 325; wing-strokes of,
82; worn-out breast-feather of, 44;
Huxley, definition of palaeontology, 2;
Thomas, quoted, 2
Hyoid, 111; of eagle, 114 Layers of cells in embryo, 470 248; crests of, 267; eggs of, 436, 461;
feet of, 368; flight of, 329; flight of,
compared with that of insect, 165;
length of intestines in, 140; nest of
Ruby-throated, 449; number of eggs
laid by, 443; tongue of, 127; wing
of, 320, 321, 325; wing-strokes of,
82; worn-out breast-feather of, 44;
Huxley, definition of palaeontology, 2;
Thomas, quoted, 2
Hyoid, 111; of eagle, 114 Layers of cells in embryo, 470
Leg, correlation with neck in Flamingo,
281; in Swan, 282; framework of,
98-102; function of, 98; of Ostrich,
100; skeleton of Ostrich, 99; human,
99; scales of, 368; spurs on, 382, 383;
of Bald Eagle, 375; of Cassowary,
394; of Flamingo, 388; of Golden
Eagle, 375; of Secretary-bird, 378,
379 Leg, correlation with neck in Flamingo,
281; in Swan, 282; framework of,
98-102; function of, 98; of Ostrich,
100; skeleton of Ostrich, 99; human,
99; scales of, 368; spurs on, 382, 383;
of Bald Eagle, 375; of Cassowary,
394; of Flamingo, 388; of Golden
Eagle, 375; of Secretary-bird, 378,
379
Life in the egg, 462-479; post-embry-
onic, 480
Limbs, origin of, 96, 97; evolution of,
96, 97; of embryo chick, 474, 475
Lobed toes, of coots, 387; of phalaropes,
384
ocust birds, method of feeding, 150-
152; relation of habits to food, 152
Loon, in winter haunts, 6; neck of, 277
Lory, tongue of 125
Lungs, evolution of, 178, 179, 180;
character of, 174, 175; cf. N Owl, aggressive coloration of Snowy,
309, 313; cause of downiness in
feathers, 35; colour phases of Screech,
314; eggs of Screech, 441; food of
Barn, 158; food and feeding habits
of Burrowing, 162; food of Elf, 158;
food of Strenuous, 158; food-pel-
lets ejected by, 132, 133; foot of
Burrowing, 373; foot of Snowy, 375;
iris of eye of, 256; nest of Burrow-
ing, 442; orbit cf Barred, 255; posi-
tion of eyes in Barn, 252, 253, 254;
sight of Barred, 211 Neck, of Flamingo, 281; Heron, 281;
Loon, 277; Snake-bird, 282, 283;
Swan, 282 Swan, 282
Neck vertebrae, cf. with Giraffe, 73, 75;
cf. with man, 72; cf. with reptile,
73 73
Nerves, action of, 198; cerebral, 202 Nerves, action of, 198; cerebral, 202
Nervous system, 196, 198; of Pigeon,
197; reflex, 197, 198 197; reflex, 197, 198
Nest, of Mourning Dove, 439; Mallard
Duck, 447, 448; Laughing Gull, 437;
Great Blue Herons, 450; Humming-
birds, 448; Anna Hummingbird,
443; Ruby-throated Hummingbird,
449; Junco, 434; Kingfisher, 444;
Nighthawk, 455; Ostrich, 453, 454;
Burrowing Owl, 442; California Part-
ridge, 438; Black Skimmer, 454,
450, 4595 Tern, 453, 456, 457; Hairy
Woodpecker, 444, 445 sight of Barred, 211
Owls, bills of, 242; colour of eggs of,
451; foot of, 373, 374; number of
eggs laid by, 440
Ovary of fowl, 464
Ovenbird, 365, 366 sight of Barred, 211
Owls, bills of, 242; colour of eggs of,
451; foot of, 373, 374; number of
eggs laid by, 440
Ovary of fowl, 464
Ovenbird, 365, 366 eggs laid by, 440
Ovary of fowl, 464
Ovenbird, 365, 366 443; Ruby-throated Hummingbird,
449; Junco, 434; Kingfisher, 444;
Nighthawk, 455; Ostrich, 453, 454;
Burrowing Owl, 442; California Part-
ridge, 438; Black Skimmer, 454,
450, 4595 Tern, 453, 456, 457; Hairy
Woodpecker, 444, 445 Ovary of fowl, 464
Ovenbird, 365, 366 Oyster-catcher, bill of, 238, 240; food
of, 153 Oyster-catcher, bill of, 238, 240; food
of, 153 wing in youn
Osteology, 62 Osprey, food of, 155, 156; foot of, 347;
method of fishing of, 154; use of
wing in young, 322; wing of, 351 Woodpecker, 444, 445
Nictitating membane, of alligator, 215;
Eagle, 214; human eye, 215; Brown
Thrasher, 213 Woodpecker, 444, 445
Nictitating membane, of alligator, 215;
Eagle, 214; human eye, 215; Brown
Thrasher, 213 Paradise, King of Saxony Bird of, 276;
269; Six-shafted Bird of, 267, 269,
Superb Bird of, 277; Twelve-wired
Bird of, 348
Parallelism, in feeding habits, 163, 164 Paradise, King of Saxony Bird of, 276;
269; Six-shafted Bird of, 267, 269,
Superb Bird of, 277; Twelve-wired
Bird of, 348
Parallelism, in feeding habits, 163, 164
Paramecium, 463; keels, 83, 84
Parrakeet, tail of Grass, 408 Nighthawk, colour of eggs of, 454, 455;
nest of, 455; number of eggs laid
by, 442; protective coloration in,
296, 301 Parallelism, in feeding habits, 163, 164
Paramecium, 463; keels, 83, 84
Parrakeet, tail of Grass, 408 296, 301
Nostrils, of bird, 204; of daer, 204
Notochord, of Amphioxus, 66; of 296, 301
Nostrils, of bird, 204; of daer, 204
Notochord, of Amphioxus, 66; of Paramecium, 463; keels, 83,
Parrakeet, tail of Grass, 408 Parrot, bill of, 242; colour of eggs of, 445;
flight of Owl, 333; food of Kea, 161;
foot of, 372; number of eggs laid by,
441; use of wing in, 322
Partridge, California, crest of, 259; nest Notochord, of Amphioxus, 66; of
Boltenia, 67 Notochord, of Amphioxus, 66; of
Boltenia, 67 Boltenia, 67
Nuthatch, bill of, 245; foot of, 359 Boltenia, 67
Nuthatch, bill of, 245; foot of, 359 Nuthatch, bill of, 245; foot of, 359
Nutrition, 116-141 Nuthatch, bill of,
Nutrition, 116-141 Nuthatch, bill of,
Nutrition, 116-141 441; use of wing in, 322
Partridge, California, crest of, 259; nest
and eggs of, 438; Plumed, crest of,
259; Scaled, feathers of, 289, 290
Passeres, foot of, 358, 359 O O Lamprey, gill-basket of, 112
Lark, foot of horned, 367; tail of
meadow, 414; cause of colour change
in Siberian black, 295, 297 Index 492 492
189; cf. with reptiles, 193, 194;
of embryo chick, 480; energy of, in
Hummingbird, 83; impressions on
bones, 194; pectoral, 190; of skin
in Goose, 188; of skin in Penguin,
188; structure of, 193; of wing and
breast of Pigeon, 190; of wing in
Owl Parrot, 333 479; eyelashes of, 257; leg of, 100:
length of intestines, 140; neck and
vertebrae of, 69, 71; nest of, 453, 454;
number of eggs laid by, 433; pro-
tective position of, 306, 307; skele-
ton of leg, 99; specific difference in
eggs, 460; sternum of, 81; tail of,
402; toes of, 358, 394, 395, 396:
wing of, 321, 337, 338, 339 N R Rabbit, tail of, 413
Radius, 92
Ratitae, 82
Raven, bill of American, 225: feeding
on ostrich-eggs, 158; foot of, 356
Redstart, life-habits of, 363, 364
Reptiles, causes of colour in, 56; eggs
of, 428, 429; trachea of, 169
Respiration of embryo chick, 476
Rhea, number of eggs laid by, 433
Rhinoceros-bird, foot of, 357
Ribs, 77-79; abdominal, of Archaeop-
teryx and of reptiles, 79; embryo,
473, 474; function of, 77 79;
Hatteria Lizard, 77; Screamer, 77;
significance of numerous, 78 Pigment, colour caused by, 54, 55 Pigment, colour caused by, 54, 55
Pipit, effect of climate on, 295, 296;
foot of, 367, 368 Raven, bill of American, 225: feeding
on ostrich-eggs, 158; foot of, 356
Redstart, life-habits of, 363, 364 foot of, 367, 368
Phalarope, feet of, 384
Pheasant, Argus, colour on ostrich-eggs, 158; foot of, 356
Redstart, life-habits of, 363, 364
Reptiles, causes of colour in, 56; eggs Phalarope, feet of, 384
Pheasant, Argus, colour pattern of, 58;
Eared, head of, 267, 268; feet of, 380;
flight of, 323, 324; number of eggs
laid by Peacock, 440; ruff of Lady
Amherst, 277, 278; spurs of Peacock,
383; tail of Peacock, 419, 420;
tail of Reeves, 416, 418; wing of, 327 Redstart, life-habits of, 363, 364
Reptiles, causes of colour in, 56; eggs
of, 428, 429; trachea of, 169
Respiration of embryo chick, 476
Rhea, number of eggs laid by, 433
Rhinoceros-bird, foot of, 357 Ribs, 77-79; abdominal, of Archaeop-
teryx and of reptiles, 79; embryo,
473, 474; function of, 77 79;
Hatteria Lizard, 77; Screamer, 77;
significance of numerous, 78 tail of Reeves, 416, 418; wing of, 327
Phororachus, skull of, 16; structure of,
13, 14 13, 14
Plantain-eater, green pigment in, 55 Plantain-eater, green pigment in, 55
Ploughshare-bone, 400, 402, 403 Plantain-eater, green pigment in, 55
Ploughshare-bone, 400, 402, 403 Ploughshare-bone, 400, 402, 403
Plover, Crook-billed, 240, 241; feet of,
384; nest and eggs of Killdeer, 431,
432
Pollen, used as food, 144, 145
Powder-down, 38; of Great White Heron,
3? significance of numerous, 78
Road-runner, food of, 157; foot of,
373; tail of, 413 Ploughshare-bone, 400, 402, 403
Plover, Crook-billed, 240, 241; feet of,
384; nest and eggs of Killdeer, 431, 373; tail of, 413
Robin, glottis of nestling, 166; moult
of, 43; .spots in young, 316, 317 432
Pollen, used as food, 144, 145
Powder-down, 38; of Great White Heron,
3? Sandgrouse, feet of, 383
Sandpipers, protective coloration in,
299; semipalmated, feet of, 384 O 188; tail of Black-footed,
406; wing of, 321, 341, 342, 343 in, 299; of eggs, 44
Proventriculus, 135 Proventriculus, 135
Ptarmigan, feet of, 381; food of, 144;
moult of, 48, 50, 51, 52; protective
coloration in, 298, 300, 309
Pterodactyls, 2, 9; weight of, 285
Pterylosis, 39, 40; of nestling Crow, 39
Pubis, 89, 90; in embryo bird, 90
Purlin, iris of eye of, 256 Ptarmigan, feet of, 381; food of, 144;
moult of, 48, 50, 51, 52; protective
coloration in, 298, 300, 309 406; wing of, 321, 341, 342, 343
Pepper, effect on plumage of, 292 Pterodactyls, 2, 9; weight of, 285
Pterylosis, 39, 40; of nestling Crow, 39
Pubis, 89, 90; in embryo bird, 90
Purlin, iris of eye of, 256 Pepper, effect on plumage of, 292
Petrel; body of, 286; ejecting oil from
crop, 131 crop, 131
Pickerel, aggressive coloration in, 310 Pickerel, aggressive coloration in, 310
Pigeon, Blood-breasted, 306, 308; cir-
culatory system of, 183; Crowned,
crest of, 260, 261 ; Double-crested, 270;
extreme development of crop of, 130;
gizzard of Fruit, 138; nervous system
of, 197; number of eggs laid by, 440;
symptoms of flight in, 350, 352; tail
of, 402; tail of Fan-tail, 404; wings
and breast-muscles of, 190; wing-
feather of, 35; X-ray photograph
of, 63 R of, 43; .spots in young, 316, 317
Rock-jumper, body-feathers of, 288;
feet of, 393 Powder-down, 38; of Great White Heron,
Prairie hen, use of air-sacs in, 117
Precocial nestling, 30 Ruff of, Lady Amherst Pheasant, 277,
278; Ruff, 277, 279, 280; Ruffed
Grouse, 277; Superb Bird of Paradise,
277
Ruff, feather cloak of, 277, 279, 280 Prairie hen, use of air-sacs
Precocial nestling, 30
Precoracoid in embryo, 475 Prairie hen, use of air-sacs in, 117
Precocial nestling, 30
Precoracoid in embryo, 475 Precoracoid in embryo, 475 Primaries, moult of, in Bob- white, 43;
English Sparrow, 46; Mallard, 47;
Robin, 43 Ruff, feather cloak of, 277, 279, 280 Pigment, colour caused by, 54, 55 O 259; Scaled, feathers of,
Passeres, foot of, 358, 359
Peacock, feather of, 20; Ind Peacock, feather of, 20; India, crest of,
259; Java, crest of, 258; ribs of, 78;
spurs of, 382, 383; tail of, 422, 423;
train of, 422, 423
Peafowl, tail of, 402 Peacock, feather of, 20; India, crest of,
259; Java, crest of, 258; ribs of, 78;
spurs of, 382, 383; tail of, 422, 423;
train of, 422, 423
Peafowl, tail of, 402
Pectoral girdle, see Shoulder-girdle. train of, 422, 423
Peafowl, tail of, 402 Peafowl, tail of, 402 Osprey, food of, 155, 156; foot of, 347;
method of fishing of, 154; use of
wing in young, 322; wing of, 351 Pectoral girdle, see Shoulder-girdle. Pelican, bill of, 228, 229, 330; c Pectoral girdle, see Shoulder-girdle. Pelican, bill of, 228, 229, 330; colour
of eggs of, 448; downy stage of, 54;
foot of, 355; full-grown, 59; glottis
of, 166; half-grown, 57; method of
fishing of, 152; nestling of Brown, 21;
newly hatched, 30; tail of, 408, 409;
tongue of, 120, 121 Osteology, 62
Ostrich, body-feathers of, 288; cause of
downiness in feather, 35; colour of
eggs, 453, 454; compared with Moa,
13; cross-section of wing-bone, 175;
eggs of, 436, 453; embryo (40-day), Index migan, 298, 300, 309; Quail, 299,
300; Sandpipers, 299; Sparrows, 299;
Black-necked Swan, 305, 306; Sooty
Tern, 300, 303; Thayer's experiments
in, 299; of eggs, 446-459
Proventriculus, 135 Pelvic girdle, see Thigh-girdle. Pelvic girdle, see Thigh-girdle. Pelvis, compared with that of reptiles,
74; vertebra? in, 74 74; vertebra? in, 74
Penguin, aggressive coloration in. 309,
310; body-leathers of, 289; eggs of,
435; fat of, 280; food of, 153; method
of fishing of, 154; moult of, 51; skin-
muscles in. Q Quadrate, 109, 110; in reptiles, 111 Quadrate, 109, 110; in reptiles, 111
Quail, feet of, 380; protective colora-
tion in, 299, 300
Quezal, sham tail of, 421, 422 Quadrate, 109, 110; in reptiles, 111
Quail, feet of, 380; protective colora-
tion in, 299, 300
Quezal, sham tail of, 421, 422 Quadrate, 109, 110; in reptiles, 111
Quail, feet of, 380; protective colora-
tion in, 299, 300 tion in, 299, 300
Quezal, sham tail of, 421, 422 Robin, 43
Primaries, of Albatross, 320, 321, 325;
Cassowary, 321, 337, 338; Hum-
mingbird, 320, 321, 325; Ostriches,
321; Penguins, 321 ^ Salivary glands, original function of,
117, 118; in Swiftlets, 119; Swifts,
119; Woodpeckers, 118
Salmon, eggs of, 427
Sandgrouse, feet of, 383
Sandpipers, protective coloration in,
299; semipalmated, feet of, 384 Salivary glands, original function of,
117, 118; in Swiftlets, 119; Swifts,
119; Woodpeckers, 118
Salmon, eggs of, 427
Sandgrouse, feet of, 383 321; Penguins, 321 ^
Protective coloration, in Brown Creeper,
303, 304; Flickers, 302, 303, 304;
Ivory Gull, 309, 311; Laughing Gull,
304, 306; Motmot, 303; Nighthawk,
296, 301; Ostrich, 306, 307; Ptar- Sandgrouse, feet of, 383
Sandpipers, protective coloration in,
299; semipalmated, feet of, 384 Index 494 Sapsucker, tongue of, 124 Snipe, bill of Dowitcher, 241; fighting,
see Ruff. Snowshoes of Grouse, 380, 381
Snowflake, colour of change in, 297 Snipe, bill of Dowitcher, 241; fighting,
see Ruff. Snowshoes of Grouse, 380, 381 Sapsucker, tongue of, 124
Scales, of carp, 289, 290; on legs of
birds, 356, 368 Snowshoes of Grouse, 380, 381 Snowflake, colour of change in, 297 birds, 356, 368
Scapula, 84; position in various ani-
mals, 88 Sparrow, albinism in English 314;
crop of English, 129; effect of cli-
mate on song, 292, 295; effect of
humidity on White-throated, 291,
294; flight of, 328; moult of wing-
leathers of, 46; moult of body-
feathers of, 52, 53; neck of White-
throated, 74, protective coloration
in, 299; stomach-glands of, 135;
tail in moult of, 409; tail of Vesper,
414 mals, 88
Scent-glands, of birds, 286, 287; of
mammals, 286 mammals, 286
Screamer, Crested, 330; flight of: 329 Screamer, Crested, 330; flight of: 329
Sea-birds, colour of eggs of, 455 Screamer, Crested, 330; flight of: 329
Sea-birds, colour of eggs of, 455 Sea-birds, colour of egg
Sea-eagles, food of, 157 Sea-eagles, food of, 157
Sea-urchins, used as food, 145 Sea-urchins, used as food, 145
Secondaries, number in Albatross, 320,
321, 325; in Hummingbird, 320, 321,
325 325
Secretary-bird, 378, 379; food of, 157
Seeds, 427 Specialization of feet, 358 Seeds, 427
Segmentation of egg, 465-470
Senses, Spoonbill, bill of, 236, 239; use of wing
in a, 350 Segmentation of egg, 465-470
Senses,
hearing, 215-218; in Geese, 216
sight, 207-215; in Barred Owl, 211;
Eagle, 208: Woodcock, 208
smell, 203-206; in Vultures, 205
taste, 218, 219
touch, 219, 222; in Apteryx, 219;
Woodcock, 219, 220, 222 Spurs of fowls, 382; of Peacock, 382,
383; Peacock Pheasant, 383; struc-
ture of, 382
Squid, used as food, 150, 153
Starling, food of Red-winged, 162, 163; Spurs of fowls, 382; of Peacock, 382,
383; Peacock Pheasant, 383; struc-
ture of, 382 Squid, used as food, 150, 153 Starling, food of Red-winged, 162, 163;
Wattled, see Locust-bird Wattled, see Locust-bird
Sternum, function of, 81; value in
classification of, 80; of Flamingo, 80;
Ostrich, 81
Stilt, 237 Sternum, function of, 81; value in
classification of, 80; of Flamingo, 80;
Ostrich, 81
Stilt, 237
Stomach, 134-140; of Chicken (young),
135; English Sparrow, 135; Snake-
bird, 139
Storks, food of, 157
Sugar-bird, food of, 144
Sulphur-tyrant, food of, 161
Swallow, food of, 360; tail of Barn, 405,
407 Woodcock, 219, 220, 222
Seriema, 17; eyelashes of, 257; habits
of, 15; relation to Phororachus, 15 of, 15; relation to Phororachus, 15
Shark, back-bone of, 68; gill-bars of,
113; skull of, 104; tooth of, 20 of, 15; relation to Phororachus, 15
Shark, back-bone of, 68; gill-bars of,
113; skull of, 104; tooth of, 20 Shore-birds, number of eggs laid by,
436 Shoulder-girdle, 84-88, 85; bones com-
posing, 84; evolution of, 86, 87;
of fish, 86, 87 of fish, 86, 87
Shrike, Puff-back, feathers in, 287 Sulphur-tyrant, food of, 161
Swallow, food of, 360; tail of Barn, 405,
407 Sulphur-tyrant, food of, 161
Swallow, food of, 360; tail of Barn, 405,
407 Shrike, Puff-back, feathers in, 287
Size, as correlated with distribution,
295 Swan, foot of Black-necked, 390;
neck of, 282; protective coloration
in Black-necked, 305, 306; sleeping
position of, 345; specific difference in
egg-shells, 460; trachea of Trumpeter,
170; weight of Trumpeter, 285;
wing-bone of Black, 175; use of
wing in defence, 346
Swift, bill of Chimney, 244, 245, 246; Skate, egg of, 428, 429 Skate, egg of, 428, 429
Skeleton, of man, leg of, 99; Ostrich,
leg of, 99; Pigeon, shown by X-ray
photograph, 63; Rooster, compared
with contour of body, 65; ways of
preparing, 64 preparing, 64
Skimmer, bill of, 231, 232; Black, nest
and eggs of, 454, 456, 459; colour
of eggs in Black, 454, 455, 456, 459
Skin, layers of, 19; products of, 20 Swift, bill of Chimney, 244, 245, 246;
foot of, 368, 369; nest of, 118; tail
of Chimney, 410, 412; wing-feathers
of, 42
Swiftlets, nest of, 119; use of nests, Swift, bill of Chimney, 244, 245, 246;
foot of, 368, 369; nest of, 118; tail
of Chimney, 410, 412; wing-feathers
of, 42
Swiftlets, nest of, 119; use of nests, of eggs in Black, 454, 455, 456, 459
Skin, layers of, 19; products of, 20 Skin, layers of, 19; products of, 20
Skull, 103-115; bones of, 107; evolu-
tion of, 104-106; origin of, 103-104;
section through, 201; alligator, 105;
Eagle, 106; Hawk, 199, 200; Heron,
199, 200; shark, 104
Sleep, birds in, 344, 355
Sloth, head of, 210
Snail used as food, 148, 453
Snake, egg of, used as food, 157, 429
Snakebird, bill of, 228, 229; body-
feathers of, 289; foot of, 391; method
of fishing of, 154; neck of, 282, 283;
stomach of, 139 Swiftlets, nest of, 119; use of nests,
120
Swim-bladder of fishes, 178, 179 Swim-bladder of fishes, 178, 179
Swimming compared with flight, 327 Swim-bladder of fishes, 178, 179
Swimming compared with flight, 327 Syrinx, drum of Mallard, 168, 170;
structure of, 172 Swiftlets, nest of, 119; use of nests,
120 teryx Tail, of Archaeopteryx, 398, 399, 400,
401; evolution of, 398-403; feathers
of, 402, 403; lack of, in Emeu, 402,
404; in Cassowary, 402 Tail, of Archaeopteryx, 398, 399, 400,
401; evolution of, 398-403; feathers
of, 402, 403; lack of, in Emeu, 402,
404; in Cassowary, 402 Snake, egg of, used as food, 157, 429
Snakebird, bill of, 228, 229; body-
feathers of, 289; foot of, 391; method
of fishing of, 154; neck of, 282, 283;
stomach of, 139 Index 495 Thrush, crest of Laughing, 263; effect
of humidity on Wood, 294
Tibia, 98 Thrush, crest of Laughing, 263; effect
of humidity on Wood, 294
Tibia, 98 Tail,
num number of feathers in Archaeopteryx,
400; Cormorant, 400; Duck, 400;
Ostrich, 402; Peafowl, 402; Fan-
tail Pigeon, 402 Tinamou, 334; egg-shell of, 460; flight
of, 333-336; tail of, 409 tail Pigeon, 402
ornaments of, 414-421; ploughshare-
bone of, 400; sham, of Quezal, 421,
422; sh m, of Peacock, 422, 423 of, 333-336; tail of, 409
Toes, of Cassowary, 394, 396; Chuck-
will's-widow, 369; Coot, 387; Donkey,
394, 395; embryo, 474; Ballinule,
385,386; Grebe, 391; Ruffed Grouse,
380, 381; Heron, 101; comb on
Heron, 387; 388; perching function
of hind, 359; horse, 358; Wood Ibis,
385; jaguar, 101; young kangaroo,
394, 395; European House Martin,
381; monkey, 358; number of, 353;
Osprey, 374; Ostrich, 358, 394, 395,
396; Snowy Owl, 375; Penguin, 391,
392; Plover, 384; Phalarope, 384;
Ptarmigan, 381; Rhinoceros-bird,
357; Sand-grouse, 383; Semipalmated
Sandpiper, 384; Snakebird, 391;
Vulture, 377; Zygodactyl, 369, 370,
371, 372 422; sh m, of Peacock, 422, 423
vertebrae of Bald Eagle, 403; embryo
bird, 403; Ostrich, 402 bird, 403; Ostrich, 402
use of, 405; voluntary decoration of,
424, 425, 425; of creeper, 410, 411;
deer, 413; Pin-tail Duck, 415;
Blue Duck, 406; Emeu-wren, 415;
Least Flycatcher, 412; Japanese
long-tailed fowl, 417, 419; Jungle
Fowl, 418; Boat-tailed Crackle,
418; Green Heron, 413; Junco,
414; Meadow Lark, 414; Lyre-
bird, 420, 421; Mexican Motmot,
424, 425, 425; Murre, 406, 407;
Grass Parrakeat, 408; Peacock, 422,
423; Pelican, 408, 409; Black-
footed Penguin, 406; Fan-tail Pigeon
404; Peacock Pheasant, 419, 420;
Reeves Pheasant, 416, 418; Rabbit,
413; Road-runner, 413; Sparrow
in moult, 409; Vesper Sparrow,
414; Barn Swallow, 405, 407; Tern,
407, 409; Chimney Swift, 410, 412;
Tinamous, 409; Wagtail, 413;
whale, 405; Paradise Whydah-
finch, 415, 416; Woodhewers, 410;
Woodpeckers, 410, 411, 412 Tongue, of Cockatoo, 125; Ducks and
Geese, 120, 121, 122; Flamingo, 126,
127; Flicker, 123, 126; function of
fleshy teeth on, 120; Goldfinch, 125;
Honey-creeper, 127, 129; Humming-
bird, 127; Lory, 125; Owls, Larks, and
Swifts, 122; Pelican, 120, 121; Sap-
sucker, 124; Toucan, 126; Wood-
peckers, 122 peckers, 122
Toucan, bill of, 243, 244; tongue of, 126
Towhee, iris of, 256
Trachea, of amphibian, 169; of cranes, Woodpeckers, 410, 411, 412
Tail-coverts of Quezal, 421, 422; of
peacock, 422, 423 Trachea, of amphibian, 169; of cranes,
170, 171; cf. teryx with oesophagus, 167;
of duck, 168; of flamingo, 168, 169;
of reptiles, 169; of Sparrow, 169; struc-
ture, of, 168, 169; of Trumpeter Swan,
170
396, 397 Trachea, of amphibian, 169; of cranes,
170, 171; cf. with oesophagus, 167;
of duck, 168; of flamingo, 168, 169;
of reptiles, 169; of Sparrow, 169; struc-
ture, of, 168, 169; of Trumpeter Swan,
170
396, 397 peacock, 422, 423
Tailor-bird, bill of, 245 peacock, 422, 423
Tailor-bird, bill of, 245 Tailor-bird, bill of, 245
Tanager, colour change in moult of, 294
Tarpon, scale of, 20
Tarsus, 99 Tailor-bird, bill of, 245
Tanager, colour change in moult of, 294 Tracks, of Cassowary, 396; of Dino aur,
396, 397
Triceratops, beak of, 226; brain of, 200 Tarsus, 99
Teeth, in embryo Tern, 476; origin of,
23, 114 23, 114
Temperature, of bird, 186; of man, 186
Tendons, of foot, 191, 192; strength of, Triceratops, beak of, 226; brain of, 200
Turbinal bones, 205 Temperature, of bird, 186; of man, 186
Tendons, of foot, 191, 192; strength of,
illustrated, 193; use in perching of,
195 Turkey, breast-ornament of Wild, 280
Turtle, beak of Snapping, 224; egg of,
428, 429 Turkey, breast-ornament of Wild, 280
Turtle, beak of Snapping, 224; egg of,
428, 429 Tern, bill of, 231, 232; colour of eggs of,
454, 456, 457; eggs of Stilt and, 452;
method of fishing of, 154; nest of, 453,
456, 457; protective coloration in
Sooty, 300, 303; tail of, 407, 409;
teeth in embryo, 476; wing of, 352
Thales, 12 Tern, bill of, 231, 232; colour of eggs of,
454, 456, 457; eggs of Stilt and, 452;
method of fishing of, 154; nest of, 453,
456, 457; protective coloration in
Sooty, 300, 303; tail of, 407, 409;
teeth in embryo, 476; wing of, 352
Thales, 12 " Two Bird-Lovsrs in Mexico," quota-
tion from, 301 X dactyls, 285
Whale, food of, 148; tail of, compared
with that of bird, 405 U Ulna, 2
Umbrella-bird, crest of, 264, 266
Uncinate processes, 77 Ulna, 2
Umbrella-bird, crest of, 264, 266
Uncinate processes, 77 teeth in embryo, 476; wing of, 352
Thales, 12 Thayer, experiments of Abbott, Thigh-bone, see Femur Thigh-bone, see Femur
Thigh-girdle, 88-91; bones composing,
88, 89; of embryo bird, cf. X-ray photograph of Pigeon, 63
Y with that of bird, 405
Whydah-finch, tail of Paradise, 415, 416 Whydah-finch, tail of Pa
Windpipe, see Trachea U with Dino-
saur, 90 Thrasher, eyelids of Brown, 212, 213;
nictitating membrane of, 213 Thrasher, eyelids of Brown, 212, 213;
nictitating membrane of, 213 Index 496 496
Vertebrae (continued)
function of, 196; Jabiru, neck
vertebra of, 72; Ostrich, 69, 71;
Ostrich-tail, 402
pelvic, of American Flamingo, 76;
alligator (young), 76; Bald Eagle, 76
of shark, 68; of tail, 75
Vireo, 144; iris of eye of, 256
Vorticlla, 463, 464 Wing of, Albatross, 320, 321, 325, 332-
Great Auk, 339, 340; Racor-billed
Auk, 339,340; Cassowary, 321,337,
338; Young Catbird, 322; Condor,
324, 326, 332; Steamer Duck, 337;
Snowy Egret, 350; West African
Goatsucker, 359; Canada Goose, 346;
Young Canada Goose, 322; Spur-
winged Goose, 346, 347; Ruffed
Grouse,347; Herring Gull, 328; Young
Green Heron, 320, 323; Great White
Heron, 321; Sleeping Heron, 344;
Hummingbird, 320, 321,325; Mallard,
47; Murrelets, 339; Young Osprey,
322, 351; Ostrich, 321,337,338,339;
Twelve-wired Bird of Paradise, 348;
Parrot, 322; Parrot Owl, 333; Pen-
guin, 321, 341, 342, 343; Pheasant,
323, 324, 327; Screamer, 346; Black
Skimmers, 324; English Sparrow, 46;
Spoonbill, 350; Trumpeter Swan, 346;
Tern, 352; Bush Warbler, 347; Wood-
cock, 348, 349; Turkey Vulture, 331
Wish-bone, see Clavicle Wing of, Albatross, 320, 321, 325, 332-
Great Auk, 339, 340; Racor-billed
Auk, 339,340; Cassowary, 321,337,
338; Young Catbird, 322; Condor,
324, 326, 332; Steamer Duck, 337;
Snowy Egret, 350; West African
Goatsucker, 359; Canada Goose, 346;
Young Canada Goose, 322; Spur-
winged Goose, 346, 347; Ruffed
Grouse,347; Herring Gull, 328; Young
Green Heron, 320, 323; Great White
Heron, 321; Sleeping Heron, 344;
Hummingbird, 320, 321,325; Mallard,
47; Murrelets, 339; Young Osprey,
322, 351; Ostrich, 321,337,338,339;
Twelve-wired Bird of Paradise, 348;
Parrot, 322; Parrot Owl, 333; Pen-
guin, 321, 341, 342, 343; Pheasant,
323, 324, 327; Screamer, 346; Black
Skimmers, 324; English Sparrow, 46;
Spoonbill, 350; Trumpeter Swan, 346;
Tern, 352; Bush Warbler, 347; Wood-
cock, 348, 349; Turkey Vulture, 331
Wish-bone, see Clavicle of shark, 68; of tail, 75
Vireo, 144; iris of eye of, 256 Vireo, 144; iris of
Vorticlla, 463, 464 Vorticlla, 463, 464
Vulture, colour in young Turkey, 317;
flight of, 328, 330, 331, 332; King,
head of, 272, 273; sense of sight in,
205; toes of, 377; Turkey, in flight, Y Wing, bones of, 95; change of function
in, 337; compared with arm of man,
94; evolution of, 91-97, 319; feather
divisions of, 320; framework of, 91-
97; noise of, 345, 347; spurs of, 346,
347; strokes of Hummingbird, 82;
use of, in defence, 337 Yellow-throat, life-habits of Maryland,
362
Z W Wading birds, number of eggs laid by,
438 Wagtail, tail of, 413
Walrus, brain of, 200 Wagtail, tail of, 413
Walrus, brain of, 200 Walrus, brain of, 200
Warbler, life-habits of Black-and-white,
366; Black-throated Green, 366; Mag-
nolia, 366; Myrtle, 364; Pine, 366;
Worm-eating, 163; Yellow Palm, 363 Wish-bone, see Clavicle
Woodcock, bill of, 222; eyes of, 221, 256;
sense of sight in, 208; sense of touch
in, 219, 221, 222; wing-song of, 348
Woodhewer, tail of, 410 Worm-eating, 163; Yellow Palm, 363
Warbler-bush, wings of, 347 Warbler-bush, wings of, 347
Warblers, evolution of, 361-367 Warblers, evolution of, 361-367
Water-birds, number of eggs laid by,
436, 437, 438 Woodhewer, tail of, 410
Woodpecker, tail of 410, 411, 412;
nest and eggs of Hairy, 445; char-
acter of egg-shell of, 460; colou of
eggs of, 444; function of saliva in,
118; tongue of, 122; bill of, 245;
crest of, 264; flight of, 329; feet of,
370 Woodhewer, tail of, 410
Woodpecker, tail of 410, 411, 412;
nest and eggs of Hairy, 445; char-
acter of egg-shell of, 460; colou of
eggs of, 444; function of saliva in,
118; tongue of, 122; bill of, 245;
crest of, 264; flight of, 329; feet of,
370 436, 437, 438
Wattles, of Bell-bird, 273; Condor, 271;
Wild Turkey, 273, 275; King Vulture,
272, 273 272, 273
Waxbill, African, abnormal number of
eggs laid by, 444 eggs laid by, 444
Web of toes, of Flamingo, 388; Sea-
birds, 389; Semipalmated Sandpiper,
384 Wren, Marsh, colour of eggs of, 446;
tail of, 411 Weight of birds, 285, 286; of Ptero-
dactyls, 285 tail of, 411
Wrist-bones of bird, 92; of man, 92 with that of bird, 405
Whydah-finch, tail of Paradise, 415, 416
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NMR Structure of Hsp12, a Protein Induced by and Required for Dietary Restriction-Induced Lifespan Extension in Yeast
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PloS one
| 2,012
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cc-by
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Abstract Dietary restriction (DR) extends lifespan in yeast, worms, flies and mammals, suggesting that it may act via conserved
processes. However, the downstream mechanisms by which DR increases lifespan remain unclear. We used a gel based
proteomic strategy to identify proteins whose expression was induced by DR in yeast and thus may correlate with longevity. One protein up-regulated by DR was Hsp12, a small heat shock protein induced by various manipulations known to retard
ageing. Lifespan extension by growth on 0.5% glucose (DR) was abolished in an hsp12D strain, indicating that Hsp12 is
essential for the longevity effect of DR. In contrast, deletion of HSP12 had no effect on growth under DR conditions or
a variety of environmental stresses, indicating that the effect of Hsp12 on lifespan is not due to increased general stress
resistance. Unlike other small heat shock proteins, recombinant Hsp12 displayed negligible in vitro molecular chaperone
activity, suggesting that its cellular function does not involve preventing protein aggregation. NMR analysis indicated that
Hsp12 is monomeric and intrinsically unfolded in solution, but switches to a 4-helical conformation upon binding to
membrane-mimetic SDS micelles. The structure of micelle-bound Hsp12 reported here is consistent with its recently
proposed function as a membrane-stabilising ‘lipid chaperone’. Taken together, our data suggest that DR-induced Hsp12
expression contributes to lifespan extension, possibly via membrane alterations. Citation: Herbert AP, Riesen M, Bloxam L, Kosmidou E, Wareing BM, et al. (2012) NMR Structure of Hsp12, a Protein Induced by and Required for Dietary
Restriction-Induced Lifespan Extension in Yeast. PLoS ONE 7(7): e41975. doi:10.1371/journal.pone.0041975 Editor: Matt Kaeberlein, University of Washington, United States of America Editor: Matt Kaeberlein, University of Washington, United States of America Received May 13, 2012; Accepted June 26, 2012; Published July 27, 2012 Received May 13, 2012; Accepted June 26, 2012; Published July 27, 2012 rbert et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
tion, and reproduction in any medium, provided the original author and source are credited. Copyright: 2012 Herbert et al. This is an open-access article distributed under the terms of the Creative Commons Attribut
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Abstract Funding: This work was supported by a Biotechnology and Biological Sciences Research Council research grant to AM and SRP (Grant Ref ERA16235; funder’s
URL http://www.bbsrc.ac.uk/home/home.aspx), a Wellcome Trust project grant to AM and LYL (Grant Ref 090077/Z/09/Z; funder’s URL http://www.wellcome.ac. uk/), and Wellcome Trust Prize Studentships to MR and LB. The funders had no role in study design, data collection and analysis, decision to publish, or
preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. Competing Interests: The authors have declared that no competing interests exist. * E-mail: amorgan@liverpool.ac.uk . These authors contributed equally to this work. . These authors contributed equally to this work. NMR Structure of Hsp12, a Protein Induced by and
Required for Dietary Restriction-Induced Lifespan
Extension in Yeast Andrew P. Herbert1,2., Miche`le Riesen1., Leanne Bloxam1, Effie Kosmidou1, Brian M. Wareing1,
James R. Johnson1, Marie M. Phelan2, Stephen R. Pennington3, Lu-Yun Lian2, Alan Morgan1*
1 Department of Cellular and Molecular Physiology, Institute of Translational Medicine, University of Liverpool, Liverpool, United Kingdom, 2 NMR Centre for Structur
Biology, Institute of Integrative Biology, University of Liverpool, Liverpool, United Kingdom, 3 University College Dublin Conway Institute of Biomedical and Biomolecula
Research, University College Dublin, Dublin, Ireland PLoS ONE | www.plosone.org Results To validate these protein expression changes, we
prepared extracts from yeast containing chromosomally-tagged
GFP fusion constructs [18] and performed western blots using
a GFP antibody. Examples of proteins selectively induced by DR
(GFP-Hxk1) or high osmolarity (GFP-Ctt1) are shown in Fig. S1B. Of the proteins identified as being induced by both interventions,
specific bands of the predicted size could not be reproducibly
detected for GFP-Rtc3 or –Rgi1; whereas GFP-Eno1 expression
was not altered by DR. However, GFP-Hsp12 was confirmed to
be induced by both DR and high osmolarity (Fig. S1B). To rule
out any artefactual effect of the GFP tag, we raised an antiserum
against the N-terminus of Hsp12 and used this in western blots of
wild type cells. This revealed a band of the expected size
(,12 kDa), which was increased in intensity upon growth in DR
and high osmolarity conditions and which was not present in an
isogenic hsp12 deletion strain (Fig. S1C), thus confirming the
increased expression of Hsp12 under conditions of enhanced
longevity. In addition to possessing anti-aggregation properties, small heat
shock proteins are often large homo-oligomeric assemblies of
folded subunits. To further investigate the possible function of
Hsp12,
we
determined
its
solution
structure
using
NMR. Recombinant Hsp12 expressed in E. coli was monomeric. The
15N-1H HSQC spectrum showed poor resonance dispersion in the
proton dimension, which suggested that Hsp12 is intrinsically
disordered in aqueous buffer (Fig. 3A). Recently published circular
dichroism studies have shown that Hsp12 gains significant helical
content upon binding to lipid or SDS micelles [16], we therefore
examined the effect of varying SDS concentrations. The 15N-1H
HSQC spectra of Hsp12 showed a dose-dependent increase in
dispersion in response to SDS, indicating that Hsp12 adopts
a folded conformation upon micelle binding (Fig. 3B). Having
determined the optimal SDS concentration for NMR, we then
characterised the temperature-dependence of Hsp12 in the
presence (Fig. S4A) and absence (Fig. S4B) of SDS. This resulted
in linear resonance dispersion until 45uC, above which some
resonances deviated from a straight line in the presence of SDS,
indicating heat-induced unfolding. These optimised conditions
were then used to assign the residues of SDS-bound 15N/13C-
labelled Hsp12 (Fig. 3C). To determine if Hsp12 is causally linked to DR-induced lifespan
extension, we performed replicative lifespan analysis. This was
done by determining the number of daughter cells removed by
micromanipulation from individual virgin mother cells [19]. Results To identify proteins that are induced by DR, we analysed
extracts from BY4741 yeast cells grown under standard (2%
glucose) or DR (0.5% glucose) conditions by 2-D gel electropho-
resis. Initial experiments using wide range (pH 3–10) gels revealed
no obvious reproducible changes in protein spot abundance,
indicating that DR does not cause gross proteomic alterations
(Fig. 1A). However, using pH 5.3–6.5 and pH 3–5.6 zoom gels,
we could resolve several proteins that were barely detectable in
control conditions, yet clearly induced in DR conditions (Fig. 1B,
C). Mass spectrometry was used to identify these differentially
expressed proteins as Eno1, Hxk1, Hsp12, Rtc3, Rgi1, Sbp1 and
Yef3 (Table S1). Various small heat shock proteins have been shown to be
‘holdase’ molecular chaperones that bind to denaturing proteins
and prevent their aggregation. To determine if Hsp12 had such
activity, we investigated the ability of recombinant purified Hsp12
to prevent aggregation of the model substrate, insulin, using the
method of Haslbeck et al. [23]. Addition of DTT reduces the
disulphide bonds between the A and B chains of insulin, causing
aggregation; whereas in the absence of DTT, insulin remains
stable (Fig. S3A). DTT-induced insulin aggregation was greatly
reduced by recombinant GST-fusion proteins of the known
chaperones, yeast Hsp26 [23] and mammalian cysteine string
protein (CSP) [24]; but not by CaBP1s, used as a control for
a protein of similar size to Hsp12 with no known or predicted
chaperone functions (Fig. S3B). However, GST-Hsp12 was similar
to GST-CaBP1s in terms of ability to prevent insulin aggregation. The differences in chaperone activity for GST-Hsp12 and GST-
Hsp26 were then assessed in a dose-dependent manner. This
revealed that GST-Hsp26 has approximately 100-fold higher anti-
aggregation activity than GST-Hsp12 (Fig. S3C), indicating that
Hsp12 has very low, if any, intrinsic chaperone activity. (
)
To help pinpoint DR-induced proteins that play a causal role in
mediating lifespan extension, as opposed to those whose expression
patterns are merely coincidental, we then analysed proteins that
were induced by high osmolarity (Fig. S1). Our rationale was
based on the observation that lifespan extension by DR and high
osmolarity act via a common downstream pathway [17]; and
hence the crucial effector proteins are likely to be common to both
interventions. Proteins induced by high osmolarity comprised
Ctt1, Eno1, Fba1, Hsp12, Hsp26, Hsp31, Lys9, Rtc3, Rgi1 and
Oye2 (Table S1). Hsp12 Structure and Function Hsp12 determined here is consistent with its recently proposed
function as a membrane-binding ‘lipid chaperone’ [16], which
may in turn explain Hsp12’s ability to modulate various
phenotypes and cellular functions. Hsp12 determined here is consistent with its recently proposed
function as a membrane-binding ‘lipid chaperone’ [16], which
may in turn explain Hsp12’s ability to modulate various
phenotypes and cellular functions. Stress resistance correlates positively with lifespan in various
model organisms and DR may represent a mild stress that extends
lifespan via a hormesis-like mechanism [20]. We therefore
investigated if deletion of HSP12 reduced resistance to environ-
mental stresses. For comparison, we also included sir2 and fob1
deletion strains in this analysis, as deletion of SIR2 and FOB1 is
known to decrease and increase replicative lifespan respectively
[21,22]. There was no detectable difference between the ability of
BY4741 wild type and the deletion mutants to grow under a wide
variety of stress conditions, including DR and other stresses that
increase Hsp12 expression (Fig. S2). We therefore conclude that
Hsp12 does not contribute to general stress resistance. Introduction in increased ribosomal DNA (rDNA) silencing and a consequent
reduction in rDNA recombination [6,9,10,11]. An alternative
theory is that DR extends lifespan in a Sir2-independent manner
by inhibition of the Tor and Sch9 kinase signalling pathways
[12,13]. In this latter model, the downstream molecular mechan-
isms effecting longevity are not entirely clear, but may include
reduced ribosomal protein biogenesis [13,14] as well as reduced
rDNA recombination [15]. It is commonly accepted that similar fundamental cellular
processes modulate ageing in most eukaryotes [1]. Evidence to
support this idea comes from studies of dietary restriction (DR), i.e. underfeeding without malnutrition [2]. DR extends lifespan in
most
model
organisms,
including
yeast,
worms,
flies,
and
mammals [3,4], suggesting that it may act via conserved longevity
mechanisms. Studies using the budding yeast, Saccharomyces
cerevisiae, have been at the forefront of recent efforts to understand
the molecular mechanism of action of DR. Reducing the
concentration of glucose in yeast growth media from the standard
2% to 0.5% or below increases both replicative and chronological
lifespan in multiple genetic backgrounds and has been suggested to
be a model of DR [5,6,7,8]. To shed light on the mechanisms by which DR extends yeast
lifespan, we set out to use an unbiased strategy to identify proteins
whose expression is induced by DR and thus correlate with
longevity. Hsp12, a small heat shock protein whose cellular
functions are unclear, was identified by this approach and was
found to be essential for lifespan extension by DR. At the
molecular level, Hsp12 was found to be unlike other small heat
shock proteins, in that it is monomeric and intrinsically unfolded in
solution and has negligible in vitro chaperone activity. Upon
binding to membrane-mimetic SDS micelles, Hsp12 switches to
a 4-helical conformation. The NMR structure of micelle-bound Although the effect of glucose limitation in extending yeast
replicative lifespan is not disputed, its mechanism of action
remains the subject of considerable debate. One popular model
postulates that DR increases lifespan by activating the NAD-
dependent histone deacetylase, Sir2, or its homologues, resulting July 2012 | Volume 7 | Issue 7 | e41975 PLoS ONE | www.plosone.org 1 Hsp12 Structure and Function PLoS ONE | www.plosone.org Results Helix III
is not as stable as the other four helices, as revealed by the lower
number of daNi,i+3, dabi,i+3 connectivities for this helix and more
variation in its length compared with the other three helices
together with a high RMSD value of 0.465 (Table S2). The
experimentally-determined structural data correspond well with prediction using the AGADIR programme [26], which shows that
the region between 52–63 has a lower helical propensity compared
with the other three helical regions. Extensive analysis of residual
dipolar couplings using stretched acrylamide gels revealed no
evidence of long-range interactions between the individual helices,
indicating that Hsp12 does not form a stably-folded structure. g
p
y
We generated a model of the tertiary structure of Hsp12 using
CYANA. The ensemble presented (Fig. 5 and Fig. S5) highlights
the flexibility of the a-helices relative to one another. The four a-
helices can be more clearly identified in the representative model
in Fig. 6A, with the 4th and most C-terminal helix represented in
yellow/red. Analysis of the charge distribution reveals each a-helix
to be broadly amphipathic, with hydrophobic (green) residues
lying on one face and charged (red) residues on the opposite face
(Fig. 6 B,C). In addition, the residues flanking each a-helix also
tend to be charged. This suggests that hydrophobic residues of
Hsp12 insert into the lipidic component of membranes, while the
charged (mainly positive) residues interact with negatively charged
head groups and project away from the membrane. A Ramachan-
dran plot of the data is presented in Fig. S6. Overall, the NMR
data indicate that Hsp12 is intrinsically unstructured in aqueous
solution, but switches to a dynamic 4-helical conformation upon
membrane binding. Figure 2. HSP12 is essential for lifespan extension by DR. (A) The
replicative lifespan of the wild type BY4741 strain grown under
standard (2% glucose) and DR (0.5% glucose) conditions was de-
termined manually on YPD plates by micromanipulation. Mean and
median lifespan were 21 and 22, respectively, (n = 55) for 2% glucose;
and 31 and 35, respectively, (n = 59) for 0.5% glucose. DR produced
a significant increase in lifespan in wild type cells (log-rank test,
P,0.001) (B) Mean and median lifespan of the isogenic hsp12D strain
was determined as above and found to be 26 and 28, respectively,
(n = 39) for 2% glucose; and 25 and 24, respectively, (n = 42) for 0.5%
glucose. Results DR did not increase lifespan in hsp12D cells (log-rank test,
P
0 5) Figure 2. HSP12 is essential for lifespan extension by DR. (A) The
replicative lifespan of the wild type BY4741 strain grown under
standard (2% glucose) and DR (0.5% glucose) conditions was de-
termined manually on YPD plates by micromanipulation. Mean and
median lifespan were 21 and 22, respectively, (n = 55) for 2% glucose;
and 31 and 35, respectively, (n = 59) for 0.5% glucose. DR produced
a significant increase in lifespan in wild type cells (log-rank test,
P,0.001) (B) Mean and median lifespan of the isogenic hsp12D strain
was determined as above and found to be 26 and 28, respectively,
(n = 39) for 2% glucose; and 25 and 24, respectively, (n = 42) for 0.5%
glucose. DR did not increase lifespan in hsp12D cells (log-rank test,
P.0.5). doi:10.1371/journal.pone.0041975.g002 Results Wild
type BY4741 cells exhibited a mean lifespan of 21 (95% CI: 19–23)
on standard (2% glucose) media, which was significantly increased
to 31 (95% CI: 28–35) under DR (0.5% glucose) conditions
(Fig. 2A). Deletion of HSP12 did not reduce longevity under
standard conditions, but rather resulted in a small increase in
mean lifespan to 26 (95% CI: 23–29). Strikingly, however, the
ability of DR to increase longevity was abolished in the hsp12D
strain, which exhibited a mean lifespan of 25 (95% CI: 22–28)
under DR conditions (Fig. 2B). These data suggest that the impact
of Hsp12 on cellular ageing is complex: the low-level expression
observed in standard media has a small negative effect on lifespan,
whereas high Hsp12 levels induced by DR are essential for the
increase in longevity caused by this intervention. Analysis of the backbone dynamics of Hsp12 in the presence of
SDS revealed relatively long T1 relaxation values compared to T2
(Fig. 4 A,C,E), suggesting restricted mobility in the majority of the
polypeptide. In contrast, T1 and T2 values were similar in the
absence of SDS (Fig. 4 B,D,F), suggesting that the protein is highly
dynamic in solution, but is structured on micelles. Consistent with
this, analysis of the assigned chemical shifts in Hsp12 using CSI
[25] suggested that micelle binding induces the formation of four
a-helices (Fig. 4G). These a-helices cover the majority of the
polypeptide and comprise residues F9-A16 (Helix I), Q22-A41 PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 PLoS ONE | www.plosone.org 2 Hsp12 Structure and Function PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 July 2012 | Volume 7 | Issue 7 | e41975 3 Hsp12 Structure and Function Figure 1. DR induces expression of a relatively small number of proteins. Wild type BY4741 yeast cells were grown in standard (2% glucose)
and DR (0.5% glucose) conditions before lysis and separation of proteins by 2-D electrophoresis. Wide-range (pH 3–10) gels revealed no obvious
reproducible differences in protein expression, as illustrated by representative gels shown in panel (A). Narrow pH range gels (pH 3–5.6 and 5.3–6.5)
revealed changes in protein spots, which were identified by mass spectrometry. Selected identified proteins are indicated by arrows in panels (B) and
(C). doi:10.1371/journal.pone.0041975.g001 doi:10.1371/journal.pone.0041975.g001 (Helix II), V52-G63 (Helix III) and L74-E94 (Helix IV). Discussion The mechanisms by which DR extends yeast replicative lifespan
remain unclear and controversial. The data presented here show
that DR (via growth in 0.5% glucose) does not induce gross
changes in overall protein abundance, but does result in up-
regulation of a limited number of proteins. This is consistent with
earlier global microarray analysis that revealed changes in mRNA
expression for only 133 genes (around 2% of total ORFs) in
response to growth in 0.5% glucose [9]. Some of the proteins we
found to be upregulated by DR (e.g. Rgi1, Hxk1) are associated
with metabolic adaptation from fermentation to respiratory
growth; whereas the reason for up-regulation of other proteins
(e.g. Hsp12, Rtc3) is less obvious. Both Hsp12 and Rtc3 are also
induced by high osmolarity (Fig. S1), which is thought to extend
lifespan via the same downstream mechanisms as DR [17],
consistent with the idea that DR is a chronic mild stress that
increases longevity via hormesis. Indeed, HSP12 mRNA levels are
increased in response to diverse environmental stresses, including
heat-, osmotic- and oxidative stress [27,28]. HSP12 is also among
the top 10 up-regulated genes in response to non-environmental,
genetically-mediated impairment of nuclear proteostasis [29] and
telomere capping [30]. Figure 2. HSP12 is essential for lifespan extension by DR. (A) The
replicative lifespan of the wild type BY4741 strain grown under
standard (2% glucose) and DR (0.5% glucose) conditions was de-
termined manually on YPD plates by micromanipulation. Mean and
median lifespan were 21 and 22, respectively, (n = 55) for 2% glucose;
and 31 and 35, respectively, (n = 59) for 0.5% glucose. DR produced
a significant increase in lifespan in wild type cells (log-rank test,
P,0.001) (B) Mean and median lifespan of the isogenic hsp12D strain
was determined as above and found to be 26 and 28, respectively,
(n = 39) for 2% glucose; and 25 and 24, respectively, (n = 42) for 0.5%
glucose. DR did not increase lifespan in hsp12D cells (log-rank test,
P.0.5). Figure 2. HSP12 is essential for lifespan extension by DR. (A) The
replicative lifespan of the wild type BY4741 strain grown under
standard (2% glucose) and DR (0.5% glucose) conditions was de-
termined manually on YPD plates by micromanipulation. Mean and
median lifespan were 21 and 22, respectively, (n = 55) for 2% glucose;
and 31 and 35, respectively, (n = 59) for 0.5% glucose. Yeast Cell Lysis and Protein Quantification Single yeast colonies were grown in 5 ml of YP media
containing the appropriate concentration of glucose overnight at
30uC. The next day, 4 ml of this starter culture was inoculated into
200 ml of YPD media YP media containing the appropriate
concentration of glucose and incubated at 30uC C until an OD600
value of 0.6 was obtained. The glucose concentration remaining in
the culture media fell by only 0.1% during this period (i.e., from
0.5% to 0.4%), as determined by glucose oxidase assays. At this
point, the culture was centrifuged at 8000 g and washed 3 times
with distilled water before freezing at 280uC. The equivalent of
a 5-ml culture in a frozen pellet was resuspended in 150 ml lysis
buffer (7 M urea, 2 M thiourea, 2% (w/v) CHAPS, 1% (w/v)
DTT, 0.8% (w/v) Pharmalyte, 1 Roche protease inhibitor tablet
per 10 ml), transferred to a 2 ml cryogenic vial and subjected to
glass bead lysis in a mikrodismembrator for 10 min at 1800 rpm at
4uC. The resulting lysate was cleared by centrifugation at
13000 rpm for 10 min at 4uC. The protein concentration of each
lysate was normalised by running samples together on a 1-D mini-
gel, staining the gel with Coomassie blue and quantifying the total
amount of protein in each track by densitometry. These values
were then used to ensure that the same amounts of protein were
loaded for each sample to be compared in subsequent 2-D gel
electrophoresis or western blotting. In summary, we have shown that Hsp12 is induced by DR, is
required for lifespan extension by DR, and have solved the
structure of the protein in the free- and micelle-bound state. Although Hsp12 does not have obvious homologues outside of
fungi, down-regulation of small heat shock proteins by RNAi also
affects ageing in C. elegans [33], consistent with a conserved
modulatory role for sHSPs in longevity. Hsp12 displays low-level
homology to two mammalian proteins: ageing-associated protein 2
(also known as HIP/ST13; NCBI reference NP_003923) and a-
synuclein (NCBI reference P37840). Intriguingly, a-synuclein is
also an intrinsically disordered protein in solution that becomes a-
helical upon binding to anionic phospholipids and SDS micelles
[34]. It is conceivable that these shared properties may have
functional implications, as it has been suggested that neuronal a-
synuclein organises and stabilises lipids in synaptic vesicle
membranes [35]. 2-D Gel Electrophoresis and Protein Identification Yeast lysates were applied at equal protein concentrations to
18 cm IPG strips (GE Healthcare) and allowed to rehydrate for
a minimum of 10 hours. The IPG strips were then run on an IEF
Multiphor II electrophoresis unit (GE Healthcare) at 20uC for
1 min at 500 V, then for 7 hours at 3500 V. Following this, IPG
strips were overlayed on SDS-PAGE gels, sealed using warm
agarose, and run using an Ettan Dalt II system (GE Healthcare) at
2.5 W per gel for 30 min and then 20 W per gel at 25uC until the
dye front had reached the bottom of the gel. Immediately after
SDS-PAGE, the gels were fixed in a solution of 40% v/v MeOH/
7% (v/v) acetic acid for a minimum of 2 hours and then stained
with colloidal Coomassie blue stain (GE Healthcare). Stained gels Discussion All other materials
were obtained from Sigma-Aldrich. supplied by Sigma Genosys (Havenhill, UK), genomic DNA
isolation kits were from Invitrogen (Paisley, UK); and PCR
enzymes/reagents were from Promega (Southampton, UK). Anti-
GFP antibody was obtained from AbCam (Cambridge, UK) and
custom-generated Hsp12 antiserum was supplied by Genosphere
Biotechnologies (Paris, France). Materials for gel electrophoresis
were obtained from GE Healthcare or BioRad. All other materials
were obtained from Sigma-Aldrich. Hsp26. It therefore seems more likely that Hsp12’s recently
discovered function as a membrane stabilising ‘lipid chaperone’
[16] underlies its effect on lifespan. Indeed, Ssd1 has been shown
to increase replicative lifespan in a Sir2-independent manner by
increasing plasma membrane stability [31]. The transition from
unfolded Hsp12 in solution to a dynamic 4-helical lipid-bound
conformation observed here may similarly contribute to mem-
brane stability under DR conditions, which in turn may be
functionally relevant to increasing lifespan. However, it remains
possible that unfolded Hsp12 in the nucleus and cytosol performs
membrane-binding-independent functions that impact on replica-
tive lifespan during DR. Yeast Strains Deletion strains with the appropriate ORF replaced by the
KanMX4 cassette in the BY4741 background (MATa his3D1
leu2D0 met15 D0 ura3D0) [36] were used for gel electrophoresis,
lifespan analysis and stress assays and were obtained from
Invitrogen (Paisley, UK). GFP-labelled strains in the BY4741
background were used to validate mass spectrometry identifica-
tions and were obtained from Invitrogen. All mutants were
confirmed by PCR using gene-specific and KanMX primers. p
g
The structure of Hsp12 bound to SDS micelles described here
provides insight into how Hsp12 may interact with cellular
membranes. This involves a switch from an intrinsically disordered
conformation in solution to a structured, 4-helical lipid-bound
conformation, as recently proposed [16]. Our extensive NMR
analysis using stretched acrylamide gels failed to reveal interactions
between the four a-helices of Hsp12, suggesting a dynamic
structure whereby each a-helix is free to move independently
within a 2-dimensional lipid bilayer. The lack of a stable folded
tertiary structure could be due to the observed instability of helix 3. While our work was being prepared for publication, the NMR
structure of micelle-bound Hsp12 was independently reported
[32]. This shows a similar 4-helical conformation in the presence
of SDS, although some differences are apparent in the secondary
structural models of the individual helices (F9-K18, Y25-G42,
Q54-S59 and D76-V100 versus F9-A16, Q22-A41, V52-G63 and
L74-E94 in our study). We propose that the dynamic nature of
Hsp12’s four a-helices we report here could be important for its
membrane
stabilising
function
[16],
as
this
would
enable
simultaneous linkage of several distinct membrane sub-domains
via a single protein tether. Yeast Cell Lysis and Protein Quantification It is tempting to speculate that enhanced
membrane stabilisation by functional homologues of Hsp12 may
be relevant to DR-induced longevity in higher organisms. Discussion DR produced
a significant increase in lifespan in wild type cells (log-rank test,
P,0.001) (B) Mean and median lifespan of the isogenic hsp12D strain
was determined as above and found to be 26 and 28, respectively,
(n = 39) for 2% glucose; and 25 and 24, respectively, (n = 42) for 0.5%
glucose. DR did not increase lifespan in hsp12D cells (log-rank test,
P.0.5). Despite the strong induction of HSP12 gene expression in
response to diverse stresses, deletion of HSP12 does not generally
affect sensitivity to these stresses. Significantly, however, we found
that lifespan extension by growth on 0.5% glucose was abolished
in an hsp12D strain, indicating that Hsp12 is essential for the
longevity effect of DR. What mechanism(s) could explain this
action of Hsp12 on lifespan? One tempting explanation is that, like
other small heat shock proteins, Hsp12 prevents the aggregation of
a variety of cellular proteins during DR-induced stress. However,
we found that Hsp12 has negligible anti-aggregation properties in
vitro, in contrast to well established ‘holdase’ co-chaperones such as doi:10.1371/journal.pone.0041975.g002 July 2012 | Volume 7 | Issue 7 | e41975 4 PLoS ONE | www.plosone.org Hsp12 Structure and Function PLoS ONE | www.plosone.org
5
July PLoS ONE | www.plosone.org
5
July 2012 | Volume 7 | Issue 7 | e41975 PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 PLoS ONE | www.plosone.org 5 Hsp12 Structure and Function Figure 3. Hsp12 is unstructured in solution, but folds in the presence of SDS. (A) 1H-15N HSQC spectrum of Hsp12 in aqueous solution at
298 K. The spectrum shows only sharp peaks with random coil shifts indicating the absence of any structured regions. (B) 1H-15N HSQC spectrum of
Hsp12 at 303 K in the presence of increasing concentrations of SDS (0, 1, 2, 5, 8 mM Red -. Blue). SDS causes a considerable increase in the amount
of chemical shift dispersion implying increased levels of folded material/regions. (C) Assigned 1H-15N HSQC spectrum of Hsp12 at 318 K in the
presence of 100 mM SDS. doi:10.1371/journal.pone.0041975.g003 p
doi:10.1371/journal.pone.0041975.g003 supplied by Sigma Genosys (Havenhill, UK), genomic DNA
isolation kits were from Invitrogen (Paisley, UK); and PCR
enzymes/reagents were from Promega (Southampton, UK). Anti-
GFP antibody was obtained from AbCam (Cambridge, UK) and
custom-generated Hsp12 antiserum was supplied by Genosphere
Biotechnologies (Paris, France). Materials for gel electrophoresis
were obtained from GE Healthcare or BioRad. Materials and Methods Chemicals and Reagents July 2012 | Volume 7 | Issue 7 | e41975 Chemicals and Reagents Chemicals and Reagents
Materials for yeast culture were obtained from Sigma-Aldrich
(Poole, UK) and Foremedium (Norwich, UK). PCR primers were g
Materials for yeast culture were obtained from Sigma-Aldrich
(Poole, UK) and Foremedium (Norwich, UK). PCR primers were PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 6 Hsp12 Structure and Function PLoS ONE | www.plosone.org
7
July 2012 | PLoS ONE | www.plosone.org
7
July 2012 | Volume 7 | PLoS ONE | www.plosone.org 7 July 2012 | Volume 7 | Issue 7 | e41975 7 PLoS ONE | www.plosone.org Hsp12 Structure and Function Hsp12 Structure and Function Figure 4. Backbone dynamics and chemical shift-based secondary structure of Hsp12. T1, T2 and T1/T2 relaxation values are shown for
Hsp12 in the presence (A,C,E) and absence (B,D,F) of 100 mM SDS at 318 K. T1 and T2 relaxation times for micelle-bound (A,C) Hsp12 show significant
variation; contrasting with the similar relaxation values observed for free Hsp12 (B,D). Micelle-bound Hsp12 (E) shows grouped variations in the T1/T2
values ranging from approximately 1.5 to 14, indicating a wide range of mobility and a clear differentiation of secondary structure elements; whereas
the free form (F) shows consistent values of around 2, indicating a completely unstructured protein. (G) The assigned chemical shifts at 318 K in
100 mM SDS expressed as deviation from random coil are shown aligned with the primary sequence and the positions of the a-helices. doi:10.1371/journal.pone.0041975.g004 Western Blotting were imaged with a GS-710 Imaging Densitometer (BioRad) and
protein spot changes analysed by eye and using PD Quest software
(BioRad). Protein spots of interest were excised from gels, dried
under vacuum, then re-hydrated in 25 mM NH4CO3 containing
5 ng/ml trypsin overnight at 37uC. Tryptic peptides
were
resuspended 1:1 in matrix solution (10 mg/ml HCCA in 50%
(v/v) ethanol, 50% (v/v) acetonitrile, 0.001% (v/v) trifluoroacetic
acid) and 1 ml of this mixture (25% of the total digest) was spiked
with 50 fmol of ACTH peptide, loaded onto a MALDI target
(Waters/Micromass Massprep workstation) and analysed on
a MALDI-TOF mass spectrometer (Waters/Micromass M@
LDI). Tryptic peptides were identified by peptide mass fingerprint
matching using MASCOT (UniProt release 2.6) allowing for 1
missed cleavage with a mass accuracy of 0.25 Da. were imaged with a GS-710 Imaging Densitometer (BioRad) and
protein spot changes analysed by eye and using PD Quest software
(BioRad). Protein spots of interest were excised from gels, dried
under vacuum, then re-hydrated in 25 mM NH4CO3 containing
5 ng/ml trypsin overnight at 37uC. Tryptic peptides
were
resuspended 1:1 in matrix solution (10 mg/ml HCCA in 50%
(v/v) ethanol, 50% (v/v) acetonitrile, 0.001% (v/v) trifluoroacetic
acid) and 1 ml of this mixture (25% of the total digest) was spiked
with 50 fmol of ACTH peptide, loaded onto a MALDI target
(Waters/Micromass Massprep workstation) and analysed on
a MALDI-TOF mass spectrometer (Waters/Micromass M@
LDI). Tryptic peptides were identified by peptide mass fingerprint
matching using MASCOT (UniProt release 2.6) allowing for 1
missed cleavage with a mass accuracy of 0.25 Da. Yeast lysates prepared as above were run on 1-D SDS-PAGE
gels and transferred to nitrocellulose. Blots were probed with either
anti-GFP antibody (AbCam, Cambridge, UK) or with custom-
generated antiserum (Genosphere Biotechnologies, Paris, France)
raised against a synthetic peptide corresponding to the N-terminal
14 residues of Hsp12 with an additional cysteine for conjugation
(sequence: MSDAGRKGFGEKASC). Blots were visualised by
enhanced chemiluminescence and imaged with a BioRad Chemi-
Doc XRS imager (BioRad). Lifespan Analysis This was performed as described previously [19]. Briefly, strains
were grown at 30uC until they reached an OD600 of 0.6–1.0. One Figure 5. Ensemble of structures calculated for micelle-bound Hsp12 overlaid on each of the four helices. Ensemble of twenty
structures overlaid on helices I (A), II (B), III (C) and IV (D). No long-range interactions were detected and so the helices appear free to move
independently with no overall fold being evident. doi:10.1371/journal.pone.0041975.g005 Figure 5. Ensemble of structures calculated for micelle-bound Hsp12 overlaid on each of the four helices. Ensemble of twenty
structures overlaid on helices I (A), II (B), III (C) and IV (D). No long-range interactions were detected and so the helices appear free to move
independently with no overall fold being evident. doi:10.1371/journal.pone.0041975.g005 PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 8 Hsp12 Structure and Function Figure 6. Helical properties of micelle-bound Hsp12. (A) The four a-helices are represented as ribbons and colour coded from the N-termin
(blue) to the C-terminus (red) in a representative structure. (B,C) Analysis of charge distribution with hydrophobic residues labelled green a
charged residues labelled red in both ribbon (B) and surface (C) representation, illustrating the amphipathic nature of Hsp12. Structures we
generated using Chimera. doi:10.1371/journal.pone.0041975.g006
PLoS ONE | www.plosone.org
9
July 2012 | Volume 7 | Issue 7 | e419 Figure 6. Helical properties of micelle-bound Hsp12. (A) The four a-helices are represented as ribbons and colour coded from the N-terminus
(blue) to the C-terminus (red) in a representative structure. (B,C) Analysis of charge distribution with hydrophobic residues labelled green and
charged residues labelled red in both ribbon (B) and surface (C) representation, illustrating the amphipathic nature of Hsp12. Structures were
generated using Chimera. doi:10.1371/journal.pone.0041975.g006 Figure 6. Helical properties of micelle-bound Hsp12. (A) The four a-helices are represented as ribbons and colour coded from the N-terminu
(blue) to the C-terminus (red) in a representative structure. (B,C) Analysis of charge distribution with hydrophobic residues labelled green an
charged residues labelled red in both ribbon (B) and surface (C) representation, illustrating the amphipathic nature of Hsp12. Structures we
generated using Chimera. doi:10.1371/journal.pone.0041975.g006 Figure 6. Helical properties of micelle-bound Hsp12. (A) The four a-helices are represented as ribbons and colour coded from the N-terminus
(blue) to the C-terminus (red) in a representative structure. PLoS ONE | www.plosone.org Aggregation Assay This was based on the method of Haslbeck et al. [23]. Insulin
was added to a final concentration of 45 mM in 40 mM HEPES-
KOH, pH 7.5 in a 96-well plate in the presence or absence of
recombinant proteins. Aggregation was induced by the addition of
1.5 ml of 1 M DTT in a final volume of 100 ml, or by adding
1.5 ml of water as a no-aggregation control. Measurements were
taken
in
an
Emax
microplate
reader
(Molecular
Devices,
Sunnyvale, USA) at 405 nm every 10 min for 3 hours, then one
final reading was obtained the next day after agitating the plate to
disperse any large aggregates. The anti-aggregation efficiency of
each recombinant protein was calculated as follows. For each
individual assay, the highest absorbance reading for the aggrega-
tion control (insulin + DTT) was assumed to reflect total
aggregation of insulin (26 mg per well) and set as 100%. Anti- NMR Methods NMR samples were prepared in 10 mM PO4
32, 40 mM NaCl,
2 mM NaN3, pH 6.5 in 90% (v/v) H2O/10% (v/v) D2O or 100%
D2O, at a final protein concentration of approximately 450 mM. Hsp12 samples in the presence of SDS were prepared using the
same buffer with the addition of 100 mM SDS. Spectra in the
absence and presence of SDS were acquired at, respectively,
303 K and 318 K, on Bruker Avance III 600 and 800 MHz
spectrometers equipped with cryogenic triple resonance probes. All NMR spectra were processed with TopSpin (Bruker) and
analysed using the CCPN Analysis package [38]. Sequence-
specific backbone and side-chain resonance assignment of Hsp12
was made using 3-D HNCA, HN(CA)CB, HN(CO)CA, HNCO,
CBCA(CO)NH,
HBHANH,
HBHA(CO)NH
and
HCCH-
TOCSY experiments. Assignment of aromatic side-chain residues
was made using 2-D [1H-13C] HSQC and homonuclear
1H
NOESY and TOCSY spectra. Interproton distance restraints were
obtained from NOEs derived from 3-D
15N- and
13C-edited
NOESY-HSQC experiments. The secondary structure for the
Hsp12 protein ensemble was determined using the STRIDE
algorithm [39]. The basis for secondary structure is a combination
of backbone torsion angle and location of hydrogen bonds. Furthermore, using the ensemble for secondary structure de-
termination enabled the location of the helices to be identified
based on twenty structures rather than one. RMSD values for the
ensemble were determined by global rmsd fit using MOLMOL
[40] which determines the average pairwise alignment RMSD for
all pairs in the ensemble. IPAP 1H 15N RDC data was collected
unaligned and aligned using the stretched gel method [41] on an
800 MHz spectrometer equipped with TCI cryoprobe. For each
sample a 600 ml preparation of gel mix (4–8% acrylamide),
containing a final concentration of 100–200 mM protein, was used
and quadrupole splitting measured to estimate the degree of
alignment. RDCs were collected on a 4.0% acrylamide gel of
initial 6 mm diameter compressed into a 4.6 mm diameter tube
with quadrupole splitting of 1.5 Hz containing 150 mM
15N
Hsp12 and buffer conditions: 100 mM SDS, 10mM PO4
32,
40 mM NaCl, pH 4.5. Couplings were measured using CCPN
analysis and a purpose-built Perl script, before direct incorporation
into structure calculation using CYANA [42]. Coordinates have
been deposited in the RCSB protein databank (PDB accession
number: 4AXP) and the Biological Magnetic Resonance Bank
(BMRB accession number: 18523). Recombinant Proteins GST-Hsp12 and GST-Hsp26 were constructed using the
Invitrogen Gateway cloning protocol. Primers for amplifying
Hsp12 and 26 from yeast genomic DNA were designed from the
ORF sequences of the required genes fused with the correspond-
ing attB1 and attB2 primer sequences. The destination vector used
for N-terminal GST-fusion, pG-GEX6p-B1, was made in-house. Recombinant GST-fusion proteins were expressed in BL21 (DE3)
E. coli, lysed using a One Shot cell disrupter (Constant Systems,
Daventry, UK) and purified using glutathione-Sepharose as
previously described [37]. p
y
For NMR studies, the ORF of Hsp12 was codon optimised for
E. coli expression and then synthesised de novo by GeneArt
(Invitrogen). The resulting construct was then cloned into the pE-
SumoProKan vector (Invitrogen) and expressed in BL21 (DE3) E. coli in M9 medium containing 1 g/L 15NH4Cl as the sole nitrogen
source, with either 4 g/L 12C-glucose or 13C-glucose as the sole
carbon source, to produce, respectively, [U-15N]- or [U-15N,13C]-
Hsp12. Recombinant His-SUMO-tagged Hsp12 was purified
using HisTrap immobilised metal affinity columns (GE Health-
care) via imidazole elution. The SUMO-specific protease, ULP1,
was used to cleave Hsp12 (with no vector-derived residues) from
the His-SUMO tag, which was removed by passage through
a second immobilised metal affinity column. The resulting Hsp12
protein was then dialysed against 10 mM KHPO3, 40 mM NaCl,
pH 6.5 and concentrated prior to use in NMR. Stress Assays Single colonies were grown overnight in 5 ml liquid YP media
containing 2% (w/v) glucose. The next morning, cultures were
diluted to OD600 = 1 in sterile H2O and then serially diluted ten-
fold five times in sterile H2O. The serial dilutions were then
spotted with a replica plater onto YP media containing the
indicated stressors. Plates were incubated at 30uC for 2 to 4 days
and then imaged with a BioRad ChemiDoc XRS imager
(BioRad). Hsp12 Structure and Function Hsp12 Structure and Function microlitre of culture was streaked onto plates and left at 30uC for
1–2 h. After this time, cell doublets were moved to uninhabited
regions of the plate. When these budded again, (newly formed)
virgin yeast cells were removed by micromanipulation to a new
location. All future buds produced by these daughter cells were
micromanipulated away and catalogued. The plates were in-
cubated at 30uC during working hours, and moved to 4uC
overnight. Lifespan was defined as number of daughter cells
removed from the mother cell. All lifespans were observed at least
twice. Statistical significance was assessed using the log rank test
and deemed significant at P,0.05. aggregation activity was then calculated by subtracting the
aggregated material in each condition from 100%, converting to
mg insulin and then dividing by the respective amount of each
GST-tagged protein to determine the anti-aggregation properties
per mg recombinant protein. Lifespan Analysis (B,C) Analysis of charge distribution with hydrophobic residues labelled green and
charged residues labelled red in both ribbon (B) and surface (C) representation, illustrating the amphipathic nature of Hsp12. Structures were
generated using Chimera. doi:10.1371/journal.pone.0041975.g006 PLoS ONE | www.plosone.org PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 9 PLoS ONE | www.plosone.org References 1. Guarente L, Kenyon C (2000) Genetic pathways that regulate ageing in model
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Temperature optimisation of Hsp12. 1H-15N
HSQC spectrum of Hsp12 in the presence of 100 mM SDS (A) or
in aqueous solution (B) at different temperatures (298, 303, 308,
313, 318, 323 K, Blue -. Red). Increases in temperature are
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not undergo significant unfolding even up to temperatures of
323 K. (TIF) Figure S5
Twenty structures calculated for micelle-
bound Hsp12 tiled individually. Structures were generated
using chimera. (TIF) Figure S2
HSP12 is not required for general stress
resistance. Overnight cultures of BY4741 wild type and deletion
strains were serially diluted and then spotted with a replica plater
onto YPD plates containing 2% glucose at 30uC unless indicated
otherwise. Plates were incubated at 30uC for 2 to 4 days and then
imaged in a BioRad Universal Hood II Imager (BioRad). (TIF) Figure S6
Ramachandran plot. The percentage of ordered
residues in the presence of 100 mM SDS at 45uC was 80.9% in
most favoured regions, 18.1% in additionally allowed regions,
0.6% in generously allowed regions and 0.4% in disallowed
regions. (TIF) Figure S3
Recombinant Hsp12 has negligible in vitro
chaperone activity. (A) 45 mM insulin was supplemented with
1.5 ml water (open squares) or 1 M DTT (black squares) and
aggregation over time at room temperature was measured at A405
in a microplate reader. (B) Aggregation assays were performed as
above in the presence or absence of the indicated GST-fusion
proteins. Anti-aggregation activity is shown as the amount of
insulin in mg which is prevented from aggregation by 1 mg of
recombinant protein. Data shown are pooled from multiple
experiments (n = 7 for GST-Hsp12; n = 8 for GST-Hsp26; n = 4
for GST-CSP; n = 4 for GST-CaBP1s). The difference between
GST-Hsp12 and GST-CaBP1s was deemed significant at P,0.05
using a Student’s t-test. (C) Dose-response curves of recombinant
GST-Hsp12 and GST-Hsp26. GST-Hsp26 greatly reduces insulin
aggregation, whereas GST-Hsp12 has mimimal effect. (TIF) Table S1
Mass spectrometry identifications of proteins
induced by dietary restriction and high osmolarity. Information from the Saccharomyces Genome Database (SGD) is
presented along with peptide mass fingerprinting data obtained for
each identified protein. (XLS) Table S2
Average RMSD values for Hsp12 helices. RMSD values calculated from the mean CYANA coordinates
for helices I-IV are shown. (DOC) Author Contributions Conceived and designed the experiments: AM SRP AH MR LYL. Performed the experiments: AH MR LB EK BMW. Analyzed the data: AM SRP AH MR LYL. Contributed reagents/materials/analysis tools:
JRJ MMP. Wrote the paper: AM. Supporting Information Figure
S1
Hsp12
is
induced
by
multiple
lifespan-
extending interventions. (A) Wild type BY4741 yeast cells
were grown in standard (2% glucose) and high osmolarity (20%
glucose) conditions before lysis and separation of proteins by 2-D
electrophoresis on narrow pH range gels (pH 3.5–6 and 5.3–6.5). Selected spot changes identified by mass spectrometry are
indicated by arrows. (B) S288c yeast cells expressing chromoso-
mally GFP-tagged fusions of selected proteins identified as being
induced by DR or high osmolarity were grown in 0.5%, 2% and
20% glucose, separated by 1-D SDS-PAGE and western blotted
with anti-GFP antiserum. (C) Wild type BY4741 yeast grown in
0.5%, 2% and 20% glucose (left panel); or wild type and isogenic PLoS ONE | www.plosone.org July 2012 | Volume 7 | Issue 7 | e41975 10 Hsp12 Structure and Function hsp12D strains grown in 0.05%, 0.5% and 2% glucose (right panel)
were separated by 1-D SDS-PAGE and western blotted with anti-
Hsp12 antiserum. (TIF) hsp12D strains grown in 0.05%, 0.5% and 2% glucose (right panel)
were separated by 1-D SDS-PAGE and western blotted with anti-
Hsp12 antiserum. (TIF) Hsp12 Structure and Function 27. Gasch AP, Spellman PT, Kao CM, Carmel-Harel O, Eisen MB, et al. (2000)
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Msn4p are required for the induction of subsets of high osmolarity glycerol
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coated microtitre plates: increased throughput, reproducibility and speed
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analysis of the response to uncapped telomeres in budding yeast reveals a novel
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CCPN data model for NMR spectroscopy: development of a software pipeline. Proteins 59: 687–696. 39. Heinig M, Frishman D (2004) STRIDE: a web server for secondary structure
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myces cerevisiae, in aqueous solution where it is intrinsically disordered and in
detergent micelles where it is locally alpha-helical. J Biol Chem 286: 43447–
43453. 41. References Lin SJ, Kaeberlein M, Andalis AA, Sturtz LA, Defossez PA, et al. (2002) Calorie
restriction extends Saccharomyces cerevisiae lifespan by increasing respiration. Nature 418: 344–348. 22. Defossez P-A, Prusty R, Kaeberlein M, Lin S-J, Ferrigno P, et al. (1999)
Elimination of replication block protein Fob1 extends the life span of yeast
mother cells. Molecular Cell 3: 447–455. 10. Anderson RM, Bitterman KJ, Wood JG, Medvedik O, Sinclair DA (2003)
Nicotinamide and PNC1 govern lifespan extension by calorie restriction in
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Hsp26: a temperature-regulated chaperone. Embo J 18: 6744–6751. 11. Lamming DW, Latorre-Esteves M, Medvedik O, Wong SN, Tsang FA, et al. (2005) HST2 mediates SIR2-independent life-span extension by calorie
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span extension by calorie restriction in yeast. PLoS Biol 2: E296. span extension by calorie restriction in yeast. PLoS Biol 2: E296 13. Kaeberlein M, Powers RW, 3rd, Steffen KK, Westman EA, Hu D, et al. (2005)
Regulation of yeast replicative life span by TOR and Sch9 in response to
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alpha-helices: local motifs, long-range electrostatics, ionic-strength dependence
and prediction of NMR parameters. J Mol Biol 284: 173–191. PLoS ONE | www.plosone.org 11 July 2012 | Volume 7 | Issue 7 | e41975 July 2012 | Volume 7 | Issue 7 | e41975 Hsp12 Structure and Function Chou JJ, Gaemers S, Howder B, Louis JM, Bax A (2001) A simple apparatus for
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Monitoring Glacier Calving using Underwater Sound Abstract. Climate shifts are particularly conspicuous in the Arctic. Satellite and terrestrial observations show significant in-
creases in the melting and breakup of Arctic tidewater glaciers and their influence on sea level rise. Increasing melt rates are
creating an urgency to better understand the link between atmospheric and oceanic conditions and glacier frontal ablation
through iceberg calving and melting. Elucidating this link requires a combination of short and long-time scale measurements
of terminus activity. Recent work has demonstrated the potential of using underwater sound to quantify the time and scale
5
of calving events to yield integrated estimates of ice mass loss (Glowacki and Deane, 2020). Here, we present estimates of Abstract. Climate shifts are particularly conspicuous in the Arctic. Satellite and terrestrial observations show significant in-
creases in the melting and breakup of Arctic tidewater glaciers and their influence on sea level rise. Increasing melt rates are
creating an urgency to better understand the link between atmospheric and oceanic conditions and glacier frontal ablation
through iceberg calving and melting. Elucidating this link requires a combination of short and long-time scale measurements of terminus activity. Recent work has demonstrated the potential of using underwater sound to quantify the time and scale
5
of calving events to yield integrated estimates of ice mass loss (Glowacki and Deane, 2020). Here, we present estimates of
subaerial calving flux using underwater sound recorded at Hansbreen, Svalbard in September 2013 combined with an algo-
rithm for the automatic detection of calving events. The method is compared with ice calving volumes estimated from geodetic
measurements of the movement of the glacier terminus and an analysis of satellite images. The total volume of above-water 5 10 calving during the 26 days of acoustical observation is estimated to be 1.7 ± 0.7 × 107 m3, whereas the subaerial calving flux
10
estimated by traditional methods is 7 ± 2 × 106 m3. The results suggest that passive cryoacoustics is a viable technique for
long-term monitoring of mass loss from marine-terminating glaciers. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Monitoring Glacier Calving using Underwater Sound
Jarosław T˛egowski1,*, Oskar Glowacki2,*, Michał Ciepły3, Małgorzata Błaszczyk3, Jacek Jania3,
Mateusz Moskalik2, Philippe Blondel4, and Grant B. Deane5
1Institute of Oceanography, University of Gdansk, Gdynia, Poland
2Institute of Geophysics, Polish Academy of Sciences, Warsaw, Poland
3Faculty of Natural Sciences, Institute of Earth Sciences, University of Silesia in Katowice, Sosnowiec, Poland
4Department of Physics, University of Bath, Bath, UK
5Scripps Institution of Oceanography, UCSD, La Jolla, California, USA
*These authors contributed equally to this work. Correspondence: Oskar Glowacki (oglowacki@igf.edu.pl), Grant B. Deane (gdeane@ucsd.edu) 1
Introduction The loss in the mass of the Greenland ice sheet, which increased from 41 ± 17 Gt yr−1 in 1990–2000 to 286 ± 20 Gt yr−1 in
2010-2018 (Mouginot et al., 2019), is of great concern. Greenland’s contribution to sea level rise by the end of the 21st century
15
is estimated to be 90 ± 50 and 32 ± 17 mm for RCP8.5 and RCP2.6 greenhouse gas concentration scenarios, respectively
(Goelzer et al., 2020); these estimates exclude peripheral glaciers and ice caps. Moreover, recent studies have shown that the
accelerated break-up and melting of Antarctica – the largest reservoir of fresh water on Earth – will be responsible for a sea
level rise of at least 0.5 cm yr−1 by 2100 (Paolo et al., 2015; DeConto et al., 2021). Despite these concerning numbers, the loss of land-based ice in the coming decades could be under-predicted. For example, Greenland and Antarctic ice sheets likely have
20
tipping points at around 1.5 −2.0◦C of a mean global temperature increase compared to the pre-industrial era (Pattyn et al.,
2018; Pattyn and Morlighem, 2020; Noël et al., 2021). The severe consequences of sea level rise and current uncertainties in 1 1 Calving is a complex process that depends on a broad range of factors, including but not limited to (i) changes in
glacier surface velocity along the flow line (longitudinal stretching), (ii) submarine melting of the terminus (undercutting),
and (iii) geometry of the glacier-bay system (van der Veen, 2002; Benn et al., 2007b). This complexity makes the formulation
of a general model of how calving changes in response to warming conditions a challenging task that requires long-term, temporally-resolved data sets (e.g., van der Veen, 2002; Benn et al., 2007a). 40
Passive cryoacoustics, the use of ambient sounds to study ice-ocean interactions in polar regions, has the potential to provide
the long-term, temporally-resolved time series required for calving model development and monitoring the long-term stability
of glaciers and ice sheets (see discussion in Glowacki and Deane, 2020), provided that the geophysical signals of relevance
can be recovered from the properties of the sound. The field of ambient noise oceanography is well-established and has been used to recover signals such as wind speed and precipitation, for example. The use of passive acoustics in polar regions is a
45
relatively new field, still under development (Glowacki et al., 2015; Pettit et al., 2015; Deane et al., 2019; Yun et al., 2021;
Podolskiy et al., 2022). A recent milestone has been the demonstration that the mass of a calving iceberg can be estimated from its underwater
impact noise with the sea surface (Glowacki and Deane, 2020). The method exploits the observed power law relationship impact noise with the sea surface (Glowacki and Deane, 2020). The method exploits the observed power law relationship
between the kinetic energy of a falling ice block immediately prior to impact and the resulting sound energy produced. The
50
kinetic energy of an impacting block can be estimated from the sound energy radiated provided that propagation losses between
the block and the recording location can be estimated. Because the drop height of icebergs is limited by the vertical extent of
the terminus, the estimate of impact energy can then be converted into a block mass using what Glowacki and Deane (2020)
call a "mass-weighted average drop height", ˆh. between the kinetic energy of a falling ice block immediately prior to impact and the resulting sound energy produced. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. models of glacial retreat suggest that long-term monitoring of glacial stability is both important and urgent (Straneo et al.,
2019). There are many challenges to the long-term monitoring of glaciers and ice shelves. Polar regions are remote and plunged
25
into darkness for up to 6 months a year. Moreover, capturing the episodic character of ice breakup usually requires sub-second
temporal resolution over year-long time scales. Despite a suite of current techniques, including satellite technology (Smith
et al., 2020; Podgórski and P˛etlicki, 2020), camera observations (Medrzycka et al., 2016; How et al., 2019), terrestrial laser
scanning (P˛etlicki and Kinnard, 2016; Podgórski et al., 2018), ground-based radar imaging (Xie et al., 2018; Walter et al., 25 2020), sonar scans (Sugiyama et al., 2019; Sutherland et al., 2019), and seismic records (Köhler et al., 2016; Podolskiy and
30
Walter, 2016), continuous and long-term observations of the scale and rapidity of ice loss in logistically challenging polar
regions are still urgently required (Straneo et al., 2013). There are 3 major processes driving mass loss from marine-terminating glaciers: surface melting, submarine melting, and
calving, which is the mechanical loss of ice from the edges of glaciers or ice shelves (Benn et al., 2007b). Calving and 2020), sonar scans (Sugiyama et al., 2019; Sutherland et al., 2019), and seismic records (Köhler et al., 2016; Podolskiy and
30
Walter, 2016), continuous and long-term observations of the scale and rapidity of ice loss in logistically challenging polar
regions are still urgently required (Straneo et al., 2013). There are 3 major processes driving mass loss from marine-terminating glaciers: surface melting, submarine melting, and
calving, which is the mechanical loss of ice from the edges of glaciers or ice shelves (Benn et al., 2007b). Calving and regions are still urgently required (Straneo et al., 2013). There are 3 major processes driving mass loss from marine-terminating glaciers: surface melting, submarine melting, and
calving, which is the mechanical loss of ice from the edges of glaciers or ice shelves (Benn et al., 2007b). Calving and submarine melting - the two mechanisms of frontal ablation - are very difficult to quantify separately (Truffer and Motyka,
35
2016). 2
Study Site The site for this study is Hansbreen, a retreating, grounded, polythermal tidewater glacier terminating in Hornsund Fjord,
60
Svalbard. Its calving mode, believed to be driven by melt undercutting and many relatively small calving events from above
the water surface (P˛etlicki et al., 2015), provides a challenging test case for camera and satellite estimates of mass flux. A schematic of the study site is shown in Fig. 1. Hansbreen covers an area of around 54 km2, is more than 15 km long
(Błaszczyk et al., 2013), and has a 1.5 km-wide active calving front with an average height of around 40 m (Błaszczyk et al., 2009, 2021), which is approximately 30–40% of the total ice thickness. The glacier surface flow is dominated by basal motion
65
in the ablation area (Vieli et al., 2004) and the mean annual flow velocity near the terminus and its calving flux is estimated to
be 177 my−1 and 35×106 m3y−1, respectively (Błaszczyk et al., 2019). Both glacial behavior (P˛etlicki et al., 2015; Błaszczyk et al., 2021) and the propagation of calving sounds (Glowacki et al.,
2016) are sensitive to the space and time-varying thermohaline structure of water masses in the bay, which were characterized with CTD casts. The water temperature and salinity in the center of the bay ranged from -1.8 °C to more than 2.0 °C and from
70
30 PSU to almost 35 PSU during 2015 and 2016 (Moskalik et al., 2018). Sound propagation also depends on water depth,
which drops to almost 100 m along a transect parallel to the glacier terminus (see Fig. 1A). The morphology of the bay varies
greatly, with numerous moraines and flat areas created during the retreat of Hansbreen (Moskalik et al., 2018). with CTD casts. The water temperature and salinity in the center of the bay ranged from -1.8 °C to more than 2.0 °C and from
70
30 PSU to almost 35 PSU during 2015 and 2016 (Moskalik et al., 2018). Sound propagation also depends on water depth,
which drops to almost 100 m along a transect parallel to the glacier terminus (see Fig. 1A). The morphology of the bay varies
greatly, with numerous moraines and flat areas created during the retreat of Hansbreen (Moskalik et al., 2018). https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. events and the removal of contaminating signals, such as iceberg disintegration. Acoustic estimates of the ice volume loss are
compared with concurrent camera and satellite observations. events and the removal of contaminating signals, such as iceberg disintegration. Acoustic estimates of the ice volume loss are
compared with concurrent camera and satellite observations. The
50
kinetic energy of an impacting block can be estimated from the sound energy radiated provided that propagation losses between
the block and the recording location can be estimated. Because the drop height of icebergs is limited by the vertical extent of
the terminus, the estimate of impact energy can then be converted into a block mass using what Glowacki and Deane (2020)
call a "mass-weighted average drop height", ˆh. Here we demonstrate the use of passive cryoacoustics to monitor the subaerial calving flux from a tidewater glacier in
55
southwestern Spitsbergen over 26 days of continuous measurements using algorithms for the automatic detection of calving Here we demonstrate the use of passive cryoacoustics to monitor the subaerial calving flux from a tidewater glacier in
55
southwestern Spitsbergen over 26 days of continuous measurements using algorithms for the automatic detection of calving 2 2 3
Automatic Detection of Calving Events Glowacki and Deane (2020) have shown that the mass of individual icebergs can be estimated from the sound they produce
75
when they impact the sea surface. In that study, 169 subaerial calving events from Hansbreen were manually identified in
time-lapse photography and simultaneous recordings of the ambient sound, both made roughly 1 km from the glacier terminus. Despite significant variability in the total sound energy produced by an impact event for icebergs of comparable size, Monte
Carlo simulations of ensemble time-series show that accurate estimates of subaerial calving flux can be made if sufficient
events are averaged (40 events for 20% standard error for Hansbreen). 80 events are averaged (40 events for 20% standard error for Hansbreen). 80
The manual identification of calving events using time-lapse photography and underwater sound is accurate but too la-
bor intensive to be seriously considered for long-term data sets. This issue is addressed here by developing algorithms for
the automatic detection of ice impact noise in the ambient sound recordings and rejection of interfering events, such as the
disintegration of icebergs in the bay to obtain accurate estimates of ice loss from the terminus of Hansbreen due to calving. The manual identification of calving events using time-lapse photography and underwater sound is accurate but too la-
bor intensive to be seriously considered for long-term data sets. This issue is addressed here by developing algorithms for
the automatic detection of ice impact noise in the ambient sound recordings and rejection of interfering events, such as the
disintegration of icebergs in the bay to obtain accurate estimates of ice loss from the terminus of Hansbreen due to calving. The automatic detection of calving events is based on the observation that the ambient sound field in a glacial bay is
85
composed of two distinct categories of sound sources: 1. sources that radiate noise with statistical properties that are stationary
on the time-scale of a calving event (such as melting glacier ice, for example) and 2. transient sources, like calving, superposed The automatic detection of calving events is based on the observation that the ambient sound field in a glacial bay is
85
composed of two distinct categories of sound sources: 1. sources that radiate noise with statistical properties that are stationary
on the time-scale of a calving event (such as melting glacier ice, for example) and 2. transient sources, like calving, superposed 3 3 The sound power, Pb, between a lower and upper frequency band, f0 = 30 Hz to f1 = 100 Hz is then computed
95
using The signal processing scheme for event detection works as follows. A 10 minute segment of ambient sound is analyzed into
a spectrogram. The sound power, Pb, between a lower and upper frequency band, f0 = 30 Hz to f1 = 100 Hz is then computed
95
using Pb = B(Ω) ⊗
M(Θ) ⊗
f1
Z
f0
Pxxdf
,
(1) Pb = B(Ω) ⊗
M(Θ) ⊗
f1
Z
f0
Pxxdf
, (1) where Pxx is the power spectral density estimate for the noise segment and the operators M(Θ) and B(Ω) are the filters
described below. The selection of frequency limits is motivated by previous analyses of the calving noise presented in Glowacki and Deane (2020) and Glowacki (2020) (see Appendix A: Materials and Methods for more details). The integral in equation
100
1 is calculated using the trapezoidal rule. The filter M(Θ) is a median filter of order Θ = 7 and is useful for eliminating loud,
impulsive noise sources that are too short to be calving noise. A boxcar filter B(Ω) of length Ω= 32 points is applied to further
reduce the effects of impulsive noise sources and random variations in noise power. and Deane (2020) and Glowacki (2020) (see Appendix A: Materials and Methods for more details). The integral in equation
100
1 is calculated using the trapezoidal rule. The filter M(Θ) is a median filter of order Θ = 7 and is useful for eliminating loud,
impulsive noise sources that are too short to be calving noise. A boxcar filter B(Ω) of length Ω= 32 points is applied to further
reduce the effects of impulsive noise sources and random variations in noise power. A baseline power for the noise in the absence of calving, Pbase, is taken to be the median of the probability density distri-
bution of Pb. A threshold of detection, Pthres, is then selected to be 1.5 times the largest value of Pb with relative likelihood
105 A baseline power for the noise in the absence of calving, Pbase, is taken to be the median of the probability density distri-
bution of Pb. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. on this stationary background. During periods when noise in the bay is dominated by ice melting and calving, the signal during
a 10-minute data segment has the form of a wandering baseline punctuated at random intervals by hill-shaped increases as
l i
t
Th l
l
d
i i
f th b
li
i
ti
d
t
h
i th
t
f i
lti
th
90 on this stationary background. During periods when noise in the bay is dominated by ice melting and calving, the signal during
a 10-minute data segment has the form of a wandering baseline punctuated at random intervals by hill-shaped increases as
calving events occur. The level and noisiness of the baseline varies over time due to changes in the rate of ice melting on the
90
glacier terminus, proximity to the hydrophone of icebergs drifting in the bay, and variable acoustic propagation through the
bay. These varying conditions are unpredictable and require the selection of a baseline level and event detection threshold that
is adaptable on this stationary background. During periods when noise in the bay is dominated by ice melting and calving, the signal during
a 10-minute data segment has the form of a wandering baseline punctuated at random intervals by hill-shaped increases as calving events occur. The level and noisiness of the baseline varies over time due to changes in the rate of ice melting on the
90
glacier terminus, proximity to the hydrophone of icebergs drifting in the bay, and variable acoustic propagation through the
bay. These varying conditions are unpredictable and require the selection of a baseline level and event detection threshold that
is adaptable. calving events occur. The level and noisiness of the baseline varies over time due to changes in the rate of ice melting on the
90
glacier terminus, proximity to the hydrophone of icebergs drifting in the bay, and variable acoustic propagation through the
bay. These varying conditions are unpredictable and require the selection of a baseline level and event detection threshold that
is adaptable. 90 The signal processing scheme for event detection works as follows. A 10-minute segment of ambient sound is analyzed into
a spectrogram. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. A threshold of detection, Pthres, is then selected to be 1.5 times the largest value of Pb with relative likelihood
105 ϕ(Pthres) > ϕ(Pbase)e−β,
(2)
where β = 5. This constant plays a central role in detector performance and is referred to as the ’detection factor’ (see Appendix
A: Materials and Methods). ϕ(Pthres) > ϕ(Pbase)e−β, (2) ϕ(
thres)
ϕ(
base)
,
( )
where β = 5. This constant plays a central role in detector performance and is referred to as the ’detection factor’ (see Appendix
A: Materials and Methods). where β = 5. This constant plays a central role in detector performance and is referred to as the ’detection factor’ (see Appendix
A: Materials and Methods). Calving events are identified as continuous periods of time when Pb exceeds Pthres. The detection algorithm is based on the
110
idea that calving events are relatively rare within 10-minute intervals and periods of time containing many successive events
will drive the algorithm to select only the most energetic of them (see Appendix A: Materials and Methods). Calving events are identified as continuous periods of time when Pb exceeds Pthres. The detection algorithm is based on the
110
idea that calving events are relatively rare within 10-minute intervals and periods of time containing many successive events
will drive the algorithm to select only the most energetic of them (see Appendix A: Materials and Methods). 110 An example of the detector output is shown in Fig. 2. Figure 2A shows 1 minute of ambient sound energy containing
a calving event, presented in dB as a function of frequency and time. Figure 2B shows the output from the band-pass and median filters, the baseline level, the threshold level, and detection of the calving event. The detector was run on the 26 days of
115
recordings, resulting in the detection of 4258 events (see Appendix A: Materials and Methods for a discussion of how sensitive
the algorithm is to the choice of detection factor). The primary output of the event detector are the event start and end times, t0
and t1. 4 4 4
Interfering Event Removal The problem remains of identifying events that were detected as calving but which are not. Visual inspection of noise spec-
120
trograms combined with listening to the corresponding sound recordings revealed two major sources of interfering sounds: (1)
the disintegration of icebergs close to the acoustic buoy, which sounds like a calving event because the iceberg sheds blocks
of ice into the water as it disintegrates, and (2) water movement around the hydrophone (flow noise). Other interfering events
include calving from the terminus with highly-energetic contacts between the falling iceberg and the terminus, icebergs collid- ing with other ice pieces that occupy the sea surface along the terminus, and calving events involving ice block rotation in the
125
air, resulting in large contact area between the iceberg and sea surface leading to atypical sound production. Although all these
events represent a challenge for the technique, the most concerning are the flow noise and disintegrating icebergs, which can
appear to be very large calving events. The interfering events must be removed from the calving inventory before the ice mass
loss is estimated. To deal with the misclassification problem, we take two steps. ing with other ice pieces that occupy the sea surface along the terminus, and calving events involving ice block rotation in the
125
air, resulting in large contact area between the iceberg and sea surface leading to atypical sound production. Although all these
events represent a challenge for the technique, the most concerning are the flow noise and disintegrating icebergs, which can
appear to be very large calving events. The interfering events must be removed from the calving inventory before the ice mass
loss is estimated. To deal with the misclassification problem, we take two steps. First, we take advantage of the fact that submarine melting of glacier ice also generates underwater noise due to the impulsive
130
release of pressurized air bubbles into the water (Urick, 1971; Deane et al., 2014; Pettit et al., 2015). Glowacki et al. (2018)
have demonstrated that the melt noise has an α-stable distribution. Importantly, the characteristic exponent of the α-stable
distribution indicates proximity to a melting source. When the melting iceberg is close to the receiver, the underwater noise at
frequencies 1—10 kHz is more impulsive and the exponent α deviates from 2. 4
Interfering Event Removal Here, we assume the following: if α < 1.95 for First, we take advantage of the fact that submarine melting of glacier ice also generates underwater noise due to the impulsive
130
release of pressurized air bubbles into the water (Urick, 1971; Deane et al., 2014; Pettit et al., 2015). Glowacki et al. (2018)
have demonstrated that the melt noise has an α-stable distribution. Importantly, the characteristic exponent of the α-stable
distribution indicates proximity to a melting source. When the melting iceberg is close to the receiver, the underwater noise at
frequencies 1—10 kHz is more impulsive and the exponent α deviates from 2. Here, we assume the following: if α < 1.95 for First, we take advantage of the fact that submarine melting of glacier ice also generates underwater noise due to the impulsive
130
release of pressurized air bubbles into the water (Urick, 1971; Deane et al., 2014; Pettit et al., 2015). Glowacki et al. (2018)
have demonstrated that the melt noise has an α-stable distribution. Importantly, the characteristic exponent of the α-stable
distribution indicates proximity to a melting source. When the melting iceberg is close to the receiver, the underwater noise at
frequencies 1—10 kHz is more impulsive and the exponent α deviates from 2. Here, we assume the following: if α < 1.95 for the noise segment immediately preceding the calving noise, there is a high likelihood that the corresponding event detected by
135
the automated algorithm is in fact an iceberg disintegrating close to the hydrophone. Such events are discarded. In this case,
875 events were removed from the inventory. Second the noise segments are occasionally contaminated with flow noise which originates from water turbulence advected the noise segment immediately preceding the calving noise, there is a high likelihood that the corresponding event detected by
135
the automated algorithm is in fact an iceberg disintegrating close to the hydrophone. Such events are discarded. In this case,
875 events were removed from the inventory. Second, the noise segments are occasionally contaminated with flow noise, which originates from water turbulence advected
past the hydrophone and is most pronounced at frequencies below ∼50 Hz. An overlap in frequency between the calving 875 events were removed from the inventory. Second, the noise segments are occasionally contaminated with flow noise, which originates from water turbulence advected
past the hydrophone and is most pronounced at frequencies below ∼50 Hz. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. The total number of submarine calving events in the present inventory is not expected to be large. Reported statistics of
submarine calving from other study sites show that such events are typically much less common than subaerial calving (Minowa
et al., 2018; Köhler et al., 2019; How et al., 2019). Glowacki (2022) estimated that submarine calving events at Hansbreen
are 8–10 times less frequent than subaerial events. It must be noted, however, that the low rate of occurrence of submarine
155
calving is offset by the fact that submarine events often produce the biggest icebergs (Warren et al., 1995; Motyka, 1997;
O’Neel et al., 2007; Glowacki, 2022). Consequently, attributing this component of the total calving flux to subaerial events
may introduce an error disproportionately large relative to their number. In response to this problem, a new semi-automatic
method for distinguishing submarine and subaerial calving events was recently proposed (Glowacki, 2022). However, there is no way at the present time to classify calving styles from sound recordings using fully automated methods. The time and
160
frequency structure of the underwater noise from submarine and subaerial calving differs significantly (Glowacki, 2022); for
example, the underwater noise from submarine calving typically has longer duration, with individual sound-source mechanisms
separated by the quiescent periods. As a result, a universal calving detector should (i) take into account the differences in time
and frequency structure of the calving noise and (ii) be validated with high-frequency time-lapse images synchronized with long-term acoustic recordings; this task is beyond the scope of the present study. Keeping this limitation in mind, the section
165
that follows will explain how the subaerial calving flux can be estimated acoustically, with an assumption that all detected
events are subaerial. 4
Interfering Event Removal An overlap in frequency between the calving detector band (30–100 Hz) and the flow noise (2–50 Hz) can result in false classifications of the episodes of intense water flow
140
around the hydrophone as calving events. To deal with this issue, we discard all events for which the ratio between the average
power spectral density at 2–30 Hz and 50–1000 Hz is higher than or equal to 1. This procedure is based on the observation that
calving noise extends into the 50–1000 Hz band, whereas flow noise is most pronounced at frequencies below 30 Hz. Using
these criteria, 328 misclassifications were identified and manually validated by listening to the selected noise segments and visual inspections of spectrograms. In total, 1128 events were discarded due to the fact that 75 events were identified as both
145
flow noise and iceberg disintegration. One issue remains, which is the fact that some fraction of the event signals found come from submarine calving events. Submarine calving events are a problem because the parameters in the power law relationship between the energy of noise
production and kinetic energy of block impact are for subaerial events and will not apply to submarine events (Glowacki and Deane, 2020). Thus any submarine events included in the calving inventory will not have their volume calculated correctly. 150
Moreover, even if their volume were calculated correctly, they are not detected by the camera observations for this study. Deane, 2020). Thus any submarine events included in the calving inventory will not have their volume calculated correctly. 150
Moreover, even if their volume were calculated correctly, they are not detected by the camera observations for this study. 5 5 5
Estimating Ice Volume Loss Following event detection, the next step is to compute the total noise energy radiated by the calving event at a standard reference
distance of 1 m, Eac,imp. This step requires the calculation of the time and frequency integrated energy of the calving noise. 170
The energy of the impact sound radiated by a falling ice block can be expressed in terms of the observed sound energy by
accounting for propagation effects and the addition of background noise using: Eac,imp =
4π
ρwcw
f1
Z
f0
t0+∆t
Z
t0
Pxx dt
−
t0
Z
t0−∆t
Pxx dt
× 10
−T L(f)
10
df,
(3) (3) (3) where ρw is the water density, cw is the sound speed, f0 = 30 Hz and f1 = 100 Hz are lower and upper frequency limits, t0
is the start time of the calving event determined from the event detector, ∆t = t1 −t0 is the calving signal duration, where t1
5 where ρw is the water density, cw is the sound speed, f0 = 30 Hz and f1 = 100 Hz are lower and upper frequency limits, t0 where ρw is the water density, cw is the sound speed, f0
30 Hz and f1
100 Hz are lower and upper frequency limits, t0
is the start time of the calving event determined from the event detector, ∆t = t1 −t0 is the calving signal duration, where t1
175
is the end time of the calving signal, TL is the frequency-dependent transmission loss between the hydrophone and the point
of ice block impact (in decibels). The acoustic signal is integrated over the surface area of a unit sphere (4π) to obtain total
noise energy in joules. The two integrals over time,
R t0+∆t
t0
Pxx dt and
R t0
t0−∆t Pxx dt, are the frequency spectral densities of
the calving noise plus background noise, and background noise alone, respectively. The transmission loss is calculated using
the standard numerical code RAM (Collins, 1993) (see Appendix A: Materials and Methods for further details). 180 is the start time of the calving event determined from the event detector, ∆t = t1 −t0 is the calving signal duration, where t1
175
is the end time of the calving signal, TL is the frequency-dependent transmission loss between the hydrophone and the point
of ice block impact (in decibels). https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. In the next step, the kinetic energy of the falling block of ice, Eimp, is calculated from the total impact sound energy radiated
into the bay, Eac,imp, using the power law relationship: Eimp = γEκ
ac,imp + ζ,
(4)
where γ = 1.45 × 109, κ = 0.18 and ζ = −2.66 × 109 are constants determined using the calibration dataset collected in 2016 (4) Eimp = γEκ
ac,imp + ζ, Eimp = γEκ
ac,imp + ζ, where γ = 1.45 × 109, κ = 0.18 and ζ = −2.66 × 109 are constants determined using the calibration dataset collected in 2016
(Glowacki and Deane (2020); see Appendix A: Materials and Methods for details). Finally, the iceberg volume is calculated as 185 (Glowacki and Deane (2020); see Appendix A: Materials and Methods for details). Finally, the iceberg volume is calculated as 185 (Glowacki and Deane (2020); see Appendix A: Materials and Methods for details). Finally,
5 V = M
ρi
= Eimp
ρigˆh
,
(5) V = M
ρi
= Eimp
ρigˆh
, (5) (5) V = ρi
=
p
ρigˆh
, where M is the iceberg mass, ρi = 917 kg m−3 is assumed ice density, g = 9.81 m s−2 is the acc where M is the iceberg mass, ρi = 917 kg m−3 is assumed ice density, g = 9.81 m s−2 is the acceleration due to gravity and
ˆh
20 7
i
h
i h d
d
h i h
i
d f
h
i
f H
b
(Gl
ki
d D
2020) 6
Results
190 The cumulative subaerial ice volume loss over the 26 days of observation is 1.7±0.7×107 m3 from
the acoustic measurements and 7±2×106 m3 from satellite data combined with stake measurements of the glacier velocity. The
discrepancy may be due to several reasons. First, any submarine calving events are treated as subaerial in the acoustic analysis. Consequently, the ice volume loss from acoustics is almost certainly overestimated. This issue has been addressed recently by
205
Glowacki (2022) and can be incorporated into a long-term analysis once a method is developed for the automatic detection of
submarine calving. Second, the power law relationship in Eq. 4 is based on the calibration dataset that does not cover the full
range of impact noise energies observed in 2013 data (see Fig. A2 and discussion in Appendix A: Materials and Methods). Third, the location of the hydrophone – far from the terminus and in shallow water – was unfavorable because the propagation Consequently, the ice volume loss from acoustics is almost certainly overestimated. This issue has been addressed recently by
205
Glowacki (2022) and can be incorporated into a long-term analysis once a method is developed for the automatic detection of
submarine calving. Second, the power law relationship in Eq. 4 is based on the calibration dataset that does not cover the full
range of impact noise energies observed in 2013 data (see Fig. A2 and discussion in Appendix A: Materials and Methods). Third, the location of the hydrophone – far from the terminus and in shallow water – was unfavorable because the propagation Consequently, the ice volume loss from acoustics is almost certainly overestimated. This issue has been addressed recently by
205
Glowacki (2022) and can be incorporated into a long-term analysis once a method is developed for the automatic detection of
submarine calving. Second, the power law relationship in Eq. 4 is based on the calibration dataset that does not cover the full
range of impact noise energies observed in 2013 data (see Fig. A2 and discussion in Appendix A: Materials and Methods). Third, the location of the hydrophone – far from the terminus and in shallow water – was unfavorable because the propagation conditions of the calving noise are very different for different segments of the terminus (see the variable bathymetry along the
210
black dashed lines in Fig. 5
Estimating Ice Volume Loss The acoustic signal is integrated over the surface area of a unit sphere (4π) to obtain total
noise energy in joules. The two integrals over time,
R t0+∆t
t0
Pxx dt and
R t0
t0−∆t Pxx dt, are the frequency spectral densities of
the calving noise plus background noise, and background noise alone, respectively. The transmission loss is calculated using the standard numerical code RAM (Collins, 1993) (see Appendix A: Materials and Methods for further details). 180 6 6
Results
190 Figure 3 shows a comparison of two time series: (i) subaerial calving flux derived from iceberg impact noise and (ii) terminus
area loss estimated from camera observations over the 26 days of analysis (see Appendix A: Materials and Methods). The ice
volume loss was calculated every 3 hours, which is the sampling time of the camera system. Both data streams show variability
between samples and between each other. This is to be expected; Glowacki and Deane (2020) demonstrated that the acoustic Figure 3 shows a comparison of two time series: (i) subaerial calving flux derived from iceberg impact noise and (ii) terminus
area loss estimated from camera observations over the 26 days of analysis (see Appendix A: Materials and Methods). The ice
volume loss was calculated every 3 hours, which is the sampling time of the camera system. Both data streams show variability
between samples and between each other. This is to be expected; Glowacki and Deane (2020) demonstrated that the acoustic method is accurate if sufficient calving events are averaged but significant variability is observed in the signal between blocks
195
of comparable volume. Moreover, camera observations are sensitive not only to weather conditions but also to the location of
calving events due to the irregular shape of the terminus and oblique angle of view (see Fig. 1B). However, there is a clear
relationship between the acoustic measurements of subaerial calving fluxes and the time-lapse observation of the terminus area
loss, despite the limitations of both methods. The Pearson’s correlation coefficient between the two data streams smoothed with a 1-day running average is 0.6. 200
The question now is: how do the acoustic measurements of the total subaerial calving flux compare to results obtained with
more traditional methods? The cumulative subaerial ice volume loss over the 26 days of observation is 1.7±0.7×107 m3 from
the acoustic measurements and 7±2×106 m3 from satellite data combined with stake measurements of the glacier velocity. The
discrepancy may be due to several reasons. First, any submarine calving events are treated as subaerial in the acoustic analysis. with a 1-day running average is 0.6. 200
The question now is: how do the acoustic measurements of the total subaerial calving flux compare to results obtained with
more traditional methods? https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. (Glowacki and Deane, 2020). Finally, stake measurements of the ice velocity that were used to calculate subaerial calving flux
from satellite images are from August 2013, which is a month before the acoustic data were collected. It is likely that Hansbreen
accelerated in September due to heavy rainfall events; such ‘autumn acceleration’ has been reported for other tidewater glaciers
in Svalbard (e.g., Luckman et al., 2015; Schellenberger et al., 2015). Moreover, stake measurements do not capture ice velocity
5
variations along the glacier terminus. The results of the experiment presented here are encouraging, given the limitations listed
above and completely independent nature of the methods used to make the measurements. (Glowacki and Deane, 2020). Finally, stake measurements of the ice velocity that were used to calculate subaerial calving flux
from satellite images are from August 2013, which is a month before the acoustic data were collected. It is likely that Hansbreen
accelerated in September due to heavy rainfall events; such ‘autumn acceleration’ has been reported for other tidewater glaciers
in Svalbard (e.g., Luckman et al., 2015; Schellenberger et al., 2015). Moreover, stake measurements do not capture ice velocity
5
variations along the glacier terminus. The results of the experiment presented here are encouraging, given the limitations listed
above and completely independent nature of the methods used to make the measurements. 215 6
Results
190 1A), which has a large effect on the transmission losses of low-frequency sounds in a glacial bay 7 7 7
Long-term Monitoring of Subaerial Calving Fluxes The study of Glowacki and Deane (2020) demonstrated the potential for underwater ambient sound in glacial terminal bays
to provide estimates of subaerial calving flux. Guided by this new opportunity, an impact-to-noise conversion model has been
220
applied here to a roughly 1-month long continuous recording of ambient sound in the bay of Hansbreen in Svalbard to es-
timate subaerial calving flux. Good agreement is found between acoustic measurements of ice volume loss and the camera
observations of the terminus area loss. The cumulative subarial calving flux estimated using the acoustic approach is higher
than the ice loss derived using a pair of satellite images combined with stake measurements of the glacier velocity. Clearly, the acoustic method requires further improvements, but the results are encouraging, especially as acoustic recorders are increas-
225
ingly deployed in the Arctic for other studies, such as studies of biodiversity or human impacts like shipping, and they can be
harnessed to also provide measurements of any neighboring glaciers. This study reinforces the idea that passive underwater
acoustics offers a new and viable method for monitoring mass loss from marine-terminating glaciers and ice shelves. The development of algorithms for the automated detection of calving events and the removal of interfering events, such
as disintegrating icebergs and flow noise, creates the possibility of monitoring the stability of tidewater glacier termini and
230
ice shelves over long time scales. We envision a scenario where a number of relatively inexpensive autonomous underwa-
ter recording systems are deployed over year-long intervals around the termini of selected glaciers on a recurrent schedule. The analysis of the resulting signals would eventually allow glacial stability to be monitored over decadal and longer time
scales. Concurrent measurements of relevant environmental drivers, such as water temperature, insolation and precipitation for
example, could provide important insights into the processes controlling terminus ablation. 235 as disintegrating icebergs and flow noise, creates the possibility of monitoring the stability of tidewater glacier termini and
230
ice shelves over long time scales. We envision a scenario where a number of relatively inexpensive autonomous underwa-
ter recording systems are deployed over year-long intervals around the termini of selected glaciers on a recurrent schedule. The analysis of the resulting signals would eventually allow glacial stability to be monitored over decadal and longer time
scales. 7
Long-term Monitoring of Subaerial Calving Fluxes Concurrent measurements of relevant environmental drivers, such as water temperature, insolation and precipitation for example, could provide important insights into the processes controlling terminus ablation. 235
Personal experience making long-term recordings in the Arctic has taught us that not all deployed recording systems are
recovered. It is therefore essential to keep the unit cost of recording systems low to allow for as much redundancy as possible
within a fixed budget. Power consumption and data storage are two important drivers of cost. Requirements for both of these
factors can be reduced by recording ambient sound on a fixed schedule or recording whenever the sound level exceeds a preset example, could provide important insights into the processes controlling terminus ablation. 235
Personal experience making long-term recordings in the Arctic has taught us that not all deployed recording systems are
recovered. It is therefore essential to keep the unit cost of recording systems low to allow for as much redundancy as possible
within a fixed budget. Power consumption and data storage are two important drivers of cost. Requirements for both of these
factors can be reduced by recording ambient sound on a fixed schedule or recording whenever the sound level exceeds a preset
threshold. These two possibilities will now be discussed. 240 threshold. These two possibilities will now be discussed. 240
Figure 4 shows the effect of recording coverage on the estimated total number of calving events (A) and cumulative ice
volume loss (B) over the study period. A continuous acoustic record corresponds to ∆= 0%. Periods of active recording
change from 1 hour per day (≈4%) to full coverage. Final estimates are derived proportionally, i.e. values obtained for 10%
and 20% coverage are multiplied by 10 and 5, respectively. The specific hours of recordings during a day are selected repeatedly threshold. These two possibilities will now be discussed. 240
Figure 4 shows the effect of recording coverage on the estimated total number of calving events (A) and cumulative ice
volume loss (B) over the study period. A continuous acoustic record corresponds to ∆= 0%. Periods of active recording
change from 1 hour per day (≈4%) to full coverage. Final estimates are derived proportionally, i.e. values obtained for 10%
and 20% coverage are multiplied by 10 and 5, respectively. The specific hours of recordings during a day are selected repeatedly threshold. These two possibilities will now be discussed. Exploiting this fact through the use of event-driven recording systemsi could result in significant power savings for autonomous recorders. 255 7
Long-term Monitoring of Subaerial Calving Fluxes 240
Figure 4 shows the effect of recording coverage on the estimated total number of calving events (A) and cumulative ice
volume loss (B) over the study period. A continuous acoustic record corresponds to ∆= 0%. Periods of active recording
change from 1 hour per day (≈4%) to full coverage. Final estimates are derived proportionally, i.e. values obtained for 10%
and 20% coverage are multiplied by 10 and 5, respectively. The specific hours of recordings during a day are selected repeatedly 8 8
Concluding Remarks This first automated analysis of underwater sound recorded near the terminus of a tidewater glacier, compared with camera
and satellite observations, demonstrates the feasibility of using cryoacoustics to monitor subaerial calving fluxes over extended
periods. Challenges remain. The method has been verified for Hansbreen, a glacier of a scale common to Spitsbergen, but the glaciers in Greenland can be factors of 10 or more larger in both horizontal and vertical scales. Moreover, calving mode and
260
propagation conditions will vary between glaciers and there will be variability in levels of ice coverage in the terminus bay,
which depend on calving rate, circulatory flow (long, narrow fjords are more prone to melange buildup), and sea ice coverage. Some or all of these factors may need to be accounted for when applying cryoacoustics to quantify subaerial calving flux
between tidewater glaciers. Cryoacoustics meets some of the important requirements for long-term monitoring of subaerial calving fluxes. The high
265
rate of signal acquisition ensures that every calving event can be detected, provided its signal is above the detection threshold
and below the rejection threshold. Although the uncertainty in the volume estimate for a single event is high, the total volume
estimate becomes precise if a sufficient number of events are accumulated (Glowacki and Deane, 2020). With current tech-
nological limitations, it is not possible to take time-lapse photographs with high enough frame rates to ensure that all events are captured. Moreover, cryoacoustics is insensitive to lighting conditions and relatively insensitive to weather conditions. For
270
example, the dominant spectral component of noise generated by rain lies well above the ’calving band’ (Pumphrey and Crum,
1988). Both cryoacoustics and cryoseismology provide temporally-resolved, continuous records of calving activity that are ex-
pected to be relatively insensitive to changing environmental conditions (Podolskiy and Walter, 2016; Deane et al., 2019). are captured. Moreover, cryoacoustics is insensitive to lighting conditions and relatively insensitive to weather conditions. For
270
example, the dominant spectral component of noise generated by rain lies well above the ’calving band’ (Pumphrey and Crum,
1988). Both cryoacoustics and cryoseismology provide temporally-resolved, continuous records of calving activity that are ex-
pected to be relatively insensitive to changing environmental conditions (Podolskiy and Walter, 2016; Deane et al., 2019). https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. using all possible combinations, creating a distribution of outputs for the chosen value of coverage. The event count is much
245
less sensitive to the recording schedule than the subaerial calving flux. Nevertheless, one standard deviation of the estimated
ice volume loss at 30% coverage is within 10% of the reference value. This result demonstrates the possibility of significant
reduction in recording time (by 70%) with relatively low increase in the resulting error. using all possible combinations, creating a distribution of outputs for the chosen value of coverage. The event count is much
245
less sensitive to the recording schedule than the subaerial calving flux. Nevertheless, one standard deviation of the estimated
ice volume loss at 30% coverage is within 10% of the reference value. This result demonstrates the possibility of significant
reduction in recording time (by 70%) with relatively low increase in the resulting error. Figure 5 shows the percentage of time occupied by calving noise during the experiment and the acoustic estimate of daily
calving rate. On average, calving events were active for only about 1% of the time; this corresponds to around 120 events per
250
day. However, we expect that the calving detector may split single calving events into several detections. This is especially
likely in the case of free falls of highly disintegrated icebergs. Therefore, we assume that the real daily calving rate is likely
lower than the acoustic estimate. Although calving activity changes significantly with seasons and geographical location, it is
still expected to be a small fraction of the total record. Exploiting this fact through the use of event-driven recording systems
could result in significant power savings for autonomous recorders. 255 calving rate. On average, calving events were active for only about 1% of the time; this corresponds to around 120 events per
250
day. However, we expect that the calving detector may split single calving events into several detections. This is especially
likely in the case of free falls of highly disintegrated icebergs. Therefore, we assume that the real daily calving rate is likely
lower than the acoustic estimate. Although calving activity changes significantly with seasons and geographical location, it is
still expected to be a small fraction of the total record. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. out the burden of a glacier-by-glacier calibration. Moreover, it may be possible to extract other geophysical signals of interest,
such as terminus melt rate, from cryoacoustic records (Pettit et al., 2015; Deane et al., 2019). Cryoacoustics, combined with other remote sensing methods, such as photogrammetry, cryoseismology and satellite obser-
vations, can form the core of a long-term monitoring system for subaerial calving fluxes from glaciers and ice shelves. 280 Data availability. The acoustic data used in this study are available upon request from J.T. (jaroslaw.tegowski@ug.edu.pl) Data availability. The acoustic data used in this study are available upon request from J.T. (jaroslaw.tegowski@ug.edu.pl) A1
Acoustic and Camera Observations Nevertheless, the image analysis provides useful information about the overall calving activity at Hansbreen during the
295
study period. frame. Nevertheless, the image analysis provides useful information about the overall calving activity at Hansbreen during the
295
study period. 8
Concluding Remarks Cryoacoustics has the additional benefit that the physics of sound production, which stems from the entrainment of bubbles
275
around falling blocks of ice and collective oscillations of bubble plumes (Glowacki, 2020), is expected to apply broadly across
calving glaciers in polar regions. If true, this will enable subaerial calving flux to be extracted from cryoacoustic signals with- Cryoacoustics has the additional benefit that the physics of sound production, which stems from the entrainment of bubbles
275
around falling blocks of ice and collective oscillations of bubble plumes (Glowacki, 2020), is expected to apply broadly across
calving glaciers in polar regions. If true, this will enable subaerial calving flux to be extracted from cryoacoustic signals with- Cryoacoustics has the additional benefit that the physics of sound production, which stems from the entrainment of bubbles
275
around falling blocks of ice and collective oscillations of bubble plumes (Glowacki, 2020), is expected to apply broadly across
calving glaciers in polar regions. If true, this will enable subaerial calving flux to be extracted from cryoacoustic signals with- 9 A2
Satellite Observations The volume of ice lost from Hansbreen through subaerial calving was estimated from satellite imagery and GPS observations
of the glacier surface movement. subaerial calving flux is estimated using Q = ¯Uice + UfrontLH, where Q is the volumetric The volume of ice lost from Hansbreen through subaerial calving was estimated from satellite imagery and GPS observations
of the glacier surface movement. subaerial calving flux is estimated using Q = ¯Uice + UfrontLH, where Q is the volumetric
flux of icebergs, ¯Uice is the mean ice flow velocity, Ufront is the front retreat/advance velocity, and L and H respectively are
300
the cliff length and height above sea level. The length of Hansbreen’s active cliff and the average front retreat per day have
been extracted from two multispectral Landsat 8 satellite images (2013-08-24 and 2013-09-25). Cliff height above sea level
was determined in 2015 using a Riegl VZ-6000 high resolution 3D laser scanner. The glacier surface velocity was estimated
using a mass balance stake mounted near the calving front (designated GPS in Fig. A1). Changes in stake position were
measured with precise dGPS in August 2013. The subaerial calving flux estimated over the 26 days of acoustic measurements
305 of the glacier surface movement. subaerial calving flux is estimated using Q = Uice + UfrontLH, where Q is the volumetric
flux of icebergs, ¯Uice is the mean ice flow velocity, Ufront is the front retreat/advance velocity, and L and H respectively are
300
the cliff length and height above sea level. The length of Hansbreen’s active cliff and the average front retreat per day have
been extracted from two multispectral Landsat 8 satellite images (2013-08-24 and 2013-09-25). Cliff height above sea level
was determined in 2015 using a Riegl VZ-6000 high resolution 3D laser scanner. The glacier surface velocity was estimated
using a mass balance stake mounted near the calving front (designated GPS in Fig. A1). Changes in stake position were flux of icebergs, ¯Uice is the mean ice flow velocity, Ufront is the front retreat/advance velocity, and L and H respectively are
300
the cliff length and height above sea level. The length of Hansbreen’s active cliff and the average front retreat per day have
been extracted from two multispectral Landsat 8 satellite images (2013-08-24 and 2013-09-25). Cliff height above sea level
was determined in 2015 using a Riegl VZ-6000 high resolution 3D laser scanner. A1
Acoustic and Camera Observations Acoustic and camera data were collected around Hansbreen from 5 – 30 September, 2013. Noise recordings were made with a
285
High Tech. Inc. HTI-96-MIN omnidirectional hydrophone mounted on the seafloor at a depth of 22 m and approximately 1.95
km from the glacier terminus. Data were sampled at a frequency of 32 kHz with 16 bits of dynamic range. Photographs of the
terminus were taken every 3 hours using a Canon D1000 time-lapse camera located 160 m above sea level and 0.8 to 2 km
from the glacier terminus (see Fig. 1). Acoustic and camera data were collected around Hansbreen from 5 – 30 September, 2013. Noise recordings were made with a
285
High Tech. Inc. HTI-96-MIN omnidirectional hydrophone mounted on the seafloor at a depth of 22 m and approximately 1.95
km from the glacier terminus. Data were sampled at a frequency of 32 kHz with 16 bits of dynamic range. Photographs of the
terminus were taken every 3 hours using a Canon D1000 time-lapse camera located 160 m above sea level and 0.8 to 2 km
from the glacier terminus (see Fig. 1). The images were analyzed using a differences technique to estimate the area of ice lost from the cliff face through calving
290
when lighting and weather conditions permitted. Changes at the calving front associated with calving events were found by
comparing two consecutive time-lapse images. The terminus area loss was then estimated using known ice cliff heights at
given locations. This methodology is sensitive to the variability of the terminus shape along its edge due to the oblique angle of
the camera view. Moreover, we are aware that the glacier can calve more than once at the same location during a 3-hour time The images were analyzed using a differences technique to estimate the area of ice lost from the cliff face through calving
290
when lighting and weather conditions permitted. Changes at the calving front associated with calving events were found by
comparing two consecutive time-lapse images. The terminus area loss was then estimated using known ice cliff heights at
given locations. This methodology is sensitive to the variability of the terminus shape along its edge due to the oblique angle of
the camera view. Moreover, we are aware that the glacier can calve more than once at the same location during a 3-hour time frame. A3
Calving Impact Energy Estimation The calving impact energy was estimated from the noise energy using a power-law relationship model shown in Eq. 4. The
parameters of the model are based on the calibration dataset presented in Glowacki and Deane (2020) that includes time-
315
lapse images and underwater noise recordings of 169 calving events observed in 2016 at Hansbreen. However, the power-law
model proposed by Glowacki and Deane (2020) was improved through the re-analysis of the calibration dataset. The following
changes/modifications were applied: The calving impact energy was estimated from the noise energy using a power-law relationship model shown in Eq. 4. The
parameters of the model are based on the calibration dataset presented in Glowacki and Deane (2020) that includes time-
315
lapse images and underwater noise recordings of 169 calving events observed in 2016 at Hansbreen. However, the power-law
model proposed by Glowacki and Deane (2020) was improved through the re-analysis of the calibration dataset. The following
changes/modifications were applied: 1. Subsequent to publication, it was discovered that the acoustic recordings contained a small DC offset, which created a
minor bias in the event energy estimates. This issue has been corrected here by running the acoustic recordings through
a (30 - 100) Hz band-pass filter, the lower frequency of which is roughly equal to the low-frequency cut-off of the
waveguide comprised of the sea surface and seafloor of the terminus bay. 2. The Bellhop ray-tracing model was used in Glowacki and Deane (2020) to calculate the loss of acoustic energy between
the iceberg/water impact locations and the hydrophone. Here, the transmission loss of the calving noise was calculated
using the parabolic equation model RAM (Collins, 1993) that is more recommended for low-frequency problems (Jensen
et al., 2011). The grain size of the sediments and source depth were set to ϕ = 5 and Zs = 5 m, respectively. The
attenuation, sound speed and density of the sediments were calculated from the grain size using formulas proposed by
Hamilton (1972) and Hamilton and Bachman (1982). Moreover, in the present study, transmission losses were evaluated
for a whole range of frequencies between 30 and 100 Hz, while a single nominal source frequency of 50 Hz was used in
Glowacki and Deane (2020). 330 3. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. the glacier surface velocity field was estimated using offset tracking from TerraSAR satellite radar images (2012-12-15 and
2012-12-26). subaerial calving flux was then estimated within 40 m sections of the glacier front to allow for variations in
velocity and cliff height (see the grey profile in Fig. A1). The total volume of icebergs calved during the observation period is
estimated to be (6±1)×106 m3. It should be borne in mind, however, that TerraSAR images were collected in December 2012
0
and not during the study period. Nevertheless, the results show good agreement with subaerial calving flux estimated using
stake measurements. the glacier surface velocity field was estimated using offset tracking from TerraSAR satellite radar images (2012-12-15 and
2012-12-26). subaerial calving flux was then estimated within 40 m sections of the glacier front to allow for variations in
velocity and cliff height (see the grey profile in Fig. A1). The total volume of icebergs calved during the observation period is
estimated to be (6±1)×106 m3. It should be borne in mind, however, that TerraSAR images were collected in December 2012
0
and not during the study period. Nevertheless, the results show good agreement with subaerial calving flux estimated using
stake measurements. A2
Satellite Observations The glacier surface velocity was estimated
using a mass balance stake mounted near the calving front (designated GPS in Fig. A1). Changes in stake position were measured with precise dGPS in August 2013. The subaerial calving flux estimated over the 26 days of acoustic measurements
305
is (7 ± 2) × 106 m3, i.e. ca. 0.27 × 106 m3 d−1. This is the result used for comparison with the acoustic data. As a crosscheck, 10 Leaving aside the causes, the lack of the highest impact noise energies in
the calibration dataset results in extrapolation of the power-law model in the analysis of data collected in 2013; unfortunately, the extrapolation applies to calving events that contribute the most to the subaerial calving flux. This is one possible explanation
355
of the discrepancy between calving fluxes derived from traditional methods and the acoustic technique. The improvement of the power-law model in Eq. 4 requires long-term acoustic measurements close to different glaciers
combined with measurements of ice block volumes with optical techniques (e.g., time-lapse photography, terrestrial laser
scanning). Collecting calibration data that would include a whole range of styles and magnitudes of calving events is critical
for the usefulness of the acoustic technique in the monitoring of subaerial calving fluxes. 360 the extrapolation applies to calving events that contribute the most to the subaerial calving flux. This is one possible explanation
355
of the discrepancy between calving fluxes derived from traditional methods and the acoustic technique. The improvement of the power-law model in Eq. 4 requires long-term acoustic measurements close to different glaciers
combined with measurements of ice block volumes with optical techniques (e.g., time-lapse photography, terrestrial laser
scanning). Collecting calibration data that would include a whole range of styles and magnitudes of calving events is critical
for the usefulness of the acoustic technique in the monitoring of subaerial calving fluxes. 360 scanning). Collecting calibration data that would include a whole range of styles and magnitudes of calving events is critical
for the usefulness of the acoustic technique in the monitoring of subaerial calving fluxes. 360 A3
Calving Impact Energy Estimation In Glowacki and Deane (2020), the parameters of the power-law relationship between the impact energy and noise
energy were estimated by performing a linear least-mean-squares analysis of log-transformed variables; the impact noise
energy was used as a dependent variable to estimate the impact-to-noise conversion coefficients. Here, the block impact
energy was used as a dependent variable to minimize the error in the subaerial calving flux. Moreover, the coefficients of
the power-law relationship were estimated using a non-linear regression model and non-transformed variables (function
fitnlm in Matlab). 335 Problems remain. Most importantly, the calibration dataset is limited to 169 calving events and certainly do not represent the
full range of possible values of the impact noise energy at Hansbreen. As a consequence, the power-law model – that is based 11 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Second, the 2016 data do not include calving events with Eac,imp > 4000 joules either. The lack of the largest events in the
calibration dataset may be due to several reasons: (i) Hansbreen may have been in different melting regimes in 2013 and 2016,
which caused differences in calving styles (e.g., differences in the water depth at the terminus, glacier surface velocity, water
content, etc.), (ii) co-occurrence of large-scale calving events and bad weather/lighting conditions that prevented estimation of
350
impact energy from time-lapse images in the calibration dataset, (iii) underestimated transmission loss for the acoustic data
collected in 2013 because of the different location of the hydrophone compared to 2016 leading to a different propagation path
(note the shallow sill close to the recorder in Fig. 1A). Leaving aside the causes, the lack of the highest impact noise energies in
the calibration dataset results in extrapolation of the power-law model in the analysis of data collected in 2013; unfortunately,
the extrapolation applies to calving events that contribute the most to the subaerial calving flux. This is one possible explanation
355
f h di
b
l i
fl
d i
d f
di i
l
h d
d h
i
h i content, etc.), (ii) co-occurrence of large-scale calving events and bad weather/lighting conditions that prevented estimation of
350
impact energy from time-lapse images in the calibration dataset, (iii) underestimated transmission loss for the acoustic data
collected in 2013 because of the different location of the hydrophone compared to 2016 leading to a different propagation path
(note the shallow sill close to the recorder in Fig. 1A). Leaving aside the causes, the lack of the highest impact noise energies in
the calibration dataset results in extrapolation of the power-law model in the analysis of data collected in 2013; unfortunately, content, etc.), (ii) co-occurrence of large-scale calving events and bad weather/lighting conditions that prevented estimation of
350
impact energy from time-lapse images in the calibration dataset, (iii) underestimated transmission loss for the acoustic data
collected in 2013 because of the different location of the hydrophone compared to 2016 leading to a different propagation path
(note the shallow sill close to the recorder in Fig. 1A). A4
Calving Event Detection Algorithm The calving event detection algorithm depends on the selection of an event detection factor, β, and a frequency band over
which the ambient sound is filtered before integration over time. Figure A3 shows a summary of how the number of detected calving events varies with the detection factor β; interfering
events were not removed in this analysis. As might be expected, the total number of events decreases rapidly with increasing
365
detection factor because increasing β results in the exclusion of the lowest-energy events. The results presented in Fig. 3 were
generated using β = 5, which was found to be an optimal selection in terms of limiting detection of non-calving events and
minimizing the loss of true calving events. The selection of β consisted of the visual analysis of noise spectrograms and audible
inspection of sound recordings performed for a number of detections. g
y
g
β;
g
events were not removed in this analysis. As might be expected, the total number of events decreases rapidly with increasing
365
detection factor because increasing β results in the exclusion of the lowest-energy events. The results presented in Fig. 3 were
generated using β = 5, which was found to be an optimal selection in terms of limiting detection of non-calving events and
minimizing the loss of true calving events. The selection of β consisted of the visual analysis of noise spectrograms and audible
inspection of sound recordings performed for a number of detections. The detection factor selected here for the terminal bay of Hansbreen is likely not the optimal detection factor for other
370
environments or geometries for the terminus and recording station. This is because the detector performance is sensitive to the The detection factor selected here for the terminal bay of Hansbreen is likely not the optimal detection factor for other
370
environments or geometries for the terminus and recording station. This is because the detector performance is sensitive to the 12 The calving detector in the present
form has two major weaknesses: (1) subaerial and submarine calving events cannot be distinguished and treated separately
and (2) certainly there are still some unexplored interfering events that either cause false detections or periodically lower the
signal-to-noise ratio, which makes calving detection more difficult (i.e., some calving events remain undetected). The latter baseline noise level. 385
Here we use a simple but robust algorithm for the detection of subaerial calving events. The calving detector in the present
form has two major weaknesses: (1) subaerial and submarine calving events cannot be distinguished and treated separately
and (2) certainly there are still some unexplored interfering events that either cause false detections or periodically lower the
signal-to-noise ratio, which makes calving detection more difficult (i.e., some calving events remain undetected). The latter baseline noise level. 385
Here we use a simple but robust algorithm for the detection of subaerial calving events. The calving detector in the present
form has two major weaknesses: (1) subaerial and submarine calving events cannot be distinguished and treated separately
and (2) certainly there are still some unexplored interfering events that either cause false detections or periodically lower the
signal-to-noise ratio, which makes calving detection more difficult (i.e., some calving events remain undetected). The latter issue can be partly addressed with the use of more sophisticated detection techniques. See, for example, recent work in the
390
cryoseismology community by Carr et al. (2020) and Köhler et al. (2022). However, we speculate that better understanding
of different (non-calving) sound-source mechanisms and their underwater acoustic signatures is required before more sophis-
ticated algorithms are implemented. Examples may include but are not limited to interactions between floating growlers and
icebergs, ice fracturing events, water outflows from subglacial conduits, coastline landslides, and activity of marine mammals. issue can be partly addressed with the use of more sophisticated detection techniques. See, for example, recent work in the
390
cryoseismology community by Carr et al. (2020) and Köhler et al. (2022). However, we speculate that better understanding
of different (non-calving) sound-source mechanisms and their underwater acoustic signatures is required before more sophis-
ticated algorithms are implemented. Examples may include but are not limited to interactions between floating growlers and
icebergs, ice fracturing events, water outflows from subglacial conduits, coastline landslides, and activity of marine mammals. Most of these “interfering” noise sources can be investigated with meteorological and oceanographic measurements, high-
395
frequency photographic images, laser scans, or radar scans synchronized with acoustic recordings. For example, Deane et al. (2014) reported that interactions of surface gravity waves with underside of ice ledges at the periphery of icebergs are sources
of underwater noise emission below 500 Hz. Some sound-source mechanisms, like underwater fracturing events, require novel
observational techniques or scaled laboratory experiments. Most of these “interfering” noise sources can be investigated with meteorological and oceanographic measurements, high-
395
frequency photographic images, laser scans, or radar scans synchronized with acoustic recordings. For example, Deane et al. (2014) reported that interactions of surface gravity waves with underside of ice ledges at the periphery of icebergs are sources
of underwater noise emission below 500 Hz. Some sound-source mechanisms, like underwater fracturing events, require novel
observational techniques or scaled laboratory experiments. The problem remains with detection of submarine events. An inclusion of submarine events would be beneficial for two
400
reasons: (i) to find out how frequent these events are and (ii) to provide more accurate estimates of subaerial calving fluxes
by rejecting submarine events from the analysis. Moreover, it should not be ruled out that the methodology for estimating
submarine calving fluxes from sound recordings made in glacial bays could be developed. If so, it would be possible to derive
both subaerial and submarine calving fluxes using acoustic techniques. Recent work by Glowacki (2022) demonstrated that The problem remains with detection of submarine events. An inclusion of submarine events would be beneficial for two
400
reasons: (i) to find out how frequent these events are and (ii) to provide more accurate estimates of subaerial calving fluxes
by rejecting submarine events from the analysis. Moreover, it should not be ruled out that the methodology for estimating
submarine calving fluxes from sound recordings made in glacial bays could be developed. If so, it would be possible to derive
both subaerial and submarine calving fluxes using acoustic techniques. Recent work by Glowacki (2022) demonstrated that the two calving modes can be acoustically distinguished using parameters of the log-normal distribution of the calving noise
405
combined with calving signal duration. A semi-automatic detection of start and end times of calving noise was used. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. signal-to-noise ratio at the hydrophone location and activity of other sources, such as nearby icebergs, which will vary from
site to site. It is noted in the main text that the event detection algorithm is based on the assumption that a good estimate for the
background noise is the median noise level in the detector frequency band. This assumption holds well when calving events are
375
relatively infrequent within an observational interval selected for analysis (10 minutes here) but may bias the detector toward
the exclusion of low energy events if too many events occur within the interval. The validity of the assumption was tested
by computing the background noise level iteratively: detected events are removed from the record and a baseline noise level
recomputed in successive passes through the data segment. The results of this analysis for change in number of events detected, total acoustic energy at the receiver and summed event durations are shown in Fig. A4. The total percentage change in acoustic
380
energy of all detected events is less than 3%, which is not a significant source of error given other, larger uncertainties in the
analysis, such as hydrophone calibration and propagation loss. The percentage change in number of events is similarly minor. There is a larger impact on the duration of events because of two effects: 1. increasing the baseline noise level increases the
detection threshold and decreases event duration and 2. the division of a single event into 2 or more events is sensitive to the total acoustic energy at the receiver and summed event durations are shown in Fig. A4. The total percentage change in acoustic
380
energy of all detected events is less than 3%, which is not a significant source of error given other, larger uncertainties in the
analysis, such as hydrophone calibration and propagation loss. The percentage change in number of events is similarly minor. There is a larger impact on the duration of events because of two effects: 1. increasing the baseline noise level increases the
detection threshold and decreases event duration and 2. the division of a single event into 2 or more events is sensitive to the baseline noise level. 385
Here we use a simple but robust algorithm for the detection of subaerial calving events. is co-architect of the sound inversion methodology and detection algorithm, participated
415
in field campaign, and participated in manuscript preparation. p
p
p p p
p
pi
p g
p
p
pi
p g
and participated in manuscript preparation. G.B.D. is co-architect of the sound inversion methodology and detection algorithm, participated
415
in field campaign, and participated in manuscript preparation. and participated in manuscript preparation. G.B.D. is co-architect of the sound inversion methodology and detection algorithm, participated
415
in field campaign, and participated in manuscript preparation. and participated in manuscript preparation. G.B.D. is co-architect of the sound inversion methodology and detection algorithm, participated
415
in field campaign, and participated in manuscript preparation. However, the two calving modes can be acoustically distinguished using parameters of the log-normal distribution of the calving noise
405
combined with calving signal duration. A semi-automatic detection of start and end times of calving noise was used. However, 13 Acknowledgements. This work has been supported by the National Science Centre, Poland (grants 2011/03/B/ST10/04275 and 2021/43/D/ST10/0
Ministry of Science and Higher Education of Poland (grant 1621/MOB/V/2017/0 and subsidy for the Institute of Geophysics, Polish Academy
of Sciences), U.S. Office of Naval Research, Ocean Acoustics Division (grant N00014-17-1-2633), USA National Science Foundation Office
420
of Polar Programs (grant OPP-1748265), and Research Council of Norway (Arctic Field Grant, RIS ID: 6133). We gratefully acknowledge
the support of the staff at the Polish Polar Research Station in Hornsund (PPS). CTD data were collected under the oceanographic monitoring
of the PPS (available at https://dataportal.igf.edu.pl/). TerraSAR-X data provided by German Aerospace Center (DLR, project LAN2787).
Landsat-8 data made available by the United States Geological Survey (USGS). https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. the problem remains of how to automatically select t0 and t1 for submarine calving events; this would require long-term
calibration dataset of synchronized high-frequency time-lapse images and noise recordings. Such data are not available for
calving events observed in 2013. Consequently, the analysis of submarine events was out of the scope of this study. the problem remains of how to automatically select t0 and t1 for submarine calving events; this would require long-term
calibration dataset of synchronized high-frequency time-lapse images and noise recordings. Such data are not available for
calving events observed in 2013. Consequently, the analysis of submarine events was out of the scope of this study. Author contributions. J.T. conceived the project, led the 2013 field expedition to Svalbard and participated in data analysis and interpre-
410
tation, and participated in manuscript preparation. O.G. is co-architect of the sound inversion methodology, performed data analysis and
interpretation, and participated in manuscript preparation. M.C. collected and processed the photographic data of terminus ablation. M.B. collected and processed satellite and stick data of glacier velocity and ice mass loss. J.J. provided scientific oversight into data analysis and
participated in manuscript preparation. M.M. participated in the field campaigns and provided CTD data. P.B. participated in field campaigns Author contributions. J.T. conceived the project, led the 2013 field expedition to Svalbard and participated in data analysis and interpre-
410
tation, and participated in manuscript preparation. O.G. is co-architect of the sound inversion methodology, performed data analysis and
interpretation, and participated in manuscript preparation. M.C. collected and processed the photographic data of terminus ablation. M.B. collected and processed satellite and stick data of glacier velocity and ice mass loss. J.J. provided scientific oversight into data analysis and
participated in manuscript preparation. M.M. participated in the field campaigns and provided CTD data. P.B. participated in field campaigns
and participated in manuscript preparation. G.B.D. is co-architect of the sound inversion methodology and detection algorithm, participated
415
in field campaign, and participated in manuscript preparation. collected and processed satellite and stick data of glacier velocity and ice mass loss. J.J. provided scientific oversight into data analysis and
participated in manuscript preparation. M.M. participated in the field campaigns and provided CTD data. P.B. participated in field campaigns
and participated in manuscript preparation. G.B.D. References
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and Park, Y.: Quantifying Soundscapes in the Ross Sea, Antarctica Using Long-Term Autonomous Hydroacoustic Monitoring Systems, Yun, S., Lee, W. S., Dziak, R. P., Roche, L., Matsumoto, H., Lau, T.-K., Sremba, A., Mellinger, D. K., Haxel, J. H., Kang, S.-G., Hong, J. K.,
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c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure 1. (A) Map of the study site and (B) example image of the Hansbreen terminus. Locations of the time-lapse camera and the acoustic
buoy are marked with yellow and white font, respectively. The black dashed lines show different propagation transects that were used for
estimating the noise transmission loss. Landsat 8 satellite data collected on 11 September 2013, courtesy of the US Geological Survey,
Department of the Interior. Bathymetric data provided by (1) the Norwegian Hydrographic Service under the permit no. 13/G722, issued to
the Institute of Geophysics Polish Academy of Sciences, and (2) the Faculty of Natural Sciences, Institute of Earth Sciences, University of
Silesia in Katowice, Sosnowiec, Poland (Błaszczyk et al., 2021). Figure 1. (A) Map of the study site and (B) example image of the Hansbreen terminus. Locations of the time-lapse camera and the acoustic
buoy are marked with yellow and white font, respectively. The black dashed lines show different propagation transects that were used for
estimating the noise transmission loss. Landsat 8 satellite data collected on 11 September 2013, courtesy of the US Geological Survey,
Department of the Interior. Bathymetric data provided by (1) the Norwegian Hydrographic Service under the permit no. 13/G722, issued to
the Institute of Geophysics Polish Academy of Sciences, and (2) the Faculty of Natural Sciences, Institute of Earth Sciences, University of
Silesia in Katowice, Sosnowiec, Poland (Błaszczyk et al., 2021). 19 Figure 2. Calving detector explained. Top: A spectrogram of the sound produced by a calving event within the background noise radiated
by the melting terminus of Hansbreen. The frequencies f0 and f1 on the right hand side of the plot denote the lower and upper limits of
the detection frequencies (see text for details). Bottom: Noise power from the spectrogram above, integrated from f0 to f1 and filtered. The
calving event is detected when the noise power exceeds the power threshold, Pthres, as annotated by t0 and t1. The threshold power is based
th b
li
P
hi h i d t
i
d f
t ti ti
l
l
i
f th
d (
t
t f
d t il )
https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure 2. Calving detector explained. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Top: A spectrogram of the sound produced by a calving event within the background noise radiated
by the melting terminus of Hansbreen. The frequencies f0 and f1 on the right hand side of the plot denote the lower and upper limits of
the detection frequencies (see text for details). Bottom: Noise power from the spectrogram above, integrated from f0 to f1 and filtered. The
calving event is detected when the noise power exceeds the power threshold, Pthres, as annotated by t0 and t1. The threshold power is based
on the baseline power, Pbase, which is determined from a statistical analysis of the sound (see text for details). 20 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure 3. A comparison of ice volume loss derived from iceberg impact noise and terminus area loss from camera observations. The ice
volume loss was calculated every 3 hours, which is the sampling time of the camera system. The thick lines show the two data streams
smoothed with 1-day running average. Figure 3. A comparison of ice volume loss derived from iceberg impact noise and terminus area loss from camera observations. The ice
volume loss was calculated every 3 hours, which is the sampling time of the camera system. The thick lines show the two data streams
smoothed with 1-day running average. 21 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure 4. The effect of sampling coverage on acoustic estimates of (A) event count and (B) ice volume loss. Recording periods range from
1 hour per day to continuous. The reference value of ∆= 0% is for full coverage. Figure 4. The effect of sampling coverage on acoustic estimates of (A) event count and (B) ice volume loss. Recording periods range from
1 hour per day to continuous. The reference value of ∆= 0% is for full coverage. 22 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure 5. (blue) Percentage of time occupied by calving noise over the study period and (orange) the acoustic estimate of daily calving rate. Figure 5. (blue) Percentage of time occupied by calving noise over the study period and (orange) the acoustic estimate of daily calving rate. Figure 5. (blue) Percentage of time occupied by calving noise over the study period and (orange) the acoustic estimate of daily calving rate. Figure 5. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. (blue) Percentage of time occupied by calving noise over the study period and (orange) the acoustic estimate of daily calving rate. 23 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure A1. Data used in the subaerial calving flux calculations. Horizontal velocity map from offset tracking on repeat TerraSAR satellite
radar images (2012-12-15 and 2012-12-26), retreat of Hansbreen cliff and localization of stake position with dGPS measurements between
August 5 and August 24, 2013. The grey profile with dots shows division of the ice cliff into 40 m lengths, along which precise calculation
of subaerial calving flux were made. Figure A1. Data used in the subaerial calving flux calculations. Horizontal velocity map from offset tracking on repeat TerraSAR satellite
radar images (2012-12-15 and 2012-12-26), retreat of Hansbreen cliff and localization of stake position with dGPS measurements between
August 5 and August 24, 2013. The grey profile with dots shows division of the ice cliff into 40 m lengths, along which precise calculation
of subaerial calving flux were made. 24 24 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure A2. (A) Histograms and (B) empirical cumulative distribution functions (ECDFs) of log-transformed impact noise energies calculated
for calibration data (2016; orange) and acoustic measurements used in this study (2013; blue). Figure A2. (A) Histograms and (B) empirical cumulative distribution functions (ECDFs) of log-transformed impact noise energies calculated
for calibration data (2016; orange) and acoustic measurements used in this study (2013; blue). 25 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure A3. Relationship between detection factor β and the number of detected calving events. Interfering events are not removed in this
analysis. Figure A3. Relationship between detection factor β and the number of detected calving events. Interfering events are not removed
analysis. Figure A3. Relationship between detection factor β and the number of detected calving events. Interfering events are not removed in this
analysis. Figure A3. Relationship between detection factor β and the number of detected calving events. Interfering events are not removed in this
analysis. 26 https://doi.org/10.5194/egusphere-2023-115
Preprint. Discussion started: 9 February 2023
c⃝Author(s) 2023. CC BY 4.0 License. Figure A4. Change in number of events detected (N), total acoustic energy at the receiver (E) and summed event durations (D) versus
iteration number for the background noise estimate. Figure A4. Change in number of events detected (N), total acoustic energy at the receiver (E) and summed event durations (D) versus
iteration number for the background noise estimate. Figure A4. Change in number of events detected (N), total acoustic energy at the receiver (E) and summed event durations (D) versus
iteration number for the background noise estimate. 27 27
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2. Unsere Studie: Design und Methoden
Die diesem Buch zu Grunde liegende empirische Untersuchung hatte ein
spezifisch aufgefächertes qualitatives Design (siehe Abbildung 1 ), wobei Expert:inneninterviews und Organisationsfallstudien die wesentlichen Stützpfeiler waren. Zu Beginn stand die theoretische Auseinandersetzung mit den
Rahmenbedingungen, unter denen organisierte Interaktionsarbeit in den von
uns betrachteten Branchen gegenwärtig (auch schon vor Corona) stattfindet.
Überprüft und ergänzt wurde diese durch 15 Expert:innengespräche (teils
mit zwei Personen) (siehe Kap. 3.1und 3.5) (vgl. Gläser/Laudel 2010; Bogner/
Littig/Menz 2009, 2014; Kaiser 2014). Diese wurden 2020 mit 21 Branchenkenner:innen aus Verbänden und Gewerkschaften geführt. Themen waren u.a. die
Sichtweisen auf Dynamiken interner Steuerung (Management), ferner wahrgenommene Einflüsse externer Steuerung (Politik, Markt) und schließlich
in der Branche beobachtete Unsicherheiten sowie der emotionale Umgang
damit. Für die Sektoren Weiterbildung und JBH wurde zusätzlich eine Fokusgruppendiskussion mit Expert:innen geführt, da diese Branche unter den für
das Forschungsprojekt relevanten Kontextgesichtspunkten wenig beforscht
ist.
Ein wesentlicher Baustein des Projekts waren Organisationsfallstudien1
in vier ausgewählten gemeinnützigen Einrichtungen in deutschen Großstädten (mit variierenden Profilen und einer gewissen regionalen Varianz von
Nord-, Süd- und Ostdeutschland), die schon seit längerem in diesem Segment
tätig sind (weitere Charakteristiken siehe Kap. 3.2) und überwiegend in 2021
erhoben wurden. Unser Sample bestand jeweils aus einer Organisation in den
Sub-Sektoren, also: je eine Organisation im Bereich der Jugendberufshilfe,
1
Sie orientiert sich an geläufigen Ansätzen der ›Case-Study‹-Forschung und fokussiert
komplexe Strukturzusammenhänge und Prozessverläufe, wobei Kontextbezug, Offenheit und Multiperspektivität wesentliche Gütemaßstäbe darstellen (vgl. Pflüger/
Pongratz/Trinczek 2017).
26
Organisierte Zerrissenheit
der Weiterbildung für Ältere, der stationären und der ambulanten Pflege.
Jede Organisationsfallstudie basierte auf Einzelinterviews (in wenigen Fällen
auch Interviews mit zwei Personen gleichzeitig), einer Gruppendiskussion
sowie Dokumentenanalysen. Pro Einrichtung wurden sechs bis elf ein- bis
zweieinhalbstündige problemzentrierte Interviews mit narrativen Anteilen
durchgeführt (vgl. Witzel 1985; zu dieser Kombination Scheibelhofer 2008),
dabei ging es in einem ersten Teil um das Erleben des beruflichen Alltags
und eventueller Herausforderungen (in Bezug auf Rahmenbedingungen, Management, Organisation und Nutzer:innen), in dem zweiten Teil wurde dann
stärker auf emotionale Belastungen fokussiert. Insgesamt haben wir in dieser
Weise 36 Personen interviewt. Befragt wurden Beschäftigte unterschiedlicher
Qualifikationsstufen und Funktionsbereiche, Leitungskräfte (auf der unteren,
mittleren und höchsten Managementebene) und Personalvertreter:innen.2
Hinzu kam für jede Einrichtung eine etwa zweistündige Gruppendiskussion (vgl. Bohnsack/Przyborski/Schäffer 2010; Lamnek 2006; Kühn/Koschel
2018) mit vier bis acht Betriebsangehörigen, in deren Mittelpunkt das Thema
Solidarität auf den Ebenen Betrieb (inklusive der Erfahrungen mit Nutzer:innen), Branche und Gesellschaft bzw. Wohlfahrtsstaat stand. Mit Ausnahme
der organisationsinternen Gruppendiskussion in der Jugendberufshilfe wurde in den anderen Fallstudien mit diesem Instrument vor allem das mittlere
Management erreicht. Durch unseren gewählten Feldzugang über die Geschäftsleitung der jeweiligen Einrichtungen haben diese auch die Befragten
für die Einzelgespräche und die Gruppendiskussion ausgewählt, unter Beachtung unserer Maßgabe, dass wir verschiedene Funktionsbereiche befragen
wollten. Dabei waren die Organisationen auch darauf angewiesen, dass Beschäftigte sich dafür bereit erklärten. Die Gesamtzahl der einzeln oder in
2
So wurden im Pflegesektor Pflegehelferinnen, examinierte Pflegefachkräfte, Betreuungskräfte, Hauswirtschafts- und Reinigungspersonal befragt. Befragte in Leitungsfunktionen waren Teamleitungen, Pflegedienstleitungen, Heimleitungen und oberste Geschäftsführungen. Im Weiterbildungssektor befragten wir gewerbliche Ausbilder:innen in Berufsausbildungsmaßnahmen, Job-Coaches und akademisch qualifizierte Sozialpädagog:innen. Befragte in Leitungsfunktionen waren hier Bereichsleitungen (z.B. für Jugendliche oder Erwachsene), Standortleitungen und Geschäftsführungen von Zweigstellen. Der Frauenanteil im gesamten Sample der Fallstudien liegt
bei 75 %. Im Sample waren alle Altersgruppen vertreten. Bei der Darstellung der Befunde wird, wo es relevant erschien, auf deren Zugehörigkeit jeweils hingewiesen. Vertreten waren ebenfalls Personen mit Migrationsgeschichte.
2. Unsere Studie: Design und Methoden
Gruppendiskussionen Befragten aller vier Fallstudien beträgt insgesamt 56
Personen.
Zu den Organisationsfallstudien gehörte auch die Sammlung und Sichtung schriftlicher Artefakte, die einer Dokumentenanalyse (vgl. Wolff 2008)
unterzogen wurden. Dabei haben wir die Organisationen um interne Dokumente gebeten – etwa Geschäftsberichte oder Tätigkeits- und Qualitäts(management)berichte – und öffentlich zugängliche Daten (z.B. Internetauftritt,
Zeitungen, Flyer) gesammelt.
Abbildung 1: Methodisches Design
Eigene Darstellung
27
28
Organisierte Zerrissenheit
Das mannigfaltige gewonnene Material aus den Organisationsfallstudien wurde im Rekurs auf die qualitative Inhaltsanalyse (vgl. Mayring 2015)
ausgewertet. Dies diente dazu, Informationen zu ordnen bzw. zu sichern
und das Gesamtmaterial besser überblicken zu können. Manifeste Aussagen
wurden paraphrasiert und mithilfe eines Softwareprogramms kodiert und
inhaltsanalytisch ausgewertet, was schließlich in Interviewportraits für jedes Einzelinterview mündete, in denen die von den Befragten dargestellten
Themen zusammengefasst sowie besonders prägnante Zitatstellen eingefügt
wurden (siehe Befunde in Kap. 3.2 und 3.3). Durch diese intensive Auseinandersetzung mit dem Material war es möglich, relevante Schlüsselstellen
vor allem in den Einzelinterviews zu identifizieren, welche dann rekonstruktiv/interpretierend analysiert wurden, um sich so einen besseren Zugang
zum latenten emotionalen Gehalt zu erschließen und Zusammenhänge zu
Solidaritätskonstrukten im oben genannten Verständnis herausarbeiten zu
können (siehe exemplarische Feinanalysen in Kap. 3.4). Ging es in der Inhaltsanalyse noch darum, was genau gesagt wurde, untersuchten wir angelehnt
an rekonstruktive Verfahren für einzelne Passagen sequenzanalytisch, was
die sprachlichen Ausgestaltungen über latente Sinnkonstruktionen verraten.
Dabei orientierten wir uns an den Grundprinzipien rekonstruktiv-hermeneutischer Analyse, welche im integrativen Basisverfahren nach Kruse (2015)
beschrieben sind; Stellen, die besonders aufschlussreich für das emotionale
Erleben wirkten, wurden also einer mikroskopischen Feinanalyse unterzogen
(vgl. ebd., S. 475ff.; auch Kleres 2011, 2015 in Bezug auf Emotionen).
Nach diesen Tiefenanalysen wird der Blick in die Breite der untersuchten
Branchen auf Basis der Erkenntnisse aus den Expert:innengesprächen gerichtet (siehe Kap. 3.5). Dies dient der Einordnung und Plausibilisierung der Befunde aus den Organisationsfallstudien.
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Ein Mentoring-Programm mit Coaching-Anteilen für die Ausbildung von Lehrpersonen
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Niggli, Alois
Ein Mentoring-Programm mit Coaching-Anteilen für die Ausbildung von
Lehrpersonen
Beiträge zur Lehrerbildung 19 (2001) 2, S. 244-250
Quellenangabe/ Reference:
Niggli, Alois: Ein Mentoring-Programm mit Coaching-Anteilen für die Ausbildung von Lehrpersonen In: Beiträge zur Lehrerbildung 19 (2001) 2, S. 244-250 - URN: urn:nbn:de:0111-pedocs-134580 - DOI:
10.25656/01:13458
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in Kooperation mit / in cooperation with:
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244
Beiträge zur Lehrerbildung, 19 (2), 2001
Ein Mentoring-Programm mit Coaching-Anteilen für
die Ausbildung von Lehrpersonen
Alois Niggli
In einer professionsorientierten Ausbildung für künftige Lehrkräfte ist der subjektive Austausch von Wissen grundlegend, weil gesicherte Handlungsanleitungen
nicht vermittelt werden können. Mentorpersonen fällt deshalb die Aufgabe zu, entsprechende Förderungsbeziehungen aufzubauen. Andererseits gelten sie auch als
Teil des Systems, das mittels Mentoring-Programmen seine Interessen implementieren will. Diese beiden präskriptiven Ausrichtungen konfligieren nicht selten und
lassen sich wohl nie optimal gegeneinander ausbalancieren. Mit dem folgenden
Modell wird der Anspruch vertreten, zum Umgang mit dieser Spannung einen
Beitrag zu leisten. Vorgeschlagen werden drei strategische Bearbeitungsbereiche.
Die Entwicklung einer beruflichen Handlungskompetenz ist ein schwieriges Unterfangen, wenn man jeweils nie genau wissen kann, ~b man nun richtig oder falsch gehandelt hat. Alle, die mit Kindern und Jugendlichen umgehen, wissen, dass Unterrichts- und Erziehungsarbeit jedoch nie sicheren Erfolg zeitigt. Jede Lehre realisiert
sich in Form einer Beziehungspraxis. Sachliche Ziele lassen sich infolgedessen immer
nur tendenziell erreichen (vgl. Combe, 1997, S. 10.). Studierende in Lehrerbildungsgängen haben nicht selten Schwierigkeiten, diese komplexe Situation zu akzeptieren. Legitimerweise verlangen sie von ihrer Ausbildung Gewissheit. Sie möchten
die Aufgaben, vor die sie im institutionellen Kontext von Schule gestellt werden, erfolgreich bewältigen können. Wenn berufliches Wissen jedoch nie sicheren Erfolg
verbürgt, dann kann professionelles Handeln nicht durch deduktive Anwendung von
erworbenem Fachwissen der wissenschaftlichen Disziplin gekennzeichnet sein. Die
Lehrerbildung ist grundsätzlich nicht in der Lage, die eingeforderte Sicherheit zu
vermitteln. Aus diesem Grunde sollte Lernen wenn immer möglich reflexiv und situiert arrangiert werden. Interne Regulationen müssen bedeutsam werden. Mentoringl
ist eine Möglichkeit, diesem Anspruch nahe zu kommen. In untenstehender
Abbildung (vgl. Abb. 1) wird ein sogenanntes 3-Wege-Modell zum Mentoring vorgestellt, das den genannten Grundvoraussetzungen Rechnung trägt. Im theoretischen
Kontext einer- Ausbildung zur Profession wird auf Seiten der Lernenden eine
wissensorientierte Kompetenz- und eine Steuerungsebene der Person unterschieden.
Auf beiden Ebenen begleiten Mentorinnen und Mentoren Lernende in ihrer
Entwicklung. Diese Begleitung erfolgt im Rahmen einer förderlichen Beziehung.
Auf der Kompetenzebene entsteht eine professionelle Entwicklung aus der Auseinandersetzung mit dem erwähnten Nicht-Perfektsein, wenn dieses nicht nur wahrgenommen, sondern im Rahmen der in der Ausbildung vertretenen Kompentenzprofile
auch als bearbeitbar erlebt wird (s. Bauer, 1998, S. 355). Mentorinnen und Mentoren
unterstützen diese Bearbeitbarkeit. Dabei müssen sie auch das Kompetenzpmfil der
' Mentor (oder Mentes) ist in der griechischen Mythologie der Freund des Odysseus. Er kümmert sich
während der Irrfahrten des Königs um seinen Sohn Telemach, und wird zur emotionalen und intellektuellen Leitfigur des Jünglings. In dieser Eigenschaft wird er ein Vorbild für Erzieher und zum 'Vater'
des modernen 'Mentoring' in Wirtschaft und Wissenschaft (Strasser & Schliesselberger, 2000, S. 13).
Begleiten und beraten in der berufspraktischen Ausbildung http://www.bzl-online.ch
245
Organisation glaubwürdig verkörpern. Sie können nicht einfach unterstützen, was
Lernende für richtig halten. Im Modell werden zwei Wissensbereiche unterschieden,
die gemeinsam bearbeitet werden können: der Tätigkeitsbereich ('knowing how'), in
welchem Können manifest wird und das Erklärungswissen ('knowing what') (vgl.
auch Heitzmann & Messner, 2001, S. 14). Den Entwicklungsaufgaben in diesen zwei
Wissensbereichen werden im Rahmen eines Mentoring spezifische Förderungsmodi
(als Gesprächstypen) zugeordnet, nämlich (a) das Feedback erweiternde Praxisgespräch und (b) das reflexive Praxisgespräch. Die beiden Wissensformen bedingen
jeweils eine andere Behandlung, weil Erklärungswissen erst in Können transformiert
werden muss.
Wahrnehmung des Unterrichtsgeschehens
(Spontanfeedback)
Weg 1
I
erweiterndes
Praxisgespräch
I
Weg 2
kraxisgespracn
Kompetenzebene der Unterrichtssituation
)
I(
I Personale Steuerungsebene
Abbildung 1: Modell des 3-Wege Mentoring (3WM)
Diese Entwicklung hat andererseits aber auch über den Aufbau eines beruflichen
Selbst2 zu erfolgen. Unter dem beruflichen Selbst wird in Anlehnung an Bauer
(1998, S. 344) ein Bewusstsein verstanden, das die persönlichen Entwicklungsaufgaben steuert, die jedem künftigen Pädagogen gestellt sind. Auf dieser Ebene setzen
Lernende ihre Ziele selbst. Sie sind Gegenstand der Selbstwahrnehmung, denn durch
Dieser Begriff mag zwar wenig präzise sein. Allerdings eröffnet er Anschluss an Entwicklungsaufgaben, welche die Person in ihrem So-Sein betreffen und nicht lediglich einzelne berufliche Kompetenzen kennzeichnen.
246
Beiträge zur Lehrerbildung, 19 (2), 2001
die erlebten Unsicherheiten werden sie in ihrem bemflichen Selbstverständnis ständig
herausgefordert. Lernende müssen sich neue Ziele setzen, ihr Wissen erneuern, generell ihre Entwicklung zur Profession steuern. Im vorliegenden Mentoringkonzept wird
diese Selbststeuerung durch Coaching begleitet. Durch diese Unterstützung soll das
individuelle Potential bewusst und zugänglich gemacht werden, damit derldie
Coachee aktiv und möglichst erfolgreich die ausbildungsbezogenen Herausfordemngen bewältigen kann. Coaching wird im vorliegenden Modell auf diese eher non-direktive Beziehung eingeschränkt.
Eine Mentorin oder ein Mentor muss sich darüber im Klaren sein, auf welchem
Weg Lernende jeweils begleitet werden sollen.
Weg I : Entwicklung im Tätigkeitsbereich
Aufgmnd der unsicheren Resultate erziehenschen Handelns sind angehende Lehrkräfte in ihrer Ausbildung auf Hinweise über die Wirkung ihrer unterrichtlichen Tätigkeit in hohem Masse angewiesen. Diese Informationen sind für ihre Selbsteinschätzung und vor allem für ihre Selbstwirksamkeit (Hertramph & Henmann, 1999)
von nicht zu unterschätzender Bedeutung. Der Lehrer(innen)bemf zeichnet sich generell durch ein hohes Mass an Selbstdefinition bei gleichzeitig institutionell bedingter
'Rückmeldearmut' aus (Schaefers & Koch, 2000, S. 614). Referenzpunkt auf Weg 1
ist deshalb die konkret wahrnehmbare Untemchtstätigkeit. Sie wird als 'knowinghow' (Ryle, 1969) gesehen. Wegleitend sind praxiswirksame (im weiteren Sinne
technische) Regeln, die auf intensiven Erfahmngen beruhen. Dieses Handlungswissen ist für eine aussenstehende Beobachter-Mentorperson nie in seiner Gesamtheit,
sondern lediglich an der Oberfläche als 'Skill' wahrnehmbar. Dabei handelt es sich
um Basishandlungen, die sich im Kontext eines Ganzen ereignen, von dem sie selber
Teil sind. Je nach Kontextbezug kann man eher geschlossene oder offenere Skills
unterscheiden. Geschlossene Skills haben wenig Kontext. Sie sind algorithmisch beschreib- und lernbar (z.B.: 'Einen verständlichen Kurzvortrag halten'). Offene, komplexe Skills haben mehr Kontext (z. B.: Kooperativen Unterricht adäquat einsetzen).
Sie beinhalten unterschiedliche Absichten, die unterschiedliche Funktionen erfüllen
können (s. Tomlinson, 1998, S. 15).
Die Behandlung von 'Skill-Ausschnitten' der beobachtbaren Untemchtstätigkeit
kann in der Ausbildungspraxis über konkretes Feedback erfolgen. Von Feedbackerweiterung kann deshalb gesprochen werden, weil Lehrpersonen über ihre Selbstwahmehmung des Untemchts immer auch spontanem Feedback ausgesetzt sind. Die
Rückmeldung der Mentorperson orientiert sich darüber hinaus jedoch an vereinbarten
praxiswirksamen Standards. Vereinbart sind Standards deshalb, weil Wirksamkeit
nicht 'objektiv' begründbar ist. Sie unterscheiden sich in ihrer Konkretheit, je nachdem ob offene oder eher geschlossene Skills beobachtet werden sollen. Mit ihrem
Feedback sollten Mentorpersonen zu allererst versuchen, das Können der Lernenden
herauszuschälen. Übereinstimmende Wahrnehmungen können dann zu mehr Gewissheit über erfolgreiche oder gegebenenfalls über zu verändernde Strategien verhelfen.
Diskrepanzen bedürfen der anschliessenden Klämng. Diese kann in einem reflexiven
Praxisgespräch herbeigeführt werden.
Begleiten und beraten in der berufspraktischen Ausbildung
Die Befolgung bestimmter Regeln im Rahmen des 'knowing how' kann darüber
hinaus manchmal intuitiv geschehen. Das obgenannte Untemchtsverhalten kann dann
die Form eines impliziten 'tacit knowledge' (Neuweg, 1999) haben, dessen Hebung
ins Bewusstsein (s. Weg 2) als 'knowing what' besonderer Anstrengung bedarf. Dies
ist eine Voraussetzung, dass Verändemngen überhaupt erst in Angriff genommen
werden.
Weg 2: Entwicklung des Erklärungswissens
'
Referenzpunkt auf Weg 2 ist das Wissen, das mehr oder weniger losgelöst von der
abgelaufenen Tätigkeit existiert. Geht es um dieses Hintergmndwissen, dann muss
ein Gespräch anders stmktunert sein, als wenn man sich über die Wahrnehmung
konkreter Tätigkeiten einigt. Nach der Terminologie von Ryle (1969) handelt es sich
bei diesem Hintergmndwissen um 'knowing what' (Erklämngswissen über die Praxis). Dieses Erklämngswissen kann aus unterschiedlichen Quellen stammen (vgl.
Shulman, 1986, 1987). Es umfasst u.a. sowohl nomologisches Wissen der wissenschaftlichen Disziplinen wie auch das professionelle Wissen kompetenter Lehrpersonen.
Das zugeordnete Reflexive Praxisgespräch zielt auf verständigung über das Erklärungswissen. Reflexion soll zu einer Ausdifferenziemng des Hintergmndwissens
beitragen. Es hat sich als praktikabel erwiesen, entsprechende Gespräche wie folgt zu
stmkturieren (vgi. Abb. 2).
1. Beschreibung
I
Beschreibuna
der
.- .
I Informieren
- - - -
2. Analyse
1 Konfrontieren
3. Synthese
1
4. Optionen
1 Rekonstruieren I Schlussfol~erun-
klären
Ereignisse auf die
man sich bezieht
festlegen
I
Mentoriin und
Mentee
1
I
Mentee
Mentoriin
I
Mentoriin und
Mentee
i. d. R. Mentee
-
Abbildung 2: Phasen im Reflexiven Praxisgespräch
Vorerst wird die wahrgenommene Realität von beiden Partnern deskriptiv dargestellt.
Die Mentoridder Mentor gibt anschliessend Anstösse zur Klärung von Hintergmndwissen (z.B. 'Welche Vorannahmen haben Sie zum Vorwissen der Lernenden getroffen?). Die Realität wird dabei unter dem Anspmch untersucht, selbst neues Erklämngswissen zu generieren. Inspirationen der Mentorperson beim Konfrontieren können Offerten sein, die von den Handelnden angenommen oder auch verworfen werden können. Die Geltung entsprechender Erklärungsansätze ist allein in der Intersubjektivität der Verständigung begründet. In der Synthesephase wird das Hintergrundwissen allenfalls neu stmkturiert (z. B. 'Gibt es Dinge, die Sie anders sehen als
zu Beginn des Gesprächs?'). Erst dann werden mögliche neue Zielsetzungen vereinbart. Dies ist deshalb notwendig, weil Reflexion allein für die bemfliche Entwicklung
nicht hinreichend ist. Praxiswirksam wird das erzeugte Wissen erst im Kontext der
Beiträge zur Lehrerbildung, 19 (Z), 2001
248
konkreten Unterrichtstätigkeit. Darüber sollen die Lernenden jedoch Feedback erhalten und nicht nur reflektieren müssen.
Weg 3: Entwicklung des Professionellen Selbst
Bei den oben beschriebenen Wissenskomplexen handelt es sich um funktionale Anforderungen, die notwendig sind, um einen Beruf kompetent auszuführen. Daneben
ist eine Lehrkraft aber quasi immer auch auf einer persönlichen Steuerungsebene, in
ihrem beruflichen Selbst herausgefordert (vgl. Bauer, 1998). Es geht dabei nicht primär um Qualifizierung, sondern um Subjektwerdung. Konkret bedeutet dies U. a.:
- sich als Veränderungsprojekt zu verstehen und entsprechend zu handeln,
- eigenes Lernen reflektiert zu begleiten,
- Lernanlässe selbst festzustellen, zu steuern und in Gang zu setzen.
Diese Selbstbeobachtung hat überprüfende Funktion. Sie sollte nicht dazu führen,
sich auf Unzulänglichkeiten zu fixieren. Ansätze, die sich unter dem Sammelbegriff
'Coaching' etabliert haben, können diesen Prozess begleiten und unterstützen. Das
zugeordnete Coaching-Gespräch fördert somit das Verständnis 'über mich selbst'.
Dieser Erkenntnisprozess wird in zwei Schleifen strukturiert (vgl. Abb. 3).
Tatsachen
Situations-
Hintergründe
Ausgangslagel
Problem
Ziele, Prinzipien
Alternativen
notwendigen Bewusstheit
und Erkenntnis für
seinelihre Situation einen
"guten" Entscheid treffen
Diagnose
Entscheidung
Begleiten und beraten in d e r berufspraktischen Ausbildung
249
sen zu strukturieren. Sein wichtigstes Arbeitsinstrument findet der Coach in der Fragetechnik der offenen Frage.
Können Lehrerbildnerinnen und Lehrerbildner die berufliche Entwicklung Studierender auf den vorgeschlagenen drei Wegen jedoch überhaupt fördern? Diese Frage
zielt in letzter Konsequenz auf die Effizienz der im Modell vertretenen Massnahmen.
In diesem Problemkontext hat die theoretische Arbeit von Vygotsky in den letzten
Jahren zunehmende Beachtung gefunden (s. Reiman, 1999). Der Prozess der kognitiven Konstruktion vollzieht sich nach Vygotskys Annahmen nicht von selbst. Notwendig sind soziale Interaktionen. Entwicklung resultiert damit aus ständigen Anforderungen der Gesellschaft an das Individuum. Aus diesen Grundannahmen leitet sich
ab, dass Mentorpersonen eine aktive Rolle zu spielen haben. Kompetente ExpertenPersonen können die Entwicklung fördern, indem sie angemessene Anforderungen
stellen. Mit dem vorgeschlagenen 3-Wege-Modell lassen sich diese Anforderungen
bereichsspezifisch differenzieren und moderieren. Die Herausforderung für Ausbildnerinnen und Ausbildner wird deshalb darin zu sehen sein, auf den drei vorgeschlagenen Wegen Unterstützung bei der Bewältigung kognitiver Verunsicherung einerseits.zu gewähren, andererseits kognitives Ungleichgewicht aber auch auszulösen.
Das angemessene Pendeln zwischen diesen Polen könnte letzten Endes eine Kunst
sein, die hohe diagnostische Sensibilität voraussetzt. Es ist eingebettet in ein Coaching, das ein Bemühen unterstützt, das der Selbstentwicklung Gestalt geben soll.
Feedback annehmen, reflektieren, sich beruflich entwickeln zu müssen stellt für Lernende immer auch eine Zumutung dar, nämlich die Zumutung, 'die Welt, meine Tatigkeiten und mich selbst anders zu sehen und zu deuten als ich es gerne möchte'
(Reichenbach, 2000, S. 803). Mentorinnen und Mentoren begleiten die Lernenden
dabei als kritische Freunde.
Literatur
Konsequenzen
Abbildung 3: Erkenntnisprozessim Coaching (Furter, 2000)
Der Erkenntnisprozess geht von einer aktuellen Frage-/Problemstellung in der Gegenwart aus und verläuft in einer 'Schleife' in die Vergangenheit, um die gewordene
Situation und deren Ursachen zu verstehen. Der Weg kehrt zum/zur Coachee als Entscheiderlin zurück: die Konklusion ist nun in seinendihrem Bewusstsein und für
ihnlsie relevant. Der Prozess der Entscheidungsfindung nimmt seinen Ausgang bei
Zielen und Prinzipien, die derldie Coachee festlegt und macht jetzt eine 'Schleife' in
die Zukunft: wie könnten die künftige(n) Situation(en) aussehen, und welches wären
die Konsequenzen? Der Weg kehrt dann wiederum zundzur Coachee zurück, derldie
schliesslich eine Entscheidung trifft. Die Aufgabe des Coachs besteht nun darin,
denldie Coachee durch diese Schlaufen zu begleiten und dabei die verschiedenen Pha-
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