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Fuel consumption and emissions reductions
The hybrid vehicle typically achieves greater fuel economy and lower emissions than conventional internal combustion engine vehicles (ICEVs), resulting in fewer emissions being generated. These savings are primarily achieved by three elements of a typical hybrid design:
Relying on both the engine and the electric motors for peak power needs, resulting in a smaller engine size more for average usage rather than peak power usage. A smaller engine can have fewer internal losses and lower weight.
Having significant battery storage capacity to store and reuse recaptured energy, especially in stop-and-go traffic typical of the city driving cycle.
Recapturing significant amounts of energy during braking that are normally wasted as heat. This regenerative braking reduces vehicle speed by converting some of its kinetic energy into electricity, depending upon the power rating of the motor/generator;
Other techniques that are not necessarily 'hybrid' features, but that are frequently found on hybrid vehicles include:
Using Atkinson cycle engines instead of Otto cycle engines for improved fuel economy.
Shutting down the engine during traffic stops or while coasting or during other idle periods.
Improving aerodynamics; (part of the reason that SUVs get such bad fuel economy is the drag on the car. A box-shaped car or truck has to exert more force to move through the air causing more stress on the engine making it work harder). Improving the shape and aerodynamics of a car is a good way to help better the fuel economy and also improve vehicle handling at the same time.
Using low rolling resistance tires (tires were often made to give a quiet, smooth ride, high grip, etc., but efficiency was a lower priority). Tires cause mechanical drag, once again making the engine work harder, consuming more fuel. Hybrid cars may use special tires that are more inflated than regular tires and stiffer or by choice of carcass structure and rubber compound have lower rolling resistance while retaining acceptable grip, and so improving fuel economy whatever the power source.
Powering the a/c, power steering, and other auxiliary pumps electrically as and when needed; this reduces mechanical losses when compared with driving them continuously with traditional engine belts.
These features make a hybrid vehicle particularly efficient for city traffic where there are frequent stops, coasting, and idling periods. In addition noise emissions are reduced, particularly at idling and low operating speeds, in comparison to conventional engine vehicles. For continuous high-speed highway use, these features are much less useful in reducing emissions. | Hybrid vehicle | Wikipedia | 495 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
Hybrid vehicle emissions
Hybrid vehicle emissions today are getting close to or even lower than the recommended level set by the EPA (Environmental Protection Agency). The recommended levels they suggest for a typical passenger vehicle should be equated to 5.5 metric tons of . The three most popular hybrid vehicles, Honda Civic, Honda Insight and Toyota Prius, set the standards even higher by producing 4.1, 3.5, and 3.5 tons showing a major improvement in carbon dioxide emissions. Hybrid vehicles can reduce air emissions of smog-forming pollutants by up to 90% and cut carbon dioxide emissions in half.
An increase in fossil fuels is needed to build hybrid vehicles versus conventional cars. This increase is more than offset by reduced emissions when running the vehicle.
Hybrid emissions have been understated when comparing certification cycles to real-world driving. In one study using real-world driving data, it was shown they use on average 120 g of per km instead of the 44 g per km in official tests.
Toyota states that three Hybrid vehicles equal one battery electric vehicle in reduction effect from carbon neutrality viewpoint which means reducing emissions to zero throughout the entire life cycle of a product, starting from procurement of raw materials, manufacturing, and transportation to use, recycling, and disposal.
Environmental impact of hybrid car battery
Though hybrid cars consume less fuel than conventional cars, there is still an issue regarding the environmental damage of the hybrid car battery. Today, most hybrid car batteries are Lithium-ion, which has higher energy density than nickel–metal hydride batteries and is more environmentally friendly than lead-based batteries which constitute the bulk of petrol car starter batteries today.
There are many types of batteries. Some are far more toxic than others. Lithium-ion is the least toxic of the batteries mentioned above. | Hybrid vehicle | Wikipedia | 359 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
The toxicity levels and environmental impact of nickel metal hydride batteries—the type previously used in hybrids—are much lower than batteries like lead acid or nickel cadmium according to one source. Another source claims nickel metal hydride batteries are much more toxic than lead batteries, also that recycling them and disposing of them safely is difficult. In general various soluble and insoluble nickel compounds, such as nickel chloride and nickel oxide, have known carcinogenic effects in chick embryos and rats. The main nickel compound in NiMH batteries is nickel oxyhydroxide (NiOOH), which is used as the positive electrode. However Nickel Metal Hydride Batteries have fallen out of favour in hybrid vehicles as various lithium-ion chemistries have become more mature to market.
The lithium-ion battery has become a market leader in this segment due to its high energy density, stability, and cost when compared to other technologies. A market leader in this area is Panasonic with their partnership with Tesla
The lithium-ion batteries are appealing because they have the highest energy density of any rechargeable batteries and can produce a voltage more than three times that of nickel–metal hydride battery cell while simultaneously storing large quantities of electricity as well. The batteries also produce higher output (boosting vehicle power), higher efficiency (avoiding wasteful use of electricity), and provides excellent durability, compared with the life of the battery being roughly equivalent to the life of the vehicle. Additionally, the use of lithium-ion batteries reduces the overall weight of the vehicle and also achieves improved fuel economy of 30% better than petro-powered vehicles with a consequent reduction in CO2 emissions helping to prevent global warming.
Lithium-ion batteries are also safer to recycle, with Volkswagen Group pioneering processes to recycle lithium-ion batteries; this is also being chased by various other large companies, such as BMW, Audi, Mercedes-Benz and Tesla. The main goal within many of these companies is to combat disinformation about the nature of lithium batteries, primarily that they are not recyclable, which primarily stem from articles discussing the difficulties of recycling. | Hybrid vehicle | Wikipedia | 445 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
Charging
There are two different levels of charging in plug-in hybrids. Level one charging is the slower method as it uses a 120 V/15 A single-phase grounded outlet. Level two is a faster method; existing Level 2 equipment offers charging from 208 V or 240 V (at up to 80 A, 19.2 kW). It may require dedicated equipment and a connection installation for home or public units. The optimum charging window for lithium-ion batteries is 3–4.2 V. Recharging with a 120-volt household outlet takes several hours, a 240-volt charger takes 1–4 hours, and a quick charge takes approximately 30 minutes to achieve 80% charge. Three important factors—distance on charge, cost of charging, and time to charge
In order for hybrids to run on electrical power, the car must perform the action of braking in order to generate some electricity. The electricity then gets discharged most effectively when the car accelerates or climbs up an incline.
In 2014, hybrid electric car batteries can run on solely electricity for 70–130 miles (110–210 km) on a single charge. Hybrid battery capacity currently ranges from 4.4 kWh to 85 kWh on a fully electric car. On a hybrid car, the battery packs currently range from 0.6 kWh to 2.4 kWh representing a large difference in use of electricity in hybrid cars.
Raw materials increasing costs
There is an impending increase in the costs of many rare materials used in the manufacture of hybrid cars. For example, the rare-earth element dysprosium is required to fabricate many of the advanced electric motors and battery systems in hybrid propulsion systems. Neodymium is another rare earth metal which is a crucial ingredient in high-strength magnets that are found in permanent magnet electric motors.
Nearly all the rare-earth elements in the world come from China, and many analysts believe that an overall increase in Chinese electronics manufacturing will consume this entire supply by 2012. In addition, export quotas on Chinese rare-earth elements have resulted in an unknown amount of supply.
A few non-Chinese sources such as the advanced Hoidas Lake project in northern Canada as well as Mount Weld in Australia are currently under development; however, the barriers to entry are high and require years to go online. | Hybrid vehicle | Wikipedia | 470 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
How hybrid-electric vehicles work
Hybrid-electric vehicles (HEVs) combine the advantage of gasoline engines and electric motors. The key areas for efficiency or performance gains are regenerative braking, dual power sources, and less idling.
Regenerative braking. The electric motor normally converts electricity into physical motion. Used in reverse as a generator, it can also convert physical motion into electricity. This both slows the car (braking) and recharges (regenerates) the batteries.
Dual power. Power can come from either the engine, motor, or both depending on driving circumstances. Additional power to assist the engine in accelerating or climbing might be provided by the electric motor. Or more commonly, a smaller electric motor provides all of the power for low-speed driving conditions and is augmented by the engine at higher speeds.
Automatic start/shutoff. It automatically shuts off the engine when the vehicle comes to a stop and restarts it when the accelerator is pressed down. This automation is much simpler with an electric motor. Also, see dual power above.
Alternative green vehicles
Other types of green vehicles include other vehicles that go fully or partly on alternative energy sources than fossil fuel. Another option is to use alternative fuel composition (i.e. biofuels) in conventional fossil fuel-based vehicles, making them go partly on renewable energy sources.
Other approaches include personal rapid transit, a public transportation concept that offers automated on-demand non-stop transportation, on a network of specially built guideways.
Marketing
Adoption
Automakers spend around $US8 million in marketing Hybrid vehicles each year. With combined effort from many car companies, the Hybrid industry has sold millions of Hybrids.
Hybrid car companies like Toyota, Honda, Ford, and BMW have pulled together to create a movement of Hybrid vehicle sales pushed by Washington lobbyists to lower the world's emissions and become less reliant on our petroleum consumption. | Hybrid vehicle | Wikipedia | 383 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
In 2005, sales went beyond 200,000 Hybrids, but in retrospect that only reduced the global use for gasoline consumption by 200,000 gallons per day—a tiny fraction of the 360 million gallons used per day. According to Bradley Berman author of Driving Change—One Hybrid at a time, "cold economics shows that in real dollars, except for a brief spike in the 1970s, gas prices have remained remarkably steady and cheap. Fuel continues to represent a small part of the overall cost of owning and operating a personal vehicle". Other marketing tactics include greenwashing which is the "unjustified appropriation of environmental virtue."
Temma Ehrenfeld explained in an article by Newsweek. Hybrids may be more efficient than many other gasoline motors as far as gasoline consumption is concerned but as far as being green and good for the environment is completely inaccurate.
Hybrid car companies have a long time to go if they expect to really go green. According to Harvard business professor Theodore Levitt states "managing products" and "meeting customers' needs", "you must adapt to consumer expectations and anticipation of future desires." This means people buy what they want, if they want a fuel efficient car they buy a Hybrid without thinking about the actual efficiency of the product. This "green myopia" as Ottman calls it, fails because marketers focus on the greenness of the product and not on the actual effectiveness.
Researchers and analysts say people are drawn to the new technology, as well as the convenience of fewer fill-ups. Secondly, people find it rewarding to own the better, newer, flashier, and so-called greener car.
Misleading advertising
In 2019 the term self-charging hybrid became prevalent in advertising, though cars referred to by this name do not offer any different functionality than a standard hybrid electric vehicle provides. The only self-charging effect is in energy recovery via regenerative braking, which is also true of plug-in hybrids, fuel cell electric vehicles and battery electric vehicles.
In January 2020, using this term has been prohibited in Norway, for misleading advertising by Toyota and Lexus. "Our claim is based on the fact that customers never have to charge the battery of their vehicle, as it is recharged during the vehicle use. There is no intention to mislead customers, on the contrary: the point is to clearly explain the difference with plug-in hybrid vehicles." | Hybrid vehicle | Wikipedia | 487 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
Adoption rate
While the adoption rate for hybrids in the US is small today (2.2% of new car sales in 2011), this compares with a 17.1% share of new car sales in Japan in 2011, and it has the potential to be very large over time as more models are offered and incremental costs decline due to learning and scale benefits. However, forecasts vary widely. For instance, Bob Lutz, a long-time skeptic of hybrids, indicated he expects hybrids "will never comprise more than 10% of the US auto market." Other sources also expect hybrid penetration rates in the US will remain under 10% for many years.
More optimistic views as of 2006 include predictions that hybrids would dominate new car sales in the US and elsewhere over the next 10 to 20 years. Another approach, taken by Saurin Shah, examines the penetration rates (or S-curves) of four analogs (historical and current) to hybrid and electrical vehicles in an attempt to gauge how quickly the vehicle stock could be hybridized and/or electrified in the United States. The analogs are (1) the electric motors in US factories in the early 20th century, (2) diesel-electric locomotives on US railways in the 1920–1945 period, (3) a range of new automotive features/technologies introduced in the US over the past fifty years, and 4) e-bike purchases in China over the past few years. These analogs collectively suggest it would take at least 30 years for hybrid and electric vehicles to capture 80% of the US passenger vehicle stock.
The EPA expects the combined market share of new gasoline hybrid light-duty vehicles to reach 13.6% for the 2023 model year from 10.2% in the 2022 model year.
European Union 2020 regulation standards
The European Parliament, Council, and European Commission have reached an agreement which is aimed at reducing the average CO2 passenger car emissions to 95 g/km by 2020, according to a European Commission press release.
According to the release, the key details of the agreement are as follows: | Hybrid vehicle | Wikipedia | 421 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
Emissions target: The agreement will reduce average CO2 emissions from new cars to 95 g/km from 2020, as proposed by the commission. This is a 40% reduction from the mandatory 2015 target of 130 g/km. The target is an average for each manufacturer's new car fleet; it allows OEMs to build some vehicles that emit less than the average and some that emit more.
2025 target: The commission is required to propose a further emissions reduction target by the end-2015 to take effect in 2025. This target will be in line with the EU's long-term climate goals.
Super credits for low-emission vehicles: The Regulation will give manufacturers additional incentives to produce cars with CO2 emissions of 50 g/km or less (which will be electric or plug-in hybrid cars). Each of these vehicles will be counted as two vehicles in 2020, 1.67 in 2021, 1.33 in 2022, and then as one vehicle from 2023 onwards. These super credits will help manufacturers further reduce the average emissions of their new car fleet. However, to prevent the scheme from undermining the environmental integrity of the legislation, there will be a 2.5 g/km cap per manufacturer on the contribution that super credits can make to their target in any year. | Hybrid vehicle | Wikipedia | 266 | 157736 | https://en.wikipedia.org/wiki/Hybrid%20vehicle | Technology | Basics_7 | null |
Permafrost () is soil or underwater sediment which continuously remains below for two years or more: the oldest permafrost had been continuously frozen for around 700,000 years. Whilst the shallowest permafrost has a vertical extent of below a meter (3 ft), the deepest is greater than . Similarly, the area of individual permafrost zones may be limited to narrow mountain summits or extend across vast Arctic regions. The ground beneath glaciers and ice sheets is not usually defined as permafrost, so on land, permafrost is generally located beneath a so-called active layer of soil which freezes and thaws depending on the season.
Around 15% of the Northern Hemisphere or 11% of the global surface is underlain by permafrost, covering a total area of around . This includes large areas of Alaska, Canada, Greenland, and Siberia. It is also located in high mountain regions, with the Tibetan Plateau being a prominent example. Only a minority of permafrost exists in the Southern Hemisphere, where it is consigned to mountain slopes like in the Andes of Patagonia, the Southern Alps of New Zealand, or the highest mountains of Antarctica.
Permafrost contains large amounts of dead biomass that have accumulated throughout millennia without having had the chance to fully decompose and release their carbon, making tundra soil a carbon sink. As global warming heats the ecosystem, frozen soil thaws and becomes warm enough for decomposition to start anew, accelerating the permafrost carbon cycle. Depending on conditions at the time of thaw, decomposition can release either carbon dioxide or methane, and these greenhouse gas emissions act as a climate change feedback. The emissions from thawing permafrost will have a sufficient impact on the climate to impact global carbon budgets. It is difficult to accurately predict how much greenhouse gases the permafrost releases because of the different thaw processes are still uncertain. There is widespread agreement that the emissions will be smaller than human-caused emissions and not large enough to result in runaway warming. Instead, the annual permafrost emissions are likely comparable with global emissions from deforestation, or to annual emissions of large countries such as Russia, the United States or China. | Permafrost | Wikipedia | 458 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Apart from its climate impact, permafrost thaw brings more risks. Formerly frozen ground often contains enough ice that when it thaws, hydraulic saturation is suddenly exceeded, so the ground shifts substantially and may even collapse outright. Many buildings and other infrastructure were built on permafrost when it was frozen and stable, and so are vulnerable to collapse if it thaws. Estimates suggest nearly 70% of such infrastructure is at risk by 2050, and that the associated costs could rise to tens of billions of dollars in the second half of the century. Furthermore, between 13,000 and 20,000 sites contaminated with toxic waste are present in the permafrost, as well as the natural mercury deposits, which are all liable to leak and pollute the environment as the warming progresses. Lastly, concerns have been raised about the potential for pathogenic microorganisms surviving the thaw and contributing to future pandemics. However, this is considered unlikely, and a scientific review on the subject describes the risks as "generally low".
Classification and extent
Permafrost is soil, rock or sediment that is frozen for more than two consecutive years. In practice, this means that permafrost occurs at a mean annual temperature of or below. In the coldest regions, the depth of continuous permafrost can exceed . It typically exists beneath the so-called active layer, which freezes and thaws annually, and so can support plant growth, as the roots can only take hold in the soil that's thawed. Active layer thickness is measured during its maximum extent at the end of summer: as of 2018, the average thickness in the Northern Hemisphere is ~, but there are significant regional differences. Northeastern Siberia, Alaska and Greenland have the most solid permafrost with the lowest extent of active layer (less than on average, and sometimes only ), while southern Norway and the Mongolian Plateau are the only areas where the average active layer is deeper than , with the record of . The border between active layer and permafrost itself is sometimes called permafrost table. | Permafrost | Wikipedia | 427 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Around 15% of Northern Hemisphere land that is not completely covered by ice is directly underlain by permafrost; 22% is defined as part of a permafrost zone or region. This is because only slightly more than half of this area is defined as a continuous permafrost zone, where 90%–100% of the land is underlain by permafrost. Around 20% is instead defined as discontinuous permafrost, where the coverage is between 50% and 90%. Finally, the remaining <30% of permafrost regions consists of areas with 10%–50% coverage, which are defined as sporadic permafrost zones, and some areas that have isolated patches of permafrost covering 10% or less of their area. Most of this area is found in Siberia, northern Canada, Alaska and Greenland. Beneath the active layer annual temperature swings of permafrost become smaller with depth. The greatest depth of permafrost occurs right before the point where geothermal heat maintains a temperature above freezing. Above that bottom limit there may be permafrost with a consistent annual temperature—"isothermal permafrost".
Continuity of coverage
Permafrost typically forms in any climate where the mean annual air temperature is lower than the freezing point of water. Exceptions are found in humid boreal forests, such as in Northern Scandinavia and the North-Eastern part of European Russia west of the Urals, where snow acts as an insulating blanket. Glaciated areas may also be exceptions. Since all glaciers are warmed at their base by geothermal heat, temperate glaciers, which are near the pressure melting point throughout, may have liquid water at the interface with the ground and are therefore free of underlying permafrost. "Fossil" cold anomalies in the geothermal gradient in areas where deep permafrost developed during the Pleistocene persist down to several hundred metres. This is evident from temperature measurements in boreholes in North America and Europe.
Discontinuous permafrost | Permafrost | Wikipedia | 420 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
The below-ground temperature varies less from season to season than the air temperature, with mean annual temperatures tending to increase with depth due to the geothermal crustal gradient. Thus, if the mean annual air temperature is only slightly below , permafrost will form only in spots that are sheltered (usually with a northern or southern aspect, in the north and south hemispheres respectively) creating discontinuous permafrost. Usually, permafrost will remain discontinuous in a climate where the mean annual soil surface temperature is between . In the moist-wintered areas mentioned before, there may not even be discontinuous permafrost down to . Discontinuous permafrost is often further divided into extensive discontinuous permafrost, where permafrost covers between 50 and 90 percent of the landscape and is usually found in areas with mean annual temperatures between , and sporadic permafrost, where permafrost cover is less than 50 percent of the landscape and typically occurs at mean annual temperatures between .
In soil science, the sporadic permafrost zone is abbreviated SPZ and the extensive discontinuous permafrost zone DPZ. Exceptions occur in un-glaciated Siberia and Alaska where the present depth of permafrost is a relic of climatic conditions during glacial ages where winters were up to colder than those of today.
Continuous permafrost
At mean annual soil surface temperatures below the influence of aspect can never be sufficient to thaw permafrost and a zone of continuous permafrost (abbreviated to CPZ) forms. A line of continuous permafrost in the Northern Hemisphere represents the most southern border where land is covered by continuous permafrost or glacial ice. The line of continuous permafrost varies around the world northward or southward due to regional climatic changes. In the southern hemisphere, most of the equivalent line would fall within the Southern Ocean if there were land there. Most of the Antarctic continent is overlain by glaciers, under which much of the terrain is subject to basal melting. The exposed land of Antarctica is substantially underlain with permafrost, some of which is subject to warming and thawing along the coastline. | Permafrost | Wikipedia | 459 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Alpine permafrost
A range of elevations in both the Northern and Southern Hemisphere are cold enough to support perennially frozen ground: some of the best-known examples include the Canadian Rockies, the European Alps, Himalaya and the Tien Shan. In general, it has been found that extensive alpine permafrost requires mean annual air temperature of , though this can vary depending on local topography, and some mountain areas are known to support permafrost at . It is also possible for subsurface alpine permafrost to be covered by warmer, vegetation-supporting soil.
Alpine permafrost is particularly difficult to study, and systematic research efforts did not begin until the 1970s. Consequently, there remain uncertainties about its geography As recently as 2009, permafrost had been discovered in a new area – Africa's highest peak, Mount Kilimanjaro ( above sea level and approximately 3° south of the equator). In 2014, a collection of regional estimates of alpine permafrost extent had established a global extent of . Yet, by 2014, alpine permafrost in the Andes has not been fully mapped, although its extent has been modeled to assess the amount of water bound up in these areas.
Subsea permafrost
Subsea permafrost occurs beneath the seabed and exists in the continental shelves of the polar regions. These areas formed during the last Ice Age, when a larger portion of Earth's water was bound up in ice sheets on land and when sea levels were low. As the ice sheets melted to again become seawater during the Holocene glacial retreat, coastal permafrost became submerged shelves under relatively warm and salty boundary conditions, compared to surface permafrost. Since then, these conditions led to the gradual and ongoing decline of subsea permafrost extent. Nevertheless, its presence remains an important consideration for the "design, construction, and operation of coastal facilities, structures founded on the seabed, artificial islands, sub-sea pipelines, and wells drilled for exploration and production". Subsea permafrost can also overlay deposits of methane clathrate, which were once speculated to be a major climate tipping point in what was known as a clathrate gun hypothesis, but are now no longer believed to play any role in projected climate change. | Permafrost | Wikipedia | 477 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Past extent of permafrost
At the Last Glacial Maximum, continuous permafrost covered a much greater area than it does today, covering all of ice-free Europe south to about Szeged (southeastern Hungary) and the Sea of Azov (then dry land) and East Asia south to present-day Changchun and Abashiri. In North America, only an extremely narrow belt of permafrost existed south of the ice sheet at about the latitude of New Jersey through southern Iowa and northern Missouri, but permafrost was more extensive in the drier western regions where it extended to the southern border of Idaho and Oregon. In the Southern Hemisphere, there is some evidence for former permafrost from this period in central Otago and Argentine Patagonia, but was probably discontinuous, and is related to the tundra. Alpine permafrost also occurred in the Drakensberg during glacial maxima above about .
Manifestations
Base depth
Permafrost extends to a base depth where geothermal heat from the Earth and the mean annual temperature at the surface achieve an equilibrium temperature of . This base depth of permafrost can vary wildly – it is less than a meter (3 ft) in the areas where it is shallowest, yet reaches in the northern Lena and Yana River basins in Siberia. Calculations indicate that the formation time of permafrost greatly slows past the first several metres. For instance, over half a million years was required to form the deep permafrost underlying Prudhoe Bay, Alaska, a time period extending over several glacial and interglacial cycles of the Pleistocene.
Base depth is affected by the underlying geology, and particularly by thermal conductivity, which is lower for permafrost in soil than in bedrock. Lower conductivity leaves permafrost less affected by the geothermal gradient, which is the rate of increasing temperature with respect to increasing depth in the Earth's interior. It occurs as the Earth's internal thermal energy is generated by radioactive decay of unstable isotopes and flows to the surface by conduction at a rate of ~47 terawatts (TW). Away from tectonic plate boundaries, this is equivalent to an average heat flow of 25–30 °C/km (124–139 °F/mi) near the surface.
Massive ground ice | Permafrost | Wikipedia | 481 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
When the ice content of a permafrost exceeds 250 percent (ice to dry soil by mass) it is classified as massive ice. Massive ice bodies can range in composition, in every conceivable gradation from icy mud to pure ice. Massive icy beds have a minimum thickness of at least 2 m and a short diameter of at least 10 m. First recorded North American observations of this phenomenon were by European scientists at Canning River (Alaska) in 1919. Russian literature provides an earlier date of 1735 and 1739 during the Great North Expedition by P. Lassinius and Khariton Laptev, respectively. Russian investigators including I.A. Lopatin, B. Khegbomov, S. Taber and G. Beskow had also formulated the original theories for ice inclusion in freezing soils.
While there are four categories of ice in permafrost – pore ice, ice wedges (also known as vein ice), buried surface ice and intrasedimental (sometimes also called constitutional) ice – only the last two tend to be large enough to qualify as massive ground ice. These two types usually occur separately, but may be found together, like on the coast of Tuktoyaktuk in western Arctic Canada, where the remains of Laurentide Ice Sheet are located.
Buried surface ice may derive from snow, frozen lake or sea ice, aufeis (stranded river ice) and even buried glacial ice from the former Pleistocene ice sheets. The latter hold enormous value for paleoglaciological research, yet even as of 2022, the total extent and volume of such buried ancient ice is unknown. Notable sites with known ancient ice deposits include Yenisei River valley in Siberia, Russia as well as Banks and Bylot Island in Canada's Nunavut and Northwest Territories. Some of the buried ice sheet remnants are known to host thermokarst lakes. | Permafrost | Wikipedia | 386 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Intrasedimental or constitutional ice has been widely observed and studied across Canada. It forms when subterranean waters freeze in place, and is subdivided into intrusive, injection and segregational ice. The latter is the dominant type, formed after crystallizational differentiation in wet sediments, which occurs when water migrates to the freezing front under the influence of van der Waals forces. This is a slow process, which primarily occurs in silts with salinity less than 20% of seawater: silt sediments with higher salinity and clay sediments instead have water movement prior to ice formation dominated by rheological processes. Consequently, it takes between 1 and 1000 years to form intrasedimental ice in the top 2.5 meters of clay sediments, yet it takes between 10 and 10,000 years for peat sediments and between 1,000 and 1,000,000 years for silt sediments.
Landforms
Permafrost processes such as thermal contraction generating cracks which eventually become ice wedges and solifluction – gradual movement of soil down the slope as it repeatedly freezes and thaws – often lead to the formation of ground polygons, rings, steps and other forms of patterned ground found in arctic, periglacial and alpine areas. In ice-rich permafrost areas, melting of ground ice initiates thermokarst landforms such as thermokarst lakes, thaw slumps, thermal-erosion gullies, and active layer detachments. Notably, unusually deep permafrost in Arctic moorlands and bogs often attracts meltwater in warmer seasons, which pools and freezes to form ice lenses, and the surrounding ground begins to jut outward at a slope. This can eventually result in the formation of large-scale land forms around this core of permafrost, such as palsas – long (), wide () yet shallow (< tall) peat mounds – and the even larger pingos, which can be high and in diameter.
Ecology
Only plants with shallow roots can survive in the presence of permafrost. Black spruce tolerates limited rooting zones, and dominates flora where permafrost is extensive. Likewise, animal species which live in dens and burrows have their habitat constrained by the permafrost, and these constraints also have a secondary impact on interactions between species within the ecosystem. | Permafrost | Wikipedia | 482 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
While permafrost soil is frozen, it is not completely inhospitable to microorganisms, though their numbers can vary widely, typically from 1 to 1000 million per gram of soil.
The permafrost carbon cycle (Arctic Carbon Cycle) deals with the transfer of carbon from permafrost soils to terrestrial vegetation and microbes, to the atmosphere, back to vegetation, and finally back to permafrost soils through burial and sedimentation due to cryogenic processes. Some of this carbon is transferred to the ocean and other portions of the globe through the global carbon cycle. The cycle includes the exchange of carbon dioxide and methane between terrestrial components and the atmosphere, as well as the transfer of carbon between land and water as methane, dissolved organic carbon, dissolved inorganic carbon, particulate inorganic carbon and particulate organic carbon.
Most of the bacteria and fungi found in permafrost cannot be cultured in the laboratory, but the identity of the microorganisms can be revealed by DNA-based techniques. For instance, analysis of 16S rRNA genes from late Pleistocene permafrost samples in eastern Siberia's Kolyma Lowland revealed eight phylotypes, which belonged to the phyla Actinomycetota and Pseudomonadota. "Muot-da-Barba-Peider", an alpine permafrost site in eastern Switzerland, was found to host a diverse microbial community in 2016. Prominent bacteria groups included phylum Acidobacteriota, Actinomycetota, AD3, Bacteroidota, Chloroflexota, Gemmatimonadota, OD1, Nitrospirota, Planctomycetota, Pseudomonadota, and Verrucomicrobiota, in addition to eukaryotic fungi like Ascomycota, Basidiomycota, and Zygomycota. In the presently living species, scientists observed a variety of adaptations for sub-zero conditions, including reduced and anaerobic metabolic processes.
Construction on permafrost | Permafrost | Wikipedia | 430 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
There are only two large cities in the world built in areas of continuous permafrost (where the frozen soil forms an unbroken, below-zero sheet) and both are in Russia – Norilsk in Krasnoyarsk Krai and Yakutsk in the Sakha Republic. Building on permafrost is difficult because the heat of the building (or pipeline) can spread to the soil, thawing it. As ice content turns to water, the ground's ability to provide structural support is weakened, until the building is destabilized. For instance, during the construction of the Trans-Siberian Railway, a steam engine factory complex built in 1901 began to crumble within a month of operations for these reasons. Additionally, there is no groundwater available in an area underlain with permafrost. Any substantial settlement or installation needs to make some alternative arrangement to obtain water.
A common solution is placing foundations on wood piles, a technique pioneered by Soviet engineer Mikhail Kim in Norilsk. However, warming-induced change of friction on the piles can still cause movement through creep, even as the soil remains frozen. The Melnikov Permafrost Institute in Yakutsk found that pile foundations should extend down to to avoid the risk of buildings sinking. At this depth the temperature does not change with the seasons, remaining at about .
Two other approaches are building on an extensive gravel pad (usually thick); or using anhydrous ammonia heat pipes. The Trans-Alaska Pipeline System uses heat pipes built into vertical supports to prevent the pipeline from sinking and the Qingzang railway in Tibet employs a variety of methods to keep the ground cool, both in areas with frost-susceptible soil. Permafrost may necessitate special enclosures for buried utilities, called "utilidors".
Impacts of climate change
Increasing active layer thickness
Globally, permafrost warmed by about between 2007 and 2016, with stronger warming observed in the continuous permafrost zone relative to the discontinuous zone. Observed warming was up to in parts of Northern Alaska (early 1980s to mid-2000s) and up to in parts of the Russian European North (1970–2020). This warming inevitably causes permafrost to thaw: active layer thickness has increased in the European and Russian Arctic across the 21st century and at high elevation areas in Europe and Asia since the 1990s.
Between 2000 and 2018, the average active layer thickness had increased from ~ to ~, at an average annual rate of ~. | Permafrost | Wikipedia | 511 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
In Yukon, the zone of continuous permafrost might have moved poleward since 1899, but accurate records only go back 30 years. The extent of subsea permafrost is decreasing as well; as of 2019, ~97% of permafrost under Arctic ice shelves is becoming warmer and thinner.
Based on high agreement across model projections, fundamental process understanding, and paleoclimate evidence, it is virtually certain that permafrost extent and volume will continue to shrink as the global climate warms, with the extent of the losses determined by the magnitude of warming.
Permafrost thaw is associated with a wide range of issues, and International Permafrost Association (IPA) exists to help address them. It convenes International Permafrost Conferences and maintains Global Terrestrial Network for Permafrost, which undertakes special projects such as preparing databases, maps, bibliographies, and glossaries, and coordinates international field programmes and networks.
Climate change feedback
As recent warming deepens the active layer subject to permafrost thaw, this exposes formerly stored carbon to biogenic processes which facilitate its entrance into the atmosphere as carbon dioxide and methane. Because carbon emissions from permafrost thaw contribute to the same warming which facilitates the thaw, it is a well-known example of a positive climate change feedback. Permafrost thaw is sometimes included as one of the major tipping points in the climate system due to the exhibition of local thresholds and its effective irreversibility. However, while there are self-perpetuating processes that apply on the local or regional scale, it is debated as to whether it meets the strict definition of a global tipping point as in aggregate permafrost thaw is gradual with warming. | Permafrost | Wikipedia | 365 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
In the northern circumpolar region, permafrost contains organic matter equivalent to 1400–1650 billion tons of pure carbon, which was built up over thousands of years. This amount equals almost half of all organic material in all soils, and it is about twice the carbon content of the atmosphere, or around four times larger than the human emissions of carbon between the start of the Industrial Revolution and 2011. Further, most of this carbon (~1,035 billion tons) is stored in what is defined as the near-surface permafrost, no deeper than below the surface. However, only a fraction of this stored carbon is expected to enter the atmosphere. In general, the volume of permafrost in the upper 3 m of ground is expected to decrease by about 25% per of global warming, yet even under the RCP8.5 scenario associated with over of global warming by the end of the 21st century, about 5% to 15% of permafrost carbon is expected to be lost "over decades and centuries".
The exact amount of carbon that will be released due to warming in a given permafrost area depends on depth of thaw, carbon content within the thawed soil, physical changes to the environment, and microbial and vegetation activity in the soil. Notably, estimates of carbon release alone do not fully represent the impact of permafrost thaw on climate change. This is because carbon can be released through either aerobic or anaerobic respiration, which results in carbon dioxide (CO2) or methane (CH4) emissions, respectively. While methane lasts less than 12 years in the atmosphere, its global warming potential is around 80 times larger than that of CO2 over a 20-year period and about 28 times larger over a 100-year period. While only a small fraction of permafrost carbon will enter the atmosphere as methane, those emissions will cause 40–70% of the total warming caused by permafrost thaw during the 21st century. Much of the uncertainty about the eventual extent of permafrost methane emissions is caused by the difficulty of accounting for the recently discovered abrupt thaw processes, which often increase the fraction of methane emitted over carbon dioxide in comparison to the usual gradual thaw processes. | Permafrost | Wikipedia | 464 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Another factor which complicates projections of permafrost carbon emissions is the ongoing "greening" of the Arctic. As climate change warms the air and the soil, the region becomes more hospitable to plants, including larger shrubs and trees which could not survive there before. Thus, the Arctic is losing more and more of its tundra biomes, yet it gains more plants, which proceed to absorb more carbon. Some of the emissions caused by permafrost thaw will be offset by this increased plant growth, but the exact proportion is uncertain. It is considered very unlikely that this greening could offset all of the emissions from permafrost thaw during the 21st century, and even less likely that it could continue to keep pace with those emissions after the 21st century. Further, climate change also increases the risk of wildfires in the Arctic, which can substantially accelerate emissions of permafrost carbon.
Impact on global temperatures
Altogether, it is expected that cumulative greenhouse gas emissions from permafrost thaw will be smaller than the cumulative anthropogenic emissions, yet still substantial on a global scale, with some experts comparing them to emissions caused by deforestation. The IPCC Sixth Assessment Report estimates that carbon dioxide and methane released from permafrost could amount to the equivalent of 14–175 billion tonnes of carbon dioxide per of warming. For comparison, by 2019, annual anthropogenic emissions of carbon dioxide alone stood around 40 billion tonnes. A major review published in the year 2022 concluded that if the goal of preventing of warming was realized, then the average annual permafrost emissions throughout the 21st century would be equivalent to the year 2019 annual emissions of Russia. Under RCP4.5, a scenario considered close to the current trajectory and where the warming stays slightly below , annual permafrost emissions would be comparable to year 2019 emissions of Western Europe or the United States, while under the scenario of high global warming and worst-case permafrost feedback response, they would approach year 2019 emissions of China. | Permafrost | Wikipedia | 419 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Fewer studies have attempted to describe the impact directly in terms of warming. A 2018 paper estimated that if global warming was limited to , gradual permafrost thaw would add around to global temperatures by 2100, while a 2022 review concluded that every of global warming would cause and from abrupt thaw by the year 2100 and 2300. Around of global warming, abrupt (around 50 years) and widespread collapse of permafrost areas could occur, resulting in an additional warming of .
Thaw-induced ground instability
As the water drains or evaporates, soil structure weakens and sometimes becomes viscous until it regains strength with decreasing moisture content. One visible sign of permafrost degradation is the random displacement of trees from their vertical orientation in permafrost areas. Global warming has been increasing permafrost slope disturbances and sediment supplies to fluvial systems, resulting in exceptional increases in river sediment. On the other hands, disturbance of formerly hard soil increases drainage of water reservoirs in northern wetlands. This can dry them out and compromise the survival of plants and animals used to the wetland ecosystem.
In high mountains, much of the structural stability can be attributed to glaciers and permafrost. As climate warms, permafrost thaws, decreasing slope stability and increasing stress through buildup of pore-water pressure, which may ultimately lead to slope failure and rockfalls. Over the past century, an increasing number of alpine rock slope failure events in mountain ranges around the world have been recorded, and some have been attributed to permafrost thaw induced by climate change. The 1987 Val Pola landslide that killed 22 people in the Italian Alps is considered one such example. In 2002, massive rock and ice falls (up to 11.8 million m3), earthquakes (up to 3.9 Richter), floods (up to 7.8 million m3 water), and rapid rock-ice flow to long distances (up to 7.5 km at 60 m/s) were attributed to slope instability in high mountain permafrost. | Permafrost | Wikipedia | 420 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Permafrost thaw can also result in the formation of frozen debris lobes (FDLs), which are defined as "slow-moving landslides composed of soil, rocks, trees, and ice". This is a notable issue in the Alaska's southern Brooks Range, where some FDLs measured over in width, in height, and in length by 2012. As of December 2021, there were 43 frozen debris lobes identified in the southern Brooks Range, where they could potentially threaten both the Trans Alaska Pipeline System (TAPS) corridor and the Dalton Highway, which is the main transport link between the Interior Alaska and the Alaska North Slope.
Infrastructure
As of 2021, there are 1162 settlements located directly atop the Arctic permafrost, which host an estimated 5 million people. By 2050, permafrost layer below 42% of these settlements is expected to thaw, affecting all their inhabitants (currently 3.3 million people). Consequently, a wide range of infrastructure in permafrost areas is threatened by the thaw. By 2050, it's estimated that nearly 70% of global infrastructure located in the permafrost areas would be at high risk of permafrost thaw, including 30–50% of "critical" infrastructure. The associated costs could reach tens of billions of dollars by the second half of the century. Reducing greenhouse gas emissions in line with the Paris Agreement is projected to stabilize the risk after mid-century; otherwise, it'll continue to worsen. | Permafrost | Wikipedia | 309 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
In Alaska alone, damages to infrastructure by the end of the century would amount to $4.6 billion (at 2015 dollar value) if RCP8.5, the high-emission climate change scenario, were realized. Over half stems from the damage to buildings ($2.8 billion), but there's also damage to roads ($700 million), railroads ($620 million), airports ($360 million) and pipelines ($170 million). Similar estimates were done for RCP4.5, a less intense scenario which leads to around by 2100, a level of warming similar to the current projections. In that case, total damages from permafrost thaw are reduced to $3 billion, while damages to roads and railroads are lessened by approximately two-thirds (from $700 and $620 million to $190 and $220 million) and damages to pipelines are reduced more than ten-fold, from $170 million to $16 million. Unlike the other costs stemming from climate change in Alaska, such as damages from increased precipitation and flooding, climate change adaptation is not a viable way to reduce damages from permafrost thaw, as it would cost more than the damage incurred under either scenario.
In Canada, Northwest Territories have a population of only 45,000 people in 33 communities, yet permafrost thaw is expected to cost them $1.3 billion over 75 years, or around $51 million a year. In 2006, the cost of adapting Inuvialuit homes to permafrost thaw was estimated at $208/m2 if they were built at pile foundations, and $1,000/m2 if they didn't. At the time, the average area of a residential building in the territory was around 100 m2. Thaw-induced damage is also unlikely to be covered by home insurance, and to address this reality, territorial government currently funds Contributing Assistance for Repairs and Enhancements (CARE) and Securing Assistance for Emergencies (SAFE) programs, which provide long- and short-term forgivable loans to help homeowners adapt. It is possible that in the future, mandatory relocation would instead take place as the cheaper option. However, it would effectively tear the local Inuit away from their ancestral homelands. Right now, their average personal income is only half that of the median NWT resident, meaning that adaptation costs are already disproportionate for them. | Permafrost | Wikipedia | 499 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
By 2022, up to 80% of buildings in some Northern Russia cities had already experienced damage. By 2050, the damage to residential infrastructure may reach $15 billion, while total public infrastructure damages could amount to 132 billion. This includes oil and gas extraction facilities, of which 45% are believed to be at risk.
Outside of the Arctic, Qinghai–Tibet Plateau (sometimes known as "the Third Pole"), also has an extensive permafrost area. It is warming at twice the global average rate, and 40% of it is already considered "warm" permafrost, making it particularly unstable. Qinghai–Tibet Plateau has a population of over 10 million people – double the population of permafrost regions in the Arctic – and over 1 million m2 of buildings are located in its permafrost area, as well as 2,631 km of power lines, and 580 km of railways. There are also 9,389 km of roads, and around 30% are already sustaining damage from permafrost thaw. Estimates suggest that under the scenario most similar to today, SSP2-4.5, around 60% of the current infrastructure would be at high risk by 2090 and simply maintaining it would cost $6.31 billion, with adaptation reducing these costs by 20.9% at most. Holding the global warming to would reduce these costs to $5.65 billion, and fulfilling the optimistic Paris Agreement target of would save a further $1.32 billion. In particular, fewer than 20% of railways would be at high risk by 2100 under , yet this increases to 60% at , while under SSP5-8.5, this level of risk is met by mid-century.
Release of toxic pollutants
For much of the 20th century, it was believed that permafrost would "indefinitely" preserve anything buried there, and this made deep permafrost areas popular locations for hazardous waste disposal. In places like Canada's Prudhoe Bay oil field, procedures were developed documenting the "appropriate" way to inject waste beneath the permafrost. This means that as of 2023, there are ~4500 industrial facilities in the Arctic permafrost areas which either actively process or store hazardous chemicals. Additionally, there are between 13,000 and 20,000 sites which have been heavily contaminated, 70% of them in Russia, and their pollution is currently trapped in the permafrost. | Permafrost | Wikipedia | 507 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
About a fifth of both the industrial and the polluted sites (1000 and 2200–4800) are expected to start thawing in the future even if the warming does not increase from its 2020 levels. Only about 3% more sites would start thawing between now and 2050 under the climate change scenario consistent with the Paris Agreement goals, RCP2.6, but by 2100, about 1100 more industrial facilities and 3500 to 5200 contaminated sites are expected to start thawing even then. Under the very high emission scenario RCP8.5, 46% of industrial and contaminated sites would start thawing by 2050, and virtually all of them would be affected by the thaw by 2100.
Organochlorines and other persistent organic pollutants are of a particular concern, due to their potential to repeatedly reach local communities after their re-release through biomagnification in fish. At worst, future generations born in the Arctic would enter life with weakened immune systems due to pollutants accumulating across generations.
A notable example of pollution risks associated with permafrost was the 2020 Norilsk oil spill, caused by the collapse of diesel fuel storage tank at Norilsk-Taimyr Energy's thermal power plant No. 3. It spilled 6,000 tonnes of fuel into the land and 15,000 into the water, polluting Ambarnaya, Daldykan and many smaller rivers on Taimyr Peninsula, even reaching lake Pyasino, which is a crucial water source in the area. State of emergency at the federal level was declared. The event has been described as the second-largest oil spill in modern Russian history. | Permafrost | Wikipedia | 342 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Another issue associated with permafrost thaw is the release of natural mercury deposits. An estimated 800,000 tons of mercury are frozen in the permafrost soil. According to observations, around 70% of it is simply taken up by vegetation after the thaw. However, if the warming continues under RCP8.5, then permafrost emissions of mercury into the atmosphere would match the current global emissions from all human activities by 2200. Mercury-rich soils also pose a much greater threat to humans and the environment if they thaw near rivers. Under RCP8.5, enough mercury will enter the Yukon River basin by 2050 to make its fish unsafe to eat under the EPA guidelines. By 2100, mercury concentrations in the river will double. Contrastingly, even if mitigation is limited to RCP4.5 scenario, mercury levels will increase by about 14% by 2100, and will not breach the EPA guidelines even by 2300.
Revival of ancient organisms
Microorganisms
Bacteria are known for being able to remain dormant to survive adverse conditions, and viruses are not metabolically active outside of host cells in the first place. This has motivated concerns that permafrost thaw could free previously unknown microorganisms, which may be capable of infecting either humans or important livestock and crops, potentially resulting in damaging epidemics or pandemics. Further, some scientists argue that horizontal gene transfer could occur between the older, formerly frozen bacteria, and modern ones, and one outcome could be the introduction of novel antibiotic resistance genes into the genome of current pathogens, exacerbating what is already expected to become a difficult issue in the future.
At the same time, notable pathogens like influenza and smallpox appear unable to survive being thawed, and other scientists argue that the risk of ancient microorganisms being both able to survive the thaw and to threaten humans is not scientifically plausible. Likewise, some research suggests that antimicrobial resistance capabilities of ancient bacteria would be comparable to, or even inferior to modern ones. | Permafrost | Wikipedia | 424 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Plants
In 2012, Russian researchers proved that permafrost can serve as a natural repository for ancient life forms by reviving a sample of Silene stenophylla from 30,000-year-old tissue found in an Ice Age squirrel burrow in the Siberian permafrost. This is the oldest plant tissue ever revived. The resultant plant was fertile, producing white flowers and viable seeds. The study demonstrated that living tissue can survive ice preservation for tens of thousands of years.
History of scientific research
Between the middle of the 19th century and the middle of the 20th century, most of the literature on basic permafrost science and the engineering aspects of permafrost was written in Russian. One of the earliest written reports describing the existence of permafrost dates to 1684, when well excavation efforts in Yakutsk were stumped by its presence. A significant role in the initial permafrost research was played by Alexander von Middendorff (1815–1894) and Karl Ernst von Baer, a Baltic German scientist at the University of Königsberg, and a member of the St Petersburg Academy of Sciences. Baer began publishing works on permafrost in 1838 and is often considered the "founder of scientific permafrost research." Baer laid the foundation for modern permafrost terminology by compiling and analyzing all available data on ground ice and permafrost.
Baer is also known to have composed the world's first permafrost textbook in 1843, "materials for the study of the perennial ground-ice", written in his native language. However, it was not printed then, and a Russian translation wasn't ready until 1942. The original German textbook was believed to be lost until the typescript from 1843 was discovered in the library archives of the University of Giessen. The 234-page text was available online, with additional maps, preface and comments. Notably, Baer's southern limit of permafrost in Eurasia drawn in 1843 corresponds well with the actual southern limit verified by modern research. | Permafrost | Wikipedia | 422 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
Beginning in 1942, Siemon William Muller delved into the relevant Russian literature held by the Library of Congress and the U.S. Geological Survey Library so that he was able to furnish the government an engineering field guide and a technical report about permafrost by 1943. That report coined the English term as a contraction of permanently frozen ground, in what was considered a direct translation of the Russian term (). In 1953, this translation was criticized by another USGS researcher Inna Poiré, as she believed the term had created unrealistic expectations about its stability: more recently, some researchers have argued that "perpetually refreezing" would be a more suitable translation. The report itself was classified (as U.S. Army. Office of the Chief of Engineers, Strategic Engineering Study, no. 62, 1943), until a revised version was released in 1947, which is regarded as the first North American treatise on the subject.
Between 11 and 15 November 1963, the First International Conference on Permafrost took place on the grounds of Purdue University in the American town of West Lafayette, Indiana. It involved 285 participants (including "engineers, manufacturers and builders" who attended alongside the researchers) from a range of countries (Argentina, Austria, Canada, Germany, Great Britain, Japan, Norway, Poland, Sweden, Switzerland, the US and the USSR). This marked the beginning of modern scientific collaboration on the subject. Conferences continue to take place every five years. During the Fourth conference in 1983, a special meeting between the "Big Four" participant countries (US, USSR, China, and Canada) officially created the International Permafrost Association.
In recent decades, permafrost research has attracted more attention than ever due to its role in climate change. Consequently, there has been a massive acceleration in published scientific literature. Around 1990, almost no papers were released containing the words "permafrost" and "carbon": by 2020, around 400 such papers were published every year. | Permafrost | Wikipedia | 407 | 157774 | https://en.wikipedia.org/wiki/Permafrost | Physical sciences | Glaciology | null |
A meteor shower is a celestial event in which a number of meteors are observed to radiate, or originate, from one point in the night sky. These meteors are caused by streams of cosmic debris called meteoroids entering Earth's atmosphere at extremely high speeds on parallel trajectories. Most meteors are smaller than a grain of sand, so almost all of them disintegrate and never hit the Earth's surface. Very intense or unusual meteor showers are known as meteor outbursts and meteor storms, which produce at least 1,000 meteors an hour, most notably from the Leonids. The Meteor Data Centre lists over 900 suspected meteor showers of which about 100 are well established. Several organizations point to viewing opportunities on the Internet. NASA maintains a daily map of active meteor showers.
Historical developments
A meteor shower in August 1583 was recorded in the Timbuktu manuscripts.
In the modern era, the first great meteor storm was the Leonids of November 1833. One estimate is a peak rate of over one hundred thousand meteors an hour, but another, done as the storm abated, estimated more than two hundred thousand meteors during the 9 hours of the storm, over the entire region of North America east of the Rocky Mountains. American Denison Olmsted (1791–1859) explained the event most accurately. After spending the last weeks of 1833 collecting information, he presented his findings in January 1834 to the American Journal of Science and Arts, published in January–April 1834, and January 1836. He noted the shower was of short duration and was not seen in Europe, and that the meteors radiated from a point in the constellation of Leo. He speculated the meteors had originated from a cloud of particles in space. Work continued, yet coming to understand the annual nature of showers though the occurrences of storms perplexed researchers.
The actual nature of meteors was still debated during the 19th century. Meteors were conceived as an atmospheric phenomenon by many scientists (Alexander von Humboldt, Adolphe Quetelet, Julius Schmidt) until the Italian astronomer Giovanni Schiaparelli ascertained the relation between meteors and comets in his work " | Meteor shower | Wikipedia | 438 | 157819 | https://en.wikipedia.org/wiki/Meteor%20shower | Physical sciences | Planetary science | null |
Viola is a genus of flowering plants in the violet family Violaceae. It is the largest genus in the family, containing over 680 species. Most species are found in the temperate Northern Hemisphere; however, some are also found in widely divergent areas such as Hawaii, Australasia, and the Andes.
Some Viola species are perennial plants, some are annual plants, and a few are small shrubs. Many species, varieties and cultivars are grown in gardens for their ornamental flowers. In horticulture, the term pansy is normally used for those multi-colored large-flowered cultivars which are raised annually or biennially from seed and used extensively in bedding. The terms viola and violet are normally reserved for small-flowered annuals or perennials, including the wild species.
Description
Annual or perennial caulescent or acaulescent (with or without a visible plant stem above the ground) herbs, shrubs or very rarely treelets. In acaulescent taxa the foliage and flowers appear to rise from the ground. The remainder have short stems with foliage and flowers produced in the axils of the leaves (axillary).
Viola typically have heart-shaped or reniform (kidney-shaped), scalloped leaves, though a number have linear or palmate leaves. The simple leaves of plants with either habit are arranged alternately; the acaulescent species produce basal rosettes. Plants always have leaves with stipules that are often leaf-like.
The flowers of the vast majority of the species are strongly zygomorphic with bilateral symmetry and solitary, but occasionally form cymes. The flowers are formed from five petals; four are upswept or fan-shaped with two per side, and there is one, broad, lobed lower petal pointing downward. This petal may be slightly or much shorter than the others and is weakly differentiated. The shape of the petals and placement defines many species, for example, some species have a "spur" on the end of each petal while most have a spur on the lower petal. The spur may vary from scarcely exserted (projecting) to very long, such as in Viola rostrata. | Viola (plant) | Wikipedia | 446 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Solitary flowers end long stalks with a pair of bracteoles. The flowers have five sepals that persist after blooming, and in some species the sepals enlarge after blooming. The corolla ranges from white to yellow, orange or various shades of blue and violet or multicolored, often blue and yellow, with or without a yellow throat.
The flowers have five free stamens with short free filaments that are oppressed against the ovary, with a dorsal connective appendage that is large, entire and oblong to ovate. Only the lower two stamens are calcarate (possessing nectary spurs that are inserted on the lowest petal into the spur or a pouch). The styles are filiform (threadlike) or clavate (clubshaped), thickened at their tip, being globose to rostellate (beaked). The stigmas are head-like, narrowed or often beaked. The flowers have a superior ovary with one cell, which has three placentae, containing many ovules.
After flowering, fruit capsules are produced that are thick walled, with few to many seeds per carpel, and dehisce (split open) by way of three valves. On drying, the capsules may eject seeds with considerable force to distances of several meters. The nutlike seeds, which are obovoid to globose, are typically arillate (with a specialized outgrowth) and have straight embryos, flat cotyledons, and soft fleshy endosperm that is oily.
Phytochemistry
One characteristic of some Viola is the elusive scent of their flowers; along with terpenes, a major component of the scent is a ketone compound called ionone, which temporarily desensitizes the receptors of the nose, thus preventing any further scent being detected from the flower until the nerves recover.
Taxonomy | Viola (plant) | Wikipedia | 404 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
History
First formally described by Carl Linnaeus in 1753 with 19 species, the genus Viola bears his botanical authority, L. When Jussieu established the hierarchical system of families (1789), he placed Viola in the Cisti (rock roses), though by 1811 he suggested Viola be separated from these. However, in 1802 Batsch had already established a separate family, which he called Violariae based on Viola as the type genus, with seven other genera. Although Violariae continued to be used by some authors, such as Bentham and Hooker in 1862 (as Violarieae), most authors adopted the alternative name Violaceae, first proposed by de Lamarck and de Candolle in 1805, and Gingins (1823) and Saint-Hilaire (1824). However de Candolle also used Violarieae in his 1824 Prodromus.
Phylogeny
Viola is one of about 25 genera and about 600 species in the large eudicot family Violaceae, divided into subfamilies and tribes. While most genera are monotypic, Viola is a very large genus, variously circumscribed as having between 500 and 600 species. Historically it was placed in subfamily Violoideae, tribe Violeae. But these divisions have been shown to be artificial and not monophyletic. Molecular phylogenetic studies show that Viola occurs in Clade I of the family, as Viola, Schweiggeria, Noisettia and Allexis, in which Schweiggeria and Noisettia are monotypic and form a sister group to Viola. | Viola (plant) | Wikipedia | 323 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Subdivision
Viola is a large genus that has traditionally been treated in sections. One of these was that of Gingins (1823), based on stigma morphology, with five sections (Nomimium, Dischidium, Chamaemelanium, Melanium, Leptidium). The extensive taxonomic studies of Wilhelm Becker, culminating in his 1925 conspectus, resulted in 14 sections and many infrasectional groups. The largest and most diverse, being section Viola, with 17 subsections. In addition to subsections, series were also described. Alternatively, some authors have preferred to subdivide the genus into subgenera. Subsequent treatments were by Gershoy (1934) and Clausen (1964), using subsections and series. These were all based on morphological characteristics. Subsequent studies using molecular phylogenetic methods, such as that of Ballard et al. (1998) have shown that many of these traditional divisions are not monophyletic, the problem being related to a high degree of hybridization. In particular section Nomimium was dismembered into several new sections and transferring part of it to section Viola. Section Viola s. lat. is represented by four sections, Viola sensu stricto, Plagiostigma s. str., Nosphinium sensu lato. and the V. spathulata group. In that analysis, the S American sections appear to be the basal groups, starting with Rubellium, then Leptidium. However, the exact phylogenetic relationships remain unresolved, as a consequence many different taxonomic nomenclatures are in use, including groupings referred to as Grex. Marcussen et al. place the five S American sections, Andinium, Leptidium, Tridens, Rubellium and Chilenium at the base of the phylogenetic tree, in that order. These are followed by the single Australian section, Erpetion, as sister group to Chilenium, the northern hemisphere sections and finally the single African section, V. abyssinica. These sections are morphologically, chromosomally, and geographically distinct.
Sections
Seventeen sections are recognized, listed alphabetically (approximate no. species); | Viola (plant) | Wikipedia | 449 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Sect. Andinium W.Becker (113) S America
Sect. Chamaemelanium Ging. s.lat. (61) N America, northeast Asia (includes Dischidium, Orbiculares)
Subsect. Chamaemelanium
Subsect. Nudicaules
Subsect. Nuttalianae
Sect. Chilenium W.Becker (8) southern S America
Sect. Danxiaviola W. B. Liao et Q. Fan (1) China
Sect. Delphiniopsis W.Becker (3) western Eurasia: southern Spain; Balkans
Sect. Erpetion (Banks) W.Becker (11–18) eastern Australia; Tasmania
Sect. Leptidium Ging. (19) S America
Sect. Melanium Ging. (125) western Eurasia (pansies)
Sect. Nosphinium W.Becker s.lat. (31–50) N, C and northern S America; Beringia; Hawaii
Sect. nov. A (V. abyssinica group) (1–3) Africa: equatorial high mountains
Sect. nov. B (V. spathulata group) (7–9) western and central Asia: northern Iraq to Mongolia
Sect. Plagiostigma Godr. (120) northern hemisphere (includes Diffusae)
Grex Primulifolia
Sect. Rubellium W.Becker (3–6) S America: Chile
Sect. Sclerosium W.Becker (1–4) northeastern Africa to southwestern Asia
Sect. Tridens W.Becker (2) southern S America
Sect. Viola s.str. (Rostellatae nom. illeg.) (75) northern hemisphere (violets) (includes Repentes)
Subsect. Rostratae Kupffer (W.Becker)
Subsect. Viola
Sect. Xylinosium W.Becker (3–4) Mediterranean region | Viola (plant) | Wikipedia | 406 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Species
The genus includes dog violets, a group of scentless species which are the most common Viola in many areas, sweet violet (Viola odorata) (named from its sweet scent), and many other species whose common name includes the word "violet". But not other "violets": Neither Streptocarpus sect. Saintpaulia ("African violets", Gesneriaceae) nor Erythronium dens-canis ("dogtooth violets", Liliaceae) are related to Viola.
List of selected species
Section Danxiaviola
Viola hybanthoides
Section Delphiniopsis
Viola cazorlensis
Viola delphinantha
Viola kosaninii
Section Erpetion
Viola banksii – Australian native violet, ivy-leaved violet
Viola hederacea – Australian native violet, ivy-leaved violet
Section Leptidium
Viola stipularis
Section Melanium (pansies)
Viola arvensis – field pansy
Viola bicolor
Viola pedunculata – yellow pansy, Pacific coast.
Viola bertolonii
Viola calcarata
Viola cheiranthifolia – Teide violet
Viola cornuta
Viola lutea
Viola tricolor – wild pansy, heartsease
Section Nosphinium
Viola pedata
Section A (V. abyssinica group)
Viola abyssinica
Section B (V. spathulata group)
Viola spathulata
Section Plagiostigma
Viola epipsila
Section Rubellium
Viola capillaris
Viola portalesia
Viola rubella
Section Sclerosium
Viola cinerea
Section Tridens
Viola tridentata – mountain violet
Section Viola (violets)
Viola canina – heath dog violet
Viola hirta – hairy violet
Viola labradorica – alpine violet
Viola odorata – sweet violet
Viola persicifolia – fen violet
Viola riviniana – common dog violet
Viola rostrata – long-spurred violet
Viola sororia – common blue violet, hooded violet
Section Xylinosium
Viola decumbens
Evolution and biogeography
One fossil seed of †Viola rimosa has been extracted from borehole samples of the Middle Miocene fresh water deposits in Nowy Sacz Basin, West Carpathians, Poland. The genus is thought to have arisen in S America, most likely the Andes. | Viola (plant) | Wikipedia | 468 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Genetics
Habitat fragmentation has been shown to have minimal effect on the genetic diversity and gene flow of the North American woodland violet Viola pubescens. This may be partially attributed to the ability of Viola pubescens to continue to persist within a largely agricultural matrix. This trend of unexpectedly high genetic diversity is also observed in Viola palmensis, a Canary Island endemic known only from a 15 square kilometer range on La palma island. High levels of genetic diversity within these species indicate that these plants are outcrossing, even though many violet species can produce many clonal offspring throughout the year via cleistogamous flowers. Plants that produce copious amounts of clonal seeds from cleistogamous flowers often experience increased levels of inbreeding. These reportedly high rates of outcrossing and genetic diversity indicate that these violets are strong competitors for pollinators during the early spring when they are in bloom and that those pollinators can travel considerable distances between often fragmented populations.
Distribution and habitat
The worldwide northern temperate distribution of the genus distinguishes it from the remaining largely tropical Violaceae genera, restricted to either Old World or New World species, while in the tropics the distribution is primarily in high mountainous areas. Centres of diversity occur mainly in the northern hemisphere, in mountainous regions of eastern Asia, Melanesia, and southern Europe, but also occur in the Andes and the southern Patagonian cone of South America. One of the highest species concentrations is in the former USSR. Australia is home to a number of Viola species, including Viola hederacea, Viola betonicifolia and Viola banksii, first collected by Joseph Banks and Daniel Solander on the Cook voyage to Botany Bay. | Viola (plant) | Wikipedia | 339 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Ecology
Viola species are used as food plants by the larvae of some Lepidoptera species, including the giant leopard moth, large yellow underwing, lesser broad-bordered yellow underwing, high brown fritillary, small pearl-bordered fritillary, pearl-bordered fritillary, regal fritillary, cardinal, and Setaceous Hebrew character. The larvae of many fritilary butterfly species use violets as an obligate host plant, although these butterflies do not always ovaposit directly onto violets. While the ecology of this genera is extremely diverse, violets are mainly pollinated by members within the orders Diptera and Hymenoptera. Showy flowers are produced in early spring, and clonal cleistogamous flowers are produced from late spring until the end of the growing season under favorable conditions. Cleistogamy allows plants to produce offspring year round and have more chances for establishment. This system is especially important in violets, as these plants are often weak competitors for pollination due to their small size.
Many violet species exhibit two modes of seed dispersal. Once seed capsules have matured, seeds are dispelled around the plant through explosive dehiscence. Viola pedata seeds have been reported being dispersed distances of up to 5 meters away from the parent plant. Often, seeds are then further dispersed by ants through a process called myrmecochory. Violets whose seeds are dispersed this way have specialized structures on the exterior of the seeds called elaiosomes. This interaction allows violet seed to germinate and establish in a protected, stable environment.
Many violet seeds exhibit physiological dormancy and require some period of cold stratification to induce germination under ex situ conditions. Rates of germination are often quite poor, especially when seeds are stored for extended periods of time. In North American habitat restoration, native violets are in high demand due to their relationship with the aforementioned fritillary butterflies. | Viola (plant) | Wikipedia | 400 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Violet species occupy a diverse array of habitats, from bogs (Viola lanceolata) to dry hill prairies (V. pedata) to woodland understories (V. labradorica). While many of these species are indicators of high quality habitat, some violets are capable of thriving in a human altered landscape. Two species of zinc violet (V. calaminaria and V. guestphalica) are capable of living in soils severely contaminated with heavy metals. Many violets form relationships with arbuscular mycorrhizal fungi, and in the case of the zinc violets, this allows them to tolerate such highly contaminated soils.
Flowering is often profuse, and may last for much of the spring and summer. Viola are most often spring-blooming with chasmogamous flowers that have well-developed petals pollinated by insects. Many species also produce self-pollinated cleistogamous flowers in summer and autumn that do not open and lack petals. In some species the showy chasmogamous flowers are infertile (e.g.,Viola sororia).
Horticultural uses
The international registration authority for the genus is the American Violet Society, where growers register new Viola cultivars. A coding system is used for cultivar description of ten horticultural divisions, such as Violet (Vt) and Violetta (Vtta). Examples include Viola 'Little David' (Vtta) and Viola 'Königin Charlotte' (Vt).
In this system violets (Vt) are defined as "stoloniferous perennials with small, highly fragrant, self-coloured purple, blue or white flowers in late winter and early spring".
Species and cultivars
Many species, varieties and cultivars are grown in gardens for their ornamental flowers. In horticulture the term pansy is normally used for those multi-colored, large-flowered cultivars which are raised annually or biennially from seed and used extensively in bedding. The terms viola and violet are normally reserved for small-flowered annuals or perennials, including the wild species.
Cultivars of Viola cornuta, Viola cucullata, and Viola odorata, are commonly grown from seed. Other species often grown include Viola labradorica, Viola pedata, and Viola rotundifolia. | Viola (plant) | Wikipedia | 478 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
The modern garden pansy (V. × wittrockiana) is a plant of complex hybrid origin involving at least three species, V. tricolor (wild pansy or heartsease), V. altaica, and V. lutea (mountain pansy). The hybrid horned pansy (V. × williamsii) originates from hybridization involving garden pansy and Viola cornuta.
Bedding plants
In 2005 in the United States, Viola cultivars (including pansies) were one of the top three bedding plant crops and 111 million dollars worth of flats of Viola were produced for the bedding flower market. Pansies and violas used for bedding are generally raised from seed, and F1 hybrid seed strains have been developed which produce compact plants of reasonably consistent flower coloring and appearance. Bedding plants are usually discarded after one growing season.
Perennial cultivars
There are hundreds of perennial viola and violetta cultivars; many of these do not breed true from seed and therefore have to be propagated from cuttings. Violettas can be distinguished from violas by the lack of ray markings on their petals. The following cultivars, of mixed or uncertain parentage, have gained the Royal Horticultural Society's Award of Garden Merit:
'Aspasia'
'Clementina'
'Huntercombe Purple'
'Jackanapes'
'Molly Sanderson'
'Moonlight'
'Nellie Britton'
Other popular examples include:
'Ardross Gem' (viola)
'Blackjack'
'Buttercup' (violetta)
'Columbine' (viola)
'Dawn' (violetta)
'Etain' (viola)
'Irish Molly' (viola)
'Maggie Mott' (viola)
'Martin' (viola)
'Rebecca' (violetta)
'Vita' (viola)
'Zoe' (violetta)
Other uses
Culinary | Viola (plant) | Wikipedia | 380 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
When newly opened, Viola flowers may be used to decorate salads or in stuffings for poultry or fish. Soufflés, cream, and similar desserts can be flavoured with essence of Viola flowers. The young leaves are edible raw or cooked as a mild-tasting leaf vegetable. The flowers and leaves of the cultivar 'Rebecca', one of the Violetta violets, have a distinct vanilla flavor with hints of wintergreen. The pungent perfume of some varieties of V. odorata adds inimitable sweetness to desserts, fruit salads, and teas while the mild pea flavor of V. tricolor combines equally well with sweet or savory foods, like grilled meats and steamed vegetables. The heart-shaped leaves of V. odorata provide a free source of greens throughout a long growing season, while the petals are used for fragrant flavoring in milk puddings and ice cream or in salads and as garnishes.
A candied violet or crystallized violet is a flower, usually of Viola odorata, preserved by a coating of egg white and crystallised sugar. Alternatively, hot syrup is poured over the fresh flower (or the flower is immersed in the syrup) and stirred until the sugar recrystallizes and has dried. This method is still used for rose petals and was applied to orange flowers in the past (when almonds or orange peel are treated this way they are called pralines). Candied violets are still made commercially in Toulouse, France, where they are known as violettes de Toulouse. They are used as decorating cakes or trifles or included in aromatic desserts.
The French are also known for their violet syrup, most commonly made from an extract of violets. In the United States, this French violet syrup is used to make violet scones and marshmallows. Viola essence flavours the liqueurs Creme Yvette, Creme de Violette, and Parfait d'Amour. It is also used in confectionery, such as Parma Violets and C. Howard's Violet candies. | Viola (plant) | Wikipedia | 434 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Medicinal
Many Viola species contain antioxidants called anthocyanins. Fourteen anthocyanins from V. yedoensis and V. prionantha have been identified. Some anthocyanins show strong antioxidant activities. Most violas tested and many other plants of the family Violaceae contain cyclotides, which have a diverse range of in vitro biological activities when isolated from the plant, including uterotonic, anti-HIV, antimicrobial, and insecticidal activities. Viola canescens, a species from India, exhibited in vitro activity against Trypanosoma cruzi.
Viola has been evaluated in different clinical indications in human studies. A double blind clinical trial showed that the adjuvant use of Viola odorata syrup with short-acting β-agonists can improve the cough suppression in children with asthma. In another study intranasal administration of Viola odorata extract oil showed to be effective in patients with insomnia. Topical use of an herbal formulation containing Viola tricolor extract also showed promising effects in patients with mild-to-moderate atopic dermatitis.
Perfume
Viola odorata is used as a source for scents in the perfume industry. Violet is known to have a 'flirty' scent as its fragrance comes and goes. Ionone is present in the flowers, which turns off the ability for humans to smell the fragrant compound for moments at a time.
Cultural associations
Birth
Violet is the traditional birth flower for February in English tradition.
Geographical territories
In the United States, the common blue violet Viola sororia is the state flower of Illinois, Rhode Island, New Jersey and Wisconsin. In Canada, the Viola cucullata is the provincial flower of New Brunswick, adopted in 1936. In the United Kingdom, Viola riviniana is the county flower of Lincolnshire.
Lesbian and bisexual culture
Violets became symbolically associated with romantic love between women. This connection originates from fragments of a poem by Sappho about a lost love, in which she describes her as "Close by my side you put around yourself [many wreaths] of violets and roses." In another poem, Sappho describes her lost love as wearing "violet tiaras, braided rosebuds, dill and crocus twined around" her neck. In 1926, one of the first plays to involve a lesbian relationship, La Prisonnière by Édouard Bourdet, used a bouquet of violets to signify lesbian love.
Tributes | Viola (plant) | Wikipedia | 510 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
Violets, and badges depicting them,
were sold in fund-raising efforts in Australia and New Zealand on and around Violet Day in commemoration of the lost soldiers of World War I. | Viola (plant) | Wikipedia | 36 | 157841 | https://en.wikipedia.org/wiki/Viola%20%28plant%29 | Biology and health sciences | Malpighiales | null |
An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system.
In higher organisms, the eye is a complex optical system that collects light from the surrounding environment, regulates its intensity through a diaphragm, focuses it through an adjustable assembly of lenses to form an image, converts this image into a set of electrical signals, and transmits these signals to the brain through neural pathways that connect the eye via the optic nerve to the visual cortex and other areas of the brain.
Eyes with resolving power have come in ten fundamentally different forms, classified into compound eyes and non-compound eyes. Compound eyes are made up of multiple small visual units, and are common on insects and crustaceans. Non-compound eyes have a single lens and focus light onto the retina to form a single image. This type of eye is common in mammals, including humans.
The simplest eyes are pit eyes. They are eye-spots which may be set into a pit to reduce the angle of light that enters and affects the eye-spot, to allow the organism to deduce the angle of incoming light.
Eyes enable several photo response functions that are independent of vision. In an organism that has more complex eyes, retinal photosensitive ganglion cells send signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect circadian adjustment and to the pretectal area to control the pupillary light reflex.
Overview
Complex eyes distinguish shapes and colours. The visual fields of many organisms, especially predators, involve large areas of binocular vision for depth perception. In other organisms, particularly prey animals, eyes are located to maximise the field of view, such as in rabbits and horses, which have monocular vision. | Eye | Wikipedia | 384 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
The first proto-eyes evolved among animals about the time of the Cambrian explosion. The last common ancestor of animals possessed the biochemical toolkit necessary for vision, and more advanced eyes have evolved in 96% of animal species in six of the ~35 main phyla. In most vertebrates and some molluscs, the eye allows light to enter and project onto a light-sensitive layer of cells known as the retina. The cone cells (for colour) and the rod cells (for low-light contrasts) in the retina detect and convert light into neural signals which are transmitted to the brain via the optic nerve to produce vision. Such eyes are typically spheroid, filled with the transparent gel-like vitreous humour, possess a focusing lens, and often an iris. Muscles around the iris change the size of the pupil, regulating the amount of light that enters the eye and reducing aberrations when there is enough light. The eyes of most cephalopods, fish, amphibians and snakes have fixed lens shapes, and focusing is achieved by telescoping the lens in a similar manner to that of a camera.
The compound eyes of the arthropods are composed of many simple facets which, depending on anatomical detail, may give either a single pixelated image or multiple images per eye. Each sensor has its own lens and photosensitive cell(s). Some eyes have up to 28,000 such sensors arranged hexagonally, which can give a full 360° field of vision. Compound eyes are very sensitive to motion. Some arthropods, including many Strepsiptera, have compound eyes of only a few facets, each with a retina capable of creating an image. With each eye producing a different image, a fused, high-resolution image is produced in the brain.
The mantis shrimp has the world's most complex colour vision system. It has detailed hyperspectral colour vision.
Trilobites, now extinct, had unique compound eyes. Clear calcite crystals formed the lenses of their eyes. They differ in this from most other arthropods, which have soft eyes. The number of lenses in such an eye varied widely; some trilobites had only one while others had thousands of lenses per eye. | Eye | Wikipedia | 470 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
In contrast to compound eyes, simple eyes have a single lens. Jumping spiders have one pair of large simple eyes with a narrow field of view, augmented by an array of smaller eyes for peripheral vision. Some insect larvae, like caterpillars, have a type of simple eye (stemmata) which usually provides only a rough image, but (as in sawfly larvae) can possess resolving powers of 4 degrees of arc, be polarization-sensitive, and capable of increasing its absolute sensitivity at night by a factor of 1,000 or more. Ocelli, some of the simplest eyes, are found in animals such as some of the snails. They have photosensitive cells but no lens or other means of projecting an image onto those cells. They can distinguish between light and dark but no more, enabling them to avoid direct sunlight. In organisms dwelling near deep-sea vents, compound eyes are adapted to see the infra-red light produced by the hot vents, allowing the creatures to avoid being boiled alive.
Types
There are ten different eye layouts. Eye types can be categorised into "simple eyes", with one concave photoreceptive surface, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface. "Simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behaviour or environment. The only limitations specific to eye types are that of resolution—the physics of compound eyes prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, so are better suited to dark-dwelling creatures.
Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being ciliated (as in the vertebrates) or rhabdomeric. These two groups are not monophyletic; the Cnidaria also possess ciliated cells, and some gastropods and annelids possess both.
Some organisms have photosensitive cells that do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms. These are not considered eyes because they lack enough structure to be considered an organ, and do not produce an image.
Every technological method of capturing an optical image that humans commonly use occurs in nature, with the exception of zoom and Fresnel lenses. | Eye | Wikipedia | 503 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Non-compound eyes
Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in vertebrates, cephalopods, annelids, crustaceans and Cubozoa.
Pit eyes
Pit eyes, also known as stemmata, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eye-spot, to allow the organism to deduce the angle of incoming light. Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eyes". They are small, comprising up to about 100 cells covering about 100 μm. The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material.
Pit vipers have developed pits that function as eyes by sensing thermal infra-red radiation, in addition to their optical wavelength eyes like those of other vertebrates (see infrared sensing in snakes). However, pit organs are fitted with receptors rather different from photoreceptors, namely a specific transient receptor potential channel (TRP channels) called TRPV1. The main difference is that photoreceptors are G-protein coupled receptors but TRP are ion channels.
Spherical lens eye
The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered—hence increasing the resolution obtainable. The most basic form, seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges; this decreases the focal length and thus allows a sharp image to form on the retina. This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration. Such a non-homogeneous lens is necessary for the focal length to drop from about 4 times the lens radius, to 2.5 radii.
So-called under-focused lens eyes, found in gastropods and polychaete worms, have eyes that are intermediate between lens-less cup eyes and real camera eyes. Also box jellyfish have eyes with a spherical lens, cornea and retina, but the vision is blurry. | Eye | Wikipedia | 503 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Heterogeneous eyes have evolved at least nine times: four or more times in gastropods, once in the copepods, once in the annelids, once in the cephalopods, and once in the chitons, which have aragonite lenses. No extant aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".
This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimise the effect of eye motion while the animal moves, most such eyes have stabilising eye muscles.
The ocelli of insects bear a simple lens, but their focal point usually lies behind the retina; consequently, those can not form a sharp image. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field; this fast response is further accelerated by the large nerve bundles which rush the information to the brain. Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity. This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).
Multiple lenses
Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, Copilia, has two lenses in each eye, arranged like those in a telescope. Such arrangements are rare and poorly understood, but represent an alternative construction.
Multiple lenses are seen in some hunters such as eagles and jumping spiders, which have a refractive cornea: these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.
Refractive cornea | Eye | Wikipedia | 456 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
In the eyes of most mammals, birds, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air. In general, the lens is not spherical. Spherical lenses produce spherical aberration. In refractive corneas, the lens tissue is corrected with inhomogeneous lens material (see Luneburg lens), or with an aspheric shape. Flattening the lens has a disadvantage; the quality of vision is diminished away from the main line of focus. Thus, animals that have evolved with a wide field-of-view often have eyes that make use of an inhomogeneous lens.
As mentioned above, a refractive cornea is only useful out of water. In water, there is little difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures that have returned to the water—penguins and seals, for example—lose their highly curved cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strongly focusing cornea.
A unique feature of most mammal eyes is the presence of eyelids which wipe the eye and spread tears across the cornea to prevent dehydration. These eyelids are also supplemented by the presence of eyelashes, multiple rows of highly innervated and sensitive hairs which grow from the eyelid margins to protect the eye from fine particles and small irritants such as insects.
Reflector eyes
An alternative to a lens is to line the inside of the eye with "mirrors", and reflect the image to focus at a central point. The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.
Many small organisms such as rotifers, copepods and flatworms use such organs, but these are too small to produce usable images. Some larger organisms, such as scallops, also use reflector eyes. The scallop Pecten has up to 100 millimetre-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses. | Eye | Wikipedia | 456 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of guanine crystals.
Compound eyes
A compound eye may consist of thousands of individual photoreceptor units or ommatidia (ommatidium, singular). The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarisation of light. Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained (assuming that they do not function as phased arrays). This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require very large compound eyes, around in radius.
Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes are common in arthropods, annelids and some bivalved molluscs. Compound eyes in arthropods grow at their margins by the addition of new ommatidia.
Apposition eyes
Apposition eyes are the most common form of eyes and are presumably the ancestral form of compound eyes. They are found in all arthropod groups, although they may have evolved more than once within this phylum. Some annelids and bivalves also have apposition eyes. They are also possessed by Limulus, the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point. (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.)
Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information. The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ommatidium. | Eye | Wikipedia | 504 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Superposition eyes
The second type is named the superposition eye. The superposition eye is divided into three types:
refracting,
reflecting and
parabolic superposition
The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This type of compound eye, for which a minimal size exists below which effective superposition cannot occur, is normally found in nocturnal insects, because it can create images up to 1000 times brighter than equivalent apposition eyes, though at the cost of reduced resolution. In the parabolic superposition compound eye type, seen in arthropods such as mayflies, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also have a transparent gap but use corner mirrors instead of lenses.
Parabolic superposition
This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.
Other
Another kind of compound eye, found in males of Order Strepsiptera, employs a series of simple eyes—eyes having one opening that provides light for an entire image-forming retina. Several of these eyelets together form the strepsipteran compound eye, which is similar to the 'schizochroal' compound eyes of some trilobites. Because each eyelet is a simple eye, it produces an inverted image; those images are combined in the brain to form one unified image. Because the aperture of an eyelet is larger than the facets of a compound eye, this arrangement allows vision under low light levels. | Eye | Wikipedia | 416 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Good fliers such as flies or honey bees, or prey-catching insects such as praying mantis or dragonflies, have specialised zones of ommatidia organised into a fovea area which gives acute vision. In the acute zone, the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution. The black spot that can be seen on the compound eyes of such insects, which always seems to look directly at the observer, is called a pseudopupil. This occurs because the ommatidia which one observes "head-on" (along their optical axes) absorb the incident light, while those to one side reflect it.
There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid shrimp, Dioptromysis paucispinosa. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialised retina. The resulting eye is a mixture of a simple eye within a compound eye.
Another version is a compound eye often referred to as "pseudofaceted", as seen in Scutigera. This type of eye consists of a cluster of numerous ommatidia on each side of the head, organised in a way that resembles a true compound eye.
The body of Ophiocoma wendtii, a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons. The tube feet of sea urchins contain photoreceptor proteins, which together act as a compound eye; they lack screening pigments, but can detect the directionality of light by the shadow cast by its opaque body. | Eye | Wikipedia | 435 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Nutrients
The ciliary body is triangular in horizontal section and is coated by a double layer, the ciliary epithelium. The inner layer is transparent and covers the vitreous body, and is continuous from the neural tissue of the retina. The outer layer is highly pigmented, continuous with the retinal pigment epithelium, and constitutes the cells of the dilator muscle.
The vitreous is the transparent, colourless, gelatinous mass that fills the space between the lens of the eye and the retina lining the back of the eye. It is produced by certain retinal cells. It is of rather similar composition to the cornea, but contains very few cells (mostly phagocytes which remove unwanted cellular debris in the visual field, as well as the hyalocytes of Balazs of the surface of the vitreous, which reprocess the hyaluronic acid), no blood vessels, and 98–99% of its volume is water (as opposed to 75% in the cornea) with salts, sugars, vitrosin (a type of collagen), a network of collagen type II fibres with the mucopolysaccharide hyaluronic acid, and also a wide array of proteins in micro amounts. Amazingly, with so little solid matter, it tautly holds the eye.
Evolution
Photoreception is phylogenetically very old, with various theories of phylogenesis. The common origin (monophyly) of all animal eyes is now widely accepted as fact. This is based upon the shared genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye believed to have evolved some 650-600 million years ago, and the PAX6 gene is considered a key factor in this. The majority of the advancements in early eyes are believed to have taken only a few million years to develop, since the first predator to gain true imaging would have touched off an "arms race" among all species that did not flee the photopic environment. Prey animals and competing predators alike would be at a distinct disadvantage without such capabilities and would be less likely to survive and reproduce. Hence multiple eye types and subtypes developed in parallel (except those of groups, such as the vertebrates, that were only forced into the photopic environment at a late stage). | Eye | Wikipedia | 497 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Eyes in various animals show adaptation to their requirements. For example, the eye of a bird of prey has much greater visual acuity than a human eye, and in some cases can detect ultraviolet radiation. The different forms of eye in, for example, vertebrates and molluscs are examples of parallel evolution, despite their distant common ancestry. Phenotypic convergence of the geometry of cephalopod and most vertebrate eyes creates the impression that the vertebrate eye evolved from an imaging cephalopod eye, but this is not the case, as the reversed roles of their respective ciliary and rhabdomeric opsin classes and different lens crystallins show.
The very earliest "eyes", called eye-spots, were simple patches of photoreceptor protein in unicellular animals. In multicellular beings, multicellular eyespots evolved, physically similar to the receptor patches for taste and smell. These eyespots could only sense ambient brightness: they could distinguish light and dark, but not the direction of the light source.
Through gradual change, the eye-spots of species living in well-lit environments depressed into a shallow "cup" shape. The ability to slightly discriminate directional brightness was achieved by using the angle at which the light hit certain cells to identify the source. The pit deepened over time, the opening diminished in size, and the number of photoreceptor cells increased, forming an effective pinhole camera that was capable of dimly distinguishing shapes. However, the ancestors of modern hagfish, thought to be the protovertebrate, were evidently pushed to very deep, dark waters, where they were less vulnerable to sighted predators, and where it is advantageous to have a convex eye-spot, which gathers more light than a flat or concave one. This would have led to a somewhat different evolutionary trajectory for the vertebrate eye than for other animal eyes.
The thin overgrowth of transparent cells over the eye's aperture, originally formed to prevent damage to the eyespot, allowed the segregated contents of the eye chamber to specialise into a transparent humour that optimised colour filtering, blocked harmful radiation, improved the eye's refractive index, and allowed functionality outside of water. The transparent protective cells eventually split into two layers, with circulatory fluid in between that allowed wider viewing angles and greater imaging resolution, and the thickness of the transparent layer gradually increased, in most species with the transparent crystallin protein. | Eye | Wikipedia | 506 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
The gap between tissue layers naturally formed a biconvex shape, an optimally ideal structure for a normal refractive index. Independently, a transparent layer and a nontransparent layer split forward from the lens: the cornea and iris. Separation of the forward layer again formed a humour, the aqueous humour. This increased refractive power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood vessels, more circulation, and larger eye sizes.
Relationship to life requirements
Eyes are generally adapted to the environment and life requirements of the organism which bears them. For instance, the distribution of photoreceptors tends to match the area in which the highest acuity is required, with horizon-scanning organisms, such as those that live on the African plains, having a horizontal line of high-density ganglia, while tree-dwelling creatures which require good all-round vision tend to have a symmetrical distribution of ganglia, with acuity decreasing outwards from the centre.
Of course, for most eye types, it is impossible to diverge from a spherical form, so only the density of optical receptors can be altered. In organisms with compound eyes, it is the number of ommatidia rather than ganglia that reflects the region of highest data acquisition. Optical superposition eyes are constrained to a spherical shape, but other forms of compound eyes may deform to a shape where more ommatidia are aligned to, say, the horizon, without altering the size or density of individual ommatidia. Eyes of horizon-scanning organisms have stalks so they can be easily aligned to the horizon when this is inclined, for example, if the animal is on a slope. | Eye | Wikipedia | 348 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
An extension of this concept is that the eyes of predators typically have a zone of very acute vision at their centre, to assist in the identification of prey. In deep water organisms, it may not be the centre of the eye that is enlarged. The hyperiid amphipods are deep water animals that feed on organisms above them. Their eyes are almost divided into two, with the upper region thought to be involved in detecting the silhouettes of potential prey—or predators—against the faint light of the sky above. Accordingly, deeper water hyperiids, where the light against which the silhouettes must be compared is dimmer, have larger "upper-eyes", and may lose the lower portion of their eyes altogether. In the giant Antarctic isopod Glyptonotus a small ventral compound eye is physically completely separated from the much larger dorsal compound eye. Depth perception can be enhanced by having eyes which are enlarged in one direction; distorting the eye slightly allows the distance to the object to be estimated with a high degree of accuracy.
Acuity is higher among male organisms that mate in mid-air, as they need to be able to spot and assess potential mates against a very large backdrop. On the other hand, the eyes of organisms which operate in low light levels, such as around dawn and dusk or in deep water, tend to be larger to increase the amount of light that can be captured.
It is not only the shape of the eye that may be affected by lifestyle. Eyes can be the most visible parts of organisms, and this can act as a pressure on organisms to have more transparent eyes at the cost of function.
Eyes may be mounted on stalks to provide better all-round vision, by lifting them above an organism's carapace; this also allows them to track predators or prey without moving the head.
Physiology
Visual acuity | Eye | Wikipedia | 374 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Visual acuity, or resolving power, is "the ability to distinguish fine detail" and is the property of cone cells. It is often measured in cycles per degree (CPD), which measures an angular resolution, or how much an eye can differentiate one object from another in terms of visual angles. Resolution in CPD can be measured by bar charts of different numbers of white/black stripe cycles. For example, if each pattern is 1.75 cm wide and is placed at 1 m distance from the eye, it will subtend an angle of 1 degree, so the number of white/black bar pairs on the pattern will be a measure of the cycles per degree of that pattern. The highest such number that the eye can resolve as stripes, or distinguish from a grey block, is then the measurement of visual acuity of the eye.
For a human eye with excellent acuity, the maximum theoretical resolution is 50 CPD (1.2 arcminute per line pair, or a 0.35 mm line pair, at 1 m). A rat can resolve only about 1 to 2 CPD. A horse has higher acuity through most of the visual field of its eyes than a human has, but does not match the high acuity of the human eye's central fovea region.
Spherical aberration limits the resolution of a 7 mm pupil to about 3 arcminutes per line pair. At a pupil diameter of 3 mm, the spherical aberration is greatly reduced, resulting in an improved resolution of approximately 1.7 arcminutes per line pair. A resolution of 2 arcminutes per line pair, equivalent to a 1 arcminute gap in an optotype, corresponds to 20/20 (normal vision) in humans.
However, in the compound eye, the resolution is related to the size of individual ommatidia and the distance between neighbouring ommatidia. Physically these cannot be reduced in size to achieve the acuity seen with single lensed eyes as in mammals. Compound eyes have a much lower acuity than vertebrate eyes.
Colour perception | Eye | Wikipedia | 426 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
"Colour vision is the faculty of the organism to distinguish lights of different spectral qualities." All organisms are restricted to a small range of electromagnetic spectrum; this varies from creature to creature, but is mainly between wavelengths of 400 and 700 nm. This is a rather small section of the electromagnetic spectrum, probably reflecting the submarine evolution of the organ: water blocks out all but two small windows of the EM spectrum, and there has been no evolutionary pressure among land animals to broaden this range.
The most sensitive pigment, rhodopsin, has a peak response at 500 nm. Small changes to the genes coding for this protein can tweak the peak response by a few nm; pigments in the lens can also filter incoming light, changing the peak response. Many organisms are unable to discriminate between colours, seeing instead in shades of grey; colour vision necessitates a range of pigment cells which are primarily sensitive to smaller ranges of the spectrum. In primates, geckos, and other organisms, these take the form of cone cells, from which the more sensitive rod cells evolved. Even if organisms are physically capable of discriminating different colours, this does not necessarily mean that they can perceive the different colours; only with behavioural tests can this be deduced.
Most organisms with colour vision can detect ultraviolet light. This high energy light can be damaging to receptor cells. With a few exceptions (snakes, placental mammals), most organisms avoid these effects by having absorbent oil droplets around their cone cells. The alternative, developed by organisms that had lost these oil droplets in the course of evolution, is to make the lens impervious to UV light—this precludes the possibility of any UV light being detected, as it does not even reach the retina.
Rods and cones
The retina contains two major types of light-sensitive photoreceptor cells used for vision: the rods and the cones.
Rods cannot distinguish colours, but are responsible for low-light (scotopic) monochrome (black-and-white) vision; they work well in dim light as they contain a pigment, rhodopsin (visual purple), which is sensitive at low light intensity, but saturates at higher (photopic) intensities. Rods are distributed throughout the retina but there are none at the fovea and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina. | Eye | Wikipedia | 502 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Cones are responsible for colour vision. They require brighter light to function than rods require. In humans, there are three types of cones, maximally sensitive to long-wavelength, medium-wavelength, and short-wavelength light (often referred to as red, green, and blue, respectively, though the sensitivity peaks are not actually at these colours). The colour seen is the combined effect of stimuli to, and responses from, these three types of cone cells. Cones are mostly concentrated in and near the fovea. Only a few are present at the sides of the retina. Objects are seen most sharply in focus when their images fall on the fovea, as when one looks at an object directly. Cone cells and rods are connected through intermediate cells in the retina to nerve fibres of the optic nerve. When rods and cones are stimulated by light, they connect through adjoining cells within the retina to send an electrical signal to the optic nerve fibres. The optic nerves send off impulses through these fibres to the brain.
Pigmentation
The pigment molecules used in the eye are various, but can be used to define the evolutionary distance between different groups, and can also be an aid in determining which are closely related—although problems of convergence do exist.
Opsins are the pigments involved in photoreception. Other pigments, such as melanin, are used to shield the photoreceptor cells from light leaking in from the sides. The opsin protein group evolved long before the last common ancestor of animals, and has continued to diversify since. | Eye | Wikipedia | 319 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
There are two types of opsin involved in vision; c-opsins, which are associated with ciliary-type photoreceptor cells, and r-opsins, associated with rhabdomeric photoreceptor cells. The eyes of vertebrates usually contain ciliary cells with c-opsins, and (bilaterian) invertebrates have rhabdomeric cells in the eye with r-opsins. However, some ganglion cells of vertebrates express r-opsins, suggesting that their ancestors used this pigment in vision, and that remnants survive in the eyes. Likewise, c-opsins have been found to be expressed in the brain of some invertebrates. They may have been expressed in ciliary cells of larval eyes, which were subsequently resorbed into the brain on metamorphosis to the adult form. C-opsins are also found in some derived bilaterian-invertebrate eyes, such as the pallial eyes of the bivalve molluscs; however, the lateral eyes (which were presumably the ancestral type for this group, if eyes evolved once there) always use r-opsins. Cnidaria, which are an outgroup to the taxa mentioned above, express c-opsins—but r-opsins are yet to be found in this group. Incidentally, the melanin produced in the cnidaria is produced in the same fashion as that in vertebrates, suggesting the common descent of this pigment.
Additional images | Eye | Wikipedia | 314 | 157898 | https://en.wikipedia.org/wiki/Eye | Biology and health sciences | Biology | null |
Genetic recombination (also known as genetic reshuffling) is the exchange of genetic material between different organisms which leads to production of offspring with combinations of traits that differ from those found in either parent. In eukaryotes, genetic recombination during meiosis can lead to a novel set of genetic information that can be further passed on from parents to offspring. Most recombination occurs naturally and can be classified into two types: (1) interchromosomal recombination, occurring through independent assortment of alleles whose loci are on different but homologous chromosomes (random orientation of pairs of homologous chromosomes in meiosis I); & (2) intrachromosomal recombination, occurring through crossing over.
During meiosis in eukaryotes, genetic recombination involves the pairing of homologous chromosomes. This may be followed by information transfer between the chromosomes. The information transfer may occur without physical exchange (a section of genetic material is copied from one chromosome to another, without the donating chromosome being changed) (see SDSA – Synthesis Dependent Strand Annealing pathway in Figure); or by the breaking and rejoining of DNA strands, which forms new molecules of DNA (see DHJ pathway in Figure).
Recombination may also occur during mitosis in eukaryotes where it ordinarily involves the two sister chromosomes formed after chromosomal replication. In this case, new combinations of alleles are not produced since the sister chromosomes are usually identical. In meiosis and mitosis, recombination occurs between similar molecules of DNA (homologous sequences). In meiosis, non-sister homologous chromosomes pair with each other so that recombination characteristically occurs between non-sister homologues. In both meiotic and mitotic cells, recombination between homologous chromosomes is a common mechanism used in DNA repair.
Gene conversion – the process during which homologous sequences are made identical also falls under genetic recombination.
Genetic recombination and recombinational DNA repair also occurs in bacteria and archaea, which use asexual reproduction.
Recombination can be artificially induced in laboratory (in vitro) settings, producing recombinant DNA for purposes including vaccine development. | Genetic recombination | Wikipedia | 476 | 158005 | https://en.wikipedia.org/wiki/Genetic%20recombination | Biology and health sciences | Genetics | Biology |
V(D)J recombination in organisms with an adaptive immune system is a type of site-specific genetic recombination that helps immune cells rapidly diversify to recognize and adapt to new pathogens.
Synapsis
During meiosis, synapsis (the pairing of homologous chromosomes) ordinarily precedes genetic recombination.
Mechanism
Genetic recombination is catalyzed by many different enzymes. Recombinases are key enzymes that catalyse the strand transfer step during recombination. RecA, the chief recombinase found in Escherichia coli, is responsible for the repair of DNA double strand breaks (DSBs). In yeast and other eukaryotic organisms there are two recombinases required for repairing DSBs. The RAD51 protein is required for mitotic and meiotic recombination, whereas the DNA repair protein, DMC1, is specific to meiotic recombination. In the archaea, the ortholog of the bacterial RecA protein is RadA.
Bacterial recombination
Bacteria regularly undergo genetic recombination in three main ways:
Transformation, the uptake of exogenous DNA from the surrounding environment.
Transduction, the virus-mediated transfer of DNA between bacteria.
Conjugation, the transfer of DNA from one bacterium to another via cell-to-cell contact.
Sometimes a strand of DNA is transferred into the target cell but fails to be copied as the target divides. This is called an abortive transfer.
Chromosomal crossover
In eukaryotes, recombination during meiosis is facilitated by chromosomal crossover. The crossover process leads to offspring having different combinations of genes from those of their parents, and can occasionally produce new chimeric alleles. The shuffling of genes brought about by genetic recombination produces increased genetic variation. It also allows sexually reproducing organisms to avoid Muller's ratchet, in which the genomes of an asexual population tend to accumulate more deleterious mutations over time than beneficial or reversing mutations. | Genetic recombination | Wikipedia | 435 | 158005 | https://en.wikipedia.org/wiki/Genetic%20recombination | Biology and health sciences | Genetics | Biology |
Chromosomal crossover involves recombination between the paired chromosomes inherited from each of one's parents, generally occurring during meiosis. During prophase I (pachytene stage) the four available chromatids are in tight formation with one another. While in this formation, homologous sites on two chromatids can closely pair with one another, and may exchange genetic information.
Because there is a small probability of recombination at any location along a chromosome, the frequency of recombination between two locations depends on the distance separating them. Therefore, for genes sufficiently distant on the same chromosome, the amount of crossover is high enough to destroy the correlation between alleles.
Tracking the movement of genes resulting from crossovers has proven quite useful to geneticists. Because two genes that are close together are less likely to become separated than genes that are farther apart, geneticists can deduce roughly how far apart two genes are on a chromosome if they know the frequency of the crossovers. Geneticists can also use this method to infer the presence of certain genes. Genes that typically stay together during recombination are said to be linked. One gene in a linked pair can sometimes be used as a marker to deduce the presence of the other gene. This is typically used to detect the presence of a disease-causing gene.
The recombination frequency between two loci observed is the crossing-over value. It is the frequency of crossing over between two linked gene loci (markers), and depends on the distance between the genetic loci observed. For any fixed set of genetic and environmental conditions, recombination in a particular region of a linkage structure (chromosome) tends to be constant, and the same is then true for the crossing-over value which is used in the production of genetic maps.
Gene conversion | Genetic recombination | Wikipedia | 380 | 158005 | https://en.wikipedia.org/wiki/Genetic%20recombination | Biology and health sciences | Genetics | Biology |
In gene conversion, a section of genetic material is copied from one chromosome to another, without the donating chromosome being changed. Gene conversion occurs at high frequency at the actual site of the recombination event during meiosis. It is a process by which a DNA sequence is copied from one DNA helix (which remains unchanged) to another DNA helix, whose sequence is altered. Gene conversion has often been studied in fungal crosses where the 4 products of individual meioses can be conveniently observed. Gene conversion events can be distinguished as deviations in an individual meiosis from the normal 2:2 segregation pattern (e.g. a 3:1 pattern).
Nonhomologous recombination
Recombination can occur between DNA sequences that contain no sequence homology. This can cause chromosomal translocations, sometimes leading to cancer.
In B cells
B cells of the immune system perform genetic recombination, called immunoglobulin class switching. It is a biological mechanism that changes an antibody from one class to another, for example, from an isotype called IgM to an isotype called IgG.
Genetic engineering
In genetic engineering, recombination can also refer to artificial and deliberate recombination of disparate pieces of DNA, often from different organisms, creating what is called recombinant DNA. A prime example of such a use of genetic recombination is gene targeting, which can be used to add, delete or otherwise change an organism's genes. This technique is important to biomedical researchers as it allows them to study the effects of specific genes. Techniques based on genetic recombination are also applied in protein engineering to develop new proteins of biological interest.
Examples include Restriction enzyme mediated integration, Gibson assembly and Golden Gate Cloning. | Genetic recombination | Wikipedia | 368 | 158005 | https://en.wikipedia.org/wiki/Genetic%20recombination | Biology and health sciences | Genetics | Biology |
Recombinational repair
DNA damages caused by a variety of exogenous agents (e.g. UV light, X-rays, chemical cross-linking agents) can be repaired by homologous recombinational repair (HRR). These findings suggest that DNA damages arising from natural processes, such as exposure to reactive oxygen species that are byproducts of normal metabolism, are also repaired by HRR. In humans, deficiencies in the gene products necessary for HRR during meiosis likely cause infertility In humans, deficiencies in gene products necessary for HRR, such as BRCA1 and BRCA2, increase the risk of cancer (see DNA repair-deficiency disorder).
In bacteria, transformation is a process of gene transfer that ordinarily occurs between individual cells of the same bacterial species. Transformation involves integration of donor DNA into the recipient chromosome by recombination. This process appears to be an adaptation for repairing DNA damages in the recipient chromosome by HRR. Transformation may provide a benefit to pathogenic bacteria by allowing repair of DNA damage, particularly damages that occur in the inflammatory, oxidizing environment associated with infection of a host.
When two or more viruses, each containing lethal genomic damages, infect the same host cell, the virus genomes can often pair with each other and undergo HRR to produce viable progeny. This process, referred to as multiplicity reactivation, has been studied in lambda and T4 bacteriophages, as well as in several pathogenic viruses. In the case of pathogenic viruses, multiplicity reactivation may be an adaptive benefit to the virus since it allows the repair of DNA damages caused by exposure to the oxidizing environment produced during host infection. | Genetic recombination | Wikipedia | 356 | 158005 | https://en.wikipedia.org/wiki/Genetic%20recombination | Biology and health sciences | Genetics | Biology |
The lipid bilayer (or phospholipid bilayer) is a thin polar membrane made of two layers of lipid molecules. These membranes form a continuous barrier around all cells. The cell membranes of almost all organisms and many viruses are made of a lipid bilayer, as are the nuclear membrane surrounding the cell nucleus, and membranes of the membrane-bound organelles in the cell. The lipid bilayer is the barrier that keeps ions, proteins and other molecules where they are needed and prevents them from diffusing into areas where they should not be. Lipid bilayers are ideally suited to this role, even though they are only a few nanometers in width, because they are impermeable to most water-soluble (hydrophilic) molecules. Bilayers are particularly impermeable to ions, which allows cells to regulate salt concentrations and pH by transporting ions across their membranes using proteins called ion pumps.
Biological bilayers are usually composed of amphiphilic phospholipids that have a hydrophilic phosphate head and a hydrophobic tail consisting of two fatty acid chains. Phospholipids with certain head groups can alter the surface chemistry of a bilayer and can, for example, serve as signals as well as "anchors" for other molecules in the membranes of cells. Just like the heads, the tails of lipids can also affect membrane properties, for instance by determining the phase of the bilayer. The bilayer can adopt a solid gel phase state at lower temperatures but undergo phase transition to a fluid state at higher temperatures, and the chemical properties of the lipids' tails influence at which temperature this happens. The packing of lipids within the bilayer also affects its mechanical properties, including its resistance to stretching and bending. Many of these properties have been studied with the use of artificial "model" bilayers produced in a lab. Vesicles made by model bilayers have also been used clinically to deliver drugs. | Lipid bilayer | Wikipedia | 408 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
The structure of biological membranes typically includes several types of molecules in addition to the phospholipids comprising the bilayer. A particularly important example in animal cells is cholesterol, which helps strengthen the bilayer and decrease its permeability. Cholesterol also helps regulate the activity of certain integral membrane proteins. Integral membrane proteins function when incorporated into a lipid bilayer, and they are held tightly to the lipid bilayer with the help of an annular lipid shell. Because bilayers define the boundaries of the cell and its compartments, these membrane proteins are involved in many intra- and inter-cellular signaling processes. Certain kinds of membrane proteins are involved in the process of fusing two bilayers together. This fusion allows the joining of two distinct structures as in the acrosome reaction during fertilization of an egg by a sperm, or the entry of a virus into a cell. Because lipid bilayers are fragile and invisible in a traditional microscope, they are a challenge to study. Experiments on bilayers often require advanced techniques like electron microscopy and atomic force microscopy.
Structure and organization
When phospholipids are exposed to water, they self-assemble into a two-layered sheet with the hydrophobic tails pointing toward the center of the sheet. This arrangement results in two 'leaflets' that are each a single molecular layer. The center of this bilayer contains almost no water and excludes molecules like sugars or salts that dissolve in water. The assembly process and maintenance are driven by aggregation of hydrophobic molecules (also called the hydrophobic effect). This complex process includes non-covalent interactions such as van der Waals forces, electrostatic and hydrogen bonds.
Cross-section analysis
The lipid bilayer is very thin compared to its lateral dimensions. If a typical mammalian cell (diameter ~10 micrometers) were magnified to the size of a watermelon (~1 ft/30 cm), the lipid bilayer making up the plasma membrane would be about as thick as a piece of office paper. Despite being only a few nanometers thick, the bilayer is composed of several distinct chemical regions across its cross-section. These regions and their interactions with the surrounding water have been characterized over the past several decades with x-ray reflectometry, neutron scattering, and nuclear magnetic resonance techniques. | Lipid bilayer | Wikipedia | 481 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
The first region on either side of the bilayer is the hydrophilic headgroup. This portion of the membrane is completely hydrated and is typically around 0.8-0.9 nm thick. In phospholipid bilayers the phosphate group is located within this hydrated region, approximately 0.5 nm outside the hydrophobic core. In some cases, the hydrated region can extend much further, for instance in lipids with a large protein or long sugar chain grafted to the head. One common example of such a modification in nature is the lipopolysaccharide coat on a bacterial outer membrane.
Next to the hydrated region is an intermediate region that is only partially hydrated. This boundary layer is approximately 0.3 nm thick. Within this short distance, the water concentration drops from 2M on the headgroup side to nearly zero on the tail (core) side. The hydrophobic core of the bilayer is typically 3-4 nm thick, but this value varies with chain length and chemistry. Core thickness also varies significantly with temperature, in particular near a phase transition.
Asymmetry
In many naturally occurring bilayers, the compositions of the inner and outer membrane leaflets are different. In human red blood cells, the inner (cytoplasmic) leaflet is composed mostly of phosphatidylethanolamine, phosphatidylserine and phosphatidylinositol and its phosphorylated derivatives. By contrast, the outer (extracellular) leaflet is based on phosphatidylcholine, sphingomyelin and a variety of glycolipids. In some cases, this asymmetry is based on where the lipids are made in the cell and reflects their initial orientation. The biological functions of lipid asymmetry are imperfectly understood, although it is clear that it is used in several different situations. For example, when a cell undergoes apoptosis, the phosphatidylserine — normally localised to the cytoplasmic leaflet — is transferred to the outer surface: There, it is recognised by a macrophage that then actively scavenges the dying cell. | Lipid bilayer | Wikipedia | 454 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Lipid asymmetry arises, at least in part, from the fact that most phospholipids are synthesised and initially inserted into the inner monolayer: those that constitute the outer monolayer are then transported from the inner monolayer by a class of enzymes called flippases. Other lipids, such as sphingomyelin, appear to be synthesised at the external leaflet. Flippases are members of a larger family of lipid transport molecules that also includes floppases, which transfer lipids in the opposite direction, and scramblases, which randomize lipid distribution across lipid bilayers (as in apoptotic cells). In any case, once lipid asymmetry is established, it does not normally dissipate quickly because spontaneous flip-flop of lipids between leaflets is extremely slow.
It is possible to mimic this asymmetry in the laboratory in model bilayer systems. Certain types of very small artificial vesicle will automatically make themselves slightly asymmetric, although the mechanism by which this asymmetry is generated is very different from that in cells. By utilizing two different monolayers in Langmuir-Blodgett deposition or a combination of Langmuir-Blodgett and vesicle rupture deposition it is also possible to synthesize an asymmetric planar bilayer. This asymmetry may be lost over time as lipids in supported bilayers can be prone to flip-flop. However, it has been reported that lipid flip-flop is slow compare to cholesterol and other smaller molecules.
It has been reported that the organization and dynamics of the lipid monolayers in a bilayer are coupled. For example, introduction of obstructions in one monolayer can slow down the lateral diffusion in both monolayers. In addition, phase separation in one monolayer can also induce phase separation in other monolayer even when other monolayer can not phase separate by itself.
Phases and phase transitions | Lipid bilayer | Wikipedia | 413 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
At a given temperature a lipid bilayer can exist in either a liquid or a gel (solid) phase. All lipids have a characteristic temperature at which they transition (melt) from the gel to liquid phase. In both phases the lipid molecules are prevented from flip-flopping across the bilayer, but in liquid phase bilayers a given lipid will exchange locations with its neighbor millions of times a second. This random walk exchange allows lipid to diffuse and thus wander across the surface of the membrane.Unlike liquid phase bilayers, the lipids in a gel phase bilayer have less mobility.
The phase behavior of lipid bilayers is determined largely by the strength of the attractive Van der Waals interactions between adjacent lipid molecules. Longer-tailed lipids have more area over which to interact, increasing the strength of this interaction and, as a consequence, decreasing the lipid mobility. Thus, at a given temperature, a short-tailed lipid will be more fluid than an otherwise identical long-tailed lipid. Transition temperature can also be affected by the degree of unsaturation of the lipid tails. An unsaturated double bond can produce a kink in the alkane chain, disrupting the lipid packing. This disruption creates extra free space within the bilayer that allows additional flexibility in the adjacent chains. An example of this effect can be noted in everyday life as butter, which has a large percentage saturated fats, is solid at room temperature while vegetable oil, which is mostly unsaturated, is liquid.
Most natural membranes are a complex mixture of different lipid molecules. If some of the components are liquid at a given temperature while others are in the gel phase, the two phases can coexist in spatially separated regions, rather like an iceberg floating in the ocean. This phase separation plays a critical role in biochemical phenomena because membrane components such as proteins can partition into one or the other phase and thus be locally concentrated or activated. One particularly important component of many mixed phase systems is cholesterol, which modulates bilayer permeability, mechanical strength, and biochemical interactions. | Lipid bilayer | Wikipedia | 438 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Surface chemistry
While lipid tails primarily modulate bilayer phase behavior, it is the headgroup that determines the bilayer surface chemistry. Most natural bilayers are composed primarily of phospholipids, but sphingolipids and sterols such as cholesterol are also important components. Of the phospholipids, the most common headgroup is phosphatidylcholine (PC), accounting for about half the phospholipids in most mammalian cells. PC is a zwitterionic headgroup, as it has a negative charge on the phosphate group and a positive charge on the amine but, because these local charges balance, no net charge.
Other headgroups are also present to varying degrees and can include phosphatidylserine (PS) phosphatidylethanolamine (PE) and phosphatidylglycerol (PG). These alternate headgroups often confer specific biological functionality that is highly context-dependent. For instance, PS presence on the extracellular membrane face of erythrocytes is a marker of cell apoptosis, whereas PS in growth plate vesicles is necessary for the nucleation of hydroxyapatite crystals and subsequent bone mineralization. Unlike PC, some of the other headgroups carry a net charge, which can alter the electrostatic interactions of small molecules with the bilayer.
Biological roles | Lipid bilayer | Wikipedia | 298 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Containment and separation
The primary role of the lipid bilayer in biology is to separate aqueous compartments from their surroundings. Without some form of barrier delineating “self” from “non-self”, it is difficult to even define the concept of an organism or of life. This barrier takes the form of a lipid bilayer in all known life forms except for a few species of archaea that utilize a specially adapted lipid monolayer. It has even been proposed that the very first form of life may have been a simple lipid vesicle with virtually its sole biosynthetic capability being the production of more phospholipids. The partitioning ability of the lipid bilayer is based on the fact that hydrophilic molecules cannot easily cross the hydrophobic bilayer core, as discussed in Transport across the bilayer below. The nucleus, mitochondria and chloroplasts have two lipid bilayers, while other sub-cellular structures are surrounded by a single lipid bilayer (such as the plasma membrane, endoplasmic reticula, Golgi apparatus and lysosomes). See Organelle.
Prokaryotes have only one lipid bilayer - the cell membrane (also known as the plasma membrane). Many prokaryotes also have a cell wall, but the cell wall is composed of proteins or long chain carbohydrates, not lipids. In contrast, eukaryotes have a range of organelles including the nucleus, mitochondria, lysosomes and endoplasmic reticulum. All of these sub-cellular compartments are surrounded by one or more lipid bilayers and, together, typically comprise the majority of the bilayer area present in the cell. In liver hepatocytes for example, the plasma membrane accounts for only two percent of the total bilayer area of the cell, whereas the endoplasmic reticulum contains more than fifty percent and the mitochondria a further thirty percent.
Signaling | Lipid bilayer | Wikipedia | 421 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
The most familiar form of cellular signaling is likely synaptic transmission, whereby a nerve impulse that has reached the end of one neuron is conveyed to an adjacent neuron via the release of neurotransmitters. This transmission is made possible by the action of synaptic vesicles which are, inside the cell, loaded with the neurotransmitters to be released later. These loaded vesicles fuse with the cell membrane at the pre-synaptic terminal and their contents are released into the space outside the cell. The contents then diffuse across the synapse to the post-synaptic terminal.
Lipid bilayers are also involved in signal transduction through their role as the home of integral membrane proteins. This is an extremely broad and important class of biomolecule. It is estimated that up to a third of the human proteome are membrane proteins. Some of these proteins are linked to the exterior of the cell membrane. An example of this is the CD59 protein, which identifies cells as “self” and thus inhibits their destruction by the immune system. The HIV virus evades the immune system in part by grafting these proteins from the host membrane onto its own surface. Alternatively, some membrane proteins penetrate all the way through the bilayer and serve to relay individual signal events from the outside to the inside of the cell. The most common class of this type of protein is the G protein-coupled receptor (GPCR). GPCRs are responsible for much of the cell's ability to sense its surroundings and, because of this important role, approximately 40% of all modern drugs are targeted at GPCRs.
In addition to protein- and solution-mediated processes, it is also possible for lipid bilayers to participate directly in signaling. A classic example of this is phosphatidylserine-triggered phagocytosis. Normally, phosphatidylserine is asymmetrically distributed in the cell membrane and is present only on the interior side. During programmed cell death a protein called a scramblase equilibrates this distribution, displaying phosphatidylserine on the extracellular bilayer face. The presence of phosphatidylserine then triggers phagocytosis to remove the dead or dying cell.
Characterization methods | Lipid bilayer | Wikipedia | 476 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
The lipid bilayer is a difficult structure to study because it is so thin and fragile. To overcome these limitations, techniques have been developed to allow investigations of its structure and function.
Electrical measurements
Electrical measurements are a straightforward way to characterize an important function of a bilayer: its ability to segregate and prevent the flow of ions in solution. By applying a voltage across the bilayer and measuring the resulting current, the resistance of the bilayer is determined. This resistance is typically quite high (108 Ohm-cm2 or more) since the hydrophobic core is impermeable to charged species. The presence of even a few nanometer-scale holes results in a dramatic increase in current. The sensitivity of this system is such that even the activity of single ion channels can be resolved.
Fluorescence microscopy
A lipid bilayer cannot be seen with a traditional microscope because it is too thin, so researchers often use fluorescence microscopy. A sample is excited with one wavelength of light and observed in another, so that only fluorescent molecules with a matching excitation and emission profile will be seen. A natural lipid bilayer is not fluorescent, so at least one fluorescent dye needs to be attached to some of the molecules in the bilayer. Resolution is usually limited to a few hundred nanometers, which is unfortunately much larger than the thickness of a lipid bilayer.
Electron microscopy
Electron microscopy offers a higher resolution image. In an electron microscope, a beam of focused electrons interacts with the sample rather than a beam of light as in traditional microscopy. In conjunction with rapid freezing techniques, electron microscopy has also been used to study the mechanisms of inter- and intracellular transport, for instance in demonstrating that exocytotic vesicles are the means of chemical release at synapses.
Nuclear magnetic resonance spectroscopy
31P-Nuclear magnetic resonance spectroscopy is widely used for studies of phospholipid bilayers and biological membranes in native conditions. The analysis of 31P-NMR spectra of lipids could provide a wide range of information about lipid bilayer packing, phase transitions (gel phase, physiological liquid crystal phase, ripple phases, non bilayer phases), lipid head group orientation/dynamics, and elastic properties of pure lipid bilayer and as a result of binding of proteins and other biomolecules.
Atomic force microscopy | Lipid bilayer | Wikipedia | 478 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
A new method to study lipid bilayers is Atomic force microscopy (AFM). Rather than using a beam of light or particles, a very small sharpened tip scans the surface by making physical contact with the bilayer and moving across it, like a record player needle. AFM is a promising technique because it has the potential to image with nanometer resolution at room temperature and even under water or physiological buffer, conditions necessary for natural bilayer behavior. Utilizing this capability, AFM has been used to examine dynamic bilayer behavior including the formation of transmembrane pores (holes) and phase transitions in supported bilayers. Another advantage is that AFM does not require fluorescent or isotopic labeling of the lipids, since the probe tip interacts mechanically with the bilayer surface. Because of this, the same scan can image both lipids and associated proteins, sometimes even with single-molecule resolution. AFM can also probe the mechanical nature of lipid bilayers.
Dual polarisation interferometry
Lipid bilayers exhibit high levels of birefringence where the refractive index in the plane of the bilayer differs from that perpendicular by as much as 0.1 refractive index units. This has been used to characterise the degree of order and disruption in bilayers using dual polarisation interferometry to understand mechanisms of protein interaction.
Quantum chemical calculations
Lipid bilayers are complicated molecular systems with many degrees of freedom. Thus, atomistic simulation of membrane and in particular ab initio calculations of its properties is difficult and computationally expensive. Quantum chemical calculations has recently been successfully performed to estimate dipole and quadrupole moments of lipid membranes.
Transport across the bilayer
Passive diffusion | Lipid bilayer | Wikipedia | 354 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Most polar molecules have low solubility in the hydrocarbon core of a lipid bilayer and, as a consequence, have low permeability coefficients across the bilayer. This effect is particularly pronounced for charged species, which have even lower permeability coefficients than neutral polar molecules. Anions typically have a higher rate of diffusion through bilayers than cations. Compared to ions, water molecules actually have a relatively large permeability through the bilayer, as evidenced by osmotic swelling. When a cell or vesicle with a high interior salt concentration is placed in a solution with a low salt concentration it will swell and eventually burst. Such a result would not be observed unless water was able to pass through the bilayer with relative ease. The anomalously large permeability of water through bilayers is still not completely understood and continues to be the subject of active debate. Small uncharged apolar molecules diffuse through lipid bilayers many orders of magnitude faster than ions or water. This applies both to fats and organic solvents like chloroform and ether. Regardless of their polar character larger molecules diffuse more slowly across lipid bilayers than small molecules.
Ion pumps and channels
Two special classes of protein deal with the ionic gradients found across cellular and sub-cellular membranes in nature- ion channels and ion pumps. Both pumps and channels are integral membrane proteins that pass through the bilayer, but their roles are quite different. Ion pumps are the proteins that build and maintain the chemical gradients by utilizing an external energy source to move ions against the concentration gradient to an area of higher chemical potential. The energy source can be ATP, as is the case for the Na+-K+ ATPase. Alternatively, the energy source can be another chemical gradient already in place, as in the Ca2+/Na+ antiporter. It is through the action of ion pumps that cells are able to regulate pH via the pumping of protons.
In contrast to ion pumps, ion channels do not build chemical gradients but rather dissipate them in order to perform work or send a signal. Probably the most familiar and best studied example is the voltage-gated Na+ channel, which allows conduction of an action potential along neurons. All ion pumps have some sort of trigger or “gating” mechanism. In the previous example it was electrical bias, but other channels can be activated by binding a molecular agonist or through a conformational change in another nearby protein. | Lipid bilayer | Wikipedia | 510 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Endocytosis and exocytosis
Some molecules or particles are too large or too hydrophilic to pass through a lipid bilayer. Other molecules could pass through the bilayer but must be transported rapidly in such large numbers that channel-type transport is impractical. In both cases, these types of cargo can be moved across the cell membrane through fusion or budding of vesicles. When a vesicle is produced inside the cell and fuses with the plasma membrane to release its contents into the extracellular space, this process is known as exocytosis. In the reverse process, a region of the cell membrane will dimple inwards and eventually pinch off, enclosing a portion of the extracellular fluid to transport it into the cell. Endocytosis and exocytosis rely on very different molecular machinery to function, but the two processes are intimately linked and could not work without each other. The primary mechanism of this interdependence is the large amount of lipid material involved. In a typical cell, an area of bilayer equivalent to the entire plasma membrane travels through the endocytosis/exocytosis cycle in about half an hour.
Exocytosis in prokaryotes: Membrane vesicular exocytosis, popularly known as membrane vesicle trafficking, a Nobel prize-winning (year, 2013) process, is traditionally regarded as a prerogative of eukaryotic cells. This myth was however broken with the revelation that nanovesicles, popularly known as bacterial outer membrane vesicles, released by gram-negative microbes, translocate bacterial signal molecules to host or target cells to carry out multiple processes in favour of the secreting microbe e.g., in host cell invasion and microbe-environment interactions, in general.
Electroporation
Electroporation is the rapid increase in bilayer permeability induced by the application of a large artificial electric field across the membrane. Experimentally, electroporation is used to introduce hydrophilic molecules into cells. It is a particularly useful technique for large highly charged molecules such as DNA, which would never passively diffuse across the hydrophobic bilayer core. Because of this, electroporation is one of the key methods of transfection as well as bacterial transformation. It has even been proposed that electroporation resulting from lightning strikes could be a mechanism of natural horizontal gene transfer.
Mechanics | Lipid bilayer | Wikipedia | 497 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Lipid bilayers are large enough structures to have some of the mechanical properties of liquids or solids. The area compression modulus Ka, bending modulus Kb, and edge energy , can be used to describe them. Solid lipid bilayers also have a shear modulus, but like any liquid, the shear modulus is zero for fluid bilayers. These mechanical properties affect how the membrane functions. Ka and Kb affect the ability of proteins and small molecules to insert into the bilayer, and bilayer mechanical properties have been shown to alter the function of mechanically activated ion channels. Bilayer mechanical properties also govern what types of stress a cell can withstand without tearing. Although lipid bilayers can easily bend, most cannot stretch more than a few percent before rupturing.
As discussed in the Structure and organization section, the hydrophobic attraction of lipid tails in water is the primary force holding lipid bilayers together. Thus, the elastic modulus of the bilayer is primarily determined by how much extra area is exposed to water when the lipid molecules are stretched apart. It is not surprising given this understanding of the forces involved that studies have shown that Ka varies strongly with osmotic pressure but only weakly with tail length and unsaturation. Because the forces involved are so small, it is difficult to experimentally determine Ka. Most techniques require sophisticated microscopy and very sensitive measurement equipment.
In contrast to Ka, which is a measure of how much energy is needed to stretch the bilayer, Kb is a measure of how much energy is needed to bend or flex the bilayer. Formally, bending modulus is defined as the energy required to deform a membrane from its intrinsic curvature to some other curvature. Intrinsic curvature is defined by the ratio of the diameter of the head group to that of the tail group. For two-tailed PC lipids, this ratio is nearly one so the intrinsic curvature is nearly zero. If a particular lipid has too large a deviation from zero intrinsic curvature it will not form a bilayer and will instead form other phases such as micelles or inverted micelles. Addition of small hydrophilic molecules like sucrose into mixed lipid lamellar liposomes made from galactolipid-rich thylakoid membranes destabilises bilayers into the micellar phase. | Lipid bilayer | Wikipedia | 472 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
is a measure of how much energy it takes to expose a bilayer edge to water by tearing the bilayer or creating a hole in it. The origin of this energy is the fact that creating such an interface exposes some of the lipid tails to water, but the exact orientation of these border lipids is unknown. There is some evidence that both hydrophobic (tails straight) and hydrophilic (heads curved around) pores can coexist.
Fusion
Fusion is the process by which two lipid bilayers merge, resulting in one connected structure. If this fusion proceeds completely through both leaflets of both bilayers, a water-filled bridge is formed and the solutions contained by the bilayers can mix. Alternatively, if only one leaflet from each bilayer is involved in the fusion process, the bilayers are said to be hemifused. Fusion is involved in many cellular processes, in particular in eukaryotes, since the eukaryotic cell is extensively sub-divided by lipid bilayer membranes. Exocytosis, fertilization of an egg by sperm activation, and transport of waste products to the lysozome are a few of the many eukaryotic processes that rely on some form of fusion. Even the entry of pathogens can be governed by fusion, as many bilayer-coated viruses have dedicated fusion proteins to gain entry into the host cell. | Lipid bilayer | Wikipedia | 288 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
There are four fundamental steps in the fusion process. First, the involved membranes must aggregate, approaching each other to within several nanometers. Second, the two bilayers must come into very close contact (within a few angstroms). To achieve this close contact, the two surfaces must become at least partially dehydrated, as the bound surface water normally present causes bilayers to strongly repel. The presence of ions, in particular divalent cations like magnesium and calcium, strongly affects this step. One of the critical roles of calcium in the body is regulating membrane fusion. Third, a destabilization must form at one point between the two bilayers, locally distorting their structures. The exact nature of this distortion is not known. One theory is that a highly curved "stalk" must form between the two bilayers. Proponents of this theory believe that it explains why phosphatidylethanolamine, a highly curved lipid, promotes fusion. Finally, in the last step of fusion, this point defect grows and the components of the two bilayers mix and diffuse away from the site of contact.
The situation is further complicated when considering fusion in vivo since biological fusion is almost always regulated by the action of membrane-associated proteins. The first of these proteins to be studied were the viral fusion proteins, which allow an enveloped virus to insert its genetic material into the host cell (enveloped viruses are those surrounded by a lipid bilayer; some others have only a protein coat). Eukaryotic cells also use fusion proteins, the best-studied of which are the SNAREs. SNARE proteins are used to direct all vesicular intracellular trafficking. Despite years of study, much is still unknown about the function of this protein class. In fact, there is still an active debate regarding whether SNAREs are linked to early docking or participate later in the fusion process by facilitating hemifusion. | Lipid bilayer | Wikipedia | 398 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
In studies of molecular and cellular biology it is often desirable to artificially induce fusion. The addition of polyethylene glycol (PEG) causes fusion without significant aggregation or biochemical disruption. This procedure is now used extensively, for example by fusing B-cells with myeloma cells. The resulting “hybridoma” from this combination expresses a desired antibody as determined by the B-cell involved, but is immortalized due to the melanoma component. Fusion can also be artificially induced through electroporation in a process known as electrofusion. It is believed that this phenomenon results from the energetically active edges formed during electroporation, which can act as the local defect point to nucleate stalk growth between two bilayers.
Model systems
Lipid bilayers can be created artificially in the lab to allow researchers to perform experiments that cannot be done with natural bilayers. They can also be used in the field of Synthetic Biology, to define the boundaries of artificial cells. These synthetic systems are called model lipid bilayers. There are many different types of model bilayers, each having experimental advantages and disadvantages. They can be made with either synthetic or natural lipids. Among the most common model systems are:
Black lipid membranes (BLM)
Supported lipid bilayers (SLB)
Vesicles
Droplet Interface Bilayers (DIBs)
Commercial applications
To date, the most successful commercial application of lipid bilayers has been the use of liposomes for drug delivery, especially for cancer treatment. (Note- the term “liposome” is in essence synonymous with “vesicle” except that vesicle is a general term for the structure whereas liposome refers to only artificial not natural vesicles) The basic idea of liposomal drug delivery is that the drug is encapsulated in solution inside the liposome then injected into the patient. These drug-loaded liposomes travel through the system until they bind at the target site and rupture, releasing the drug. In theory, liposomes should make an ideal drug delivery system since they can isolate nearly any hydrophilic drug, can be grafted with molecules to target specific tissues and can be relatively non-toxic since the body possesses biochemical pathways for degrading lipids. | Lipid bilayer | Wikipedia | 477 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
The first generation of drug delivery liposomes had a simple lipid composition and suffered from several limitations. Circulation in the bloodstream was extremely limited due to both renal clearing and phagocytosis. Refinement of the lipid composition to tune fluidity, surface charge density, and surface hydration resulted in vesicles that adsorb fewer proteins from serum and thus are less readily recognized by the immune system. The most significant advance in this area was the grafting of polyethylene glycol (PEG) onto the liposome surface to produce “stealth” vesicles, which circulate over long times without immune or renal clearing.
The first stealth liposomes were passively targeted at tumor tissues. Because tumors induce rapid and uncontrolled angiogenesis they are especially “leaky” and allow liposomes to exit the bloodstream at a much higher rate than normal tissue would. More recently work has been undertaken to graft antibodies or other molecular markers onto the liposome surface in the hope of actively binding them to a specific cell or tissue type. Some examples of this approach are already in clinical trials.
Another potential application of lipid bilayers is the field of biosensors. Since the lipid bilayer is the barrier between the interior and exterior of the cell, it is also the site of extensive signal transduction. Researchers over the years have tried to harness this potential to develop a bilayer-based device for clinical diagnosis or bioterrorism detection. Progress has been slow in this area and, although a few companies have developed automated lipid-based detection systems, they are still targeted at the research community. These include Biacore (now GE Healthcare Life Sciences), which offers a disposable chip for utilizing lipid bilayers in studies of binding kinetics and Nanion Inc., which has developed an automated patch clamping system.
A supported lipid bilayer (SLB) as described above has achieved commercial success as a screening technique to measure the permeability of drugs. This parallel artificial membrane permeability assay (PAMPA) technique measures the permeability across specifically formulated lipid cocktail(s) found to be highly correlated with Caco-2 cultures, the gastrointestinal tract, blood–brain barrier and skin.
History | Lipid bilayer | Wikipedia | 477 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
By the early twentieth century scientists had come to believe that cells are surrounded by a thin oil-like barrier, but the structural nature of this membrane was not known. Two experiments in 1925 laid the groundwork to fill in this gap. By measuring the capacitance of erythrocyte solutions, Hugo Fricke determined that the cell membrane was 3.3 nm thick.
Although the results of this experiment were accurate, Fricke misinterpreted the data to mean that the cell membrane is a single molecular layer. Prof. Dr. Evert Gorter (1881–1954) and F. Grendel of Leiden University approached the problem from a different perspective, spreading the erythrocyte lipids as a monolayer on a Langmuir-Blodgett trough. When they compared the area of the monolayer to the surface area of the cells, they found a ratio of two to one. Later analyses showed several errors and incorrect assumptions with this experiment but, serendipitously, these errors canceled out and from this flawed data Gorter and Grendel drew the correct conclusion- that the cell membrane is a lipid bilayer.
This theory was confirmed through the use of electron microscopy in the late 1950s. Although he did not publish the first electron microscopy study of lipid bilayers J. David Robertson was the first to assert that the two dark electron-dense bands were the headgroups and associated proteins of two apposed lipid monolayers. In this body of work, Robertson put forward the concept of the “unit membrane.” This was the first time the bilayer structure had been universally assigned to all cell membranes as well as organelle membranes. | Lipid bilayer | Wikipedia | 348 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Around the same time, the development of model membranes confirmed that the lipid bilayer is a stable structure that can exist independent of proteins. By “painting” a solution of lipid in organic solvent across an aperture, Mueller and Rudin were able to create an artificial bilayer and determine that this exhibited lateral fluidity, high electrical resistance and self-healing in response to puncture, all of which are properties of a natural cell membrane. A few years later, Alec Bangham showed that bilayers, in the form of lipid vesicles, could also be formed simply by exposing a dried lipid sample to water. This demonstrated that lipid bilayers form spontaneously via self assembly and do not require a patterned support structure. In 1977, a totally synthetic bilayer membrane was prepared by Kunitake and Okahata, from a single organic compound, didodecyldimethylammonium bromide. This showed that the bilayer membrane was assembled by the intermolecular forces. | Lipid bilayer | Wikipedia | 204 | 158011 | https://en.wikipedia.org/wiki/Lipid%20bilayer | Biology and health sciences | Cell parts | Biology |
Sepsis is a potentially life-threatening condition that arises when the body's response to infection causes injury to its own tissues and organs.
This initial stage of sepsis is followed by suppression of the immune system. Common signs and symptoms include fever, increased heart rate, increased breathing rate, and confusion. There may also be symptoms related to a specific infection, such as a cough with pneumonia, or painful urination with a kidney infection. The very young, old, and people with a weakened immune system may not have any symptoms that are specific to their infection, and their body temperature may be low or normal instead of constituting a fever. Severe sepsis causes poor organ function or blood flow. The presence of low blood pressure, high blood lactate, or low urine output may suggest poor blood flow. Septic shock is low blood pressure due to sepsis that does not improve after fluid replacement.
Sepsis is caused by many organisms including bacteria, viruses, and fungi. Common locations for the primary infection include the lungs, brain, urinary tract, skin, and abdominal organs. Risk factors include being very young or old, a weakened immune system from conditions such as cancer or diabetes, major trauma, and burns. Previously, a sepsis diagnosis required the presence of at least two systemic inflammatory response syndrome (SIRS) criteria in the setting of presumed infection. In 2016, a shortened sequential organ failure assessment score (SOFA score), known as the quick SOFA score (qSOFA), replaced the SIRS system of diagnosis. qSOFA criteria for sepsis include at least two of the following three: increased breathing rate, change in the level of consciousness, and low blood pressure. Sepsis guidelines recommend obtaining blood cultures before starting antibiotics; however, the diagnosis does not require the blood to be infected. Medical imaging is helpful when looking for the possible location of the infection. Other potential causes of similar signs and symptoms include anaphylaxis, adrenal insufficiency, low blood volume, heart failure, and pulmonary embolism. | Sepsis | Wikipedia | 416 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Sepsis requires immediate treatment with intravenous fluids and antimicrobial medications. Ongoing care and stabilization often continues in an intensive care unit. If an adequate trial of fluid replacement is not enough to maintain blood pressure, then the use of medications that raise blood pressure becomes necessary. Mechanical ventilation and dialysis may be needed to support the function of the lungs and kidneys, respectively. A central venous catheter and an arterial catheter may be placed for access to the bloodstream and to guide treatment. Other helpful measurements include cardiac output and superior vena cava oxygen saturation. People with sepsis need preventive measures for deep vein thrombosis, stress ulcers, and pressure ulcers unless other conditions prevent such interventions. Some people might benefit from tight control of blood sugar levels with insulin. The use of corticosteroids is controversial, with some reviews finding benefit, and others not.
Disease severity partly determines the outcome. The risk of death from sepsis is as high as 30%, while for severe sepsis it is as high as 50%, and the risk of death from septic shock is 80%. Sepsis affected about 49 million people in 2017, with 11 million deaths (1 in 5 deaths worldwide). In the developed world, approximately 0.2 to 3 people per 1000 are affected by sepsis yearly, resulting in about a million cases per year in the United States. Rates of disease have been increasing. Some data indicate that sepsis is more common among males than females, however, other data show a greater prevalence of the disease among women. Descriptions of sepsis date back to the time of Hippocrates.
Signs and symptoms
In addition to symptoms related to the actual cause, people with sepsis may have a fever, low body temperature, rapid breathing, a fast heart rate, confusion, and edema. Early signs include a rapid heart rate, decreased urination, and high blood sugar. Signs of established sepsis include confusion, metabolic acidosis (which may be accompanied by a faster breathing rate that leads to respiratory alkalosis), low blood pressure due to decreased systemic vascular resistance, higher cardiac output, and disorders in blood-clotting that may lead to organ failure. Fever is the most common presenting symptom in sepsis, but fever may be absent in some people such as the elderly or those who are immunocompromised. | Sepsis | Wikipedia | 493 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
The drop in blood pressure seen in sepsis can cause lightheadedness and is part of the criteria for septic shock.
Oxidative stress is observed in septic shock, with circulating levels of copper and vitamin C being decreased.
Diastolic blood pressure falls during the early stages of sepsis, causing a widening/increasing of pulse pressure, which is the difference between the systolic and diastolic blood pressures. If sepsis becomes severe and hemodynamic compromise advances, the systolic pressure also decreases, causing a narrowing/decreasing of pulse pressure. A pulse pressure of over 70 mmHg in patients with sepsis is correlated with an increased chance of survival. A widened pulse pressure is also correlated with an increased chance that someone with sepsis will benefit from and respond to IV fluids.
Cause
Infections leading to sepsis are usually bacterial but may be fungal, parasitic, or viral. Gram-positive bacteria were the primary cause of sepsis before the introduction of antibiotics in the 1950s. After the introduction of antibiotics, gram-negative bacteria became the predominant cause of sepsis from the 1960s to the 1980s. After the 1980s, gram-positive bacteria, most commonly staphylococci, are thought to cause more than 50% of cases of sepsis. Other commonly implicated bacteria include Streptococcus pyogenes, Escherichia coli, Pseudomonas aeruginosa, and Klebsiella species. Fungal sepsis accounts for approximately 5% of severe sepsis and septic shock cases; the most common cause of fungal sepsis is an infection by Candida species of yeast, a frequent hospital-acquired infection. The most common causes for parasitic sepsis are Plasmodium (which leads to malaria), Schistosoma and Echinococcus.
The most common sites of infection resulting in severe sepsis are the lungs, the abdomen, and the urinary tract. Typically, 50% of all sepsis cases start as an infection in the lungs. In one-third to one-half of cases, the source of infection is unclear.
Pathophysiology | Sepsis | Wikipedia | 433 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Sepsis is caused by a combination of factors related to the particular invading pathogen(s) and the status of the immune system of the host. The early phase of sepsis characterized by excessive inflammation (sometimes resulting in a cytokine storm) may be followed by a prolonged period of decreased functioning of the immune system. Either of these phases may prove fatal. On the other hand, systemic inflammatory response syndrome (SIRS) occurs in people without the presence of infection, for example, in those with burns, polytrauma, or the initial state in pancreatitis and chemical pneumonitis. However, sepsis also causes similar response to SIRS.
Microbial factors
Bacterial virulence factors, such as glycocalyx and various adhesins, allow colonization, immune evasion, and establishment of disease in the host. Sepsis caused by gram-negative bacteria is thought to be largely due to a response by the host to the lipid A component of lipopolysaccharide, also called endotoxin. Sepsis caused by gram-positive bacteria may result from an immunological response to cell wall lipoteichoic acid. Bacterial exotoxins that act as superantigens also may cause sepsis. Superantigens simultaneously bind major histocompatibility complex and T-cell receptors in the absence of antigen presentation. This forced receptor interaction induces the production of pro-inflammatory chemical signals (cytokines) by T-cells. | Sepsis | Wikipedia | 305 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
There are a number of microbial factors that may cause the typical septic inflammatory cascade. An invading pathogen is recognized by its pathogen-associated molecular patterns (PAMPs). Examples of PAMPs include lipopolysaccharides and flagellin in gram-negative bacteria, muramyl dipeptide in the peptidoglycan of the gram-positive bacterial cell wall, and CpG bacterial DNA. These PAMPs are recognized by the pattern recognition receptors (PRRs) of the innate immune system, which may be membrane-bound or cytosolic. There are four families of PRRs: the toll-like receptors, the C-type lectin receptors, the NOD-like receptors, and the RIG-I-like receptors. Invariably, the association of a PAMP and a PRR will cause a series of intracellular signalling cascades. Consequentially, transcription factors such as nuclear factor-kappa B and activator protein-1, will up-regulate the expression of pro-inflammatory and anti-inflammatory cytokines.
Host factors
Upon detection of microbial antigens, the host systemic immune system is activated. Immune cells not only recognise pathogen-associated molecular patterns but also damage-associated molecular patterns from damaged tissues. An uncontrolled immune response is then activated because leukocytes are not recruited to the specific site of infection, but instead, they are recruited all over the body. Then, an immunosuppression state ensues when the proinflammatory T helper cell 1 (TH1) is shifted to TH2, mediated by interleukin 10, which is known as "compensatory anti-inflammatory response syndrome". The apoptosis (cell death) of lymphocytes further worsens the immunosuppression. Neutrophils, monocytes, macrophages, dendritic cells, CD4+ T cells, and B cells all undergo apoptosis, whereas regulatory T cells are more apoptosis-resistant. Subsequently, multiple organ failure ensues because tissues are unable to use oxygen efficiently due to inhibition of cytochrome c oxidase. | Sepsis | Wikipedia | 450 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Inflammatory responses cause multiple organ dysfunction syndrome through various mechanisms as described below. Increased permeability of the lung vessels causes leaking of fluids into alveoli, which results in pulmonary edema and acute respiratory distress syndrome (ARDS). Impaired utilization of oxygen in the liver impairs bile salt transport, causing jaundice (yellowish discoloration of the skin). In kidneys, inadequate oxygenation results in tubular epithelial cell injury (of the cells lining the kidney tubules), and thus causes acute kidney injury (AKI). Meanwhile, in the heart, impaired calcium transport, and low production of adenosine triphosphate (ATP), can cause myocardial depression, reducing cardiac contractility and causing heart failure. In the gastrointestinal tract, increased permeability of the mucosa alters the microflora, causing mucosal bleeding and paralytic ileus. In the central nervous system, direct damage of the brain cells and disturbances of neurotransmissions causes altered mental status. Cytokines such as tumor necrosis factor, interleukin 1, and interleukin 6 may activate procoagulation factors in the cells lining blood vessels, leading to endothelial damage. The damaged endothelial surface inhibits anticoagulant properties as well as increases antifibrinolysis, which may lead to intravascular clotting, the formation of blood clots in small blood vessels, and multiple organ failure.
The low blood pressure seen in those with sepsis is the result of various processes, including excessive production of chemicals that dilate blood vessels such as nitric oxide, a deficiency of chemicals that constrict blood vessels such as vasopressin, and activation of ATP-sensitive potassium channels. In those with severe sepsis and septic shock, this sequence of events leads to a type of circulatory shock known as distributive shock.
Diagnosis
Early diagnosis is necessary to properly manage sepsis, as the initiation of rapid therapy is key to reducing deaths from severe sepsis. Some hospitals use alerts generated from electronic health records to bring attention to potential cases as early as possible. | Sepsis | Wikipedia | 450 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Within the first three hours of suspected sepsis, diagnostic studies should include white blood cell counts, measuring serum lactate, and obtaining appropriate cultures before starting antibiotics, so long as this does not delay their use by more than 45 minutes. To identify the causative organism(s), at least two sets of blood cultures using bottles with media for aerobic and anaerobic organisms are necessary. At least one should be drawn through the skin and one through each vascular access device (such as an IV catheter) that has been in place for more than 48 hours. Bacteria are present in the blood in only about 30% of cases. Another possible method of detection is by polymerase chain reaction. If other sources of infection are suspected, cultures of these sources, such as urine, cerebrospinal fluid, wounds, or respiratory secretions, also should be obtained, as long as this does not delay the use of antibiotics.
Within six hours, if blood pressure remains low despite initial fluid resuscitation of 30 mL/kg, or if initial lactate is ≥ four mmol/L (36 mg/dL), central venous pressure and central venous oxygen saturation should be measured. Lactate should be re-measured if the initial lactate was elevated. Evidence for point of care lactate measurement over usual methods of measurement, however, is poor.
Within twelve hours, it is essential to diagnose or exclude any source of infection that would require emergent source control, such as a necrotizing soft tissue infection, an infection causing inflammation of the abdominal cavity lining, an infection of the bile duct, or an intestinal infarction. A pierced internal organ (free air on an abdominal X-ray or CT scan), an abnormal chest X-ray consistent with pneumonia (with focal opacification), or petechiae, purpura, or purpura fulminans may indicate the presence of an infection.
Definitions | Sepsis | Wikipedia | 401 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Previously, SIRS criteria had been used to define sepsis. If the SIRS criteria are negative, it is very unlikely the person has sepsis; if it is positive, there is just a moderate probability that the person has sepsis. According to SIRS, there were different levels of sepsis: sepsis, severe sepsis, and septic shock. The definition of SIRS is shown below:
SIRS is the presence of two or more of the following: abnormal body temperature, heart rate, respiratory rate, or blood gas, and white blood cell count.
Sepsis is defined as SIRS in response to an infectious process.
Severe sepsis is defined as sepsis with sepsis-induced organ dysfunction or tissue hypoperfusion (manifesting as hypotension, elevated lactate, or decreased urine output). Severe sepsis is an infectious disease state associated with multiple organ dysfunction syndrome (MODS)
Septic shock is severe sepsis plus persistently low blood pressure, despite the administration of intravenous fluids.
In 2016 a new consensus was reached to replace screening by systemic inflammatory response syndrome (SIRS) with the sequential organ failure assessment (SOFA score) and the abbreviated version (qSOFA). The three criteria for the qSOFA score include a respiratory rate greater than or equal to 22 breaths per minute, systolic blood pressure 100 mmHg or less, and altered mental status. Sepsis is suspected when 2 of the qSOFA criteria are met. The SOFA score was intended to be used in the intensive care unit (ICU) where it is administered upon admission to the ICU and then repeated every 48 hours, whereas the qSOFA could be used outside the ICU. Some advantages of the qSOFA score are that it can be administered quickly and does not require labs. However, the American College of Chest Physicians (CHEST) raised concerns that qSOFA and SOFA criteria may lead to delayed diagnosis of serious infection, leading to delayed treatment. Although SIRS criteria can be too sensitive and not specific enough in identifying sepsis, SOFA also has its limitations and is not intended to replace the SIRS definition. qSOFA has also been found to be poorly sensitive though decently specific for the risk of death with SIRS possibly better for screening. NOTE - Surviving Sepsis Campaign 2021 Guidelines recommend "against using qSOFA compared with SIRS, NEWS, or MEWS as a single screening tool for sepsis or septic shock".
End-organ dysfunction | Sepsis | Wikipedia | 511 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Examples of end-organ dysfunction include the following:
Lungs: acute respiratory distress syndrome (ARDS) (PaO2/FiO2 ratio < 300), different ratio in pediatric acute respiratory distress syndrome
Brain: encephalopathy symptoms including agitation, confusion, and coma; causes may include ischemia, bleeding, formation of blood clots in small blood vessels, microabscesses, multifocal necrotizing leukoencephalopathy
Liver: disruption of protein synthetic function manifests acutely as progressive disruption of blood clotting due to an inability to synthesize clotting factors and disruption of metabolic functions leads to impaired bilirubin metabolism, resulting in elevated unconjugated serum bilirubin levels
Kidney: low urine output or no urine output, electrolyte abnormalities, or volume overload
Heart: systolic and diastolic heart failure, likely due to chemical signals that depress myocyte function, cellular damage, manifest as a troponin leak (although not necessarily ischemic in nature) | Sepsis | Wikipedia | 217 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
More specific definitions of end-organ dysfunction exist for SIRS in pediatrics.
Cardiovascular dysfunction (after fluid resuscitation with at least 40 mL/kg of crystalloid)
hypotension with blood pressure < 5th percentile for age or systolic blood pressure < 2 standard deviations below normal for age, or
vasopressor requirement, or
Two of the following criteria:
unexplained metabolic acidosis with base deficit > 5 mEq/L
lactic acidosis: serum lactate 2 times the upper limit of normal
oliguria (urine output )
prolonged capillary refill > 5 seconds
core to peripheral temperature difference
Respiratory dysfunction (in the absence of a cyanotic heart defect or a known chronic respiratory disease)
the ratio of the arterial partial pressure of oxygen to the fraction of oxygen in the gases inspired (PaO2/FiO2) < 300 (the definition of acute lung injury), or
arterial partial pressure of carbon dioxide (PaCO2) > 65 torr (20 mmHg) over baseline PaCO2 (evidence of hypercapnic respiratory failure), or
supplemental oxygen requirement of greater than FiO2 0.5 to maintain oxygen saturation ≥ 92%
Neurologic dysfunction
Glasgow Coma Score (GCS) ≤ 11, or
altered mental status with a drop in GCS of 3 or more points in a person with developmental delay/intellectual disability
Hematologic dysfunction
platelet count or 50% drop from the maximum in chronically thrombocytopenic, or
international normalized ratio (INR) > 2
Disseminated intravascular coagulation
Kidney dysfunction
serum creatinine ≥ 2 times the upper limit of normal for age or 2-fold increase in baseline creatinine in people with chronic kidney disease
Liver dysfunction (only applicable to infants > 1 month)
total serum bilirubin ≥ 4 mg/dL, or
alanine aminotransferase (ALT) ≥ 2 times the upper limit of normal
Consensus definitions, however, continue to evolve, with the latest expanding the list of signs and symptoms of sepsis to reflect clinical bedside experience. | Sepsis | Wikipedia | 435 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Biomarkers
Biomarkers can help with diagnosis because they can point to the presence or severity of sepsis, although their exact role in the management of sepsis remains undefined. A 2013 review concluded moderate-quality evidence exists to support the use of the procalcitonin level as a method to distinguish sepsis from non-infectious causes of SIRS. The same review found the sensitivity of the test to be 77% and the specificity to be 79%. The authors suggested that procalcitonin may serve as a helpful diagnostic marker for sepsis, but cautioned that its level alone does not definitively make the diagnosis. More current literature recommends utilizing the PCT to direct antibiotic therapy for improved antibiotic stewardship and better patient outcomes.
A 2012 systematic review found that soluble urokinase-type plasminogen activator receptor (SuPAR) is a nonspecific marker of inflammation and does not accurately diagnose sepsis. This same review concluded, however, that SuPAR has prognostic value, as higher SuPAR levels are associated with an increased rate of death in those with sepsis. Serial measurement of lactate levels (approximately every 4 to 6 hours) may guide treatment and is associated with lower mortality in sepsis.
Differential diagnosis
The differential diagnosis for sepsis is broad and has to examine (to exclude) the non-infectious conditions that may cause the systemic signs of SIRS: alcohol withdrawal, acute pancreatitis, burns, pulmonary embolism, thyrotoxicosis, anaphylaxis, adrenal insufficiency, and neurogenic shock. Hyperinflammatory syndromes such as hemophagocytic lymphohistiocytosis (HLH) may have similar symptoms and are on the differential diagnosis.
Neonatal sepsis
In common clinical usage, neonatal sepsis refers to a bacterial blood stream infection in the first month of life, such as meningitis, pneumonia, pyelonephritis, or gastroenteritis, but neonatal sepsis also may be due to infection with fungi, viruses, or parasites. Criteria with regard to hemodynamic compromise or respiratory failure are not useful because they present too late for intervention.
Management | Sepsis | Wikipedia | 467 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Early recognition and focused management may improve the outcomes of sepsis. Current professional recommendations include several actions ("bundles") to be followed as soon as possible after diagnosis. Within the first three hours, someone with sepsis should have received antibiotics, and intravenous fluids if there is evidence of either low blood pressure or other evidence for inadequate blood supply to organs (as evidenced by a raised level of lactate); blood cultures also should be obtained within this period. After six hours the blood pressure should be adequate, close monitoring of blood pressure and blood supply to organs should be in place, and the lactate should be measured again if initially it was raised. A related bundle, the "Sepsis Six", is in widespread use in the United Kingdom; this requires the administration of antibiotics within an hour of recognition, blood cultures, lactate, and hemoglobin determination, urine output monitoring, high-flow oxygen, and intravenous fluids.
Apart from the timely administration of fluids and antibiotics, the management of sepsis also involves surgical drainage of infected fluid collections and appropriate support for organ dysfunction. This may include hemodialysis in kidney failure, mechanical ventilation in lung dysfunction, transfusion of blood products, and drug and fluid therapy for circulatory failure. Ensuring adequate nutrition—preferably by enteral feeding, but if necessary, by parenteral nutrition—is important during prolonged illness. Medication to prevent deep vein thrombosis and gastric ulcers also may be used.
Antibiotics
Two sets of blood cultures (aerobic and anaerobic) are recommended without delaying the initiation of antibiotics. Cultures from other sites such as respiratory secretions, urine, wounds, cerebrospinal fluid, and catheter insertion sites (in situ for more than 48 hours) are recommended if infections from these sites are suspected. In severe sepsis and septic shock, broad-spectrum antibiotics (usually two, a β-lactam antibiotic with broad coverage, or broad-spectrum carbapenem combined with fluoroquinolones, macrolides, or aminoglycosides) are recommended. The choice of antibiotics is important in determining the survival of the person. Some recommend they be given within one hour of making the diagnosis, stating that for every hour of delay in the administration of antibiotics, there is an associated 6% rise in mortality. Others did not find a benefit with early administration. | Sepsis | Wikipedia | 490 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Several factors determine the most appropriate choice for the initial antibiotic regimen. These factors include local patterns of bacterial sensitivity to antibiotics, whether the infection is thought to be a hospital or community-acquired infection, and which organ systems are thought to be infected. Antibiotic regimens should be reassessed daily and narrowed if appropriate. Treatment duration is typically 7–10 days with the type of antibiotic used directed by the results of cultures. If the culture result is negative, antibiotics should be de-escalated according to the person's clinical response or stopped altogether if an infection is not present to decrease the chances that the person is infected with multiple drug resistance organisms. In case of people having a high risk of being infected with multiple drug resistant organisms such as Pseudomonas aeruginosa, Acinetobacter baumannii, the addition of an antibiotic specific to the gram-negative organism is recommended. For methicillin-resistant Staphylococcus aureus (MRSA), vancomycin or teicoplanin is recommended. For Legionella infection, addition of macrolide or fluoroquinolone is chosen. If fungal infection is suspected, an echinocandin, such as caspofungin or micafungin, is chosen for people with severe sepsis, followed by triazole (fluconazole and itraconazole) for less ill people. Prolonged antibiotic prophylaxis is not recommended in people who has SIRS without any infectious origin such as acute pancreatitis and burns unless sepsis is suspected.
Once-daily dosing of aminoglycoside is sufficient to achieve peak plasma concentration for a clinical response without kidney toxicity. Meanwhile, for antibiotics with low volume distribution (vancomycin, teicoplanin, colistin), a loading dose is required to achieve an adequate therapeutic level to fight infections. Frequent infusions of beta-lactam antibiotics without exceeding the total daily dose would help to keep the antibiotics level above minimum inhibitory concentration (MIC), thus providing a better clinical response. Giving beta-lactam antibiotics continuously may be better than giving them intermittently. Access to therapeutic drug monitoring is important to ensure adequate drug therapeutic level while at the same time preventing the drug from reaching a toxic level. | Sepsis | Wikipedia | 474 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Intravenous fluids
The Surviving Sepsis Campaign has recommended 30 mL/kg of fluid to be given in adults in the first three hours followed by fluid titration according to blood pressure, urine output, respiratory rate, and oxygen saturation with a target mean arterial pressure (MAP) of 65 mmHg. In children an initial amount of 20 mL/kg is reasonable in shock. In cases of severe sepsis and septic shock where a central venous catheter is used to measure blood pressures dynamically, fluids should be administered until the central venous pressure reaches 8–12 mmHg. Once these goals are met, the central venous oxygen saturation (ScvO2), i.e., the oxygen saturation of venous blood as it returns to the heart as measured at the vena cava, is optimized. If the ScvO2 is less than 70%, blood may be given to reach a hemoglobin of 10 g/dL and then inotropes are added until the ScvO2 is optimized. In those with acute respiratory distress syndrome (ARDS) and sufficient tissue blood fluid, more fluids should be given carefully.
Crystalloid solution is recommended as the fluid of choice for resuscitation. Albumin can be used if a large amount of crystalloid is required for resuscitation. Crystalloid solutions shows little difference with hydroxyethyl starch in terms of risk of death. Starches also carry an increased risk of acute kidney injury, and need for blood transfusion. Various colloid solutions (such as modified gelatin) carry no advantage over crystalloid. Albumin also appears to be of no benefit over crystalloids. | Sepsis | Wikipedia | 349 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Blood products
The Surviving Sepsis Campaign recommended packed red blood cells transfusion for hemoglobin levels below 70 g/L if there is no myocardial ischemia, hypoxemia, or acute bleeding. In a 2014 trial, blood transfusions to keep target hemoglobin above 70 or 90 g/L did not make any difference to survival rates; meanwhile, those with a lower threshold of transfusion received fewer transfusions in total. Erythropoietin is not recommended in the treatment of anemia with septic shock because it may precipitate blood clotting events. Fresh frozen plasma transfusion usually does not correct the underlying clotting abnormalities before a planned surgical procedure. However, platelet transfusion is suggested for platelet counts below (10 × 109/L) without any risk of bleeding, or (20 × 109/L) with a high risk of bleeding, or (50 × 109/L) with active bleeding, before planned surgery or an invasive procedure. IV immunoglobulin is not recommended because its beneficial effects are uncertain. Monoclonal and polyclonal preparations of intravenous immunoglobulin (IVIG) do not lower the rate of death in newborns and adults with sepsis. Evidence for the use of IgM-enriched polyclonal preparations of IVIG is inconsistent. On the other hand, the use of antithrombin to treat disseminated intravascular coagulation is also not useful. Meanwhile, the blood purification technique (such as hemoperfusion, plasma filtration, and coupled plasma filtration adsorption) to remove inflammatory mediators and bacterial toxins from the blood also does not demonstrate any survival benefit for septic shock.
Vasopressors
If the person has been sufficiently fluid resuscitated but the mean arterial pressure is not greater than 65 mmHg, vasopressors are recommended. Norepinephrine (noradrenaline) is recommended as the initial choice. Delaying initiation of vasopressor therapy during septic shock is associated with increased mortality. | Sepsis | Wikipedia | 435 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
Norepinephrine is often used as a first-line treatment for hypotensive septic shock because evidence shows that there is a relative deficiency of vasopressin when shock continues for 24 to 48 hours. Norepinephrine raises blood pressure through a vasoconstriction effect, with little effect on stroke volume and heart rate. In some people, the required dose of vasopressor needed to increase the mean arterial pressure can become exceedingly high and it becomes toxic. To reduce the required dose of vasopressor, epinephrine may be added. Epinephrine is not often used as a first-line treatment for hypotensive shock because it reduces blood flow to the abdominal organs and increases lactate levels. Vasopressin can be used in septic shock because studies have shown that there is a relative deficiency of vasopressin when shock continues for 24 to 48 hours. However, vasopressin reduces blood flow to the heart, fingers/toes, and abdominal organs, resulting in a lack of oxygen supply to these tissues. Dopamine is typically not recommended. Although dopamine is useful for increasing the stroke volume of the heart, it causes more abnormal heart rhythms than norepinephrine and also has an immunosuppressive effect. Dopamine is not proven to have protective properties on the kidneys. Dobutamine can also be used in hypotensive septic shock to increase cardiac output and correct blood flow to the tissues. Dobutamine is not used as often as epinephrine due to its associated side effects, which include reducing blood flow to the gut. Additionally, dobutamine increases the cardiac output by abnormally increasing the heart rate.
Steroids
The use of steroids in sepsis is controversial. Studies do not give a clear picture as to whether and when glucocorticoids should be used. The 2016 Surviving Sepsis Campaign recommends low dose hydrocortisone only if both intravenous fluids and vasopressors are not able to adequately treat septic shock. The 2021 Surviving Sepsis Campaign recommends IV corticosteroids for adults with septic shock who have an ongoing requirement for vasopressor therapy. A 2019 Cochrane review found low-quality evidence of benefit, as did two 2019 reviews. | Sepsis | Wikipedia | 482 | 158400 | https://en.wikipedia.org/wiki/Sepsis | Biology and health sciences | Symptoms and signs | Health |
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