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An astronomer is a scientist in the field of astronomy who focuses on a specific question or field outside the scope of Earth. Astronomers observe astronomical objects, such as stars, planets, moons, comets and galaxies – in either observational (by analyzing the data) or theoretical astronomy. Examples of topics or fields astronomers study include planetary science, solar astronomy, the origin or evolution of stars, or the formation of galaxies. A related but distinct subject is physical cosmology, which studies the Universe as a whole.
Types
Astronomers typically fall under either of two main types: observational and theoretical. Observational astronomers make direct observations of celestial objects and analyze the data. In contrast, theoretical astronomers create and investigate models of things that cannot be observed. Because it takes millions to billions of years for a system of stars or a galaxy to complete a life cycle, astronomers must observe snapshots of different systems at unique points in their evolution to determine how they form, evolve, and die. They use this data to create models or simulations to theorize how different celestial objects work.
Further subcategories under these two main branches of astronomy include planetary astronomy, astrobiology, stellar astronomy, astrometry, galactic astronomy, extragalactic astronomy, or physical cosmology. Astronomers can also specialize in certain specialties of observational astronomy, such as infrared astronomy, neutrino astronomy, x-ray astronomy, and gravitational-wave astronomy.
Academic
History
Historically, astronomy was more concerned with the classification and description of phenomena in the sky, while astrophysics attempted to explain these phenomena and the differences between them using physical laws. Today, that distinction has mostly disappeared and the terms "astronomer" and "astrophysicist" are interchangeable. Professional astronomers are highly educated individuals who typically have a PhD in physics or astronomy and are employed by research institutions or universities. They spend the majority of their time working on research, although they quite often have other duties such as teaching, building instruments, or aiding in the operation of an observatory.
The American Astronomical Society, which is the major organization of professional astronomers in North America, has approximately 8,200 members (as of 2024). This number includes scientists from other fields such as physics, geology, and engineering, whose research interests are closely related to astronomy. The International Astronomical Union comprises about 12,700 members from 92 countries who are involved in astronomical research at the PhD level and beyond (as of 2024). | Astronomer | Wikipedia | 492 | 580 | https://en.wikipedia.org/wiki/Astronomer | Physical sciences | Astronomy basics | Astronomy |
Contrary to the classical image of an old astronomer peering through a telescope through the dark hours of the night, it is far more common to use a charge-coupled device (CCD) camera to record a long, deep exposure, allowing a more sensitive image to be created because the light is added over time. Before CCDs, photographic plates were a common method of observation. Modern astronomers spend relatively little time at telescopes, usually just a few weeks per year. Analysis of observed phenomena, along with making predictions as to the causes of what they observe, takes the majority of observational astronomers' time.
Activities and graduate degree training
Astronomers who serve as faculty spend much of their time teaching undergraduate and graduate classes. Most universities also have outreach programs, including public telescope time and sometimes planetariums, as a public service to encourage interest in the field.
Those who become astronomers usually have a broad background in physics, mathematics, sciences, and computing in high school. Taking courses that teach how to research, write, and present papers are part of the higher education of an astronomer, while most astronomers attain both a Master's degree and eventually a PhD degree in astronomy, physics or astrophysics.
PhD training typically involves 5-6 years of study, including completion of upper-level courses in the core sciences, a competency examination, experience with teaching undergraduates and participating in outreach programs, work on research projects under the student's supervising professor, completion of a PhD thesis, and passing a final oral exam. Throughout the PhD training, a successful student is financially supported with a stipend.
Amateur astronomers
While there is a relatively low number of professional astronomers, the field is popular among amateurs. Most cities have amateur astronomy clubs that meet on a regular basis and often host star parties. The Astronomical Society of the Pacific is the largest general astronomical society in the world, comprising both professional and amateur astronomers as well as educators from 70 different nations.
As with any hobby, most people who practice amateur astronomy may devote a few hours a month to stargazing and reading the latest developments in research. However, amateurs span the range from so-called "armchair astronomers" to the highly ambitious people who own science-grade telescopes and instruments with which they are able to make their own discoveries, create astrophotographs, and assist professional astronomers in research. | Astronomer | Wikipedia | 468 | 580 | https://en.wikipedia.org/wiki/Astronomer | Physical sciences | Astronomy basics | Astronomy |
ASCII ( ), an acronym for American Standard Code for Information Interchange, is a character encoding standard for electronic communication. ASCII codes represent text in computers, telecommunications equipment, and other devices. ASCII has just 128 code points, of which only 95 are , which severely limit its scope. The set of available punctuation had significant impact on the syntax of computer languages and text markup. ASCII hugely influenced the design of character sets used by modern computers, including Unicode which has over a million code points, but the first 128 of these are the same as ASCII.
The Internet Assigned Numbers Authority (IANA) prefers the name US-ASCII for this character encoding.
ASCII is one of the IEEE milestones.
Overview
ASCII was developed in part from telegraph code. Its first commercial use was in the Teletype Model 33 and the Teletype Model 35 as a seven-bit teleprinter code promoted by Bell data services. Work on the ASCII standard began in May 1961, with the first meeting of the American Standards Association's (ASA) (now the American National Standards Institute or ANSI) X3.2 subcommittee. The first edition of the standard was published in 1963, underwent a major revision during 1967, and experienced its most recent update during 1986. Compared to earlier telegraph codes, the proposed Bell code and ASCII were both ordered for more convenient sorting (i.e., alphabetization) of lists and added features for devices other than teleprinters.
The use of ASCII format for Network Interchange was described in 1969. That document was formally elevated to an Internet Standard in 2015.
Originally based on the (modern) English alphabet, ASCII encodes 128 specified characters into seven-bit integers as shown by the ASCII chart in this article. Ninety-five of the encoded characters are printable: these include the digits 0 to 9, lowercase letters a to z, uppercase letters A to Z, and punctuation symbols. In addition, the original ASCII specification included 33 non-printing control codes which originated with s; most of these are now obsolete, although a few are still commonly used, such as the carriage return, line feed, and tab codes.
For example, lowercase i would be represented in the ASCII encoding by binary 1101001 = hexadecimal 69 (i is the ninth letter) = decimal 105. | ASCII | Wikipedia | 504 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
Despite being an American standard, ASCII does not have a code point for the cent (¢). It also does not support English terms with diacritical marks such as résumé and jalapeño, or proper nouns with diacritical marks such as Beyoncé (although on certain devices characters could be combined with punctuation such as Tilde (~) and Backtick (`) to approximate such characters.)
History
The American Standard Code for Information Interchange (ASCII) was developed under the auspices of a committee of the American Standards Association (ASA), called the X3 committee, by its X3.2 (later X3L2) subcommittee, and later by that subcommittee's X3.2.4 working group (now INCITS). The ASA later became the United States of America Standards Institute (USASI) and ultimately became the American National Standards Institute (ANSI).
With the other special characters and control codes filled in, ASCII was published as ASA X3.4-1963, leaving 28 code positions without any assigned meaning, reserved for future standardization, and one unassigned control code. There was some debate at the time whether there should be more control characters rather than the lowercase alphabet. The indecision did not last long: during May 1963 the CCITT Working Party on the New Telegraph Alphabet proposed to assign lowercase characters to sticks 6 and 7, and International Organization for Standardization TC 97 SC 2 voted during October to incorporate the change into its draft standard. The X3.2.4 task group voted its approval for the change to ASCII at its May 1963 meeting. Locating the lowercase letters in sticks 6 and 7 caused the characters to differ in bit pattern from the upper case by a single bit, which simplified case-insensitive character matching and the construction of keyboards and printers.
The X3 committee made other changes, including other new characters (the brace and vertical bar characters), renaming some control characters (SOM became start of header (SOH)) and moving or removing others (RU was removed). ASCII was subsequently updated as USAS X3.4-1967, then USAS X3.4-1968, ANSI X3.4-1977, and finally, ANSI X3.4-1986.
Revisions | ASCII | Wikipedia | 474 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
ASA X3.4-1963
ASA X3.4-1965 (approved, but not published, nevertheless used by IBM 2260 & 2265 Display Stations and IBM 2848 Display Control)
USAS X3.4-1967
USAS X3.4-1968
ANSI X3.4-1977
ANSI X3.4-1986
ANSI X3.4-1986 (R1992)
ANSI X3.4-1986 (R1997)
ANSI INCITS 4-1986 (R2002)
ANSI INCITS 4-1986 (R2007)
INCITS 4-1986 (R2012)
INCITS 4-1986 (R2017)
INCITS 4-1986 (R2022)
In the X3.15 standard, the X3 committee also addressed how ASCII should be transmitted (least significant bit first) and recorded on perforated tape. They proposed a 9-track standard for magnetic tape and attempted to deal with some punched card formats.
Design considerations
Bit width
The X3.2 subcommittee designed ASCII based on the earlier teleprinter encoding systems. Like other character encodings, ASCII specifies a correspondence between digital bit patterns and character symbols (i.e. graphemes and control characters). This allows digital devices to communicate with each other and to process, store, and communicate character-oriented information such as written language. Before ASCII was developed, the encodings in use included 26 alphabetic characters, 10 numerical digits, and from 11 to 25 special graphic symbols. To include all these, and control characters compatible with the Comité Consultatif International Téléphonique et Télégraphique (CCITT) International Telegraph Alphabet No. 2 (ITA2) standard of 1932, FIELDATA (1956), and early EBCDIC (1963), more than 64 codes were required for ASCII.
ITA2 was in turn based on Baudot code, the 5-bit telegraph code Émile Baudot invented in 1870 and patented in 1874. | ASCII | Wikipedia | 422 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
The committee debated the possibility of a shift function (like in ITA2), which would allow more than 64 codes to be represented by a six-bit code. In a shifted code, some character codes determine choices between options for the following character codes. It allows compact encoding, but is less reliable for data transmission, as an error in transmitting the shift code typically makes a long part of the transmission unreadable. The standards committee decided against shifting, and so ASCII required at least a seven-bit code.
The committee considered an eight-bit code, since eight bits (octets) would allow two four-bit patterns to efficiently encode two digits with binary-coded decimal. However, it would require all data transmission to send eight bits when seven could suffice. The committee voted to use a seven-bit code to minimize costs associated with data transmission. Since perforated tape at the time could record eight bits in one position, it also allowed for a parity bit for error checking if desired. Eight-bit machines (with octets as the native data type) that did not use parity checking typically set the eighth bit to 0.
Internal organization
The code itself was patterned so that most control codes were together and all graphic codes were together, for ease of identification. The first two so-called ASCII sticks (32 positions) were reserved for control characters. The "space" character had to come before graphics to make sorting easier, so it became position 20hex; for the same reason, many special signs commonly used as separators were placed before digits. The committee decided it was important to support uppercase 64-character alphabets, and chose to pattern ASCII so it could be reduced easily to a usable 64-character set of graphic codes, as was done in the DEC SIXBIT code (1963). Lowercase letters were therefore not interleaved with uppercase. To keep options available for lowercase letters and other graphics, the special and numeric codes were arranged before the letters, and the letter A was placed in position 41hex to match the draft of the corresponding British standard. The digits 0–9 are prefixed with 011, but the remaining 4 bits correspond to their respective values in binary, making conversion with binary-coded decimal straightforward (for example, 5 in encoded to 0110101, where 5 is 0101 in binary). | ASCII | Wikipedia | 491 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
Many of the non-alphanumeric characters were positioned to correspond to their shifted position on typewriters; an important subtlety is that these were based on mechanical typewriters, not electric typewriters. Mechanical typewriters followed the de facto standard set by the Remington No. 2 (1878), the first typewriter with a shift key, and the shifted values of 23456789- were "#$%_&'() early typewriters omitted 0 and 1, using O (capital letter o) and l (lowercase letter L) instead, but 1! and 0) pairs became standard once 0 and 1 became common. Thus, in ASCII !"#$% were placed in the second stick, positions 1–5, corresponding to the digits 1–5 in the adjacent stick. The parentheses could not correspond to 9 and 0, however, because the place corresponding to 0 was taken by the space character. This was accommodated by removing _ (underscore) from 6 and shifting the remaining characters, which corresponded to many European typewriters that placed the parentheses with 8 and 9. This discrepancy from typewriters led to bit-paired keyboards, notably the Teletype Model 33, which used the left-shifted layout corresponding to ASCII, differently from traditional mechanical typewriters.
Electric typewriters, notably the IBM Selectric (1961), used a somewhat different layout that has become de facto standard on computers following the IBM PC (1981), especially Model M (1984) and thus shift values for symbols on modern keyboards do not correspond as closely to the ASCII table as earlier keyboards did. The /? pair also dates to the No. 2, and the ,< .> pairs were used on some keyboards (others, including the No. 2, did not shift , (comma) or . (full stop) so they could be used in uppercase without unshifting). However, ASCII split the ;: pair (dating to No. 2), and rearranged mathematical symbols (varied conventions, commonly -* =+) to :* ;+ -=. | ASCII | Wikipedia | 445 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
Some then-common typewriter characters were not included, notably ½ ¼ ¢, while ^ ` ~ were included as diacritics for international use, and < > for mathematical use, together with the simple line characters \ | (in addition to common /). The @ symbol was not used in continental Europe and the committee expected it would be replaced by an accented À in the French variation, so the @ was placed in position 40hex, right before the letter A.
The control codes felt essential for data transmission were the start of message (SOM), end of address (EOA), end of message (EOM), end of transmission (EOT), "who are you?" (WRU), "are you?" (RU), a reserved device control (DC0), synchronous idle (SYNC), and acknowledge (ACK). These were positioned to maximize the Hamming distance between their bit patterns.
Character order
ASCII-code order is also called ASCIIbetical order. Collation of data is sometimes done in this order rather than "standard" alphabetical order (collating sequence). The main deviations in ASCII order are:
All uppercase come before lowercase letters; for example, "Z" precedes "a"
Digits and many punctuation marks come before letters
An intermediate order converts uppercase letters to lowercase before comparing ASCII values.
Character set
Character groups
Control characters
ASCII reserves the first 32 code points (numbers 0–31 decimal) and the last one (number 127 decimal) for control characters. These are codes intended to control peripheral devices (such as printers), or to provide meta-information about data streams, such as those stored on magnetic tape. Despite their name, these code points do not represent printable characters (i.e. they are not characters at all, but signals). For debugging purposes, "placeholder" symbols (such as those given in ISO 2047 and its predecessors) are assigned to them. | ASCII | Wikipedia | 424 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
For example, character 0x0A represents the "line feed" function (which causes a printer to advance its paper), and character 8 represents "backspace". refers to control characters that do not include carriage return, line feed or white space as non-whitespace control characters. Except for the control characters that prescribe elementary line-oriented formatting, ASCII does not define any mechanism for describing the structure or appearance of text within a document. Other schemes, such as markup languages, address page and document layout and formatting.
The original ASCII standard used only short descriptive phrases for each control character. The ambiguity this caused was sometimes intentional, for example where a character would be used slightly differently on a terminal link than on a data stream, and sometimes accidental, for example the standard is unclear about the meaning of "delete".
Probably the most influential single device affecting the interpretation of these characters was the Teletype Model 33 ASR, which was a printing terminal with an available paper tape reader/punch option. Paper tape was a very popular medium for long-term program storage until the 1980s, less costly and in some ways less fragile than magnetic tape. In particular, the Teletype Model 33 machine assignments for codes 17 (control-Q, DC1, also known as XON), 19 (control-S, DC3, also known as XOFF), and 127 (delete) became de facto standards. The Model 33 was also notable for taking the description of control-G (code 7, BEL, meaning audibly alert the operator) literally, as the unit contained an actual bell which it rang when it received a BEL character. Because the keytop for the O key also showed a left-arrow symbol (from ASCII-1963, which had this character instead of underscore), a noncompliant use of code 15 (control-O, shift in) interpreted as "delete previous character" was also adopted by many early timesharing systems but eventually became neglected. | ASCII | Wikipedia | 418 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
When a Teletype 33 ASR equipped with the automatic paper tape reader received a control-S (XOFF, an abbreviation for transmit off), it caused the tape reader to stop; receiving control-Q (XON, transmit on) caused the tape reader to resume. This so-called flow control technique became adopted by several early computer operating systems as a "handshaking" signal warning a sender to stop transmission because of impending buffer overflow; it persists to this day in many systems as a manual output control technique. On some systems, control-S retains its meaning, but control-Q is replaced by a second control-S to resume output.
The 33 ASR also could be configured to employ control-R (DC2) and control-T (DC4) to start and stop the tape punch; on some units equipped with this function, the corresponding control character lettering on the keycap above the letter was TAPE and TAPE respectively.
Delete vs backspace
The Teletype could not move its typehead backwards, so it did not have a key on its keyboard to send a BS (backspace). Instead, there was a key marked that sent code 127 (DEL). The purpose of this key was to erase mistakes in a manually-input paper tape: the operator had to push a button on the tape punch to back it up, then type the rubout, which punched all holes and replaced the mistake with a character that was intended to be ignored. Teletypes were commonly used with the less-expensive computers from Digital Equipment Corporation (DEC); these systems had to use what keys were available, and thus the DEL character was assigned to erase the previous character. Because of this, DEC video terminals (by default) sent the DEL character for the key marked "Backspace" while the separate key marked "Delete" sent an escape sequence; many other competing terminals sent a BS character for the backspace key. | ASCII | Wikipedia | 398 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
The early Unix tty drivers, unlike some modern implementations, allowed only one character to be set to erase the previous character in canonical input processing (where a very simple line editor is available); this could be set to BS or DEL, but not both, resulting in recurring situations of ambiguity where users had to decide depending on what terminal they were using (shells that allow line editing, such as ksh, bash, and zsh, understand both). The assumption that no key sent a BS character allowed Ctrl+H to be used for other purposes, such as the "help" prefix command in GNU Emacs.
Escape
Many more of the control characters have been assigned meanings quite different from their original ones. The "escape" character (ESC, code 27), for example, was intended originally to allow sending of other control characters as literals instead of invoking their meaning, an "escape sequence". This is the same meaning of "escape" encountered in URL encodings, C language strings, and other systems where certain characters have a reserved meaning. Over time this interpretation has been co-opted and has eventually been changed.
In modern usage, an ESC sent to the terminal usually indicates the start of a command sequence, which can be used to address the cursor, scroll a region, set/query various terminal properties, and more. They are usually in the form of a so-called "ANSI escape code" (often starting with a "Control Sequence Introducer", "CSI", "") from ECMA-48 (1972) and its successors. Some escape sequences do not have introducers, like the "Reset to Initial State", "RIS" command "".
In contrast, an ESC read from the terminal is most often used as an out-of-band character used to terminate an operation or special mode, as in the TECO and vi text editors. In graphical user interface (GUI) and windowing systems, ESC generally causes an application to abort its current operation or to exit (terminate) altogether. | ASCII | Wikipedia | 425 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
End of line
The inherent ambiguity of many control characters, combined with their historical usage, created problems when transferring "plain text" files between systems. The best example of this is the newline problem on various operating systems. Teletype machines required that a line of text be terminated with both "carriage return" (which moves the printhead to the beginning of the line) and "line feed" (which advances the paper one line without moving the printhead). The name "carriage return" comes from the fact that on a manual typewriter the carriage holding the paper moves while the typebars that strike the ribbon remain stationary. The entire carriage had to be pushed (returned) to the right in order to position the paper for the next line.
DEC operating systems (OS/8, RT-11, RSX-11, RSTS, TOPS-10, etc.) used both characters to mark the end of a line so that the console device (originally Teletype machines) would work. By the time so-called "glass TTYs" (later called CRTs or "dumb terminals") came along, the convention was so well established that backward compatibility necessitated continuing to follow it. When Gary Kildall created CP/M, he was inspired by some of the command line interface conventions used in DEC's RT-11 operating system.
Until the introduction of PC DOS in 1981, IBM had no influence in this because their 1970s operating systems used EBCDIC encoding instead of ASCII, and they were oriented toward punch-card input and line printer output on which the concept of "carriage return" was meaningless. IBM's PC DOS (also marketed as MS-DOS by Microsoft) inherited the convention by virtue of being loosely based on CP/M, and Windows in turn inherited it from MS-DOS. | ASCII | Wikipedia | 371 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
Requiring two characters to mark the end of a line introduces unnecessary complexity and ambiguity as to how to interpret each character when encountered by itself. To simplify matters, plain text data streams, including files, on Multics used line feed (LF) alone as a line terminator. The tty driver would handle the LF to CRLF conversion on output so files can be directly printed to terminal, and NL (newline) is often used to refer to CRLF in UNIX documents. Unix and Unix-like systems, and Amiga systems, adopted this convention from Multics. On the other hand, the original Macintosh OS, Apple DOS, and ProDOS used carriage return (CR) alone as a line terminator; however, since Apple later replaced these obsolete operating systems with their Unix-based macOS (formerly named OS X) operating system, they now use line feed (LF) as well. The Radio Shack TRS-80 also used a lone CR to terminate lines.
Computers attached to the ARPANET included machines running operating systems such as TOPS-10 and TENEX using CR-LF line endings; machines running operating systems such as Multics using LF line endings; and machines running operating systems such as OS/360 that represented lines as a character count followed by the characters of the line and which used EBCDIC rather than ASCII encoding. The Telnet protocol defined an ASCII "Network Virtual Terminal" (NVT), so that connections between hosts with different line-ending conventions and character sets could be supported by transmitting a standard text format over the network. Telnet used ASCII along with CR-LF line endings, and software using other conventions would translate between the local conventions and the NVT. The File Transfer Protocol adopted the Telnet protocol, including use of the Network Virtual Terminal, for use when transmitting commands and transferring data in the default ASCII mode. This adds complexity to implementations of those protocols, and to other network protocols, such as those used for E-mail and the World Wide Web, on systems not using the NVT's CR-LF line-ending convention. | ASCII | Wikipedia | 440 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
End of file/stream
The PDP-6 monitor, and its PDP-10 successor TOPS-10, used control-Z (SUB) as an end-of-file indication for input from a terminal. Some operating systems such as CP/M tracked file length only in units of disk blocks, and used control-Z to mark the end of the actual text in the file. For these reasons, EOF, or end-of-file, was used colloquially and conventionally as a three-letter acronym for control-Z instead of SUBstitute. The end-of-text character (ETX), also known as control-C, was inappropriate for a variety of reasons, while using control-Z as the control character to end a file is analogous to the letter Z's position at the end of the alphabet, and serves as a very convenient mnemonic aid. A historically common and still prevalent convention uses the ETX character convention to interrupt and halt a program via an input data stream, usually from a keyboard.
The Unix terminal driver uses the end-of-transmission character (EOT), also known as control-D, to indicate the end of a data stream.
In the C programming language, and in Unix conventions, the null character is used to terminate text strings; such null-terminated strings can be known in abbreviation as ASCIZ or ASCIIZ, where here Z stands for "zero".
Table of codes
Control code table
Other representations might be used by specialist equipment, for example ISO 2047 graphics or hexadecimal numbers.
Printable character table
At the time of adoption, the codes 20hex to 7Ehex would cause the printing of a visible character (a glyph), and thus were designated "printable characters". These codes represent letters, digits, punctuation marks, and a few miscellaneous symbols. There are 95 printable characters in total.
The empty space between words, as produced by the space bar of a keyboard, is character code 20hex. Since the space character is visible in printed text it considered a "printable character", even though it is unique in having no visible glyph. It is listed in the printable character table, as per the ASCII standard, instead of in the control character table.
Code 7Fhex corresponds to the non-printable "delete" (DEL) control character and is listed in the control character table. | ASCII | Wikipedia | 502 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
Earlier versions of ASCII used the up arrow instead of the caret (5Ehex) and the left arrow instead of the underscore (5Fhex).
Usage
ASCII was first used commercially during 1963 as a seven-bit teleprinter code for American Telephone & Telegraph's TWX (TeletypeWriter eXchange) network. TWX originally used the earlier five-bit ITA2, which was also used by the competing Telex teleprinter system. Bob Bemer introduced features such as the escape sequence. His British colleague Hugh McGregor Ross helped to popularize this work according to Bemer, "so much so that the code that was to become ASCII was first called the Bemer–Ross Code in Europe". Because of his extensive work on ASCII, Bemer has been called "the father of ASCII".
On March 11, 1968, US President Lyndon B. Johnson mandated that all computers purchased by the United States Federal Government support ASCII, stating:
I have also approved recommendations of the Secretary of Commerce [Luther H. Hodges] regarding standards for recording the Standard Code for Information Interchange on magnetic tapes and paper tapes when they are used in computer operations.
All computers and related equipment configurations brought into the Federal Government inventory on and after July 1, 1969, must have the capability to use the Standard Code for Information Interchange and the formats prescribed by the magnetic tape and paper tape standards when these media are used.
ASCII was the most common character encoding on the World Wide Web until December 2007, when UTF-8 encoding surpassed it; UTF-8 is backward compatible with ASCII.
Variants and derivations
As computer technology spread throughout the world, different standards bodies and corporations developed many variations of ASCII to facilitate the expression of non-English languages that used Roman-based alphabets. One could class some of these variations as "ASCII extensions", although some misuse that term to represent all variants, including those that do not preserve ASCII's character-map in the 7-bit range. Furthermore, the ASCII extensions have also been mislabelled as ASCII.
7-bit codes
From early in its development, ASCII was intended to be just one of several national variants of an international character code standard. | ASCII | Wikipedia | 479 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
Other international standards bodies have ratified character encodings such as ISO 646 (1967) that are identical or nearly identical to ASCII, with extensions for characters outside the English alphabet and symbols used outside the United States, such as the symbol for the United Kingdom's pound sterling (£); e.g. with code page 1104. Almost every country needed an adapted version of ASCII, since ASCII suited the needs of only the US and a few other countries. For example, Canada had its own version that supported French characters.
Many other countries developed variants of ASCII to include non-English letters (e.g. é, ñ, ß, Ł), currency symbols (e.g. £, ¥), etc. | ASCII | Wikipedia | 155 | 586 | https://en.wikipedia.org/wiki/ASCII | Technology | Software development: General | null |
In mathematics and statistics, the arithmetic mean ( ), arithmetic average, or just the mean or average (when the context is clear) is the sum of a collection of numbers divided by the count of numbers in the collection. The collection is often a set of results from an experiment, an observational study, or a survey. The term "arithmetic mean" is preferred in some mathematics and statistics contexts because it helps distinguish it from other types of means, such as geometric and harmonic.
In addition to mathematics and statistics, the arithmetic mean is frequently used in economics, anthropology, history, and almost every academic field to some extent. For example, per capita income is the arithmetic average income of a nation's population.
While the arithmetic mean is often used to report central tendencies, it is not a robust statistic: it is greatly influenced by outliers (values much larger or smaller than most others). For skewed distributions, such as the distribution of income for which a few people's incomes are substantially higher than most people's, the arithmetic mean may not coincide with one's notion of "middle". In that case, robust statistics, such as the median, may provide a better description of central tendency.
Definition
The arithmetic mean of a set of observed data is equal to the sum of the numerical values of each observation, divided by the total number of observations. Symbolically, for a data set consisting of the values , the arithmetic mean is defined by the formula:
(For an explanation of the summation operator, see summation.)
In simpler terms, the formula for the arithmetic mean is:
For example, if the monthly salaries of employees are , then the arithmetic mean is:
If the data set is a statistical population (i.e., consists of every possible observation and not just a subset of them), then the mean of that population is called the population mean and denoted by the Greek letter . If the data set is a statistical sample (a subset of the population), it is called the sample mean (which for a data set is denoted as ).
The arithmetic mean can be similarly defined for vectors in multiple dimensions, not only scalar values; this is often referred to as a centroid. More generally, because the arithmetic mean is a convex combination (meaning its coefficients sum to ), it can be defined on a convex space, not only a vector space. | Arithmetic mean | Wikipedia | 487 | 612 | https://en.wikipedia.org/wiki/Arithmetic%20mean | Mathematics | Statistics | null |
History
The statistician Churchill Eisenhart, senior researcher fellow at the U. S. National Bureau of Standards, traced the history of the arithmetic mean in detail. In the modern age it started to be used as a way of combining various observations that should be identical, but were not such as estimates of the direction of magnetic north.
In 1635 the mathematician Henry Gellibrand described as “meane” the midpoint of a lowest and highest number, not quite the arithmetic mean. In 1668, a person known as “DB” was quoted in the Transactions of the Royal Society describing “taking the mean” of five values:
Motivating properties
The arithmetic mean has several properties that make it interesting, especially as a measure of central tendency. These include:
If numbers have mean , then . Since is the distance from a given number to the mean, one way to interpret this property is by saying that the numbers to the left of the mean are balanced by the numbers to the right. The mean is the only number for which the residuals (deviations from the estimate) sum to zero. This can also be interpreted as saying that the mean is translationally invariant in the sense that for any real number , .
If it is required to use a single number as a "typical" value for a set of known numbers , then the arithmetic mean of the numbers does this best since it minimizes the sum of squared deviations from the typical value: the sum of . The sample mean is also the best single predictor because it has the lowest root mean squared error. If the arithmetic mean of a population of numbers is desired, then the estimate of it that is unbiased is the arithmetic mean of a sample drawn from the population.
The arithmetic mean is independent of scale of the units of measurement, in the sense that So, for example, calculating a mean of liters and then converting to gallons is the same as converting to gallons first and then calculating the mean. This is also called first order homogeneity.
Additional properties
The arithmetic mean of a sample is always between the largest and smallest values in that sample.
The arithmetic mean of any amount of equal-sized number groups together is the arithmetic mean of the arithmetic means of each group.
Contrast with median | Arithmetic mean | Wikipedia | 457 | 612 | https://en.wikipedia.org/wiki/Arithmetic%20mean | Mathematics | Statistics | null |
The arithmetic mean may be contrasted with the median. The median is defined such that no more than half the values are larger, and no more than half are smaller than it. If elements in the data increase arithmetically when placed in some order, then the median and arithmetic average are equal. For example, consider the data sample . The mean is , as is the median. However, when we consider a sample that cannot be arranged to increase arithmetically, such as , the median and arithmetic average can differ significantly. In this case, the arithmetic average is , while the median is . The average value can vary considerably from most values in the sample and can be larger or smaller than most.
There are applications of this phenomenon in many fields. For example, since the 1980s, the median income in the United States has increased more slowly than the arithmetic average of income.
Generalizations
Weighted average
A weighted average, or weighted mean, is an average in which some data points count more heavily than others in that they are given more weight in the calculation. For example, the arithmetic mean of and is , or equivalently . In contrast, a weighted mean in which the first number receives, for example, twice as much weight as the second (perhaps because it is assumed to appear twice as often in the general population from which these numbers were sampled) would be calculated as . Here the weights, which necessarily sum to one, are and , the former being twice the latter. The arithmetic mean (sometimes called the "unweighted average" or "equally weighted average") can be interpreted as a special case of a weighted average in which all weights are equal to the same number ( in the above example and in a situation with numbers being averaged).
Continuous probability distributions | Arithmetic mean | Wikipedia | 352 | 612 | https://en.wikipedia.org/wiki/Arithmetic%20mean | Mathematics | Statistics | null |
If a numerical property, and any sample of data from it, can take on any value from a continuous range instead of, for example, just integers, then the probability of a number falling into some range of possible values can be described by integrating a continuous probability distribution across this range, even when the naive probability for a sample number taking one certain value from infinitely many is zero. In this context, the analog of a weighted average, in which there are infinitely many possibilities for the precise value of the variable in each range, is called the mean of the probability distribution. The most widely encountered probability distribution is called the normal distribution; it has the property that all measures of its central tendency, including not just the mean but also the median mentioned above and the mode (the three Ms), are equal. This equality does not hold for other probability distributions, as illustrated for the log-normal distribution here.
Angles
Particular care is needed when using cyclic data, such as phases or angles. Taking the arithmetic mean of 1° and 359° yields a result of 180°.
This is incorrect for two reasons:
Firstly, angle measurements are only defined up to an additive constant of 360° ( or , if measuring in radians). Thus, these could easily be called 1° and -1°, or 361° and 719°, since each one of them produces a different average.
Secondly, in this situation, 0° (or 360°) is geometrically a better average value: there is lower dispersion about it (the points are both 1° from it and 179° from 180°, the putative average).
In general application, such an oversight will lead to the average value artificially moving towards the middle of the numerical range. A solution to this problem is to use the optimization formulation (that is, define the mean as the central point: the point about which one has the lowest dispersion) and redefine the difference as a modular distance (i.e., the distance on the circle: so the modular distance between 1° and 359° is 2°, not 358°).
Symbols and encoding
The arithmetic mean is often denoted by a bar (vinculum or macron), as in .
Some software (text processors, web browsers) may not display the "x̄" symbol correctly. For example, the HTML symbol "x̄" combines two codes — the base letter "x" plus a code for the line above ( ̄ or ¯). | Arithmetic mean | Wikipedia | 509 | 612 | https://en.wikipedia.org/wiki/Arithmetic%20mean | Mathematics | Statistics | null |
In some document formats (such as PDF), the symbol may be replaced by a "¢" (cent) symbol when copied to a text processor such as Microsoft Word. | Arithmetic mean | Wikipedia | 35 | 612 | https://en.wikipedia.org/wiki/Arithmetic%20mean | Mathematics | Statistics | null |
Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all tetrapods excluding the amniotes (tetrapods with an amniotic membrane, such as modern reptiles, birds and mammals). All extant (living) amphibians belong to the monophyletic subclass Lissamphibia, with three living orders: Anura (frogs and toads), Urodela (salamanders), and Gymnophiona (caecilians). Evolved to be mostly semiaquatic, amphibians have adapted to inhabit a wide variety of habitats, with most species living in freshwater, wetland or terrestrial ecosystems (such as riparian woodland, fossorial and even arboreal habitats). Their life cycle typically starts out as aquatic larvae with gills known as tadpoles, but some species have developed behavioural adaptations to bypass this.
Young amphibians generally undergo metamorphosis from an aquatic larval form with gills to an air-breathing adult form with lungs. Amphibians use their skin as a secondary respiratory interface and some small terrestrial salamanders and frogs lack lungs and rely entirely on their skin. They are superficially similar to reptiles like lizards, but unlike reptiles and other amniotes, require access to water bodies to breed. With their complex reproductive needs and permeable skins, amphibians are often ecological indicators to habitat conditions; in recent decades there has been a dramatic decline in amphibian populations for many species around the globe. | Amphibian | Wikipedia | 339 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The earliest amphibians evolved in the Devonian period from tetrapodomorph sarcopterygians (lobe-finned fish with articulated limb-like fins) that evolved primitive lungs, which were helpful in adapting to dry land. They diversified and became ecologically dominant during the Carboniferous and Permian periods, but were later displaced in terrestrial environments by early reptiles and basal synapsids (predecessors of mammals). The origin of modern lissamphibians, which first appeared during the Early Triassic, around 250 million years ago, has long been contentious. The most popular hypothesis is that they likely originated from temnospondyls, the most diverse group of prehistoric amphibians, during the Permian period. Another hypothesis is that they emerged from lepospondyls. A fourth group of lissamphibians, the Albanerpetontidae, became extinct around 2 million years ago.
The number of known amphibian species is approximately 8,000, of which nearly 90% are frogs. The smallest amphibian (and vertebrate) in the world is a frog from New Guinea (Paedophryne amauensis) with a length of just . The largest living amphibian is the South China giant salamander (Andrias sligoi), but this is dwarfed by prehistoric temnospondyls such as Mastodonsaurus which could reach up to in length. The study of amphibians is called batrachology, while the study of both reptiles and amphibians is called herpetology.
Classification
The word amphibian is derived from the Ancient Greek term (), which means 'both kinds of life', meaning 'of both kinds' and meaning 'life'. The term was initially used as a general adjective for animals that could live on land or in water, including seals and otters. Traditionally, the class Amphibia includes all tetrapod vertebrates that are not amniotes. Amphibia in its widest sense () was divided into three subclasses, two of which are extinct: | Amphibian | Wikipedia | 433 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Subclass Lepospondyli† (A potentially polyphyletic Late Paleozoic group of small forms, likely more closely related to amniotes than Lissamphibia)
Subclass Temnospondyli† (diverse Late Paleozoic and early Mesozoic grade, some of which were large predators)
Subclass Lissamphibia (all modern amphibians, including frogs, toads, salamanders, newts and caecilians)
Salientia (frogs, toads and relatives): Early Triassic to present—7,360 current species in 53 families. Modern (crown group) salientians are described via the name Anura.
Caudata (salamanders, newts and relatives): Late Triassic to present—764 current species in 9 families. Modern (crown group) caudatans are described via the name Urodela.
Gymnophiona (caecilians and relatives): Late Triassic to present—215 current species in 10 families. The name Apoda is also sometimes used for caecilians.
Allocaudata† (Albanerpetontidae) Middle Jurassic – Early Pleistocene
These three subclasses do not include all extinct amphibians. Other extinct amphibian groups include Embolomeri (Late Paleozoic large aquatic predators), Seymouriamorpha (semiaquatic to terrestrial Permian forms related to amniotes)[citation needed], among others. Names such as Tetrapoda and Stegocephalia encompass the entirety of amphibian-grade tetrapods, while Reptiliomorpha or Anthracosauria are variably used to describe extinct amphibians more closely related to amniotes than to lissamphibians.
The actual number of species in each group depends on the taxonomic classification followed. The two most common systems are the classification adopted by the website AmphibiaWeb, University of California, Berkeley, and the classification by herpetologist Darrel Frost and the American Museum of Natural History, available as the online reference database "Amphibian Species of the World". The numbers of species cited above follows Frost and the total number of known (living) amphibian species as of March 31, 2019, is exactly 8,000, of which nearly 90% are frogs. | Amphibian | Wikipedia | 485 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
With the phylogenetic classification, the taxon Labyrinthodontia has been discarded as it is a polyparaphyletic group without unique defining features apart from shared primitive characteristics. Classification varies according to the preferred phylogeny of the author and whether they use a stem-based or a node-based classification. Traditionally, amphibians as a class are defined as all tetrapods with a larval stage, while the group that includes the common ancestors of all living amphibians (frogs, salamanders and caecilians) and all their descendants is called Lissamphibia. The phylogeny of Paleozoic amphibians is uncertain, and Lissamphibia may possibly fall within extinct groups, like the Temnospondyli (traditionally placed in the subclass Labyrinthodontia) or the Lepospondyli, and in some analyses even in the amniotes. This means that advocates of phylogenetic nomenclature have removed a large number of basal Devonian and Carboniferous amphibian-type tetrapod groups that were formerly placed in Amphibia in Linnaean taxonomy, and included them elsewhere under cladistic taxonomy. If the common ancestor of amphibians and amniotes is included in Amphibia, it becomes a paraphyletic group.
All modern amphibians are included in the subclass Lissamphibia, which is usually considered a clade, a group of species that have evolved from a common ancestor. The three modern orders are Anura (the frogs), Caudata (or Urodela, the salamanders), and Gymnophiona (or Apoda, the caecilians). It has been suggested that salamanders arose separately from a temnospondyl-like ancestor, and even that caecilians are the sister group of the advanced reptiliomorph amphibians, and thus of amniotes. Although the fossils of several older proto-frogs with primitive characteristics are known, the oldest "true frog", with hopping adaptations is Prosalirus bitis, from the Early Jurassic Kayenta Formation of Arizona. It is anatomically very similar to modern frogs. The oldest known caecilians are Funcusvermis gilmorei (from the Late Triassic) and Eocaecilia micropodia (from the Early Jurassic), both from Arizona. The earliest salamander is Beiyanerpeton jianpingensis from the Late Jurassic of northeastern China. | Amphibian | Wikipedia | 509 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Authorities disagree as to whether Salientia is a superorder that includes the order Anura, or whether Anura is a sub-order of the order Salientia. The Lissamphibia are traditionally divided into three orders, but an extinct salamander-like family, the Albanerpetontidae, is now considered part of Lissamphibia alongside the superorder Salientia. Furthermore, Salientia includes all three recent orders plus the Triassic proto-frog, Triadobatrachus.
Evolutionary history
The first major groups of amphibians developed in the Devonian period, around 370 million years ago, from lobe-finned fish which were similar to the modern coelacanth and lungfish. These ancient lobe-finned fish had evolved multi-jointed leg-like fins with digits that enabled them to crawl along the sea bottom. Some fish had developed primitive lungs that help them breathe air when the stagnant pools of the Devonian swamps were low in oxygen. They could also use their strong fins to hoist themselves out of the water and onto dry land if circumstances so required. Eventually, their bony fins would evolve into limbs and they would become the ancestors to all tetrapods, including modern amphibians, reptiles, birds, and mammals. Despite being able to crawl on land, many of these prehistoric tetrapodomorph fish still spent most of their time in the water. They had started to develop lungs, but still breathed predominantly with gills. | Amphibian | Wikipedia | 306 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Many examples of species showing transitional features have been discovered. Ichthyostega was one of the first primitive amphibians, with nostrils and more efficient lungs. It had four sturdy limbs, a neck, a tail with fins and a skull very similar to that of the lobe-finned fish, Eusthenopteron. Amphibians evolved adaptations that allowed them to stay out of the water for longer periods. Their lungs improved and their skeletons became heavier and stronger, better able to support the weight of their bodies on land. They developed "hands" and "feet" with five or more digits; the skin became more capable of retaining body fluids and resisting desiccation. The fish's hyomandibula bone in the hyoid region behind the gills diminished in size and became the stapes of the amphibian ear, an adaptation necessary for hearing on dry land. An affinity between the amphibians and the teleost fish is the multi-folded structure of the teeth and the paired supra-occipital bones at the back of the head, neither of these features being found elsewhere in the animal kingdom. | Amphibian | Wikipedia | 234 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
At the end of the Devonian period (360 million years ago), the seas, rivers and lakes were teeming with life while the land was the realm of early plants and devoid of vertebrates, though some, such as Ichthyostega, may have sometimes hauled themselves out of the water. It is thought they may have propelled themselves with their forelimbs, dragging their hindquarters in a similar manner to that used by the elephant seal. In the early Carboniferous (360 to 323 million years ago), the climate was relatively wet and warm. Extensive swamps developed with mosses, ferns, horsetails and calamites. Air-breathing arthropods evolved and invaded the land where they provided food for the carnivorous amphibians that began to adapt to the terrestrial environment. There were no other tetrapods on the land and the amphibians were at the top of the food chain, with some occupying ecological positions currently held by crocodiles. Though equipped with limbs and the ability to breathe air, most still had a long tapering body and strong tail. Others were the top land predators, sometimes reaching several metres in length, preying on the large insects of the period and the many types of fish in the water. They still needed to return to water to lay their shell-less eggs, and even most modern amphibians have a fully aquatic larval stage with gills like their fish ancestors. It was the development of the amniotic egg, which prevents the developing embryo from drying out, that enabled the reptiles to reproduce on land and which led to their dominance in the period that followed.
After the Carboniferous rainforest collapse amphibian dominance gave way to reptiles, and amphibians were further devastated by the Permian–Triassic extinction event. During the Triassic Period (252 to 201 million years ago), the reptiles continued to out-compete the amphibians, leading to a reduction in both the amphibians' size and their importance in the biosphere. According to the fossil record, Lissamphibia, which includes all modern amphibians and is the only surviving lineage, may have branched off from the extinct groups Temnospondyli and Lepospondyli at some period between the Late Carboniferous and the Early Triassic. The relative scarcity of fossil evidence precludes precise dating, but the most recent molecular study, based on multilocus sequence typing, suggests a Late Carboniferous/Early Permian origin for extant amphibians. | Amphibian | Wikipedia | 510 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The origins and evolutionary relationships between the three main groups of amphibians is a matter of debate. A 2005 molecular phylogeny, based on rDNA analysis, suggests that salamanders and caecilians are more closely related to each other than they are to frogs. It also appears that the divergence of the three groups took place in the Paleozoic or early Mesozoic (around 250 million years ago), before the breakup of the supercontinent Pangaea and soon after their divergence from the lobe-finned fish. The briefness of this period, and the swiftness with which radiation took place, would help account for the relative scarcity of primitive amphibian fossils. There are large gaps in the fossil record, the discovery of the dissorophoid temnospondyl Gerobatrachus from the Early Permian in Texas in 2008 provided a missing link with many of the characteristics of modern frogs. Molecular analysis suggests that the frog–salamander divergence took place considerably earlier than the palaeontological evidence indicates. One study suggested that the last common ancestor of all modern amphibians lived about 315 million years ago, and that stereospondyl temnospondyls are the closest relatives to the caecilians. However, most studies support a single monophyletic origin of all modern amphibians within the dissorophoid temnospondyls.
As they evolved from lunged fish, amphibians had to make certain adaptations for living on land, including the need to develop new means of locomotion. In the water, the sideways thrusts of their tails had propelled them forward, but on land, quite different mechanisms were required. Their vertebral columns, limbs, limb girdles and musculature needed to be strong enough to raise them off the ground for locomotion and feeding. Terrestrial adults discarded their lateral line systems and adapted their sensory systems to receive stimuli via the medium of the air. They needed to develop new methods to regulate their body heat to cope with fluctuations in ambient temperature. They developed behaviours suitable for reproduction in a terrestrial environment. Their skins were exposed to harmful ultraviolet rays that had previously been absorbed by the water. The skin changed to become more protective and prevent excessive water loss. | Amphibian | Wikipedia | 474 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Characteristics
The superclass Tetrapoda is divided into four classes of vertebrate animals with four limbs. Reptiles, birds and mammals are amniotes, the eggs of which are either laid or carried by the female and are surrounded by several membranes, some of which are impervious. Lacking these membranes, amphibians require water bodies for reproduction, although some species have developed various strategies for protecting or bypassing the vulnerable aquatic larval stage. They are not found in the sea with the exception of one or two frogs that live in brackish water in mangrove swamps; the Anderson's salamander meanwhile occurs in brackish or salt water lakes. On land, amphibians are restricted to moist habitats because of the need to keep their skin damp.
Modern amphibians have a simplified anatomy compared to their ancestors due to paedomorphosis, caused by two evolutionary trends: miniaturization and an unusually large genome, which result in a slower growth and development rate compared to other vertebrates. Another reason for their size is associated with their rapid metamorphosis, which seems to have evolved only in the ancestors of Lissamphibia; in all other known lines the development was much more gradual. Because a remodeling of the feeding apparatus means they do not eat during the metamorphosis, the metamorphosis has to go faster the smaller the individual is, so it happens at an early stage when the larvae are still small. (The largest species of salamanders do not go through a metamorphosis.) Amphibians that lay eggs on land often go through the whole metamorphosis inside the egg. An anamniotic terrestrial egg is less than 1 cm in diameter due to diffusion problems, a size which puts a limit on the amount of posthatching growth. | Amphibian | Wikipedia | 375 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The smallest amphibian (and vertebrate) in the world is a microhylid frog from New Guinea (Paedophryne amauensis) first discovered in 2012. It has an average length of and is part of a genus that contains four of the world's ten smallest frog species. The largest living amphibian is the Chinese giant salamander (Andrias davidianus) but this is a great deal smaller than the largest amphibian that ever existed—the extinct Prionosuchus, a crocodile-like temnospondyl dating to 270 million years ago from the middle Permian of Brazil. The largest frog is the African Goliath frog (Conraua goliath), which can reach and weigh .
Amphibians are ectothermic (cold-blooded) vertebrates that do not maintain their body temperature through internal physiological processes. Their metabolic rate is low and as a result, their food and energy requirements are limited. In the adult state, they have tear ducts and movable eyelids, and most species have ears that can detect airborne or ground vibrations. They have muscular tongues, which in many species can be protruded. Modern amphibians have fully ossified vertebrae with articular processes. Their ribs are usually short and may be fused to the vertebrae. Their skulls are mostly broad and short, and are often incompletely ossified. Their skin contains little keratin and lacks scales, apart from a few fish-like scales in certain caecilians. The skin contains many mucous glands and in some species, poison glands (a type of granular gland). The hearts of amphibians have three chambers, two atria and one ventricle. They have a urinary bladder and nitrogenous waste products are excreted primarily as urea. Most amphibians lay their eggs in water and have aquatic larvae that undergo metamorphosis to become terrestrial adults. Amphibians breathe by means of a pump action in which air is first drawn into the buccopharyngeal region through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat. They supplement this with gas exchange through the skin.
Anura | Amphibian | Wikipedia | 463 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The order Anura (from the Ancient Greek a(n)- meaning "without" and oura meaning "tail") comprises the frogs and toads. They usually have long hind limbs that fold underneath them, shorter forelimbs, webbed toes with no claws, no tails, large eyes and glandular moist skin. Members of this order with smooth skins are commonly referred to as frogs, while those with warty skins are known as toads. The difference is not a formal one taxonomically and there are numerous exceptions to this rule. Members of the family Bufonidae are known as the "true toads". Frogs range in size from the Goliath frog (Conraua goliath) of West Africa to the Paedophryne amauensis, first described in Papua New Guinea in 2012, which is also the smallest known vertebrate. Although most species are associated with water and damp habitats, some are specialised to live in trees or in deserts. They are found worldwide except for polar areas.
Anura is divided into three suborders that are broadly accepted by the scientific community, but the relationships between some families remain unclear. Future molecular studies should provide further insights into their evolutionary relationships. The suborder Archaeobatrachia contains four families of primitive frogs. These are Ascaphidae, Bombinatoridae, Discoglossidae and Leiopelmatidae which have few derived features and are probably paraphyletic with regard to other frog lineages. The six families in the more evolutionarily advanced suborder Mesobatrachia are the fossorial Megophryidae, Pelobatidae, Pelodytidae, Scaphiopodidae and Rhinophrynidae and the obligatorily aquatic Pipidae. These have certain characteristics that are intermediate between the two other suborders. Neobatrachia is by far the largest suborder and includes the remaining families of modern frogs, including most common species. Approximately 96% of the over 5,000 extant species of frog are neobatrachians.
Caudata | Amphibian | Wikipedia | 431 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The order Caudata (from the Latin cauda meaning "tail") consists of the salamanders—elongated, low-slung animals that mostly resemble lizards in form. This is a symplesiomorphic trait and they are no more closely related to lizards than they are to mammals. Salamanders lack claws, have scale-free skins, either smooth or covered with tubercles, and tails that are usually flattened from side to side and often finned. They range in size from the Chinese giant salamander (Andrias davidianus), which has been reported to grow to a length of , to the diminutive Thorius pennatulus from Mexico which seldom exceeds in length. Salamanders have a mostly Laurasian distribution, being present in much of the Holarctic region of the northern hemisphere. The family Plethodontidae is also found in Central America and South America north of the Amazon basin; South America was apparently invaded from Central America by about the start of the Miocene, 23 million years ago. Urodela is a name sometimes used for all the extant species of salamanders. Members of several salamander families have become paedomorphic and either fail to complete their metamorphosis or retain some larval characteristics as adults. Most salamanders are under long. They may be terrestrial or aquatic and many spend part of the year in each habitat. When on land, they mostly spend the day hidden under stones or logs or in dense vegetation, emerging in the evening and night to forage for worms, insects and other invertebrates.
The suborder Cryptobranchoidea contains the primitive salamanders. A number of fossil cryptobranchids have been found, but there are only three living species, the Chinese giant salamander (Andrias davidianus), the Japanese giant salamander (Andrias japonicus) and the hellbender (Cryptobranchus alleganiensis) from North America. These large amphibians retain several larval characteristics in their adult state; gills slits are present and the eyes are unlidded. A unique feature is their ability to feed by suction, depressing either the left side of their lower jaw or the right. The males excavate nests, persuade females to lay their egg strings inside them, and guard them. As well as breathing with lungs, they respire through the many folds in their thin skin, which has capillaries close to the surface. | Amphibian | Wikipedia | 509 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The suborder Salamandroidea contains the advanced salamanders. They differ from the cryptobranchids by having fused prearticular bones in the lower jaw, and by using internal fertilisation. In salamandrids, the male deposits a bundle of sperm, the spermatophore, and the female picks it up and inserts it into her cloaca where the sperm is stored until the eggs are laid. The largest family in this group is Plethodontidae, the lungless salamanders, which includes 60% of all salamander species. The family Salamandridae includes the true salamanders and the name "newt" is given to members of its subfamily Pleurodelinae.
The third suborder, Sirenoidea, contains the four species of sirens, which are in a single family, Sirenidae. Members of this order are eel-like aquatic salamanders with much reduced forelimbs and no hind limbs. Some of their features are primitive while others are derived. Fertilisation is likely to be external as sirenids lack the cloacal glands used by male salamandrids to produce spermatophores and the females lack spermathecae for sperm storage. Despite this, the eggs are laid singly, a behaviour not conducive for external fertilisation.
Gymnophiona | Amphibian | Wikipedia | 285 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The order Gymnophiona (from the Greek gymnos meaning "naked" and ophis meaning "serpent") or Apoda comprises the caecilians. These are long, cylindrical, limbless animals with a snake- or worm-like form. The adults vary in length from 8 to 75 centimetres (3 to 30 inches) with the exception of Thomson's caecilian (Caecilia thompsoni), which can reach . A caecilian's skin has a large number of transverse folds and in some species contains tiny embedded dermal scales. It has rudimentary eyes covered in skin, which are probably limited to discerning differences in light intensity. It also has a pair of short tentacles near the eye that can be extended and which have tactile and olfactory functions. Most caecilians live underground in burrows in damp soil, in rotten wood and under plant debris, but some are aquatic. Most species lay their eggs underground and when the larvae hatch, they make their way to adjacent bodies of water. Others brood their eggs and the larvae undergo metamorphosis before the eggs hatch. A few species give birth to live young, nourishing them with glandular secretions while they are in the oviduct. Caecilians have a mostly Gondwanan distribution, being found in tropical regions of Africa, Asia and Central and South America.
Anatomy and physiology
Skin
The integumentary structure contains some typical characteristics common to terrestrial vertebrates, such as the presence of highly cornified outer layers, renewed periodically through a moulting process controlled by the pituitary and thyroid glands. Local thickenings (often called warts) are common, such as those found on toads. The outside of the skin is shed periodically mostly in one piece, in contrast to mammals and birds where it is shed in flakes. Amphibians often eat the sloughed skin. Caecilians are unique among amphibians in having mineralized dermal scales embedded in the dermis between the furrows in the skin. The similarity of these to the scales of bony fish is largely superficial. Lizards and some frogs have somewhat similar osteoderms forming bony deposits in the dermis, but this is an example of convergent evolution with similar structures having arisen independently in diverse vertebrate lineages. | Amphibian | Wikipedia | 485 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Amphibian skin is permeable to water. Gas exchange can take place through the skin (cutaneous respiration) and this allows adult amphibians to respire without rising to the surface of water and to hibernate at the bottom of ponds. To compensate for their thin and delicate skin, amphibians have evolved mucous glands, principally on their heads, backs and tails. The secretions produced by these help keep the skin moist. In addition, most species of amphibian have granular glands that secrete distasteful or poisonous substances. Some amphibian toxins can be lethal to humans while others have little effect. The main poison-producing glands, the parotoids, produce the neurotoxin bufotoxin and are located behind the ears of toads, along the backs of frogs, behind the eyes of salamanders and on the upper surface of caecilians.
The skin colour of amphibians is produced by three layers of pigment cells called chromatophores. These three cell layers consist of the melanophores (occupying the deepest layer), the guanophores (forming an intermediate layer and containing many granules, producing a blue-green colour) and the lipophores (yellow, the most superficial layer). The colour change displayed by many species is initiated by hormones secreted by the pituitary gland. Unlike bony fish, there is no direct control of the pigment cells by the nervous system, and this results in the colour change taking place more slowly than happens in fish. A vividly coloured skin usually indicates that the species is toxic and is a warning sign to predators.
Skeletal system and locomotion
Amphibians have a skeletal system that is structurally homologous to other tetrapods, though with a number of variations. They all have four limbs except for the legless caecilians and a few species of salamander with reduced or no limbs. The bones are hollow and lightweight. The musculoskeletal system is strong to enable it to support the head and body. The bones are fully ossified and the vertebrae interlock with each other by means of overlapping processes. The pectoral girdle is supported by muscle, and the well-developed pelvic girdle is attached to the backbone by a pair of sacral ribs. The ilium slopes forward and the body is held closer to the ground than is the case in mammals. | Amphibian | Wikipedia | 506 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In most amphibians, there are four digits on the fore foot and five on the hind foot, but no claws on either. Some salamanders have fewer digits and the amphiumas are eel-like in appearance with tiny, stubby legs. The sirens are aquatic salamanders with stumpy forelimbs and no hind limbs. The caecilians are limbless. They burrow in the manner of earthworms with zones of muscle contractions moving along the body. On the surface of the ground or in water they move by undulating their body from side to side.
In frogs, the hind legs are larger than the fore legs, especially so in those species that principally move by jumping or swimming. In the walkers and runners the hind limbs are not so large, and the burrowers mostly have short limbs and broad bodies. The feet have adaptations for the way of life, with webbing between the toes for swimming, broad adhesive toe pads for climbing, and keratinised tubercles on the hind feet for digging (frogs usually dig backwards into the soil). In most salamanders, the limbs are short and more or less the same length and project at right angles from the body. Locomotion on land is by walking and the tail often swings from side to side or is used as a prop, particularly when climbing. In their normal gait, only one leg is advanced at a time in the manner adopted by their ancestors, the lobe-finned fish. Some salamanders in the genus Aneides and certain plethodontids climb trees and have long limbs, large toepads and prehensile tails. In aquatic salamanders and in frog tadpoles, the tail has dorsal and ventral fins and is moved from side to side as a means of propulsion. Adult frogs do not have tails and caecilians have only very short ones.
Salamanders use their tails in defence and some are prepared to jettison them to save their lives in a process known as autotomy. Certain species in the Plethodontidae have a weak zone at the base of the tail and use this strategy readily. The tail often continues to twitch after separation which may distract the attacker and allow the salamander to escape. Both tails and limbs can be regenerated. Adult frogs are unable to regrow limbs but tadpoles can do so.
Circulatory system | Amphibian | Wikipedia | 498 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Amphibians have a juvenile stage and an adult stage, and the circulatory systems of the two are distinct. In the juvenile (or tadpole) stage, the circulation is similar to that of a fish; the two-chambered heart pumps the blood through the gills where it is oxygenated, and is spread around the body and back to the heart in a single loop. In the adult stage, amphibians (especially frogs) lose their gills and develop lungs. They have a heart that consists of a single ventricle and two atria. When the ventricle starts contracting, deoxygenated blood is pumped through the pulmonary artery to the lungs. Continued contraction then pumps oxygenated blood around the rest of the body. Mixing of the two bloodstreams is minimized by the anatomy of the chambers.
Nervous and sensory systems
The nervous system is basically the same as in other vertebrates, with a central brain, a spinal cord, and nerves throughout the body. The amphibian brain is relatively simple but broadly the same structurally as in reptiles, birds and mammals. Their brains are elongated, except in caecilians, and contain the usual motor and sensory areas of tetrapods. The pineal body, known to regulate sleep patterns in humans, is thought to produce the hormones involved in hibernation and aestivation in amphibians. | Amphibian | Wikipedia | 282 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Tadpoles retain the lateral line system of their ancestral fishes, but this is lost in terrestrial adult amphibians. Many aquatic salamanders and some caecilians possess electroreceptors called ampullary organs (completely absent in anurans), that allow them to locate objects around them when submerged in water. The ears are well developed in frogs. There is no external ear, but the large circular eardrum lies on the surface of the head just behind the eye. This vibrates and sound is transmitted through a single bone, the stapes, to the inner ear. Only high-frequency sounds like mating calls are heard in this way, but low-frequency noises can be detected through another mechanism. There is a patch of specialized haircells, called papilla amphibiorum, in the inner ear capable of detecting deeper sounds. Another feature, unique to frogs and salamanders, is the columella-operculum complex adjoining the auditory capsule which is involved in the transmission of both airborne and seismic signals. The ears of salamanders and caecilians are less highly developed than those of frogs as they do not normally communicate with each other through the medium of sound.
The eyes of tadpoles lack lids, but at metamorphosis, the cornea becomes more dome-shaped, the lens becomes flatter, and eyelids and associated glands and ducts develop. The adult eyes are an improvement on invertebrate eyes and were a first step in the development of more advanced vertebrate eyes. They allow colour vision and depth of focus. In the retinas are green rods, which are receptive to a wide range of wavelengths.
Digestive and excretory systems
Many amphibians catch their prey by flicking out an elongated tongue with a sticky tip and drawing it back into the mouth before seizing the item with their jaws. Some use inertial feeding to help them swallow the prey, repeatedly thrusting their head forward sharply causing the food to move backwards in their mouth by inertia. Most amphibians swallow their prey whole without much chewing so they possess voluminous stomachs. The short oesophagus is lined with cilia that help to move the food to the stomach and mucus produced by glands in the mouth and pharynx eases its passage. The enzyme chitinase produced in the stomach helps digest the chitinous cuticle of arthropod prey. | Amphibian | Wikipedia | 502 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Amphibians possess a pancreas, liver and gall bladder. The liver is usually large with two lobes. Its size is determined by its function as a glycogen and fat storage unit, and may change with the seasons as these reserves are built or used up. Adipose tissue is another important means of storing energy and this occurs in the abdomen (in internal structures called fat bodies), under the skin and, in some salamanders, in the tail.
There are two kidneys located dorsally, near the roof of the body cavity. Their job is to filter the blood of metabolic waste and transport the urine via ureters to the urinary bladder where it is stored before being passed out periodically through the cloacal vent. Larvae and most aquatic adult amphibians excrete the nitrogen as ammonia in large quantities of dilute urine, while terrestrial species, with a greater need to conserve water, excrete the less toxic product urea. Some tree frogs with limited access to water excrete most of their metabolic waste as uric acid.
Urinary bladder
Most aquatic and semi-aquatic amphibians have a membranous skin which allows them to absorb water directly through it. Some semi-aquatic animals also have similarly permeable bladder membrane. As a result, they tend to have high rates of urine production to offset this high water intake, and have urine which is low in dissolved salts. The urinary bladder assists such animals to retain salts. Some aquatic amphibian such as Xenopus do not reabsorb water, to prevent excessive water influx. For land-dwelling amphibians, dehydration results in reduced urine output.
The amphibian bladder is usually highly distensible and among some land-dwelling species of frogs and salamanders may account for between 20% and 50% of their total body weight. Urine flows from the kidneys through the ureters into the bladder and is periodically released from the bladder to the cloaca.
Respiratory system | Amphibian | Wikipedia | 415 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The lungs in amphibians are primitive compared to those of amniotes, possessing few internal septa and large alveoli, and consequently having a comparatively slow diffusion rate for oxygen entering the blood. Ventilation is accomplished by buccal pumping. Most amphibians, however, are able to exchange gases with the water or air via their skin. To enable sufficient cutaneous respiration, the surface of their highly vascularised skin must remain moist to allow the oxygen to diffuse at a sufficiently high rate. Because oxygen concentration in the water increases at both low temperatures and high flow rates, aquatic amphibians in these situations can rely primarily on cutaneous respiration, as in the Titicaca water frog and the hellbender salamander. In air, where oxygen is more concentrated, some small species can rely solely on cutaneous gas exchange, most famously the plethodontid salamanders, which have neither lungs nor gills. Many aquatic salamanders and all tadpoles have gills in their larval stage, with some (such as the axolotl) retaining gills as aquatic adults.
Reproduction
For the purpose of reproduction, most amphibians require fresh water although some lay their eggs on land and have developed various means of keeping them moist. A few (e.g. Fejervarya raja) can inhabit brackish water, but there are no true marine amphibians. There are reports, however, of particular amphibian populations unexpectedly invading marine waters. Such was the case with the Black Sea invasion of the natural hybrid Pelophylax esculentus reported in 2010.
Several hundred frog species in adaptive radiations (e.g., Eleutherodactylus, the Pacific Platymantis, the Australo-Papuan microhylids, and many other tropical frogs), however, do not need any water for breeding in the wild. They reproduce via direct development, an ecological and evolutionary adaptation that has allowed them to be completely independent from free-standing water. Almost all of these frogs live in wet tropical rainforests and their eggs hatch directly into miniature versions of the adult, passing through the tadpole stage within the egg. Reproductive success of many amphibians is dependent not only on the quantity of rainfall, but the seasonal timing. | Amphibian | Wikipedia | 468 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In the tropics, many amphibians breed continuously or at any time of year. In temperate regions, breeding is mostly seasonal, usually in the spring, and is triggered by increasing day length, rising temperatures or rainfall. Experiments have shown the importance of temperature, but the trigger event, especially in arid regions, is often a storm. In anurans, males usually arrive at the breeding sites before females and the vocal chorus they produce may stimulate ovulation in females and the endocrine activity of males that are not yet reproductively active.
In caecilians, fertilisation is internal, the male extruding an intromittent organ, the , and inserting it into the female cloaca. The paired Müllerian glands inside the male cloaca secrete a fluid which resembles that produced by mammalian prostate glands and which may transport and nourish the sperm. Fertilisation probably takes place in the oviduct.
The majority of salamanders also engage in internal fertilisation. In most of these, the male deposits a spermatophore, a small packet of sperm on top of a gelatinous cone, on the substrate either on land or in the water. The female takes up the sperm packet by grasping it with the lips of the cloaca and pushing it into the vent. The spermatozoa move to the spermatheca in the roof of the cloaca where they remain until ovulation which may be many months later. Courtship rituals and methods of transfer of the spermatophore vary between species. In some, the spermatophore may be placed directly into the female cloaca while in others, the female may be guided to the spermatophore or restrained with an embrace called amplexus. Certain primitive salamanders in the families Sirenidae, Hynobiidae and Cryptobranchidae practice external fertilisation in a similar manner to frogs, with the female laying the eggs in water and the male releasing sperm onto the egg mass. | Amphibian | Wikipedia | 421 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
With a few exceptions, frogs use external fertilisation. The male grasps the female tightly with his forelimbs either behind the arms or in front of the back legs, or in the case of Epipedobates tricolor, around the neck. They remain in amplexus with their cloacae positioned close together while the female lays the eggs and the male covers them with sperm. Roughened nuptial pads on the male's hands aid in retaining grip. Often the male collects and retains the egg mass, forming a sort of basket with the hind feet. An exception is the granular poison frog (Oophaga granulifera) where the male and female place their cloacae in close proximity while facing in opposite directions and then release eggs and sperm simultaneously. The tailed frog (Ascaphus truei) exhibits internal fertilisation. The "tail" is only possessed by the male and is an extension of the cloaca and used to inseminate the female. This frog lives in fast-flowing streams and internal fertilisation prevents the sperm from being washed away before fertilisation occurs. The sperm may be retained in storage tubes attached to the oviduct until the following spring.
Most frogs can be classified as either prolonged or explosive breeders. Typically, prolonged breeders congregate at a breeding site, the males usually arriving first, calling and setting up territories. Other satellite males remain quietly nearby, waiting for their opportunity to take over a territory. The females arrive sporadically, mate selection takes place and eggs are laid. The females depart and territories may change hands. More females appear and in due course, the breeding season comes to an end. Explosive breeders on the other hand are found where temporary pools appear in dry regions after rainfall. These frogs are typically fossorial species that emerge after heavy rains and congregate at a breeding site. They are attracted there by the calling of the first male to find a suitable place, perhaps a pool that forms in the same place each rainy season. The assembled frogs may call in unison and frenzied activity ensues, the males scrambling to mate with the usually smaller number of females. | Amphibian | Wikipedia | 451 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
There is a direct competition between males to win the attention of the females in salamanders and newts, with elaborate courtship displays to keep the female's attention long enough to get her interested in choosing him to mate with. Some species store sperm through long breeding seasons, as the extra time may allow for interactions with rival sperm.
Unisexual reproduction
Unisexual female mole salamanders (genus Ambystoma) are common in the Great Lakes region of North America. These salamanders are the oldest known unisexual vertebrate lineage, having emerged about 5 million years ago. Genome exchange can sometimes occur between the unisexual female Ambystoma and males from sympatric sexual species.
Life cycle
Most amphibians go through metamorphosis, a process of significant morphological change after birth. In typical amphibian development, eggs are laid in water and larvae are adapted to an aquatic lifestyle. Frogs, toads and salamanders all hatch from the egg as larvae with external gills. Metamorphosis in amphibians is regulated by thyroxine concentration in the blood, which stimulates metamorphosis, and prolactin, which counteracts thyroxine's effect. Specific events are dependent on threshold values for different tissues. Because most embryonic development is outside the parental body, it is subject to many adaptations due to specific environmental circumstances. For this reason tadpoles can have horny ridges instead of teeth, whisker-like skin extensions or fins. They also make use of a sensory lateral line organ similar to that of fish. After metamorphosis, these organs become redundant and will be reabsorbed by controlled cell death, called apoptosis. The variety of adaptations to specific environmental circumstances among amphibians is wide, with many discoveries still being made.
Eggs | Amphibian | Wikipedia | 383 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In the egg, the embryo is suspended in perivitelline fluid and surrounded by semi-permeable gelatinous capsules, with the yolk mass providing nutrients. As the larvae hatch, the capsules are dissolved by enzymes secreted from gland at the tip of the snout. The eggs of some salamanders and frogs contain unicellular green algae. These penetrate the jelly envelope after the eggs are laid and may increase the supply of oxygen to the embryo through photosynthesis. They seem to both speed up the development of the larvae and reduce mortality. In the wood frog (Rana sylvatica), the interior of the globular egg cluster has been found to be up to warmer than its surroundings, which is an advantage in its cool northern habitat.
The eggs may be deposited singly, in cluster or in long strands. Sites for laying eggs include water, mud, burrows, debris and on plants or under logs or stones. The greenhouse frog (Eleutherodactylus planirostris) lays eggs in small groups in the soil where they develop in about two weeks directly into juvenile frogs without an intervening larval stage. The tungara frog (Physalaemus pustulosus) builds a floating nest from foam to protect its eggs. First a raft is built, then eggs are laid in the centre, and finally a foam cap is overlaid. The foam has anti-microbial properties. It contains no detergents but is created by whipping up proteins and lectins secreted by the female.
Larvae | Amphibian | Wikipedia | 318 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The eggs of amphibians are typically laid in water and hatch into free-living larvae that complete their development in water and later transform into either aquatic or terrestrial adults. In many species of frog and in most lungless salamanders (Plethodontidae), direct development takes place, the larvae growing within the eggs and emerging as miniature adults. Many caecilians and some other amphibians lay their eggs on land, and the newly hatched larvae wriggle or are transported to water bodies. Some caecilians, the alpine salamander (Salamandra atra) and some of the African live-bearing toads (Nectophrynoides spp.) are viviparous. Their larvae feed on glandular secretions and develop within the female's oviduct, often for long periods. Other amphibians, but not caecilians, are ovoviviparous. The eggs are retained in or on the parent's body, but the larvae subsist on the yolks of their eggs and receive no nourishment from the adult. The larvae emerge at varying stages of their growth, either before or after metamorphosis, according to their species. The toad genus Nectophrynoides exhibits all of these developmental patterns among its dozen or so members. Amphibian larvae are known as tadpoles. They have thick, rounded bodies with powerful muscular tails. | Amphibian | Wikipedia | 297 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Frogs
Unlike in other amphibians, frog tadpoles do not resemble adults. The free-living larvae are normally fully aquatic, but the tadpoles of some species (such as Nannophrys ceylonensis) are semi-terrestrial and live among wet rocks. Tadpoles have cartilaginous skeletons, gills for respiration (external gills at first, internal gills later), lateral line systems and large tails that they use for swimming. Newly hatched tadpoles soon develop gill pouches that cover the gills. These internal gills and operculum are not homologous with those of fish, and are only found in tadpoles as both salamanders and caecilians have external gills only. Combined with buccal pumping the internal gills has allowed tadpoles to adopt a filter feeding lifestyle, even if several species have since evolved other types of feeding strategies. The lungs develop early and are used as accessory breathing organs, the tadpoles rising to the water surface to gulp air. Some species complete their development inside the egg and hatch directly into small frogs. These larvae do not have gills but instead have specialised areas of skin through which respiration takes place. While tadpoles do not have true teeth, in most species, the jaws have long, parallel rows of small keratinized structures called keradonts surrounded by a horny beak. Front legs are formed under the gill sac and hind legs become visible a few days later.
Iodine and T4 (over stimulate the spectacular apoptosis [programmed cell death] of the cells of the larval gills, tail and fins) also stimulate the evolution of nervous systems transforming the aquatic, vegetarian tadpole into the terrestrial, carnivorous frog with better neurological, visuospatial, olfactory and cognitive abilities for hunting. | Amphibian | Wikipedia | 371 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In fact, tadpoles developing in ponds and streams are typically herbivorous. Pond tadpoles tend to have deep bodies, large caudal fins and small mouths; they swim in the quiet waters feeding on growing or loose fragments of vegetation. Stream dwellers mostly have larger mouths, shallow bodies and caudal fins; they attach themselves to plants and stones and feed on the surface films of algae and bacteria. They also feed on diatoms, filtered from the water through the gills, and stir up the sediment at bottom of the pond, ingesting edible fragments. They have a relatively long, spiral-shaped gut to enable them to digest this diet. Some species are carnivorous at the tadpole stage, eating insects, smaller tadpoles and fish. Young of the Cuban tree frog (Osteopilus septentrionalis) can occasionally be cannibalistic, the younger tadpoles attacking a larger, more developed tadpole when it is undergoing metamorphosis.
At metamorphosis, rapid changes in the body take place as the lifestyle of the frog changes completely. The spiral-shaped mouth with horny tooth ridges is reabsorbed together with the spiral gut. The animal develops a large jaw, and its gills disappear along with its gill sac. Eyes and legs grow quickly, and a tongue is formed. There are associated changes in the neural networks such as development of stereoscopic vision and loss of the lateral line system. All this can happen in about a day. A few days later, the tail is reabsorbed, due to the higher thyroxine concentration required for this to take place.
Salamanders | Amphibian | Wikipedia | 339 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
At hatching, a typical salamander larva has eyes without lids, teeth in both upper and lower jaws, three pairs of feathery external gills, and a long tail with dorsal and ventral fins. The forelimbs may be partially developed and the hind limbs are rudimentary in pond-living species but may be rather more developed in species that reproduce in moving water. Pond-type larvae often have a pair of balancers, rod-like structures on either side of the head that may prevent the gills from becoming clogged up with sediment. Both of these are able to breed. Some have larvae that never fully develop into the adult form, a condition known as neoteny. Neoteny occurs when the animal's growth rate is very low and is usually linked to adverse conditions such as low water temperatures that may change the response of the tissues to the hormone thyroxine. as well as lack of food. There are fifteen species of obligate neotenic salamanders, including species of Necturus, Proteus and Amphiuma, and many examples of facultative ones, such as the northwestern salamander (Ambystoma gracile) and the tiger salamander (A. tigrinum) that adopt this strategy under appropriate environmental circumstances.
Lungless salamanders in the family Plethodontidae are terrestrial and lay a small number of unpigmented eggs in a cluster among damp leaf litter. Each egg has a large yolk sac and the larva feeds on this while it develops inside the egg, emerging fully formed as a juvenile salamander. The female salamander often broods the eggs. In the genus Ensatinas, the female has been observed to coil around them and press her throat area against them, effectively massaging them with a mucous secretion. | Amphibian | Wikipedia | 379 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In newts and salamanders, metamorphosis is less dramatic than in frogs. This is because the larvae are already carnivorous and continue to feed as predators when they are adults so few changes are needed to their digestive systems. Their lungs are functional early, but the larvae do not make as much use of them as do tadpoles. Their gills are never covered by gill sacs and are reabsorbed just before the animals leave the water. Other changes include the reduction in size or loss of tail fins, the closure of gill slits, thickening of the skin, the development of eyelids, and certain changes in dentition and tongue structure. Salamanders are at their most vulnerable at metamorphosis as swimming speeds are reduced and transforming tails are encumbrances on land. Adult salamanders often have an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water phase, prolactin is the required hormone, and for adaptation to the land phase, thyroxine. External gills do not return in subsequent aquatic phases because these are completely absorbed upon leaving the water for the first time.
Caecilians
Most terrestrial caecilians that lay eggs do so in burrows or moist places on land near bodies of water. The development of the young of Ichthyophis glutinosus, a species from Sri Lanka, has been much studied. The eel-like larvae hatch out of the eggs and make their way to water. They have three pairs of external red feathery gills, a blunt head with two rudimentary eyes, a lateral line system and a short tail with fins. They swim by undulating their body from side to side. They are mostly active at night, soon lose their gills and make sorties onto land. Metamorphosis is gradual. By the age of about ten months they have developed a pointed head with sensory tentacles near the mouth and lost their eyes, lateral line systems and tails. The skin thickens, embedded scales develop and the body divides into segments. By this time, the caecilian has constructed a burrow and is living on land. | Amphibian | Wikipedia | 442 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In the majority of species of caecilians, the young are produced by viviparity. Typhlonectes compressicauda, a species from South America, is typical of these. Up to nine larvae can develop in the oviduct at any one time. They are elongated and have paired sac-like gills, small eyes and specialised scraping teeth. At first, they feed on the yolks of the eggs, but as this source of nourishment declines they begin to rasp at the ciliated epithelial cells that line the oviduct. This stimulates the secretion of fluids rich in lipids and mucoproteins on which they feed along with scrapings from the oviduct wall. They may increase their length sixfold and be two-fifths as long as their mother before being born. By this time they have undergone metamorphosis, lost their eyes and gills, developed a thicker skin and mouth tentacles, and reabsorbed their teeth. A permanent set of teeth grow through soon after birth.
Gills are only necessarily during embryonic development, and in species that give birth the offspring is born after gill degeneration. In egg laying caecilians the gills are either reabsorbed before hatching, or, in species that hatch with gill remnants still present, short lived and only leaves behind a gill slit. For species with scales under their skin, the scales does not form before during metamorphosis.
The ringed caecilian (Siphonops annulatus) has developed a unique adaptation for the purposes of reproduction. The progeny feed on a skin layer that is specially developed by the adult in a phenomenon known as maternal dermatophagy. The brood feed as a batch for about seven minutes at intervals of approximately three days which gives the skin an opportunity to regenerate. Meanwhile, they have been observed to ingest fluid exuded from the maternal cloaca.
Parental care
The care of offspring among amphibians has been little studied but, in general, the larger the number of eggs in a batch, the less likely it is that any degree of parental care takes place. Nevertheless, it is estimated that in up to 20% of amphibian species, one or both adults play some role in the care of the young. Those species that breed in smaller water bodies or other specialised habitats tend to have complex patterns of behaviour in the care of their young. | Amphibian | Wikipedia | 508 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Many woodland salamanders lay clutches of eggs under dead logs or stones on land. The black mountain salamander (Desmognathus welteri) does this, the mother brooding the eggs and guarding them from predation as the embryos feed on the yolks of their eggs. When fully developed, they break their way out of the egg capsules and disperse as juvenile salamanders. The male hellbender, a primitive salamander, excavates an underwater nest and encourages females to lay there. The male then guards the site for the two or three months before the eggs hatch, using body undulations to fan the eggs and increase their supply of oxygen. | Amphibian | Wikipedia | 141 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
The male Colostethus subpunctatus, a tiny frog, protects the egg cluster which is hidden under a stone or log. When the eggs hatch, the male transports the tadpoles on his back, stuck there by a mucous secretion, to a temporary pool where he dips himself into the water and the tadpoles drop off. The male midwife toad (Alytes obstetricans) winds egg strings round his thighs and carries the eggs around for up to eight weeks. He keeps them moist and when they are ready to hatch, he visits a pond or ditch and releases the tadpoles. The female gastric-brooding frog (Rheobatrachus spp.) reared larvae in her stomach after swallowing either the eggs or hatchlings; however, this stage was never observed before the species became extinct. The tadpoles secrete a hormone that inhibits digestion in the mother whilst they develop by consuming their very large yolk supply. The pouched frog (Assa darlingtoni) lays eggs on the ground. When they hatch, the male carries the tadpoles around in brood pouches on his hind legs. The aquatic Surinam toad (Pipa pipa) raises its young in pores on its back where they remain until metamorphosis. The granular poison frog (Oophaga granulifera) is typical of a number of tree frogs in the poison dart frog family Dendrobatidae. Its eggs are laid on the forest floor and when they hatch, the tadpoles are carried one by one on the back of an adult to a suitable water-filled crevice such as the axil of a leaf or the rosette of a bromeliad. The female visits the nursery sites regularly and deposits unfertilised eggs in the water and these are consumed by the tadpoles. | Amphibian | Wikipedia | 385 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Genetics and genomics
Amphibians are notable among vertebrates for their diversity of chromosomes and genomes. The karyotypes (chromosomes) have been determined for at least 1,193 (14.5%) of the ≈8,200 known (diploid) species, including 963 anurans, 209 salamanders, and 21 caecilians. Generally, the karyotypes of diploid amphibians are characterized by 20–26 bi-armed chromosomes. Amphibians have also very large genomes compared to other taxa of vertebrates and corresponding variation in genome size (C-value: picograms of DNA in haploid nuclei). The genome sizes range from 0.95 to 11.5 pg in frogs, from 13.89 to 120.56 pg in salamanders, and from 2.94 to 11.78 pg in caecilians.
The large genome sizes have prevented whole-genome sequencing of amphibians although a number of genomes have been published recently. The 1.7GB draft genome of Xenopus tropicalis was the first to be reported for amphibians in 2010. Compared to some salamanders this frog genome is tiny. For instance, the genome of the Mexican axolotl turned out to be 32 Gb, which is more than 10 times larger than the human genome (3GB).
Feeding and diet
With a few exceptions, adult amphibians are predators, feeding on virtually anything that moves that they can swallow. The diet mostly consists of small prey that do not move too fast such as beetles, caterpillars, earthworms and spiders. The sirens (Siren spp.) often ingest aquatic plant material with the invertebrates on which they feed and a Brazilian tree frog (Xenohyla truncata) includes a large quantity of fruit in its diet. The Mexican burrowing toad (Rhinophrynus dorsalis) has a specially adapted tongue for picking up ants and termites. It projects it with the tip foremost whereas other frogs flick out the rear part first, their tongues being hinged at the front. | Amphibian | Wikipedia | 436 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Food is mostly selected by sight, even in conditions of dim light. Movement of the prey triggers a feeding response. Frogs have been caught on fish hooks baited with red flannel and green frogs (Rana clamitans) have been found with stomachs full of elm seeds that they had seen floating past. Toads, salamanders and caecilians also use smell to detect prey. This response is mostly secondary because salamanders have been observed to remain stationary near odoriferous prey but only feed if it moves. Cave-dwelling amphibians normally hunt by smell. Some salamanders seem to have learned to recognize immobile prey when it has no smell, even in complete darkness.
Amphibians usually swallow food whole but may chew it lightly first to subdue it. They typically have small hinged pedicellate teeth, a feature unique to amphibians. The base and crown of these are composed of dentine separated by an uncalcified layer and they are replaced at intervals. Salamanders, caecilians and some frogs have one or two rows of teeth in both jaws, but some frogs (Rana spp.) lack teeth in the lower jaw, and toads (Bufo spp.) have no teeth. In many amphibians there are also vomerine teeth attached to a facial bone in the roof of the mouth.
The tiger salamander (Ambystoma tigrinum) is typical of the frogs and salamanders that hide under cover ready to ambush unwary invertebrates. Other amphibians, such as the Bufo spp. toads, actively search for prey, while the Argentine horned frog (Ceratophrys ornata) lures inquisitive prey closer by raising its hind feet over its back and vibrating its yellow toes. Among leaf litter frogs in Panama, frogs that actively hunt prey have narrow mouths and are slim, often brightly coloured and toxic, while ambushers have wide mouths and are broad and well-camouflaged. Caecilians do not flick their tongues, but catch their prey by grabbing it with their slightly backward-pointing teeth. The struggles of the prey and further jaw movements work it inwards and the caecilian usually retreats into its burrow. The subdued prey is gulped down whole. | Amphibian | Wikipedia | 473 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
When they are newly hatched, frog larvae feed on the yolk of the egg. When this is exhausted some move on to feed on bacteria, algal crusts, detritus and raspings from submerged plants. Water is drawn in through their mouths, which are usually at the bottom of their heads, and passes through branchial food traps between their mouths and their gills where fine particles are trapped in mucus and filtered out. Others have specialised mouthparts consisting of a horny beak edged by several rows of labial teeth. They scrape and bite food of many kinds as well as stirring up the bottom sediment, filtering out larger particles with the papillae around their mouths. Some, such as the spadefoot toads, have strong biting jaws and are carnivorous or even cannibalistic.
Vocalization
The calls made by caecilians and salamanders are limited to occasional soft squeaks, grunts or hisses and have not been much studied. A clicking sound sometimes produced by caecilians may be a means of orientation, as in bats, or a form of communication. Most salamanders are considered voiceless, but the California giant salamander (Dicamptodon ensatus) has vocal cords and can produce a rattling or barking sound. Some species of salamander emit a quiet squeak or yelp if attacked. | Amphibian | Wikipedia | 280 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Frogs are much more vocal, especially during the breeding season when they use their voices to attract mates. The presence of a particular species in an area may be more easily discerned by its characteristic call than by a fleeting glimpse of the animal itself. In most species, the sound is produced by expelling air from the lungs over the vocal cords into one or more air sacs in the throat or at the corner of the mouth. This may distend like a balloon and acts as a resonator, helping to transfer the sound to the atmosphere, or the water at times when the animal is submerged. The main vocalisation is the male's loud advertisement call which seeks to both encourage a female to approach and discourage other males from intruding on its territory. This call is modified to a quieter courtship call on the approach of a female or to a more aggressive version if a male intruder draws near. Calling carries the risk of attracting predators and involves the expenditure of much energy. Other calls include those given by a female in response to the advertisement call and a release call given by a male or female during unwanted attempts at amplexus. When a frog is attacked, a distress or fright call is emitted, often resembling a scream. The usually nocturnal Cuban tree frog (Osteopilus septentrionalis) produces a rain call when there is rainfall during daylight hours.
Territorial behaviour
Little is known of the territorial behaviour of caecilians, but some frogs and salamanders defend home ranges. These are usually feeding, breeding or sheltering sites. Males normally exhibit such behaviour though in some species, females and even juveniles are also involved. Although in many frog species, females are larger than males, this is not the case in most species where males are actively involved in territorial defence. Some of these have specific adaptations such as enlarged teeth for biting or spines on the chest, arms or thumbs. | Amphibian | Wikipedia | 386 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
In salamanders, defence of a territory involves adopting an aggressive posture and if necessary attacking the intruder. This may involve snapping, chasing and sometimes biting, occasionally causing the loss of a tail. The behaviour of red back salamanders (Plethodon cinereus) has been much studied. 91% of marked individuals that were later recaptured were within a metre (yard) of their original daytime retreat under a log or rock. A similar proportion, when moved experimentally a distance of , found their way back to their home base. The salamanders left odour marks around their territories which averaged in size and were sometimes inhabited by a male and female pair. These deterred the intrusion of others and delineated the boundaries between neighbouring areas. Much of their behaviour seemed stereotyped and did not involve any actual contact between individuals. An aggressive posture involved raising the body off the ground and glaring at the opponent who often turned away submissively. If the intruder persisted, a biting lunge was usually launched at either the tail region or the naso-labial grooves. Damage to either of these areas can reduce the fitness of the rival, either because of the need to regenerate tissue or because it impairs its ability to detect food.
In frogs, male territorial behaviour is often observed at breeding locations; calling is both an announcement of ownership of part of this resource and an advertisement call to potential mates. In general, a deeper voice represents a heavier and more powerful individual, and this may be sufficient to prevent intrusion by smaller males. Much energy is used in the vocalization and it takes a toll on the territory holder who may be displaced by a fitter rival if he tires. There is a tendency for males to tolerate the holders of neighbouring territories while vigorously attacking unknown intruders. Holders of territories have a "home advantage" and usually come off better in an encounter between two similar-sized frogs. If threats are insufficient, chest to chest tussles may take place. Fighting methods include pushing and shoving, deflating the opponent's vocal sac, seizing him by the head, jumping on his back, biting, chasing, splashing, and ducking him under the water.
Defence mechanisms | Amphibian | Wikipedia | 447 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Amphibians have soft bodies with thin skins, and lack claws, defensive armour, or spines. Nevertheless, they have evolved various defence mechanisms to keep themselves alive. The first line of defence in salamanders and frogs is the mucous secretion that they produce. This keeps their skin moist and makes them slippery and difficult to grip. The secretion is often sticky and distasteful or toxic. Snakes have been observed yawning and gaping when trying to swallow African clawed frogs (Xenopus laevis), which gives the frogs an opportunity to escape. Caecilians have been little studied in this respect, but the Cayenne caecilian (Typhlonectes compressicauda) produces toxic mucus that has killed predatory fish in a feeding experiment in Brazil. In some salamanders, the skin is poisonous. The rough-skinned newt (Taricha granulosa) from North America and other members of its genus contain the neurotoxin tetrodotoxin (TTX), the most toxic non-protein substance known and almost identical to that produced by pufferfish. Handling the newts does not cause harm, but ingestion of even the most minute amounts of the skin is deadly. In feeding trials, fish, frogs, reptiles, birds and mammals were all found to be susceptible. The only predators with some tolerance to the poison are certain populations of common garter snake (Thamnophis sirtalis).
In locations where both snake and salamander co-exist, the snakes have developed immunity through genetic changes and they feed on the amphibians with impunity. Coevolution occurs with the newt increasing its toxic capabilities at the same rate as the snake further develops its immunity. Some frogs and toads are toxic, the main poison glands being at the side of the neck and under the warts on the back. These regions are presented to the attacking animal and their secretions may be foul-tasting or cause various physical or neurological symptoms. Altogether, over 200 toxins have been isolated from the limited number of amphibian species that have been investigated. | Amphibian | Wikipedia | 439 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Poisonous species often use bright colouring to warn potential predators of their toxicity. These warning colours tend to be red or yellow combined with black, with the fire salamander (Salamandra salamandra) being an example. Once a predator has sampled one of these, it is likely to remember the colouration next time it encounters a similar animal. In some species, such as the fire-bellied toad (Bombina spp.), the warning colouration is on the belly and these animals adopt a defensive pose when attacked, exhibiting their bright colours to the predator. The frog Allobates zaparo is not poisonous, but mimics the appearance of other toxic species in its locality, a strategy that may deceive predators.
Many amphibians are nocturnal and hide during the day, thereby avoiding diurnal predators that hunt by sight. Other amphibians use camouflage to avoid being detected. They have various colourings such as mottled browns, greys and olives to blend into the background. Some salamanders adopt defensive poses when faced by a potential predator such as the North American northern short-tailed shrew (Blarina brevicauda). Their bodies writhe and they raise and lash their tails which makes it difficult for the predator to avoid contact with their poison-producing granular glands. A few salamanders will autotomise their tails when attacked, sacrificing this part of their anatomy to enable them to escape. The tail may have a constriction at its base to allow it to be easily detached. The tail is regenerated later, but the energy cost to the animal of replacing it is significant.
Some frogs and toads inflate themselves to make themselves look large and fierce, and some spadefoot toads (Pelobates spp) scream and leap towards the attacker. Giant salamanders of the genus Andrias, as well as Ceratophrine and Pyxicephalus frogs possess sharp teeth and are capable of drawing blood with a defensive bite. The blackbelly salamander (Desmognathus quadramaculatus) can bite an attacking common garter snake (Thamnophis sirtalis) two or three times its size on the head and often manages to escape. | Amphibian | Wikipedia | 468 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Cognition
In amphibians, there is evidence of habituation, associative learning through both classical and instrumental learning, and discrimination abilities. Amphibians are widely considered to be sentient, able to feel emotions such as anxiety and fear.
In one experiment, when offered live fruit flies (Drosophila virilis), salamanders chose the larger of 1 vs 2 and 2 vs 3. Frogs can distinguish between low numbers (1 vs 2, 2 vs 3, but not 3 vs 4) and large numbers (3 vs 6, 4 vs 8, but not 4 vs 6) of prey. This is irrespective of other characteristics, i.e. surface area, volume, weight and movement, although discrimination among large numbers may be based on surface area.
Conservation
Dramatic declines in amphibian populations, including population crashes and mass localized extinction, have been noted since the late 1980s from locations all over the world, and amphibian declines are thus perceived to be one of the most critical threats to global biodiversity. In 2004, the International Union for Conservation of Nature (IUCN) reported stating that currently birds, mammals, and amphibians extinction rates were at minimum 48 times greater than natural extinction rates—possibly 1,024 times higher. In 2006, there were believed to be 4,035 species of amphibians that depended on water at some stage during their life cycle. Of these, 1,356 (33.6%) were considered to be threatened and this figure is likely to be an underestimate because it excludes 1,427 species for which there was insufficient data to assess their status. A number of causes are believed to be involved, including habitat destruction and modification, over-exploitation, pollution, introduced species, global warming, endocrine-disrupting pollutants, destruction of the ozone layer (ultraviolet radiation has shown to be especially damaging to the skin, eyes, and eggs of amphibians), and diseases like chytridiomycosis. However, many of the causes of amphibian declines are still poorly understood, and are a topic of ongoing discussion.
Food webs and predation
Any decline in amphibian numbers will affect the patterns of predation. The loss of carnivorous species near the top of the food chain will upset the delicate ecosystem balance and may cause dramatic increases in opportunistic species. | Amphibian | Wikipedia | 487 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Predators that feed on amphibians are affected by their decline. The western terrestrial garter snake (Thamnophis elegans) in California is largely aquatic and depends heavily on two species of frog that are decreasing in numbers, the Yosemite toad (Bufo canorus) and the mountain yellow-legged frog (Rana muscosa), putting the snake's future at risk. If the snake were to become scarce, this would affect birds of prey and other predators that feed on it. Meanwhile, in the ponds and lakes, fewer frogs means fewer tadpoles. These normally play an important role in controlling the growth of algae and also forage on detritus that accumulates as sediment on the bottom. A reduction in the number of tadpoles may lead to an overgrowth of algae, resulting in depletion of oxygen in the water when the algae later die and decompose. Aquatic invertebrates and fish might then die and there would be unpredictable ecological consequences.
Pollution and pesticides
The decline in amphibian and reptile populations has led to an awareness of the effects of pesticides on reptiles and amphibians. In the past, the argument that amphibians or reptiles were more susceptible to any chemical contamination than any land aquatic vertebrate was not supported by research until recently. Amphibians and reptiles have complex life cycles, live in different climate and ecological zones, and are more vulnerable to chemical exposure. Certain pesticides, such as organophosphates, neonicotinoids, and carbamates, react via cholinesterase inhibition. Cholinesterase is an enzyme that causes the hydrolysis of acetylcholine, an excitatory neurotransmitter that is abundant in the nervous system. AChE inhibitors are either reversible or irreversible, and carbamates are safer than organophosphorus insecticides, which are more likely to cause cholinergic poisoning. Reptile exposure to an AChE inhibitory pesticide may result in disruption of neural function in reptiles. The buildup of these inhibitory effects on motor performance, such as food consumption and other activities. | Amphibian | Wikipedia | 440 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Conservation and protection strategies
The Amphibian Specialist Group of the IUCN is spearheading efforts to implement a comprehensive global strategy for amphibian conservation. Amphibian Ark is an organization that was formed to implement the ex-situ conservation recommendations of this plan, and they have been working with zoos and aquaria around the world, encouraging them to create assurance colonies of threatened amphibians. One such project is the Panama Amphibian Rescue and Conservation Project that built on existing conservation efforts in Panama to create a country-wide response to the threat of chytridiomycosis.
Another measure would be to stop exploitation of frogs for human consumption. In the Middle East, a growing appetite for eating frog legs and the consequent gathering of them for food was already linked to an increase in mosquitoes and thus has direct consequences for human health. | Amphibian | Wikipedia | 172 | 621 | https://en.wikipedia.org/wiki/Amphibian | Biology and health sciences | Biology | null |
Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthetic eukaryotes, which include species from multiple distinct clades. Such organisms range from unicellular microalgae such as Chlorella, Prototheca and the diatoms, to multicellular macroalgae such as the giant kelp, a large brown alga which may grow up to in length. Most algae are aquatic organisms and lack many of the distinct cell and tissue types, such as stomata, xylem and phloem that are found in land plants. The largest and most complex marine algae are called seaweeds. In contrast, the most complex freshwater forms are the Charophyta, a division of green algae which includes, for example, Spirogyra and stoneworts. Algae that are carried passively by water are plankton, specifically phytoplankton.
Algae constitute a polyphyletic group since they do not include a common ancestor, and although their chlorophyll-bearing plastids seem to have a single origin (from symbiogenesis with cyanobacteria), they were acquired in different ways. Green algae are a prominent examples of algae that have primary chloroplasts derived from endosymbiont cyanobacteria. Diatoms and brown algae are examples of algae with secondary chloroplasts derived from endosymbiotic red algae, which they acquired via phagocytosis. Algae exhibit a wide range of reproductive strategies, from simple asexual cell division to complex forms of sexual reproduction via spores. | Algae | Wikipedia | 344 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Algae lack the various structures that characterize plants (which evolved from freshwater green algae), such as the phyllids (leaf-like structures) and rhizoids of bryophytes (non-vascular plants), and the roots, leaves and other xylemic/phloemic organs found in tracheophytes (vascular plants). Most algae are autotrophic, although some are mixotrophic, deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy or phagotrophy. Some unicellular species of green algae, many golden algae, euglenids, dinoflagellates, and other algae have become heterotrophs (also called colorless or apochlorotic algae), sometimes parasitic, relying entirely on external energy sources and have limited or no photosynthetic apparatus. Some other heterotrophic organisms, such as the apicomplexans, are also derived from cells whose ancestors possessed chlorophyllic plastids, but are not traditionally considered as algae. Algae have photosynthetic machinery ultimately derived from cyanobacteria that produce oxygen as a byproduct of splitting water molecules, unlike other organisms that conduct anoxygenic photosynthesis such as purple and green sulfur bacteria. Fossilized filamentous algae from the Vindhya basin have been dated to 1.6 to 1.7 billion years ago.
Because of the wide range of algae types, they have increasingly different industrial and traditional applications in human society. Traditional seaweed farming practices have existed for thousands of years and have strong traditions in East Asia food cultures. More modern algaculture applications extend the food traditions for other applications, including cattle feed, using algae for bioremediation or pollution control, transforming sunlight into algae fuels or other chemicals used in industrial processes, and in medical and scientific applications. A 2020 review found that these applications of algae could play an important role in carbon sequestration to mitigate climate change while providing lucrative value-added products for global economies.
Etymology and study
The singular is the Latin word for 'seaweed' and retains that meaning in English. The etymology is obscure. Although some speculate that it is related to Latin , 'be cold', no reason is known to associate seaweed with temperature. A more likely source is , 'binding, entwining'. | Algae | Wikipedia | 500 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
The Ancient Greek word for 'seaweed' was (), which could mean either the seaweed (probably red algae) or a red dye derived from it. The Latinization, , meant primarily the cosmetic rouge. The etymology is uncertain, but a strong candidate has long been some word related to the Biblical (), 'paint' (if not that word itself), a cosmetic eye-shadow used by the ancient Egyptians and other inhabitants of the eastern Mediterranean. It could be any color: black, red, green, or blue.
The study of algae is most commonly called phycology (); the term algology is falling out of use.
Classifications
One definition of algae is that they "have chlorophyll as their primary photosynthetic pigment and lack a sterile covering of cells around their reproductive cells". On the other hand, the colorless Prototheca under Chlorophyta are all devoid of any chlorophyll. Although cyanobacteria are often referred to as "blue-green algae", most authorities exclude all prokaryotes, including cyanobacteria, from the definition of algae.
The algae contain chloroplasts that are similar in structure to cyanobacteria. Chloroplasts contain circular DNA like that in cyanobacteria and are interpreted as representing reduced endosymbiotic cyanobacteria. However, the exact origin of the chloroplasts is different among separate lineages of algae, reflecting their acquisition during different endosymbiotic events. The table below describes the composition of the three major groups of algae. Their lineage relationships are shown in the figure in the upper right. Many of these groups contain some members that are no longer photosynthetic. Some retain plastids, but not chloroplasts, while others have lost plastids entirely.
Phylogeny based on plastid not nucleocytoplasmic genealogy:
Linnaeus, in Species Plantarum (1753), the starting point for modern botanical nomenclature, recognized 14 genera of algae, of which only four are currently considered among algae. In Systema Naturae, Linnaeus described the genera Volvox and Corallina, and a species of Acetabularia (as Madrepora), among the animals. | Algae | Wikipedia | 479 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
In 1768, Samuel Gottlieb Gmelin (1744–1774) published the Historia Fucorum, the first work dedicated to marine algae and the first book on marine biology to use the then new binomial nomenclature of Linnaeus. It included elaborate illustrations of seaweed and marine algae on folded leaves.
W. H. Harvey (1811–1866) and Lamouroux (1813) were the first to divide macroscopic algae into four divisions based on their pigmentation. This is the first use of a biochemical criterion in plant systematics. Harvey's four divisions are: red algae (Rhodospermae), brown algae (Melanospermae), green algae (Chlorospermae), and Diatomaceae.
At this time, microscopic algae were discovered and reported by a different group of workers (e.g., O. F. Müller and Ehrenberg) studying the Infusoria (microscopic organisms). Unlike macroalgae, which were clearly viewed as plants, microalgae were frequently considered animals because they are often motile. Even the nonmotile (coccoid) microalgae were sometimes merely seen as stages of the lifecycle of plants, macroalgae, or animals.
Although used as a taxonomic category in some pre-Darwinian classifications, e.g., Linnaeus (1753), de Jussieu (1789), Lamouroux (1813), Harvey (1836), Horaninow (1843), Agassiz (1859), Wilson & Cassin (1864), in further classifications, the "algae" are seen as an artificial, polyphyletic group. | Algae | Wikipedia | 346 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Throughout the 20th century, most classifications treated the following groups as divisions or classes of algae: cyanophytes, rhodophytes, chrysophytes, xanthophytes, bacillariophytes, phaeophytes, pyrrhophytes (cryptophytes and dinophytes), euglenophytes, and chlorophytes. Later, many new groups were discovered (e.g., Bolidophyceae), and others were splintered from older groups: charophytes and glaucophytes (from chlorophytes), many heterokontophytes (e.g., synurophytes from chrysophytes, or eustigmatophytes from xanthophytes), haptophytes (from chrysophytes), and chlorarachniophytes (from xanthophytes).
With the abandonment of plant-animal dichotomous classification, most groups of algae (sometimes all) were included in Protista, later also abandoned in favour of Eukaryota. However, as a legacy of the older plant life scheme, some groups that were also treated as protozoans in the past still have duplicated classifications (see ambiregnal protists). | Algae | Wikipedia | 282 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Some parasitic algae (e.g., the green algae Prototheca and Helicosporidium, parasites of metazoans, or Cephaleuros, parasites of plants) were originally classified as fungi, sporozoans, or protistans of incertae sedis, while others (e.g., the green algae Phyllosiphon and Rhodochytrium, parasites of plants, or the red algae Pterocladiophila and Gelidiocolax mammillatus, parasites of other red algae, or the dinoflagellates Oodinium, parasites of fish) had their relationship with algae conjectured early. In other cases, some groups were originally characterized as parasitic algae (e.g., Chlorochytrium), but later were seen as endophytic algae. Some filamentous bacteria (e.g., Beggiatoa) were originally seen as algae. Furthermore, groups like the apicomplexans are also parasites derived from ancestors that possessed plastids, but are not included in any group traditionally seen as algae.
Evolution
Algae are polyphyletic thus their origin cannot be traced back to single hypothetical common ancestor. It is thought that they came into existence when photosynthetic coccoid cyanobacteria got phagocytized by a unicellular heterotrophic eukaryote (a protist), giving rise to double-membranous primary plastids. Such symbiogenic events (primary symbiogenesis) are believed to have occurred more than 1.5 billion years ago during the Calymmian period, early in Boring Billion, but it is difficult to track the key events because of so much time gap. Primary symbiogenesis gave rise to three divisions of archaeplastids, namely the Viridiplantae (green algae and later plants), Rhodophyta (red algae) and Glaucophyta ("grey algae"), whose plastids further spread into other protist lineages through eukaryote-eukaryote predation, engulfments and subsequent endosymbioses (secondary and tertiary symbiogenesis). This process of serial cell "capture" and "enslavement" explains the diversity of photosynthetic eukaryotes. | Algae | Wikipedia | 494 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Recent genomic and phylogenomic approaches have significantly clarified plastid genome evolution, the horizontal movement of endosymbiont genes to the "host" nuclear genome, and plastid spread throughout the eukaryotic tree of life.
Relationship to land plants
Fossils of isolated spores suggest land plants may have been around as long as 475 million years ago (mya) during the Late Cambrian/Early Ordovician period, from sessile shallow freshwater charophyte algae much like Chara, which likely got stranded ashore when riverine/lacustrine water levels dropped during dry seasons. These charophyte algae probably already developed filamentous thalli and holdfasts that superficially resembled plant stems and roots, and probably had an isomorphic alternation of generations. They perhaps evolved some 850 mya and might even be as early as 1 Gya during the late phase of the Boring Billion.
Morphology
A range of algal morphologies is exhibited, and convergence of features in unrelated groups is common. The only groups to exhibit three-dimensional multicellular thalli are the reds and browns, and some chlorophytes. Apical growth is constrained to subsets of these groups: the florideophyte reds, various browns, and the charophytes. The form of charophytes is quite different from those of reds and browns, because they have distinct nodes, separated by internode 'stems'; whorls of branches reminiscent of the horsetails occur at the nodes. Conceptacles are another polyphyletic trait; they appear in the coralline algae and the Hildenbrandiales, as well as the browns.
Most of the simpler algae are unicellular flagellates or amoeboids, but colonial and nonmotile forms have developed independently among several of the groups. Some of the more common organizational levels, more than one of which may occur in the lifecycle of a species, are
Colonial: small, regular groups of motile cells
Capsoid: individual non-motile cells embedded in mucilage
Coccoid: individual non-motile cells with cell walls
Palmelloid: nonmotile cells embedded in mucilage
Filamentous: a string of connected nonmotile cells, sometimes branching
Parenchymatous: cells forming a thallus with partial differentiation of tissues | Algae | Wikipedia | 484 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
In three lines, even higher levels of organization have been reached, with full tissue differentiation. These are the brown algae,—some of which may reach 50 m in length (kelps)—the red algae, and the green algae. The most complex forms are found among the charophyte algae (see Charales and Charophyta), in a lineage that eventually led to the higher land plants. The innovation that defines these nonalgal plants is the presence of female reproductive organs with protective cell layers that protect the zygote and developing embryo. Hence, the land plants are referred to as the Embryophytes.
Turfs
The term algal turf is commonly used but poorly defined. Algal turfs are thick, carpet-like beds of seaweed that retain sediment and compete with foundation species like corals and kelps, and they are usually less than 15 cm tall. Such a turf may consist of one or more species, and will generally cover an area in the order of a square metre or more. Some common characteristics are listed:
Algae that form aggregations that have been described as turfs include diatoms, cyanobacteria, chlorophytes, phaeophytes and rhodophytes. Turfs are often composed of numerous species at a wide range of spatial scales, but monospecific turfs are frequently reported.
Turfs can be morphologically highly variable over geographic scales and even within species on local scales and can be difficult to identify in terms of the constituent species.
Turfs have been defined as short algae, but this has been used to describe height ranges from less than 0.5 cm to more than 10 cm. In some regions, the descriptions approached heights which might be described as canopies (20 to 30 cm).
Physiology
Many algae, particularly species of the Characeae, have served as model experimental organisms to understand the mechanisms of the water permeability of membranes, osmoregulation, salt tolerance, cytoplasmic streaming, and the generation of action potentials. Plant hormones are found not only in higher plants, but in algae, too.
Symbiotic algae
Some species of algae form symbiotic relationships with other organisms. In these symbioses, the algae supply photosynthates (organic substances) to the host organism providing protection to the algal cells. The host organism derives some or all of its energy requirements from the algae. Examples are:
Lichens | Algae | Wikipedia | 508 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Lichens are defined by the International Association for Lichenology to be "an association of a fungus and a photosynthetic symbiont resulting in a stable vegetative body having a specific structure". The fungi, or mycobionts, are mainly from the Ascomycota with a few from the Basidiomycota. In nature, they do not occur separate from lichens. It is unknown when they began to associate. One or more mycobiont associates with the same phycobiont species, from the green algae, except that alternatively, the mycobiont may associate with a species of cyanobacteria (hence "photobiont" is the more accurate term). A photobiont may be associated with many different mycobionts or may live independently; accordingly, lichens are named and classified as fungal species. The association is termed a morphogenesis because the lichen has a form and capabilities not possessed by the symbiont species alone (they can be experimentally isolated). The photobiont possibly triggers otherwise latent genes in the mycobiont.
Trentepohlia is an example of a common green alga genus worldwide that can grow on its own or be lichenised. Lichen thus share some of the habitat and often similar appearance with specialized species of algae (aerophytes) growing on exposed surfaces such as tree trunks and rocks and sometimes discoloring them.
Coral reefs
Coral reefs are accumulated from the calcareous exoskeletons of marine invertebrates of the order Scleractinia (stony corals). These animals metabolize sugar and oxygen to obtain energy for their cell-building processes, including secretion of the exoskeleton, with water and carbon dioxide as byproducts. Dinoflagellates (algal protists) are often endosymbionts in the cells of the coral-forming marine invertebrates, where they accelerate host-cell metabolism by generating sugar and oxygen immediately available through photosynthesis using incident light and the carbon dioxide produced by the host. Reef-building stony corals (hermatypic corals) require endosymbiotic algae from the genus Symbiodinium to be in a healthy condition. The loss of Symbiodinium from the host is known as coral bleaching, a condition which leads to the deterioration of a reef.
Sea sponges | Algae | Wikipedia | 505 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Endosymbiontic green algae live close to the surface of some sponges, for example, breadcrumb sponges (Halichondria panicea). The alga is thus protected from predators; the sponge is provided with oxygen and sugars which can account for 50 to 80% of sponge growth in some species.
Life cycle
Rhodophyta, Chlorophyta, and Heterokontophyta, the three main algal divisions, have life cycles which show considerable variation and complexity. In general, an asexual phase exists where the seaweed's cells are diploid, a sexual phase where the cells are haploid, followed by fusion of the male and female gametes. Asexual reproduction permits efficient population increases, but less variation is possible. Commonly, in sexual reproduction of unicellular and colonial algae, two specialized, sexually compatible, haploid gametes make physical contact and fuse to form a zygote. To ensure a successful mating, the development and release of gametes is highly synchronized and regulated; pheromones may play a key role in these processes. Sexual reproduction allows for more variation and provides the benefit of efficient recombinational repair of DNA damages during meiosis, a key stage of the sexual cycle. However, sexual reproduction is more costly than asexual reproduction. Meiosis has been shown to occur in many different species of algae.
Numbers
The Algal Collection of the US National Herbarium (located in the National Museum of Natural History) consists of approximately 320,500 dried specimens, which, although not exhaustive (no exhaustive collection exists), gives an idea of the order of magnitude of the number of algal species (that number remains unknown). Estimates vary widely. For example, according to one standard textbook, in the British Isles, the UK Biodiversity Steering Group Report estimated there to be 20,000 algal species in the UK. Another checklist reports only about 5,000 species. Regarding the difference of about 15,000 species, the text concludes: "It will require many detailed field surveys before it is possible to provide a reliable estimate of the total number of species ..." | Algae | Wikipedia | 450 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Regional and group estimates have been made, as well:
5,000–5,500 species of red algae worldwide
"some 1,300 in Australian Seas"
400 seaweed species for the western coastline of South Africa, and 212 species from the coast of KwaZulu-Natal. Some of these are duplicates, as the range extends across both coasts, and the total recorded is probably about 500 species. Most of these are listed in List of seaweeds of South Africa. These exclude phytoplankton and crustose corallines.
669 marine species from California (US)
642 in the check-list of Britain and Ireland
and so on, but lacking any scientific basis or reliable sources, these numbers have no more credibility than the British ones mentioned above. Most estimates also omit microscopic algae, such as phytoplankton.
The most recent estimate suggests 72,500 algal species worldwide.
Distribution
The distribution of algal species has been fairly well studied since the founding of phytogeography in the mid-19th century. Algae spread mainly by the dispersal of spores analogously to the dispersal of cryptogamic plants by spores. Spores can be found in a variety of environments: fresh and marine waters, air, soil, and in or on other organisms. Whether a spore is to grow into an adult organism depends on the species and the environmental conditions where the spore lands.
The spores of freshwater algae are dispersed mainly by running water and wind, as well as by living carriers. However, not all bodies of water can carry all species of algae, as the chemical composition of certain water bodies limits the algae that can survive within them. Marine spores are often spread by ocean currents. Ocean water presents many vastly different habitats based on temperature and nutrient availability, resulting in phytogeographic zones, regions, and provinces.
To some degree, the distribution of algae is subject to floristic discontinuities caused by geographical features, such as Antarctica, long distances of ocean or general land masses. It is, therefore, possible to identify species occurring by locality, such as "Pacific algae" or "North Sea algae". When they occur out of their localities, hypothesizing a transport mechanism is usually possible, such as the hulls of ships. For example, Ulva reticulata and U. fasciata travelled from the mainland to Hawaii in this manner. | Algae | Wikipedia | 488 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Mapping is possible for select species only: "there are many valid examples of confined distribution patterns." For example, Clathromorphum is an arctic genus and is not mapped far south of there. However, scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."
Ecology
Algae are prominent in bodies of water, common in terrestrial environments, and are found in unusual environments, such as on snow and ice. Seaweeds grow mostly in shallow marine waters, under deep; however, some such as Navicula pennata have been recorded to a depth of . A type of algae, Ancylonema nordenskioeldii, was found in Greenland in areas known as the 'Dark Zone', which caused an increase in the rate of melting ice sheet. The same algae was found in the Italian Alps, after pink ice appeared on parts of the Presena glacier.
The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live suspended in the water column (phytoplankton) provide the food base for most marine food chains. In very high densities (algal blooms), these algae may discolor the water and outcompete, poison, or asphyxiate other life forms.
Algae can be used as indicator organisms to monitor pollution in various aquatic systems. In many cases, algal metabolism is sensitive to various pollutants. Due to this, the species composition of algal populations may shift in the presence of chemical pollutants. To detect these changes, algae can be sampled from the environment and maintained in laboratories with relative ease.
On the basis of their habitat, algae can be categorized as: aquatic (planktonic, benthic, marine, freshwater, lentic, lotic), terrestrial, aerial (subaerial), lithophytic, halophytic (or euryhaline), psammon, thermophilic, cryophilic, epibiont (epiphytic, epizoic), endosymbiont (endophytic, endozoic), parasitic, calcifilic or lichenic (phycobiont). | Algae | Wikipedia | 453 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Cultural associations
In classical Chinese, the word is used both for "algae" and (in the modest tradition of the imperial scholars) for "literary talent". The third island in Kunming Lake beside the Summer Palace in Beijing is known as the Zaojian Tang Dao (藻鑒堂島), which thus simultaneously means "Island of the Algae-Viewing Hall" and "Island of the Hall for Reflecting on Literary Talent".
Cultivation
Seaweed farming
Bioreactors
Uses
Agar
Agar, a gelatinous substance derived from red algae, has a number of commercial uses. It is a good medium on which to grow bacteria and fungi, as most microorganisms cannot digest agar.
Alginates
Alginic acid, or alginate, is extracted from brown algae. Its uses range from gelling agents in food, to medical dressings. Alginic acid also has been used in the field of biotechnology as a biocompatible medium for cell encapsulation and cell immobilization. Molecular cuisine is also a user of the substance for its gelling properties, by which it becomes a delivery vehicle for flavours.
Between 100,000 and 170,000 wet tons of Macrocystis are harvested annually in New Mexico for alginate extraction and abalone feed.
Energy source
To be competitive and independent from fluctuating support from (local) policy on the long run, biofuels should equal or beat the cost level of fossil fuels. Here, algae-based fuels hold great promise, directly related to the potential to produce more biomass per unit area in a year than any other form of biomass. The break-even point for algae-based biofuels is estimated to occur by 2025.
Fertilizer
For centuries, seaweed has been used as a fertilizer; George Owen of Henllys writing in the 16th century referring to drift weed in South Wales:
Today, algae are used by humans in many ways; for example, as fertilizers, soil conditioners, and livestock feed. Aquatic and microscopic species are cultured in clear tanks or ponds and are either harvested or used to treat effluents pumped through the ponds. Algaculture on a large scale is an important type of aquaculture in some places. Maerl is commonly used as a soil conditioner.
As food | Algae | Wikipedia | 477 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Algae are used as foods in many countries: China consumes more than 70 species, including fat choy, a cyanobacterium considered a vegetable; Japan, over 20 species such as nori and aonori; Ireland, dulse; Chile, cochayuyo. Laver is used to make laverbread in Wales, where it is known as . In Korea, green laver is used to make .
Three forms of algae used as food:
Chlorella: This form of alga is found in freshwater and contains photosynthetic pigments in its chloroplast.
Klamath AFA: A subspecies of Aphanizomenon flos-aquae found wild in many bodies of water worldwide but harvested only from Upper Klamath Lake, Oregon.
Spirulina: Known otherwise as a cyanobacterium (a prokaryote or a "blue-green alga")
The oils from some algae have high levels of unsaturated fatty acids. Some varieties of algae favored by vegetarianism and veganism contain the long-chain, essential omega-3 fatty acids, docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA). Fish oil contains the omega-3 fatty acids, but the original source is algae (microalgae in particular), which are eaten by marine life such as copepods and are passed up the food chain.
Pollution control
Sewage can be treated with algae, reducing the use of large amounts of toxic chemicals that would otherwise be needed.
Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested, the enriched algae can be used as fertilizer.
Aquaria and ponds can be filtered using algae, which absorb nutrients from the water in a device called an algae scrubber, also known as an algae turf scrubber. | Algae | Wikipedia | 389 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Agricultural Research Service scientists found that 60–90% of nitrogen runoff and 70–100% of phosphorus runoff can be captured from manure effluents using a horizontal algae scrubber, also called an algal turf scrubber (ATS). Scientists developed the ATS, which consists of shallow, 100-foot raceways of nylon netting where algae colonies can form, and studied its efficacy for three years. They found that algae can readily be used to reduce the nutrient runoff from agricultural fields and increase the quality of water flowing into rivers, streams, and oceans. Researchers collected and dried the nutrient-rich algae from the ATS and studied its potential as an organic fertilizer. They found that cucumber and corn seedlings grew just as well using ATS organic fertilizer as they did with commercial fertilizers. Algae scrubbers, using bubbling upflow or vertical waterfall versions, are now also being used to filter aquaria and ponds.
Polymers
Various polymers can be created from algae, which can be especially useful in the creation of bioplastics. These include hybrid plastics, cellulose-based plastics, poly-lactic acid, and bio-polyethylene. Several companies have begun to produce algae polymers commercially, including for use in flip-flops and in surf boards.
Bioremediation
The alga Stichococcus bacillaris has been seen to colonize silicone resins used at archaeological sites; biodegrading the synthetic substance.
Pigments
The natural pigments (carotenoids and chlorophylls) produced by algae can be used as alternatives to chemical dyes and coloring agents.
The presence of some individual algal pigments, together with specific pigment concentration ratios, are taxon-specific: analysis of their concentrations with various analytical methods, particularly high-performance liquid chromatography, can therefore offer deep insight into the taxonomic composition and relative abundance of natural algae populations in sea water samples.
Stabilizing substances
Carrageenan, from the red alga Chondrus crispus, is used as a stabilizer in milk products.
Additional images | Algae | Wikipedia | 433 | 633 | https://en.wikipedia.org/wiki/Algae | Biology and health sciences | Biology | null |
Analysis of variance (ANOVA) is a collection of statistical models and their associated estimation procedures (such as the "variation" among and between groups) used to analyze the differences between groups. It uses F-test by comparing variance between groups and taking noise, or assumed normal distribution of group, into consideration by dividing by variance between elements in a group. ANOVA was developed by the statistician Ronald Fisher. ANOVA is based on the law of total variance, where the observed variance in a particular variable is partitioned into components attributable to different sources of variation. In its simplest form, ANOVA provides a statistical test of whether two or more population means are equal, and therefore generalizes the t-test beyond two means. In other words, the ANOVA is used to test the difference between two or more means.
History
While the analysis of variance reached fruition in the 20th century, antecedents extend centuries into the past according to Stigler. These include hypothesis testing, the partitioning of sums of squares, experimental techniques and the additive model. Laplace was performing hypothesis testing in the 1770s. Around 1800, Laplace and Gauss developed the least-squares method for combining observations, which improved upon methods then used in astronomy and geodesy. It also initiated much study of the contributions to sums of squares. Laplace knew how to estimate a variance from a residual (rather than a total) sum of squares. By 1827, Laplace was using least squares methods to address ANOVA problems regarding measurements of atmospheric tides. Before 1800, astronomers had isolated observational errors resulting
from reaction times (the "personal equation") and had developed methods of reducing the errors. The experimental methods used in the study of the personal equation were later accepted by the emerging field of psychology which developed strong (full factorial) experimental methods to which randomization and blinding were soon added. An eloquent non-mathematical explanation of the additive effects model was available in 1885. | Analysis of variance | Wikipedia | 401 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Ronald Fisher introduced the term variance and proposed its formal analysis in a 1918 article on theoretical population genetics, The Correlation Between Relatives on the Supposition of Mendelian Inheritance. His first application of the analysis of variance to data analysis was published in 1921, Studies in Crop Variation I. This divided the variation of a time series into components representing annual causes and slow deterioration. Fisher's next piece, Studies in Crop Variation II, written with Winifred Mackenzie and published in 1923, studied the variation in yield across plots sown with different varieties and subjected to different fertiliser treatments. Analysis of variance became widely known after being included in Fisher's 1925 book Statistical Methods for Research Workers.
Randomization models were developed by several researchers. The first was published in Polish by Jerzy Neyman in 1923. | Analysis of variance | Wikipedia | 163 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Example
The analysis of variance can be used to describe otherwise complex relations among variables. A dog show provides an example. A dog show is not a random sampling of the breed: it is typically limited to dogs that are adult, pure-bred, and exemplary. A histogram of dog weights from a show is likely to be rather complicated, like the yellow-orange distribution shown in the illustrations. Suppose we wanted to predict the weight of a dog based on a certain set of characteristics of each dog. One way to do that is to explain the distribution of weights by dividing the dog population into groups based on those characteristics. A successful grouping will split dogs such that (a) each group has a low variance of dog weights (meaning the group is relatively homogeneous) and (b) the mean of each group is distinct (if two groups have the same mean, then it isn't reasonable to conclude that the groups are, in fact, separate in any meaningful way).
In the illustrations to the right, groups are identified as X1, X2, etc. In the first illustration, the dogs are divided according to the product (interaction) of two binary groupings: young vs old, and short-haired vs long-haired (e.g., group 1 is young, short-haired dogs, group 2 is young, long-haired dogs, etc.). Since the distributions of dog weight within each of the groups (shown in blue) has a relatively large variance, and since the means are very similar across groups, grouping dogs by these characteristics does not produce an effective way to explain the variation in dog weights: knowing which group a dog is in doesn't allow us to predict its weight much better than simply knowing the dog is in a dog show. Thus, this grouping fails to explain the variation in the overall distribution (yellow-orange).
An attempt to explain the weight distribution by grouping dogs as pet vs working breed and less athletic vs more athletic would probably be somewhat more successful (fair fit). The heaviest show dogs are likely to be big, strong, working breeds, while breeds kept as pets tend to be smaller and thus lighter. As shown by the second illustration, the distributions have variances that are considerably smaller than in the first case, and the means are more distinguishable. However, the significant overlap of distributions, for example, means that we cannot distinguish X1 and X2 reliably. Grouping dogs according to a coin flip might produce distributions that look similar. | Analysis of variance | Wikipedia | 507 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
An attempt to explain weight by breed is likely to produce a very good fit. All Chihuahuas are light and all St Bernards are heavy. The difference in weights between Setters and Pointers does not justify separate breeds. The analysis of variance provides the formal tools to justify these intuitive judgments. A common use of the method is the analysis of experimental data or the development of models. The method has some advantages over correlation: not all of the data must be numeric and one result of the method is a judgment in the confidence in an explanatory relationship.
Classes of models
There are three classes of models used in the analysis of variance, and these are outlined here.
Fixed-effects models
The fixed-effects model (class I) of analysis of variance applies to situations in which the experimenter applies one or more treatments to the subjects of the experiment to see whether the response variable values change. This allows the experimenter to estimate the ranges of response variable values that the treatment would generate in the population as a whole.
Random-effects models
Random-effects model (class II) is used when the treatments are not fixed. This occurs when the various factor levels are sampled from a larger population. Because the levels themselves are random variables, some assumptions and the method of contrasting the treatments (a multi-variable generalization of simple differences) differ from the fixed-effects model.
Mixed-effects models
A mixed-effects model (class III) contains experimental factors of both fixed and random-effects types, with appropriately different interpretations and analysis for the two types.
Example
Teaching experiments could be performed by a college or university department to find a good introductory textbook, with each text considered a treatment. The fixed-effects model would compare a list of candidate texts. The random-effects model would determine whether important differences exist among a list of randomly selected texts. The mixed-effects model would compare the (fixed) incumbent texts to randomly selected alternatives.
Defining fixed and random effects has proven elusive, with multiple competing definitions.
Assumptions
The analysis of variance has been studied from several approaches, the most common of which uses a linear model that relates the response to the treatments and blocks. Note that the model is linear in parameters but may be nonlinear across factor levels. Interpretation is easy when data is balanced across factors but much deeper understanding is needed for unbalanced data. | Analysis of variance | Wikipedia | 471 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Textbook analysis using a normal distribution
The analysis of variance can be presented in terms of a linear model, which makes the following assumptions about the probability distribution of the responses:
Independence of observations – this is an assumption of the model that simplifies the statistical analysis.
Normality – the distributions of the residuals are normal.
Equality (or "homogeneity") of variances, called homoscedasticity—the variance of data in groups should be the same.
The separate assumptions of the textbook model imply that the errors are independently, identically, and normally distributed for fixed effects models, that is, that the errors () are independent and
Randomization-based analysis
In a randomized controlled experiment, the treatments are randomly assigned to experimental units, following the experimental protocol. This randomization is objective and declared before the experiment is carried out. The objective random-assignment is used to test the significance of the null hypothesis, following the ideas of C. S. Peirce and Ronald Fisher. This design-based analysis was discussed and developed by Francis J. Anscombe at Rothamsted Experimental Station and by Oscar Kempthorne at Iowa State University. Kempthorne and his students make an assumption of unit treatment additivity, which is discussed in the books of Kempthorne and David R. Cox.
Unit-treatment additivity
In its simplest form, the assumption of unit-treatment additivity states that the observed response from experimental unit when receiving treatment can be written as the sum of the unit's response and the treatment-effect , that is
The assumption of unit-treatment additivity implies that, for every treatment , the th treatment has exactly the same effect on every experiment unit.
The assumption of unit treatment additivity usually cannot be directly falsified, according to Cox and Kempthorne. However, many consequences of treatment-unit additivity can be falsified. For a randomized experiment, the assumption of unit-treatment additivity implies that the variance is constant for all treatments. Therefore, by contraposition, a necessary condition for unit-treatment additivity is that the variance is constant.
The use of unit treatment additivity and randomization is similar to the design-based inference that is standard in finite-population survey sampling. | Analysis of variance | Wikipedia | 465 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Derived linear model
Kempthorne uses the randomization-distribution and the assumption of unit treatment additivity to produce a derived linear model, very similar to the textbook model discussed previously. The test statistics of this derived linear model are closely approximated by the test statistics of an appropriate normal linear model, according to approximation theorems and simulation studies. However, there are differences. For example, the randomization-based analysis results in a small but (strictly) negative correlation between the observations. In the randomization-based analysis, there is no assumption of a normal distribution and certainly no assumption of independence. On the contrary, the observations are dependent!
The randomization-based analysis has the disadvantage that its exposition involves tedious algebra and extensive time. Since the randomization-based analysis is complicated and is closely approximated by the approach using a normal linear model, most teachers emphasize the normal linear model approach. Few statisticians object to model-based analysis of balanced randomized experiments.
Statistical models for observational data
However, when applied to data from non-randomized experiments or observational studies, model-based analysis lacks the warrant of randomization. For observational data, the derivation of confidence intervals must use subjective models, as emphasized by Ronald Fisher and his followers. In practice, the estimates of treatment-effects from observational studies generally are often inconsistent. In practice, "statistical models" and observational data are useful for suggesting hypotheses that should be treated very cautiously by the public.
Summary of assumptions
The normal-model based ANOVA analysis assumes the independence, normality, and homogeneity of variances of the residuals. The randomization-based analysis assumes only the homogeneity of the variances of the residuals (as a consequence of unit-treatment additivity) and uses the randomization procedure of the experiment. Both these analyses require homoscedasticity, as an assumption for the normal-model analysis and as a consequence of randomization and additivity for the randomization-based analysis.
However, studies of processes that change variances rather than means (called dispersion effects) have been successfully conducted using ANOVA. There are no necessary assumptions for ANOVA in its full generality, but the F-test used for ANOVA hypothesis testing has assumptions and practical
limitations which are of continuing interest. | Analysis of variance | Wikipedia | 478 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Problems which do not satisfy the assumptions of ANOVA can often be transformed to satisfy the assumptions.
The property of unit-treatment additivity is not invariant under a "change of scale", so statisticians often use transformations to achieve unit-treatment additivity. If the response variable is expected to follow a parametric family of probability distributions, then the statistician may specify (in the protocol for the experiment or observational study) that the responses be transformed to stabilize the variance. Also, a statistician may specify that logarithmic transforms be applied to the responses which are believed to follow a multiplicative model.
According to Cauchy's functional equation theorem, the logarithm is the only continuous transformation that transforms real multiplication to addition.
Characteristics
ANOVA is used in the analysis of comparative experiments, those in which only the difference in outcomes is of interest. The statistical significance of the experiment is determined by a ratio of two variances. This ratio is independent of several possible alterations to the experimental observations: Adding a constant to all observations does not alter significance. Multiplying all observations by a constant does not alter significance. So ANOVA statistical significance result is independent of constant bias and scaling errors as well as the units used in expressing observations. In the era of mechanical calculation it was common to subtract a constant from all observations (when equivalent to dropping leading digits) to simplify data entry. This is an example of data coding.
Algorithm
The calculations of ANOVA can be characterized as computing a number of means and variances, dividing two variances and comparing the ratio to a handbook value to determine statistical significance. Calculating a treatment effect is then trivial: "the effect of any treatment is estimated by taking the difference between the mean of the observations which receive the treatment and the general mean".
Partitioning of the sum of squares | Analysis of variance | Wikipedia | 378 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
ANOVA uses traditional standardized terminology. The definitional equation of sample variance is , where the divisor is called the degrees of freedom (DF), the summation is called
the sum of squares (SS), the result is called the mean square (MS) and the squared terms are deviations from the sample mean. ANOVA estimates 3 sample variances: a total variance based on all the observation deviations from the grand mean, an error variance based on all the observation deviations from their appropriate treatment means, and a treatment variance. The treatment variance is based on the deviations of treatment means from the grand mean, the result being multiplied by the number of observations in each treatment to account for the difference between the variance of observations and the variance of means.
The fundamental technique is a partitioning of the total sum of squares SS into components related to the effects used in the model. For example, the model for a simplified ANOVA with one type of treatment at different levels.
The number of degrees of freedom DF can be partitioned in a similar way: one of these components (that for error) specifies a chi-squared distribution which describes the associated sum of squares, while the same is true for "treatments" if there is no treatment effect.
The F-test
The F-test is used for comparing the factors of the total deviation. For example, in one-way, or single-factor ANOVA, statistical significance is tested for by comparing the F test statistic
where MS is mean square, is the number of treatments and is the total number of cases
to the F-distribution with being the numerator degrees of freedom and the denominator degrees of freedom. Using the F-distribution is a natural candidate because the test statistic is the ratio of two scaled sums of squares each of which follows a scaled chi-squared distribution.
The expected value of F is (where is the treatment sample size) which is 1 for no treatment effect. As values of F increase above 1, the evidence is increasingly inconsistent with the null hypothesis. Two apparent experimental methods of increasing F are increasing the sample size and reducing the error variance by tight experimental controls. | Analysis of variance | Wikipedia | 439 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
There are two methods of concluding the ANOVA hypothesis test, both of which produce the same result:
The textbook method is to compare the observed value of F with the critical value of F determined from tables. The critical value of F is a function of the degrees of freedom of the numerator and the denominator and the significance level (α). If F ≥ FCritical, the null hypothesis is rejected.
The computer method calculates the probability (p-value) of a value of F greater than or equal to the observed value. The null hypothesis is rejected if this probability is less than or equal to the significance level (α).
The ANOVA F-test is known to be nearly optimal in the sense of minimizing false negative errors for a fixed rate of false positive errors (i.e. maximizing power for a fixed significance level). For example, to test the hypothesis that various medical treatments have exactly the same effect, the F-test's p-values closely approximate the permutation test's p-values: The approximation is particularly close when the design is balanced. Such permutation tests characterize tests with maximum power against all alternative hypotheses, as observed by Rosenbaum. The ANOVA F-test (of the null-hypothesis that all treatments have exactly the same effect) is recommended as a practical test, because of its robustness against many alternative distributions.
Extended algorithm
ANOVA consists of separable parts; partitioning sources of variance and hypothesis testing can be used individually. ANOVA is used to support other statistical tools. Regression is first used to fit more complex models to data, then ANOVA is used to compare models with the objective of selecting simple(r) models that adequately describe the data. "Such models could be fit without any reference to ANOVA, but ANOVA tools could then be used to make some sense of the fitted models, and to test hypotheses about batches of coefficients." "[W]e think of the analysis of variance as a way of understanding and structuring multilevel models—not as an alternative to regression but as a tool for summarizing complex high-dimensional inferences ..."
For a single factor | Analysis of variance | Wikipedia | 456 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
The simplest experiment suitable for ANOVA analysis is the completely randomized experiment with a single factor. More complex experiments with a single factor involve constraints on randomization and include completely randomized blocks and Latin squares (and variants: Graeco-Latin squares, etc.). The more complex experiments share many of the complexities of multiple factors.
There are some alternatives to conventional one-way analysis of variance, e.g.: Welch's heteroscedastic F test, Welch's heteroscedastic F test with trimmed means and Winsorized variances, Brown-Forsythe test, Alexander-Govern test, James second order test and Kruskal-Wallis test, available in onewaytests R
It is useful to represent each data point in the following form, called a statistical model:
where
i = 1, 2, 3, ..., R
j = 1, 2, 3, ..., C
μ = overall average (mean)
τj = differential effect (response) associated with the j level of X; this assumes that overall the values of τj add to zero (that is, )
εij = noise or error associated with the particular ij data value
That is, we envision an additive model that says every data point can be represented by summing three quantities: the true mean, averaged over all factor levels being investigated, plus an incremental component associated with the particular column (factor level), plus a final component associated with everything else affecting that specific data value.
For multiple factors
ANOVA generalizes to the study of the effects of multiple factors. When the experiment includes observations at all combinations of levels of each factor, it is termed factorial. Factorial experiments are more efficient than a series of single factor experiments and the efficiency grows as the number of factors increases. Consequently, factorial designs are heavily used. | Analysis of variance | Wikipedia | 380 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
The use of ANOVA to study the effects of multiple factors has a complication. In a 3-way ANOVA with factors x, y and z, the ANOVA model includes terms for the main effects (x, y, z) and terms for interactions (xy, xz, yz, xyz).
All terms require hypothesis tests. The proliferation of interaction terms increases the risk that some hypothesis test will produce a false positive by chance. Fortunately, experience says that high order interactions are rare.
The ability to detect interactions is a major advantage of multiple factor ANOVA. Testing one factor at a time hides interactions, but produces apparently inconsistent experimental results.
Caution is advised when encountering interactions; Test interaction terms first and expand the analysis beyond ANOVA if interactions are found. Texts vary in their recommendations regarding the continuation of the ANOVA procedure after encountering an interaction. Interactions complicate the interpretation of experimental data. Neither the calculations of significance nor the estimated treatment effects can be taken at face value. "A significant interaction will often mask the significance of main effects." Graphical methods are recommended to enhance understanding. Regression is often useful. A lengthy discussion of interactions is available in Cox (1958). Some interactions can be removed (by transformations) while others cannot.
A variety of techniques are used with multiple factor ANOVA to reduce expense. One technique used in factorial designs is to minimize replication (possibly no replication with support of analytical trickery) and to combine groups when effects are found to be statistically (or practically) insignificant. An experiment with many insignificant factors may collapse into one with a few factors supported by many replications.
Associated analysis
Some analysis is required in support of the design of the experiment while other analysis is performed after changes in the factors are formally found to produce statistically significant changes in the responses. Because experimentation is iterative, the results of one experiment alter plans for following experiments.
Preparatory analysis
The number of experimental units
In the design of an experiment, the number of experimental units is planned to satisfy the goals of the experiment. Experimentation is often sequential.
Early experiments are often designed to provide mean-unbiased estimates of treatment effects and of experimental error. Later experiments are often designed to test a hypothesis that a treatment effect has an important magnitude; in this case, the number of experimental units is chosen so that the experiment is within budget and has adequate power, among other goals. | Analysis of variance | Wikipedia | 488 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Reporting sample size analysis is generally required in psychology. "Provide information on sample size and the process that led to sample size decisions." The analysis, which is written in the experimental protocol before the experiment is conducted, is examined in grant applications and administrative review boards.
Besides the power analysis, there are less formal methods for selecting the number of experimental units. These include graphical methods based on limiting the probability of false negative errors, graphical methods based on an expected variation increase (above the residuals) and methods based on achieving a desired confidence interval.
Power analysis
Power analysis is often applied in the context of ANOVA in order to assess the probability of successfully rejecting the null hypothesis if we assume a certain ANOVA design, effect size in the population, sample size and significance level. Power analysis can assist in study design by determining what sample size would be required in order to have a reasonable chance of rejecting the null hypothesis when the alternative hypothesis is true.
Effect size
Several standardized measures of effect have been proposed for ANOVA to summarize the strength of the association between a predictor(s) and the dependent variable or the overall standardized difference of the complete model. Standardized effect-size estimates facilitate comparison of findings across studies and disciplines. However, while standardized effect sizes are commonly used in much of the professional literature, a non-standardized measure of effect size that has immediately "meaningful" units may be preferable for reporting purposes.
Model confirmation
Sometimes tests are conducted to determine whether the assumptions of ANOVA appear to be violated. Residuals are examined or analyzed to confirm homoscedasticity and gross normality. Residuals should have the appearance of (zero mean normal distribution) noise when plotted as a function of anything including time and
modeled data values. Trends hint at interactions among factors or among observations.
Follow-up tests
A statistically significant effect in ANOVA is often followed by additional tests. This can be done in order to assess which groups are different from which other groups or to test various other focused hypotheses. Follow-up tests are often distinguished in terms of whether they are "planned" (a priori) or "post hoc." Planned tests are determined before looking at the data, and post hoc tests are conceived only after looking at the data (though the term "post hoc" is inconsistently used). | Analysis of variance | Wikipedia | 469 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
The follow-up tests may be "simple" pairwise comparisons of individual group means or may be "compound" comparisons (e.g., comparing the mean pooling across groups A, B and C to the mean of group D). Comparisons can also look at tests of trend, such as linear and quadratic relationships, when the independent variable involves ordered levels. Often the follow-up tests incorporate a method of adjusting for the multiple comparisons problem.
Follow-up tests to identify which specific groups, variables, or factors have statistically different means include the Tukey's range test, and Duncan's new multiple range test. In turn, these tests are often followed with a Compact Letter Display (CLD) methodology in order to render the output of the mentioned tests more transparent to a non-statistician audience.
Study designs
There are several types of ANOVA. Many statisticians base ANOVA on the design of the experiment, especially on the protocol that specifies the random assignment of treatments to subjects; the protocol's description of the assignment mechanism should include a specification of the structure of the treatments and of any blocking. It is also common to apply ANOVA to observational data using an appropriate statistical model. | Analysis of variance | Wikipedia | 248 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Some popular designs use the following types of ANOVA:
One-way ANOVA is used to test for differences among two or more independent groups (means), e.g. different levels of urea application in a crop, or different levels of antibiotic action on several different bacterial species, or different levels of effect of some medicine on groups of patients. However, should these groups not be independent, and there is an order in the groups (such as mild, moderate and severe disease), or in the dose of a drug (such as 5 mg/mL, 10 mg/mL, 20 mg/mL) given to the same group of patients, then a linear trend estimation should be used. Typically, however, the one-way ANOVA is used to test for differences among at least three groups, since the two-group case can be covered by a t-test. When there are only two means to compare, the t-test and the ANOVA F-test are equivalent; the relation between ANOVA and t is given by .
Factorial ANOVA is used when there is more than one factor.
Repeated measures ANOVA is used when the same subjects are used for each factor (e.g., in a longitudinal study).
Multivariate analysis of variance (MANOVA) is used when there is more than one response variable.
Cautions
Balanced experiments (those with an equal sample size for each treatment) are relatively easy to interpret; unbalanced experiments offer more complexity. For single-factor (one-way) ANOVA, the adjustment for unbalanced data is easy, but the unbalanced analysis lacks both robustness and power. For more complex designs the lack of balance leads to further complications. "The orthogonality property of main effects and interactions present in balanced data does not carry over to the unbalanced case. This means that the usual analysis of variance techniques do not apply. Consequently, the analysis of unbalanced factorials is much more difficult than that for balanced designs." In the general case, "The analysis of variance can also be applied to unbalanced data, but then the sums of squares, mean squares, and F-ratios will depend on the order in which the sources of variation are considered."
ANOVA is (in part) a test of statistical significance. The American Psychological Association (and many other organisations) holds the view that simply reporting statistical significance is insufficient and that reporting confidence bounds is preferred. | Analysis of variance | Wikipedia | 501 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
Generalizations
ANOVA is considered to be a special case of linear regression which in turn is a special case of the general linear model. All consider the observations to be the sum of a model (fit) and a residual (error) to be minimized.
The Kruskal-Wallis test and the Friedman test are nonparametric tests which do not rely on an assumption of normality.
Connection to linear regression
Below we make clear the connection between multi-way ANOVA and linear regression.
Linearly re-order the data so that -th observation is associated with a response and factors where denotes the different factors and is the total number of factors. In one-way ANOVA and in two-way ANOVA . Furthermore, we assume the -th factor has levels, namely . Now, we can one-hot encode the factors into the dimensional vector .
The one-hot encoding function is defined such that the -th entry of is
The vector is the concatenation of all of the above vectors for all . Thus, . In order to obtain a fully general -way interaction ANOVA we must also concatenate every additional interaction term in the vector and then add an intercept term. Let that vector be .
With this notation in place, we now have the exact connection with linear regression. We simply regress response against the vector . However, there is a concern about identifiability. In order to overcome such issues we assume that the sum of the parameters within each set of interactions is equal to zero. From here, one can use F-statistics or other methods to determine the relevance of the individual factors.
Example
We can consider the 2-way interaction example where we assume that the first factor has 2 levels and the second factor has 3 levels.
Define if and if , i.e. is the one-hot encoding of the first factor and is the one-hot encoding of the second factor.
With that,
where the last term is an intercept term. For a more concrete example suppose that
Then, | Analysis of variance | Wikipedia | 409 | 634 | https://en.wikipedia.org/wiki/Analysis%20of%20variance | Mathematics | Statistics | null |
In organic chemistry, an alkane, or paraffin (a historical trivial name that also has other meanings), is an acyclic saturated hydrocarbon. In other words, an alkane consists of hydrogen and carbon atoms arranged in a tree structure in which all the carbon–carbon bonds are single. Alkanes have the general chemical formula . The alkanes range in complexity from the simplest case of methane (), where n = 1 (sometimes called the parent molecule), to arbitrarily large and complex molecules, like pentacontane () or 6-ethyl-2-methyl-5-(1-methylethyl) octane, an isomer of tetradecane ().
The International Union of Pure and Applied Chemistry (IUPAC) defines alkanes as "acyclic branched or unbranched hydrocarbons having the general formula , and therefore consisting entirely of hydrogen atoms and saturated carbon atoms". However, some sources use the term to denote any saturated hydrocarbon, including those that are either monocyclic (i.e. the cycloalkanes) or polycyclic, despite them having a distinct general formula (e.g. cycloalkanes are ).
In an alkane, each carbon atom is sp3-hybridized with 4 sigma bonds (either C–C or C–H), and each hydrogen atom is joined to one of the carbon atoms (in a C–H bond). The longest series of linked carbon atoms in a molecule is known as its carbon skeleton or carbon backbone. The number of carbon atoms may be considered as the size of the alkane.
One group of the higher alkanes are waxes, solids at standard ambient temperature and pressure (SATP), for which the number of carbon atoms in the carbon backbone is greater than about 17.
With their repeated – units, the alkanes constitute a homologous series of organic compounds in which the members differ in molecular mass by multiples of 14.03 u (the total mass of each such methylene-bridge unit, which comprises a single carbon atom of mass 12.01 u and two hydrogen atoms of mass ~1.01 u each). | Alkane | Wikipedia | 466 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
Methane is produced by methanogenic bacteria and some long-chain alkanes function as pheromones in certain animal species or as protective waxes in plants and fungi. Nevertheless, most alkanes do not have much biological activity. They can be viewed as molecular trees upon which can be hung the more active/reactive functional groups of biological molecules.
The alkanes have two main commercial sources: petroleum (crude oil) and natural gas.
An alkyl group is an alkane-based molecular fragment that bears one open valence for bonding. They are generally abbreviated with the symbol for any organyl group, R, although Alk is sometimes used to specifically symbolize an alkyl group (as opposed to an alkenyl group or aryl group).
Structure and classification
Ordinarily the C-C single bond distance is .
Saturated hydrocarbons can be linear, branched, or cyclic. The third group is sometimes called cycloalkanes. Very complicated structures are possible by combining linear, branch, cyclic alkanes.
Isomerism | Alkane | Wikipedia | 218 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
Alkanes with more than three carbon atoms can be arranged in various ways, forming structural isomers. The simplest isomer of an alkane is the one in which the carbon atoms are arranged in a single chain with no branches. This isomer is sometimes called the n-isomer (n for "normal", although it is not necessarily the most common). However, the chain of carbon atoms may also be branched at one or more points. The number of possible isomers increases rapidly with the number of carbon atoms. For example, for acyclic alkanes:
C1: methane only
C2: ethane only
C3: propane only
C4: 2 isomers: butane and isobutane
C5: 3 isomers: pentane, isopentane, and neopentane
C6: 5 isomers: hexane, 2-methylpentane, 3-methylpentane, 2,2-dimethylbutane, and 2,3-dimethylbutane
C7: 9 isomers: heptane, 2-methylhexane, 3-methylhexane, 2,2-dimethylpentane, 2,3-dimethylpentane, 2,4-dimethylpentane, 3,3-dimethylpentane, 3-ethylpentane, 2,2,3-trimethylbutane
C8: 18 isomers: octane, 2-methylheptane, 3-methylheptane, 4-methylheptane, 2,2-dimethylhexane, 2,3-dimethylhexane, 2,4-dimethylhexane, 2,5-dimethylhexane, 3,3-dimethylhexane, 3,4-dimethylhexane, 3-ethylhexane, 2,2,3-trimethylpentane, 2,2,4-trimethylpentane, 2,3,3-trimethylpentane, 2,3,4-trimethylpentane, 3-ethyl-2-methylpentane, 3-ethyl-3-methylpentane, 2,2,3,3-tetramethylbutane
C9: 35 isomers
C10: 75 isomers | Alkane | Wikipedia | 508 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
C12: 355 isomers
C32: 27,711,253,769 isomers
C60: 22,158,734,535,770,411,074,184 isomers, many of which are not stable | Alkane | Wikipedia | 50 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
Branched alkanes can be chiral. For example, 3-methylhexane and its higher homologues are chiral due to their stereogenic center at carbon atom number 3. The above list only includes differences of connectivity, not stereochemistry. In addition to the alkane isomers, the chain of carbon atoms may form one or more rings. Such compounds are called cycloalkanes, and are also excluded from the above list because changing the number of rings changes the molecular formula. For example, cyclobutane and methylcyclopropane are isomers of each other (C4H8), but are not isomers of butane (C4H10).
Branched alkanes are more thermodynamically stable than their linear (or less branched) isomers. For example, the highly branched 2,2,3,3-tetramethylbutane is about 1.9 kcal/mol more stable than its linear isomer, n-octane.
Nomenclature
The IUPAC nomenclature (systematic way of naming compounds) for alkanes is based on identifying hydrocarbon chains. Unbranched, saturated hydrocarbon chains are named systematically with a Greek numerical prefix denoting the number of carbons and the suffix "-ane".
In 1866, August Wilhelm von Hofmann suggested systematizing nomenclature by using the whole sequence of vowels a, e, i, o and u to create suffixes -ane, -ene, -ine (or -yne), -one, -une, for the hydrocarbons CnH2n+2, CnH2n, CnH2n−2, CnH2n−4, CnH2n−6. In modern nomenclature, the first three specifically name hydrocarbons with single, double and triple bonds; while "-one" now represents a ketone.
Linear alkanes | Alkane | Wikipedia | 403 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
Straight-chain alkanes are sometimes indicated by the prefix "n-" or "n-"(for "normal") where a non-linear isomer exists. Although this is not strictly necessary and is not part of the IUPAC naming system, the usage is still common in cases where one wishes to emphasize or distinguish between the straight-chain and branched-chain isomers, e.g., "n-butane" rather than simply "butane" to differentiate it from isobutane. Alternative names for this group used in the petroleum industry are linear paraffins or n-paraffins.
The first eight members of the series (in terms of number of carbon atoms) are named as follows:
methane CH4 – one carbon and 4 hydrogen
ethane C2H6 – two carbon and 6 hydrogen
propane C3H8 – three carbon and 8 hydrogen
butane C4H10 – four carbon and 10 hydrogen
pentane C5H12 – five carbon and 12 hydrogen
hexane C6H14 – six carbon and 14 hydrogen
heptane C7H16 – seven carbons and 16 hydrogen
octane C8H18 – eight carbons and 18 hydrogen
The first four names were derived from methanol, ether, propionic acid and butyric acid. Alkanes with five or more carbon atoms are named by adding the suffix -ane to the appropriate numerical multiplier prefix with elision of any terminal vowel (-a or -o) from the basic numerical term. Hence, pentane, C5H12; hexane, C6H14; heptane, C7H16; octane, C8H18; etc. The numeral prefix is generally Greek; however, alkanes with a carbon atom count ending in nine, for example nonane, use the Latin prefix non-.
Branched alkanes
Simple branched alkanes often have a common name using a prefix to distinguish them from linear alkanes, for example n-pentane, isopentane, and neopentane.
IUPAC naming conventions can be used to produce a systematic name. | Alkane | Wikipedia | 448 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
The key steps in the naming of more complicated branched alkanes are as follows:
Identify the longest continuous chain of carbon atoms
Name this longest root chain using standard naming rules
Name each side chain by changing the suffix of the name of the alkane from "-ane" to "-yl"
Number the longest continuous chain in order to give the lowest possible numbers for the side-chains
Number and name the side chains before the name of the root chain
If there are multiple side chains of the same type, use prefixes such as "di-" and "tri-" to indicate it as such, and number each one.
Add side chain names in alphabetical (disregarding "di-" etc. prefixes) order in front of the name of the root chain
Saturated cyclic hydrocarbons
Though technically distinct from the alkanes, this class of hydrocarbons is referred to by some as the "cyclic alkanes." As their description implies, they contain one or more rings.
Simple cycloalkanes have a prefix "cyclo-" to distinguish them from alkanes. Cycloalkanes are named as per their acyclic counterparts with respect to the number of carbon atoms in their backbones, e.g., cyclopentane (C5H10) is a cycloalkane with 5 carbon atoms just like pentane (C5H12), but they are joined up in a five-membered ring. In a similar manner, propane and cyclopropane, butane and cyclobutane, etc.
Substituted cycloalkanes are named similarly to substituted alkanes – the cycloalkane ring is stated, and the substituents are according to their position on the ring, with the numbering decided by the Cahn–Ingold–Prelog priority rules.
Trivial/common names
The trivial (non-systematic) name for alkanes is 'paraffins'. Together, alkanes are known as the 'paraffin series'. Trivial names for compounds are usually historical artifacts. They were coined before the development of systematic names, and have been retained due to familiar usage in industry. Cycloalkanes are also called naphthenes. | Alkane | Wikipedia | 470 | 639 | https://en.wikipedia.org/wiki/Alkane | Physical sciences | Hydrocarbons | null |
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