{{Short description|Reason for a change under natural selection; in physiology, what a system does}} {{good article}} {{Use dmy dates|date=April 2023}} {{CS1 config|mode=cs1}} In evolutionary biology, '''function''' is the reason some object or process occurred in a system that evolved through natural selection. That reason is typically that it achieves some result, such as that chlorophyll helps to capture the energy of sunlight in photosynthesis. Hence, the organism that contains it is more likely to survive and reproduce, in other words the function increases the organism's fitness. A characteristic that assists in evolution is called an adaptation; other characteristics may be non-functional spandrels, though these in turn may later be co-opted by evolution to serve new functions.
In biology, function has been defined in many ways. In physiology, it is simply what an organ, tissue, cell or molecule does.
In the philosophy of biology, talk of function inevitably suggests some kind of teleological purpose, even though natural selection operates without any goal for the future. All the same, biologists often use teleological language as a shorthand for function. In contemporary philosophy of biology, there are three major accounts of function in the biological world: theories of causal role, selected effect, and goal contribution.
==In pre-evolutionary biology== {{further|Four causes|Aristotle's biology}}
In physiology, a function is an activity or process carried out by a system in an organism, such as sensation or locomotion in an animal.<ref name = Fletcher>{{cite book |author=Fletcher, John |date=1837 |title=On the functions of organized beings, and their arrangement |work=Rudiments of physiology, Part 2. On life, as manifested in irritation |publisher=John Carfrae & Son |pages=[https://archive.org/details/b21301608/page/n206 1]–15 |url=https://archive.org/details/b21301608}}</ref> This concept of function as opposed to form (respectively Aristotle's ''ergon'' and ''morphê''<ref>{{cite book |author=Tipton, Jason A. |date=2014 |title=Philosophical Biology in Aristotle's ''Parts of Animals'' |publisher=Springer |page=33 |url=https://books.google.com/books?id=zrG8BAAAQBAJ&pg=PA33 |isbn=978-3-319-01421-0}} citing ''The Parts of Animals'' 640–641.</ref>) was central in biological explanations in classical antiquity. In more modern times it formed part of the 1830 Cuvier–Geoffroy debate, where Cuvier argued that an animal's structure was driven by its functional needs, while Geoffroy proposed that each animal's structure was modified from a common plan.<ref>{{cite book |author-link=E. S. Russell |author=Russell, Edward Stewart |date=1916 |title=Form and Function: A Contribution to the History of Animal Morphology |publisher=John Murray |url=https://www.biodiversitylibrary.org/bibliography/3747#/summary |access-date=2018-01-14 |archive-date=2023-06-12 |archive-url=https://web.archive.org/web/20230612153307/https://www.biodiversitylibrary.org/bibliography/3747#/summary |url-status=live }}</ref><ref>{{cite book |author=Asma, S. T. |date=1996 |title=Following form and function: A philosophical archaeology of life science |publisher=Northwestern University Press |url=https://books.google.com/books?id=deLaAAAAMAAJ|isbn=9780810113978 }}</ref><ref>{{cite book |author-link=Agnes Arber |author=Arber, Agnes |date=1950 |title=The Natural Philosophy of Plant Form |publisher=Cambridge University Press |url=https://books.google.com/books?id=Dvc8AAAAIAAJ}}</ref>
==In evolutionary biology== Function can be defined in a variety of ways,<ref>{{cite book |last=Toepfer |first=G. |year=2011 |chapter=Funktion |title=Historisches Wörterbuch der Biologie. Geschichte und Theorie der biologischen Grundbegriffe |language=de |publisher=Metzler |volume=1 |page=644 |url=http://www.zfl-berlin.org/tl_files/zfl/downloads/personen/toepfer/Histor_WoeBuch_Biologie/Band1.pdf |access-date=2018-05-04 |archive-date=2023-06-12 |archive-url=https://web.archive.org/web/20230612153313/https://www.zfl-berlin.org/tl_files/zfl/downloads/personen/toepfer/Histor_WoeBuch_Biologie/Band1.pdf |url-status=live }}</ref><ref>{{cite web |last=Toepfer |first=G. |title=Function |work=BioConcepts: The Origin and Definition of Biological Concepts |publisher=Das Zentrum für Literatur- und Kulturforschung Berlin |url=http://www.biological-concepts.com/views/search.php?term=559&listed=y |access-date=4 May 2018 |archive-date=12 June 2023 |archive-url=https://web.archive.org/web/20230612153312/http://www.biological-concepts.com/views/search.php?term=559&listed=y |url-status=dead }}</ref> including as adaptation,<ref name=Berkeley/> as contributing to evolutionary fitness,<ref name=Zimmer/> in animal behaviour,<ref name=Hladky/> and, as discussed below, also as some kind of causal role or goal in the philosophy of biology.<ref name=Stanford/>
===Adaptation=== {{main|Adaptation}}
A functional characteristic is known in evolutionary biology as an adaptation, and the research strategy for investigating whether a character is adaptive is known as adaptationism. Although assuming that a character is functional may be helpful in research, some characteristics of organisms are non-functional, formed as accidental spandrels, side effects of neighbouring functional systems.<ref name=Berkeley>{{cite web |title=Understanding Evolution: Qualifying as an adaptation |url=https://evolution.berkeley.edu/evolibrary/article/side_0_0/qualifying_01 |publisher=University of California at Berkeley |access-date=29 July 2016 |archive-date=7 November 2017 |archive-url=https://web.archive.org/web/20171107112919/https://evolution.berkeley.edu/evolibrary/article/side_0_0/qualifying_01 |url-status=live }}</ref>
===Natural selection=== [[File:Chlorophyll-a-3D-vdW.png|thumb|Chlorophyll molecule has a function in photosynthesis.]] {{main|Natural selection}}
From the point of view of natural selection, biological functions exist to contribute to fitness, increasing the chance that an organism will survive to reproduce.<ref name=Zimmer>{{cite book |last1=Zimmer |first1=Carl |author-link1=Carl Zimmer |last2=Emlen |first2=Douglas J. |author-link2=Douglas Emlen |year=2013 |title=Evolution: Making Sense of Life |edition=1st |publisher=Roberts and Company Publishers |isbn=978-1-936221-17-2}}</ref><ref>{{cite book |title=Strickberger's Evolution |edition=4th |author1=Hall, Brian K. |author2=Hallgrímsson, Benedikt |publisher=Jones and Bartlett |date=2008 |pages=4–6 |url=https://books.google.com/books?id=jrDD3cyA09kC&pg=PA4|isbn=9781449647223 }}</ref> For example, the function of chlorophyll in a plant is to capture the energy of sunlight for photosynthesis,<ref>{{cite web | url=http://biology.clc.uc.edu/courses/bio104/photosyn.htm | title=Photosynthesis | author=Carter, J. Stein | publisher=University of Cincinnati | year=1996 | url-status=dead | archive-url=https://web.archive.org/web/20130629204107/http://biology.clc.uc.edu/courses/bio104/photosyn.htm | archive-date =2013-06-29 }}</ref> which contributes to evolutionary success.<ref name="Shih2015">{{cite journal |last1=Shih |first1=Patrick M. |title=Photosynthesis and early Earth |journal=Current Biology |volume=25 |issue=19 |year=2015 |pages=R855–R859 |doi=10.1016/j.cub.2015.04.046 |pmid=26439346 |quote=Photosynthesis has been instrumental in the success of life on Earth|doi-access=free |bibcode=2015CBio...25.R855S }}</ref>
==In ethology== {{main article|Tinbergen's four questions}}
The ethologist Niko Tinbergen named four questions, based on Aristotle's Four Causes,<ref name=Hladky>{{cite journal |author1=Hladký, V. |author2=Havlíček, J. |date=2013 |url=http://ishe.org/wp-content/uploads/2015/04/HEB_2013_28_4_3-11.pdf |title=Was Tinbergen an Aristotelian? Comparison of Tinbergen's Four Whys and Aristotle's Four Causes |journal=Human Ethology Bulletin |volume=28 |issue=4 |pages=3–11 |access-date=2017-04-05 |archive-date=2017-04-04 |archive-url=https://web.archive.org/web/20170404215824/http://ishe.org/wp-content/uploads/2015/04/HEB_2013_28_4_3-11.pdf |url-status=live }}</ref> that a biologist could ask to help explain a behaviour, though they have been generalised to a wider scope. 1) Mechanism: What mechanisms cause the animal to behave as it does? 2) Ontogeny: What developmental mechanisms in the animal's embryology (and its youth, if it learns) created the structures that cause the behaviour? 3) Function/adaptation: What is the evolutionary function of the behaviour? 4) Evolution: What is the phylogeny of the behaviour, or in other words, when did it first appear in the evolutionary history of the animal? The questions are interdependent, so that, for example, adaptive function is constrained by embryonic development.<ref>{{cite web |title=Sociobiology |url=https://plato.stanford.edu/entries/sociobiology/ |publisher=Stanford Encyclopedia of Philosophy |access-date=4 April 2017 |date=11 November 2013 |archive-date=7 August 2019 |archive-url=https://web.archive.org/web/20190807134513/https://plato.stanford.edu/entries/sociobiology/ |url-status=live }}</ref><ref>{{Cite journal | last1=Tinbergen | first1=N. | author-link=Nikolaas Tinbergen | title=On aims and methods of Ethology | doi=10.1111/j.1439-0310.1963.tb01161.x | journal=Zeitschrift für Tierpsychologie | volume=20 | issue=4 | pages=410–433 | year=1963| bibcode=1963Ethol..20..410T }}</ref><ref>{{cite web |url=http://www-personal.umich.edu/~nesse/Nesse-Tinbergen4Q.PDF |title=The Four Areas of Biology |access-date=2 September 2018 |archive-date=18 January 2012 |archive-url=https://web.archive.org/web/20120118055035/http://www-personal.umich.edu/~nesse/Nesse-Tinbergen4Q.pdf |url-status=dead }}</ref><ref>{{cite web |url=http://homepage.uibk.ac.at/~c720126/humanethologie/ws/medicus/block1/inhalt.html |title=The Four Areas of Biology |access-date=2 September 2018 |archive-date=14 May 2017 |archive-url=https://web.archive.org/web/20170514163716/http://homepage.uibk.ac.at/~c720126/humanethologie/ws/medicus/block1/inhalt.html |url-status=live }}</ref>
==In philosophy of biology== {{main |Teleology in biology}} [[File:Springbok pronk.jpg|thumb|upright|"Behaviour with a purpose": a young springbok stotting.<ref name=Stanford/><ref name=Caro>{{cite journal | title=The functions of stotting in Thomson's gazelles: Some tests of the predictions | author=Caro, TM |author-link=Tim Caro | journal=Animal Behaviour | year=1986 | volume=34 | issue=3 | pages=663–684 | doi=10.1016/S0003-3472(86)80052-5| s2cid=53155678 }}</ref> A philosopher of biology might argue that this has the function of signalling to predators, helping the springbok to survive and allowing it to reproduce.<ref name=Stanford/>]]
Function is not the same as purpose in the teleological sense, that is, possessing conscious mental intention to achieve a goal. In the philosophy of biology, evolution is a blind process which has no 'goal' for the future. For example, a tree does not grow flowers for any purpose, but does so simply because it has evolved to do so. To say 'a tree grows flowers to attract pollinators' would be incorrect if the 'to' implies purpose. A function describes what something ''does'', not what its 'purpose' is. However, teleological language is often used by biologists as a shorthand way of describing function, even though its applicability is disputed.<ref name=Stanford>{{cite web |title=Teleological Notions in Biology |url=https://plato.stanford.edu/entries/teleology-biology/ |website=Stanford Encyclopedia of Philosophy |access-date=28 July 2016 |date=18 May 2003 |archive-date=13 March 2020 |archive-url=https://web.archive.org/web/20200313041857/https://plato.stanford.edu/entries/teleology-biology/ |url-status=live }}</ref>
In contemporary philosophy of biology, there are three major accounts of function in the biological world: theories of causal role,<ref name=Cummins1975/> selected effect,<ref name=Millikan1989/> and goal contribution.<ref name=Boorse1977/>
=== Causal role ===
Causal role theories of biological function trace their origin back to a 1975 paper by Robert Cummins.<ref name=Cummins1975>{{cite journal |last=Cummins |first=Robert |date=1975 |title=Functional Analysis |journal=The Journal of Philosophy |volume=72 |issue=20 |pages=741–765 |jstor=2024640|doi=10.2307/2024640 }}</ref> Cummins defines the functional role of a component of a system to be the causal effect that the component has on the larger containing system. For example, the heart has the actual causal role of pumping blood in the circulatory system; therefore, the function of the heart is to pump blood. This account has been objected to on the grounds that it is too loose a notion of function. For example, the heart also has the causal effect of producing a sound, but we would not consider producing sound to be the function of the heart.<ref>{{cite journal |last1=Amundson |first1=Ron |last2=Lauder |first2=George |date=1994 |title=Function Without Purpose |journal=Biology and Philosophy |volume=9 |issue=4 |pages=443–469 |doi=10.1007/BF00850375|s2cid=4997676 }}</ref><ref>{{cite journal |last=Craver |first=Carl F. |date=2001 |title=Role Functions, Mechanisms, and Hierarchy |journal=Philosophy of Science |volume=68 |issue=1 |pages=53–74 |jstor=3081024|doi=10.1086/392866 |s2cid=35230404 }}</ref>
=== Selected effect === {{further|Natural selection}}
Selected effect theories of biological functions hold that the function of a biological trait is the function that the trait was selected for, as argued by Ruth Millikan.<ref name=Millikan1989>{{cite journal |last=Millikan |first=Ruth |date=1989 |title=In Defense of Proper Functions |journal=Philosophy of Science |volume=56 |issue=2 |pages=288–302 |jstor=187875|doi=10.1086/289488 |s2cid=120910145 |doi-access=free }}</ref> For example, the function of the heart is pumping blood, for that is the action for which the heart was selected for by evolution. In other words, pumping blood is ''the reason'' that the heart has evolved. This account has been criticized for being too restrictive a notion of function. It is not always clear which behavior has contributed to the selection of a trait, as biological traits can have functions, even if they have not been selected for. Beneficial mutations are initially not selected for, but they do have functions.<ref>{{Cite journal |last=Neander |first=Karen |date=1991 |title=Functions as Selected Effects: The Conceptual Analyst's Defense |journal=Philosophy of Science |volume=58 |issue=2 |pages=168–184 |jstor=187457|doi=10.1086/289610 |s2cid=122866072 }}</ref>
=== Goal contribution ===
Goal contribution theories seek to carve a middle ground between causal role and selected effect theories, as with Boorse (1977).<ref name=Boorse1977>{{cite journal |last=Boorse |first=Christopher |date=1977 |title=Health as a Theoretical Concept |journal=Philosophy of Science |volume=44 |issue=4 |pages=542–573 |jstor=186939|doi=10.1086/288768 |citeseerx=10.1.1.486.2236 |s2cid=146325798 }}</ref> Boorse defines the function of a biological trait to be the statistically typical causal contribution of that trait to survival and reproduction. So for example, zebra stripes were sometimes said to work by confusing predators. This role of zebra stripes would contribute to the survival and reproduction of zebras, and that is why confusing predators would be said to be the function of zebra stripes. Under this account, whether or not a particular causal role of a trait is its function depends on whether that causal role contributes to the survival and reproduction of that organism.<ref>{{cite journal |last1=Bigelow |first1=John |last2=Pargetter |first2=Robert |date=1987 |title=Functions |journal=The Journal of Philosophy |volume=84 |issue=4 |pages=181–196 |doi=10.2307/2027157|jstor=2027157 }}</ref>
==See also== *Preadaptation
==References== {{reflist|30em}}
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{{DEFAULTSORT:Function (Biology)}} Category:Evolutionary biology terminology