Biological Effect
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In
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary biolo ...
, function is the reason some object or process occurred in a system that
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
through
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
. That reason is typically that it achieves some result, such as that
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
helps to capture the energy of sunlight in
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
. 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 In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
; other characteristics may be non-functional
spandrels A spandrel is a roughly triangular space, usually found in pairs, between the top of an arch and a rectangular frame, between the tops of two adjacent arches, or one of the four spaces between a circle within a square. They are frequently fil ...
, though these in turn may later be co-opted by evolution to serve new functions. In
biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
, function has been defined in many ways. In
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
, it is simply what an organ, tissue, cell or molecule does. In the
philosophy of biology The philosophy of biology is a subfield of philosophy of science, which deals with epistemology, epistemological, metaphysics, metaphysical, and ethics, ethical issues in the biological and biomedical sciences. Although philosophers of science and ...
, talk of function inevitably suggests some kind of
teleological Teleology (from , and )Partridge, Eric. 1977''Origins: A Short Etymological Dictionary of Modern English'' London: Routledge, p. 4187. or finalityDubray, Charles. 2020 912Teleology. In ''The Catholic Encyclopedia'' 14. New York: Robert Applet ...
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

In
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
, a function is an activity or process carried out by a system in an
organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...
, such as
sensation Sensation (psychology) refers to the processing of the senses by the sensory system. Sensation or sensations may also refer to: In arts and entertainment In literature *Sensation (fiction), a fiction writing mode *Sensation novel, a British ...
or locomotion in an animal. This concept of function as opposed to
form Form is the shape, visual appearance, or configuration of an object. In a wider sense, the form is the way something happens. Form may also refer to: *Form (document), a document (printed or electronic) with spaces in which to write or enter dat ...
(respectively Aristotle's ''ergon'' and ''morphê'') was central in biological explanations in
classical antiquity Classical antiquity, also known as the classical era, classical period, classical age, or simply antiquity, is the period of cultural History of Europe, European history between the 8th century BC and the 5th century AD comprising the inter ...
. 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.


In evolutionary biology

Function can be defined in a variety of ways, including as adaptation, as contributing to evolutionary fitness, in animal behaviour, and, as discussed below, also as some kind of causal role or goal in the philosophy of biology.


Adaptation

A functional characteristic is known in
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary biolo ...
as an
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
, and the research strategy for investigating whether a character is adaptive is known as
adaptationism Adaptationism is a scientific perspective on evolution that focuses on accounting for the products of evolution as collections of adaptive traits, each a product of natural selection with some adaptive rationale. A formal alternative would be to ...
. Although assuming that a character is functional may be helpful in research, some characteristics of organisms are non-functional, formed as accidental
spandrels A spandrel is a roughly triangular space, usually found in pairs, between the top of an arch and a rectangular frame, between the tops of two adjacent arches, or one of the four spaces between a circle within a square. They are frequently fil ...
, side effects of neighbouring functional systems.


Natural selection

From the point of view of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
, biological functions exist to contribute to fitness, increasing the chance that an organism will survive to
reproduce Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: asexual and sexual. In asexual reprod ...
. For example, the function of
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
in a plant is to capture the energy of sunlight for
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
, which contributes to evolutionary success.


In ethology

The ethologist
Niko Tinbergen Nikolaas "Niko" Tinbergen ( , ; 15 April 1907 – 21 December 1988) was a Dutch biologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine with Karl von Frisch and Konrad Lorenz for their discoveries concerning th ...
named four questions, based on
Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
's
Four Causes The four causes or four explanations are, in Aristotelianism, Aristotelian thought, categories of questions that explain "the why's" of something that exists or changes in nature. The four causes are the: #Material, material cause, the #Formal, f ...
, that a biologist could ask to help explain a
behaviour Behavior (American English) or behaviour (British English) is the range of actions of Individual, individuals, organisms, systems or Artificial intelligence, artificial entities in some environment. These systems can include other systems or or ...
, though they have been generalised to a wider scope. 1) Mechanism: What mechanisms
cause Causality is an influence by which one event, process, state, or object (''a'' ''cause'') contributes to the production of another event, process, state, or object (an ''effect'') where the cause is at least partly responsible for the effect, ...
the animal to behave as it does? 2)
Ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the ovum, egg to adult. The term can also be used to refer to t ...
: 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 A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
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.


In philosophy of biology

Function is not the same as purpose in the
teleological Teleology (from , and )Partridge, Eric. 1977''Origins: A Short Etymological Dictionary of Modern English'' London: Routledge, p. 4187. or finalityDubray, Charles. 2020 912Teleology. In ''The Catholic Encyclopedia'' 14. New York: Robert Applet ...
sense, that is, possessing conscious mental intention to achieve a goal. In the
philosophy of biology The philosophy of biology is a subfield of philosophy of science, which deals with epistemology, epistemological, metaphysics, metaphysical, and ethics, ethical issues in the biological and biomedical sciences. Although philosophers of science and ...
,
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
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
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s to attract
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
s' 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. 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.


Causal role

Causal role theories of biological function trace their origin back to a 1975 paper by Robert Cummins. 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.


Selected effect

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. 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
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
s are initially not selected for, but they do have functions.


Goal contribution

Goal contribution theories seek to carve a middle ground between causal role and selected effect theories, as with Boorse (1977). 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.


See also

* Preadaptation


References

{{DEFAULTSORT:Function (Biology) Evolutionary biology terminology