Function (biology)

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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.

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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

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. [1] This concept of function as opposed to form (respectively Aristotle's ergon and morphê [2] ) 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. [3] [4] [5]

In evolutionary biology

Function can be defined in a variety of ways, [6] [7] including as adaptation, [8] as contributing to evolutionary fitness, [9] in animal behaviour, [10] and, as discussed below, also as some kind of causal role or goal in the philosophy of biology. [11]

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. [8]

Natural selection

Chlorophyll molecule has a function in photosynthesis. Chlorophyll-a-3D-vdW.png
Chlorophyll molecule has a function in photosynthesis.

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. [9] [12] For example, the function of chlorophyll in a plant is to capture the energy of sunlight for photosynthesis, [13] which contributes to evolutionary success. [14]

In ethology

The ethologist Niko Tinbergen named four questions, based on Aristotle's Four Causes, [10] 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. [15] [16] [17] [18]

In philosophy of biology

"Behaviour with a purpose": a young springbok stotting. 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. Springbok pronk.jpg
"Behaviour with a purpose": a young springbok stotting. 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.

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. [11]

In contemporary philosophy of biology, there are three major accounts of function in the biological world: theories of causal role, [20] selected effect, [21] and goal contribution. [22]

Causal role

Causal role theories of biological function trace their origin back to a 1975 paper by Robert Cummins. [20] 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. [23] [24]

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. [21] 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. [25]

Goal contribution

Goal contribution theories seek to carve a middle ground between causal role and selected effect theories, as with Boorse (1977). [22] 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. [26]

See also

Related Research Articles

<span class="mw-page-title-main">Natural selection</span> Mechanism of evolution by differential survival and reproduction of individuals

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 heritable traits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which is intentional, whereas natural selection is not.

Sociobiology is a field of biology that aims to examine and explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of human societies, sociobiology is closely allied to evolutionary anthropology, human behavioral ecology, evolutionary psychology, and sociology.

Zoology is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one of the primary branches of biology. The term is derived from Ancient Greek ζῷον, zōion ('animal'), and λόγος, logos.

<span class="mw-page-title-main">Teleology</span> Thinking in terms of destiny or purpose

Teleology or finality is a branch of causality giving the reason or an explanation for something as a function of its end, its purpose, or its goal, as opposed to as a function of its cause.

<span class="mw-page-title-main">Vestigiality</span> Evolution keeping organs no longer needed

Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on comparison with homologous features in related species. The emergence of vestigiality occurs by normal evolutionary processes, typically by loss of function of a feature that is no longer subject to positive selection pressures when it loses its value in a changing environment. The feature may be selected against more urgently when its function becomes definitively harmful, but if the lack of the feature provides no advantage, and its presence provides no disadvantage, the feature may not be phased out by natural selection and persist across species.

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 population during that process. Thirdly, it is a phenotypic trait or adaptive trait, with a functional role in each individual organism, that is maintained and has evolved through natural selection.

<span class="mw-page-title-main">Proximate and ultimate causation</span> Event which is closest to, or immediately responsible for causing, some observed result

A proximate cause is an event which is closest to, or immediately responsible for causing, some observed result. This exists in contrast to a higher-level ultimate cause which is usually thought of as the "real" reason something occurred.

Exaptation and the related term co-option describe a shift in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. Exaptations are common in both anatomy and behaviour.

Teleonomy is the quality of apparent purposefulness and of goal-directedness of structures and functions in living organisms brought about by natural processes like natural selection. The term derives from two Greek words, τέλος, from τελε-, and νόμος nomos ("law"). Teleonomy is sometimes contrasted with teleology, where the latter is understood as a purposeful goal-directedness brought about through human or divine intention. Teleonomy is thought to derive from evolutionary history, adaptation for reproductive success, and/or the operation of a program. Teleonomy is related to programmatic or computational aspects of purpose.

The philosophy of biology is a subfield of philosophy of science, which deals with epistemological, metaphysical, and ethical issues in the biological and biomedical sciences. Although philosophers of science and philosophers generally have long been interested in biology, philosophy of biology only emerged as an independent field of philosophy in the 1960s and 1970s, associated with the research of David Hull. Philosophers of science then began paying increasing attention to biology, from the rise of Neodarwinism in the 1930s and 1940s to the discovery of the structure of DNA in 1953 to more recent advances in genetic engineering. Other key ideas include the reduction of all life processes to biochemical reactions, and the incorporation of psychology into a broader neuroscience.

Adaptationism is the Darwinian view that many physical and psychological traits of organisms are evolved adaptations. Pan-adaptationism is the strong form of this, deriving from the early 20th century modern synthesis, that all traits are adaptations, a view now shared by only a few biologists.

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Cultural selection theory is the study of cultural change modelled on theories of evolutionary biology. Cultural selection theory has so far never been a separate discipline. However it has been proposed that human culture exhibits key Darwinian evolutionary properties, and "the structure of a science of cultural evolution should share fundamental features with the structure of the science of biological evolution". In addition to Darwin's work the term historically covers a diverse range of theories from both the sciences and the humanities including those of Lamark, politics and economics e.g. Bagehot, anthropology e.g. Edward B. Tylor, literature e.g. Ferdinand Brunetière, evolutionary ethics e.g. Leslie Stephen, sociology e.g. Albert Keller, anthropology e.g. Bronislaw Malinowski, Biosciences e.g. Alex Mesoudi, geography e.g. Richard Ormrod, sociobiology and biodiversity e.g. E.O. Wilson, computer programming e.g. Richard Brodie, and other fields e.g. Neoevolutionism, and Evolutionary archaeology.

<span class="mw-page-title-main">Four causes</span> Topic in Aristotelian philosophy

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<span class="mw-page-title-main">Structuralism (biology)</span> Attempt to explain evolution by forces other than natural selection

Biological or process structuralism is a school of biological thought that objects to an exclusively Darwinian or adaptationist explanation of natural selection such as is described in the 20th century's modern synthesis. It proposes instead that evolution is guided differently, basically by more or less physical forces which shape the development of an animal's body, and sometimes implies that these forces supersede selection altogether.

Biological constraints are factors which make populations resistant to evolutionary change. One proposed definition of constraint is "A property of a trait that, although possibly adaptive in the environment in which it originally evolved, acts to place limits on the production of new phenotypic variants." Constraint has played an important role in the development of such ideas as homology and body plans.

Tinbergen's four questions, named after 20th century biologist Nikolaas Tinbergen, are complementary categories of explanations for animal behaviour. These are also commonly referred to as levels of analysis. It suggests that an integrative understanding of behaviour must include ultimate (evolutionary) explanations, in particular:

<span class="mw-page-title-main">Evolutionary physiology</span> Study of changes in physiological characteristics

Evolutionary physiology is the study of the biological evolution of physiological structures and processes; that is, the manner in which the functional characteristics of individuals in a population of organisms have responded to natural selection across multiple generations during the history of the population. It is a sub-discipline of both physiology and evolutionary biology. Practitioners in the field come from a variety of backgrounds, including physiology, evolutionary biology, ecology, and genetics.

<span class="mw-page-title-main">Outline of evolution</span>

The following outline is provided as an overview of and topical guide to evolution:

<span class="mw-page-title-main">Teleology in biology</span> Use of language of goal-directedness in the context of evolutionary adaptation

Teleology in biology is the use of the language of goal-directedness in accounts of evolutionary adaptation, which some biologists and philosophers of science find problematic. The term teleonomy has also been proposed. Before Darwin, organisms were seen as existing because God had designed and created them; their features such as eyes were taken by natural theology to have been made to enable them to carry out their functions, such as seeing. Evolutionary biologists often use similar teleological formulations that invoke purpose, but these imply natural selection rather than actual goals, whether conscious or not. Biologists and religious thinkers held that evolution itself was somehow goal-directed (orthogenesis), and in vitalist versions, driven by a purposeful life force. With evolution working by natural selection acting on inherited variation, the use of teleology in biology has attracted criticism, and attempts have been made to teach students to avoid teleological language.

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