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 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]
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]
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]
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]
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]
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 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 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 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]
Ecology is the natural science of the relationships among living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely related sciences of biogeography, evolutionary biology, genetics, ethology, and natural history.
Evolutionary psychology is a theoretical approach in psychology that examines cognition and behavior from a modern evolutionary perspective. It seeks to identify human psychological adaptations with regards to the ancestral problems they evolved to solve. In this framework, psychological traits and mechanisms are either functional products of natural and sexual selection or non-adaptive by-products of other adaptive traits.
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.
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.
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.
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.
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 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 or raison d'etre. A formal alternative would be to look at the products of evolution as the result of neutral evolution, in terms of structural constraints, or in terms of a mixture of factors including natural selection.
Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompasses all types of environmentally induced changes that may or may not be permanent throughout an individual's lifespan.
The four causes or four explanations are, in Aristotelian thought, four fundamental types of answer to the question "why?" in analysis of change or movement in nature: the material, the formal, the efficient, and the final. Aristotle wrote that "we do not have knowledge of a thing until we have grasped its why, that is to say, its cause." While there are cases in which classifying a "cause" is difficult, or in which "causes" might merge, Aristotle held that his four "causes" provided an analytical scheme of general applicability.
Human nature comprises the fundamental dispositions and characteristics—including ways of thinking, feeling, and acting—that humans are said to have naturally. The term is often used to denote the essence of humankind, or what it 'means' to be human. This usage has proven to be controversial in that there is dispute as to whether or not such an essence actually exists.
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:
An organism is defined in a medical dictionary as any 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 been proposed to define what is an organism. Among the commonest is that an organism has autonomous reproduction, growth, and metabolism. This would exclude viruses, despite that fact that they evolve like organisms. Other problematic cases include colonial organisms; a colony of eusocial insects is organised adaptively, and has germ-soma specialisation, with some insects reproducing, others not, like cells in an animal's body. The body of a siphonophore, a jelly-like marine animal, is composed of organism-like zooids, but the whole structure looks and functions much like an animal such as a jellyfish, the parts collaborating to provide the functions of the colonial organism.
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. Some 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.
Alternatives to Darwinian evolution have been proposed by scholars investigating biology to explain signs of evolution and the relatedness of different groups of living things. The alternatives in question do not deny that evolutionary changes over time are the origin of the diversity of life, nor that the organisms alive today share a common ancestor from the distant past ; rather, they propose alternative mechanisms of evolutionary change over time, arguing against mutations acted on by natural selection as the most important driver of evolutionary change.
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