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Exaptation



 
 
Exaptation, cooption, and preadaptation are related terms referring to shifts in the function of a trait during evolution
Evolution

In biology, evolution is change in the heritability trait of a population of organisms from one generation to the next. These changes are caused by a combination of three main processes: variation, reproduction, and selection....
. 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 behavior. Bird feathers are a classic example: initially these evolved for temperature regulation, but later were adapted for flight.






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Exaptation, cooption, and preadaptation are related terms referring to shifts in the function of a trait during evolution
Evolution

In biology, evolution is change in the heritability trait of a population of organisms from one generation to the next. These changes are caused by a combination of three main processes: variation, reproduction, and selection....
. 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 behavior. Bird feathers are a classic example: initially these evolved for temperature regulation, but later were adapted for flight. Interest in exaptation relates to both the process and product of evolution: the process that creates complex traits and the product that may be imperfectly designed.

History and definitions

The idea that the function of a trait might shift during its evolutionary history originated with Charles Darwin
Charles Darwin

Charles Robert Darwin Royal Society was an English people natural history who realised and presented compelling evidence that all species of life have evolution over time from common descent, through the process he called natural selection....
 (1859, ch. 6). For many years the phenomenon was labeled “preadaptation.” Unfortunately, the term suggests forethought, which is contrary to a basic principle of natural selection.

The idea had been explored by several scholars when in 1982 Gould
Stephen Jay Gould

Stephen Jay Gould was a prominent American Paleontology, Evolution, and History of science. He was also one of the most influential and widely read writers of popular science of his generation....
 and Vrba introduced the term “exaptation”. Unfortunately for subsequent discussions, this definition had two categories with different implications for the role of adaptation.
(1) A character, previously shaped by natural selection for a particular function (an adaptation), is coopted for a new use—cooptation. (2) A character whose origin cannot be ascribed to the direct action of natural selection (a nonaptation), is coopted for a current use—cooptation. Gould and Vrba (1982, Table 1)
The definitions are silent as to whether exaptations had been shaped by natural selection after cooption, although Gould and Vrba cite examples (e.g., feathers) of traits shaped after cooption.

To avoid these ambiguities, Buss, et al. (1998) suggested the term “co-opted adaptation,” which is limited to traits that evolved after cooption. However, the commonly-used terms of ”exaptation” and “cooption” are ambiguous in this regard.

Examples

Of the many examples of exaptations, here are two involving familiar traits. A multi-stage example involves human hands, which evolved to facilitate tool use and which are an exaptation of primate hands that were used for grasping tree branches. Those primate hands, in turn, were an exaptation of front legs that were used for locomotion on the ground, and those legs were an exaptation of the fins of fish, which were used for locomotion in the water. As this lineage exploited different niches—water, land, trees, and tool-use on the ground—natural selection reshaped its limbs.

A behavioral example pertains to subdominant wolves licking the mouths of alpha wolves as a sign of submissiveness. (Similarly, dogs, which are domesticated wolves, lick the faces of their human owners.) This trait can be explained as an exaptation of wolf pups licking the faces of adults to encourage them to regurgitate food.

Implications


Evolution of complex traits

One of the challenges to Darwin’s theory of evolution was explaining how complex structures could evolve gradually, given that their incipient forms may have been inadequate to serve any function. As Mivart
George Jackson Mivart

File:St George Jackson Mivart.jpgSt. George Jackson Mivart PhD MD Fellow of the Royal Society was an England biologist. He is famous for starting as an ardent believer in natural selection who later became one of its fiercest critics....
 (a critic of Darwin) pointed out, 5 percent of a bird wing would not be functional. The incipient form of complex traits would not have survived long enough to evolve to a useful form.

As Darwin elaborated on in the last edition of The Origin of Species
The Origin of Species

Charles Darwin's On the Origin of Species is a seminal work in scientific literature and a landmark work in evolutionary biology. The book's full title is On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life....
, many complex traits evolved from earlier traits that had served different functions. By trapping air, primitive wings would have enabled birds to efficiently regulate their temperature, in part, by lifting up their feathers when too warm. Individual animals with more of this functionality would have left more offspring, resulting in the spread of this trait.

Eventually, feathers became sufficiently large that they enabled some individuals to glide. These individuals would leave more offspring, resulting in the spread of this trait because it served a second function, that of locomotion. Hence, the evolution of bird wings can be explained by a shifting in function from the regulation of temperature to flight.

Jury-rigged design

Darwin explained how the traits of living organisms are well-designed for their environment, but he also recognized that many traits are imperfectly designed. They appear to have been made from available material, that is, jury-rigged. Understanding exaptations may suggest hypotheses regarding subtleties in the adaptation. For instance, that feathers evolved initially for thermal regulation may help to explain some of their features unrelated to flight (Buss et al., 1998).

Some of the chemical pathways for physical pain and pain from social exclusion overlap (MacDonald and Leary, 2005). The physical pain system may have been co-opted to motivate social animals to respond to threats to their inclusion in the group. The physical-social pain overlap helps to explain the paradox that people who are threatened with social exclusion sometimes react with physical violence.

See also


  • Atavism
    Atavism

    The term atavism denotes the tendency to revert to ancestral type. An atavism is an evolutionary throwback, such as traits reappearing which had disappeared generations ago....
  • Vestigiality
  • Spandrel (biology)
    Spandrel (biology)

    Spandrel is a term used in evolution describing a phenotype characteristic that is considered to have developed during evolution as a side-effect of an adaptation, rather than arising from natural selection....


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