, and preadaptation
are related terms referring to shifts 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 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
(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 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.
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.
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
(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, 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.
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.
- Buss, David M., Martie G. Haselton, Todd K. Shackelford, et al. (1998) “Adaptations, Exaptations, and Spandrels,” American Psychologist, 53 (May):533–548. http://www.sscnet.ucla.edu/comm/haselton/webdocs/spandrels.html
- Darwin, Charles (1859) The Origin of Species, London, ch. 6, section “Modes of Transition.” http://www.infidels.org/library/historical/charles_darwin/origin_of_species/Chapter6.html.
- Ehrlich, Paul, and Marcus Feldman (2003) “Genes and Culture: What Creates Our Behavioral Phenome?,” Current Anthropology, 44 (February):87–107. Included are comments and a reply.
- Gould, Stephen Jay, and Elizabeth S. Vrba (1982), "Exaptation — a missing term in the science of form," Paleobiology 8 (1): 4–15.
- Gould, Stephen Jay (1991) “Exaptation: A Crucial tool for Evolutionary Psychology,” Journal of Social Issues 47: 43–65.
- Jacob, Francois (1977) “Evolution and Tinkering,” Science 196 (June 10): 1161–1166.
- MacDonald, Geoff, and Mark R. Leary (2005) “Why Does Social Exclusion Hurt? The Relationship between Social and Physical Pain,” Psychological Bulletin 131 (2): 202–223.
- Mayr, Ernst (1982) The Growth of Biological Thought: Diversity, Evolution, and Inheritance, Harvard University Press, ISBN 0-674-36445-7.
- http://wiki.cotch.net/index.php/Citations_of_cooption, which points to additional webpages.