Adaptationism

Adaptationism is a set of methods in the evolutionary sciences for distinguishing the products of adaptation from traits that arise through other processes. It is employed in fields such as ethology and evolutionary psychology that are concerned with identifying adaptations. George Williams' Adaptation and Natural Selection was highly influential in its development, defining some of the heuristics, such as complex functional design, used to identify adaptations.

Debate

Adaptationism is sometimes characterized by critics as an unsubstantiated assumption that all or most traits are optimal adaptations. Critics (most notably Richard Lewontin and Stephen Jay Gould) contend that the adaptationists (John Maynard Smith, W.D. Hamilton and Richard Dawkins being frequent examples) have over-emphasized the power of natural selection to shape individual traits to an evolutionary optimum, and ignored the role of developmental constraints, and other factors to explain extant morphological and behavioural traits.

Adaptationism could also be characterized as an approach to studying evolution of form and function that attempts to frame the existence and persistence of traits on the scenario that each of them arose independently due to how that trait improved the reproductive success of the organism's ancestors. Adaptationism is also a description of "folk biology" where non-experts see that, in general, organisms have an amazing array of adaptations, then apply this principle too broadly and describe everything as adaptive.

Criteria of calling something adaptive

If and only if a trait fulfills the following criteria will evolutionary biologists in general declare the trait an adaptation:

1. The trait is a variation of an earlier form.
2. The trait is heritable through the transmission of genes.
3. The trait enhances reproductive success.

Adaptationists too agree with these rules, but their opponents maintain adaptationists are sometimes too eager to take an imaginative leap where the evidence is spotty or ambiguous.

Evolution cannot form perfect organisms for numerous reasons. Foremost is that some elements of an organism's physiology are subject to constraints that environmental pressure cannot alter.

Anatomical Constraints

Anatomical constraints are features of organism's anatomy that are prevented from change by being constrained in some way. When organisms diverge from a common ancestor and inherit certain characteristics which become modified by natural selection of mutant phenotypes, it is as if some are traits are locked in place and are unable to change in certain ways. Some textbook anatomical constraints often include examples of structures that connect parts of the body together though a physical link.

These links are hard if not impossible to break because evolution usually requires that anatomy be formed by small consecutive modifications in populations through generations. Dr. Randolf Nesse, in his book: Why We Get sick explains that the "blind spot" in the vertebrate eye caused by the nerve fibers running in front of the retina and blocking vision in parts. He says that natural selection has come up with an elaborate work-around of the eyes wobbling back-and-forth to correct for this, but vertebrates have not found the solution embodied in cephalopod eyes, where the optic nerve does not obstruct the view. He goes on to say that a likely reason for this is that in order for the nerve to migrate over evolutionary time it would have to be disconnected or "cut" at some point to complete the trip. This disconnection would not allow for the message to go from the eye to the brain and this would render those mutant individuals blind, so making the transition virtually impossible. See also: Evolution of the eye.

Other examples often cited for the same reason are the cranial nerves in tetrapods. Some of the best derived models of early vertebrate evolution have been sharks, skates, and rays (collectively chondrichthyes). In chondrichthyes the cranial nerves run from the part of the brain that interprets sensory information, and radiate out towards the organs that produce those sensations. In tetrapods however, and mammals in particular, the nerves take an elaborate winding path through the cranium. They go up down and around structures that evolved after the common ancestor with sharks. This is because when their locations shifted over time each stage had to remain connected. See also: Cranial nerves.

It is not possible to have a number of generations of organisms that are blind (as in the vertebrate eye example) or generations with areas that are not "innervated", nor is it possible that in transition the "spermatochord" becomes severed, then continue to migrate to reach around the pelvis. These would not work because the intermediates with severed connections would be poorly suited for their environments and would fail to reproduce. So these links are maintained throughout millions of years simply because departure from the status quo is inviable.

Genetic Constraints

Genetic constraints provide another force that inhibits the effectiveness of natural selection. Examples include pleiotropy, epistasis, and polygenic traits.

With pleiotropy a hypothetical gene would be selected by natural selection or other related mechanisms. The trait would be selected for its effects on reproductive success. This is well established as how traits evolve. However, some genes control multiple traits. Selection that influences epistasis is a case where the regulation or expression of one gene, depends on one or several others. This is true for a good number of genes though to differing extents. The reason why this leads to muddied responses is that selection for a trait that is epistatically based can mean that an allele for a gene that is epistatic when selected would happen to affect others. This leads to the coregulation of others for a reason other than there is an adaptive quality to each of those traits. Like with pleiotropy, traits could reach fixation in a population as a by-product of selection for another.

In the context of development the difference between pleiotropy and epistasis is not so clear but at the genetic level the distinction is more clear. With these traits as being by-products of others it can ultimately be said that these traits evolved by not that they necessarily represent adaptations.

Polygenic traits are traits that are controlled by a number of different genes. Counter to what is commonly taught in high school and introductory biology classes with beginner's Mendelian Genetics, traits are rarely controlled by one single discrete, either/or allele. Things like human height vary with a great range because this trait is controlled by several different genes.

To drastically change some quantitative trait controlled by many genes, this could require mutations in more than one gene or changes in regulation of more than one gene. This means that mutations effect these systems more on a one-at-a-time basis. Because of this

Adaptationists are accused by their critics of using ad-hoc "just-so stories" to make their theories unfalsifiable. The critics, in turn, have often been accused of attacking straw men, rather than the actual views of supposed adaptationists.

Adaptationist rebuttals

Adaptationist researchers respond by asserting that they, too, follow George Williams' depiction of adaptation as an "onerous concept" that should only be applied in light of strong evidence. This evidence can be generally characterized as the successful prediction of novel phenomena based on the hypothesis that design details of adaptations should fit a complex evolved design to respond to a specific set of selection pressures. In evolutionary psychology, researchers such as David Buss contend that the bulk of research findings that were uniquely predicted through adaptationist hypothesizing comprise evidence of the methods' validity.

Adaptationists such as Steven Pinker have also suggested that the debate has a strong ad hominem component. Some suggest that the controversy over the relative importance of various factors would be a quiet debate over subtleties if the critics were less prone to caricaturing their opponents^{[citation needed]}.

See also

Evolutionary biology portal

• Adaptation
• Gene-centered view of evolution
• Spandrel
• Beneficial acclimation hypothesis

References

• Cronin, H. (1992). The Ant and the Peacock: Altruism and Sexual Selection from Darwin to Today. Cambridge: Cambridge University Press. ISBN 0-521-32937-X. 
• Gould, S.J.; Lewontin, R.C. (1979). "The spandrels of San Marco and the Panglossian paradigm: A critique of the adaptationist programme". Proceedings of the Royal Society of London B 205 (1161): 581–598. doi:10.1098/rspb.1979.0086. PMID 42062. 
• Lewontin, R.C. (1979). "Sociobiology as an adaptationist program". Behavioral Science 24 (1): 5–14. doi:10.1002/bs.3830240103. PMID 435219. 
• Lewontin, R.C. (1993). Biology as Ideology: The Doctrine of DNA. New York: Harper Collins. ISBN 0-06-097519-9. 
• Maynard Smith, J. (1988). Did Darwin get it right? Essays on games, sex and evolution. London: Penguin books. ISBN 0-14-023013-0. 
• Orzack, S.H.; & Sober, E.R., eds. (2001). Adaptationism and Optimality. Cambridge: Cambridge University Press. ISBN 0-521-59166-X. 
• Sober, E. (1998). "Six Sayings about Adaptationism". In D. Hull and M. Ruse (eds). The Philosophy of Biology. Oxford: Oxford University Press. ISBN 0-19-875213-x. 

External links

• Information from "Deep Ethology" course website, by Neil Greenberg
• Tooby & Cosmides comments on Maynard Smith's New York Review of Books piece on Gould et al.