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Adaptationism and the Logic of Research Questions: How to Think Clearly About Evolutionary Causes

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Abstract

This article discusses various dangers that accompany the supposedly benign methods in behavioral evolutionary biology and evolutionary psychology that fall under the framework of “methodological adaptationism.” A “Logic of Research Questions” is proposed that aids in clarifying the reasoning problems that arise due to the framework under critique. The live, and widely practiced, “evolutionary factors” framework is offered as the key comparison and alternative. The article goes beyond the traditional critique of Stephen Jay Gould and Richard C. Lewontin, to present problems such as the disappearance of evidence, the mishandling of the null hypothesis, and failures in scientific reasoning, exemplified by a case from human behavioral ecology. In conclusion the paper shows that “methodological adaptationism” does not deserve its benign reputation.

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Notes

  1. See Rose and Lauder (1996) for some examples of the application of this evolutionary factors approach. Or Martins (2000), for some methodological details.

  2. Symons gives a variety of reasons against using correlation with fitness for detecting adaptations, and prefers evidence of design (1990). Thornhill (1990) provides another perspective of behavioral adaptationists.

  3. See Lewens (2009) for an extensive categorization of types of adaptationism. Amundson (2001) and Sansom (2003) have also emphasized the multiple nature of adaptationist questions and answers, but not in the way I do here.

  4. Martin Kreitman introduced a technique using DNA sequence data that same year that can create the statistical tests to discriminate between selection and drift (1983). Thank you to Michael Dietrich for highlighting this sequence.

  5. Note that it is not always true that to identify a trait’s function is to identify its selection pressure: commonly, for example, we have multilevel selection, such as family and kin selection, and there are multiple processes responsible for the trait’s form and function. The trait itself does not tell us how to describe its selection pressure, although the investigating biologist may play favorites about which process to privilege in his or her explanations (for examples, see Wade 2016).

  6. Buss et al. (1998) argued that this category is really “adaptations” in their destructive analysis of exaptation, while Reeve and Sherman argued that the past selective history of a trait should not be included in the notion of adaptation, which is based instead on current utility, exactly backwards from the Gould and Vrba’s, and many others’ definitions: “ask why certain traits predominate over conceivable others in nature, irrespective of the precise historical pathways leading to their predominance, and then infer evolutionary causation based on current utility” (1993, p. 1; in contrast, see Burian 1992 and West-Eberhard 1992; Lloyd and Gould ([2002]2014)).

  7. Godfrey-Smith offers three general categories of “adaptationism,” including, besides “methodological adaptationism,” “empirical adaptationism” and “explanatory adaptationism.” We will not be dealing with these others, except to note that the evolutionary factors framework is independent of any commitment regarding empirical (or “metaphysical”) adaptationism. That is, it does not matter how many adaptations actually exist in the world, with regard to the relative superiority of the framework in researching those adaptations and related traits. See also Lewens (2009).

  8. I would like to thank the first reviewer from Biological Theory for discussion on this issue, and for posing these questions.

  9. Andrews et al. claim that, “The point of disagreement [concerning adaptationism] centers around the probative value of the evidentiary standards that adaptationists use to classify a trait as an adaptation” (2002a, p. 493).

  10. Note that Williams, here, is just as strict, or stricter, than Gould and Lewontin in his requirements for assigning the status of “adaptation” to a trait. There is an open question regarding how to read Williams on this topic (Lloyd 2013).

  11. In contrast, Seger and Stubblefield (1996) find that the bias towards functions is what limits the number of traits that we can treat as adaptations, e.g., in clutch size in birds or in various life history traits. Thanks to Steve Downes for pointing this out.

  12. I do not mean to deny the common point about the division of scientific labor, by saying that it is a good idea for some to start by asking about the function of a trait. It would be more useful for a phylogeneticist to start by asking whether a trait is ancestral or derived, and more useful for a developmental biologist to ask how the trait is developed in the organism. (Thanks to James Griesemer.) See Beatty (1987). The question is: is it useful for anyone to be a methodological adaptationist rather than following an evolutionary factors approach?

  13. But note that there is a problem with our running definitions of “adaptation” and "function": the first generation feature, arising from exaptations, byproducts, spandrels, or any source—e.g., a change that provides additional protection, enables association with a new food source, or otherwise brings a new niche into existence — does not yet have a “function” under our chosen definition, because it has not yet had a chance to be selected. Thus, in the first generation, we cannot tell whether it is an exaptation or an adaptation, just that it is an aptation (Gould and Vrba 1982). See discussion of Reeve and Sherman’s definition of “adaptation,” based on current utility rather than historical function (1993) in footnote 6 and “The Onerous Burden of Proof” section, above. An approach from developmental byproducts and novelties might clarify the arena of problems. Thank you to Stuart Newman for this example.

  14. Thanks to Archie Fields III and Carla Fehr for discussion of this issue.

  15. Thanks to Michael Wade for this example.

  16. Interestingly, Symons identifies himself as an “adaptationist,” but an analysis of his research shows that he is not a methodological adaptationist in the Mayrian sense used in this paper, but rather an “adaptationist” following the path of the evolutionary factors framework’s first questions (see Symons 1990).

  17. While apparently most often these tissues involve primarily the total clitoris and lower vaginal areas, Barry Komisaruk and his colleagues had noted that the cervix could serve as a center of orgasmic pleasure in some women under appropriate conditions of stimulation (2006; Kinsey et al. 1953). Komisaruk et al. have more recently shown that the human vagina, cervix, and clitoris are innervated by different afferent pathways, which project to different areas in the sensory cortex in the brain (2011). More research is necessary to understand these aspects of female orgasm more fully.

  18. Gould says: “Elisabeth Lloyd, a philosopher of science at the University of California at San Diego, has just completed a critical study of explanations recently proposed by evolutionary biologists for the origins and significance of female orgasm. Nearly all these proposals follow the lamentable tradition of speculative storytelling in the a priori adaptationist mode” (1987, p. 17).

  19. By “evolution,” Barash means “selection,” in context. This mistake is discussed in the next section.

  20. Thanks to Eduoard Machery for discussion. See Meehl (1954) for corrections to similar confusions in the comparison between statistical and causal hypotheses.

  21. This language comes from Thompson Clarke, “The Legacy of Skepticism,” (Clarke 1972).

  22. It may be that modern circumstances somehow block the historically relevant correlations from appearing in the data; however, the authors argue against such an interpretation (Zietsch and Santtila 2013).

  23. This low rate is apparently explained by an anatomical correlation between the structure of the genitals and the rate of orgasm with intercourse. Those women with a longer distance between clitoris and urinary meatus have  reliably many fewer orgasms with intercourse than those women with a shorter distance (Wallen and Lloyd 2011). Note again that the occurrence of orgasm is not correlated with fitness measures (Zietsch and Santtila 2011, 2013), so these different distances cannot be interpreted functionally, under the present information. Only Hrdy’s, of the current theories, presents female orgasm as anything but a present adaptation.

  24. See Mitchell (2002) for an excellent critique of Sherman’s similar “levels of analysis” approach to evolutionary explanations. Ian Jamieson rejected the “levels” approach (1989, p. 696).

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Acknowledgments

Thank you to Arnold and Maxine Tanis for their support of my research over many years. I owe thanks to many biologists and philosophers for discussion about the topic of this paper, including especially the following: The Biology Studies Reading Group at IU, Colin Allen, Linnda Caporael, Janet Collett, Michael Dietrich, Stephen Downes, Marcus Feldman, Stephen Jay Gould (with whom I discussed the germs of ideas of this article), Jim Griesemer, Chris Haufe, Ryan Ketcham, Roy Levin, Richard Lewontin, Daniel Lindquist, Alan Love, Eduoard Machery, Gordon McOuat, Roberta Millstein, Stuart Newman, Elizabeth and Rudy Raff, Elliott Sober, Donald Symons, Michael Wade, Michael Weisberg, and two anonymous referees for this journal. Please forgive me, those I have not listed due to my faulty memory!

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    Elisabeth A. Lloyd

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Lloyd, E.A. Adaptationism and the Logic of Research Questions: How to Think Clearly About Evolutionary Causes. Biol Theory 10, 343–362 (2015). https://doi.org/10.1007/s13752-015-0214-2

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