Abstract
Two controversies exist regarding the appropriate characterization of hierarchical and adaptive evolution in natural populations. In biology, there is the Wright–Fisher controversy over the relative roles of random genetic drift, natural selection, population structure, and interdemic selection in adaptive evolution begun by Sewall Wright and Ronald Aylmer Fisher. There is also the Units of Selection debate, spanning both the biological and the philosophical literature and including the impassioned group-selection debate. Why do these two discourses exist separately, and interact relatively little? We postulate that the reason for this schism can be found in the differing focus of each controversy, a deep difference itself determined by distinct general styles of scientific research guiding each discourse. That is, the Wright–Fisher debate focuses on adaptive process, and tends to be instructed by the mathematical modeling style, while the focus of the Units of Selection controversy is adaptive product, and is typically guided by the function style. The differences between the two discourses can be usefully tracked by examining their interpretations of two contested strategies for theorizing hierarchical selection: horizontal and vertical averaging.
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Notes
Wright acknowledged that three dimensional landscapes are themselves a heuristic device and that there were many dimensions along which selection might act. More recent work (e.g., Gavrilets 1999) proposes that, in very high dimensional landscapes, all peaks are connected by ridges, so that the need for traversing fitness valleys by drift essentially disappears. Whether nature traverses valleys or ridges, however, remains an open question.
There are additional genetic and environmental influences, not mentioned (e.g., Wolf et al. 2000).
For further discussion of Fisher’s realism, see, e.g., Frank and Slatkin (1992). In interpreting Fisher’s Fundamental Theorem of natural selection, they highlight “Fisher’s ecological, holistic view, and the very reasonable interpretation of clutch size that follows from this view.” (94).
Brandon’s characterization: “If A renders B statistically irrelevant with respect to outcome E but not vice versa, then A is a better causal explainer of E than is B” (1990, 83).
As opposed to dispositional fitness.
The functions literature is central to the philosophy of biology. A detailed investigation of this literature would be necessary for understanding the function style in general and for comprehending, for instance, Sober and Wilson's views on group functionality (e.g., Kitcher 1993; Allen et al. 1998; Ariew et al. 2002; Buller 1999). Standard analyses focus on the functions of individuals and their parts; interlocutors in this discourse have unfortunately not addressed group functionality in any detail. We believe that it would be fruitful to examine the concept of group function, especially as it relates to concepts of group adaptation and design.
This could be legitimately interpreted as a concern with adaptive process as well as adaptive product. However, we wish to stress that adaptive process requires fleshing out the evolutionary ecological details, something the predictive and explanatory theory of evolutionary genetics arguably does in a much less abstracted way than the Units of Selection debates.
At least one outstanding issue, inspired by Gould and Lewontin (1979), is the potential danger of what could be called group adaptationism.
There are some exceptions to this pattern. For instance, Simpson’s Paradox as used in the Units debate is mathematical, but the mathematics is used to make a conceptual point.
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Acknowledgments
Peter Godfrey-Smith, MJS Hodge, Elisabeth Lloyd, Lucas McGranahan, Fabrizzio McManus Guerrero, Amir Najmi, Elliott Sober, Kim Sterelny, and two anonymous reviewers, provided useful feedback on earlier drafts. RGW was supported in part from a Faculty Research Grant from the Academic Senate Committe on Research, University of California, Santa Cruz, and by the Center for Models of Life, Niels Bohr Insitute, and the Center for the Philosophy of Nature and Science Studies, Copenhagen University; MJW thanks support from the NIH grant 5R01GM65414–4; CCD appreciates support from an NSF Graduate Research Fellowship.
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Winther, R.G., Wade, M.J. & Dimond, C.C. Pluralism in evolutionary controversies: styles and averaging strategies in hierarchical selection theories. Biol Philos 28, 957–979 (2013). https://doi.org/10.1007/s10539-013-9378-y
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DOI: https://doi.org/10.1007/s10539-013-9378-y