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Some Criticism of the Contextual Approach, and a Few Proposals

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Abstract

The contextual approach is a prominent framework for thinking about group selection. Here, I highlight ambiguity about what the contextual approach is. Then, I discuss problematic entailments the contextual approach has for what processes count as group selection—entailments more troublesome than typically noted. However, Sober and Wilson’s version of the Price approach, which is the main alternative to the contextual approach, is problematic too: it leads to an underappreciated paradox called the vanishing selection problem and thereby generates the wrong qualitative account of whether group selection is occurring in a certain family of cases. In response, I develop an account of group selection that can deal with the counterexamples to both the contextual approach and the Price approach. I then discuss the role that contextual analysis can continue to play in the discussion of individual fitness and metapopulation evolution.

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Notes

  1. As I explain in detail below, the Price approach is use of the Price equation and a particular endorsement of what processes count as “group selection.”

  2. This article is concerned with a debate among those who are sympathetic to multilevel selection-1 (MLS1). There is no dearth of criticism of MLS1, and while I address some of this below, largely I will assume that that framework’s account of “group fitness” is legitimate.

  3. A variant of this approach is called “neighborhood analysis,” wherein the “group” to which the organism belongs does not include the individual itself (Nunney 1985). Because of space constraints, the distinction between neighborhood analysis and contextual analysis will largely be ignored in this paper.

  4. I here assume discrete generations, perfect fidelity of transmission of phenotypes from parents to offspring, and that z ij and Z j are the only factors of individual fitness.

  5. Throughout, I will assume causal or statistical inference is not misleading in these examples, such that a nonzero estimate for β 1 or β 2 means there really is an individual- or group-level cause of individual fitness, respectively, as “cause” is articulated in, for instance, Pearl (2009). I say more about causation below.

  6. I assume that in this last quote Okasha meant to say the “contextual approach” instead of “contextual analysis,” for contextual analysis itself does not define group selection.

  7. Note, the fitness of the individuals in a group at time t can influence the composition of the group at time t + 1.

  8. Here and throughout, a group’s fitness is the mean fitness of the members of a group. This is what group fitness means in MLS1.

  9. For thorough discussion of soft selection, see Wallace (1975) and Van Dyken (2010).

  10. Perhaps the idea of MLS1 “group selection” is theoretically incoherent too, since its notion of “group fitness” diverges from standard accounts of “individual fitness.” I disagree, but this paper is not the place to discuss this issue.

  11. Perhaps Goodnight et al. (2008) note this too when they write: “However, contextual analysis works equally well with spatially structured or poorly mixed continuous populations when neighbors have sustained interactions influencing fitness” (p. 36, my emphasis).

  12. Earnshaw (2015) argues that soft selection can legitimately be thought of as group selection because at least there is selection among the members of the groups. I disagree, but even if we grant Earnshaw this claim, it is irrelevant to the present counterexample, because there are no fitness differences at any level.

  13. However, as I have noted, I believe the contextual approach’s judgment about soft selection is sufficient reason to reject the approach too.

  14. For instance, a contextualist can say that in a case of hard selection the members of each group “interact” with each other, even though they do not affect each other’s fitness. All that is mean by “interact” is that the investigator has some reason to lump together these organisms in the set whose members determine β 2 . That contextualists have this permissive account of “group” is evidenced by the fact that they talk of “groups” in hard selection (e.g., Heisler and Damuth 1987; Goodnight et al. 1992; Okasha 2006).

  15. In fact, this is the definition of “group” that Sober and Wilson use (Sober and Wilson 1998; Sober 2011).

  16. It is not clear to me that Sober and Wilson are as committed to the Price equation as Okasha suggests, but I will grant Okasha this point.

  17. This distinction between QUAL and QUAN comes from Sober (2011). His definitions of QUAL and QUAN are slightly broader, but not in a way that affects this discussion.

  18. Perhaps this judgment of hard selection is another fixed point. But I will not assume it is here, though my below definition of group selection does not classify hard selection as group selection.

  19. This fact is ignored in Earnshaw’s criticism of the Price approach (Earnshaw 2015).

  20. It follows trivially from this that it gets QUAN wrong sometimes too.

  21. I assume here that individuals reproduce with perfect fidelity, though one can construct the same scenario in which this is not the case. I have chosen not to for purposes of simplicity.

  22. John Basl and I independently discovered this issue, though he well before I.

  23. Note, this will not be a paradox if one is thinking of the situation in terms of MLS2, not MLS1.

  24. This is because my definition says, “for at least one of these groups, its composition affects the fitness of at least some of its members.”.

  25. Since z ij is a logical and mereological component of Z j , and such relationships can be misleading about causal structure (Hausman and Woodward 1999), it may be better to use the neighborhood analysis version of contextual analysis.

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Acknowledgments

This work was supported by a Junior Visiting Fellowship at The KLI Institute, in Klosterneuburg, Austria. I thank Ann-Sophie Barwich, John Basl, Hayley Clatterbuck, Steven Frank, Bruce Glymour, Charles Goodnight, Dan Hausman, Olivier Morin, Samir Okasha, Elliott Sober, Reuben Stern, Mike Titelbaum, and two anonymous reviewers for comments on earlier versions of this paper.

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McLoone, B. Some Criticism of the Contextual Approach, and a Few Proposals. Biol Theory 10, 116–124 (2015). https://doi.org/10.1007/s13752-015-0209-z

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