Abstract
Mayr’s proximate–ultimate distinction has received renewed interest in recent years. Here we discuss its role in arguments about the relevance of developmental to evolutionary biology. We show that two recent critiques of the proximate–ultimate distinction fail to explain why developmental processes in particular should be of interest to evolutionary biologists. We trace these failures to a common problem: both critiques take the proximate–ultimate distinction to neglect specific causal interactions in nature. We argue that this is implausible, and that the distinction should instead be understood in the context of explanatory abstractions in complete causal models of evolutionary change. Once the debate is reframed in this way, the proximate–ultimate distinction’s role in arguments against the theoretical significance of evo-devo is seen to rely on a generally implicit premise: that the variation produced by development is abundant, small and undirected. We show that a “lean version” of the proximate–ultimate distinction can be maintained even when this isotropy assumption does not hold. Finally, we connect these considerations to biological practice. We show that the investigation of developmental constraints in evolutionary transitions has long relied on a methodology which foregrounds the explanatory role of developmental processes. It is, however, entirely compatible with the lean version of the proximate–ultimate distinction.
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Notes
It has recently been suggested by several authors that ultimate questions should be restricted to issues of adaptive rationale or “what for?” (Haig 2013; Gardner 2013). To some extent, this is a dispute about preferred linguistic usage. However, in the spirit of Gould and Lewontin (1979) we prefer to include all evolutionary processes in the ultimate category. This includes selection, drift, developmental constraints, and more. Which of these processes are the most relevant should then be investigated as an empirical matter on a case by case basis.
Similar arguments could be made for other processes that have an effect on variational isotropy, such as variation in mutation rates (Hodgkinson and Eyre-Walker 2011).
Gould (1989) argued valiantly that “constraint” should be understood not only in the negative sense of limiting the power of selection, but also in the positive sense of channeling evolutionary transitions in particular adaptive directions. It seems that this usage has not been widely adopted; the more recent term “developmental drive” (Arthur 2001) appears to have been received more favorably for the positive case. Both constraints and drive, however, have the same conceptual foundation: If the variation available to selection is strongly structured by developmental mechanisms, then evolutionary transitions will need to be explained by asking both which variants were selectively advantageous and which variants were produced in the first place.
We thank Günter Wagner for suggesting this example.
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Acknowledgments
An early version of this paper was presented at the second meeting of the European Society for Evolutionary Developmental Biology in Ghent, Belgium, in 2008. We are particularly grateful to the following for comments on earlier drafts of the paper: Mark Jonas, Kärin Nickelsen, Tim Räz, and the members of the Lake Geneva Biology Interest Group (LG-BIG). We have also greatly benefitted from the comments of an anonymous referee for Biology & Philosophy.
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Scholl, R., Pigliucci, M. The proximate–ultimate distinction and evolutionary developmental biology: causal irrelevance versus explanatory abstraction. Biol Philos 30, 653–670 (2015). https://doi.org/10.1007/s10539-014-9427-1
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DOI: https://doi.org/10.1007/s10539-014-9427-1