This paper analyzes recent attempts to reject reproduction with lineage formation as a necessary condition for evolution by means of natural selection (Bouchard in Philos Sci 75(5):560–570, 2008; Stud Hist Philos Sci Part C Stud Hist Philos Biol Biomed Sci 42(1):106–114, 2011; Bourrat in Biol Philos 29(4):517–538, 2014; Br J Philos Sci 66(4):883–903, 2015; Charbonneau in Philos Sci 81(5):727–740, 2014; Doolittle and Inkpen in Proc Natl Acad Sci 115(16):4006–4014, 2018). Building on the strengths of these attempts and avoiding their pitfalls, it is argued that a robust formulation of evolution by natural selection without reproduction can be established. The main contribution of this paper is a reformulation of Lewontin’s three principles (Lewontin in Annu Rev Ecol Syst 1:1–18, 1970) stating that minimal evolution by natural selection occurs when two conditions are met in a population: fitness-related variation and memory (population-level inheritance). Paradigmatic evolution by natural selection, which can generate adaptations, takes place when an additional condition is present, namely regeneration.
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I provide only a superficial analysis of Godfrey-Smith’s very rich and insightful treatment of the concept of reproduction. This overview should suffice to fulfill the aim of this section, that is, to highlight the consensus according to which reproduction is necessary for ENS to obtain and to identify an abstract definition of reproduction that can apply despite the variety of ways in which it has been defined.
While the expression “origin explanation” has indeed been coined by Godfrey-Smith, it should be noted that Neander (1995a, b) similarly highlighted the creative power of natural selection, in cases of cumulative (rather than single-step) selection. Her claim sparked a debate with Sober (1995), who famously emphasized the negative power of selection (Sober 1984).
As it is shown shortly, I reject Bouchard’s claim that the notion of population needs to be replaced by the concept of ensemble. I keep using the concept of population even when discussing his work in order to simplify the analysis.
In a recent paper (Bourrat 2019), Bourrat further theorizes heritability by developing a causation (interventionist) account of it. While the paper is extremely interesting and has great practical value, the issue tackled therein is orthogonal to the more abstract and conceptual debate broached in this paper.
Charbonneau explicitly acknowledges that the resulting analysis is not a comprehensive in regard to the complexity of reproduction and the role it plays in the biological world. However, he claims that focusing on those two aspects allows for sufficient precision relative to its role as a necessary condition for ENS. This nuance is completely endorsed and reiterated in the context of this paper.
Given the aim of the paper, one might think I am suggesting that the evolutionary success of an entity should only be calculated in regard to its persisting capacities. It is not the case. Calculating fitness in regard to reproduction and survivability is an extremely powerful way to make evolutionary predictions. This is true even if reproduction is but one evolutionary strategy among others that generate success. In the thought experiment now being discussed, reproduction could be taken into account in the evolutionary success as calculated by the machine before a new event of creation. Hence, there will sometimes be causal input by parent on the offspring, and sometimes not. This is coherent with the claim that reproduction is neither a defining trait of units of selection nor necessary for ENS to occur.
As this paper is focused on the notion of reproduction and hence on those of memory and regeneration, it leaves the notion of fitness somewhat unspecified. Nonetheless, building on the work of Bouchard and Bourrat, the abstract definition provided here fulfills the needs of a general account of ENS. Further specifications of parameters can be added to fulfill the needs of specific empirical inquiries, when required (e.g. Roughgarden et al. 2018).
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I wish to thank Mathieu Charbonneau, Pierrick Bourrat, Frédéric Bouchard, Éric Bapteste and Cassandre Ville for their unwavering support and for their help with the paper. I am also grateful to a few anonymous referees for their great insights and comments.
The Social Sciences and Humanities Research Council of Canada provided financial support for this research (file number: 767-2018-1113).
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Papale, F. Evolution by means of natural selection without reproduction: revamping Lewontin’s account. Synthese (2020). https://doi.org/10.1007/s11229-020-02729-6
- Natural selection
- Unit of selection