Abstract
For evolution by natural selection to occur it is classically admitted that the three ingredients of variation, difference in fitness and heredity are necessary and sufficient. In this paper, I show using simple individual-based models, that evolution by natural selection can occur in populations of entities in which neither heredity nor reproduction are present. Furthermore, I demonstrate by complexifying these models that both reproduction and heredity are predictable Darwinian products (i.e. complex adaptations) of populations initially lacking these two properties but in which new variation is introduced via mutations. Later on, I show that replicators are not necessary for evolution by natural selection, but rather the ultimate product of such processes of adaptation. Finally, I assess the value of these models in three relevant domains for Darwinian evolution.
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Notes
I borrow this term and idea from Okasha (2006) and his discussion on the levels of selection in which he explains that multicellular organisms once taken for granted by individual level selection theory have been “endogeneized” by the multilevel selection theory, which makes of multicellularity a product of natural selection.
A synonym for ahistorical adaptation is “adaptiveness”.
I distinguish procreation from “progeneration” used by Griesemer (2000) which is in my terminology, procreation with material overlap.
The software NETLOGO 5.02 has been used. Two advantages of individual-based models (a microscopic approach to modeling) over macroscopic approaches are their flexibility (which makes them appealing for testing new hypotheses) and that they allow for each individual in the simulation to be unique. These two features make individual-based modeling ideal for the purpose of this article.
In Model 2 we assume that the mutation has already occurred. Mutation is thus exogenous to the model.
In Model 2 and subsequent, procreation is always asexual.
Another way to make the distinction is by using the well-known distinction between determinable and determinate (Sanford 2011). Heredity of is a determinable property while heredity on is a determinate property.
A similar mutation could appear and affect fertility.
This definition of replicators leads to a form of replication similar to the one argued for by Nanay (2011) who claims that replication is about properties rather than entities.
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Acknowledgments
I am thankful to Samuel Baron, Michael Duncan, Paul Griffiths, Johann Hariman, Adam Hochman, Robyn Kath, Arnon Levy, Kristie Miller, Mark Olson, Maureen O’Malley, Susanna Saracco, Kim Sterelny, Simon Varey, Elena Walsh, Michael Weisberg and two anonymous referees for their comments on an earlier version of this paper. I am especially grateful to Michael Duncan who proofread the English of the paper. I would also like to thank Peter Godfrey-Smith for discussions on this subject. This research was supported under Australian Research Council’s Discovery Projects funding scheme DP0878650 and an International Postgraduate Research Scholarship from the University of Sydney.
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Bourrat, P. From survivors to replicators: evolution by natural selection revisited. Biol Philos 29, 517–538 (2014). https://doi.org/10.1007/s10539-013-9383-1
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DOI: https://doi.org/10.1007/s10539-013-9383-1