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Capturing the superorganism: a formal theory of group adaptation
Article first published online: 29 JAN 2009
DOI: 10.1111/j.1420-9101.2008.01681.x
© 2009 The Authors. Journal Compilation © 2009 European Society For Evolutionary Biology
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How to Cite
GARDNER, A. and GRAFEN, A. (2009), Capturing the superorganism: a formal theory of group adaptation. Journal of Evolutionary Biology, 22: 659–671. doi: 10.1111/j.1420-9101.2008.01681.x
Publication History
- Issue published online: 13 MAR 2009
- Article first published online: 29 JAN 2009
- Received 23 October 2008; revised 21 November 2008; accepted 25 November 2008
- Abstract
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Keywords:
- formal Darwinism;
- group as maximizing agent;
- group selection;
- inclusive fitness;
- individuality;
- kin selection;
- levels of selection;
- major transitions;
- natural selection;
- Price equation
Abstract
Adaptation is conventionally regarded as occurring at the level of the individual organism. However, in recent years there has been a revival of interest in the possibility for group adaptations and superorganisms. Here, we provide the first formal theory of group adaptation. In particular: (1) we clarify the distinction between group selection and group adaptation, framing the former in terms of gene frequency change and the latter in terms of optimization; (2) we capture the superorganism in the form of a ‘group as maximizing agent’ analogy that links an optimization program to a model of a group-structured population; (3) we demonstrate that between-group selection can lead to group adaptation, but only in rather special circumstances; (4) we provide formal support for the view that between-group selection is the best definition for ‘group selection’; and (5) we reveal that mechanisms of conflict resolution such as policing cannot be regarded as group adaptations.