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J Evol Biol. 2009 Apr;22(4):659-71. doi: 10.1111/j.1420-9101.2008.01681.x. Epub 2009 Jan 20.

Capturing the superorganism: a formal theory of group adaptation.

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Institute of Evolutionary Biology, University of Edinburgh, King's Buildings, Edinburgh, UK.


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.

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