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Science 8 December 2006:
Vol. 314. no. 5805, pp. 1560 - 1563
DOI: 10.1126/science.1133755
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Review

Five Rules for the Evolution of Cooperation

Martin A. Nowak

Cooperation is needed for evolution to construct new levels of organization. Genomes, cells, multicellular organisms, social insects, and human society are all based on cooperation. Cooperation means that selfish replicators forgo some of their reproductive potential to help one another. But natural selection implies competition and therefore opposes cooperation unless a specific mechanism is at work. Here I discuss five mechanisms for the evolution of cooperation: kin selection, direct reciprocity, indirect reciprocity, network reciprocity, and group selection. For each mechanism, a simple rule is derived that specifies whether natural selection can lead to cooperation.

Program for Evolutionary Dynamics, Department of Organismic and Evolutionary Biology, and Department of Mathematics, Harvard University, Cambridge, MA 02138, USA.

E-mail: martin_nowak{at}harvard.edu

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Science. ISSN 0036-8075 (print), 1095-9203 (online)