Theory of Cooperation

Abstract

Cooperation is defined as any adaptation that has evolved, at least in part, to increase the reproductive success of the actor's social partners. Inclusive fitness theory reveals that cooperation can be favoured by natural selection owing to either direct fitness benefits (mutually beneficial cooperation) or indirect fitness benefits (altruistic cooperation). Direct fitness benefits can arise as a simple by‐product of cooperation, or else owing to the existence of enforcement mechanisms, which may be fixed or conditioned according to the individual's cooperative behaviour. Indirect fitness benefits can arise when cooperation occurs between genetically similar individuals, as a consequence of limited dispersal, kin discrimination or greenbeard mechanisms. These theoretical mechanisms are illustrated with empirical examples, from laboratory experiments to field studies.

Key Concepts

  • The function of Darwinian adaptation is to maximise the organism's inclusive fitness.
  • Inclusive fitness describes how well an organism transmits copies of its genes to future generations.
  • Direct fitness is the part of inclusive fitness that comes from the organism's own reproductive success.
  • Indirect fitness is the part of inclusive fitness that comes from the reproductive success of the organism's genetic relatives.
  • Cooperation is any adaptation whose function is, at least in part, to increase the reproductive success of a social partner.
  • Cooperation is mutually beneficial if the actor also benefits and altruistic if the actor suffers a net loss of reproductive success.
  • Mutually beneficial cooperation is favoured by direct fitness benefits.
  • Direct fitness benefits can arise as a by‐product or owing to enforcement mechanisms.
  • Altruistic cooperation is favoured by indirect fitness benefits.
  • Indirect fitness benefits can arise as a consequence of limited dispersal, kin discrimination or greenbeard mechanisms.

Keywords: altruism; Hamilton's rule; inclusive fitness; kin selection; social evolution

Figure 1. Inclusive fitness is the sum of direct fitness and indirect fitness. Social behaviours affect the reproductive success of individuals beyond the actor. The impact on the actor's own reproductive success is the direct fitness effect. The impact on the reproductive success of social partners, weighted by the relatedness of the actor to the recipient, is the indirect fitness effect. In particular, inclusive fitness does not include all the reproductive success of relatives, only that which is due to the behaviour of the actor (yellow hands). In addition, inclusive fitness does not include any of the reproductive success of the actor that is due to the actions of its social partners (blue hands).Reproduced from West et al.2007a © Elsevier.
Figure 2. Mechanisms generating direct fitness benefits for cooperation. (a) By‐product benefit. Helping a social partner may lead to increases in the overall fitness of all individuals in a social group; for example, in situations where larger social groups offer better protection against predators, the actor could benefit from helping its neighbours to reproduce (group augmentation). (b) Conditional enforcement (e.g. reciprocity). Helping may lead to a change in the behaviour of the recipient or a third party, in a way that leads to an overall direct fitness benefit for the actor; for example, direct reciprocity ensures that cooperators receive more cooperation than cheats, giving a direct benefit to cooperation even if each act involves an immediate cost. (c) Unconditional enforcement (e.g. policing). Helping may be the only option available to an individual, if the possibility for it to behave selfishly is ruled out by a successful system of policing.
Figure 3. Cooperation and punishment in humans. Humans show higher levels of cooperation in economic games, when there are opportunities to punish individuals who do not cooperate (Fehr and Gächter, ). Thanks to E. Fehr for data and O. Henderson for illustration. Reproduced from West et al.2007a © Elsevier.
Figure 4. Mechanisms generating indirect fitness benefits for cooperation. (a) Limited dispersal. If individuals do not move far during their lifetime, then they will tend to be surrounded by kin (shaded) and hence even indiscriminate altruism could be directed primarily towards kin rather than nonkin. (b) Kin discrimination. For example, if the actor can remember those individuals it shared a nest with when young and discriminate these kin (shaded) from nonkin (unshaded) after leaving the nest, then cooperation can be directed primarily towards genetic relatives. (c) Greenbeard. If the gene‐controlling altruism is also associated with a phenotypic marker, such as a green beard, then green‐bearded individuals can identify which of their neighbours carries a copy of the gene. Altruism directed at genetic relatives can be favoured by natural selection, even if these are not genealogical relatives (kin).
Figure 5. Kin discrimination in long‐tailed tits. Ninety‐four percent of helpers prefer to help at nests containing related chicks when they have the choice of where to invest their efforts (Russell and Hatchwell, ). Thanks to O. Henderson for illustration.Reproduced from West et al.2007a © Elsevier.
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Further Reading

Bourke AFG (2011) Principles of Social Evolution. Oxford: Oxford University Press.

Frank SA (1998) Foundations of Social Evolution Theory. Princeton, NJ: Princeton University Press.

Hamilton WD (1996) Narrow Roads of Geneland. Vol 1: Evolution of Social Behaviour. Oxford: Freeman.

Maynard Smith J and Szathmáry E (1995) The Major Transitions in Evolution. Oxford: Oxford University Press.

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Gardner, Andy, Griffin, Ashleigh S, and West, Stuart A(Nov 2016) Theory of Cooperation. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0021910.pub2]