An interesting modeling article by McNamara et al. suggests a novel evolutionary mechanism based on a positive coevolutionary feedback between cooperativeness and choosiness. If individuals vary in their degree of cooperativeness, and if they can decide whether or not to continue interacting with each other on the basis of their respective levels of cooperativeness, then cooperation can gradually evolve from an uncooperative state. When an individual's cooperativeness is used by other individuals as a choice criterion, there can be competition to be more generous than others (competetive altruism). The evolution of cooperation between non-relatives can then be driven by a positive feedback between increasing levels of cooperativeness and choosiness. In this model, individual behavioural differences are the key to the evolution of cooperation. Because the model does not invoke complex mechanisms such as negotiation behaviour, it can be applied to a wide range of species.
The model calculations use an infinite population where, in each of a discrete series of time steps (rounds), pairs of individuals engage in a game that can be described as a social dilemma. Each individual is characterized by two traits: a cooperativeness trait x, which specifies the amount of effort that the individual devotes to generating benefits available (at least in part) to its co-player; and a choosiness trait y, which specifies the minimum degree of cooperativeness that the focal individual is prepared to accept from its co-player. The traits x and y are genetically determined and are not adjusted in response to the co-player's behaviour. Thus, unlike in many models in which flexible effort adjustment is a key ingredient1, individuals in their model are consistent in their degree of cooperativeness.
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