Whence tit-for-tat?

In theoretical and empirical studies of the evolution of cooperation, the tit-for-tat strategy (i.e. cooperate unless your partner did not cooperate in the previous interaction) is widely considered to be of central importance. Nevertheless, surprisingly little is known about the conditions in which tit-for-tat appears and disappears across generations in a population of interacting individuals. Here, we apply a newly developed classifier-system model (EvA) in addressing this issue when the key features of interactions are caricatured using the iterated prisoner's dilemma game. Our simple representation of behavioural strategies as algorithms composed of two interacting rules allowed us to determine conditions in which tit- for-tat can replace non-cooperative strategies and vice versa. Using direct game-theoretic analysis and simulations with the EvA model, we determined that no strategy is evolutionarily stable, but larger population sizes and longer sequences of interactions between individuals can yield transient dominance by tit-for-tat. Genetic drift among behaviourally equivalent strategies is the key mechanism underlying this dominance. Our analysis suggests that tit-for-tat could be important in nature for cognitively simple organisms of limited memory capacity, in strongly kin-selected or group-selected populations, when interaction sequences between individuals are relatively short, in moderate-sized populations of widely interacting individuals and when defectors appear in the population with moderate frequency.