In theoretical and empirical studies of the evolution of cooperation,
the tit-for-tat strategy (i.e. cooperate unless your partner did not c
ooperate in the previous interaction) is widely considered to be of ce
ntral importance. Nevertheless, surprisingly little is known about the
conditions in which tit-for-tat appears and disappears across generat
ions in a population of interacting individuals. Here, we apply a newl
y developed classifier-system model (EvA) in addressing this issue whe
n the key features of interactions are caricatured using the iterated
prisoner's dilemma game. Our simple representation of behavioural stra
tegies as algorithms composed of two interacting rules allowed us to d
etermine conditions in which tit-for-tat can replace noncooperative st
rategies and vice versa. Using direct game-theoretic analysis and simu
lations with the EvA model, we determined that no strategy is evolutio
narily stable, but larger population sizes and longer sequences of int
eractions between individuals can yield transient dominance by tit-for
-tat. Genetic drift among behaviourally equivalent strategies is the k
ey 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-selecte
d populations, when interaction sequences between individuals are rela
tively short, in moderate-sized populations of widely interacting indi
viduals and when defectors appear in the population with moderate freq
uency.