Evolution of cooperation through indirect reciprocity.

How can cooperation through indirect reciprocity evolve and what would it be like? This problem has previously been studied by simulating evolution in a small group of interacting individuals, assuming no gene flow between groups. In these simulations, certain 'image scoring' strategies were found to be the most successful. However, analytical arguments show that it would not be in an individual's interest to use these strategies. Starting with this puzzle, we investigate indirect reciprocity in simulations based on an island model. This has an advantage in that the role of genetic drift can be examined. Our results show that the image scoring strategies depend on very strong drift or a very small cost of giving help. As soon as these factors are absent, selection eliminates image scoring. We also consider other possibilities for the evolution of indirect reciprocity. In particular, we find that the strategy of aiming for 'good standing' has superior properties. It can be an evolutionarily stable strategy and, even if not, it usually beats image scoring. Furthermore, by introducing quality variation among individuals into the model, we show that the standing strategy can be quality revealing, adding a new dimension to indirect reciprocity. Finally, we discuss general problems with currently popular modelling styles.