Abstract: Cooperation among unrelated individuals can evolve not only via reciprocal altruism but also via trait-group selection or by-product mutualism (or some combination of all three categories). Therefore the (iterated) prisoner's dilemma is an insufficient paradigm for studying the evolution of cooperation. We replace this game by the cooperator's dilemma, which is more versatile because it enables all three categories of cooperative behavior to be examined within the framework of a single theory. Controlled studies of cooperation among fish provide examples of each category of cooperation. Specifically, we describe reciprocal altruism among simultaneous hermaphrodites that swap egg parcels, group-selected cooperation among fish that inspect dangerous predators and by-product mutualism in the cooperative foraging of coral-reef fish.

Abstract: Sixteen domestic dogs (Canis familiaris) were tested in a familiar context in a series of 1-min trials on how well they obeyed after being told by their owner to lie down. Food was used in 1/3 of all trials, and during the trial the owner engaged in 1 of 5 activities. The dogs behaved differently depending on the owner's attention to them. When being watched by the owner, the dogs stayed lying down most often and/or for the longest time compared with when the owner read a book, watched TV, turned his or her back on them, or left the room. These results indicate that the dogs sensed the attentional state of their owners by judging observable behavioral cues such as eye contact and eye, head, and body orientation.

Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.