Ultimate and proximate mechanisms of reciprocal altruism in rats.

The reciprocal exchange of goods and services among social partners is a conundrum in evolutionary biology because of its proneness to cheating, but also the behavioral and cognitive mechanisms involved in such mutual cooperation are hotly debated. Extreme viewpoints range from the assumption that, at the proximate level, observed cases of "direct reciprocity" can be merely explained by basic instrumental and Pavlovian association processes, to the other extreme implying that "cultural factors" must be involved, as is often attributed to reciprocal cooperation among humans. Here we argue that neither one nor the other extreme conception is likely to explain proximate mechanisms underlying reciprocal altruism in animals. In particular, we outline that Pavlovian association processes are not sufficient to explain the documented reciprocal cooperation among Norway rats, as has been recently argued.

Generalized reciprocity in rats.

The evolution of cooperation among nonrelatives has been explained by direct, indirect, and strong reciprocity. Animals should base the decision to help others on expected future help, which they may judge from past behavior of their partner. Although many examples of cooperative behavior exist in nature where reciprocity may be involved, experimental evidence for strategies predicted by direct reciprocity models remains controversial; and indirect and strong reciprocity have been found only in humans so far. Here we show experimentally that cooperative behavior of female rats is influenced by prior receipt of help, irrespective of the identity of the partner. Rats that were trained in an instrumental cooperative task (pulling a stick in order to produce food for a partner) pulled more often for an unknown partner after they were helped than if they had not received help before. This alternative mechanism, called generalized reciprocity, requires no specific knowledge about the partner and may promote the evolution of cooperation among unfamiliar nonrelatives.

What sets the odds of winning and losing?

Social experience influences the outcome of conflicts such that winners are more likely to win again and losers will more likely lose again, even against different opponents. Although winner and loser effects prevail throughout the animal kingdom and crucially influence social structures, the ultimate and proximate causes for their existence remain unknown. We propose here that two hypotheses are particularly important among the potential adaptive explanations: the 'social-cue hypothesis', which assumes that victory and defeat leave traces that affect the decisions of subsequent opponents; and the 'self-assessment hypothesis', which assumes that winners and losers gain information about their own relative fighting ability in the population. We discuss potential methodologies for experimental tests of the adaptive nature of winner and loser effects.

Evolution of cooperation by generalized reciprocity.

The evolution of cooperation by direct reciprocity requires that individuals recognize their present partner and remember the outcome of their last encounter with that specific partner. Direct reciprocity thus requires advanced cognitive abilities. Here, we demonstrate that if individuals repeatedly interact within small groups with different partners in a two person Prisoner's Dilemma, cooperation can emerge and also be maintained in the absence of such cognitive capabilities. It is sufficient for an individual to base their decision of whether or not to cooperate on the outcome of their last encounter--even if it was with a different partner.