Direct reciprocity in animals: The roles of bonding and affective processes

The presence of direct reciprocity in animals is a debated topic, because, despite its evolutionary plausibility, it is believed to be uncommon. Some authors claim that stable reciprocal exchanges require sophisticated cognition which has acted as a constraint on its evolution across species. In contrast, a more recent trend of research has focused on the possibility that direct reciprocity occurs within long-term bonds and relies on simple as well as more complex affective mechanisms such as emotional book-keeping, rudimentary and higher forms of empathy, and inequity aversion, among others. First, we present evidence supporting the occurrence of long-term reciprocity in the context of existing bonds in social birds and mammals. Second, we discuss the evidence for affective responses which, modulated by bonding, may underlie altruistic behaviours in different species. We conclude that the mechanisms that may underlie reciprocal exchanges are diverse, and that some act in interaction with bonding processes. From simple associative learning in social contexts, through emotional contagion and behavioural mimicry, to empathy and a sense of fairness, widespread and diverse social affective mechanisms may explain why direct reciprocity may not be a rare phenomenon among social vertebrates.