Social bonds facilitate cooperative resource sharing in wild chimpanzees.

Why share when access to benefits is uncertain is crucial to our understanding of the evolution of humans' extensive cooperation. Here, we investigated some of the different human sharing hypotheses and potential neuroendocrine mechanisms, in one of our closest living relatives, chimpanzees. The strongest predictor of sharing across food types was the presence of enduring and mutually preferred grooming partners, more than harassment, direct signalling, or trade. Moreover, urinary oxytocin levels were higher after the sharing of both individually and jointly acquired resources compared with controls. We conclude that the emotional connection inherent in social bonds was a key factor determining sharing patterns, with the oxytocinergic system potentially facilitating long-term cooperative exchanges. Testing for the role of social bonds in increasing predictability of sharing behaviour, a feature frequently overlooked, may help us to identify the evolutionary drivers of resource sharing and mechanisms that sustain delayed reciprocity between non-kin.

Urinary oxytocin levels in relation to post-conflict affiliations in wild male chimpanzees (Pan troglodytes verus).

Many animals living in social groups have evolved behaviors to resolve conflicts between group members, behaviors thought crucial for maintaining stable group life. Several hypotheses, based mainly on observational data, aim to explain how post-conflict (PC) affiliations, such as reconciliation and consolation, resolve conflicts by restoring relationships and/or alleviating anxiety. To examine a potential endocrinological mechanism of PC affiliations, we used an experimental-like procedure to investigate whether the oxytocinergic system is activated during naturally observed reconciliations, receiving bystander PC affiliations and aggressions not followed by PC affiliations in wild male chimpanzees. We compared urinary oxytocin (uOT) levels after reconciliations, receiving bystander PC affiliations or aggressions without affiliations with two control conditions: affiliations without previous aggression and after time periods without social interactions. We furthermore tested the 'valuable relationship' hypothesis of reconciliation, as well as the influence of relationship quality between individuals engaged in each of the three behavioral conditions involving aggression on uOT levels. We found that the probability to reconcile a conflict increased with increasing relationship quality between opponents, thus our results support the 'valuable relationship' hypothesis. However, relationship quality did not influence uOT levels, while behavioral condition had a significant effect on uOT levels. uOT levels after reconciliations, receiving bystander PC affiliations and affiliations not related to conflicts were higher than after aggressions alone and time periods without social interactions. Overall, our results indicate that the oxytocinergic system is activated during affiliative interactions, whether occurring as reconciliation, bystander PC affiliation or affiliation alone. We conclude that the oxytocinergic system, in addition to building and maintaining social relationships, also takes part in repairing them.

Cortisol and oxytocin show independent activity during chimpanzee intergroup conflict.

The oxytocinergic system is involved in a range of functions, from attachment and social bonding to aggression and stress responses. Whether oxytocin is released in response to a stressor, shows contradictory results across species and potential contexts-dependent differences. To avoid unintended contextual changes due to experimental procedures, we tested this question non-invasively in wild chimpanzees in an ecologically valid context. We collected endogenous hormonal measures during exposure to a known natural stressor, intergroup conflict. Specifically, we tested for potential synchronous activation patterns between urinary oxytocin and cortisol in male and female chimpanzees during stressor exposure. Oxytocinergic system reactivity during chimpanzee intergroup conflict has already been established in this study population. Thus, we first investigated urinary cortisol levels during border patrol and intergroup encounter days, in comparison to another potential stressor, hunting, and control days. We found higher urinary cortisol levels during intergroup encounter days compared with control and hunting days. We then compared secretion patterns of oxytocin and cortisol in relation to increased levels of out-group contact and hostility ('out-group risk') during intergroup conflict. We found that increased 'out-group risk' was associated with higher cortisol levels, especially when involving direct visual or physical contact with rival groups. Although urinary oxytocin levels were high across intergroup conflict contexts, increasing levels of out-group risk showed no significant variation. Taken together, results indicate independent secretion of oxytocin and cortisol during chimpanzee intergroup conflict, emphasizing that stressor exposure in this context is not the main trigger of oxytocin secretion.