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The social norms of fairness and reciprocity are fundamental to cooperation and constitute core behavioral principles. Warm glow theory suggests that cooperative behavior is driven by positive emotions, whereas inequity aversion theory proposes that cooperative behavior is necessary to avoid negative feelings. However, the precise characteristics underlying the enforcement (fairness or reciprocity) and violation (unfairness or betrayal) of cooperation remain elusive. Moreover, whether the neural mechanism of cooperation as a partner or a spectator is the same remains unclear. To resolve the above issues, we summarized the findings of human cooperation neuroimaging studies through a meta-analysis. Based on our results, cooperation enforcement activates reward-related brain areas, such as the striatum and orbitofrontal cortex, only during other-cooperation. In contrast, cooperation violation is associated with the negative emotion-related insula in both self- and other-noncooperation. Thus, people expect others to cooperate rather than themselves; however, people are disgusted when cooperation is violated by themselves or others. Taken together, cooperative behavior might be mainly driven by a process designed to avoid negative emotion, which supports the inequity aversion theory but not the warm glow theory, thereby improving our understanding of cooperation theory.

Neural signatures of cooperation enforcement and violation: a coordinate-based meta-analysis