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Natural selection favors individuals to act in their own interests, implying that wild animals experience a competitive psychology. Animals in the wild also express helping behaviors, presumably at their own expense and suggestive of a more compassionate psychology. This apparent paradox can be partially explained by ultimate mechanisms that include kin selection, reciprocity, and multilevel selection, yet some theorists argue such ultimate explanations may not be sufficient and that an additional "stake in others" is necessary for altruism's evolution. We suggest this stake is the "camaraderie effect," a by-product of two highly adaptive psychological experiences: social motivation and empathy. Rodents can derive pleasure from access to others and this appetite for social rewards motivates individuals to live together, a valuable psychology when group living is adaptive. Rodents can also experience empathy, the generation of an affective state more appropriate to the situation of another compared to one's own. Empathy is not a compassionate feeling but it has useful predictive value. For instance, empathy allows an individual to feel an unperceived danger from social cues. Empathy of another's stance toward one's self would predict either social acceptance or ostracism and amplify one's physiological sensitivity to social isolation, including impaired immune responses and delayed wound healing. By contrast, altruistic behaviors would promote well-being in others and feelings of camaraderie from others, thereby improving one's own physiological well-being. Together, these affective states engender a stake in others necessary for the expression of altruistic behavior.

Social Reward and Empathy as Proximal Contributions to Altruism: The Camaraderie Effect