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Altruism—benefiting fellow group members at a cost to oneself—and parochialism—hostility toward individuals not of one's own ethnic, racial, or other group—are common human behaviors. The intersection of the two—which we term “parochial altruism”—is puzzling from an evolutionary perspective because altruistic or parochial behavior reduces one's payoffs by comparison to what one would gain by eschewing these behaviors. But parochial altruism could have evolved if parochialism promoted intergroup hostilities and the combination of altruism and parochialism contributed to success in these conflicts. Our game-theoretic analysis and agent-based simulations show that under conditions likely to have been experienced by late Pleistocene and early Holocene humans, neither parochialism nor altruism would have been viable singly, but by promoting group conflict, they could have evolved jointly.

Why do humans, uniquely among animals, cooperate in large numbers to advance projects for the common good? Contrary to the conventional wisdom in biology and economics, this generous and civic-minded behavior is widespread and cannot be explained simply by far-sighted self-interest or a desire to help close genealogical kin.In A Cooperative Species, Samuel Bowles and Herbert Gintis--pioneers in the new experimental and evolutionary science of human behavior--show that the central issue is not why selfish people act generously, but instead how genetic and cultural evolution has produced a species in which substantial numbers make sacrifices to uphold ethical norms and to help even total strangers. The authors describe how, for thousands of generations, cooperation with fellow group members has been essential to survival. Groups that created institutions to protect the civic-minded from exploitation by the selfish flourished and prevailed in conflicts with less cooperative groups. Key to this process was the evolution of social emotions such as shame and guilt, and our capacity to internalize social norms so that acting ethically became a personal goal rather than simply a prudent way to avoid punishment. Using experimental, archaeological, genetic, and ethnographic data to calibrate models of the coevolution of genes and culture as well as prehistoric warfare and other forms of group competition, A Cooperative Species provides a compelling and novel account of how humans came to be moral and cooperative.

Both laboratory and field data suggest that people punish noncooperators even in one-shot interactions. Although such “altruistic punishment” may explain the high levels of cooperation in human societies, it creates an evolutionary puzzle: existing models suggest that altruistic cooperation among nonrelatives is evolutionarily stable only in small groups. Thus, applying such models to the evolution of altruistic punishment leads to the prediction that people will not incur costs to punish others to provide benefits to large groups of nonrelatives. However, here we show that an important asymmetry between altruistic cooperation and altruistic punishment allows altruistic punishment to evolve in populations engaged in one-time, anonymous interactions. This process allows both altruistic punishment and altruistic cooperation to be maintained even when groups are large and other parameter values approximate conditions that characterize cultural evolution in the small-scale societies in which humans lived for most of our prehistory.

Recent experimental research has revealed forms of human behavior involving interaction among unrelated individuals that have proven difficult to explain in terms of kin or reciprocal altruism. One such trait, strong reciprocity is a predisposition to cooperate with others and to punish those who violate the norms of cooperation, at personal cost, even when it is implausible to expect that these costs will be repaid. We present evidence supporting strong reciprocity as a schema for predicting and understanding altruism in humans. We show that under conditions plausibly characteristic of the early stages of human evolution, a small number of strong reciprocators could invade a population of self-regarding types, and strong reciprocity is an evolutionary stable strategy. Although most of the evidence we report is based on behavioral experiments, the same behaviors are regularly described in everyday life, for example, in wage setting by firms, tax compliance, and cooperation in the protection of local environmental public goods.

Humans behave altruistically in natural settings and experiments. A possible explanation—that groups with more altruists survive when groups compete—has long been judged untenable on empirical grounds for most species. But there have been no empirical tests of this explanation for humans. My empirical estimates show that genetic differences between early human groups are likely to have been great enough so that lethal intergroup competition could account for the evolution of altruism. Crucial to this process were distinctive human practices such as sharing food beyond the immediate family, monogamy, and other forms of reproductive leveling. These culturally transmitted practices presuppose advanced cognitive and linguistic capacities, possibly accounting for the distinctive forms of altruism found in our species.Early human practices requiring language and sophisticated cognition enhanced the contribution of altruism to group survival, perhaps selecting for altruistic traits. Early human practices requiring language and sophisticated cognition enhanced the contribution of altruism to group survival, perhaps selecting for altruistic traits.