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Human cooperation is an evolutionary puzzle. Unlike other creatures, people frequently cooperate with genetically unrelated strangers, often in large groups, with people they will never meet again, and when reputation gains are small or absent. These patterns of cooperation cannot be explained by the nepotistic motives associated with the evolutionary theory of kin selection and the selfish motives associated with signalling theory or the theory of reciprocal altruism. Here we show experimentally that the altruistic punishment of defectors is a key motive for the explanation of cooperation. Altruistic punishment means that individuals punish, although the punishment is costly for them and yields no material gain. We show that cooperation flourishes if altruistic punishment is possible, and breaks down if it is ruled out. The evidence indicates that negative emotions towards defectors are the proximate mechanism behind altruistic punishment. These results suggest that future study of the evolution of human cooperation should include a strong focus on explaining altruistic punishment.

Most economic models are based on the self-interest hypothesis that assumes that material self-interest exclusively motivates all people. Experimental economists have gathered overwhelming evidence in recent years, however, that systematically refutes the self-interest hypothesis, suggesting that concerns for altruism, fairness, and reciprocity strongly motivate many people. Moreover, several theoretical papers demonstrate that the observed phenomena can be explained in a rigorous and tractable manner. These theories then induced a first wave of experimental research which offered exciting insights into both the nature of preferences and the relative performance of competing fairness theories. The purpose of this chapter is to review these developments, to point out open questions, and to suggest avenues for future research. We also discuss recent neuroeconomic evidence that is consistent with the view that many people have a taste for mutual cooperation and the punishment of norm violators. We further illustrate the powerful impact of fairness concerns on cooperation, competition, incentives, and contract design.

The existence of cooperation and social order among genetically unrelated individuals is a fundamental problem in the behavioural sciences. The prevailing approaches in biology and economics view cooperation exclusively as self-interested behaviour—unrelated individuals cooperate only if they face economic rewards or sanctions rendering cooperation a self-interested choice. Whether economic incentives are perceived as just or legitimate does not matter in these theories. Fairness-based altruism is, however, a powerful source of human cooperation. Here we show experimentally that the prevailing self-interest approach has serious shortcomings because it overlooks negative effects of sanctions on human altruism. Sanctions revealing selfish or greedy intentions destroy altruistic cooperation almost completely, whereas sanctions perceived as fair leave altruism intact. These findings challenge proximate and ultimate theories of human cooperation that neglect the distinction between fair and unfair sanctions, and they are probably relevant in all domains in which voluntary compliance matters—in relations between spouses, in the education of children, in business relations and organizations as well as in markets.

Human social interaction is strongly shaped by other-regarding preferences, that is, a concern for the welfare of others. These preferences are important for a unique aspect of human sociality—large scale cooperation with genetic strangers—but little is known about their developmental roots. Here we show that young children’s other-regarding preferences assume a particular form, inequality aversion that develops strongly between the ages of 3 and 8. At age 3–4, the overwhelming majority of children behave selfishly, whereas most children at age 7–8 prefer resource allocations that remove advantageous or disadvantageous inequality. Moreover, inequality aversion is strongly shaped by parochialism, a preference for favouring the members of one’s own social group. These results indicate that human egalitarianism and parochialism have deep developmental roots, and the simultaneous emergence of altruistic sharing and parochialism during childhood is intriguing in view of recent evolutionary theories which predict that the same evolutionary process jointly drives both human altruism and parochialism.

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.

Some of the most fundamental questions concerning our evolutionary origins, our social relations, and the organization of society are centred around issues of altruism and selfishness. Experimental evidence indicates that human altruism is a powerful force and is unique in the animal world. However, there is much individual heterogeneity and the interaction between altruists and selfish individuals is vital to human cooperation. Depending on the environment, a minority of altruists can force a majority of selfish individuals to cooperate or, conversely, a few egoists can induce a large number of altruists to defect. Current gene-based evolutionary theories cannot explain important patterns of human altruism, pointing towards the importance of both theories of cultural evolution as well as gene–culture co-evolution.

Many people voluntarily incur costs to punish violations of social norms. Evolutionary models and empirical evidence indicate that such altruistic punishment has been a decisive force in the evolution of human cooperation. We used H2 15O positron emission tomography to examine the neural basis for altruistic punishment of defectors in an economic exchange. Subjects could punish defection either symbolically or effectively. Symbolic punishment did not reduce the defector’s economic payoff, whereas effective punishment did reduce the payoff. We scanned the subjects’ brains while they learned about the defector’s abuse of trust and determined the punishment. Effective punishment, as compared with symbolic punishment, activated the dorsal striatum, which has been implicated in the processing of rewards that accrue as a result of goal-directed actions. Moreover, subjects with stronger activations in the dorsal striatum were willing to incur greater costs in order to punish. Our findings support the hypothesis that people derive satisfaction from punishing norm violations and that the activation in the dorsal striatum reflects the anticipated satisfaction from punishing defectors.

Social norms and the associated altruistic behaviours are decisive for the evolution of human cooperation1,2,3,4,5,6,7,8,9 and the maintenance of social order10, and they affect family life, politics11 and economic interactions12. However, as altruistic norm compliance and norm enforcement often emerge in the context of inter-group conflicts13,14, they are likely to be shaped by parochialism15—a preference for favouring the members of one's ethnic, racial or language group. We have conducted punishment experiments16, which allow ‘impartial’ observers to punish norm violators, with indigenous groups in Papua New Guinea. Here we show that these experiments confirm the prediction of parochialism. We found that punishers protect ingroup victims—who suffer from a norm violation—much more than they do outgroup victims, regardless of the norm violator's group affiliation. Norm violators also expect that punishers will be lenient if the latter belong to their social group. As a consequence, norm violations occur more often if the punisher and the norm violator belong to the same group. Our results are puzzling for evolutionary multi-level selection theories based on selective group extinction2,3,4,5 as well as for theories of individual selection17,18,19; they also indicate the need to explicitly examine the interactions between individuals stemming from different groups in evolutionary models.

Combining the methods of neuroscience and economics generates powerful tools for studying the brain processes behind human social interaction. We argue that hedonic interpretations of theories of social preferences provide a useful framework that generates interesting predictions and helps interpret brain activations involved in altruistic, fair and trusting behaviors. These behaviors are consistently associated with activation in reward-related brain areas, such as the striatum, and with prefrontal activity implicated in cognitive control, the processing of emotions, and integration of benefits and costs, consistent with resolution of a conflict between self-interest and other-regarding motives.