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Economic experiments have suggested that cooperative humans will altruistically match local levels of cooperation (conditional cooperation) and pay to punish non-cooperators (altruistic punishment). Evolutionary models have suggested that if altruists punish non-altruists this could favour the evolution of costly helping behaviours (cooperation) among strangers. An often-key requirement is that helping behaviours and punishing behaviours form one single conjoined trait (strong reciprocity). Previous economics experiments have provided support for the hypothesis that punishment and cooperation form one conjoined, altruistically motivated, trait. However, such a conjoined trait may be evolutionarily unstable, and previous experiments have confounded a fear of being punished with being surrounded by cooperators, two factors that could favour cooperation. Here, we experimentally decouple the fear of punishment from a cooperative environment and allow cooperation and punishment behaviour to freely separate (420 participants). We show, that if a minority of individuals is made immune to punishment, they (i) learn to stop cooperating on average despite being surrounded by high levels of cooperation, contradicting the idea of conditional cooperation and (ii) often continue to punish, 'hypocritically', showing that cooperation and punishment do not form one, altruistically motivated, linked trait.

Strong reciprocity (SR) has recently been subject to heated debate. In this debate, the "West camp" (West et al. in Evol Hum Behav 32(4):231-262, 2011), which is critical of the case for SR, and the "Laland camp" (Laland et al. in Science, 334(6062):1512-1516, 2011, Biol Philos 28(5):719-745, 2013), which is sympathetic to the case of SR, seem to take diametrically opposed positions. The West camp criticizes advocates of SR for conflating proximate and ultimate causation. SR is said to be a proximate mechanism that is put forward by its advocates as an ultimate explanation of human cooperation. The West camp thus accuses advocates of SR for not heeding Mayr's original distinction between ultimate and proximate causation. The Laland camp praises advocates of SR for revising Mayr's distinction. Advocates of SR are said to replace Mayr's uni-directional view on the relation between ultimate and proximate causes by the bi-directional one of reciprocal causation. The paper argues that both the West camp and the Laland camp misrepresent what advocates of SR are up to. The West camp is right that SR is a proximate cause of human cooperation. But rather than putting forward SR as an ultimate explanation, as the West camp argues, advocates of SR believe that SR itself is in need of ultimate explanation. Advocates of SR tend to take gene-culture co-evolutionary theory as the correct meta-theoretical framework for advancing ultimate explanations of SR. Appearances notwithstanding, gene-culture coevolutionary theory does not imply Laland et al.'s notion of reciprocal causation. "Reciprocal causation" suggests that proximate and ultimate causes interact simultaneously, while advocates of SR assume that they interact sequentially. I end by arguing that the best way to understand the debate is by disambiguating Mayr's ultimate-proximate distinction. I propose to reserve "ultimate" and "proximate" for different sorts of explanations, and to use other terms for distinguishing different kinds of causes and different parts of the total causal chain producing behavior.

Humans regularly help strangers, even when interactions are apparently unobserved and unlikely to be repeated. Such situations have been simulated in the laboratory using anonymous one-shot games (e.g., prisoner's dilemma) where the payoff matrices used make helping biologically altruistic. As in real-life, participants often cooperate in the lab in these one-shot games with non-relatives, despite that fact that helping is under negative selection under these circumstances. Two broad explanations for such behavior prevail. The "big mistake" or "mismatch" theorists argue that behavior is constrained by psychological mechanisms that evolved predominantly in the context of repeated interactions with known individuals. In contrast, the cultural group selection theorists posit that humans have been selected to cooperate in anonymous one-shot interactions due to strong between-group competition, which creates interdependence among in-group members. We present these two hypotheses before discussing alternative routes by which humans could increase their direct fitness by cooperating with strangers under natural conditions. In doing so, we explain why the standard lab games do not capture real-life in various important aspects. First, asymmetries in the cost of perceptual errors regarding the context of the interaction (one-shot vs. repeated; anonymous vs. public) might have selected for strategies that minimize the chance of making costly behavioral errors. Second, helping strangers might be a successful strategy for identifying other cooperative individuals in the population, where partner choice can turn strangers into interaction partners. Third, in contrast to the assumptions of the prisoner's dilemma model, it is possible that benefits of cooperation follow a non-linear function of investment. Non-linear benefits result in negative frequency dependence even in one-shot games. Finally, in many real-world situations individuals are able to parcel investments such that a one-shot interaction is turned into a repeated game of many decisions.

Strong reciprocity is considered here as the propensity to sacrifice resources to be kind or to punish in response to prior acts, a behavior not simply reducible to self-interest and a likely force behind human cooperation and sociality. The aim was to capture emerging signs of strong reciprocity in human ontogeny and across highly contrasted cultures. Three- and 5-year-old middle class American children (N = 162) were tested in a simple, multiple round, three-way sharing game involving the child, a generous puppet, and a stingy puppet. At the end of the game, the child was offered an opportunity to sacrifice some of her personal gains to punish one of the puppets. By 3 years, American children demonstrate a willingness to engage in costly punishment. However, only 5-year-olds show some evidence of strong reciprocity by orienting their punishment systematically toward the stingy puppet. Further analyses and three additional control conditions demonstrate that such propensity is not simply reducible to (a) straight imitation, or (b) inequity aversion. To assess the relative universality of such development, a group of 5- to 6-year-old children from rural Samoa (N = 14) were tested and compared to age and gender-matched American children. Samoan children did not manifest the same propensity toward strong reciprocity. The results are interpreted as pointing to (1) the developmental emergence of an ethical stance between 3 and 5 years of age, and (2) that the expression of such stance by young children could depend on culture.

The origin of altruism remains one of the most enduring puzzles of human behaviour. Indeed, true altruism is often thought either not to exist, or to arise merely as a miscalculation of otherwise selfish behaviour. In this paper, we argue that altruism emerges directly from the way in which distinct human decision-making systems learn about rewards. Using insights provided by neurobiological accounts of human decision-making, we suggest that reinforcement learning in game-theoretic social interactions (habitisation over either individuals or games) and observational learning (either imitative of inference based) lead to altruistic behaviour. This arises not only as a result of computational efficiency in the face of processing complexity, but as a direct consequence of optimal inference in the face of uncertainty. Critically, we argue that the fact that evolutionary pressure acts not over the object of learning ('what' is learned), but over the learning systems themselves ('how' things are learned), enables the evolution of altruism despite the direct threat posed by free-riders.

The concept of cooperative communities that enforce norm conformity through reward, as well as shaming, ridicule, and ostracism, has been central to anthropology since the work of Durkheim. Prevailing approaches from evolutionary theory explain the willingness to exert sanctions to enforce norms as self-interested behavior, while recent experimental studies suggest that altruistic rewarding and punishing-"strong reciprocity"-play an important role in promoting cooperation. This paper will use data from 308 conversations among the Ju/'hoansi (!Kung) Bushmen (a) to examine the dynamics of norm enforcement, (b) to evaluate the costs of punishment in a forager society and understand how they are reduced, and, ((c) to determine whether hypotheses that center on individual self-interest provide sufficient explanations for bearing the costs of norm enforcement, or whether there is evidence for strong reciprocity.)

This paper provides strong evidence challenging the self-interest assumption that dominates the behavioral sciences and much evolutionary thinking. The evidence indicates that many people have a tendency to voluntarily cooperate, if treated fairly, and to punish noncooperators. We call this behavioral propensity "strong reciprocity" and show empirically that it can lead to almost universal cooperation in circumstances in which purely self-interested behavior would cause a complete breakdown of cooperation. In addition, we show that people are willing to punish those who behaved unfairly towards a third person or who defected in a Prisoner's Dilemma game with a third person. This suggests that strong reciprocity is a powerful device for the enforcement of social norms involving, for example, food sharing or collective action. Strong reciprocity cannot be rationalized as an adaptive trait by the leading evolutionary theories of human cooperation (in other words, kin selection, reciprocal altruism, indirect reciprocity, and costly signaling theory). However, multilevel selection theories of cultural evolution are consistent with strong reciprocity.