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Modern theories of the evolution of human cooperation focus mainly on altruism. In contrast, we propose that humans’ species-unique forms of cooperation—as well as their species-unique forms of cognition, communication, and social life—all derive from mutualistic collaboration (with social selection against cheaters). In a first step, humans became obligate collaborative foragers such that individuals were interdependent with one another and so had a direct interest in the well-being of their partners. In this context, they evolved new skills and motivations for collaboration not possessed by other great apes (joint intentionality), and they helped their potential partners (and avoided cheaters). In a second step, these new collaborative skills and motivations were scaled up to group life in general, as modern humans faced competition from other groups. As part of this new group-mindedness, they created cultural conventions, norms, and institutions (all characterized by collective intentionality), with knowledge of a specific set of these marking individuals as members of a particular cultural group. Human cognition and sociality thus became ever more collaborative and altruistic as human individuals became ever more interdependent.

Significance Concern for the welfare of others is a key component of moral decision making and is disturbed in antisocial and criminal behavior. However, little is known about how people evaluate the costs of others’ suffering. Past studies have examined people’s judgments in hypothetical scenarios, but there is evidence that hypothetical judgments cannot accurately predict actual behavior. Here we addressed this issue by measuring how much money people will sacrifice to reduce the number of painful electric shocks delivered to either themselves or an anonymous stranger. Surprisingly, most people sacrifice more money to reduce a stranger’s pain than their own pain. This finding may help us better understand how people resolve moral dilemmas that commonly arise in medical, legal, and political decision making. Concern for the suffering of others is central to moral decision making. How humans evaluate others’ suffering, relative to their own suffering, is unknown. We investigated this question by inviting subjects to trade off profits for themselves against pain experienced either by themselves or an anonymous other person. Subjects made choices between different amounts of money and different numbers of painful electric shocks. We independently varied the recipient of the shocks (self vs. other) and whether the choice involved paying to decrease pain or profiting by increasing pain. We built computational models to quantify the relative values subjects ascribed to pain for themselves and others in this setting. In two studies we show that most people valued others’ pain more than their own pain. This was evident in a willingness to pay more to reduce others’ pain than their own and a requirement for more compensation to increase others’ pain relative to their own. This ”hyperaltruistic” valuation of others’ pain was linked to slower responding when making decisions that affected others, consistent with an engagement of deliberative processes in moral decision making. Subclinical psychopathic traits correlated negatively with aversion to pain for both self and others, in line with reports of aversive processing deficits in psychopathy. Our results provide evidence for a circumstance in which people care more for others than themselves. Determining the precise boundaries of this surprisingly prosocial disposition has implications for understanding human moral decision making and its disturbance in antisocial behavior.

When enforcing norms for cooperative behavior, human adults sometimes exhibit in-group bias. For example, third-party observers punish selfish behaviors committed by out-group members more harshly than similar behaviors committed by in-group members. Although evidence suggests that children begin to systematically punish selfish behavior around the age of 6 y, the development of in-group bias in their punishment remains unknown. Do children start off enforcing fairness norms impartially, or is norm enforcement biased from its emergence? How does bias change over development? Here, we created novel social groups in the laboratory and gave 6- and 8-year-olds the opportunity to engage in costly third-party punishment of selfish sharing behavior. We found that by age 6, punishment was already biased: Selfish resource allocations received more punishment when they were proposed by out-group members and when they disadvantaged in-group members. We also found that although costly punishment increased between ages 6 and 8, bias in punishment partially decreased. Although 8-y-olds also punished selfish out-group members more harshly, they were equally likely to punish on behalf of disadvantaged in-group and out-group members, perhaps reflecting efforts to enforce norms impartially. Taken together, our results suggest that norm enforcement is biased from its emergence, but that this bias can be partially overcome through developmental change.

Humans regulate intergroup conflict through parochial altruism; they self-sacrifice to contribute to in-group welfare and to aggress against competing out-groups. Parochial altruism has distinct survival functions, and the brain may have evolved to sustain and promote in-group cohesion and effectiveness and to ward off threatening out-groups. Here, we have linked oxytocin, a neuropeptide produced in the hypothalamus, to the regulation of intergroup conflict. In three experiments using double-blind placebo-controlled designs, male participants self-administered oxytocin or placebo and made decisions with financial consequences to themselves, their in-group, and a competing out-group. Results showed that oxytocin drives a \"tend and defend\" response in that it promoted in-group trust and cooperation, and defensive, but not offensive, aggression toward competing out-groups.

Are humans too generous? The discovery that subjects choose to incur costs to allocate benefits to others in anonymous, one-shot economic games has posed an unsolved challenge to models of economic and evolutionary rationality. Using agent-based simulations, we show that such generosity is the necessary byproduct of selection on decision systems for regulating dyadic reciprocity under conditions of uncertainty. In deciding whether to engage in dyadic reciprocity, these systems must balance (i) the costs of mistaking a one-shot interaction for a repeated interaction (hence, risking a single chance of being exploited) with (ii) the far greater costs of mistaking a repeated interaction for a one-shot interaction (thereby precluding benefits from multiple future cooperative interactions). This asymmetry builds organisms naturally selected to cooperate even when exposed to cues that they are in one-shot interactions.

Theoretical models suggest that social networks influence the evolution of cooperation, but to date there have been few experimental studies. Observational data suggest that a wide variety of behaviors may spread in human social networks, but subjects in such studies can choose to befriend people with similar behaviors, posing difficulty for causal inference. Here, we exploit a seminal set of laboratory experiments that originally showed that voluntary costly punishment can help sustain cooperation. In these experiments, subjects were randomly assigned to a sequence of different groups to play a series of single-shot public goods games with strangers; this feature allowed us to draw networks of interactions to explore how cooperative and uncooperative behaviors spread from person to person to person. We show that, in both an ordinary public goods game and in a public goods game with punishment, focal individuals are influenced by fellow group members' contribution behavior in future interactions with other individuals who were not a party to the initial interaction. Furthermore, this influence persists for multiple periods and spreads up to three degrees of separation (from person to person to person to person). The results suggest that each additional contribution a subject makes to the public good in the first period is tripled over the course of the experiment by other subjects who are directly or indirectly influenced to contribute more as a consequence. These results show experimentally that cooperative behavior cascades in human social networks.

People feel grateful when they have benefited from someone's costly, intentional, voluntary effort on their behalf. Experiencing gratitude motivates beneficiaries to repay their benefactors and to extend generosity to third parties. Expressions of gratitude also reinforce benefactors for their generosity. These social features distinguish gratitude from related emotions such as happiness and feelings of indebtedness. Evolutionary theories propose that gratitude is an adaptation for reciprocal altruism (the sequential exchange of costly benefits between nonrelatives) and, perhaps, upstream reciprocity (a pay-it-forward style distribution of an unearned benefit to a third party after one has received a benefit from another benefactor). Gratitude therefore may have played a unique role in human social evolution.

We present a simple framework that highlights the most fundamental requirement for the evolution of altruism: assortment between individuals carrying the cooperative genotype and the helping behaviours of others with which these individuals interact. We partition the fitness effects on individuals into those due to self and those due to the 'interaction environment', and show that it is the latter that is most fundamental to understanding the evolution of altruism. We illustrate that while kinship or genetic similarity among those interacting may generate a favourable structure of interaction environments, it is not a fundamental requirement for the evolution of altruism, and even suicidal aid can theoretically evolve without help ever being exchanged among genetically similar individuals. Using our simple framework, we also clarify a common confusion made in the literature between alternative fitness accounting methods (which may equally apply to the same biological circumstances) and unique causal mechanisms for creating the assortment necessary for altruism to be favoured by natural selection.

Reciprocal altruism has been the backbone of research on the evolution of altruistic behaviour towards non-kin, but recent research has begun to apply costly signalling theory to this problem. In addition to signalling resources or abilities, public generosity could function as a costly signal of cooperative intent, benefiting altruists in terms of (i) better access to cooperative relationships and (ii) greater cooperation within those relationships. When future interaction partners can choose with whom they wish to interact, this could lead to competition to be more generous than others. Little empirical work has tested for the possible existence of this 'competitive altruism'. Using a cooperative monetary game with and without opportunities for partner choice and signalling cooperative intent, we show here that people actively compete to be more generous than others when they can benefit from being chosen for cooperative partnerships, and the most generous people are correspondingly chosen more often as cooperative partners. We also found evidence for increased scepticism of altruistic signals when the potential reputational benefits for dishonest signalling were high. Thus, this work supports the hypothesis that public generosity can be a signal of cooperative intent, which people sometimes 'fake' when conditions permit it.