Reliance on functional resting-state network for stable task control predicts behavioral tendency for cooperation.

Humans display individual variability in cooperative behavior. While an ever-growing body of research has investigated the neural correlates of task-specific cooperation, the mechanisms by which situation-independent, stable differences in cooperation render behavior consistent across a wide range of situations remain elusive. Addressing this issue, we show that the individual tendency to behave in a prosocial or individualistic manner can be predicted from the functional resting-state connectome. More specifically, connections of the cinguloopercular network which supports goal-directed behavior encode cooperative tendency. Effects of virtual lesions to this network on the efficacy of information exchange throughout the brain corroborate our findings. These results shed light on the neural mechanisms underlying individualists' and prosocials' habitual social decisions by showing that reliance on the cinguloopercular task-control network predicts stable cooperative behavior. Based on this evidence, we provide a unifying framework for the interpretation of functional imaging and behavioral studies of cooperative behavior.

Costs for switching partners reduce network dynamics but not cooperative behaviour.

Social networks represent the structuring of interactions between group members. Above all, many interactions are profoundly cooperative in humans and other animals. In accordance with this natural observation, theoretical work demonstrates that certain network structures favour the evolution of cooperation. Yet, recent experimental evidence suggests that static networks do not enhance cooperative behaviour in humans. By contrast, dynamic networks do foster cooperation. However, costs associated with dynamism such as time or resource investments in finding and establishing new partnerships have been neglected so far. Here, we show that human participants are much less likely to break links when costs arise for building new links. Especially, when costs were high, the network was nearly static. Surprisingly, cooperation levels in Prisoner's Dilemma games were not affected by reduced dynamism in social networks. We conclude that the mere potential to quit collaborations is sufficient in humans to reach high levels of cooperative behaviour. Effects of self-structuring processes or assortment on the network played a minor role: participants simply adjusted their cooperative behaviour in response to the threats of losing a partner or of being expelled.

Human strategy updating in evolutionary games.

Evolutionary game dynamics describe not only frequency-dependent genetic evolution, but also cultural evolution in humans. In this context, successful strategies spread by imitation. It has been shown that the details of strategy update rules can have a crucial impact on evolutionary dynamics in theoretical models and, for example, can significantly alter the level of cooperation in social dilemmas. What kind of strategy update rules can describe imitation dynamics in humans? Here, we present a way to measure such strategy update rules in a behavioral experiment. We use a setting in which individuals are virtually arranged on a spatial lattice. This produces a large number of different strategic situations from which we can assess strategy updating. Most importantly, spontaneous strategy changes corresponding to mutations or exploration behavior are more frequent than assumed in many models. Our experimental approach to measure properties of the update mechanisms used in theoretical models will be useful for mathematical models of cultural evolution.

Local orientation and the evolution of foraging: changes in decision making can eliminate evolutionary trade-offs.

Information processing is a major aspect of the evolution of animal behavior. In foraging, responsiveness to local feeding opportunities can generate patterns of behavior which reflect or "recognize patterns" in the environment beyond the perception of individuals. Theory on the evolution of behavior generally neglects such opportunity-based adaptation. Using a spatial individual-based model we study the role of opportunity-based adaptation in the evolution of foraging, and how it depends on local decision making. We compare two model variants which differ in the individual decision making that can evolve (restricted and extended model), and study the evolution of simple foraging behavior in environments where food is distributed either uniformly or in patches. We find that opportunity-based adaptation and the pattern recognition it generates, plays an important role in foraging success, particularly in patchy environments where one of the main challenges is "staying in patches". In the restricted model this is achieved by genetic adaptation of move and search behavior, in light of a trade-off on within- and between-patch behavior. In the extended model this trade-off does not arise because decision making capabilities allow for differentiated behavioral patterns. As a consequence, it becomes possible for properties of movement to be specialized for detection of patches with more food, a larger scale information processing not present in the restricted model. Our results show that changes in decision making abilities can alter what kinds of pattern recognition are possible, eliminate an evolutionary trade-off and change the adaptive landscape.

Patch depletion, niche structuring and the evolution of co-operative foraging.

BACKGROUND: Many animals live in groups. One proposed reason is that grouping allows cooperative food finding. Group foraging models suggest that grouping could increase food finding rates, but that such group processes could be evolutionarily unstable. These models assume discrete food patches which are fully detectable. However, often animals may only be able to perceive local parts of larger-scale environmental patterns. We therefore use a spatial individual-based model where food patches are aggregates of food items beyond the scale of individual perception. We then study the evolution of foraging and grouping behavior in environments with different resource distributions. RESULTS: Our results show that grouping can evolve to increase food intake rates. Two kinds of grouping evolve: traveling pairs and opportunistic grouping, where individuals only aggregate when feeding. Grouping evolves because it allows individuals to better sense and deplete patches. Such enhanced patch depletion is particularly apparent on fragmented and partially depleted patches, which are especially difficult for solitary foragers to deplete. Solitary foragers often leave a patch prematurely because a whole patch cannot be observed directly. In groups, individuals that are still eating allow other individuals that inadvertently leave the patch, to return and continue feeding. For this information sharing a grouping tendency is sufficient and observing whether a neighbor is eating is not necessary. Grouping therefore leads to a release from individual sensing constraints and a shift in niche specialization, allowing individuals to better exploit partially depleted patches. CONCLUSIONS: The evolved group foraging can be seen as cooperative in the sense that it leads to a mutually-beneficial synergy: together individuals can achieve more than on their own. This cooperation exists as a group-level process generated by the interaction between grouping and the environment. Thus we reveal how such a synergy can originate in evolution as a side-effect of grouping via multi-level selection. Here there is no cooperative dilemma as individuals cannot avoid producing information for their neighbors. This scenario may be a useful starting point for studying the evolution of further social and cooperative complexity.

Co-evolution of behaviour and social network structure promotes human cooperation.

The ubiquity of cooperation in nature is puzzling because cooperators can be exploited by defectors. Recent theoretical work shows that if dynamic networks define interactions between individuals, cooperation is favoured by natural selection. To address this, we compare cooperative behaviour in multiple but independent repeated games between participants in static and dynamic networks. In the latter, participants could break their links after each social interaction. As predicted, we find higher levels of cooperation in dynamic networks. Through biased link breaking (i.e. to defectors) participants affected their social environment. We show that this link-breaking behaviour leads to substantial network clustering and we find primarily cooperators within these clusters. This assortment is remarkable because it occurred on top of behavioural assortment through direct reciprocity and beyond the perception of participants, and represents a self-organized pattern. Our results highlight the importance of the interaction between ecological context and selective pressures on cooperation.

Volunteering leads to rock-paper-scissors dynamics in a public goods game.

Collective efforts are a trademark of both insect and human societies. They are achieved through relatedness in the former and unknown mechanisms in the latter. The problem of achieving cooperation among non-kin has been described as the 'tragedy of the commons', prophesying the inescapable collapse of many human enterprises. In public goods experiments, initial cooperation usually drops quickly to almost zero. It can be maintained by the opportunity to punish defectors or the need to maintain good reputation. Both schemes require that defectors are identified. Theorists propose that a simple but effective mechanism operates under full anonymity. With optional participation in the public goods game, 'loners' (players who do not join the group), defectors and cooperators will coexist through rock-paper-scissors dynamics. Here we show experimentally that volunteering generates these dynamics in public goods games and that manipulating initial conditions can produce each predicted direction. If, by manipulating displayed decisions, it is pretended that defectors have the highest frequency, loners soon become most frequent, as do cooperators after loners and defectors after cooperators. On average, cooperation is perpetuated at a substantial level.

Humans choose representatives who enforce cooperation in social dilemmas through extortion.

Social dilemmas force players to balance between personal and collective gain. In many dilemmas, such as elected governments negotiating climate-change mitigation measures, the decisions are made not by individual players but by their representatives. However, the behaviour of representatives in social dilemmas has not been investigated experimentally. Here inspired by the negotiations for greenhouse-gas emissions reductions, we experimentally study a collective-risk social dilemma that involves representatives deciding on behalf of their fellow group members. Representatives can be re-elected or voted out after each consecutive collective-risk game. Selfish players are preferentially elected and are hence found most frequently in the 'representatives' treatment. Across all treatments, we identify the selfish players as extortioners. As predicted by our mathematical model, their steadfast strategies enforce cooperation from fair players who finally compensate almost completely the deficit caused by the extortionate co-players. Everybody gains, but the extortionate representatives and their groups gain the most.

Donors to charity gain in both indirect reciprocity and political reputation.

Darwinian evolution can explain human cooperative behaviour among non-kin by either direct or indirect reciprocity. In the latter case one does not expect a return for an altruistic act from the recipient as with direct reciprocity, but from another member of the social group. However, the widespread human behaviour of donating to poor people outside the social group, for example, to charity organizations, that are unlikely to reciprocate indirectly and thus are equivalent to defectors in the game is still an evolutionary puzzle. Here we show experimentally that donations made in public to a well-known relief organization resulted both in increased income (that the donors received from the members of their group) and in enhanced political reputation (they were elected to represent the interests of their group). Donations may thus function as an honest signal for one's social reliability.

A comparative analysis of spatial Prisoner's Dilemma experiments: conditional cooperation and payoff irrelevance.

We have carried out a comparative analysis of data collected in three experiments on Prisoner's Dilemmas on lattices available in the literature. We focus on the different ways in which the behavior of human subjects can be interpreted, in order to empirically narrow down the possibilities for behavioral rules. Among the proposed update dynamics, we find that the experiments do not provide significant evidence for non-innovative game dynamics such as imitate-the-best or pairwise comparison rules, whereas moody conditional cooperation is supported by the data from all three experiments. This conclusion questions the applicability of many theoretical models that have been proposed to understand human behavior in spatial Prisoner's Dilemmas. A rule compatible with all our experiments, moody conditional cooperation, suggests that there is no detectable influence of interaction networks on the emergence of cooperation in behavioral experiments.

Consistent strategy updating in spatial and non-spatial behavioral experiments does not promote cooperation in social networks.

The presence of costly cooperation between otherwise selfish actors is not trivial. A prominent mechanism that promotes cooperation is spatial population structure. However, recent experiments with human subjects report substantially lower level of cooperation then predicted by theoretical models. We analyze the data of such an experiment in which a total of 400 players play a Prisoner's Dilemma on a 4×4 square lattice in two treatments, either interacting via a fixed square lattice (15 independent groups) or with a population structure changing after each interaction (10 independent groups). We analyze the statistics of individual decisions and infer in which way they can be matched with the typical models of evolutionary game theorists. We find no difference in the strategy updating between the two treatments. However, the strategy updates are distinct from the most popular models which lead to the promotion of cooperation as shown by computer simulations of the strategy updating. This suggests that the promotion of cooperation by population structure is not as straightforward in humans as often envisioned in theoretical models.

I dare you to punish me-vendettas in games of cooperation.

Everybody has heard of neighbours, who have been fighting over some minor topic for years. The fight goes back and forth, giving the neighbours a hard time. These kind of reciprocal punishments are known as vendettas and they are a cross-cultural phenomenon. In evolutionary biology, punishment is seen as a mechanism for maintaining cooperative behaviour. However, this notion of punishment excludes vendettas. Vendettas pose a special kind of evolutionary problem: they incur high costs on individuals, i.e. costs of punishing and costs of being punished, without any benefits. Theoretically speaking, punishment should be rare in dyadic relationships and vendettas would not evolve under natural selection. In contrast, punishment is assumed to be more efficient in group environments which then can pave the way for vendettas. Accordingly, we found that under the experimental conditions of a prisoner's dilemma game, human participants punished only rarely and vendettas are scarce. In contrast, we found that participants retaliated frequently in the group environment of a public goods game. They even engaged in cost-intense vendettas (i.e. continuous retaliation), especially when the first punishment was unjustified or ambiguous. Here, punishment was mainly targeted at defectors in the beginning, but provocations led to mushrooming of counter-punishments. Despite the counter-punishing behaviour, participants were able to enhance cooperation levels in the public goods game. Few participants even seemed to anticipate the outbreak of costly vendettas and delayed their punishment to the last possible moment. Overall, our results highlight the importance of different social environments while studying punishment as a cooperation-enhancing mechanism.

How is human cooperation different?

Although cooperation is a widespread phenomenon in nature, human cooperation exceeds that of all other species with regard to the scale and range of cooperative activities. Here we review and discuss differences between humans and non-humans in the strategies employed to maintain cooperation and control free-riders. We distinguish forms of cooperative behaviour based on their influence on the immediate payoffs of actor and recipient. If the actor has immediate costs and only the recipient obtains immediate benefits, we term this investment. If the behaviour has immediate positive effects for both actor and recipient, we call this a self-serving mutually beneficial behaviour or mutual cooperation. We argue that humans, in contrast to all other species, employ a wider range of enforcement mechanisms, which allow higher levels of cooperation to evolve and stabilize among unrelated individuals and in large groups. We also discuss proximate mechanisms underlying cooperative behaviour and focus on our experimental work with humans and our closest primate relatives. Differences in the proximate mechanisms also seem to contribute to explaining humans' greater ability to cooperate and enforce cooperation.

Gossip as an alternative for direct observation in games of indirect reciprocity.

Communication about social topics is abundant in human societies, and many functions have been attributed to such gossiping. One of these proposed functions is the management of reputations. Reputation by itself has been shown to have a strong influence on cooperation dynamics in games of indirect reciprocity, and this notion helps to explain the observed high level of cooperation in humans. Here we designed a game to test a widespread assumption that gossip functions as a vector for the transmission of social information. This empirical study (with 14 groups of nine students each) focuses on the composition of gossip, information transfer by gossip, and the behavior based on gossip information. We show that gossip has a strong influence on the resulting behavior even when participants have access to the original information (i.e., direct observation) as well as gossip about the same information. Thus, it is evident that gossip has a strong manipulative potential. Furthermore, gossip about cooperative individuals is more positive than gossip about uncooperative individuals, gossip comments transmit social information successfully, and cooperation levels are higher when people encounter positive compared with negative gossip.

Stabilizing the earth's climate is not a losing game: supporting evidence from public goods experiments.

Maintaining the Earth's climate within habitable boundaries is probably the greatest "public goods game" played by humans. However, with >6 billion "players" taking part, the game seems to rule out individual altruistic behavior. Thus, climate protection is a problem of sustaining a public resource that everybody is free to overuse, a "tragedy of the commons" problem that emerges in many social dilemmas. We perform a previously undescribed type of public goods experiment with human subjects contributing to a public pool. In contrast to the standard protocol, here the common pool is not divided among the participants; instead, it is promised that the pool will be invested to encourage people to reduce their fossil fuel use. Our extensive experiments demonstrate that players can behave altruistically to maintain the Earth's climate given the right set of circumstances. We find a nonzero basic level of altruistic behavior, which is enhanced if the players are provided with expert information describing the state of knowledge in climate research. Furthermore, personal investments in climate protection increase substantially if players can invest publicly, thus gaining social reputation. This increase occurs because subjects reward other subjects' contributions to sustaining the climate, thus reinforcing their altruism. Therefore, altruism may convert to net personal benefit and to relaxing the dilemma if the gain in reputation is large enough. Our finding that people reward contributions to sustaining the climate of others is a surprising result. There are obvious ways these unexpected findings can be applied on a large scale.

Reputation helps solve the 'tragedy of the commons'.

The problem of sustaining a public resource that everybody is free to overuse-the 'tragedy of the commons'-emerges in many social dilemmas, such as our inability to sustain the global climate. Public goods experiments, which are used to study this type of problem, usually confirm that the collective benefit will not be produced. Because individuals and countries often participate in several social games simultaneously, the interaction of these games may provide a sophisticated way by which to maintain the public resource. Indirect reciprocity, 'give and you shall receive', is built on reputation and can sustain a high level of cooperation, as shown by game theorists. Here we show, through alternating rounds of public goods and indirect reciprocity games, that the need to maintain reputation for indirect reciprocity maintains contributions to the public good at an unexpectedly high level. But if rounds of indirect reciprocation are not expected, then contributions to the public good drop quickly to zero. Alternating the games leads to higher profits for all players. As reputation may be a currency that is valid in many social games, our approach could be used to test social dilemmas for their solubility.