Oxytocin and cooperation under conditions of uncertainty: The modulating role of incentives and social information

The neuropeptide Oxytocin (OT) has been implicated in many aspects of mammalian social behavior. This study investigates how OT interacts with two well-studied determinants of cooperative behavior: incentives and social information. Participants received OT or a placebo and played two economic games: a Coordination Game (with strong incentives to cooperate) and a Prisoner's Dilemma (with weak cooperative incentives). OT enhanced cooperation only when social information was present, and this effect was significantly more pronounced in the Coordination Game. When social information was lacking, OT surprisingly decreased cooperation. Consistent with the well-established role of OT in trust-building and in social cognition, social information appears to be crucial for OT to boost cooperative expectations in an interdependent social interaction that provides incentives to cooperate. When these cues are absent, OT appears to instead elicit a risk-averse strategy.     

A functional MRI study on how oxytocin affects decision making in social dilemmas: Cooperate as long as it pays off,aggress only when you think you can win

We investigate if the neuropeptide oxytocin (OT), known to moderate social behaviour, influences strategic decision making in social dilemmas by facilitating the integration of incentives and social cues. Participants (N = 29) played two economic games with different incentive structures in the fMRI scanner after receiving OT or placebo (following a double blind, within-subject design). Pictures of angry or neutral faces (the social cues) were displayed alongside the game matrices. Consistent with a priori hypotheses based on the modulatory role of OT in mesolimbic dopaminergic brain regions, the results indicate that, compared to placebo, OT significantly increases the activation of the nucleus accumbens during an assurance (coordination) game that rewards mutual cooperation. This increases appetitive motivation so that cooperative behaviour is facilitated for risk averse individuals. OT also significantly attenuates the amygdala, thereby reducing the orienting response to social cues. The corresponding change in behaviour is only apparent in the chicken (or anti-coordination) game, where aggression is incentivized but fatal if the partner also aggresses. Because of this ambiguity, decision making can be improved by additional information, and OT steers decisions in the chicken game in accordance with the valence of the facial cue: aggress when face is neutral; retreat when it is angry. Through its combined influence on amygdala and nucleus accumbens, OT improves the selection of a cooperative or aggressive strategy in function of the best match between the incentives of the game and the social cues present in the decision environment.     

False friends are worse than bitter enemies: “Altruistic” punishment of in-group members

One of the most critical features of human society is the pervasiveness of cooperation in social and economic exchanges. Moreover, social scientists have found overwhelming evidence that such cooperative behavior is likely to be directed toward in-group members. We propose that the group-based nature of cooperation includes punishment behavior. Punishment behavior is used to maintain cooperation within systems of social exchange and, thus, is directed towards members of an exchange system. Because social exchanges often take place within groups, we predict that punishment behavior is used to maintain cooperation in the punisher's group. Specifically, punishment behavior is directed toward in-group members who are found to be noncooperators. To examine this, we conducted a gift-giving game experiment with third-party punishment. The results of the experiment (N=90) support the following hypothesis: Participants who are cooperative in a gift-giving game punish noncooperative in-group members more severely than they punish noncooperative out-group members.     

Societal background influences social learning in cooperative decision making

Humans owe their ecological success to their great capacities for social learning and cooperation: learning from others helps individuals adjust to their environment and can promote cooperation in groups. Classic and recent studies indicate that the cultural organization of societies shapes the influence of social information on decision making and suggest that collectivist values (prioritizing the group relative to the individual) increase tendencies to conform to the majority. However, it is unknown whether and how societal background impacts social learning in cooperative interactions. Here we show that social learning in cooperative decision making systematically varies across two societies. We experimentally compare people's basic propensities for social learning in samples from a collectivist (China) and an individualist society (United Kingdom; total n?=?540) in a social dilemma and a coordination game. We demonstrate that Chinese participants base their cooperation decisions on information about their peers much more frequently than their British counterparts. Moreover, our results reveal remarkable societal differences in the type of peer information people consider. In contrast to the consensus view, Chinese participants tend to be substantially less majority-oriented than the British. While Chinese participants are inclined to adopt peer behavior that leads to higher payoffs, British participants tend to cooperate only if sufficiently many peers do so too. These results indicate that the basic processes underlying social transmission are not universal; rather, they vary with cultural conditions. As success-based learning is associated with selfish behavior and majority-based learning can help foster cooperation, our study suggests that in different societies social learning can play diverging roles in the emergence and maintenance of cooperation.     

Human uniqueness-self-interest and social cooperation

Humans are unique among all species of terrestrial history in both ecological dominance and individual properties. Many, or perhaps all, of the unique elements of this nonpareil status can be plausibly interpreted as evolutionary and strategic elements and consequences of the unprecedented intensity and scale of our social cooperation. Convincing explanation of this unique human social adaptation remains a central, unmet challenge to the scientific enterprise.We develop a hypothesis for the ancestral origin of expanded cooperative social behavior. Specifically, we present a game theoretic analysis demonstrating that a specific pattern of expanded social cooperation between conspecific individuals with conflicts of interest (including non-kin) can be strategically viable, but only in animals that possess a highly unusual capacity for conspecific violence (credible threat) having very specific properties that dramatically reduce the costs of coercive violence. The resulting reduced costs allow preemptive or compensated coercion to be an instantaneously self-interested behavior under diverse circumstances rather than in rare, idiosyncratic circumstances as in actors (animals) who do not have access to inexpensive coercive threat.Humans are apparently unique among terrestrial organisms in having evolved conspecific coercive capabilities that fulfill these stringent requirements. Thus, our results support the proposal that access to a novel capacity for projection of coercive threat might represent the essential initiating event for the evolution of a human-like pattern of social cooperation and the subsequent evolution of the diverse features of human uniqueness. Empirical evidence indicates that these constraints were, in fact, met only in our evolutionary lineage. The logic for the emergence of uniquely human cooperation suggested by our analysis apparently accounts simply for the human fossil record.     

Social exchange and reciprocity: confusion or a heuristic?

We propose that a “social exchange heuristic” is as important as the cheater detection mechanism for attaining mutual cooperation in social exchange. The social exchange heuristic prompts people to perceive a mixed-motive situation, such as the Prisoner's Dilemma (PD), as an Assurance Game (AG) situation in which cooperation is a personally better choice than defection insofar as the partner is cooperating as well. We demonstrate the operation of the social exchange heuristic through a comparison of the ordinary one-shot, simultaneous PD with the one-shot, sequential PD. Participants in the current experiments, involving a total of 261 volunteers, committed a logical error in the direction of favoring mutual cooperation as the situation involved more serious consequences. This result strongly suggests the operation of a domain specific “bias” that encourages pursuit of mutual cooperation in social exchange.     

Lattice models in ecology and social sciences

Random matching between individuals, or the complete-mixing model, is often assumed in analyzing evolutionary or population dynamics in ecology and game theory or other models in social sciences. Making and analyzing a model is not difficult under this simple assumption. However spatial- or network-structured populations, including the lattice model and the power-law network, are more realistic for many ecological and social phenomena than the complete-mixing model. In this review, I will show first that a lattice model can be useful in investigating the effect of neighborhood interactions on the dynamics, not only of plants and forests, but also of animal and human societies. Second, the lattice model promotes the evolution of spiteful behavior, even though it is well-known that the lattice model promotes the evolution of cooperation. Finally, different social networks result in traits, such as social norms, spreading at different speeds.     

Coveting thy neighbors fitness as a means to resolve social dilemmas

In spatial evolutionary games the fitness of each individual is traditionally determined by the payoffs it obtains upon playing the game with its neighbors. Since defection yields the highest individual benefits, the outlook for cooperators is gloomy. While network reciprocity promotes collaborative efforts, chances of averting the impending social decline are slim if the temptation to defect is strong. It is, therefore, of interest to identify viable mechanisms that provide additional support for the evolution of cooperation. Inspired by the fact that the environment may be just as important as inheritance for individual development, we introduce a simple switch that allows a player to either keep its original payoff or use the average payoff of all its neighbors. Depending on which payoff is higher, the influence of either option can be tuned by means of a single parameter. We show that, in general, taking into account the environment promotes cooperation. Yet coveting the fitness of one's neighbors too strongly is not optimal. In fact, cooperation thrives best only if the influence of payoffs obtained in the traditional way is equal to that of the average payoff of the neighborhood. We present results for the prisoner's dilemma and the snowdrift game, for different levels of uncertainty governing the strategy adoption process, and for different neighborhood sizes. Our approach outlines a viable route to increased levels of cooperative behavior in structured populations, but one that requires a thoughtful implementation.     

Synergy and discounting of cooperation in social dilemmas

The emergence and maintenance of cooperation by natural selection is an enduring conundrum in evolutionary biology, which has been studied using a variety of game theoretical models inspired by different biological situations. The most widely studied games are the Prisoner's Dilemma, the Snowdrift game and by-product mutualism for pairwise interactions, as well as Public Goods games in larger groups of interacting individuals. Here, we present a general framework for cooperation in social dilemmas in which all the traditional scenarios can be recovered as special cases. In social dilemmas, cooperators provide a benefit to the group at some cost, while defectors exploit the group by reaping the benefits without bearing the costs of cooperation. Using the concepts of discounting and synergy for describing how benefits accumulate when more than one cooperator is present in a group of interacting individuals, we recover the four basic scenarios of evolutionary dynamics given by (i) dominating defection, (ii) coexistence of defectors and cooperators, (iii) dominating cooperation and (iv) bi-stability, in which cooperators and defectors cannot invade each other. Generically, for groups of three or more interacting individuals further, more complex, dynamics can occur. Our framework provides the first unifying approach to model cooperation in different kinds of social dilemmas.     

Spatial effects in social dilemmas

Social dilemmas and the evolutionary conundrum of cooperation are traditionally studied through various kinds of game theoretical models such as the prisoner's dilemma, public goods games, snowdrift games or by-product mutualism. All of them exemplify situations which are characterized by different degrees of conflicting interests between the individuals and the community. In groups of interacting individuals, cooperators produce a common good benefitting the entire group at some cost to themselves, whereas defectors attempt to exploit the resource by avoiding the costly contributions. Based on synergistic or discounted accumulation of cooperative benefits a unifying theoretical framework was recently introduced that encompasses all games that have traditionally been studied separately (Hauert, Michor, Nowak, Doebeli, 2005. Synergy and discounting of cooperation in social dilemmas. J. Theor. Biol., in press.). Within this framework we investigate the effects of spatial structure with limited local interactions on the evolutionary fate of cooperators and defectors. The quantitative effects of space turn out to be quite sensitive to the underlying microscopic update mechanisms but, more general, we demonstrate that in prisoner's dilemma type interactions spatial structure benefits cooperation-although the parameter range is quite limited-whereas in snowdrift type interactions spatial structure may be beneficial too, but often turns out to be detrimental to cooperation.     

Motion-guided attention promotes adaptive communications during social navigation

Animals are capable of enhanced decision making through cooperation, whereby accurate decisions can occur quickly through decentralized consensus. These interactions often depend upon reliable social cues, which can result in highly coordinated activities in uncertain environments. Yet information within a crowd may be lost in translation, generating confusion and enhancing individual risk. As quantitative data detailing animal social interactions accumulate, the mechanisms enabling individuals to rapidly and accurately process competing social cues remain unresolved. Here, we model how motion-guided attention influences the exchange of visual information during social navigation. We also compare the performance of this mechanism to the hypothesis that robust social coordination requires individuals to numerically limit their attention to a set of n-nearest neighbours. While we find that such numerically limited attention does not generate robust social navigation across ecological contexts, several notable qualities arise from selective attention to motion cues. First, individuals can instantly become a local information hub when startled into action, without requiring changes in neighbour attention level. Second, individuals can circumvent speed–accuracy trade-offs by tuning their motion thresholds. In turn, these properties enable groups to collectively dampen or amplify social information. Lastly, the minority required to sway a group''s short-term directional decisions can change substantially with social context. Our findings suggest that motion-guided attention is a fundamental and efficient mechanism underlying collaborative decision making during social navigation.     

Different gain/loss sensitivity and social adaptation ability in gifted adolescents during a public goods game

Gifted adolescents are considered to have high IQs with advanced mathematical and logical performances, but are often thought to suffer from social isolation or emotional mal-adaptation to the social group. The underlying mechanisms that cause stereotypic portrayals of gifted adolescents are not well known. We aimed to investigate behavioral performance of gifted adolescents during social decision-making tasks to assess their affective and social/non-social cognitive abilities. We examined cooperation behaviors of 22 gifted and 26 average adolescents during an iterative binary public goods (PG) game, a multi-player social interaction game, and analyzed strategic decision processes that include cooperation and free-riding. We found that the gifted adolescents were more cooperative than average adolescents. Particularly, comparing the strategies for the PG game between the two groups, gifted adolescents were less sensitive to loss, yet were more sensitive to gain. Additionally, the behavioral characteristics of average adolescents, such as low trust of the group and herding behavior, were not found in gifted adolescents. These results imply that gifted adolescents have a high cognitive ability but a low ability to process affective information or to adapt in social groups compared with average adolescents. We conclude that gain/loss sensitivity and the ability to adapt in social groups develop to different degrees in average and gifted adolescents.     

The Effect of Incentives and Meta-incentives on the Evolution of Cooperation

Although positive incentives for cooperators and/or negative incentives for free-riders in social dilemmas play an important role in maintaining cooperation, there is still the outstanding issue of who should pay the cost of incentives. The second-order free-rider problem, in which players who do not provide the incentives dominate in a game, is a well-known academic challenge. In order to meet this challenge, we devise and analyze a meta-incentive game that integrates positive incentives (rewards) and negative incentives (punishments) with second-order incentives, which are incentives for other players’ incentives. The critical assumption of our model is that players who tend to provide incentives to other players for their cooperative or non-cooperative behavior also tend to provide incentives to their incentive behaviors. In this paper, we solve the replicator dynamics for a simple version of the game and analytically categorize the game types into four groups. We find that the second-order free-rider problem is completely resolved without any third-order or higher (meta) incentive under the assumption. To do so, a second-order costly incentive, which is given individually (peer-to-peer) after playing donation games, is needed. The paper concludes that (1) second-order incentives for first-order reward are necessary for cooperative regimes, (2) a system without first-order rewards cannot maintain a cooperative regime, (3) a system with first-order rewards and no incentives for rewards is the worst because it never reaches cooperation, and (4) a system with rewards for incentives is more likely to be a cooperative regime than a system with punishments for incentives when the cost-effect ratio of incentives is sufficiently large. This solution is general and strong in the sense that the game does not need any centralized institution or proactive system for incentives.     

Cooperative coordination as a social behavior

Coordinating behavior is widespread in contexts that include courtship, aggression, and cooperation for shared outcomes. The social significance of cooperative coordination (CC) is usually downplayed by learning theorists, evolutionary biologists, and game theorists in favor of an individual behavior → outcome perspective predicated on maximizing payoffs for all participants. To more closely model CC as it occurs under free-ranging conditions, pairs of rats were rewarded for coordinated shuttling within a shared chamber with unrestricted social interaction. Results show that animals learned to work together with sensitivity to the task and type of partner. Moreover, social interaction and coordination influenced both consumption of the reward solution immediately following a session and preference for cooperation, suggesting that affective states and incentives related to cooperation extend beyond the outcomes obtained. These results support field studies by showing not only how cooperation is performed but also the importance of considering how the behavior of cooperating affects outcomes and preference for cooperating. This article and some of the research described here were supported in part by Grant 96-00293 from the United States-Israel Binational Science Foundation to the author and Peter R. Killeen of Arizona State University. Richard Schuster (Ph.D. in Psychology, Harvard University 1968) has been a Lecturer in Psychology at the University of Zambia (1971–1977), with field research on antelopes, and a Lecturer at the Department of Psychology, University of Haifa, Israel (1977-present). His main research focus is on laboratory models of social behavior.     

The evolution of altruistic social preferences in human groups

In this paper, we consider three hypotheses to account for the evolution of the extraordinary capacity for large-scale cooperation and altruistic social preferences within human societies. One hypothesis is that human cooperation is built on the same evolutionary foundations as cooperation in other animal societies, and that fundamental elements of the social preferences that shape our species'' cooperative behaviour are also shared with other closely related primates. Another hypothesis is that selective pressures favouring cooperative breeding have shaped the capacity for cooperation and the development of social preferences, and produced a common set of behavioural dispositions and social preferences in cooperatively breeding primates and humans. The third hypothesis is that humans have evolved derived capacities for collaboration, group-level cooperation and altruistic social preferences that are linked to our capacity for culture. We draw on naturalistic data to assess differences in the form, scope and scale of cooperation between humans and other primates, experimental data to evaluate the nature of social preferences across primate species, and comparative analyses to evaluate the evolutionary origins of cooperative breeding and related forms of behaviour.     

On the social nature of eyes: The effect of social cues in interaction and individual choice tasks

In an experimental setting, we applied a dual strategy to better understand the effect of pictures of eyes on human behavior. First, we investigated whether the effect of eyes was limited to interaction tasks in which the subjects' decisions influenced the outcomes of other subjects. We expanded the range of tasks to include individual choice tasks in which the subjects' decisions only influenced their own outcomes. Second, we investigated whether pictures of eyes were one of many social cues or were unique in their effect. We compared the effect of pictures of eyes with the effect of a different condition in which we presented the subjects with pictures of other students (peers). Our results suggest that the effect of pictures of eyes is limited to interaction tasks and that eyes should be considered distinct from other social cues, such as reminders of peers. While pictures of eyes uniformly enhanced pro-social behavior in interaction tasks, this was not the case for reminders of peers. Furthermore, the reminders of peers led to more rational behavior in individual choice tasks, whereas the effect of pictures of eyes was limited to situations involving interaction. Combined, these findings are in line with the claim that the effect of pictures of eyes on behavior is caused by a social exchange heuristic that works to enhance mutual cooperative behavior.     

Long-term social bonds promote cooperation in the iterated Prisoner's Dilemma

Reciprocal altruism, one of the most probable explanations for cooperation among non-kin, has been modelled as a Prisoner''s Dilemma. According to this game, cooperation could evolve when individuals, who expect to play again, use conditional strategies like tit-for-tat or Pavlov. There is evidence that humans use such strategies to achieve mutual cooperation, but most controlled experiments with non-human animals have failed to find cooperation. One reason for this could be that subjects fail to cooperate because they behave as if they were to play only once. To assess this hypothesis, we conducted an experiment with monogamous zebra finches (Taeniopygia guttata) that were tested in a two-choice apparatus, with either their social partner or an experimental opponent of the opposite sex. We found that zebra finches maintained high levels of cooperation in an iterated Prisoner''s Dilemma game only when interacting with their social partner. Although other mechanisms may have contributed to the observed difference between the two treatments, our results support the hypothesis that animals do not systematically give in to the short-term temptation of cheating when long-term benefits exist. Thus, our findings contradict the commonly accepted idea that reciprocal altruism will be rare in non-human animals.     

Mutual trust and cooperation in the evolutionary hawks–doves game

Using a new dynamical network model of society in which pairwise interactions are weighted according to mutual satisfaction, we show that cooperation is the norm in the hawks–doves game when individuals are allowed to break ties with undesirable neighbors and to make new acquaintances in their extended neighborhood. Moreover, cooperation is robust with respect to rather strong strategy perturbations. We also discuss the empirical structure of the emerging networks, and the reasons that allow cooperators to thrive in the population. Given the metaphorical importance of this game for social interaction, this is an encouraging positive result as standard theory for large mixing populations prescribes that a certain fraction of defectors must always exist at equilibrium.     

Evolution of cooperation with shared costs and benefits

The quest to determine how cooperation evolves can be based on evolutionary game theory, in spite of the fact that evolutionarily stable strategies (ESS) for most non-zero-sum games are not cooperative. We analyse the evolution of cooperation for a family of evolutionary games involving shared costs and benefits with a continuum of strategies from non-cooperation to total cooperation. This cost-benefit game allows the cooperator to share in the benefit of a cooperative act, and the recipient to be burdened with a share of the cooperator's cost. The cost-benefit game encompasses the Prisoner's Dilemma, Snowdrift game and Partial Altruism. The models produce ESS solutions of total cooperation, partial cooperation, non-cooperation and coexistence between cooperation and non-cooperation. Cooperation emerges from an interplay between the nonlinearities in the cost and benefit functions. If benefits increase at a decelerating rate and costs increase at an accelerating rate with the degree of cooperation, then the ESS has an intermediate level of cooperation. The game also exhibits non-ESS points such as unstable minima, convergent-stable minima and unstable maxima. The emergence of cooperative behaviour in this game represents enlightened self-interest, whereas non-cooperative solutions illustrate the Tragedy of the Commons. Games having either a stable maximum or a stable minimum have the property that small changes in the incentive structure (model parameter values) or culture (starting frequencies of strategies) result in correspondingly small changes in the degree of cooperation. Conversely, with unstable maxima or unstable minima, small changes in the incentive structure or culture can result in a switch from non-cooperation to total cooperation (and vice versa). These solutions identify when human or animal societies have the potential for cooperation and whether cooperation is robust or fragile.     

Spatial dynamics,social norms,and the opportunity of the commons

The most important message of Levin (Ecol Res 21:328–333, 2006) is that “Ecologists and economists have much incentive for interaction.” Recent studies that account for evolutionary processes and local interactions support this view by obtaining results that run counter to conventional wisdom within resource economics. A second major message of the article is that to meet environmental challenges, humanity must develop social norms that enhance cooperative responses. Successful examples of resource management systems back up norms with economic incentives: rewards for good behavior and punishments for bad. Economic incentives are especially important if rapid and large changes in human behavior are desired.