An Evolutionary Model of Cooperation,Fairness and Altruistic Punishment in Public Good Games

We identify and explain the mechanisms that account for the emergence of fairness preferences and altruistic punishment in voluntary contribution mechanisms by combining an evolutionary perspective together with an expected utility model. We aim at filling a gap between the literature on the theory of evolution applied to cooperation and punishment, and the empirical findings from experimental economics. The approach is motivated by previous findings on other-regarding behavior, the co-evolution of culture, genes and social norms, as well as bounded rationality. Our first result reveals the emergence of two distinct evolutionary regimes that force agents to converge either to a defection state or to a state of coordination, depending on the predominant set of self- or other-regarding preferences. Our second result indicates that subjects in laboratory experiments of public goods games with punishment coordinate and punish defectors as a result of an aversion against disadvantageous inequitable outcomes. Our third finding identifies disadvantageous inequity aversion as evolutionary dominant and stable in a heterogeneous population of agents endowed initially only with purely self-regarding preferences. We validate our model using previously obtained results from three independently conducted experiments of public goods games with punishment.     

Social Influences on Inequity Aversion in Children

Adults and children are willing to sacrifice personal gain to avoid both disadvantageous and advantageous inequity. These two forms of inequity aversion follow different developmental trajectories, with disadvantageous inequity aversion emerging around 4 years and advantageous inequity aversion emerging around 8 years. Although inequity aversion is assumed to be specific to situations where resources are distributed among individuals, the role of social context has not been tested in children. Here, we investigated the influence of two aspects of social context on inequity aversion in 4- to 9-year-old children: (1) the role of the experimenter distributing rewards and (2) the presence of a peer with whom rewards could be shared. Experiment 1 showed that children rejected inequity at the same rate, regardless of whether the experimenter had control over reward allocations. This indicates that children’s decisions are based upon reward allocations between themselves and a peer and are not attempts to elicit more favorable distributions from the experimenter. Experiment 2 compared rejections of unequal reward allocations in children interacting with or without a peer partner. When faced with a disadvantageous distribution, children frequently rejected a smaller reward when a larger reward was visible, even if no partner would obtain the larger reward. This suggests that nonsocial factors partly explain disadvantageous inequity rejections. However, rejections of disadvantageous distributions were higher when the larger amount would go to a peer, indicating that social context enhances disadvantageous inequity aversion. By contrast, children rejected advantageous distributions almost exclusively in the social context. Therefore, advantageous inequity aversion appears to be genuinely social, highlighting its potential relevance for the development of fairness concerns. By comparing social and nonsocial factors, this study provides a detailed picture of the expression of inequity aversion in human ontogeny and raises questions about the function and evolution of inequity aversion in humans.     

Beneficial betrayal aversion

Many studies demonstrate the social benefits of cooperation. Likewise, recent studies convincingly demonstrate that betrayal aversion hinders trust and discourages cooperation. In this respect, betrayal aversion is unlike socially "beneficial" preferences including altruism, fairness and inequity aversion, all of which encourage cooperation and exchange. To our knowledge, other than the suggestion that it acts as a barrier to rash trust decisions, the benefits of betrayal aversion remain largely unexplored. Here we use laboratory experiments with human participants to show that groups including betrayal-averse agents achieve higher levels of reciprocity and more profitable social exchange than groups lacking betrayal aversion. These results are the first rigorous evidence on the benefits of betrayal aversion, and may help future research investigating cultural differences in betrayal aversion as well as future research on the evolutionary roots of betrayal aversion. Further, our results extend the understanding of how intentions affect social interactions and exchange and provide an effective platform for further research on betrayal aversion and its effects on human behavior.     

Fairness expectations and altruistic sharing in 15-month-old human infants

Human cooperation is a key driving force behind the evolutionary success of our hominin lineage. At the proximate level, biologists and social scientists have identified other-regarding preferences--such as fairness based on egalitarian motives, and altruism--as likely candidates for fostering large-scale cooperation. A critical question concerns the ontogenetic origins of these constituents of cooperative behavior, as well as whether they emerge independently or in an interrelated fashion. The answer to this question will shed light on the interdisciplinary debate regarding the significance of such preferences for explaining how humans become such cooperative beings. We investigated 15-month-old infants' sensitivity to fairness, and their altruistic behavior, assessed via infants' reactions to a third-party resource distribution task, and via a sharing task. Our results challenge current models of the development of fairness and altruism in two ways. First, in contrast to past work suggesting that fairness and altruism may not emerge until early to mid-childhood, 15-month-old infants are sensitive to fairness and can engage in altruistic sharing. Second, infants' degree of sensitivity to fairness as a third-party observer was related to whether they shared toys altruistically or selfishly, indicating that moral evaluations and prosocial behavior are heavily interconnected from early in development. Our results present the first evidence that the roots of a basic sense of fairness and altruism can be found in infancy, and that these other-regarding preferences develop in a parallel and interwoven fashion. These findings support arguments for an evolutionary basis--most likely in dialectical manner including both biological and cultural mechanisms--of human egalitarianism given the rapidly developing nature of other-regarding preferences and their role in the evolution of human-specific forms of cooperation. Future work of this kind will help determine to what extent uniquely human sociality and morality depend on other-regarding preferences emerging early in life.     

Competing demands of prosociality and equity in monkeys

Prosocial decisions may lead to unequal payoffs among group members. Although an aversion to inequity has been found in empirical studies of both human and nonhuman primates, the contexts previously studied typically do not involve a trade-off between prosociality and inequity. Here we investigate the apparent coexistence of these two factors, specifically the competing demands of prosociality and equity. We directly compare the responses of brown capuchin monkeys (Cebus apella) among situations where prosocial preferences conflict with equality, using a paradigm comparable to other studies of cooperation and inequity in this species. By choosing to pull a tray towards themselves, subjects rewarded themselves and/or another in conditions in which the partner either received the same or different rewards, or the subject received no reward. In unequal payoff conditions, subjects could obtain equality by choosing not to pull in the tray, so that neither individual was rewarded. The monkeys showed prosocial preferences even in situations of moderate disadvantageous inequity, preferring to pull in the tray more often when a partner was present than absent. However, when the discrepancy between rewards increased, prosocial behavior ceased.     

Human punishment is motivated by inequity aversion, not a desire for reciprocity

Humans involved in cooperative interactions willingly pay a cost to punish cheats. However, the proximate motives underpinning punitive behaviour are currently debated. Individuals who interact with cheats experience losses, but they also experience lower payoffs than the cheating partner. Thus, the negative emotions that trigger punishment may stem from a desire to reciprocate losses or from inequity aversion. Previous studies have not disentangled these possibilities. Here, we use an experimental approach to ask whether punishment is motivated by inequity aversion or by a desire for reciprocity. We show that humans punish cheats only when cheating produces disadvantageous inequity, while there is no evidence for reciprocity. This finding challenges the notion that punishment is motivated by a simple desire to reciprocally harm cheats and shows that victims compare their own payoffs with those of partners when making punishment decisions.     

Understanding Recurrent Crime as System-Immanent Collective Behavior

Containing the spreading of crime is a major challenge for society. Yet, since thousands of years, no effective strategy has been found to overcome crime. To the contrary, empirical evidence shows that crime is recurrent, a fact that is not captured well by rational choice theories of crime. According to these, strong enough punishment should prevent crime from happening. To gain a better understanding of the relationship between crime and punishment, we consider that the latter requires prior discovery of illicit behavior and study a spatial version of the inspection game. Simulations reveal the spontaneous emergence of cyclic dominance between “criminals”, “inspectors”, and “ordinary people” as a consequence of spatial interactions. Such cycles dominate the evolutionary process, in particular when the temptation to commit crime or the cost of inspection are low or moderate. Yet, there are also critical parameter values beyond which cycles cease to exist and the population is dominated either by a stable mixture of criminals and inspectors or one of these two strategies alone. Both continuous and discontinuous phase transitions to different final states are possible, indicating that successful strategies to contain crime can be very much counter-intuitive and complex. Our results demonstrate that spatial interactions are crucial for the evolutionary outcome of the inspection game, and they also reveal why criminal behavior is likely to be recurrent rather than evolving towards an equilibrium with monotonous parameter dependencies.     

Attending to the outcome of others: disadvantageous inequity aversion in male capuchin monkeys (Cebus apella)

Brosnan and de Waal [Nature 425:297-299, 2003] reported that capuchin monkeys responded negatively to unequal reward distributions between themselves and another individual when comparing their own rewards with that of their partner. It was suggested that social emotions provided the underlying motivation for such behavior and that this inequity aversion is specific to the social domain. However, alternative hypotheses such as the "frustration effect" or the "food expectation hypothesis" may provide more parsimonious explanations for Brosnan and de Waal's [Nature 425:297-299] results, while others have argued that these findings are not congruent with the Fehr-Schmidt inequity aversion model cited by the authors. The claim that inequity aversion behavior is specific to the social domain has also been questioned, as primates also develop expectations about rewards in the absence of partners, and react negatively when those expectations are violated. In this study, a modified Dictator game was used to investigate whether capuchins would exhibit either disadvantageous inequity aversion behavior or reference-dependent expectancy violation in social and nonsocial conditions, respectively. When given the choice between an equitable and an inequitable outcome, the subjects showed disadvantageous inequity aversion behavior, choosing the equitable outcome significantly more in the social condition. In the nonsocial condition, however, subjects did not show negative expectancy violation resulting from the formation of reference-dependent expectations, choosing the equitable outcome at chance levels. These results suggest that capuchins attend to differential payoffs and that they are averse to inequity, which is disadvantageous to themselves.     

Morals Matter in Economic Games

Contrary to predictions from Expected Utility Theory and Game Theory, when making economic decisions in interpersonal situations, people take the interest of others into account and express various forms of solidarity, even in one-shot interactions with anonymous strangers. Research in other-regarding behavior is dominated by behavioral economical and evolutionary biological approaches. Psychological theory building, which addresses mental processes underlying other-regarding behavior, is rare. Based on Relational Models Theory (RMT, [1]) and Relationship Regulation Theory (RRT, [2]) it is proposed that moral motives influence individuals’ decision behavior in interpersonal situations via conscious and unconscious (automatic) processes. To test our propositions we developed the ‘Dyadic Solidarity Game’ and its solitary equivalent, the ‘Self-Insurance Game’. Four experiments, in which the moral motives “Unity” and “Proportionality” were manipulated, support the propositions made. First, it was shown that consciously activated moral motives (via framing of the overall goal of the experiment) and unconsciously activated moral motives (via subliminal priming) influence other-regarding behavior. Second, this influence was only found in interpersonal, not in solitary situations. Third, by combining the analyses of the two experimental games the extent to which participants apply the Golden Rule (“treat others how you wish to be treated”) could be established. Individuals with a “Unity” motive treated others like themselves, whereas individuals with a “Proportionality” motive gave others less then they gave themselves. The four experiments not only support the assumption that morals matter in economic games, they also deliver new insights in how morals matter in economic decision making.     

Evolutionary Establishment of Moral and Double Moral Standards through Spatial Interactions

Situations where individuals have to contribute to joint efforts or share scarce resources are ubiquitous. Yet, without proper mechanisms to ensure cooperation, the evolutionary pressure to maximize individual success tends to create a tragedy of the commons (such as over-fishing or the destruction of our environment). This contribution addresses a number of related puzzles of human behavior with an evolutionary game theoretical approach as it has been successfully used to explain the behavior of other biological species many times, from bacteria to vertebrates. Our agent-based model distinguishes individuals applying four different behavioral strategies: non-cooperative individuals (“defectors”), cooperative individuals abstaining from punishment efforts (called “cooperators” or “second-order free-riders”), cooperators who punish non-cooperative behavior (“moralists”), and defectors, who punish other defectors despite being non-cooperative themselves (“immoralists”). By considering spatial interactions with neighboring individuals, our model reveals several interesting effects: First, moralists can fully eliminate cooperators. This spreading of punishing behavior requires a segregation of behavioral strategies and solves the “second-order free-rider problem”. Second, the system behavior changes its character significantly even after very long times (“who laughs last laughs best effect”). Third, the presence of a number of defectors can largely accelerate the victory of moralists over non-punishing cooperators. Fourth, in order to succeed, moralists may profit from immoralists in a way that appears like an “unholy collaboration”. Our findings suggest that the consideration of punishment strategies allows one to understand the establishment and spreading of “moral behavior” by means of game-theoretical concepts. This demonstrates that quantitative biological modeling approaches are powerful even in domains that have been addressed with non-mathematical concepts so far. The complex dynamics of certain social behaviors become understandable as the result of an evolutionary competition between different behavioral strategies.     

Neural Correlates of Advantageous and Disadvantageous Inequity in Sharing Decisions

Humans have a strong preference for fair distributions of resources. Neuroimaging studies have shown that being treated unfairly coincides with activation in brain regions involved in signaling conflict and negative affect. Less is known about neural responses involved in violating a fairness norm ourselves. Here, we investigated the neural patterns associated with inequity, where participants were asked to choose between an equal split of money and an unequal split that could either maximize their own (advantageous inequity) or another person’s (disadvantageous inequity) earnings. Choosing to divide money unequally, irrespective who benefited from the unequal distribution, was associated with activity in the dorsal anterior cingulate cortex, anterior insula and the dorsolateral prefrontal cortex. Inequity choices that maximized another person’s profits were further associated with activity in the ventral striatum and ventromedial prefrontal cortex. Taken together, our findings show evidence of a common neural pattern associated with both advantageous and disadvantageous inequity in sharing decisions and additional recruitment of neural circuitry previously linked to the computation of subjective value and reward when violating a fairness norm at the benefit of someone else.     

Punishment is strongly motivated by revenge and weakly motivated by inequity aversion

There are two broad functional explanations for second-party punishment: fitness-leveling and deterrence. The former suggests that people punish to reduce fitness differences, while the latter suggests that people punish in order to reciprocate losses and deter others from inflicting losses on them in the future. We explore the relative roles of these motivations using a pre-registered, two-player experiment with 2426 US participants from Amazon Mechanical Turk. Participants played as the “responder” and were assigned to either a Take or Augment condition. In the Take condition, the “partner” could steal money from the responder's bonus or do nothing. In the Augment condition, the partner could augment the responder's bonus by giving them money at no cost to themselves or do nothing. We also manipulated the responders' starting endowments, such that after the partner's decision, responders experienced different payoff outcomes: advantageous inequity, equality, or varying degrees of disadvantageous inequity. Responders then decided whether to pay a cost to punish the partner. Punishment was clearly influenced by theft and was most frequent when theft resulted in disadvantageous inequity. However, people also punished in the absence of theft, particularly when confronted with disadvantageous inequity. While the effect of inequity on punishment was small, our results suggest that punishment is motivated by more than just the desire to reciprocate losses. These findings highlight the multiple motivations undergirding punishment and bear directly on functional explanations for the existence of punishment in human societies.     

Gene-culture coevolution and the nature of human sociality

Human characteristics are the product of gene-culture coevolution, which is an evolutionary dynamic involving the interaction of genes and culture over long time periods. Gene-culture coevolution is a special case of niche construction. Gene-culture coevolution is responsible for human other-regarding preferences, a taste for fairness, the capacity to empathize and salience of morality and character virtues.     

Patient and impatient punishers of free-riders

Costly punishment of cheaters who contribute little or nothing to a cooperating group has been extensively studied, as an effective means to enforce cooperation. The prevailing view is that individuals use punishment to retaliate against transgressions of moral standards such as fairness or equity. However, there is much debate regarding the psychological underpinnings of costly punishment. Some authors suggest that costly punishment must be a product of humans'' capacity for reasoning, self-control and long-term planning, whereas others argue that it is the result of an impulsive, present-oriented emotional drive. Here, we explore the inter-temporal preferences of punishers in a multilateral cooperation game and show that both interpretations might be right, as we can identify two different types of punishment: punishment of free-riders by cooperators, which is predicted by patience (future orientation); and free-riders'' punishment of other free-riders, which is predicted by impatience (present orientation). Therefore, the picture is more complex as punishment by free-riders probably comes not from a reaction against a moral transgression, but instead from a competitive, spiteful drive. Thus, punishment grounded on morals may be related to lasting or delayed psychological incentives, whereas punishment triggered by competitive desires may be linked to short-run aspirations. These results indicate that the individual''s time horizon is relevant for the type of social behaviour she opts for. Integrating such differences in inter-temporal preferences and the social behaviour of agents might help to achieve a better understanding of how human cooperation and punishment behaviour has evolved.     

Antagonistic Parent-Offspring Co-Adaptation

Background

In species across taxa, offspring have means to influence parental investment (PI). PI thus evolves as an interacting phenotype and indirect genetic effects may strongly affect the co-evolutionary dynamics of offspring and parental behaviors. Evolutionary theory focused on explaining how exaggerated offspring solicitation can be understood as resolution of parent-offspring conflict, but the evolutionary origin and diversification of different forms of family interactions remains unclear.

Methodology/Principal Findings

In contrast to previous theory that largely uses a static approach to predict how “offspring individuals” and “parental individuals” should interact given conflict over PI, we present a dynamic theoretical framework of antagonistic selection on the PI individuals obtain/take as offspring and the PI they provide as parents to maximize individual lifetime reproductive success; we analyze a deterministic and a stochastic version of this dynamic framework. We show that a zone for equivalent co-adaptation outcomes exists in which stable levels of PI can evolve and be maintained despite fast strategy transitions and ongoing co-evolutionary dynamics. Under antagonistic co-adaptation, cost-free solicitation can evolve as an adaptation to emerging preferences in parents.

Conclusions/Significance

We show that antagonistic selection across the offspring and parental life-stage of individuals favors co-adapted offspring and parental behavior within a zone of equivalent outcomes. This antagonistic parent-offspring co-adaptation does not require solicitation to be costly, allows for rapid divergence and evolutionary novelty and potentially explains the origin and diversification of the observed provisioning forms in family life.     

Effect of cooperation level of group on punishment for non-cooperators: a functional magnetic resonance imaging study

Sometimes we punish non-cooperators in our society. Such behavior could be derived from aversive emotion for inequity (inequity aversion) to make non-cooperators cooperative. Thus, punishing behavior derived from inequity is believed to be important for maintaining our society. Meanwhile, our daily experiences suggest that the degree of cooperation by the members of society (cooperation level of the group) could change the punishing behavior for non-cooperators even if the inequity were equal. Such effect of the cooperation level of the group cannot be explained by simple inequity aversion. Although punishment-related brain regions have been reported in previous functional magnetic resonance imaging (fMRI) study, little is known about such regions affected by the cooperation level of the group. In the present fMRI study, we investigated the effect of the cooperation level of the group on the punishing behavior for non-cooperators and its related brain activations by a paradigm in which the degree of the cooperative state varied from low to high. Punishment-related activations were observed in brain regions such as the anterior insula, dorsolateral prefrontal cortex (DLPFC), and anterior cingulate cortex (ACC). The quantity of punishment in a high cooperation context was greater than in a low cooperation context, and activation in the right DLPFC and ACC in a high cooperation context showed greater activity than in a low cooperation context. This indicates that the cooperation level of the group, as well as aversive emotion for inequity, is the important factor of punishing behavior.     

Fairness Norms and Theory of Mind in an Ultimatum Game: Judgments,Offers, and Decisions in School-Aged Children

The sensitivity to fairness undergoes relevant changes across development. Whether such changes depend on primary inequity aversion or on sensitivity to a social norm of fairness is still debated. Using a modified version of the Ultimatum Game that creates informational asymmetries between Proposer and Responder, a previous study showed that both perceptions of fairness and fair behavior depend upon normative expectations, i.e., beliefs about what others expect one should do in a specific situation. Individuals tend to comply with the norm when risking sanctions, but disregard the norm when violations are undetectable. Using the same methodology with children aged 8–10 years, the present study shows that children''s beliefs and behaviors differ from what is observed in adults. Playing as Proposers, children show a self-serving bias only when there is a clear informational asymmetry. Playing as Responders, they show a remarkable discrepancy between their normative judgment about fair procedures (a coin toss to determine the offer) and their behavior (rejection of an unfair offer derived from the coin toss), supporting the existence of an outcome bias effect. Finally, our results reveal no influence of theory of mind on children''s decision-making behavior.     

Criminal Defectors Lead to the Emergence of Cooperation in an Experimental,Adversarial Game

While the evolution of cooperation has been widely studied, little attention has been devoted to adversarial settings wherein one actor can directly harm another. Recent theoretical work addresses this issue, introducing an adversarial game in which the emergence of cooperation is heavily reliant on the presence of “Informants,” actors who defect at first-order by harming others, but who cooperate at second-order by punishing other defectors. We experimentally study this adversarial environment in the laboratory with human subjects to test whether Informants are indeed critical for the emergence of cooperation. We find in these experiments that, even more so than predicted by theory, Informants are crucial for the emergence and sustenance of a high cooperation state. A key lesson is that successfully reaching and maintaining a low defection society may require the cultivation of criminals who will also aid in the punishment of others.     

Suffering makes you egoist: acute pain increases acceptance rates and reduces fairness during a bilateral ultimatum game

Social preferences like interpersonal altruism, fairness, reciprocity and inequity aversion are inherently linked to departures from pure self-interest. During economic interactions, for example, defectors may be punished even if this implies a cost for the punishers. This violation of canonical assumptions in economics indicates that socially oriented decisions may predominate over self-centred stances. Here we explore whether the personal experience of pain changes the balance between self-gain and socially based choices. We used laser stimulation to induce pain or a warm sensation in subjects playing a modified version of the Ultimatum Game (UG) both in the role of responder and proposer. After each shot, responders evaluated the fairness of the offer. Moreover, responders and proposers rated the intensity and unpleasantness of the sensation evoked by the laser stimulation. Results show that suffering proposers decrease fair offers and suffering responders increase their acceptance rate irrespective of economic offer. Crucially, the intensity of painful stimulation has a predictive role on Moderately Unfair offers' acceptance rates. Thus the personal experience of pain may favour the emergence of a self-centered perspective aimed at maximizing self-gain. The results suggest that bodily states play a fundamental role in higher-order interpersonal negotiations and interactions.