Learning rules of engagement for social exchange within and between groups.

Globalizing economies and long-distance trade rely on individuals from different cultural groups to negotiate agreement on what to give and take. In such settings, individuals often lack insight into what interaction partners deem fair and appropriate, potentially seeding misunderstandings, frustration, and conflict. Here, we examine how individuals decipher distinct rules of engagement and adapt their behavior to reach agreements with partners from other cultural groups. Modeling individuals as Bayesian learners with inequality aversion reveals that individuals, in repeated ultimatum bargaining with responders sampled from different groups, can be more generous than needed. While this allows them to reach agreements, it also gives rise to biased beliefs about what is required to reach agreement with members from distinct groups. Preregistered behavioral (N = 420) and neuroimaging experiments (N = 49) support model predictions: Seeking equitable agreements can lead to overly generous behavior toward partners from different groups alongside incorrect beliefs about prevailing norms of what is appropriate in groups and cultures other than one's own.

Peace as prerequisite rather than consequence of cooperation.

We take issue with Glowacki's assumption that intergroup relations are characterized by positive-sum interactions and suggest to include negative-sum interactions, and between-group independence. As such, peace may be better defined as the absence of negative-sum interactions. Rather than being a consequence of cooperation, peace emerges as a necessary but not sufficient prerequisite for positive (in)direct reciprocity between groups that, in turn, is key to social identities and cultural complexity.

Distinct neurocognitive pathways underlying creativity: An integrative approach.

By examining the shared neuro-cognitive correlates of curiosity and creativity, we better understand the brain basis of creativity. However, by only examining shared components, important neuro-cognitive correlates are overlooked. Here, we argue that any comprehensive brain model of creativity should consider multiple cognitive processes and, alongside the interplay between brain networks, also the neurochemistry and neural oscillations that underly creativity.

Oxytocin and the Punitive Hub-Dynamic Spread of Cooperation in Human Social Networks.

Human society operates on large-scale cooperation. However, individual differences in cooperativeness and incentives to free ride on others' cooperation make large-scale cooperation fragile and can lead to reduced social welfare. Thus, how individual cooperation spreads through human social networks remains puzzling from ecological, evolutionary, and societal perspectives. Here, we identify oxytocin and costly punishment as biobehavioral mechanisms that facilitate the propagation of cooperation in social networks. In three laboratory experiments (n = 870 human participants: 373 males, 497 females), individuals were embedded in heterogeneous networks and made repeated decisions with feedback in games of trust (n = 342), ultimatum bargaining (n = 324), and prisoner's dilemma with punishment (n = 204). In each heterogeneous network, individuals at central positions (hub nodes) were given intranasal oxytocin (or placebo). Giving oxytocin (vs matching placebo) to central individuals increased their trust and enforcement of cooperation norms. Oxytocin-enhanced norm enforcement, but not elevated trust, explained the spreading of cooperation throughout the social network. Moreover, grounded in evolutionary game theory, we simulated computer agents that interacted in heterogeneous networks with central nodes varying in terms of cooperation and punishment levels. Simulation results confirmed that central cooperators' willingness to punish noncooperation allowed the permeation of the network and enabled the evolution of network cooperation. These results identify an oxytocin-initiated proximate mechanism explaining how individual cooperation facilitates network-wide cooperation in human society and shed light on the widespread phenomenon of heterogeneous composition and enforcement systems at all levels of life.SIGNIFICANCE STATEMENT Human society operates on large-scale cooperation. Yet because cooperation is exploitable by free riding, how cooperation in social networks emerges remains puzzling from evolutionary and societal perspectives. Here we identify oxytocin and altruistic punishment as key factors facilitating the propagation of cooperation in human social networks. Individuals played repeated economic games in heterogeneous networks where individuals at central positions were given oxytocin or placebo. Oxytocin-enhanced cooperative norm enforcement, but not elevated trust, explained cooperation spreading throughout the social network. Evolutionary simulations confirmed that central cooperators' willingness to punish noncooperation allowed the permeation of the network and enabled the evolution of cooperation. These results identify an oxytocin-initiated proximate mechanism explaining how individual cooperation facilitates network-wide cooperation in human social networks.

Anticipatory Threat Mitigates the Breakdown of Group Cooperation.

Humans are exposed to environmental and economic threats that can profoundly affect individual survival and group functioning. Although anecdotal evidence suggests that threat exposure can increase collective action, the effects of threat on decision-making have been mainly investigated at the individual level. Here we examine how threat exposure and concomitant physiological responses modulate cooperation in small groups. Individuals (N = 105, ages 18-34 years) in groups of three were exposed to threat of electric shocks while deciding how much to contribute to a public good. Threat of shock induced a state of physiological freezing and, compared with no-threat conditions, reduced free riding and enabled groups to maintain higher cooperation over time. Exploratory analyses revealed that more cooperative responses under threat were driven by stronger baseline prosociality, suggesting that habitual prosociality is reinforced under threat. The current results support the view that human groups respond to outside threat with increased cooperation.

Experimental Games and Social Decision Making.

Experimental games model situations in which the future outcomes of individuals and groups depend on their own choices and on those of other (groups of) individuals. Games are a powerful tool to identify the neural and psychological mechanisms underlying interpersonal and group cooperation and coordination. Here we discuss recent developments in how experimental games are used and adapted, with an increased focus on repeated interactions, partner control through sanctioning, and partner (de)selection for future interactions. Important advances have been made in uncovering the neurobiological underpinnings of key factors involved in cooperation and coordination, including social preferences, cooperative beliefs, (emotion) signaling, and, in particular, reputations and (in)direct reciprocity. Emerging trends at the cross-sections of psychology, economics, and the neurosciences include an increased focus on group heterogeneities, intergroup polarization and conflict, cross-cultural differences in cooperation and norm enforcement, and neurocomputational modeling of the formation and updating of social preferences and beliefs.

Four frames and a funeral: Commentary on Bermúdez (2022).

There is much to like in Bermúdez's analysis, yet it is incomplete and at times problematic for social decision making and, by extension, interpersonal conflict. Here I explain how four frames - gains, losses, me, we - operate in conjunction and how humans gravitate toward a "me-loss" frame that, without intervention, leads to a breakdown of cooperation and an arguably tragic funeral of the commons.

When Helping Is Risky: The Behavioral and Neurobiological Trade-off of Social and Risk Preferences.

Helping other people can entail risks for the helper. For example, when treating infectious patients, medical volunteers risk their own health. In such situations, decisions to help should depend on the individual's valuation of others' well-being (social preferences) and the degree of personal risk the individual finds acceptable (risk preferences). We investigated how these distinct preferences are psychologically and neurobiologically integrated when helping is risky. We used incentivized decision-making tasks (Study 1; N = 292 adults) and manipulated dopamine and norepinephrine levels in the brain by administering methylphenidate, atomoxetine, or a placebo (Study 2; N = 154 adults). We found that social and risk preferences are independent drivers of risky helping. Methylphenidate increased risky helping by selectively altering risk preferences rather than social preferences. Atomoxetine influenced neither risk preferences nor social preferences and did not affect risky helping. This suggests that methylphenidate-altered dopamine concentrations affect helping decisions that entail a risk to the helper.

Pupil mimicry promotes trust through the theory-of-mind network.

The human eye can provide powerful insights into the emotions and intentions of others; however, how pupillary changes influence observers' behavior remains largely unknown. The present fMRI-pupillometry study revealed that when the pupils of interacting partners synchronously dilate, trust is promoted, which suggests that pupil mimicry affiliates people. Here we provide evidence that pupil mimicry modulates trust decisions through the activation of the theory-of-mind network (precuneus, temporo-parietal junction, superior temporal sulcus, and medial prefrontal cortex). This network was recruited during pupil-dilation mimicry compared with interactions without mimicry or compared with pupil-constriction mimicry. Furthermore, the level of theory-of-mind engagement was proportional to individual's susceptibility to pupil-dilation mimicry. These data reveal a fundamental mechanism by which an individual's pupils trigger neurophysiological responses within an observer: when interacting partners synchronously dilate their pupils, humans come to feel reflections of the inner states of others, which fosters trust formation.

Neurocognitive Underpinnings of Aggressive Predation in Economic Contests.

Competitions are part and parcel of daily life and require people to invest time and energy to gain advantage over others and to avoid (the risk of) falling behind. Whereas the behavioral mechanisms underlying competition are well documented, its neurocognitive underpinnings remain poorly understood. We addressed this using neuroimaging and computational modeling of individual investment decisions aimed at exploiting one's counterpart ("attack") or at protecting against exploitation by one's counterpart ("defense"). Analyses revealed that during attack relative to defense (i) individuals invest less and are less successful; (ii) computations of expected reward are strategically more sophisticated (reasoning level k = 4 vs. k = 3 during defense); (iii) ventral striatum activity tracks reward prediction errors; (iv) risk prediction errors were not correlated with neural activity in either ROI or whole-brain analyses; and (v) successful exploitation correlated with neural activity in the bilateral ventral striatum, left OFC, left anterior insula, left TPJ, and lateral occipital cortex. We conclude that, in economic contests, coming out ahead (vs. not falling behind) involves sophisticated strategic reasoning that engages both reward and value computation areas and areas associated with theory of mind.

Methylphenidate does not affect convergent and divergent creative processes in healthy adults.

An increasing number of healthy people use methylphenidate, a psychostimulant that increases dopamine and noradrenaline transmission in the brain, to help them focus over extended periods of time. While methylphenidate has been shown to facilitate some cognitive functions, like focus and distractor-resistance, the same drug might also contribute to cognitive impairment, for example, in creativity. In this study, we investigated whether acute administration of a low oral dose (20 mg) of methylphenidate affected convergent and divergent creative processes in a sample of young healthy participants. Also, we explored whether such effects depended on individual differences in ADHD symptoms and working memory capacity. Contrary to our expectations, methylphenidate did not affect participants' creative performance on any of the tasks. Also, methylphenidate effects did not depend on individual differences in trait hyperactivity-impulsivity or baseline working memory capacity. Thus, although the effects of methylphenidate on creativity might be underestimated in our study due to several methodological factors, our findings do not suggest that methylphenidate impairs people's ability to be creative.

Neurobiological Mechanisms of Responding to Injustice.

People are particularly sensitive to injustice. Accordingly, deeper knowledge regarding the processes that underlie the perception of injustice, and the subsequent decisions to either punish transgressors or compensate victims, is of important social value. By combining a novel decision-making paradigm with functional neuroimaging, we identified specific brain networks that are involved with both the perception of, and response to, social injustice, with reward-related regions preferentially involved in punishment compared with compensation. Developing a computational model of punishment allowed for disentangling the neural mechanisms and psychological motives underlying decisions of whether to punish and, subsequently, of how severely to punish. Results show that the neural mechanisms underlying punishment differ depending on whether one is directly affected by the injustice, or whether one is a third-party observer of a violation occurring to another. Specifically, the anterior insula was involved in decisions to punish following harm, whereas, in third-party scenarios, we found amygdala activity associated with punishment severity. Additionally, we used a pharmacological intervention using oxytocin, and found that oxytocin influenced participants' fairness expectations, and in particular enhanced the frequency of low punishments. Together, these results not only provide more insight into the fundamental brain mechanisms underlying punishment and compensation, but also illustrate the importance of taking an explorative, multimethod approach when unraveling the complex components of everyday decision-making.SIGNIFICANCE STATEMENT The perception of injustice is a fundamental precursor to many disagreements, from small struggles at the dinner table to wasteful conflict between cultures and countries. Despite its clear importance, relatively little is known about how the brain processes these violations. Taking an interdisciplinary approach, we combine methods from neuroscience, psychology, and economics to explore the neurobiological mechanisms involved in both the perception of injustice as well as the punishment and compensation decisions that follow. Using a novel behavioral paradigm, we identified specific brain networks, developed a computational model of punishment, and found that administrating the neuropeptide oxytocin increases the administration of low punishments of norm violations in particular. Results provide valuable insights into the fundamental neurobiological mechanisms underlying social injustice.

In-group defense, out-group aggression, and coordination failures in intergroup conflict.

Intergroup conflict persists when and because individuals make costly contributions to their group's fighting capacity, but how groups organize contributions into effective collective action remains poorly understood. Here we distinguish between contributions aimed at subordinating out-groups (out-group aggression) from those aimed at defending the in-group against possible out-group aggression (in-group defense). We conducted two experiments in which three-person aggressor groups confronted three-person defender groups in a multiround contest game (n = 276; 92 aggressor-defender contests). Individuals received an endowment from which they could contribute to their group's fighting capacity. Contributions were always wasted, but when the aggressor group's fighting capacity exceeded that of the defender group, the aggressor group acquired the defender group's remaining resources (otherwise, individuals on both sides were left with the remainders of their endowment). In-group defense appeared stronger and better coordinated than out-group aggression, and defender groups survived roughly 70% of the attacks. This low success rate for aggressor groups mirrored that of group-hunting predators such as wolves and chimpanzees (n = 1,382 cases), hostile takeovers in industry (n = 1,637 cases), and interstate conflicts (n = 2,586). Furthermore, whereas peer punishment increased out-group aggression more than in-group defense without affecting success rates (Exp. 1), sequential (vs. simultaneous) decision-making increased coordination of collective action for out-group aggression, doubling the aggressor's success rate (Exp. 2). The relatively high success rate of in-group defense suggests evolutionary and cultural pressures may have favored capacities for cooperation and coordination when the group goal is to defend, rather than to expand, dominate, and exploit.

Asymmetric conflict: Structures, strategies, and settlement.

Our target article modeled conflict within and between groups as an asymmetric game of strategy and developed a framework to explain the evolved neurobiological, psychological, and sociocultural mechanisms underlying attack and defense. Twenty-seven commentaries add insights from diverse disciplines, such as animal biology, evolutionary game theory, human neuroscience, psychology, anthropology, and political science, that collectively extend and supplement this model in three ways. Here we draw attention to the superordinate structure of attack and defense, and its subordinate means to meet the end of status quo maintenance versus change, and we discuss (1) how variations in conflict structure and power disparities between antagonists can impact strategy selection and behavior during attack and defense; (2) how the positions of attack and defense emerge endogenously and are subject to rhetoric and propaganda; and (3) how psychological and economic interventions can transform attacker-defender conflicts into coordination games that allow mutual gains and dispute resolution.

Revisiting the form and function of conflict: Neurobiological, psychological, and cultural mechanisms for attack and defense within and between groups.

Conflict can profoundly affect individuals and their groups. Oftentimes, conflict involves a clash between one side seeking change and increased gains through victory and the other side defending the status quo and protecting against loss and defeat. However, theory and empirical research largely neglected these conflicts between attackers and defenders, and the strategic, social, and psychological consequences of attack and defense remain poorly understood. To fill this void, we model (1) the clashing of attack and defense as games of strategy and reveal that (2) attack benefits from mismatching its target's level of defense, whereas defense benefits from matching the attacker's competitiveness. This suggests that (3) attack recruits neuroendocrine pathways underlying behavioral activation and overconfidence, whereas defense invokes neural networks for behavioral inhibition, vigilant scanning, and hostile attributions; and that (4) people invest less in attack than defense, and attack often fails. Finally, we propose that (5) in intergroup conflict, out-group attack needs institutional arrangements that motivate and coordinate collective action, whereas in-group defense benefits from endogenously emerging in-group identification. We discuss how games of attack and defense may have shaped human capacities for prosociality and aggression, and how third parties can regulate such conflicts and reduce their waste.

Pupil-mimicry conditions trust in partners: moderation by oxytocin and group membership.

Across species, oxytocin, an evolutionarily ancient neuropeptide, facilitates social communication by attuning individuals to conspecifics' social signals, fostering trust and bonding. The eyes have an important signalling function; and humans use their salient and communicative eyes to intentionally and unintentionally send social signals to others, by contracting the muscles around their eyes and pupils. In our earlier research, we observed that interaction partners with dilating pupils are trusted more than partners with constricting pupils. But over and beyond this effect, we found that the pupil sizes of partners synchronize and that when pupils synchronously dilate, trust is further boosted. Critically, this linkage between mimicry and trust was bound to interactions between ingroup members. The current study investigates whether these findings are modulated by oxytocin and sex of participant and partner. Using incentivized trust games with partners from ingroup and outgroup whose pupils dilated, remained static or constricted, this study replicates our earlier findings. It further reveals that (i) male participants withhold trust from partners with constricting pupils and extend trust to partners with dilating pupils, especially when given oxytocin rather than placebo; (ii) female participants trust partners with dilating pupils most, but this effect is blunted under oxytocin; (iii) under oxytocin rather than placebo, pupil dilation mimicry is weaker and pupil constriction mimicry stronger; and (iv) the link between pupil constriction mimicry and distrust observed under placebo disappears under oxytocin. We suggest that pupil-contingent trust is parochial and evolved in social species in and because of group life.

Oxytocin promotes intuitive rather than deliberated cooperation with the in-group.

A contribution to a special issue on Hormones and Human Competition. In intergroup settings, individuals prefer cooperating with their in-group, and sometimes derogate and punish out-groups. Here we replicate earlier work showing that such in-group bounded cooperation is conditioned by oxytocin and extend it by showing that oxytocin-motivated in-group cooperation is intuitive rather than deliberated. Healthy males (N=65) and females (N=129) self-administered intranasal placebo or 24IU oxytocin in a double-blind placebo-controlled between-subjects design, were assigned to a three-person in-group (that faced a 3-person out-group), and given an endowment from which they could contribute to a within-group pool (benefitting the in-group), and/or to a between-group pool (benefitting the in-group and punishing the out-group). Prior to decision-making, participants performed a Stroop Interference task that was either cognitively taxing, or not. Cognitively taxed individuals kept less to themselves and contributed more to the within-group pool. Furthermore, participants receiving placebo contributed more to the within-group pool when they were cognitively taxed rather than not; those receiving oxytocin contributed to the within-group pool regardless of cognitive taxation. Neither taxation nor treatment influenced contributions to the between-group pool, and no significant sex differences were observed. It follows that in intergroup settings (i) oxytocin increases in-group bounded cooperation, (ii) oxytocin neither reduces nor increases out-group directed spite, and (iii) oxytocin-induced in-group cooperation is independent of cognitive taxation and, therefore, likely to be intuitive rather than consciously deliberated.

Oxytocin promotes group-serving dishonesty.

To protect and promote the well-being of others, humans may bend the truth and behave unethically. Here we link such tendencies to oxytocin, a neuropeptide known to promote affiliation and cooperation with others. Using a simple coin-toss prediction task in which participants could dishonestly report their performance levels to benefit their group's outcome, we tested the prediction that oxytocin increases group-serving dishonesty. A double-blind, placebo-controlled experiment allowing individuals to lie privately and anonymously to benefit themselves and fellow group members showed that healthy males (n = 60) receiving intranasal oxytocin, rather than placebo, lied more to benefit their group, and did so faster, yet did not necessarily do so because they expected reciprocal dishonesty from fellow group members. Treatment effects emerged when lying had financial consequences and money could be gained; when losses were at stake, individuals in placebo and oxytocin conditions lied to similar degrees. In a control condition (n = 60) in which dishonesty only benefited participants themselves, but not fellow group members, oxytocin did not influence lying. Together, these findings fit a functional perspective on morality revealing dishonesty to be plastic and rooted in evolved neurobiological circuitries, and align with work showing that oxytocin shifts the decision-maker's focus from self to group interests. These findings highlight the role of bonding and cooperation in shaping dishonesty, providing insight into when and why collaboration turns into corruption.