Super-additive cooperation.

Repeated interactions provide an evolutionary explanation for one-shot human cooperation that is counterintuitive but orthodox1-3. Intergroup competition4-7 provides an explanation that is intuitive but heterodox. Here, using models and a behavioural experiment, we show that neither mechanism reliably supports cooperation. Ambiguous reciprocity, a class of strategies that is generally ignored in models of reciprocal altruism, undermines cooperation under repeated interactions. This finding challenges repeated interactions as an evolutionary explanation for cooperation in general, which further challenges the claim that repeated interactions in the past can explain one-shot cooperation in the present. Intergroup competitions also do not reliably support cooperation because groups quickly become extremely similar, which limits scope for group selection. Moreover, even if groups vary, group competitions may generate little group selection for multiple reasons. Cooperative groups, for example, may tend to compete against each other8. Whereas repeated interactions and group competitions do not support cooperation by themselves, combining them triggers powerful synergies because group competitions constrain the corrosive effect of ambiguous reciprocity. Evolved strategies often consist of cooperative reciprocity with ingroup partners and uncooperative reciprocity with outgroup partners. Results from a behavioural experiment in Papua New Guinea fit exactly this pattern. They thus suggest neither an evolutionary history of repeated interactions without group competition nor a history of group competition without repeated interactions. Instead, our results suggest social motives that evolved under the joint influence of both mechanisms.

Neural Mechanisms Underlying Individual Differences in Control-Averse Behavior.

When another person tries to control one's decisions, some people might comply, but many will feel the urge to act against that control. This control aversion can lead to suboptimal decisions and it affects social interactions in many societal domains. To date, however, it has been unclear what drives individual differences in control-averse behavior. Here, we address this issue by measuring brain activity with fMRI while healthy female and male human participants made choices that were either free or controlled by another person, with real consequences to both interaction partners. In addition, we assessed the participants' affects, social cognitions, and motivations via self-reports. Our results indicate that the social cognitions perceived distrust and lack of understanding for the other person play a key role in explaining control aversion at the behavioral level. At the neural level, we find that control-averse behavior can be explained by functional connectivity between the inferior parietal lobule and the dorsolateral prefrontal cortex, brain regions commonly associated with attention reorientation and cognitive control. Further analyses reveal that the individual strength of functional connectivity complements and partially mediates the self-reported social cognitions in explaining individual differences in control-averse behavior. These findings therefore provide valuable contributions to a more comprehensive model of control aversion.SIGNIFICANCE STATEMENT Control aversion is a prevalent phenomenon in our society. When someone tries to control their decisions, many people tend to act against the control. This can lead to suboptimal decisions such as noncompliance to medical treatments or disobeying the law. The degree to which individuals engage in control-averse behavior, however, varies significantly. Understanding the proximal mechanisms that underlie individual differences in control-averse behavior has potential policy implications, for example, when designing policies aimed at increasing compliance with vaccination recommendations, and is therefore a highly relevant research goal. Here, we identify a neural mechanism between parietal and prefrontal brain regions that can explain individual differences in control-averse behavior. This mechanism provides novel insights into control aversion beyond what is accessible through self-reports.

Intrinsic connectivity networks underlying individual differences in control-averse behavior.

When people sense that another person tries to control their decisions, some people will act against the control, whereas others will not. This individual tendency to control-averse behavior can have far-reaching consequences, such as engagement in illegal activities or noncompliance with medical treatments. Although individual differences in control-averse behavior have been well documented in behavioral studies, their neurological basis is less well understood. Here, we use a neural trait approach to examine whether individual differences in control-averse behavior might be linked to stable brain-based characteristics. To do so, we analyze the association between intrinsic connectivity networks as measured by resting state functional magnetic resonance imaging and control-averse behavior in an economic exchange game. In this game, subjects make choices that are either free or controlled by another person, with real consequences to both interaction partners. We find that the individual level of control-averse behavior can be positively predicted by intrinsic connectivity within the salience network, but not the central executive network or the default mode network. Specifically, subjects with a more prominent connectivity hub in the dorsal anterior cingulate cortex show greater levels of control-averse behavior. This finding provides the first evidence that the heterogeneity in control-averse behavior might originate in systematic differences of the stable functional brain organization.

The scent of attractiveness: levels of reproductive hormones explain individual differences in women's body odour.

Individuals are thought to have their own distinctive body odour which reportedly plays an important role in mate choice. In the present study we investigated individual differences in body odours of women and examined whether some women generally smell more attractive than others or whether odour preferences are a matter of individual taste. We then explored whether levels of reproductive hormones explain women's body odour attractiveness, to test the idea that body odour attractiveness may act as a chemosensory marker of reproductive fitness. Fifty-seven men rated body odours of 28 healthy, naturally cycling women of reproductive age. We collected all odours at peak fertility to control for menstrual cycle effects on body odour attractiveness. Women's salivary oestradiol, progesterone, testosterone and cortisol levels were assessed at the time of odour collection to test whether hormone levels explain body odour attractiveness. We found that the men highly agreed on how attractive they found women's body odours. Interestingly, women's body odour attractiveness was predicted by their oestradiol and progesterone levels: the higher a woman's levels of oestradiol and the lower her levels of progesterone, the more attractive her body odour was rated. In showing that women's body odour attractiveness is explained by levels of female reproductive hormones, but not by levels of cortisol or testosterone, we provide evidence that body odour acts as a valid cue to potential fertility.

Men's preferences for women's body odours are not associated with human leucocyte antigen.

Body odours reportedly portray information about an individual's genotype at the major histocompatibility complex (MHC, called human leucocyte antigen, HLA, in humans). While there is strong experimental support for MHC-associated mating behaviour in animals, the situation in humans is more complex. A lot of effort has been spent on testing HLA-associated odour preferences of women. To date, only very few studies have looked at HLA-linked olfactory preferences in men and these studies have revealed inconsistent results. Here, we investigate men's HLA-associated preferences for women's body odours. Importantly, and in contrast to previous studies, these odours were gathered at peak fertility (i.e. just before ovulation) when any HLA-associated odour preferences should be strongest. We scrutinized whether men's preference for women's body odours is influenced by (i) the number of shared HLA alleles between men and women, (ii) HLA heterozygosity, and (iii) the frequency of rare HLA alleles. We found that men could readily differentiate between odours they found attractive and odours they found less attractive, but that these preferences were not associated with HLA. Specifically, men did not prefer odours from women who are HLA dissimilar, HLA heterozygous, or who have rare HLA alleles. Together, these findings suggest that HLA has no effect on men's odour preferences.

The social dimension of stress reactivity: acute stress increases prosocial behavior in humans.

Psychosocial stress precipitates a wide spectrum of diseases with major public-health significance. The fight-or-flight response is generally regarded as the prototypic human stress response, both physiologically and behaviorally. Given that having positive social interactions before being exposed to acute stress plays a preeminent role in helping individuals control their stress response, engaging in prosocial behavior in response to stress (tend-and-befriend) might also be a protective pattern. Little is known, however, about the immediate social responses following stress in humans. Here we show that participants who experienced acute social stress, induced by a standardized laboratory stressor, engaged in substantially more prosocial behavior (trust, trustworthiness, and sharing) compared with participants in a control condition, who did not experience socioevaluative threat. These effects were highly specific: Stress did not affect the readiness to exhibit antisocial behavior or to bear nonsocial risks. These results show that stress triggers social approach behavior, which operates as a potent stress-buffering strategy in humans, thereby providing evidence for the tend-and-befriend hypothesis.

Parochial altruism in humans.

Social norms and the associated altruistic behaviours are decisive for the evolution of human cooperation and the maintenance of social order, and they affect family life, politics and economic interactions. However, as altruistic norm compliance and norm enforcement often emerge in the context of inter-group conflicts, they are likely to be shaped by parochialism--a preference for favouring the members of one's ethnic, racial or language group. We have conducted punishment experiments, which allow 'impartial' observers to punish norm violators, with indigenous groups in Papua New Guinea. Here we show that these experiments confirm the prediction of parochialism. We found that punishers protect ingroup victims--who suffer from a norm violation--much more than they do outgroup victims, regardless of the norm violator's group affiliation. Norm violators also expect that punishers will be lenient if the latter belong to their social group. As a consequence, norm violations occur more often if the punisher and the norm violator belong to the same group. Our results are puzzling for evolutionary multi-level selection theories based on selective group extinction as well as for theories of individual selection; they also indicate the need to explicitly examine the interactions between individuals stemming from different groups in evolutionary models.

Which hospital workers do (not) want the jab? Behavioral correlates of COVID-19 vaccine willingness among employees of Swiss hospitals.

In many countries, the current vaccination rates are stagnating, to the extent that vaccine hesitancy-the delay or refusal to take recommended vaccinations-forms a major obstacle to ending the COVID-19 pandemic. This tendency is particularly concerning when observed among healthcare workers who are opinion leaders on medical matters for their patients and peers. Our study surveys 965 employees of two large Swiss hospitals and profiles vaccine-hesitant hospital employees using not only socio-demographic characteristics, but also a comprehensive set of standard behavioral preference measures: (i) Big-5 personality traits, (ii) risk-, time- and social preferences, and (iii) perceived prevailing social norms. Using multinomial probit models and linear probability models, we find that vaccine-hesitant hospital employees are less patient and less likely to perceive vaccination as the prevailing social norm-in addition to replicating previously published socio-demographic results. Our findings are robust to a range of model specifications, as well as individual and situational covariates. Our study thus offers actionable policy implications for tailoring public-health communications to vaccine-hesitant hospital employees.

The neural signature of social norm compliance.

All known human societies establish social order by punishing violators of social norms. However, little is known about how the brain processes the punishment threat associated with norm violations. We use fMRI to study the neural circuitry behind social norm compliance by comparing a treatment in which norm violations can be punished with a control treatment in which punishment is impossible. Individuals' increase in norm compliance when punishment is possible exhibits a strong positive correlation with activations in the lateral orbitofrontal cortex and right dorsolateral prefrontal cortex. Moreover, lateral orbitofrontal cortex activity is strongly correlated with Machiavellian personality characteristics. These findings indicate a neural network involved in social norm compliance that might constitute an important basis for human sociality. Different activations of this network reveal individual differences in the behavioral response to the punishment threat and might thus provide a deeper understanding of the neurobiological sources of pathologies such as antisocial personality disorder.

Oxytocin increases trust in humans.

Trust pervades human societies. Trust is indispensable in friendship, love, families and organizations, and plays a key role in economic exchange and politics. In the absence of trust among trading partners, market transactions break down. In the absence of trust in a country's institutions and leaders, political legitimacy breaks down. Much recent evidence indicates that trust contributes to economic, political and social success. Little is known, however, about the biological basis of trust among humans. Here we show that intranasal administration of oxytocin, a neuropeptide that plays a key role in social attachment and affiliation in non-human mammals, causes a substantial increase in trust among humans, thereby greatly increasing the benefits from social interactions. We also show that the effect of oxytocin on trust is not due to a general increase in the readiness to bear risks. On the contrary, oxytocin specifically affects an individual's willingness to accept social risks arising through interpersonal interactions. These results concur with animal research suggesting an essential role for oxytocin as a biological basis of prosocial approach behaviour.

The modulation of social behavior and empathy via oral contraceptives and female sex hormones.

Oral contraceptives (OC) and endogenous female sex hormones in naturally cycling women (NC) are related to a wide range of psychological variables (eg, cognition and affect). Little research on social behavior has been done. One study documented a tendency towards more prosocial behavior in NC than OC women, but the underlying neuroendocrine mechanisms remain unknown. The sex hormones progesterone and estradiol are potential candidates. We analyzed social decision-making and social behavior in 83 healthy women (38 OC and 45 NC) via the Social Value Orientation (SVO) and in real social interactions within a paradigm adapted from behavioral economics. We also measured empathy, and collected saliva samples to quantify the basal levels of estradiol and progesterone. Our analyses revealed higher levels of prosocial behavior and emotional empathy in NC than in OC women, a finding supported by higher levels of prosocial decisions in NC than OC women in the SVO. Regarding the underlying biological mechanisms, we detected lower progesterone levels in OC than NC women. Exploratory analyses revealed a negative correlation between progesterone and trust on the trend level. We found no correlations between estradiol and behavior. Our findings provide evidence that OC modulate social behavior and initial indications of a possible modulation by progesterone. Further research is needed to replicate our findings and extend them to other social behaviors.

Wearing a mask-For yourself or for others? Behavioral correlates of mask wearing among COVID-19 frontline workers.

Human behavior can have effects on oneself and externalities on others. Mask wearing is such a behavior in the current pandemic. What motivates people to wear face masks in public when mask wearing is voluntary or not enforced? Which benefits should the policy makers rather emphasize in information campaigns-the reduced chances of getting the SARS-CoV-2 virus (benefits for oneself) or the reduced chances of transmitting the virus (benefits for others in the society)? In this paper, we link measured risk preferences and other-regarding preferences to mask wearing habits among 840 surveyed employees of two large Swiss hospitals. We find that the leading mask-wearing motivations change with age: While for older people, mask wearing habits are best explained by their self-regarding risk preferences, younger people are also motivated by other-regarding concerns. Our results are robust to different specifications including linear probability models, probit models and Lasso covariate selection models. Our findings thus allow drawing policy implications for effectively communicating public-health recommendations to frontline workers during the COVID-19 pandemic.

The nature of human altruism.

Some of the most fundamental questions concerning our evolutionary origins, our social relations, and the organization of society are centred around issues of altruism and selfishness. Experimental evidence indicates that human altruism is a powerful force and is unique in the animal world. However, there is much individual heterogeneity and the interaction between altruists and selfish individuals is vital to human cooperation. Depending on the environment, a minority of altruists can force a majority of selfish individuals to cooperate or, conversely, a few egoists can induce a large number of altruists to defect. Current gene-based evolutionary theories cannot explain important patterns of human altruism, pointing towards the importance of both theories of cultural evolution as well as gene-culture co-evolution.

Pleasant body odours, but not genetic similarity, influence trustworthiness in a modified trust game.

Identifying trustworthy partners is an important adaptive challenge for establishing mutually cooperative relationships. Previous studies have demonstrated a marked relationship between a person's attractiveness and his apparent trustworthiness (beauty premium). Kin selection theory, however, suggests that cues to kinship enhance trustworthiness. Here we directly tested predictions of the beauty premium and kin selection theory by using body odours as cues to trustworthiness. Body odours reportedly portray information about an individuals' genotype at the human leucocyte antigen system (HLA) and thus olfactory cues in body odours serve as a promising means for kin recognition. Ninety men played trust games in which they divided uneven sums of monetary units between two male trustees represented by their body odour and rated each body odour for pleasantness. Half of the odours came from HLA-similar men (suggesting closer kin) and half from HLA dissimilar men (suggesting non-kin). We found that the amount of money the players transferred was not related to HLA-similarity, but to the pleasantness of the trustee's body odour. By showing that people with more pleasant body odours are trusted more than people with unpleasant body odour we provide evidence for a "beauty-premium" that overrides any putative effect of kin.

The Trust Game for Couples (TGC): A new standardized paradigm to assess trust in romantic relationships.

Trust between couples is a prerequisite for stable and satisfactory romantic relationships. However, there has been no valid research tool to assess partner-specific trust behavior including costly investments in the trustworthiness of the romantic partner. We here present a comprehensive validation of the newly developed Trust Game for Couples (TGC) by means of various self-report and implicit relationship-related measures. The TGC operationalizes trust by measuring an individual's willingness to invest his or her own financial resources in pro-relationship attitudes of their romantic partner (collected by dichotomous responses to relationship-relevant items, e.g., answering yes to "I am absolutely sure that I love my partner"). Thirty-five healthy couples between 20 and 34 years completed the TGC in an interactive (both partners present), but anonymous setting (no information on the partner's responses revealed). Trust, as measured by the TGC, correlates positively with self-reported trust, satisfaction, and felt closeness in the relationship, but not with general interpersonal trust, confirming both its convergent and discriminant validity. In addition to explicit criteria for construct validity, implicit measures of partner valence and confidence explained variance in the TGC, demonstrating that it constitutes an economical measure of implicit and explicit ingredients of trust between couples. In sum, the TGC provides a novel, specific behavioral tool for a sensitive assessment of trust in dyadic relationships with potential for numerous research fields.

Studying the neurobiology of social interaction with transcranial direct current stimulation--the example of punishing unfairness.

Studying social behavior often requires the simultaneous interaction of many subjects. As yet, however, no painless, noninvasive brain stimulation tool existed that allowed the simultaneous affection of brain processes in many interacting subjects. Here we show that transcranial direct current stimulation (tDCS) can overcome these limits. We apply right prefrontal cathodal tDCS and show that subjects' propensity to punish unfair behavior is reduced significantly.

Effects of acute stress on social behavior in women.

Acute stress is known to increase prosocial behavior in men via a "tend-and-befriend" pattern originally proposed as a specifically female stress response alongside the fight-or-flight response. However, the direct effects of acute stress on women's social behavior have not been investigated. Applying the Trier Social Stress Test for groups (TSST-G), 94 women were confronted with either a stress or control condition. We repeatedly measured their subjective stress responses, salivary cortisol, and heart rate, and investigated their level of trust, trustworthiness, sharing, punishment and non-social risk using social decision paradigms. We detected significant increases in all stress parameters, as well as the wish for closeness during the stress condition. Acute stress exposure elevated prosocial trustworthiness and sharing without affecting non-social risk behavior. These results are in line with findings on the effects of stress in men, and further validate the tend-and-befriend pattern as one possible behavioral response during stress in humans.

Genome-wide association analyses of risk tolerance and risky behaviors in over 1 million individuals identify hundreds of loci and shared genetic influences.

Humans vary substantially in their willingness to take risks. In a combined sample of over 1 million individuals, we conducted genome-wide association studies (GWAS) of general risk tolerance, adventurousness, and risky behaviors in the driving, drinking, smoking, and sexual domains. Across all GWAS, we identified hundreds of associated loci, including 99 loci associated with general risk tolerance. We report evidence of substantial shared genetic influences across risk tolerance and the risky behaviors: 46 of the 99 general risk tolerance loci contain a lead SNP for at least one of our other GWAS, and general risk tolerance is genetically correlated ([Formula: see text] ~ 0.25 to 0.50) with a range of risky behaviors. Bioinformatics analyses imply that genes near SNPs associated with general risk tolerance are highly expressed in brain tissues and point to a role for glutamatergic and GABAergic neurotransmission. We found no evidence of enrichment for genes previously hypothesized to relate to risk tolerance.