Do marmosets care to share? Oxytocin treatment reduces prosocial behavior toward strangers

Cooperatively-breeding and socially-monogamous primates, like marmosets and humans, exhibit high levels of social tolerance and prosociality toward others. Oxytocin (OXT) generally facilitates prosocial behavior, but there is growing recognition that OXT modulation of prosocial behavior is shaped by the context of social interactions and by other motivational states such as arousal or anxiety. To determine whether prosociality varies based on social context, we evaluated whether marmoset donors (Callithrix penicillata) preferentially rewarded pairmates versus opposite-sex strangers in a prosocial food-sharing task. To examine potential links among OXT, stress systems, and prosociality, we evaluated whether pretrial cortisol levels in marmosets altered the impact of OXT on prosocial responses. Marmosets exhibited spontaneous prosociality toward others, but they did so preferentially toward strangers compared to their pairmates. When donor marmosets were treated with marmoset-specific Pro⁸-OXT, they exhibited reduced prosociality toward strangers compared to marmosets treated with saline or consensus-mammalian Leu⁸-OXT. When pretrial cortisol levels were lower, marmosets exhibited higher prosociality toward strangers. These findings demonstrate that while marmosets show spontaneous prosocial responses toward others, they do so preferentially toward opposite-sex strangers. Cooperative breeding may be associated with the expression of prosociality, but the existence of a pair-bond between marmoset partners appears to be neither necessary nor sufficient for the expression of spontaneous prosocial responses. Furthermore, high prosociality toward strangers is significantly reduced in marmosets treated with Pro⁸-OXT, suggesting that OXT does not universally enhance prosociality, but, rather OXT modulation of prosocial behavior varies depending on social context.     

Friends without benefits: When we react negatively to helpful and generous friends

Being able to identify reliable friends and allies is key to surviving and thriving in the social world. Many cooperative accounts of friendship argue that people select friends based on how helpful and generous they are. While people certainly like helpful and generous others, here we explore a context in which people might respond negatively to a friend being prosocial: When one's friend is more helpful or generous toward another friend. We argue that such preferential prosociality prompts negative reactions, even when the alternative is a friend being less prosocial overall, because giving preferentially to another friend may be viewed as a threat of potential displacement of one's own friendship. In four studies (N?=?702), we predict and find that people respond negatively to a friend who was more helpful (Studies 1–2) and generous (Studies 3–4), preferring instead that a friend be less helpful and generous overall. Importantly, this preferential prosociality was viewed as particularly negative when the recipient was another friend and was seen as much less negative when the recipient was a relative (Study 1 and 4) or a romantic partner (Study 2). We discuss the implications of these results for cooperation and alliance-based accounts of friendship.     

Preschool Children Fail Primate Prosocial Game Because of Attentional Task Demands

Various nonhuman primate species have been tested with prosocial games (i.e. derivates from dictator games) in order to better understand the evolutionary origin of proactive prosociality in humans. Results of these efforts are mixed, and it is difficult to disentangle true species differences from methodological artifacts. We tested 2- to 5-year-old children with a costly and a cost-free version of a prosocial game that differ with regard to the payoff distribution and are widely used with nonhuman primates. Simultaneously, we assessed the subjects’ level of Theory of Mind understanding. Prosocial behavior was demonstrated with the prosocial game, and did not increase with more advanced Theory of Mind understanding. However, prosocial behavior could only be detected with the costly version of the game, whereas the children failed the cost-free version that is most commonly used with nonhuman primates. A detailed comparison of the children’s behavior in the two versions of the game indicates that the failure was due to higher attentional demands of the cost-free version, rather than to a lack of prosociality per se. Our results thus show (i) that subtle differences in prosociality tasks can substantially bias the outcome and thus prevent meaningful species comparisons, and (ii) that like in nonhuman primates, prosocial behavior in human children does not require advanced Theory of Mind understanding in the present context. However, both developmental and comparative psychology accumulate increasing evidence for the multidimensionality of prosocial behaviors, suggesting that different forms of prosociality are also regulated differentially. For future efforts to understand the evolutionary origin of prosociality it is thus crucial to take this heterogeneity into account.     

Benefiting friends or dominants: prosocial choices mainly depend on rank position in long-tailed macaques (<Emphasis Type="Italic">Macaca fascicularis</Emphasis>)

Long-term observational studies in a number of animal species suggest that exchange patterns of social acts depend on long-term emotional bonds. Therefore, it is expected that the frequency of prosocial behavior will depend on the strength of such a bond. In this study we tested whether variation in relationship quality among unrelated individuals, i.e., “friends” and “nonfriends,” is predictive of the prosocial behavior of long-tailed macaques in two experiments. First, we related relationship quality to prosociality in a dyadic prosociality test, and second, we gave subjects the choice to give to either a friend or a nonfriend in a triadic choice test. We show that prosocial behavior of long-tailed macaques in the dyadic test is not related to relationship quality. When given the choice to give to either a friend or a nonfriend in the triadic test, there is a minor indication that long-tailed macaques show a preference to give to their friends, yet this indication is neither significant nor consistent. In contrast, subordinate long-tailed macaques make a more “competitive” choice and avoid giving to the individual closest in rank. Therefore, in the short-term situation of experimental tests, prosocial behavior of long-tailed macaques seems unaffected by the relationship quality of the dyad/triad tested, and the relative dominance position of these dyads/triads seems to have a much stronger effect on their prosocial behavior.     

Human prosociality from an evolutionary perspective: variation and correlations at a city-wide scale

Prosociality is a fundamental theme in all branches of the human behavioral sciences. Evolutionary theory sets an even broader stage by examining prosociality in all species, including the distinctive human capacity to cooperate in large groups of unrelated individuals. We use evolutionary theory to investigate human prosociality at the scale of a small city (Binghamton, NY), based on survey data and a direct measure of prosocial behavior. In a survey of public school students (Grades 6–12), individual prosociality correlates strongly with social support, which is a basic requirement for prosociality to succeed as a behavioral strategy in Darwinian terms. The most prosocial individuals receive social support from multiple sources (e.g., family, school, neighborhood, religion and extracurricular activities). Neighborhood social support is significant as a group-level variable in addition to an individual-level variable. The median income of a neighborhood does not directly influence individual prosociality, but only indirectly through forms of social support. Variation in neighborhood quality, as measured by the survey, corresponds to the likelihood that a stamped addressed letter dropped on the sidewalk of a given neighborhood will be mailed. We discuss the results in relation to evolutionary theory, the experimental economics literature and the social capital literature in an effort to integrate the study of human prosociality across disciplines.     

How Do Rituals Affect Cooperation?

Collective rituals have long puzzled anthropologists, yet little is known about how rituals affect participants. Our study investigated the effects of nine naturally occurring rituals on prosociality. We operationalized prosociality as (1) attitudes about fellow ritual participants and (2) decisions in a public goods game. The nine rituals varied in levels of synchrony and levels of sacred attribution. We found that rituals with synchronous body movements were more likely to enhance prosocial attitudes. We also found that rituals judged to be sacred were associated with the largest contributions in the public goods game. Path analysis favored a model in which sacred values mediate the effects of synchronous movements on prosocial behaviors. Our analysis offers the first quantitative evidence for the long-standing anthropological conjecture that rituals orchestrate body motions and sacred values to support prosociality. Our analysis, moreover, adds precision to this old conjecture with evidence of a specific mechanism: ritual synchrony increases perceptions of oneness with others, which increases sacred values to intensify prosocial behaviors.     

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.     

Giving leads to happiness in young children

Evolutionary models of cooperation require proximate mechanisms that sustain prosociality despite inherent costs to individuals. The "warm glow" that often follows prosocial acts could provide one such mechanism; if so, these emotional benefits may be observable very early in development. Consistent with this hypothesis, the present study finds that before the age of two, toddlers exhibit greater happiness when giving treats to others than receiving treats themselves. Further, children are happier after engaging in costly giving--forfeiting their own resources--than when giving the same treat at no cost. By documenting the emotionally rewarding properties of costly prosocial behavior among toddlers, this research provides initial support for the claim that experiencing positive emotions when giving to others is a proximate mechanism for human cooperation.     

On the relationship of emotional abilities and prosocial behavior

The ability to perceive and infer the meaning of facial expressions has been considered a critical component of emotional intelligence being essential for successful social functioning: Longitudinal findings suggest that the ability to recognize emotion cues is related to positive social interactions. Moreover, pronounced recognition abilities for at least some emotions facilitate prosocial behavior in everyday situations. Integrating paradigms from behavioral economics and psychometrics, we used an interdisciplinary approach to study the relationship between prosociality as trait cooperativeness and the ability to recognize emotions in others. We measured emotion recognition accuracy (ERA) using a multivariate test battery. We captured prosocial behavior in standard socio-economic games, along with spontaneous emotion expressions. Structural equation modeling revealed no significant relationship between overall ERA and prosocial behavior. However, modeling emotion-specific factors suggested that more prosocial individuals are better in recognizing fear and tend to express more spontaneous emotions during the prisoner's dilemma. In all, cooperative individuals seem to be more sensitive to the distress of others and more expressive, possibly fostering reciprocal interactions with like-minded others.     

When the selfish suffer: evidence for selective prosocial emotional and physiological responses to suffering egoists

Prosociality is fundamental to social relationships, but providing it indiscriminately risks exploitation by egoists. Past work demonstrates that individuals avoid these risks through a more selective form of prosociality, cooperating less and sharing fewer resources with egoists (e.g. Axelrod & Hamilton, 1981). The evolution of cooperation. Science, 211(4489), 1390–1396). We extend this work to explore whether individuals experience reduced prosocial affective and physiological responses to egoists in situations where they are suffering. In two studies, participants learned of a target’s egoistic or non-egoistic traits, and then encountered the target suffering. Suffering egoists evoked less compassion in others than non-egoists and elicited physiological responses that diverged from patterns associated with compassion and social engagement (reduced heart rate and greater respiratory sinus arrhythmia activity). Participants' feelings of distrust toward egoists explained these attenuated emotional and physiological responses. These results build upon studies of prosocial behavior by suggesting that individuals experience reduced prosocial emotional and physiological responses toward suffering egoists.     

A common heritable factor influences prosocial obligations across multiple domains

Although it has been shown that prosocial behaviour is heritable, it has not yet been established whether narrower aspects of prosociality are heritable, nor whether a common mechanism influences prosociality across its multiple domains. Here, we examine civic duty, work-place commitment and concern for the welfare of others with a study of prosocial obligations in 958 adult twin-pairs. Multivariate modelling indicated the existence of genetic factors underlying general prosocial obligations in females, with familial effects (genetic and shared-environment effects were indistinguishable) influencing this general mechanism in males. At the domain-specific level, modest genetic effects were observed in females for civic and work obligations, with shared-environment effects influencing welfare obligations. In males, genetic influences were observed for welfare obligation, with unique environments affecting work and civic duty.     

Fourteen-month-old infants use interpersonal synchrony as a cue to direct helpfulness

Musical behaviours such as dancing, singing and music production, which require the ability to entrain to a rhythmic beat, encourage high levels of interpersonal coordination. Such coordination has been associated with increased group cohesion and social bonding between group members. Previously, we demonstrated that this association influences even the social behaviour of 14-month-old infants. Infants were significantly more likely to display helpfulness towards an adult experimenter following synchronous bouncing compared with asynchronous bouncing to music. The present experiment was designed to determine whether interpersonal synchrony acts as a cue for 14-month-olds to direct their prosocial behaviours to specific individuals with whom they have experienced synchronous movement, or whether it acts as a social prime, increasing prosocial behaviour in general. Consistent with the previous results, infants were significantly more likely to help an experimenter following synchronous versus asynchronous movement with this person. Furthermore, this manipulation did not affect infant''s behaviour towards a neutral stranger, who was not involved in any movement experience. This indicates that synchronous bouncing acts as a social cue for directing prosociality. These results have implications for how musical engagement and rhythmic synchrony affect social behaviour very early in development.     

Multiple reputation domains and cooperative behaviour in two Latin American communities

Reputations are a ubiquitous feature of human social life, and a large literature has been dedicated to explaining the relationship between prosocial reputations and cooperation in social dilemmas. However, humans form reputations in domains other than prosociality, such as economic competency that could affect cooperation. To date, no research has evaluated the relative effects of multiple reputation domains on cooperation. To bridge this gap, we analyse how prosocial and competency reputations affect cooperation in two Latin American communities (Bwa Mawego, Dominica, and Pucucanchita, Peru) across a number of social contexts (Dominica: labour contracting, labour exchange and conjugal partnership formation; Peru: agricultural and health advice network size). First, we examine the behavioural correlates of prosocial and competency reputations. Following, we analyse whether prosocial, competency, or both reputation domains explain the flow of cooperative benefits within the two communities. Our analyses suggest that (i) although some behaviours affect both reputation domains simultaneously, each reputation domain has a unique behavioural signature; and (ii) competency reputations affect cooperation across a greater number of social contexts compared to prosocial reputations. Results are contextualized with reference to the social markets in which behaviour is embedded and a call for greater theory development is stressed.     

Prosocial behaviour emerges independent of reciprocity in cottontop tamarins

The cooperative breeding hypothesis posits that cooperatively breeding species are motivated to act prosocially, that is, to behave in ways that provide benefits to others, and that cooperative breeding has played a central role in the evolution of human prosociality. However, investigations of prosocial behaviour in cooperative breeders have produced varying results and the mechanisms contributing to this variation are unknown. We investigated whether reciprocity would facilitate prosocial behaviour among cottontop tamarins, a cooperatively breeding primate species likely to engage in reciprocal altruism, by comparing the number of food rewards transferred to partners who had either immediately previously provided or denied rewards to the subject. Subjects were also tested in a non-social control condition. Overall, results indicated that reciprocity increased food transfers. However, temporal analyses revealed that when the tamarins'' behaviour was evaluated in relation to the non-social control, results were best explained by (i) an initial depression in the transfer of rewards to partners who recently denied rewards, and (ii) a prosocial effect that emerged late in sessions independent of reciprocity. These results support the cooperative breeding hypothesis, but suggest a minimal role for positive reciprocity, and emphasize the importance of investigating proximate temporal mechanisms underlying prosocial behaviour.     

Lack of prosociality in great apes,capuchin monkeys and spider monkeys: convergent evidence from two different food distribution tasks

Prosociality can be defined as any behaviour performed to alleviate the needs of others or to improve their welfare. Prosociality has probably played an essential role in the evolution of cooperative behaviour and several studies have already investigated it in primates to understand the evolutionary origins of human prosociality. Two main tasks have been used to test prosociality in a food context. In the Platforms task, subjects can prosocially provide food to a partner by selecting a prosocial platform over a selfish one. In the Tokens task, subjects can prosocially provide food to a partner by selecting a prosocial token over a selfish one. As these tasks have provided mixed results, we used both tasks to test prosociality in great apes, capuchin monkeys and spider monkeys. Our results provided no compelling evidence of prosociality in a food context in any of the species tested. Additionally, our study revealed serious limitations of the Tokens task as it has been previously used. These results highlight the importance of controlling for confounding variables and of using multiple tasks to address inconsistencies present in the literature.     

Young Children Are More Generous When Others Are Aware of Their Actions

Adults frequently employ reputation-enhancing strategies when engaging in prosocial acts, behaving more generously when their actions are likely to be witnessed by others and even more so when the extent of their generosity is made public. This study examined the developmental origins of sensitivity to cues associated with reputationally motivated prosociality by presenting five-year-olds with the option to provide one or four stickers to a familiar peer recipient at no cost to themselves. We systematically manipulated the recipient’s knowledge of the actor’s choices in two different ways: (1) occluding the recipient’s view of both the actor and the allocation options and (2) presenting allocations in opaque containers whose contents were visible only to the actor. Children were consistently generous only when the recipient was fully aware of the donation options; in all cases in which the recipient was not aware of the donation options, children were strikingly ungenerous. These results demonstrate that five-year-olds exhibit “strategic prosociality,” behaving differentially generous as a function of the amount of information available to the recipient about their actions. These findings suggest that long before they develop a rich understanding of the social significance of reputation or are conscious of complex strategic reasoning, children behave more generously when the details of their prosocial actions are available to others.     

Task Design Influences Prosociality in Captive Chimpanzees (Pan troglodytes)

Chimpanzees confer benefits on group members, both in the wild and in captive populations. Experimental studies of how animals allocate resources can provide useful insights about the motivations underlying prosocial behavior, and understanding the relationship between task design and prosocial behavior provides an important foundation for future research exploring these animals'' social preferences. A number of studies have been designed to assess chimpanzees'' preferences for outcomes that benefit others (prosocial preferences), but these studies vary greatly in both the results obtained and the methods used, and in most cases employ procedures that reduce critical features of naturalistic social interactions, such as partner choice. The focus of the current study is on understanding the link between experimental methodology and prosocial behavior in captive chimpanzees, rather than on describing these animals'' social motivations themselves. We introduce a task design that avoids isolating subjects and allows them to freely decide whether to participate in the experiment. We explore key elements of the methods utilized in previous experiments in an effort to evaluate two possibilities that have been offered to explain why different experimental designs produce different results: (a) chimpanzees are less likely to deliver food to others when they obtain food for themselves, and (b) evidence of prosociality may be obscured by more “complex” experimental apparatuses (e.g., those including more components or alternative choices). Our results suggest that the complexity of laboratory tasks may generate observed variation in prosocial behavior in laboratory experiments, and highlights the need for more naturalistic research designs while also providing one example of such a paradigm.     

The influence of body weight on social network ties among adolescents

Evidence of negative stereotypes, prejudice and discrimination towards obese individuals has been widely documented. However, the effect of a larger body size on social network ties or friendship formations is less well understood. In this paper, we explore the extent to which higher body weight results in social marginalization of adolescents. Using data from a nationally representative sample of adolescents, we estimate endogeneity-corrected models including school-level fixed effects that account for bi-directionality and unobserved confounders to ascertain the effect of body weight on social network ties. We find that obese adolescents have fewer friends and are less socially integrated than their non-obese counterparts. We also find that such penalties in friendship networks are present among whites but not African-Americans or Hispanics, with the largest effect among white females. These results are robust to common environmental influences at the school-level and to controls for preferences, risk attitudes, low self-esteem and objective measures of physical attractiveness.     

Alcohol Use among Adolescent Youth: The Role of Friendship Networks and Family Factors in Multiple School Studies

To explore the co-evolution of friendship tie choice and alcohol use behavior among 1,284 adolescents from 12 small schools and 976 adolescents from one big school sampled in the National Longitudinal Study of Adolescent to Adult Health (AddHealth), we apply a Stochastic Actor-Based (SAB) approach implemented in the R-based Simulation Investigation for Empirical Network Analysis (RSiena) package. Our results indicate the salience of both peer selection and peer influence effects for friendship tie choice and adolescent drinking behavior. Concurrently, the main effect models indicate that parental monitoring and the parental home drinking environment affected adolescent alcohol use in the small school sample, and that parental home drinking environment affected adolescent drinking in the large school sample. In the small school sample, we detect an interaction between the parental home drinking environment and choosing friends that drink as they multiplicatively affect friendship tie choice. Our findings suggest that future research should investigate the synergistic effects of both peer and parental influences for adolescent friendship tie choices and drinking behavior. And given the tendency of adolescents to form ties with their friends'' friends, and the evidence of local hierarchy in these networks, popular youth who do not drink may be uniquely positioned and uniquely salient as the highest rank of the hierarchy to cause anti-drinking peer influences to diffuse down the social hierarchy to less popular youth. As such, future interventions should harness prosocial peer influences simultaneously with strategies to increase parental support and monitoring among parents to promote affiliation with prosocial peers.     

A mechanism for the evolution of altruism among nonkin: positive assortment through environmental feedback

The evolution of altruism often requires genetic similarity among interactors. For structured populations in which a social trait affects all group members, this entails positive assortment, meaning that cooperators and noncooperators tend to be segregated into different groups. Several authors have claimed that mechanisms other than common descent can produce positive assortment, but this claim has not been generally accepted. Here, we describe one such mechanism. The process of "environmental feedback" requires only that the cooperative trait affects the quality of the local environment and that individuals are more likely to leave low-quality than high-quality environments. We illustrate this dynamic using an agent-based spatial model of feeding restraint. Depending on parameter settings, results included both positive assortment (required for the evolution of altruism) and negative assortment (required for the evolution of spite). The mechanism of environmental feedback appears to be a general one that could play a role in the evolution of many forms of cooperation.