Adrian V. Jaeggi

On the psychology of cooperation in humans and other primates: combining the natural history and experimental evidence of prosociality

In any given species, cooperation involves prosocial acts that usually return a fitness benefit to the actor. These acts are produced by a set of psychological rules, which will be similar in related species if they have a similar natural history of cooperation. Prosocial acts can be (i) reactive, i.e. in response to specific stimuli, or (ii) proactive, i.e. occur in the absence of such stimuli. We propose that reactive prosocial acts reflect sensitivity to (i) signals or signs of need and (ii) the presence and size of an audience, as modified by (iii) social distance to the partner or partners. We examine the evidence for these elements in humans and other animals, especially non-human primates, based on the natural history of cooperation, quantified in the context of food sharing, and various experimental paradigms. The comparison suggests that humans share with their closest living relatives reactive responses to signals of need, but differ in sensitivity to signs of need and cues of being watched, as well as in the presence of proactive prosociality. We discuss ultimate explanations for these derived features, in particular the adoption of cooperative breeding as well as concern for reputation and costly signalling during human evolution.

Social learning of diet and foraging skills by wild immature Bornean orangutans: implications for culture

Studies of social learning in the wild are important to complement findings from experiments in captivity. In this field study, immature Bornean orangutans rarely foraged independently but consistently followed their mothers choices. Their diets were essentially identical to their mothers even though not all mothers had the same diet. This suggests vertical transmission of diet by enhancement. Also, immatures selectively observed their mothers during extractive foraging, which increased goal‐directed practice but not general manipulation of similar objects, suggesting observational forms of learning of complex skills. Teaching was not observed. These results are consistent with the reported presence of food traditions and skill cultures in wild orangutans. We suggest that food traditions can develop wherever association commonly allows for social learning. However, the capacity for observational learning, and thus more complex culture, is more likely to evolve among extractive foragers with prolonged association between adults and immatures. Am. J. Primatol. 72:62–71, 2010.

Tolerant food sharing and reciprocity is precluded by despotism among bonobos but not chimpanzees.

Tolerant food sharing among human foragers can largely be explained by reciprocity. In contrast, food sharing among chimpanzees and bonobos may not always reflect reciprocity, which could be explained by different dominance styles: in egalitarian societies reciprocity is expressed freely, while in more despotic groups dominants may hinder reciprocity. We tested the degree of reciprocity and the influence of dominance on food sharing among chimpanzees and bonobos in two captive groups. First, we found that chimpanzees shared more frequently, more tolerantly, and more actively than bonobos. Second, among chimpanzees, food received was the best predictor of food shared, indicating reciprocal exchange, whereas among bonobos transfers were mostly unidirectional. Third, chimpanzees had a shallower and less linear dominance hierarchy, indicating that they were less despotic than bonobos. This suggests that the tolerant and reciprocal sharing found in chimpanzees, but not bonobos, was made possible by the absence of despotism. To investigate this further, we tested the relationship between despotism and reciprocity in grooming using data from an additional five groups and five different study periods on the main groups. The results showed that i) all chimpanzee groups were less despotic and groomed more reciprocally than bonobo groups, and ii) there was a general negative correlation between despotism and grooming reciprocity across species. This indicates that an egalitarian hierarchy may be more common in chimpanzees, at least in captivity, thus fostering reciprocal exchange. We conclude that a shallow dominance hierarchy was a necessary precondition for the evolution of human-like reciprocal food sharing.

The evolution of food sharing in primates

The aim of this study is to explain the occurrence of food sharing across primates. Defined as the unresisted transfer of food, evolutionary hypotheses have to explain why possessors should relinquish food rather than keep it. While sharing with offspring can be explained by kin selection, explanations for sharing among unrelated adults are more controversial. Here we test the hypothesis that sharing occurs with social partners that have leverage over food possessors due to the opportunity for partner choice in other contexts. Thus, we predict that possessors should relinquish food to potential mates or allies, who could provide or withhold matings or coalitionary support in the future. We used phylogenetic analyses based on both maximum likelihood and Bayesian approaches in a sample of 68 primate species to test these predictions. The analyses strongly indicate that (1) sharing with offspring is predicted by the relative processing difficulty of the diet, as measured by the degree of extractive foraging, but not overall diet quality, (2) food sharing among adults only evolved in species already sharing with offspring, regardless of diet, and (3) male–female sharing co-evolved with the opportunity for female mate choice and sharing within the sexes with coalition formation. These results provide comparative support for the hypothesis that sharing is “traded” for matings and coalitionary support in the sense that these services are statistically associated and can thus be selected for. Based on this, we predict that sharing should occur in any species with opportunities for partner choice.

Reciprocity explains food sharing in humans and other primates independent of kin selection and tolerated scrounging: a phylogenetic meta-analysis

Helping, i.e. behaviour increasing the fitness of others, can evolve when directed towards kin or reciprocating partners. These predictions have been tested in the context of food sharing both in human foragers and non-human primates. Here, we performed quantitative meta-analyses on 32 independent study populations to (i) test for overall effects of reciprocity on food sharing while controlling for alternative explanations, methodological biases, publication bias and phylogeny and (ii) compare the relative effects of reciprocity, kinship and tolerated scrounging, i.e. sharing owing to costs imposed by others. We found a significant overall weighted effect size for reciprocity of r = 0.20–0.48 for the most and least conservative measure, respectively. Effect sizes did not differ between humans and other primates, although there were species differences in in-kind reciprocity and trade. The relative effect of reciprocity in sharing was similar to those of kinship and tolerated scrounging. These results indicate a significant independent contribution of reciprocity to human and primate helping behaviour. Furthermore, similar effect sizes in humans and primates speak against cognitive constraints on reciprocity. This study is the first to use meta-analyses to quantify these effects on human helping and to directly compare humans and other primates.

Begging for information: mother–offspring food sharing among wild Bornean orangutans

Transfer of solid food from mothers or other adults to dependent offspring is commonly observed in various primate species and both nutritional and informational benefits have been proposed to explain the function of such food sharing. Predictions from these hypotheses are tested using observational data on wild orangutans (Pongo pygmaeus wurmbii) at Tuanan, Central Borneo, Indonesia. In 1,145u2009hr of focal observation and 458 recorded food interactions between four pairs of females with offspring it was found that virtually all transfers were initiated by the offspring and that younger infants solicited food more often and did so for a greater variety of items than older offspring. All offspring primarily solicited food that was difficult to process, i.e., inaccessible to them. Furthermore, the amount of food solicitation was negatively correlated with ecological competence. Hence food sharing seemed to be related to an offsprings skill level, as suggested by the informational hypothesis. In contrast, offspring did not solicit high‐quality items more than low‐quality items and food sharing did not peak around the age of weaning, as predicted by the nutritional hypothesis. Mothers were usually passively tolerant, allowing offspring to take food but hardly ever provisioned. Parent–offspring conflict concerning food sharing was only observed well after weaning. Thus, by taking food directly from the mother, young orangutans were able to obtain information about the affordances and nutritional value of food items that were otherwise out of their reach and could familiarize themselves with the mothers diet. In species such as orangutans or other apes, characterized by a broad diet that requires extractive foraging, informational food transfer may be vital for an immature to acquire complex feeding skills and adult diet. Am. J. Primatol. 70:533–541, 2008.

Natural cooperators: Food sharing in humans and other primates

The study of cooperation is rich with theoretical models and laboratory experiments that have greatly advanced our knowledge of human uniqueness, but have sometimes lacked ecological validity. We therefore emphasize the need to tie discussions of human cooperation to the natural history of our species and its closest relatives, focusing on behavioral contexts best suited to reveal underlying selection pressures and evolved decision rules. Food sharing is a fundamental form of cooperation that is well‐studied across primates and is particularly noteworthy because of its central role in shaping evolved human life history, social organization, and cooperative psychology. Here we synthesize available evidence on food sharing in humans and other primates, tracing the origins of offspring provisioning, mutualism, trade, and reciprocity throughout the primate order. While primates may gain some benefits from sharing, humans, faced with more collective action problems in a risky foraging niche, expanded on primate patterns to buffer risk and recruit mates and allies through reciprocity and signaling, and established co‐evolving social norms of production and sharing. Differences in the necessity for sharing are reflected in differences in sharing psychology across species, thus helping to explain unique aspects of our evolved cooperative psychology.

Physical Activity and Modernization among Bolivian Amerindians

Background Physical inactivity is a growing public health problem, and the fourth leading risk factor for global mortality. Conversely, indigenous populations living traditional lifestyles reportedly engage in vigorous daily activity that is protective against non-communicable diseases. Here we analyze physical activity patterns among the Tsimane, forager-horticulturalists of Amazonian Bolivia with minimal heart disease and diabetes. We assess age patterns of adult activity among men and women, test whether modernization affects activity levels, and examine whether nascent obesity is associated with reduced activity. Methods and Findings A factorial method based on a large sample of behavioral observations was employed to estimate effects of age, sex, body mass index, and modernization variables on physical activity ratio (PAR), the ratio of total energy expenditure to basal metabolic rate. Accelerometry combined with heart rate monitoring was compared to the factorial method and used for nighttime sampling. Tsimane men and women display 24 hr physical activity level (PAL) of 2.02–2.15 and 1.73–1.85, respectively. Little time was spent “sedentary”, whereas most activity was light to moderate, rather than vigorous. Activity peaks by the late twenties in men, and declines thereafter, but remains constant among women after the early teens. Neither BMI, fat free mass or body fat percentage are associated with PAR. There was no negative effect of modernization on physical activity. Conclusions Tsimane display relatively high PALs typical of other subsistence populations, but of moderate intensity, and not outside the range of developed populations. Despite rapidly increasing socioeconomic change, there is little evidence that total activity has yet been affected. Overweight and obesity are more prevalent among women than men, and Spanish fluency is associated with greater obesity in women. The lack of cardiovascular disease among Tsimane is unlikely caused by activity alone; further study of diet, food intake and infectious disease is needed.

Men’s status and reproductive success in 33 nonindustrial societies: Effects of subsistence, marriage system, and reproductive strategy

Significance Much of human behavior results from a desire for social status. From an evolutionary perspective, answering the question of why we pursue status must consider how status affects reproduction, especially in nonindustrial societies with natural fertility. In a metaanalysis of 288 results from 33 nonindustrial populations, we find that status is significantly associated with men’s reproductive success, consistent with an evolved basis for status pursuit. Status hierarchies have changed dramatically throughout human history, yet we find that the association between status and reproductive success does not depend on subsistence category (foraging, horticulture, pastoralism, agriculture) or how status is measured. These findings suggest no significant increase in selection on status-enhancing traits with the domestication of plants and animals. Social status motivates much of human behavior. However, status may have been a relatively weak target of selection for much of human evolution if ancestral foragers tended to be more egalitarian. We test the “egalitarianism hypothesis” that status has a significantly smaller effect on reproductive success (RS) in foragers compared with nonforagers. We also test between alternative male reproductive strategies, in particular whether reproductive benefits of status are due to lower offspring mortality (parental investment) or increased fertility (mating effort). We performed a phylogenetic multilevel metaanalysis of 288 statistical associations between measures of male status (physical formidability, hunting ability, material wealth, political influence) and RS (mating success, wife quality, fertility, offspring mortality, and number of surviving offspring) from 46 studies in 33 nonindustrial societies. We found a significant overall effect of status on RS (r = 0.19), though this effect was significantly lower than for nonhuman primates (r = 0.80). There was substantial variation due to marriage system and measure of RS, in particular status associated with offspring mortality only in polygynous societies (r = −0.08), and with wife quality only in monogamous societies (r = 0.15). However, the effects of status on RS did not differ significantly by status measure or subsistence type: foraging, horticulture, pastoralism, and agriculture. These results suggest that traits that facilitate status acquisition were not subject to substantially greater selection with domestication of plants and animals, and are part of reproductive strategies that enhance fertility more than offspring well-being.

Evolving the neuroendocrine physiology of human and primate cooperation and collective action

While many hormones play vital roles in facilitating or reinforcing cooperative behaviour, the neurohormones underlying competitive and cooperative behaviours are largely conserved across all mammals. This raises the question of how endocrine mechanisms have been shaped by selection to produce different levels of cooperation in different species. Multiple components of endocrine physiology—from baseline hormone concentrations, to binding proteins, to the receptor sensitivity and specificity—can evolve independently and be impacted by current socio-ecological conditions or individual status, thus potentially generating a wide range of variation within and between species. Here, we highlight several neurohormones and variation in hormone receptor genes associated with cooperation, focusing on the role of oxytocin and testosterone in contexts ranging from parenting and pair-bonding to reciprocity and territorial defence. While the studies reviewed herein describe the current state of the literature with regard to hormonal modulators of cooperation and collective action, there is still a paucity of research on hormonal mechanisms that help facilitate large-scale collective action. We end by discussing several potential areas for future research.