When Given the Opportunity, Chimpanzees Maximize Personal Gain Rather than “Level the Playing Field”

We provided chimpanzees (Pan troglodytes) with the ability to improve the quality of food rewards they received in a dyadic test of inequity.We were interested to see if this provision influenced their responses and, if so, whether it was mediated by a social partner’s outcomes. We tested eight dyads using an exchange paradigm in which, depending on the condition, the chimpanzees were rewarded with either high-value (a grape) or low-value (a piece of celery) food rewards for each completed exchange. We included four conditions. In the first, “Different” condition, the subject received different, less-preferred, rewards than their partner for each exchange made (a test of inequity). In the “Unavailable” condition, high-value rewards were shown, but not given, to both chimpanzees prior to each exchange and the chimpanzees were rewarded equally with low-value rewards (a test of individual contrast). The final two conditions created equity. In these High-value and Low-value “Same” conditions both chimpanzees received the same food rewards for each exchange.Within each condition, the chimpanzees first completed ten trials in the Baseline Phase, in which the experimenter determined the rewards they received, and then ten trials in the Test Phase. In the Test Phase, the chimpanzees could exchange tokens through the aperture of a small wooden picture frame hung on their cage mesh in order to receive the high-value reward. Thus, in the Test Phase, the chimpanzees were provided with an opportunity to improve the quality of the rewards they received, either absolutely or relative to what their partner received. The chimpanzees responded in a targeted manner; in the Test Phase they attempted to maximize their returns in all conditions in which they had received low-value rewards during the Baseline Phase. Thus, the chimpanzees were apparently motivated to increase their reward regardless of their partners’, but they only used the mechanism provided when it afforded the opportunity for them to increase their rewards.We also found evidence that the chimpanzees’ responses were enhanced by social facilitation. Specifically, the chimpanzees were more likely to exchange their tokens through the frame when their test partner also did so, even in circumstances in which their reward value could not be improved. Our paradigm provided the chimpanzees with the possibility to improve the quality of rewards they received in the Test Phase. We found that refusals – to exchange tokens or to eat rewards – decreased significantly in the Test Phase compared to the Baseline Phase, where no such opportunity for improvement of outcomes existed. Thus, the chimpanzees participated more when they could improve the rewards they received

Chimpanzees’ Socially Maintained Food Preferences Indicate both Conservatism and Conformity

Chimpanzees remain fixed on a single strategy, even if a novel, more efficient, strategy is introduced. Previous studies reporting such findings have incorporated paradigms in which chimpanzees learn one behavioural method and then are shown a new one that the chimpanzees invariably do not adopt. This study provides the first evidence that chimpanzees show such conservatism even when the new method employs the identical required behaviour as the first, but for a different reward. Groups of chimpanzees could choose to exchange one of two inedible tokens; one was rewarded with a highly preferred food (grape) and the other with a less preferred food (carrot). Individuals first observed a model chimpanzee from their social group trained to choose one of the two types of tokens. In one group, this token earned a carrot, while in the other, control, group the token earned a grape. In both groups, chimpanzees conformed to the trained model’s choice. This was especially striking for those gaining the pieces of carrot; the less favoured reward. This resulted in a population-level trend of food choices, even when counter to their original, individual, preferences. Moreover, the chimpanzees’ food preferences did not change over time, demonstrating that these results were not due to a simple shift in preferences. We discuss social factors apparent in the interactions and suggest that, despite seeming to be inefficient, in chimpanzees, conformity may benefit them, possibly by assisting with the maintenance of group relations

The Ontogeny of Social Comparisons by Rhesus Macaques (Macaca mulatta)

This longitudinal study investigated the development of social contrast-negative responses to inequitable rewards-in rhesus macaques (Macaca mulatta). Although responses to inequity by humans appear universal, this is something that develops with age. Infants first recognize inequity when around 18 months old and respond to it only when they are around 3 years old. To date, however, there have been no studies of the ontogeny of the inequity response in any species other than humans. To address this, we used an exchange paradigm, in which 10 pairs of rhesus monkeys had to exchange inedible tokens with the experimenter to get food rewards that differed in quality depending on the condition. All subjects were tested first when they were an average of 17 months old and a subset, of four pairs, was tested again a year later. Subjects responded negatively to contrast-recognizing a disparity in expected, as compared to, received rewards-based on both social and individual comparisons at the older age, but not at the younger age. Similar to humans, rhesus showed a developmental trajectory to social comparison, providing the first evidence for the ontogeny of this response in a non-human species

Dissecting the Mechanisms of Squirrel Monkey (Saimiri boliviensis) Social Learning

Although the social learning abilities of monkeys have been well documented, this research has only focused on a few species. Furthermore, of those that also incorporated dissections of social learning mechanisms, the majority studied either capuchins (Cebus apella) or marmosets (Callithrix jacchus). To gain a broader understanding of how monkeys gain new skills, we tested squirrel monkeys (Saimiri boliviensis) which have never been studied in tests of social learning mechanisms. To determine whether S. boliviensis can socially learn, we ran “open diffusion” tests with monkeys housed in two social groups (N = 23). Over the course of 10 20-min sessions, the monkeys in each group observed a trained group member retrieving a mealworm from a bidirectional task (the “Slide-box”). Two thirds (67%) of these monkeys both learned how to operate the Slide-box and they also moved the door significantly more times in the direction modeled by the trained demonstrator than the alternative direction. To tease apart the underlying social learning mechanisms we ran a series of three control conditions with 35 squirrel monkeys that had no previous experience with the Slide-box. The first replicated the experimental open diffusion sessions but without the inclusion of a trained model, the second was a no-information control with dyads of monkeys, and the third was a ‘ghost’ display shown to individual monkeys. The first two controls tested for the importance of social support (mere presence effect) and the ghost display showed the affordances of the task to the monkeys. The monkeys showed a certain level of success in the group control (54% of subjects solved the task on one or more occasions) and paired controls (28% were successful) but none were successful in the ghost control.We propose that the squirrel monkeys’ learning, observed in the experimental open diffusion tests, can be best described by a combination of social learning mechanisms in concert; in this case, those mechanisms are most likely object movement reenactment and social facilitation. We discuss the interplay of these mechanisms and how they related to learning shown by other primate species

The Endowment Effect in Orangutans

The endowment effect is the tendency to, seemingly irrationally, immediately value a possessed item more than the opportunity to acquire the identical item when one does not already possess it. Although endowment effects are reported in chimpanzees (Brosnan, Jones, Lambeth, Mareno, Richardson, & Shapiro, 2007) and capuchin monkeys (Lakshminarayanan, Chen, & Santos, 2008), both species share social traits with humans that make convergence as likely an evolutionary mechanism as homology. Orangutans (Pongo spp.) provide a unique insight into the evolution of the endowment effect, along with other apparently irrational behaviors, because their less frequent social interactions and relatively more solitary social organization distinguishes them from the more gregarious apes, allowing a test of evolutionary homology. In the present study, we used pairs of both food and non-food objects, as in an earlier test on chimpanzees (Brosnan et al., 2007). We established the apes’ preferences in forced-choice tasks, then tested whether they showed an endowment effect in an exchange task, in which subjects were given one of the objects, followed by the option to exchange it for the other. Here, we report the first evidence of the endowment effect in a relatively less social primate, the orangutan. This indicates that this behavior may have evolved as a homology within the primates, rather than being due to convergent social pressures. These findings provide stronger evidence for the hypothesis that at least one bias, the endowment effect, may be common in primates and, potentially, other species

Generous Leaders and Selfish Underdogs: Pro-Sociality in Despotic Macaques

Actively granting food to a companion is called pro-social behavior and is considered to be part of altruism. Recent findings show that some non-human primates behave pro-socially. However, pro-social behavior is not expected in despotic species, since the steep dominance hierarchy will hamper pro-sociality. We show that some despotic long-tailed macaques do grant others access to food. Moreover, their dominance hierarchy determines pro-social behavior in an unexpected way: high-ranking individuals grant, while low-ranking individuals withhold their partner access to food. Surprisingly, pro-social behavior is not used by subordinates to obtain benefits from dominants, but by dominants to emphasize their dominance position. Hence, Machiavellian macaques rule not through “fear above love”, but through “be feared when needed and loved when possible”

Anything for a Cheerio: Brown Capuchins (\u3cem\u3eSapajus [Cebus] apella\u3c/em\u3e) Consistently Coordinate in an Assurance Game for Unequal Payoffs

Unequal outcomes disrupt cooperation in some situations, but this has not been tested in the context of coordination in economic games. To explore this, we tested brown capuchins (Sapajus [Cebus] apella) on a manual version of the Stag Hunt (or Assurance) Game, in which individuals sequentially chose between two options, Stag or Hare, and were rewarded according to their choices and that of their partner. Typically, coordination on Stag results in an equal highest payout, whereas coordinating on Hare results in a guaranteed equal but lower payoff and uncoordinated play results in the lowest payoff when playing Stag. We varied this structure such that one capuchin received double the rewards for the coordinated Stag outcome; thus, it was still both animals\u27 best option, but no longer equally rewarding. Despite the inequality, capuchins coordinated on Stag in 78% of trials, and neither payoff structure nor their partner\u27s choice impacted their decision. Additionally, there was no relationship between self-scratching, a measure of stress in capuchins, and choices. After completing the study, we discovered our reward, cheerios, was sufficiently valuable that in another study, capuchins never refused it, so post hoc we repeated the study using a lower value reward, banana flavored pellets. Capuchins completed only 26% of the pellet trials (compared to 98% with cheerios), constraining our ability to interpret the results, but nonetheless the monkeys showed a decrease in preference for Stag, particularly when they received fewer rewards for the coordinated Stag outcome. These results reinforce capuchins\u27 ability to find coordinated outcomes in the Stag Hunt game, but more work is needed to determine whether the monkeys did not mind the inequality or were unwilling to sacrifice a highly preferred food to rectify it. In either case, researchers should carefully consider the impact of their chosen rewards on subjects\u27 choices

How Is Chimpanzee Self-Control Influenced by Social Setting?

Self-control is often required in natural situations involving interactions with other individuals, and personal self-control can be compromised if other individuals act impulsively. In this study, we tested self-control in pairs of chimpanzees in a variety of settings where at least one chimpanzee of each pair performed an established test for self-control in which candies accumulated one at time as long as the chimpanzee did not eat any of them. When tested alone, some chimpanzees exhibited greater self-control as compared to when tested alongside a chimpanzee that independently performed the same type of test. However, when the nonfocal animal freely consumed rewards while the focal chimpanzee performed the accumulation task, the self-control of some focal chimpanzees was elevated as compared to when working alone. Finally, when the focal and nonfocal animals worked jointly on the same test and the number of rewards accumulated was dependent on both animals’ continued ability to inhibit eating the items, chimpanzees performed the same when housed together or in adjacent enclosures. On the whole, the effects of social setting were modest, but these results may relate to the literature on vicarious depletion of self-control, and they present interesting avenues for future research

Chimpanzee Autarky

Background: Economists believe that barter is the ultimate cause of social wealth—and even much of our human culture—yet little is known about the evolution and development of such behavior. It is useful to examine the circumstances under which other species will or will not barter to more fully understand the phenomenon. Chimpanzees (Pan troglodytes) are an interesting test case as they are an intelligent species, closely related to humans, and known to participate in reciprocal interactions and token economies with humans, yet they have not spontaneously developed costly barter. Methodology/Principle Findings: Although chimpanzees do engage in noncostly barter, in which otherwise value-less tokens are exchanged for food, this lack of risk is not typical of human barter. Thus, we systematically examined barter in chimpanzees to ascertain under what circumstances chimpanzees will engage in costly barter of commodities, that is, trading food items for other food items with a human experimenter. We found that chimpanzees do barter, relinquishing lower value items to obtain higher value items (and not the reverse). However, they do not trade in all beneficial situations, maintaining possession of less preferred items when the relative gains they stand to make are small. Conclusions/Significance: Two potential explanations for this puzzling behavior are that chimpanzees lack ownership norms, and thus have limited opportunity to benefit from the gains of trade, and that chimpanzees\u27 risk of defection is sufficiently high that large gains must be imminent to justify the risk. Understanding the conditions that support barter in chimpanzees may increase understanding of situations in which humans, too, do not maximize their gains