Chimpanzees and bonobos use social leverage in an ultimatum game.

The ultimatum game (UG) is widely used to investigate our sense of fairness, a key characteristic that differentiates us from our closest living relatives, bonobos and chimpanzees. Previous studies found that, in general, great apes behave as rational maximizers in the UG. Proposers tend to choose self-maximizing offers, while responders accept most non-zero offers. These studies do not rule out the possibility that apes can behave prosocially to improve the returns for themselves and others. However, this has never been well studied. In this study, we offer chimpanzee and bonobo proposers the possibility of taking into account the leverage of responders over the offers they receive. This leverage takes the form of access to alternatives for responders. We find that proposers tend to propose fairer offers when responders have the option to access alternatives. Furthermore, we find that both species use their leverage to reject unequal offers. Our results suggest that great apes mostly act as rational maximizers in an UG, yet access to alternatives can lead them to change their strategies such as not choosing the self-maximizing offer as proposers and not accepting every offer higher than zero as responders.

Gibbons exploit information about what a competitor can see.

How much nonhuman animals understand about seeing has been the focus of comparative cognition research for decades. Many social primates (and other species) are sensitive to cues about what others can and cannot see. Whether this sensitivity evolved in primates through shared descent or convergent evolution remains unclear. The current study tested gibbons-the apes that are least studied yet most distantly related to humans and one of the less social primates-in two food-competition tasks. Specifically, we presented eastern hoolock gibbons, Hoolock leuconedys, and silvery gibbons, Hylobates moloch, with a choice between a contested piece of food visible to both themselves and a human competitor and an uncontested piece visible only to themselves. Subjects successfully stole the uncontested food when the competitor turned away his body (N = 10, experiment 1) and his head (N = 9, experiment 2). However, when the head of the experimenter was oriented towards the contested piece of food, whether the competitor opened or closed his eyes made no difference. Subjects' sensitivity to body- and head-orientation cues was comparable to that of chimpanzees, rhesus macaques, and ring-tailed lemurs-species living in much larger groups than gibbons. These findings support the continuity hypothesis that sensitivity to body- and head-orientation cues is a product of shared descent among primates.

Do 7-year-old children understand social leverage?

Individuals with an advantageous position during a negotiation possess leverage over their partners. Several studies with adults have investigated how leverage can influence the coordination strategies of individuals when conflicts of interest arise. In this study, we explored how pairs of 7-year-old children solved a coordination game (based on the Snowdrift scenario) when one child had leverage over the other child. We presented a social dilemma in the form of an unequal reward distribution on a rotating tray. The rotating tray could be accessed by both children. The child who waited longer to act received the best outcome, but if both children waited too long, they would lose the rewards. In addition, one child could forgo the access to the rotating tray for an alternative option-the leverage. Although children rarely used their leverage strategically, children with access to the alternative were less likely to play the social dilemma, especially when their leverage was larger. Furthermore, children waited longer to act as the leverage decreased. Finally, children almost never failed to coordinate. The results hint to a trade-off between maximizing benefits while maintaining long-term collaboration in complex scenarios where strategies such as turn taking are hard to implement.

Chimpanzees, bonobos and children successfully coordinate in conflict situations.

Social animals need to coordinate with others to reap the benefits of group-living even when individuals' interests are misaligned. We compare how chimpanzees, bonobos and children coordinate their actions with a conspecific in a Snowdrift game, which provides a model for understanding how organisms coordinate and make decisions under conflict. In study 1, we presented pairs of chimpanzees, bonobos and children with an unequal reward distribution. In the critical condition, the preferred reward could only be obtained by waiting for the partner to act, with the risk that if no one acted, both would lose the rewards. Apes and children successfully coordinated to obtain the rewards. Children used a 'both-partner-pull' strategy and communicated during the task, while some apes relied on an 'only-one-partner-pulls' strategy to solve the task, although there were also signs of strategic behaviour as they waited for their partner to pull when that strategy led to the preferred reward. In study 2, we presented pairs of chimpanzees and bonobos with the same set-up as in study 1 with the addition of a non-social option that provided them with a secure reward. In this situation, apes had to actively decide between the unequal distribution and the alternative. In this set-up, apes maximized their rewards by taking their partners' potential actions into account. In conclusion, children and apes showed clear instances of strategic decision-making to maximize their own rewards while maintaining successful coordination.

Chimpanzees engage in competitive altruism in a triadic ultimatum game.

Partner choice promotes competition among individuals to be selected as a cooperative partner, a phenomenon referred to as competitive altruism. We explored whether chimpanzees engage in competitive altruism in a triadic Ultimatum Game where two proposers can send offers simultaneously or consecutively to a responder who can only accept one of the two competing offers. In a dyadic control condition only one proposer at a time could send an offer to the responder. Chimpanzees increased their offers across trials in the competitive triadic, but not in the dyadic control condition. Chimpanzees also increased their offers after being rejected in previous triadic trials. Furthermore, we found that chimpanzees, under specific conditions, outcompete first proposers in triadic consecutive trials before the responder could choose which offer to accept by offering more than what is expected if they acted randomly or simply offered the smallest possible amount. These results suggest that competitive altruism in chimpanzees did not emerge just as a by-product of them trying to increase over previous losses. Chimpanzees might consider how others' interactions affect their outcomes and engage in strategies to maximize their chances of being selected as cooperative partners.

Functional fixedness in chimpanzees.

Differences in the tool use of non-human primates and humans are subject of ongoing debate. In humans, representations of object functions underpin efficient tool use. Yet, representations of object functions can lead to functional fixedness, which describes the fixation on a familiar tool function leading to less efficient problem solving when the problem requires using the tool for a new function. In the current study, we examined whether chimpanzees exhibit functional fixedness. After solving a problem with a tool, chimpanzees were less efficient in solving another problem which required using the same tool with a different function compared to a control group. This fixation effect was still apparent after a period of nine months and when chimpanzees had learned about the function of a tool by observation of a conspecific. These results suggest that functional fixedness in our closest living relatives likely exists and cast doubt on the notion that stable function representations are uniquely human.

Comparative curiosity: How do great apes and children deal with uncertainty?

Humans are perhaps the most curious animals on earth, but to what extent our innate motivations for discovering new information are shared with our closest relatives remain poorly understood. To shed light on this question, we presented great apes with two experimental paradigms in which they had to initially choose between an empty opaque cup and a baited opaque cup with rewards invisible to the ape in study 1, or to choose between a transparent cup with rewards or a baited opaque cup with rewards invisible to the ape in studies 2 and 3. We also presented young children with scenarios comparable to the second paradigm (studies 4 and 5). Notably, after the initial choice phase, we presented participants with potential alternatives providing better rewards than the previously secured options. Importantly, those alternatives shared some features with the uncertain options, giving subjects the possibility to relate both options through analogical reasoning. We found that most great apes were not curious about the uncertain options. They only explored those options after they were presented with the alternatives. Children, instead, explored the uncertain options before the alternatives were presented, showing a higher degree of curiosity than the great apes. We argue that differences between children and apes mostly lay in motivational dispositions to explore the unknown.

Gibbon strategies in a food competition task.

Social primates face conflicts of interest with other partners when their individual and collective interests collide. Despite living in small, primarily bonded, groups compared to other social primates, gibbons are not exempt from these conflicts in their everyday lives. In the current task, we asked whether dyads of gibbons would solve a conflict of interest over food rewards. We presented dyads of gibbons with a situation in which they could decide whether to take an active role and pull a handle to release food rewards at a distance or take a passive role and avoid action. In this situation, the passive partner could take an advantageous position to obtain the rewards over the active partner. Gibbons participated in three conditions: a control condition with no food rewards, a test condition with indirect food rewards and a test condition with direct food rewards. In both test conditions, five rewards were released at a distance from the handle. In addition, the active individual could obtain one extra food reward from the handle in the direct food condition. We found that gibbons acted more often in the two conditions involving food rewards, and waited longer in the indirect compared to the direct food condition, thus suggesting that they understood the task contingencies. Surprisingly, we found that in a majority of dyads, individuals in the active role obtained most of the payoff compared to individuals in the passive role in both food conditions. Furthermore, in some occasions individuals in the active role did not approach the location where the food was released. These results suggest that while gibbons may strategize to maximize benefits in a competitive food task, they often allowed their partners to obtain better rewards. Our results highlight the importance of social tolerance and motivation as drivers promoting cooperation in these species.

How environmental unpredictability and harshness affect chimpanzees (Pan troglodytes) in risk-choice and temporal discounting tasks.

Animals need to adjust their decision-making strategies to the ecological challenges of their environment. Variation in ecological unpredictability and harshness thus seem to affect their decisions in the wild. In our study, we combine methods from human life history theory and previous comparative work with chimpanzees to investigate whether current variability in ecological factors such as reward unpredictability and harshness affect chimpanzees' decision-making strategies in two value-based scenarios. We presented chimpanzees with choices varying in the probability of obtaining food rewards (risk-choice task) and in their temporal availability (temporal discounting task). These scenarios were preceded by different sets of priming phases mimicking variability in resource unpredictability (predictable or unpredictable rewards) and harshness (abundant or scarce rewards). In addition, we implemented a social manipulation to explore whether variations in unpredictable gains and losses affected chimpanzees' performance in both tasks. We found that chimpanzees were only affected by the social manipulation in the risk-choice scenario. Specifically, after a period of constant food losses chimpanzees became less risk prone. We discuss how different types of negative experiences affect chimpanzees' decisions. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Establishing an infrastructure for collaboration in primate cognition research.

Inferring the evolutionary history of cognitive abilities requires large and diverse samples. However, such samples are often beyond the reach of individual researchers or institutions, and studies are often limited to small numbers of species. Consequently, methodological and site-specific-differences across studies can limit comparisons between species. Here we introduce the ManyPrimates project, which addresses these challenges by providing a large-scale collaborative framework for comparative studies in primate cognition. To demonstrate the viability of the project we conducted a case study of short-term memory. In this initial study, we were able to include 176 individuals from 12 primate species housed at 11 sites across Africa, Asia, North America and Europe. All subjects were tested in a delayed-response task using consistent methodology across sites. Individuals could access food rewards by remembering the position of the hidden reward after a 0, 15, or 30-second delay. Overall, individuals performed better with shorter delays, as predicted by previous studies. Phylogenetic analysis revealed a strong phylogenetic signal for short-term memory. Although, with only 12 species, the validity of this analysis is limited, our initial results demonstrate the feasibility of a large, collaborative open-science project. We present the ManyPrimates project as an exciting opportunity to address open questions in primate cognition and behaviour with large, diverse datasets.

Chimpanzees' understanding of social leverage.

Social primates can influence others through the control of resources. For instance, dominant male chimpanzees might allow subordinates access to mate with females in exchange for social support. However, little is known about how chimpanzees strategically use a position of leverage to maximize their own benefits. We address this question by presenting dyads of captive chimpanzee (N = 6) with a task resulting in an unequal reward distribution. To gain the higher reward each individual should wait for their partner to act. In addition, one participant had leverage: access to an alternative secure reward. By varying the presence and value of the leverage we tested whether individuals used it strategically (e.g. by waiting longer for partners to act when they had leverage in the form of alternatives). Additionally, non-social controls served to show if chimpanzees understood the social dilemma. We measured the likelihood to choose the leverage and their latencies to act. The final decision made by the chimpanzees did not differ as a function of condition (test versus non-social control) or the value of the leverage, but they did wait longer to act when the leverage was smaller-particularly in test (versus non-social control) trials suggesting that they understood the conflict of interest involved. The chimpanzees thus recognized the existence of social leverage, but did not use it strategically to maximize their rewards.

Differences in between-reinforcer value modulate the selective-value effect in great apes (Pan troglodytes, P. Paniscus, Gorilla gorilla, Pongo abelii).

We investigated how apes allocated their choices between 2 food options that varied in terms of their quantity and quality. Experiment 1 tested whether subjects preferred an AB option over an A option, where the A item is preferred to the B item (e.g., apple + carrot vs. apple). Additionally, we tested whether the length of the intertrial interval (ITI) affected subjects' choices. Five orangutans, 4 gorillas, 7 bonobos, and 10 chimpanzees received 3 types of trials: preference (A vs. B), quantity (AA vs. A), and mixed (AB vs. A where A is the preferred food). We used 3 food items that substantially differed in terms of preference (carrots, apples, and pellets). Subjects showed no overall preference for the mixed option (AB) compared with the single option (A), even though they showed clear preferences during both the preference and quantity trials. The intertrial length had no effect on choice behavior. Experiment 2 further explored apes' choices by using 3 highly preferred food items (bananas, grapes, and pellets) in 6 orangutans, 4 gorillas, 8 bonobos, and 18 chimpanzees. Unlike the results of Experiment 1, apes generally chose the mixed option. Our results indicated that apes did not show a general "selective-value" effect but chose depending on the relative value of the food items involved. Subjects were more likely to select the mixed over the single option when (a) the mixed option was composed of items that were closer in value and (b) they were compared against the less valuable item forming the mixed option.