Children cooperate more with in-group members than with out-group members in an iterated face-to-face Prisoner's Dilemma Game.

Adults are more likely to cooperate with in-group members than with out-group members in the context of social dilemmas, situations in which self-interest is in conflict with collective interest. This bias has the potential to profoundly shape human cooperation, and therefore it is important to understand when it emerges in development. Here we asked whether 6- to 9-year-old children (N = 146) preferentially cooperate with in-group members in the context of a well-studied social dilemma, the iterated Prisoner's Dilemma Game. We assigned children to minimal groups and paired them with unfamiliar same-age and same-gender peers. Consistent with our predictions, children were more likely to cooperate with in-group members than with out-group members in this minimal group context. This finding adds to the current literature on group bias in children's prosocial behavior by showing that it affects decision making in a context that calls on strategic cooperation. In addition, our analyses revealed an effect of gender, with girls more likely to cooperate than boys regardless of the group membership of their partner. Exploring this gender effect further, we found an interaction between gender and age across condition, with older girls showing less sensitivity to the group membership of their partner than younger girls and with older boys showing more sensitivity to the group membership of the partner than younger boys. Our findings suggest that risky cooperation in the face of social dilemmas is shaped by group bias during childhood, highlighting the potentially deeply rooted ties between cooperation and parochialism in humans.

Western lowland gorillas (Gorilla gorilla gorilla) do not show an aversion to inequity in a token exchange task.

Although individuals in some species refuse foods they normally accept if their partner receives a more preferred one, this is not true across all species. The cooperation hypothesis proposes that this species-level variability evolved because inequity aversion is a mechanism to identify situations in which cooperation is not paying off, and that species regularly observed cooperating should be more likely to be averse to inequity. To rule out other potential explanations of inequity aversion, we need to test the converse as well: species rarely observed cooperating, especially those phylogenetically close to more cooperative species, should be less likely to be inequity averse. To this end, we tested eight zoo-housed Western lowland gorillas (Gorilla gorilla gorilla) on a token exchange task in which subjects received either the same food reward or a less-preferred reward for the same or more effort than their partner, recording both refusals to participate in the exchange and refusals to accept the reward. Supporting the cooperation hypothesis, even with procedural differences across sessions, gorillas were significantly more likely to refuse in all conditions in which they received a low-value food reward after completing an exchange, regardless of what their partner received, suggesting that gorillas were not inequity averse, but instead would not work for a low-value reward. Additionally, gorillas were more likely to refuse later in the session; while the pattern of refusals remained unchanged after accounting for this, this suggests that species should be tested on as many trials as is practical.

Comparative performance of orangutans (Pongo spp.), gorillas (Gorilla gorilla gorilla), and drills (Mandrillus leucophaeus), in an ephemeral foraging task.

A goal of the comparative approach is to test a variety of species on the same task. Here, we examined whether the factors that helped capuchin monkeys improve their performance in a dichotomous choice task would generalize to three other primate species: orangutans, gorillas, and drill monkeys. In this task, subjects have access to two options, each resulting in an identical food, but one (the ephemeral option) is only available if it is chosen first, whereas the other one (the permanent option) is always available. Therefore, the food-maximizing solution is to choose the ephemeral option first, followed by the permanent option for an additional reward. On the original version (plate task), the options were discriminated by the color and pattern of the plates holding the food, while on two subsequent versions we used altered cues that we predicted would improve performance: (1) the color of the foods themselves (color task), which we hypothesized was relevant to primates, who choose foods rather than substrates on which foods are found when foraging, and (2) patterned cups covering the foods (cup task), which we hypothesized would help primates avoid the prepotent response associated with visible food. Like capuchins, all three species initially failed to solve the plate task. However, while orangutans improved their performance from the plate to the color task, they did not for the cup task, and only a few gorillas and no drills succeeded in either task. Unfortunately, our ability to interpret these data was obscured by differences in the subjects' level of experience with cognitive testing and practical constraints that precluded the use of completely identical procedures across species. Nonetheless, we consider what these results can tell us, and discuss the value of conducting studies across multiple sites despite unavoidable differences.

Author Correction: Children avoid inefficient but fair partners in a cooperative game.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Does nonbinding commitment promote children's cooperation in a social dilemma?

Communication is a powerful tool for promoting cooperation in adults and is considered one of the most important solutions to social dilemmas. One feature that makes communication particularly useful in cooperative contexts is that it allows people to advertise their intentions to partners. Some work suggests that adults cooperate more after making nonbinding commitments to cooperate (i.e., commitments they do not need to uphold) than when they are not allowed to communicate their intentions to their partners. However, we know little about whether nonbinding commitments play a similar role in children. We addressed this gap by testing 6- to 9-year-old children in a simultaneous version of the iterated prisoner's dilemma game. In the communication condition, children could communicate their intended decision prior to their actual choice, whereas in the silent condition, they could not communicate. Overall, children in the communication condition were no more likely to cooperate than children in the silent condition. However, in the communication condition, but not in the silent condition, children's behavior was contingent on their previous decision; they were more likely to cooperate or defect when they had previously cooperated or defected, respectively. In addition, they rarely reversed their intended decisions in the game. Our findings suggest that, although nonbinding commitment does not promote children's cooperation in general, it may encourage children to stick to their chosen strategy, perhaps for the sake of appearing consistent. More broadly, these results contribute to our emerging understanding of the ways in which children solve social dilemmas.

Children avoid inefficient but fair partners in a cooperative game.

Human adults use a range of social cues to obtain information about potential partners in cooperative contexts: we prefer partners who are competent, wealthy and generous, and those who abide by moral and social rules. One factor that carries particular weight is whether a prospective partner is fair. Here we ask whether children share this preference for fair partners and, if so, whether a prospective partner's past fair behaviour influences children's behaviour in a cooperative dilemma. Six- to nine-year-olds chose between partners who accepted or rejected resource allocations that were either strongly advantageously unequal, strongly disadvantageously unequal, or equal. They then played a one-shot Prisoner's Dilemma Game with their chosen partner. Children overwhelmingly preferred to play with the partner who accepted rather than rejected allocations. Regardless of their partner choice decisions, children tended to defect in the Prisoner's Dilemma Game, yet expected that their partners would be relatively more cooperative. Finally, children were more likely to cooperate with those they believed would cooperate. Together, these findings shed new light on the links between partner choice, fairness and cooperation in child development.

Capuchin monkeys (Cebus [sapajus] apella) show planning in a manual maze task.

The capacity for planning in nonhuman species has long been an interest of many comparative and cognitive psychologists. There is now considerable evidence that at least great apes show both motor planning and planning for future needs in various contexts and modalities. Few studies, however, have investigated planning ability in a monkey species, and most of these exceptions have used computerized procedures. To gain a broader view, in the current study, we tested capuchin monkeys' capacity for motor planning using the "paddle-box" apparatus, a manual maze task originally designed for testing planning skills in apes (Tecwyn, Thorpe, & Chappell, 2013). This consisted of a box containing different levels of paddles that subjects rotated to drop food into a tube that delivered it to the subject. To evaluate subjects' degree of skill in the task, the initial locations of the food and the final goal (i.e., the "paths" the food could take) were chosen according to the probability of retrieving the reward, starting with those that gave the highest probability of success and moving sequentially to those with the lowest probability. Most subjects solved all levels of difficulty in the task, and capuchin monkeys succeeded as a group in a generalization test with novel paths. These findings demonstrate that some monkeys, like apes, show planning in different contexts and modalities. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Eliminating the VGlut2-Dependent Glutamatergic Transmission of Parvalbumin-Expressing Neurons Leads to Deficits in Locomotion and Vocalization, Decreased Pain Sensitivity, and Increased Dominance.

The calcium-binding protein parvalbumin (PV) is a recognized marker of short-axon GABA-ergic neurons in the cortex and the hippocampus. However in addition, PV is expressed by excitatory, glutamatergic neurons in various areas of the brain and spinal cord. Depending on the location of these neurons, loading of their synaptic vesicles with glutamate is mediated by either of three vesicular glutamate transporters (VGlut): VGlut1, VGlut2, or VGlut3. Driven by our interest in one of these glutamatergic/PV-expressing cell clusters-the lateral hypothalamic parvafox nucleus-we investigated the functions of this population of neurons by the selective deletion of VGlut2 expression in PV-expressing cells according to the Cre/Lox-approach. PV-Cre;VGlut2-Lox mutant mice are phenotypically characterized by deficits in locomotion and vocalization, by a decreased thermal nociception, and by an increased social dominance. We conducted a search of the Allen Brain Atlas for regions that might co-express the genes encoding PV and VGlut2, and that might thus contribute to the manifestation of the observed phenotypes. Our survey revealed several structures that could contribute to the deficits in locomotion and vocalization, such as the red, the subthalamic and the deep cerebellar nuclei. It also disclosed that a shift in the balance of afferental glutamatergic neurotransmission to the periaqueductal gray matter might be accountable for the decrease in sensitivity to pain and for the increase in social dominance. As a whole, this study broadens the state of knowledge about PV-expressing excitatory neurons.

Comparing species decisions in a dichotomous choice task: adjusting task parameters improves performance in monkeys.

In comparative psychology, both similarities and differences among species are studied to better understand the evolution of their behavior. To do so, we first test species in tasks using similar procedures, but if differences are found, it is important to determine their underlying cause(s) (e.g., are they due to ecology, cognitive ability, an artifact of the study, and/or some other factor?). In our previous work, primates performed unexpectedly poorly on an apparently simple two-choice discrimination task based on the natural behavior of cleaner fish, while the fish did quite well. In this task, if the subjects first chose one of the options (ephemeral) they received both food items, but if they chose the other (permanent) option first, the ephemeral option disappeared. Here, we test several proposed explanations for primates' relatively poorer performance. In Study 1, we used a computerized paradigm that differed from the previous test by removing interaction with human experimenters, which may be distracting, and providing a more standardized testing environment. In Study 2, we adapted the computerized paradigm from Study 1 to be more relevant to primate ecology. Monkeys' overall performance in these adapted tasks matched the performance of the fish in the original study, showing that with the appropriate modifications they can solve the task. We discuss these results in light of comparative research, which requires balancing procedural similarity with considerations of how the details of the task or the context may influence how different species perceive and solve tasks differently.

Using photographs to study animal social cognition and behaviour: Do capuchins' responses to photos reflect reality?

Behavioural responses to photos are often used to infer what animals understand about their social environment, but are rarely validated against the same stimuli in real life. If subjects' responses to photos do not reflect responses to the same live stimuli, it is difficult to conclude what happens in reality based on photo responses alone. We compared capuchins' responses to photos versus live stimuli in an identical scenario within research cubicles. Subjects had the opportunity to approach food placed in front of an alpha group member and, in a separate condition, photos depicting the same individual. Subjects' latencies to approach food when placed in front of the real alpha negatively correlated with time subjects spent in close proximity to the alpha in their main enclosure. We therefore predicted subjects' latencies to approach food in the presence of photos would positively correlate with their latencies to approach food in the presence of the real alpha inside the cubicles, but negatively correlate with time they spent in proximity to the alpha in their enclosure. Neither prediction was supported. While not necessarily surprising, we explain why these results should be an important reminder that care is needed when interpreting results from photo studies.

Adult cleaner wrasse outperform capuchin monkeys, chimpanzees and orang-utans in a complex foraging task derived from cleaner--client reef fish cooperation.

The insight that animals' cognitive abilities are linked to their evolutionary history, and hence their ecology, provides the framework for the comparative approach. Despite primates renowned dietary complexity and social cognition, including cooperative abilities, we here demonstrate that cleaner wrasse outperform three primate species, capuchin monkeys, chimpanzees and orang-utans, in a foraging task involving a choice between two actions, both of which yield identical immediate rewards, but only one of which yields an additional delayed reward. The foraging task decisions involve partner choice in cleaners: they must service visiting client reef fish before resident clients to access both; otherwise the former switch to a different cleaner. Wild caught adult, but not juvenile, cleaners learned to solve the task quickly and relearned the task when it was reversed. The majority of primates failed to perform above chance after 100 trials, which is in sharp contrast to previous studies showing that primates easily learn to choose an action that yields immediate double rewards compared to an alternative action. In conclusion, the adult cleaners' ability to choose a superior action with initially neutral consequences is likely due to repeated exposure in nature, which leads to specific learned optimal foraging decision rules.