Social influences on ant-dipping acquisition in the wild chimpanzees (Pan troglodytes verus) of Bossou,Guinea, West Africa

We currently have little understanding of the influence of learning opportunity, whether social or environmental, and maternal role on tool-use acquisition in young wild chimpanzees. This study aims to fill this gap by focusing on the acquisition of ant-dipping among chimpanzees of Bossou, Guinea. Ant-dipping is a hazardous tool-use behaviour aimed at army ants (Dorylus spp.). Bossou chimpanzees target these ants both at nests (high risk) and trails (low risk) and employ two techniques to consume them: direct mouthing and pull-through. We present data for 13 mother–offspring pairs (1–10 years old). Mothers with young ≤5 years old dipped significantly more often at trails than at nests, thus minimizing the risk posed to themselves and their young. Infants thus benefited from better conditions to observe and practice ant-dipping. Mothers also varied greatly in their percent time spent ant-dipping and offspring differed in their learning opportunity. Our results suggest that high opportunity young started to observe and perform ant-dipping sooner and were better at ant-dipping than low opportunity young. Although mothers and weaned offspring correlated positively in their percent time spent dipping and proficiency, they did not match in technique used or tool length. Finally, we propose that the learning trajectory of young may predict individual and sex differences in adulthood. This study demonstrates the important role of mothers and learning opportunity in the acquisition of a hazardous tool-use behaviour and suggests that chimpanzee material culture is a product of a complex interaction between social processes and ecological factors.     

Chimpanzees prioritise social information over pre-existing behaviours in a group context but not in dyads

How animal communities arrive at homogeneous behavioural preferences is a central question for studies of cultural evolution. Here, we investigated whether chimpanzees (Pan troglodytes) would relinquish a pre-existing behaviour to adopt an alternative demonstrated by an overwhelming majority of group mates; in other words, whether chimpanzees behave in a conformist manner. In each of five groups of chimpanzees (N?=?37), one individual was trained on one method of opening a two-action puzzle box to obtain food, while the remaining individuals learned the alternative method. Over 5 h of open access to the apparatus in a group context, it was found that 4/5 ‘minority’ individuals explored the majority method and three of these used this new method in the majority of trials. Those that switched did so after observing only a small subset of their group, thereby not matching conventional definitions of conformity. In a further ‘Dyad’ condition, six pairs of chimpanzees were trained on alternative methods and then given access to the task together. Only one of these individuals ever switched method. The number of observations that individuals in the minority and Dyad individuals made of their untrained method was not found to influence whether or not they themselves switched to use it. In a final ‘Asocial’ condition, individuals (N?=?10) did not receive social information and did not deviate from their first-learned method. We argue that these results demonstrate an important influence of social context upon prioritisation of social information over pre-existing methods, which can result in group homogeneity of behaviour.     

Conservatism and “copy-if-better” in chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Social learning is predicted to evolve in socially living animals provided the learning process is not random but biased by certain socio-ecological factors. One bias of particular interest for the emergence of (cumulative) culture is the tendency to forgo personal behaviour in favour of relatively better variants observed in others, also known as the “copy-if-better” strategy. We investigated whether chimpanzees employ copy-if-better in a simple token-exchange paradigm controlling for individual and random social learning. After being trained on one token-type, subjects were confronted with a conspecific demonstrator who either received the same food reward as the subject (control condition) or a higher value food reward than the subject (test condition) for exchanging another token-type. In general, the chimpanzees persisted in exchanging the token-type they were trained on individually, indicating a form of conservatism consistent with previous studies. However, the chimpanzees were more inclined to copy the demonstrator in the test compared to the control condition, indicating a tendency to employ a copy-if-better strategy. We discuss the validity of our results by considering alternative explanations and relate our findings to the emergence of cumulative culture.     

Early social deprivation negatively affects social skill acquisition in chimpanzees (Pan troglodytes)

In a highly social species like chimpanzees, the process by which individuals become attuned to their social environment may be of vital importance to their chances of survival. Typically, this socialization process, defined by all acquisition experiences and fine-tuning efforts of social interaction patterns during ontogeny, occurs in large part through parental investment. In this study, we investigated whether maternal presence enhances the socialization process in chimpanzees by comparing the social interactions of orphaned and mother-reared individuals at the Chimfunshi Wildlife Orphanage Trust in Zambia. As response variables, we selected social interactions during which an elaborate level of fine-tuning is assumed to be necessary for sustaining the interaction and preventing escalation: social play. Comparing orphaned (n = 8) to sex- and age-matched mother-reared juvenile chimpanzees (n = 9), we hypothesized that the orphaned juveniles would play less frequently than the mother-reared and would be less equipped for fine-tuning social play (which we assayed by rates of aggression) because of the lack of a safe and facilitating social environment provided by the mother. First, contrary to our hypothesis, results showed that the orphaned juveniles engaged in social play more frequently than the mother-reared juveniles, although for significantly shorter amounts of time. Second, in support of our hypothesis, results showed that social play of the orphaned juveniles more often resulted in aggression than social play of the mother-reared juveniles. In conjunction, these results may indicate that, just like in humans, chimpanzee mothers provide their offspring with adequate social skills that might be of pivotal importance for future challenges like successful group-living and securing competitive fitness advantages.     

Does social environment influence learning ability in a family-living lizard?

Early developmental environment can have profound effects on individual physiology, behaviour, and learning. In birds and mammals, social isolation during development is known to negatively affect learning ability; yet in other taxa, like reptiles, the effect of social isolation during development on learning ability is unknown. We investigated how social environment affects learning ability in the family-living tree skink (Egernia striolata). We hypothesized that early social environment shapes cognitive development in skinks and predicted that skinks raised in social isolation would have reduced learning ability compared to skinks raised socially. Offspring were separated at birth into two rearing treatments: (1) raised alone or (2) in a pair. After 1 year, we quantified spatial learning ability of skinks in these rearing treatments (N = 14 solitary, 14 social). We found no effect of rearing treatment on learning ability. The number of skinks to successfully learn the task, the number of trials taken to learn the task, the latency to perform the task, and the number of errors in each trial did not differ between isolated and socially reared skinks. Our results were unexpected, yet the facultative nature of this species’ social system may result in a reduced effect of social isolation on behaviour when compared to species with obligate sociality. Overall, our findings do not provide evidence that social environment affects development of spatial learning ability in this family-living lizard.     

Experimental evidence for action imitation in killer whales (Orcinus orca)

Comparative experimental studies of imitative learning have focused mainly on primates and birds. However, cetaceans are promising candidates to display imitative learning as they have evolved in socioecological settings that have selected for large brains, complex sociality, and coordinated predatory tactics. Here we tested imitative learning in killer whales, Orcinus orca. We used a ‘do-as-other-does’ paradigm in which 3 subjects witnessed a conspecific demonstrator’s performance that included 15 familiar and 4 novel behaviours. The three subjects (1) learned the copy command signal ‘Do that’ very quickly, that is, 20 trials on average; (2) copied 100 % of the demonstrator’s familiar and novel actions; (3) achieved full matches in the first attempt for 8–13 familiar behaviours (out of 15) and for the 2 novel behaviours (out of 2) in one subject; and (4) took no longer than 8 trials to accurately copy any familiar behaviour, and no longer than 16 trials to copy any novel behaviour. This study provides experimental evidence for body imitation, including production imitation, in killer whales that is comparable to that observed in dolphins tested under similar conditions. These findings suggest that imitative learning may underpin some of the group-specific traditions reported in killer whales in the field.     

Asymmetries in the production of self-directed behavior by chimpanzees and gorillas during a computerized cognitive test

Self-directed behaviors (SDBs) are a commonly used behavioral indicator of arousal in nonhuman primates. Experimental manipulations, designed to increase arousal and uncertainty, have been used to elicit SDB production in primates. Beyond measuring rates of SDB production, researchers have also recorded their lateralized production by primates, thought to reflect laterality of hemispheric brain control and response to emotion. Although a handful of such studies exist, all have been conducted with chimpanzees. Expanding on this line of inquiry, we tested both chimpanzees (N = 3) and gorillas (N = 3) in a serial learning task presented on a touchscreen interface that incorporated both EASY (two-item list) and HARD (four-item list) versions of the task. Although SDB production by the apes did not differ across the two levels of task complexity, both species produced higher rates of SDB when they made an error, regardless of task difficulty. Furthermore, the apes made more SDB with the left hand—directed to the right side of their body (contralateral SDB) and left side of their body (ipsilateral SDB)—when they made an incorrect response. There was no difference in the rate of SDB produced with the right hand across correct compared to incorrect trials. The apes’ responses reflect previous reports that show humans are quicker at selecting negative emotional stimuli when using their left, compared to their right, hand (the reverse is true for positive stimuli). However, previous work has shown that chimpanzees are more likely to produce (contralateral) SDB with their right hand when aroused and so we discuss our results in relation to these findings and consider how they relate to the ‘right hemisphere’ and ‘valence’ models of emotional processing in apes.     

Leaf swallowing behavior in chimpanzees (Pan troglodytes): biased learning and the emergence of group level cultural differences

Demonstrating the ability to ‘copy’ the behavior of others is an important aspect in determining whether social learning occurs and whether group level differences in a given behavior represent cultural differences or not. Understanding the occurrence of this process in its natural context is essential, but can be a daunting task in the wild. In order to test the social learning hypothesis for the acquisition of leaf swallowing (LS), a self-medicative behavior associated with the expulsion of parasites, we conducted semi-naturalistic experiments on two captive groups of parasite-free, naïve chimpanzees (Pan troglodytes). Individuals in the group were systematically provided appropriate stimuli (rough hispid leaves) identical to those used by chimpanzees in the wild. Individuals initially responded in a variety of ways, ranging from total aversion to normal chewing and swallowing. Over time, however, the two groups adopted different variants for inserting and folding the leaves in the mouth prior to swallowing them (complete and partial LS), following the specific method spontaneously displayed by the first and primary LS models in their respective groups. These variants were similar to LS displayed by chimpanzees in the wild. Using the option-bias method, we found evidence for social learning leading to group-level biased transmission and group-level stabilization of these two variants. This is the first report on two distinct cultural variants innovated in response to the introduction of natural stimuli that emerged and spread spontaneously and concurrently within two adjacent groups of socially housed primates. These observations support the assertion that LS may reflect a generalized propensity for ingesting rough hispid leaves, which can be socially induced and transmitted within a group. Ingesting an adequate number of these leaves induces increased gut motility, which is responsible for the subsequent expulsion of particular parasite species in the wild. Cultural transmission and maintenance of LS within a group and associative learning by the individual of the positive consequences of this otherwise non-nutritive mode of ingestion is proposed to be the pivotal link between this feeding propensity and its maintenance as a self-medicative behavior by great apes in the wild.     

The relationship between perceived organizational support and proactive behaviour directed towards the organization

Numerous studies have shown that perceived organizational support has positive consequences on both employees and organizations. However, no study has examined the causal relationship between perceived organizational support and proactive behaviour directed towards the organization and few studies have identified potential mechanisms underlying this relationship. We first investigated the direction of causality between perceived organizational support and proactive behaviour, and hypothesized that perceived organizational support leads to proactive behaviour directed towards the organization. Second, based on social exchange and motivational perspectives, we examined two potential mechanisms underlying this relationship, i.e., felt obligation and work engagement. Employees from a Belgian Federal Public Service were invited to take part in a three-wave longitudinal survey. Using a cross-lagged panel design, our results (N = 1963) revealed that Time 2 perceived organizational support was positively related to Time 3 proactive behaviour towards the organization, whereas the reverse was not true. Additionally, our results (N = 1293) showed that Time 2 felt obligation and Time 2 work engagement mediated the positive relationship between Time 1 perceived organizational support and Time 3 proactive behaviour directed towards the organization.     

Chimpanzees demonstrate individual differences in social information use

Studies of transmission biases in social learning have greatly informed our understanding of how behaviour patterns may diffuse through animal populations, yet within-species inter-individual variation in social information use has received little attention and remains poorly understood. We have addressed this question by examining individual performances across multiple experiments with the same population of primates. We compiled a dataset spanning 16 social learning studies (26 experimental conditions) carried out at the same study site over a 12-year period, incorporating a total of 167 chimpanzees. We applied a binary scoring system to code each participant’s performance in each study according to whether they demonstrated evidence of using social information from conspecifics to solve the experimental task or not (Social Information Score—‘SIS’). Bayesian binomial mixed effects models were then used to estimate the extent to which individual differences influenced SIS, together with any effects of sex, rearing history, age, prior involvement in research and task type on SIS. An estimate of repeatability found that approximately half of the variance in SIS was accounted for by individual identity, indicating that individual differences play a critical role in the social learning behaviour of chimpanzees. According to the model that best fit the data, females were, depending on their rearing history, 15–24% more likely to use social information to solve experimental tasks than males. However, there was no strong evidence of an effect of age or research experience, and pedigree records indicated that SIS was not a strongly heritable trait. Our study offers a novel, transferable method for the study of individual differences in social learning.     

Influence of personality,age, sex,and estrous state on chimpanzee problem-solving success

Despite the importance of individual problem solvers for group- and individual-level fitness, the correlates of individual problem-solving success are still an open topic of investigation. In addition to demographic factors, such as age or sex, certain personality dimensions have also been revealed as reliable correlates of problem-solving by animals. Such correlates, however, have been little-studied in chimpanzees. To empirically test the influence of age, sex, estrous state, and different personality factors on chimpanzee problem-solving, we individually tested 36 captive chimpanzees with two novel foraging puzzles. We included both female (N = 24) and male (N = 12) adult chimpanzees (aged 14–47 years) in our sample. We also controlled for the females’ estrous state—a potential influence on cognitive reasoning—by testing cycling females both when their sexual swelling was maximally tumescent (associated with the luteinizing hormone surge of a female’s estrous cycle) and again when it was detumescent. Although we found no correlation between the chimpanzees’ success with either puzzle and their age or sex, the chimpanzees’ personality ratings did correlate with responses to the novel foraging puzzles. Specifically, male chimpanzees that were rated highly on the factors Methodical, Openness (to experience), and Dominance spent longer interacting with the puzzles. There was also a positive relationship between the latency of females to begin interacting with the two tasks and their rating on the factor Reactivity/Undependability. No other significant correlations were found, but we report tentative evidence for increased problem-solving success by the females when they had detumescent estrous swellings.     

The effect of oxytocin on biological motion perception in dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Recent studies have shown that the neuropeptide oxytocin is involved in the regulation of several complex human social behaviours. There is, however, little research on the effect of oxytocin on basic mechanisms underlying human sociality, such as the perception of biological motion. In the present study, we investigated the effect of oxytocin on biological motion perception in dogs (Canis familiaris), a species adapted to the human social environment and thus widely used to model many aspects of human social behaviour. In a within-subjects design, dogs (N = 39), after having received either oxytocin or placebo treatment, were presented with 2D projection of a moving point-light human figure and the inverted and scrambled version of the same movie. Heart rate (HR) and heart rate variability (HRV) were measured as physiological responses, and behavioural response was evaluated by observing dogs’ looking time. Subjects were also rated on the personality traits of Neuroticism and Agreeableness by their owners. As expected, placebo-pretreated (control) dogs showed a spontaneous preference for the biological motion pattern; however, there was no such preference after oxytocin pretreatment. Furthermore, following the oxytocin pretreatment female subjects looked more at the moving point-light figure than males. The individual variations along the dimensions of Agreeableness and Neuroticism also modulated dogs’ behaviour. Furthermore, HR and HRV measures were affected by oxytocin treatment and in turn played a role in subjects? looking behaviour. We discuss how these findings contribute to our understanding of the neurohormonal regulatory mechanisms of human (and non-human) social skills.     

Sex-specific cognitive–behavioural profiles emerging from individual variation in numerosity discrimination in <Emphasis Type="Italic">Gambusia affinis</Emphasis>

The relationship between an individual’s cognitive abilities and other behavioural attributes is complex, yet critical to understanding how individual differences in cognition arise. Here we use western mosquitofish, Gambusia affinis, to investigate the relationship between individual associative learning performance in numerical discrimination tests and independent measures of activity, exploration, anxiety and sociability. We found extensive and highly repeatable inter-individual variation in learning performance (r = 0.89; ICC = 0.89). Males and females exhibited similar learning performance, yet differed in sociability, activity and their relationship between learning and anxiety/exploration tendencies. Sex-specific multivariate behaviour scores successfully predicted variation in individual learning performance, whereas combined sex analyses did not. Female multivariate behaviour scores significantly predict learning performance across females (ρ = 0.80, p = 0.005) with high-performing female learners differentiated from female non-learners and low-performing learners by significant contributions of activity and sociability measures. Meanwhile, males of different learning performance levels (high-, low- and non-learners) were distinguished from each other by unique behavioural loadings of sociability, activity and anxiety/exploration scores, respectively. Our data suggest that despite convergence on learning performance, the sexes diverge in cognitive–behavioural relationships that are likely products of different sexual selection pressures.     

Social learning in Cartilaginous fish (stingrays Potamotrygon falkneri)

Social learning is considered one of the hallmarks of cognition. Observers learn from demonstrators that a particular behavior pattern leads to a specific consequence or outcome, which may be either positive or negative. In the last few years, social learning has been studied in a variety of taxa including birds and bony fish. To date, there are few studies demonstrating learning processes in cartilaginous fish. Our study shows that the cartilaginous fish freshwater stingrays (Potamotrygon falkneri) are capable of social learning and isolates the processes involved. Using a task that required animals to learn to remove a food reward from a tube, we found that observers needed significantly (P < 0.01) fewer trials to learn to extract the reward than demonstrators. Furthermore, observers immediately showed a significantly (P < 0.05) higher frequency of the most efficient “suck and undulation” strategy exhibited by the experienced demonstrators, suggesting imitation. Shedding light on social learning processes in cartilaginous fish advances the systematic comparison of cognition between aquatic and terrestrial vertebrates and helps unravel the evolutionary origins of social cognition.     

Extrinsic Religious Orientation and Well-Being: Is Their Negative Association Real or Spurious?

Turning to religion to seek its social benefits has been associated with poor psychological well-being. Researchers have concluded that endorsing this extrinsic and social orientation toward religion is inauthentic and unhealthy. However, few studies have focused on extrinsic-social religious orientation’s negative relationship with well-being, leaving open the possibility that their relationship is spurious. The present study argues that people endorsing an extrinsic-social religious orientation also perceive lower levels of social support in their lives, thus their turning to religion to fill this social void. As social support is important for healthy psychological functioning, perceived social support may be the critical third variable explaining why extrinsic-social religious orientation appears to have psychological costs. This study supported our expectations among undergraduates in two countries: the United States (N = 156) and the Republic of Ireland (N = 255). There were negative bivariate associations between extrinsic-social religious orientation and both perceived social support and emotional well-being. Accounting for the effects of perceived social support, however, reduced the association between the extrinsic-social religious orientation and well-being to non-significance. Thus, people endorsing an extrinsic and social orientation toward religion tend to have poor well-being because they perceive less supportive relationships in their lives.     

Trace classical conditioning in rainbow trout (Oncorhynchus mykiss): what do they learn?

There are two main memory systems: declarative and procedural memory. Knowledge of these two systems in fish is scarce, and controlled laboratory studies are needed. Trace classical conditioning is an experimentally tractable model of declarative memory. We tested whether rainbow trout (Oncorhynchus mykiss) can learn by trace conditioning and form stimulus–stimulus, as opposed to stimulus–response, associations. We predicted that rainbow trout trained by trace conditioning would show appetitive behaviour (conditioned response; CR) towards the conditioned stimulus (CS; light), and that the CR would be sensitive to devaluation of the unconditioned stimulus (US; food). The learning group (L, N = 14) was trained on a CS + US contingency schedule with a trace interval of 3.4 s. The control group (CtrL, N = 4) was kept on a completely random schedule. The fish that learnt were further trained as either an experimental (L, N = 6) or a memory control (CtrM, N = 3) group. The L group had the US devalued. The CtrM group received only food. No fish in the CtrL group, but nine fish from the L group conditioned to the light. When tested, five L fish changed their CRs after US devaluation, indicating learning by stimulus–stimulus association of the light with the food. CtrM fish retained their original CRs. To the best of our knowledge, this experiment is the first to show that rainbow trout can learn by trace classical conditioning. The results indicate that the fish learnt by ‘facts-learning’ rather than by reflex acquisition in this study.     

Impaired Fear Recognition and Social Anxiety Symptoms in Adolescence

This study represents the first examination of adolescent anxiety in relation to peer emotion recognition, rather than adult emotion recognition. Additionally, we examine potential mechanisms for the development of social anxiety in females. Facial emotion recognition (FER) is important for accurate social cognition, which is impaired in individuals with various disorders, including anxiety disorders. Social anxiety often onsets during adolescence, is observed more commonly in females, and is often associated with FER difficulties. Given the importance of peer interaction during adolescence, and some evidence that FER may differ as a function of the stimuli (adolescent or adult faces), we sought to study FER in relation to social anxiety symptoms using stimuli portraying adolescent faces. Male and female adolescents (N = 64) completed an online survey in which they rated 257 child and adolescent emotional faces and completed a self-report measure of social anxiety symptoms. We examined differences in emotion recognition (e.g., fear, anger, sadness) between individuals with high and low levels of social anxiety symptoms. Adolescents with high social anxiety symptoms were more likely to have problems correctly identifying fearful expressions (90.55 % accuracy) compared to adolescents with low social anxiety symptoms (96.00 % accuracy; t = 2.375, p = .021, d = 0.594), and this effect was observed exclusively in female adolescents. The observed sex difference in accurate identification of fearful faces in relation to social anxiety could suggest a potential mechanism for social anxiety development in adolescent females.     

Responses toward a trapped animal by wild bonobos at Wamba

Chimpanzees and bonobos are the closest living relatives of humans and diverged relatively recently in their phylogenetic history. However, a number of reports have suggested behavioral discrepancies between the two Pan species, such as more cooperative and tolerant social interaction and poorer tool-using repertoires in bonobos. Concerning hunting behavior and meat consumption, recent studies from the field have confirmed both behaviors not only in chimpanzees but also in bonobos. The present study reports an encounter by wild bonobos at Wamba with a duiker trapped in a snare. Bonobos interacted with the live duiker for about 10 min but did not eventually kill the animal. They showed fear responses when the duiker moved and exhibited behaviors related to anxiety and stress such as branch-drag displays and self-scratching. Although bonobos manipulated nearby saplings and parts of the snare, they did not use detached objects to make indirect contact with the duiker. Juveniles and adults of both sexes engaged in active interactions with the trapped duiker. Overall, bonobos’ behavioral responses indicated species-specific cognitive characteristics largely different from those of chimpanzees.     

Reaching around barriers: the performance of the great apes and 3–5-year-old children

Inhibitory control has been suggested as a key predictive measure of problem-solving skills in human and nonhuman animals. However, there has yet to be a direct comparison of the inhibitory skills of the nonhuman apes and their development in human children. We compared the inhibitory skills of all great ape species, including 3–5-year-old children in a detour-reaching task, which required subjects to avoid reaching directly for food and instead use an indirect reaching method to successfully obtain the food. We tested 22 chimpanzees, 18 bonobos, 18 orangutans, 6 gorillas and 42 children. Our sample included chimpanzees, bonobos and orangutans housed in zoos (N = 27) and others housed in sanctuaries in their native habitats (N = 37). Overall, orangutans were the most skilful apes, including human children. As expected older children outperformed younger children. Sanctuary chimpanzees and bonobos outperformed their zoo counterparts whereas there was no difference between the two orangutan samples. Most zoo chimpanzees and bonobos failed to solve the original task, but improved their performance with additional training, although the training method determined to a considerable extent the level of success that the apes achieved in a transfer phase. In general, the performance of the older children was far from perfect and comparable to some of the nonhuman apes tested.     

Can children break the cycle of disadvantage? Structure and agency in the transmission of education across generations

Research has shown that parents tend to pass educational advantage or disadvantage on to their children. However, little is known about the extent to which the intergenerational transmission of education involves children’s agency. In this study we drew from two traditions in sociological and social psychological theorizing—the theory of cultural and social reproduction and the theory of human agency—to examine whether agency influences children’s educational performance, and if so, whether this influence can be observed among children across social classes. We used data from the Spanish sample of the Program for International Student Assessment (N = 25,003 15-year-olds). Results indicate that the level of child agency was weakly positively related to social class, that child agency impacted on a child’s educational performance, and that the positive effect of agency on educational performance did not vary by social class. This suggests that strategies to enhance disadvantaged children’s agency may prove useful in reducing social gradients in educational performance. More generally, our findings may ignite a debate about the role that social structure and human agency play in shaping social inequality and mobility.