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Abstract Individuals vary in how they move their faces in everyday social interactions. In a first large-scale study, we measured variation in dynamic facial behaviour during social interaction and examined dyadic outcomes and impression formation. In Study 1, we recorded semi-structured video calls with 52 participants interacting with a confederate across various everyday contexts. Video clips were rated by 176 independent participants. In Study 2, we examined video calls of 1315 participants engaging in unstructured video-call interactions. Facial expressivity indices were extracted using automated Facial Action Coding Scheme analysis and measures of personality and partner impressions were obtained by self-report. Facial expressivity varied considerably across participants, but little across contexts, social partners or time. In Study 1, more facially expressive participants were more well-liked, agreeable, and successful at negotiating (if also more agreeable). Participants who were more facially competent, readable, and perceived as readable were also more well-liked. In Study 2, we replicated the findings that facial expressivity was associated with agreeableness and liking by their social partner, and additionally found it to be associated with extraversion and neuroticism. Findings suggest that facial behaviour is a stable individual difference that proffers social advantages, pointing towards an affiliative, adaptive function.

Abstract Infants across species are thought to exhibit specific facial features (termed the “baby schema”, such as a relatively bigger forehead and eyes, and protruding cheeks), with an adaptive function to induce caretaking behaviour from adults. There is abundant empirical evidence for this in humans, but, surprisingly, the existence of a baby schema in non-human animals has not been scientifically demonstrated. We investigated which facial characteristics are shared across infants in five species of great apes: humans, chimpanzees, bonobos, mountain gorillas, and Bornean orangutans. We analysed eight adult and infant faces for each species (80 images in total) using geometric morphometric analysis and machine learning. We found two principal components characterizing infant faces consistently observed across species. These included (1) relatively bigger eyes located lower in the face, (2) a rounder and vertically shorter face shape, and (3) an inverted triangular face shape. While these features are shared, human infant faces are unique in that the second characteristic (round face shape) is more pronounced, whereas the third (inverted triangular face shape) is less pronounced than other species. We also found some infantile features only found in some species. We discuss future directions to investigate the baby schema using an evolutionary approach.

Inhibitory control, the ability to override an inappropriate prepotent response, is crucial in many aspects of everyday life. However, the various paradigms designed to measure inhibitory control often suffer from a lack of systematic validation and have yielded mixed results. Thus the nature of this ability remains unclear, is it a general construct or a family of distinct sub-components? Therefore, the aim of this study was first to demonstrate the content validity and the temporal repeatability of a battery of inhibitory control tasks. Then we wanted to assess the contextual consistency of performances between these tasks to better understand the structure of inhibitory control. We tested 21 rhesus macaques (Macaca mulatta, 12 males, nine females) in a battery of touchscreen tasks assessing three main components of inhibitory control: inhibition of a distraction (using a Distraction task), inhibition of an impulsive action (using a Go/No-go task) and inhibition of a cognitive set (using a Reversal learning task). All tasks were reliable and effective at measuring the inhibition of a prepotent response. However, while there was consistency of performance between the inhibition of a distraction and the inhibition of an action, representing a response-driven basic form of inhibition, this was not found for the inhibition of a cognitive set. We argue that the inhibition of a cognitive set is a more cognitively demanding form of inhibition. This study gives a new insight in the multifaceted structure of inhibitory control and highlights the importance of a systematic validation of cognitive tasks in animal cognition.

Darwin and other pioneering scholars made comparisons between human facial signals and those of non-human primates, suggesting that they share evolutionary history. We now have tools available (the Facial Action Coding System) to make these comparisons anatomically based and standardised, as well as analytical methods to facilitate comparative studies. Here we review the evidence establishing a shared anatomical basis between the facial behaviour of human and non-human primate species, concluding which signals are probably related, and which are not. We then review the evidence for shared function and discuss the implications for understanding human communication. Where differences between humans and other species exist, we explore possible explanations and future directions for enquiry.

Inhibitory control, the ability to override a dominant response, is crucial in many aspects of everyday life. In animal studies, striking individual variations are often largely ignored and their causes rarely considered. Hence, our aims were to systematically investigate individual variability in inhibitory control, to replicate the most common causes of individual variation (age, sex and rank) and to determine if these factors had a consistent effect on three main components of inhibitory control (inhibition of a distraction, inhibition of an action, inhibition of a cognitive set). We tested 21 rhesus macaques (Macaca mulatta) in a battery of validated touchscreen tasks. We first found individual variations in all inhibitory control performances. We then demonstrated that males had poorer performances to inhibit a distraction and that middle-aged individuals exhibited poorer performance in the inhibition of a cognitive set. Hence, the factors of age and sex were not consistently associated with the main components of inhibitory control, suggesting a multi-faceted structure. The rank of the subjects did not influence any inhibitory control performances. This study adopts a novel approach for animal behaviour studies and gives new insight into the individual variability of inhibitory control which is crucial to understand its evolutionary underpinnings.

Humans are uniquely cooperative and form crucial short- and long-term social bonds between individuals that ultimately shape human societies. The need for such intense cooperation may have provided a particularly powerful selection pressure on the emotional and communicative behaviours regulating cooperative processes, such as guilt. Guilt is a social, other-oriented moral emotion that promotes relationship repair and pro-sociality. For example, people can be more lenient towards wrongdoers who display guilt than towards those who do not. Here, we examined the social consequences of guilt in a novel experimental setting with pairs of friends differing in relationship quality. Pairs of participants took part in a cooperative game with a mutual goal. We then induced guilt in one of the participants and informed the other participant of their partner's wrongdoing. We examined the outcome using a dictator game to see how they split a joint reward. We found that guilty people were motivated to repair wrongdoing regardless of friendship. Observing guilt in others led to a punishment effect and a victim of wrongdoing punished close friends who appeared guilty more so than acquaintances. We suggest, therefore, that guilt has a stronger function between close friends as the costs of relationship breakdown are greater. Relationship context, therefore, is crucial to the functional relevance of moral emotions.

Abstract Most mammalian species produce facial expressions. Historically, animal facial expressions have been considered inflexible and involuntary displays of emotional states rather than active attempts to communicate with others. In the current study, we aimed to test whether domestic dog facial expressions are subject to audience effects and/ or changes in response to an arousing stimulus (e.g. food) alone. We presented dogs with an experimental situation in which a human demonstrator was either attending to them or turned away, and varied whether she presented food or not. Dogs produced significantly more facial movements when the human was attentive than when she was not. The food, however, as a non-social but arousing stimulus, did not affect the dogs’ behaviour. The current study is therefore evidence that dogs are sensitive to the human’s attentional state when producing facial expressions, suggesting that facial expressions are not just inflexible and involuntary displays of emotional states, but rather potentially active attempts to communicate with others.

Human and non-human primates exhibit facial movements or displays to communicate with one another. The evolution of form and function of those displays could be better understood through multispecies comparisons. Anatomically based coding systems (Facial Action Coding Systems: FACS) are developed to enable such comparisons because they are standardized and systematic and aid identification of homologous expressions underpinned by similar muscle contractions. To date, FACS has been developed for humans, and subsequently modified for chimpanzees, rhesus macaques, orangutans, hylobatids, dogs, and cats. Here, we wanted to test whether the MaqFACS system developed in rhesus macaques (Macaca mulatta) could be used to code facial movements in Barbary macaques (M. sylvanus), a species phylogenetically close to the rhesus macaques. The findings show that the facial movement capacity of Barbary macaques can be reliably coded using the MaqFACS. We found differences in use and form of some movements, most likely due to specializations in the communicative repertoire of each species, rather than morphological differences.

Primate facial expressions are widely accepted as underpinned by reflexive emotional processes and not under voluntary control. In contrast, other modes of primate communication, especially gestures, are widely accepted as underpinned by intentional, goal-driven cognitive processes. One reason for this distinction is that production of primate gestures is often sensitive to the attentional state of the recipient, a phenomenon used as one of the key behavioural criteria for identifying intentionality in signal production. The reasoning is that modifying/producing a signal when a potential recipient is looking could demonstrate that the sender intends to communicate with them. Here, we show that the production of a primate facial expression can also be sensitive to the attention of the play partner. Using the orangutan (Pongo pygmaeus) Facial Action Coding System (OrangFACS), we demonstrate that facial movements are more intense and more complex when recipient attention is directed towards the sender. Therefore, production of the playface is not an automated response to play (or simply a play behaviour itself) and is instead produced flexibly depending on the context. If sensitivity to attentional stance is a good indicator of intentionality, we must also conclude that the orangutan playface is intentionally produced. However, a number of alternative, lower level interpretations for flexible production of signals in response to the attention of another are discussed. As intentionality is a key feature of human language, claims of intentional communication in related primate species are powerful drivers in language evolution debates, and thus caution in identifying intentionality is important.

Many species use facial features to identify conspecifics, which is necessary to navigate a complex social environment. The fundamental mechanisms underlying face processing are starting to be well understood in a variety of primate species. However, most studies focus on a limited subset of species tested with unfamiliar faces. As well as limiting our understanding of how widely distributed across species these skills are, this also limits our understanding of how primates process faces of individuals they know, and whether social factors (e.g. dominance and social bonds) influence how readily they recognize others. In this study, socially housed crested macaques voluntarily participated in a series of computerized matching-to-sample tasks investigating their ability to discriminate (i) unfamiliar individuals and (ii) members of their own social group. The macaques performed above chance on all tasks. Familiar faces were not easier to discriminate than unfamiliar faces. However, the subjects were better at discriminating higher ranking familiar individuals, but not unfamiliar ones. This suggests that our subjects applied their knowledge of their dominance hierarchies to the pictorial representation of their group mates. Faces of high-ranking individuals garner more social attention, and therefore might be more deeply encoded than other individuals. Our results extend the study of face recognition to a novel species, and consequently provide valuable data for future comparative studies.

Scientists studying the communication of non-human animals are often aiming to better understand the evolution of human communication, including human language. Some scientists take a phylogenetic perspective, where the goal is to trace the evolutionary history of communicative traits, while others take a functional perspective, where the goal is to understand the selection pressures underpinning specific traits. Both perspectives are necessary to fully understand the evolution of communication, but it is important to understand how the two perspectives differ and what they can and cannot tell us. Here, we suggest that integrating phylogenetic and functional questions can be fruitful in better understanding the evolution of communication. We also suggest that adopting a multimodal approach to communication might help to integrate phylogenetic and functional questions, and provide an interesting avenue for research into language evolution.

The social complexity hypothesis for communicative complexity posits that animal societies with more complex social systems require more complex communication systems. We tested the social complexity hypothesis on three macaque species that vary in their degree of social tolerance and complexity. We coded facial behavior in >3000 social interactions across three social contexts (aggressive, submissive, affiliative) in 389 animals, using the Facial Action Coding System for macaques (MaqFACS). We quantified communicative complexity using three measures of uncertainty: entropy, specificity, and prediction error. We found that the relative entropy of facial behavior was higher for the more tolerant crested macaques as compared to the less tolerant Barbary and rhesus macaques across all social contexts, indicating that crested macaques more frequently use a higher diversity of facial behavior. The context specificity of facial behavior was higher in rhesus as compared to Barbary and crested macaques, demonstrating that Barbary and crested macaques used facial behavior more flexibly across different social contexts. Finally, a random forest classifier predicted social context from facial behavior with highest accuracy for rhesus and lowest for crested, indicating there is higher uncertainty and complexity in the facial behavior of crested macaques. Overall, our results support the social complexity hypothesis.

Guilt is a complex emotion with a potentially important social function of stimulating cooperative behaviours towards and from others, but whether the feeling of guilt is associated with a recognisable pattern of nonverbal behaviour is unknown. We examined the production and perception of guilt in two different studies, with a total of 238 participants with various places of origin. Guilt was induced experimentally, eliciting patterns of movement that were associated with both the participants' self-reported feelings of guilt and judges' impressions of their guilt. Guilt was most closely associated with frowning and neck touching. While there were differences between self-reported guilt and perception of guilt the findings suggest that there are consistent patterns that could be considered a non-verbal signal of guilt in humans.

Many facilities that house captive primates play music for animal enrichment or for caregiver enjoyment. However, the impact on primates is unknown as previous studies have been inconclusive. We conducted three studies with zoo-housed chimpanzees (Pan troglodytes) and one with group-housed chimpanzees at the National Centre for Chimpanzee Care to investigate the effects of classical and pop/rock music on various variables that may be indicative of increased welfare. Study one compared the behaviour and use of space of 18 animals when silence, classical or pop/rock music was played into one of several indoor areas. Overall, chimpanzees did not actively avoid the area when music was playing but were more likely to exit the area when songs with higher beats per minute were broadcast. Chimpanzees showed significantly fewer active social behaviours when music, rather than silence, was playing. They also tended to be more active and engage in less abnormal behaviour during the music but there was no change to either self-grooming or aggression between music and silent conditions. The genre of music had no differential effects on the chimpanzees' use of space and behaviour. In the second study, continuous focal observations were carried out on three individuals with relatively high levels of abnormal behaviour. No differences in behaviour between music and silence periods were found in any of the individuals. The final two studies used devices that allowed chimpanzees to choose if they wanted to listen to music of various types or silence. Both studies showed that there were no persistent preferences for any type of music or silence. When taken together, our results do not suggest music is enriching for group-housed captive chimpanzees, but they also do not suggest that music has a negative effect on welfare.

Non-human primates use various communicative means in interactions with others. While primate gestures are commonly considered to be intentionally and flexibly used signals, facial expressions are often referred to as inflexible, automatic expressions of affective internal states. To explore whether and how non-human primates use facial expressions in specific communicative interactions, we studied five species of small apes (gibbons) by employing a newly established Facial Action Coding System for hylobatid species (GibbonFACS). We found that, despite individuals often being in close proximity to each other, in social (as opposed to non-social contexts) the duration of facial expressions was significantly longer when gibbons were facing another individual compared to non-facing situations. Social contexts included grooming, agonistic interactions and play, whereas non-social contexts included resting and self-grooming. Additionally, gibbons used facial expressions while facing another individual more often in social contexts than non-social contexts where facial expressions were produced regardless of the attentional state of the partner. Also, facial expressions were more likely 'responded to' by the partner's facial expressions when facing another individual than non-facing. Taken together, our results indicate that gibbons use their facial expressions differentially depending on the social context and are able to use them in a directed way in communicative interactions with other conspecifics.

Although previous studies of horses have investigated their facial expressions in specific contexts, e.g. pain, until now there has been no methodology available that documents all the possible facial movements of the horse and provides a way to record all potential facial configurations. This is essential for an objective description of horse facial expressions across a range of contexts that reflect different emotional states. Facial Action Coding Systems (FACS) provide a systematic methodology of identifying and coding facial expressions on the basis of underlying facial musculature and muscle movement. FACS are anatomically based and document all possible facial movements rather than a configuration of movements associated with a particular situation. Consequently, FACS can be applied as a tool for a wide range of research questions. We developed FACS for the domestic horse (Equus caballus) through anatomical investigation of the underlying musculature and subsequent analysis of naturally occurring behaviour captured on high quality video. Discrete facial movements were identified and described in terms of the underlying muscle contractions, in correspondence with previous FACS systems. The reliability of others to be able to learn this system (EquiFACS) and consistently code behavioural sequences was high--and this included people with no previous experience of horses. A wide range of facial movements were identified, including many that are also seen in primates and other domestic animals (dogs and cats). EquiFACS provides a method that can now be used to document the facial movements associated with different social contexts and thus to address questions relevant to understanding social cognition and comparative psychology, as well as informing current veterinary and animal welfare practices.

Scientists are increasing their efforts to promote public engagement with their science, but the efficacy of the methods used is often not scientifically evaluated. Here, we designed, installed and evaluated the educational impact of interactive games on touchscreens at two primate research centres based in zoo environments. The games were designed to promote interest in and understanding of primates and comparative psychology, as a scaffold towards interest in science more generally and with the intention of targeting younger individuals (under 16's). We used systematic observational techniques and questionnaires to assess the impact of the games on zoo visitors. The games facilitated increased interest in psychology and science in zoo visitors, and changed the knowledge of visitors, through demonstration of learning about specific scientific findings nested within the games. The impact of such devices was greatest on younger individuals (under 16's) as they were significantly more likely to engage with the games. On the whole, therefore, this study demonstrates that interactive devices can be successful educational tools, and adds to the growing body of evidence that conducting research on public view in zoos can have a tangible impact on public engagement with science.

How wolves were first domesticated is unknown. One hypothesis suggests that wolves underwent a process of self-domestication by tolerating human presence and taking advantage of scavenging possibilities. The puppy-like physical and behavioural traits seen in dogs are thought to have evolved later, as a byproduct of selection against aggression. Using speed of selection from rehoming shelters as a proxy for artificial selection, we tested whether paedomorphic features give dogs a selective advantage in their current environment. Dogs who exhibited facial expressions that enhance their neonatal appearance were preferentially selected by humans. Thus, early domestication of wolves may have occurred not only as wolf populations became tamer, but also as they exploited human preferences for paedomorphic characteristics. These findings, therefore, add to our understanding of early dog domestication as a complex co-evolutionary process.

Providing cognitive challenges to zoo-housed animals may provide enriching effects and subsequently enhance their welfare. Primates may benefit most from such challenges as they often face complex problems in their natural environment and can be observed to seek problem solving opportunities in captivity. However, the extent to which welfare benefits can be achieved through programmes developed primarily for cognitive research is unknown. We tested the impact of voluntary participation cognitive testing on the welfare of a socially housed group of crested macaques (Macaca nigra) at the Macaque Study Centre (Marwell Zoo). First, we compared the rate of self-directed and social behaviours on testing and non-testing days, and between conditions within testing days. Minimal differences in behaviour were found when comparing testing and non-testing days, suggesting that there was no negative impact on welfare as a result of cognitive testing. Lipsmacking behaviours were found to increase and aggressive interaction was found to decrease in the group as a result of testing. Second, social network analysis was used to assess the effect of testing on associations and interactions between individuals. The social networks showed that testing subjects increased their association with others during testing days. One interpretation of this finding could be that providing socially housed primates with an opportunity for individuals to separate from the group for short periods could help mimic natural patterns of sub-group formation and reunion in captivity. The findings suggest, therefore, that the welfare of captive primates can be improved through the use of cognitive testing in zoo environments.

Primate behavioural and cognitive research is increasingly conducted on direct public view in zoo settings. The potential of such facilities for public engagement with science is often heralded, but evidence of tangible, positive effects on public understanding is rare. Here, the effect of a new zoo-based primate research centre on visitor behaviour, learning and attitudes was assessed using a quasi-experimental design. Zoo visitors approached the primate research centre more often when a scientist was present and working with the primates, and reported greater awareness of primates (including conservation) compared to when the scientist was not present. Visitors also reported greater perceived learning when the scientist was present. Installation of information signage had no main effect on visitor attitudes or learning. Visitors who interacted with the signage, however, demonstrated increased knowledge and understanding when asked about the specific information present on the signs (which was related to the ongoing facial expression research at the research centre). The findings show that primate behaviour research centres on public view can have a demonstrable and beneficial effect on public understanding of science.