Perceptual integration of bodily and facial emotion cues in chimpanzees and humans.

For highly visual species like primates, facial and bodily emotion expressions play a crucial role in emotion perception. However, most research focuses on facial expressions, while the perception of bodily cues is still poorly understood. Using a novel comparative priming eye-tracking design, we examined whether our close primate relatives, the chimpanzees (Pan troglodytes), and humans infer emotions from bodily cues through subsequent perceptual integration with facial expressions. In experiment 1, we primed chimpanzees with videos of bodily movements of unfamiliar conspecifics engaged in social activities of opposite valence (play and fear) against neutral control scenes to examine attentional bias toward succeeding congruent or incongruent facial expressions. In experiment 2, we assessed the same attentional bias in humans yet using stimuli showing unfamiliar humans. In experiment 3, humans watched the chimpanzee stimuli of experiment 1, to examine cross-species emotion perception. Chimpanzees exhibited a persistent fear-related attention bias but did not associate bodily with congruent facial cues. In contrast, humans prioritized conspecifics' congruent facial expressions (matching bodily scenes) over incongruent ones (mismatching). Nevertheless, humans exhibited no congruency effect when viewing chimpanzee stimuli, suggesting difficulty in cross-species emotion perception. These results highlight differences in emotion perception, with humans being greatly affected by fearful and playful bodily cues and chimpanzees being strongly drawn toward fearful expressions, regardless of the preceding bodily priming cue. These data advance our understanding of the evolution of emotion signaling and the presence of distinct perceptual patterns in hominids.

Variable expression of linguistic laws in ape gesture: a case study from chimpanzee sexual solicitation.

Two language laws have been identified as consistent patterns shaping animal behaviour, both acting on the organizational level of communicative systems. Zipf's law of brevity describes a negative relationship between behavioural length and frequency. Menzerath's law defines a negative correlation between the number of behaviours in a sequence and average length of the behaviour composing it. Both laws have been linked with the information-theoretic principle of compression, which tends to minimize code length. We investigated their presence in a case study of male chimpanzee sexual solicitation gesture. We failed to find evidence supporting Zipf's law of brevity, but solicitation gestures followed Menzerath's law: longer sequences had shorter average gesture duration. Our results extend previous findings suggesting gesturing may be limited by individual energetic constraints. However, such patterns may only emerge in sufficiently large datasets. Chimpanzee gestural repertoires do not appear to manifest a consistent principle of compression previously described in many other close-range systems of communication. Importantly, the same signallers and signals were previously shown to adhere to these laws in subsets of the repertoire when used in play; highlighting that, in addition to selection on the signal repertoire, ape gestural expression appears shaped by factors in the immediate socio-ecological context.

The Association Between the Bared-Teeth Display and Social Dominance in Captive Chimpanzees (Pan troglodytes).

Humans use smiles - widely observed emotional expressions - in a variety of social situations, of which the meaning varies depending on social relationship and the context in which it is displayed. The homologue of the human smile in non-human primates - both due to morphological and functional similarities - is the bared-teeth display (BT). According to the power asymmetry hypothesis (PAH), species with strict linear dominance hierarchies are predicted to produce distinct communicative signals to avoid escalations of social conflicts. Hence, while the BT in a despotic species is predicted to be expressed from low- to high-ranking individuals, signaling submission, the BT in a tolerant species is predicted to be expressed in multiple contexts, regardless of rank. We tested this hypothesis in a group of 8 captive chimpanzees (Pan troglodytes), a species commonly characterized as rather despotic. An investigation of 11,774 dyadic social interactions revealed this chimpanzee group to have a linear dominance hierarchy, with moderate steepness. A Bayesian GLMM - used to test the effects of social contexts and rank relationships of dyads on the use of the BT display - indicated multi-contextual use of the BT which is contingent on the rank relationship. We also found that slight morphological and/or acoustic variants (i.e., silent bared-teeth and vocalized bared-teeth) of the BT display may have different communicative meanings. Our findings are in line with the prediction derived from the PAH for a moderately despotic species, and the view that the human smile originated from the primate BT display. Supplementary Information: The online version contains supplementary material available at 10.1007/s42761-022-00138-1.

Flexible signalling strategies by victims mediate post-conflict interactions in bonobos.

Compared to other animals, humans supposedly excel at voluntarily controlling and strategically displaying emotional signals. Yet, new data shows that nonhuman great apes' emotion expressions may also be subject to voluntary control. A key context to further explore this is during post-conflict (PC) periods, where signalling by distressed victims may influence bystander responses, including the offering of consolation. To address this, our study investigates the signalling behaviour of sanctuary-living bonobo victims following aggression and its relation to audience composition and PC interactions. Results show that the production of paedomorphic signals by victims (regardless of age) increased their chances of receiving consolation. In adults, the production of such signals additionally reduced the risk of renewed aggression from opponents. Signal production also increased with audience size, yet strategies differed by age: while immatures reduced signalling in proximity of close-social partners, adults did so especially after receiving consolation. These results suggest that bonobos can flexibly adjust their emotion signalling to influence the outcome of PC events, and that this tendency has a developmental trajectory. Overall, these findings highlight the potential role that flexible emotion communication played in the sociality of our last common ancestor with Pan. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Bared-teeth displays in bonobos (Pan paniscus): An assessment of the power asymmetry hypothesis.

Facial expressions are key to navigating social group life. The Power Asymmetry Hypothesis of Motivational Emancipation predicts that the type of social organization shapes the meaning of communicative displays in relation to an individual's dominance rank. The bared-teeth (BT) display represents one of the most widely observed communicative signals across primate species. Studies in macaques indicate that the BT display in despotic species is often performed unidirectionally, from low- to high-ranking individuals (signaling submission), whereas the BT display in egalitarian species is usually produced irrespective of dominance (mainly signaling affiliation and appeasement). Despite its widespread presence, research connecting BT displays to the power asymmetry hypothesis outside the Macaca genus remains scarce. To extend this knowledge, we investigated the production of BT in relation to social dominance in dyadic interactions (N = 11,377 events) of 11 captive bonobos (Pan paniscus). Although adult bonobos were more despotic than previously suggested in the literature, BT displays were produced irrespective of dominance rank. Moreover, while adults produced the BT exclusively during socio-sexual interactions, especially during periods of social tension, immature bonobos produced the BT in a wider number of contexts. As such, the results indicate that the communicative meaning of the BT display is consistent with signaling appeasement, especially in periods of social tension. Moreover, the BT display does not seem to signal social status, supporting the prediction for species with a high degree of social tolerance. These results advance our understanding of the origins of communicative signals and their relation to species' social systems.

How 2- and 4-year-old children coordinate social interactions with peers.

The Interaction Engine Hypothesis postulates that humans have a unique ability and motivation for social interaction. A crucial juncture in the ontogeny of the interaction engine could be around 2-4 years of age, but observational studies of children in natural contexts are limited. These data appear critical also for comparison with non-human primates. Here, we report on focal observations on 31 children aged 2- and 4-years old in four preschools (10 h per child). Children interact with a wide range of partners, many infrequently, but with one or two close friends. Four-year olds engage in cooperative social interactions more often than 2-year olds and fight less than 2-year olds. Conversations and playing with objects are the most frequent social interaction types in both age groups. Children engage in social interactions with peers frequently (on average 13 distinct social interactions per hour) and briefly (28 s on average) and shorter than those of great apes in comparable studies. Their social interactions feature entry and exit phases about two-thirds of the time, less frequently than great apes. The results support the Interaction Engine Hypothesis, as young children manifest a remarkable motivation and ability for fast-paced interactions with multiple partners. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.

Coordinating social action: a primer for the cross-species investigation of communicative repair.

Human joint action is inherently cooperative, manifested in the collaborative efforts of participants to minimize communicative trouble through interactive repair. Although interactive repair requires sophisticated cognitive abilities, it can be dissected into basic building blocks shared with non-human animal species. A review of the primate literature shows that interactionally contingent signal sequences are at least common among species of non-human great apes, suggesting a gradual evolution of repair. To pioneer a cross-species assessment of repair this paper aims at (i) identifying necessary precursors of human interactive repair; (ii) proposing a coding framework for its comparative study in humans and non-human species; and (iii) using this framework to analyse examples of interactions of humans (adults/children) and non-human great apes. We hope this paper will serve as a primer for cross-species comparisons of communicative breakdowns and how they are repaired. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.

Every product needs a process: unpacking joint commitment as a process across species.

Joint commitment, the feeling of mutual obligation binding participants in a joint action, is typically conceptualized as arising by the expression and acceptance of a promise. This account limits the possibilities of investigating fledgling forms of joint commitment in actors linguistically less well-endowed than adult humans. The feeling of mutual obligation is one aspect of joint commitment (the product), which emerges from a process of signal exchange. It is gradual rather than binary; feelings of mutual obligation can vary in strength according to how explicit commitments are perceived to be. Joint commitment processes are more complex than simple promising, in at least three ways. They are affected by prior joint actions, which create precedents and conventions that can be embodied in material arrangements of institutions. Joint commitment processes also arise as solutions to generic coordination problems related to opening up, maintaining and closing down joint actions. Finally, during joint actions, additional, specific commitments are made piecemeal. These stack up over time and persist, making it difficult for participants to disengage from joint actions. These complexifications open up new perspectives for assessing joint commitment across species. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.

Revisiting the human 'interaction engine': comparative approaches to social action coordination.

The evolution of language was likely facilitated by a special predisposition for social interaction, involving a set of communicative and cognitive skills summarized as the 'interaction engine'. This assemblage seems to emerge early in development, to be found universally across cultures, and to enable participation in sophisticated joint action through the addition of spoken language. Yet, new evidence on social action coordination and communication in nonhuman primates warrants an update of the interaction engine hypothesis, particularly with respect to the evolutionary origins of its specific ingredients. However, one enduring problem for comparative research results from a conceptual gulf between disciplines, rendering it difficult to test concepts derived from human interaction research in nonhuman animals. The goal of this theme issue is to make such concepts accessible for comparative research, to promote a fruitful interdisciplinary debate on social action coordination as a new arena of research, and to enable mutual fertilization between human and nonhuman interaction research. In consequence, we here consider relevant theoretical and empirical research within and beyond this theme issue to revisit the interaction engine's shared, convergently derived and uniquely derived ingredients preceding (or perhaps in the last case, succeeding) human language. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.

Evidence of joint commitment in great apes' natural joint actions.

Human joint action seems special, as it is grounded in joint commitment-a sense of mutual obligation participants feel towards each other. Comparative research with humans and non-human great apes has typically investigated joint commitment by experimentally interrupting joint actions to study subjects' resumption strategies. However, such experimental interruptions are human-induced, and thus the question remains of how great apes naturally handle interruptions. Here, we focus on naturally occurring interruptions of joint actions, grooming and play, in bonobos and chimpanzees. Similar to humans, both species frequently resumed interrupted joint actions (and the previous behaviours, like grooming the same body part region or playing the same play type) with their previous partners and at the previous location. Yet, the probability of resumption attempts was unaffected by social bonds or rank. Our data suggest that great apes experience something akin to joint commitment, for which we discuss possible evolutionary origins.

Assessing joint commitment as a process in great apes.

Many social animals interact jointly, but only humans experience a specific sense of obligation toward their co-participants, a joint commitment. However, joint commitment is not only a mental state but also a process that reveals itself in the coordination efforts deployed during entry and exit phases of joint action. Here, we investigated the presence and duration of such phases in N = 1,242 natural play and grooming interactions of captive chimpanzees and bonobos. The apes frequently exchanged mutual gaze and communicative signals prior to and after engaging in joint activities with conspecifics, demonstrating entry and exit phases comparable to those of human joint activities. Although rank effects were less clear, phases in bonobos were more moderated by friendship compared to phases in chimpanzees, suggesting bonobos were more likely to reflect patterns analogous to human "face management". This suggests that joint commitment as process was already present in our last common ancestor with Pan.

Bonobos engage in joint commitment.

Joint action is central to human nature, enabling collectives to achieve goals otherwise unreachable by individuals. It is enabled by humans' capacity to understand and engage in joint commitments. Joint commitments are evidenced when partners in interrupted joint actions reengage one another. To date, there is no clear evidence whether nonhuman animals understand joint commitment, suggesting that only humans experience it. Here, we revisit this claim by interrupting bonobos engaged in social activities. Bonobos reliably resumed the activity, and the likelihood of resumption was higher for social compared to solitary activities. Furthermore, communicative efforts deployed to suspend and resume social activities varied depending on partners' social relationships and interactive roles. Our results suggest that bonobos, like humans, engage in joint commitment and have some awareness of the social consequences of breaking it.

An exploration of Menzerath's law in wild mountain gorilla vocal sequences.

Menzerath's law, traditionally framed as a negative relationship between the size of a structure and its constituent parts (e.g. sentences with more clauses have shorter clauses), is widespread across information-coding systems ranging from human language and the vocal and gestural sequences of primates and birds, to the building blocks of DNA, genes and proteins. Here, we analysed an extensive dataset of 'close-call' sequences produced by wild mountain gorillas (Gorilla beringei beringei, no. individuals = 10, no. sequences = 2189) to determine whether, in accordance with Menzerath's law, a negative relationship existed between the number of vocal units in a sequence and the duration of its constituent units. We initially found positive evidence for this but, on closer inspection, the negative relationship was driven entirely by the difference between single- and multi-unit (two to six unit) sequences. Once single-unit sequences were excluded from the analysis, we identified a relationship in the opposite direction, with longer sequences generally composed of longer units. The close-call sequences of mountain gorillas therefore represent an intriguing example of a non-human vocal system that only partially conforms to the predictions of Menzerath's law.

Linguistic laws in chimpanzee gestural communication.

Studies testing linguistic laws outside language have provided important insights into the organization of biological systems. For example, patterns consistent with Zipf's law of abbreviation (which predicts a negative relationship between word length and frequency of use) have been found in the vocal and non-vocal behaviour of a range of animals, and patterns consistent with Menzerath's law (according to which longer sequences are made up of shorter constituents) have been found in primate vocal sequences, and in genes, proteins and genomes. Both laws have been linked to compression-the information theoretic principle of minimizing code length. Here, we present the first test of these laws in animal gestural communication. We initially did not find the negative relationship between gesture duration and frequency of use predicted by Zipf's law of abbreviation, but this relationship was seen in specific subsets of the repertoire. Furthermore, a pattern opposite to that predicted was seen in one subset of gestures-whole body signals. We found a negative correlation between number and mean duration of gestures in sequences, in line with Menzerath's law. These results provide the first evidence that compression underpins animal gestural communication, and highlight an important commonality between primate gesturing and language.

Social play as joint action: A framework to study the evolution of shared intentionality as an interactional achievement.

Social play has a complex, cooperative nature that requires substantial coordination. This has led researchers to use social games to study cognitive abilities like shared intentionality, the skill and motivation to share goals and intentions with others during joint action. We expand this proposal by considering play as a joint action and examining how shared intentionality is achieved during human joint action. We describe how humans get into, conduct, and get out of joint actions together in an orderly way, thereby constructing the state of "togetherness" characteristic of shared intentionality. These processes play out as three main phases, the opening (where participants are ratified and joint commitments are established), the main body (where progress, ongoing commitments, and possible role reversals are coordinated), and the closing (where the intention to terminate the action is coordinated and where participants take leave of each other). We use this process in humans as a framework for examining how various animal species get into, maintain, and get out of play bouts. This comparative approach constitutes an alternative measure of those species' possession of shared intentionality. Using this framework, we review the play literature on human children and different social species of mammals and birds in search of behavioral markers of shared intentionality in the coordination of play bouts. We discuss how our approach could shed light on the evolution of the special human motivation to cooperate and share psychological states with others.