Selective and prolonged attention to emotional scenes in humans and bonobos.

Perceiving emotions in others is at the foundation of higher-order social cognition. The importance of emotions is evidenced by the fact that they receive prioritized attention at early stages of processing the environment in humans and some other primates. Nevertheless, we do not fully understand how emotion modulates attention over longer durations in primates, particularly in great apes. Bonobos, one of our closest relatives, stand out in emotion processing and regulation among great apes. This makes them an interesting comparison species and a valuable model for studying the evolution of emotion perception in hominids. We investigated how bonobos and humans spontaneously attend to emotionally valent scenes in a preferential looking task using eye-tracking. With Bayesian mixed modelling, we found that bonobos and humans generally looked longer at emotional scenes, mainly of conspecifics. Moreover, while bonobos did not have a bias toward emotional human scenes, humans sustained their attention toward bonobos playing, grooming and having sex. Furthermore, when exploring an immediate bias for emotions, humans showed a bias toward affiliative human scenes, and bonobos showed a bias away from bonobos-in-distress scenes. These findings suggest that emotions modulate attention at early and later attentional stages in bonobos, similar to humans.

Merging sociality and robotics through an evolutionary perspective.

Robotics, using social mechanisms like hormonal modulation, may accelerate our understanding of core sociality principles.

Preliminary evidence that eye appearance in parrots (Psittaciformes) co-varies with latitude and altitude.

External eye appearance in avian taxa has been proposed to be driven by social and ecological functions. Recent research in primates suggests, instead, that, photoprotective functions are important drivers of external eye appearance. Using similar methods, we examined the variation in external eye appearance of 132 parrot species (Psittaciformes) in relation to their ecology and sociality. Breeding systems, flock size and sexual dimorphism, as well as species' latitude and maximum living altitude, and estimated UV-B incidence in species' ranges were used to explore the contribution of social and ecological factors in driving external eye appearance. We measured the hue and brightness of visible parts of the eye and the difference in measurements of brightness between adjacent parts of the eye. We found no link between social variables and our measurements. We did, however, find a negative association between the brightness of the inner part of the iris and latitude and altitude. Darker inner irises were more prevalent farther away from the equator and for those species living at higher altitudes. We found no link between UV-B and brightness measurements of the iris, or tissue surrounding the eye. We speculate that these results are consistent with an adaptation for visual functions. While preliminary, these results suggest that external eye appearance in parrots is influenced by ecological, but not social factors.

No immediate attentional bias towards or choice bias for male secondary sexual characteristics in Bornean orang-utans (Pongo pygmaeus).

Primate faces provide information about a range of variant and invariant traits, including some that are relevant for mate choice. For example, faces of males may convey information about their health or genetic quality through symmetry or facial masculinity. Because perceiving and processing such information may have bearing on the reproductive success of an individual, cognitive systems are expected to be sensitive to facial cues of mate quality. However, few studies have investigated this topic in non-human primate species. Orang-utans are an interesting species to test mate-relevant cognitive biases, because they are characterised by male bimaturism: some adult males are fully developed and bear conspicuous flanges on the side of their face, while other males look relatively similar to females. Here, we describe two non-invasive computerised experiments with Bornean orang-utans (Pongo pygmaeus), testing (i) immediate attention towards large flanges and symmetrical faces using a dot-probe task (N = 3 individuals; 2F) and (ii) choice bias for pictures of flanged males over unflanged males using a preference test (N = 6 individuals; 4F). In contrast with our expectations, we found no immediate attentional bias towards either large flanges or symmetrical faces. In addition, individuals did not show a choice bias for stimuli of flanged males. We did find exploratory evidence for a colour bias and energy efficiency trade-offs in the preference task. We discuss our null results and exploratory results in the context of the evolutionary history of Bornean orang-utans, and provide suggestions for a more biocentric approach to the study of orang-utan cognition.

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.

Basic Emotions or Constructed Emotions: Insights From Taking an Evolutionary Perspective.

The ongoing debate between basic emotion theories (BETs) and the theory of constructed emotion (TCE) hampers progress in the field of emotion research. Providing a new perspective, here we aim to bring the theories closer together by dissecting them according to Tinbergen's four questions to clarify a focus on their evolutionary basis. On the basis of our review of the literature, we conclude that whereas BETs focus on the evolution question of Tinbergen, the TCE is more concerned with the causation of emotion. On the survival value of emotions both theories largely agree: to provide the best reaction in specific situations. Evidence is converging on the evolutionary history of emotions but is still limited for both theories-research within both frameworks focuses heavily on the causation. We conclude that BETs and the TCE explain two different phenomena: emotion and feeling. Therefore, they seem irreconcilable but possibly supplementary for explaining and investigating the evolution of emotion-especially considering their similar answer to the question of survival value. Last, this article further highlights the importance of carefully describing what aspect of emotion is being discussed or studied. Only then can evidence be interpreted to converge toward explaining emotion.

A Cross-Cultural Comparison on Implicit and Explicit Attitudes Towards Artificial Agents.

Historically, there has been a great deal of confusion in the literature regarding cross-cultural differences in attitudes towards artificial agents and preferences for their physical appearance. Previous studies have almost exclusively assessed attitudes using self-report measures (i.e., questionnaires). In the present study, we sought to expand our knowledge on the influence of cultural background on explicit and implicit attitudes towards robots and avatars. Using the Negative Attitudes Towards Robots Scale and the Implicit Association Test in a Japanese and Dutch sample, we investigated the effect of culture and robots' body types on explicit and implicit attitudes across two experiments (total n = 669). Partly overlapping with our hypothesis, we found that Japanese individuals had a more positive explicit attitude towards robots compared to Dutch individuals, but no evidence of such a difference was found at the implicit level. As predicted, the implicit preference towards humans was moderate in both cultural groups, but in contrast to what we expected, neither culture nor robot embodiment influenced this preference. These results suggest that only at the explicit but not implicit level, cultural differences appear in attitudes towards robots. Supplementary Information: The online version contains supplementary material available at 10.1007/s12369-022-00917-7.

Facial mimicry and metacognitive judgments in emotion recognition are distinctly modulated by social anxiety and autistic traits.

Facial mimicry as well as the accurate assessment of one's performance when judging others' emotional expressions have been suggested to inform successful emotion recognition. Differences in the integration of these two information sources might explain alterations in the perception of others' emotions in individuals with Social Anxiety Disorder and individuals on the autism spectrum. Using a non-clinical sample (N = 57), we examined the role of social anxiety and autistic traits in the link between facial mimicry, or confidence in one's performance, and emotion recognition. While participants were presented with videos of spontaneous emotional facial expressions, we measured their facial muscle activity, asked them to label the expressions and indicate their confidence in accurately labelling the expressions. Our results showed that confidence in emotion recognition was lower with higher social anxiety traits even though actual recognition was not related to social anxiety traits. Higher autistic traits, in contrast, were associated with worse recognition, and a weakened link between facial mimicry and performance. Consequently, high social anxiety traits might not affect emotion recognition itself, but the top-down evaluation of own abilities in emotion recognition contexts. High autistic traits, in contrast, may be related to lower integration of sensorimotor simulations, which promote emotion recognition.

How video calls affect mimicry and trust during interactions.

Many social species, humans included, mimic emotional expressions, with important consequences for social bonding. Although humans increasingly interact via video calls, little is known about the effect of these online interactions on the mimicry of scratching and yawning, and their linkage with trust. The current study investigated whether mimicry and trust are affected by these new communication media. Using participant-confederate dyads (n = 27), we tested the mimicry of four behaviours across three different conditions: watching a pre-recorded video, online video call, and face-to-face. We measured mimicry of target behaviours frequently observed in emotional situations, yawn and scratch and control behaviours, lip-bite and face-touch. In addition, trust in the confederate was assessed via a trust game. Our study revealed that (i) mimicry and trust did not differ between face-to-face and video calls, but were significantly lower in the pre-recorded condition; and (ii) target behaviours were significantly more mimicked than the control behaviours. This negative relationship can possibly be explained by the negative connotation usually associated with the behaviours included in this study. Overall, this study showed that video calls might provide enough interaction cues for mimicry to occur in our student population and during interactions between strangers. This article is part of a discussion meeting issue 'Face2face: advancing the science of social interaction'.

Individual attractiveness preferences differentially modulate immediate and voluntary attention.

Physical attractiveness plays a crucial role in mate choice for both men and women. This is reflected in visual attention: people immediately attend towards and look longer at attractive faces, especially when they are motivated to find a partner. However, previous studies did not incorporate real-life dating decisions. Here, we aimed to combine attentional tasks with individual attractiveness ratings and a real-life mate choice context, namely a speed-dating paradigm. We investigated whether heterosexual non-committed young adults showed biases in immediate and voluntary attention towards attractive faces and preferred dating partners. In line with previous research, we found considerable individual differences in individual attractiveness preferences. Furthermore, our results showed that men had a bias towards attractive faces and preferred dating partners in the immediate attention task, while results for women were mixed. In the voluntary attention task, however, both men and women had an attentional bias towards attractive faces and preferred dating partners. Our results suggest that individual attractiveness preferences are good predictors of especially voluntary attention. We discuss these findings from an evolutionary perspective and suggest directions for future research.

Attention toward emotions is modulated by familiarity with the expressor: A comparison between bonobos and humans.

Why can humans be intolerant of, yet also be empathic toward strangers? This cardinal question can be tackled by studying emotions in our closest living relatives, bonobos. Their striking xenophilic tendencies make them an interesting model for reconstructing the socioemotional capacities of the last common ancestor of hominids. Within two dot-probe studies, we compared bonobos' and humans' attention toward scenes depicting familiar (kith and kin) or unfamiliar individuals with emotional or neutral expressions. Results show that the attention of bonobos is biased toward emotional scenes depicting unfamiliar bonobos, but not toward emotional groupmates (Study 1). In contrast, Study 2 shows that human attention is biased toward emotional rather than neutral expressions of family and friends, but not toward unfamiliar others. On the one hand, our results show that an attentional bias toward emotions is a shared phenomenon between humans and bonobos, but on the other, both species have their own unique evolutionarily informed biases. These findings support previously proposed adaptive explanations for xenophilia in bonobos that potentially biases them toward emotional expressions of unfamiliar conspecifics, and parochialism in humans, which makes them sensitive to the emotional expressions of close others. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Ecological factors are likely drivers of eye shape and colour pattern variations across anthropoid primates.

External eye appearance across primate species is diverse in shape and colouration, yet we still lack an explanation for the drivers of such diversity. Here we quantify substantial interspecific variation in eye shape and colouration across 77 primate species representing all extant genera of anthropoid primates. We reassess a series of hypotheses aiming to explain ocular variation in horizontal elongation and in colouration across species. Heavier body weight and terrestrial locomotion are associated with elongated eye outlines. Species living closer to the equator present more pigmented conjunctivae, suggesting photoprotective functions. Irises become bluer in species living further away from the equator, adding to existing literature supporting a circadian clock function for bluer irises. These results shift the current focus from communicative, to ecological factors in driving variation in external eye appearance in anthropoid primates. They also highlight the possibility that similar ecological factors contributed to selection for blue eyes in ancestral human populations living in northern latitudes.

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.

My Fear Is Not, and Never Will Be, Your Fear: On Emotions and Feelings in Animals.

Do nonhuman animals (henceforth, animals) have emotions, and if so, are these similar to ours? This opinion piece aims to add to the recent debate about this question and provides a critical re-evaluation of what can be concluded about animal and human emotions. Emotions, and their cognitive interpretation, i.e., feelings, serve important survival functions. Emotions, we believe, can exist without feelings and are unconsciously influencing our behavior more than we think, and possibly more so than feelings do. Given that emotions are expressed in body and brain, they can be inferred from these measures. We view feelings primarily as private states, which may be similar across closely related species but remain mostly inaccessible to science. Still, combining data acquired through behavioral observation with data obtained from noninvasive techniques (e.g., eyetracking, thermography, hormonal samples) and from cognitive tasks (e.g., decision-making paradigms, cognitive bias, attentional bias) provides new information about the inner states of animals, and possibly about their feelings as well. Given that many other species show behavioral, neurophysiological, hormonal, and cognitive responses to valenced stimuli equivalent to human responses, it seems logical to speak of animal emotions and sometimes even of animal feelings. At the very least, the contemporary multi-method approach allows us to get closer than ever before. We conclude with recommendations on how the field should move forward.

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.

Can third-party observers detect attraction in others based on subtle nonverbal cues?

In a series of three studies, we examined whether third-party observers can detect attraction in others based on subtle nonverbal cues. We employed video segments of dates collected from a speed-dating experiment, in which daters went on a brief (approx. 4 min) blind-date and indicated whether they would like to go on another date with their brief interaction partner or not. We asked participants to view these stimuli and indicate whether or not each couple member is attracted to their partner. Our results show that participants could not reliably detect attraction, and this ability was not influenced by the age of the observer, video segment location (beginning or middle of the date), video duration, or general emotion recognition capacity. Contrary to previous research findings, our findings suggest that third-party observers cannot reliably detect attraction in others. However, there was one exception: Recognition rose above chance level when the daters were both interested in their partners compared to when they were not interested. Supplementary Information: The online version contains supplementary material available at 10.1007/s12144-022-02927-0.

Reading Your Emotions in My Physiology? Reliable Emotion Interpretations in Absence of a Robust Physiological Resonance.

Affective states are expressed in an individual's physical appearance, ranging from facial expressions and body postures, to indicators of physiological arousal (e.g., a blush). Confirming the claimed communicative function of these markers, humans are capable of distinguishing between a variety of discrete emotion displays. In an attempt to explain the underlying mechanism, characteristic bodily changes within the observer, including physiological arousal and mimicry, have been suggested to facilitate the interpretation of an expression. The current study aims to create a holistic picture of emotion perception by (1) using three different sources of emotional information (prototypical facial expressions, bodily expressions, and subtle facial cues) and (2) measuring changes in multiple physiological signals (facial electromyography, skin conductance level, skin temperature, and pupil size). While participants clearly discriminated between perceived emotional expressions, there was no overall 1-1 correspondence with their physiological responses. Some specific but robust effects were observed. Angry facial expressions were consistently responded to with a peak in skin conductance level. Furthermore, sad body expressions were associated with a drop in skin temperature. In addition to being the best recognized expression, viewing happy faces elicited congruent facial muscle responses, which supports the potential role of embodied simulation in emotion recognition. Lastly, tears were not only rated as highly emotional intense but also evoked a peak in skin conductance level in the observer. The absence of distinct physiological responses to other expressions could be explained by the lacking functionality of affect sharing in a non-interactive experimental context. Consequentially, emotional alignment in body and mind might especially take place in real social situations, which should be considered in future research. Supplementary Information: The online version contains supplementary material available at 10.1007/s42761-021-00083-5.

Reading emotional faces in deaf and hard-of-hearing and typically hearing children.

Reading emotions from other people's facial expressions is an important skill that guides social interactions. With limited auditory input and atypical emotion socialization, deaf and hard-of-hearing (DHH) children may develop atypical processing patterns when reading emotional faces. The current study aimed at understanding whether and how DHH and typically hearing (TH) children differed at 3 emotion processing levels: gaze patterns, physiological arousal, and interpretation. Fifty-five DHH children and 72 TH children completed an emotional face matching task in which they were presented with happy, angry, fearful, and emotionally neutral faces. During the task participants' eye gazes and pupil diameter were measured by an eye-tracking device. The DHH and TH children both paid most attention to the eye region when reading emotional faces. Yet, a contrast between happy faces and nonhappy faces was observed in physiological arousal and interpretation tendency in the DHH children only: Nonhappy facial expressions were more arousing and were confused more often than happy expressions, which may reflect the DHH children being less experienced in processing nonhappy expressions due to limited access to the social environment. The results highlighted the importance of looking into the qualitative differences between typical and atypical development. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Attention Biases to Threat in Infants and Parents: Links to Parental and Infant Anxiety Dispositions.

Parent-to-child transmission of information processing biases to threat is a potential causal mechanism in the family aggregation of anxiety symptoms and traits. This study is the first to investigate the link between infants' and parents' attention bias to dynamic threat-relevant (versus happy) emotional expressions. Moreover, the associations between infant attention and anxiety dispositions in infants and parents were explored. Using a cross-sectional design, we tested 211 infants in three age groups: 5-to-7-month-olds (n = 71), 11-to-13-month-olds (n = 73), and 17-to-19-month-olds (n = 67), and 216 parents (153 mothers). Infant and parental dwell times to angry and fearful versus happy facial expressions were measured via eye-tracking. The parents also reported on their anxiety and stress. Ratings of infant temperamental fear and distress were averaged across both parents. Parents and infants tended to show an attention bias for fearful faces with marginally longer dwell times to fearful versus happy faces. Parents dwelled longer on angry versus happy faces, whereas infants showed an avoidant pattern with longer dwell times to happy versus angry expressions. There was a significant positive association between infant and parent attention to emotional expressions. Parental anxiety dispositions were not related to their own or their infant's attention bias. No significant link emerged between infants' temperament and attention bias. We conclude that an association between parental and infant attention may already be evident in the early years of life, whereas a link between anxiety dispositions and attention biases may not hold in community samples.