{"title":"The interpersonal and intrapsychic functions of the pride expression.","authors":"Jessica L Tracy, Zachary Witkower","doi":"10.1098/rstb.2025.0149","DOIUrl":"10.1098/rstb.2025.0149","url":null,"abstract":"<p><p>A large body of research suggests that pride is associated with a distinct, cross-culturally recognized and spontaneously displayed nonverbal expression. Given its apparent universality, this expression is likely to have evolved as a functional signal, benefitting both receivers and senders. Here, we focus primarily on the intrapsychic benefits received by those who express pride. We review research suggesting that the expression communicates an individual's deservedness of high status to others, and that it does so implicitly and across diverse cultural contexts. We then discuss the expression's function in the context of dual-strategies theory, which suggests that there are two distinct forms of high rank: prestige, based on earned respect, and dominance, based on threat of force. Studies suggest that the pride expression reliably and cross-culturally communicates prestige, but not dominance. Finally, we discuss functions of the pride expression that go beyond the display's signal value. In sum, the pride nonverbal expression appears to be a highly functional display that likely evolved to facilitate humans' adaptive responding to opportunities for social rank attainment. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601887/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032997","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Comparative neuroanatomy of emotion expression: from flies to humans.","authors":"Ralph Adolphs","doi":"10.1098/rstb.2025.0135","DOIUrl":"10.1098/rstb.2025.0135","url":null,"abstract":"<p><p>All animals exhibit complex behaviours that share features with those that in humans we call 'emotional expressions'. Sidestepping contentious issues about the conscious experience of emotion, their expression across species provides a wealth of detail about their emergence in evolution, the shared features that distinguish them from reflexes, and principles for their neural control. This paper provides a high-level survey of this vast field. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601884/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032949","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Luigi Baciadonna, Catherine Macri, David Baracchi, Martin Giurfa
{"title":"Exploring emotional expression in invertebrates: mechanisms, functions and phylogeny.","authors":"Luigi Baciadonna, Catherine Macri, David Baracchi, Martin Giurfa","doi":"10.1098/rstb.2025.0146","DOIUrl":"10.1098/rstb.2025.0146","url":null,"abstract":"<p><p>The idea that invertebrates may exhibit basic forms of emotion, and possibly sentience, is gaining traction as evidence of their advanced cognitive abilities continues to mount. While high-level cognition does not imply emotional states, invertebrates offer a unique window into whether such states can arise in small-brained animals. With their relatively simple yet highly structured nervous systems, they serve as tractable models for dissecting the mechanisms underlying emotional expression. To probe these processes, researchers have developed behavioural paradigms exposing different species to contexts likely to elicit emotional responses. From a phylogenetic perspective, studying emotional expression in invertebrates can reveal whether basic emotional systems are conserved across species. This review synthesizes current knowledge of non-verbal emotional expression in social insects, focusing on mechanisms and functional roles. We apply a multi-component approach to examine key building blocks shared across emotions and taxa, offering insights into their evolutionary origins. We also identify critical gaps in knowledge and propose directions for future research. By examining the proximate and ultimate causes of emotion, this framework enhances our understanding of the biological basis and evolutionary significance of emotions, promoting interdisciplinary research that bridges neuroscience, ethology and evolutionary biology. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601889/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032966","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"The play fever: individual and group factors influence the contagion of play behaviour within social hominid groups.","authors":"Ivan Norscia, Edoardo Collarini, Giada Cordoni","doi":"10.1098/rstb.2025.0144","DOIUrl":"10.1098/rstb.2025.0144","url":null,"abstract":"<p><p>Via the perception-action model and mirror neuron system, individuals replicate others' behaviour (behavioural/social contagion) and reduce prediction errors on other behaviours. Among contagious behaviours, play offers a safe context for cooperation, competition and tolerance. Thus, play contagion is interesting to better understand the evolution of social dynamics in many non-primates and primates, including humans. To our knowledge, no study has examined play contagion across social hominids nor analysed individual/group features to identify common modulators. In 2022-2025, we gathered all-occurrences data on play, affiliative and agonistic patterns in social hominids (two groups of captive chimpanzees, bonobos and gorillas; and two toddler classes of ≤ 3 years and their educators). In the 3 min period, we checked if individuals not involved in play would start playing either after the onset of a trigger play session (post-play (PP)) or in absence of any previous play session (matched-control). Play occurrence was highest during PP, informing play contagion. Contagion increased with higher male-to-female, immature-to-adult and affiliative-to-agonistic interaction ratios. Males, youngsters and individuals that were in trigger proximity (≤ 10 m) showed the highest contagion. Shared across social hominids, play contagion and its common modulators might trace back to their last common ancestor, underlying the transmission of behaviour, and possible underlying emotions, within groups. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601880/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032995","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Vocal display rules in everyday life: when is it appropriate to laugh, sigh or scream?","authors":"Roza G Kamiloğlu","doi":"10.1098/rstb.2025.0151","DOIUrl":"10.1098/rstb.2025.0151","url":null,"abstract":"<p><p>The appropriateness of common nonverbal vocalizations is guided by social norms, known as display rules. This preregistered, 7-day experience-sampling study examined how perceived appropriateness varies across situations. A total of 142 adults completed three surveys per day (2670 surveys total; 3821 vocalizations), rating appropriateness alongside situational cues and psychological situation characteristics. Laughter and sighs were most often judged appropriate, whereas screams and retches were least. Appropriateness was higher at home and in social places (e.g. parties) than in work/school or public settings (e.g. library), and higher with close ties than with weak ties or strangers. Formality and power were associated with lower appropriateness, whereas sociality and positivity were associated with higher appropriateness. Effects were vocalization-specific: for example, laughter was considered more appropriate in positive, social and brisk contexts, but less appropriate in formal and high-power contexts. By charting when and where vocalizations are judged acceptable, these findings outline situational principles of vocal display rules in everyday life. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601882/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149033019","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Expressing emotions through music.","authors":"Vesa Putkinen, Maya Rassouli, Lauri Nummenmaa","doi":"10.1098/rstb.2025.0131","DOIUrl":"10.1098/rstb.2025.0131","url":null,"abstract":"<p><p>Music evokes strong emotions across listeners although it conveys no biologically relevant information regarding the survival challenges in the environment. Here we review the neurobiology and psychology of emotional communication through music, focussing on the functions and mechanisms of musical emotional expressions in social interaction. We also extend our analysis of the communicative space beyond the pure acoustic features to the interaction of musical emotional cues with other verbal and non-verbal communication channels. Activation likelihood estimation meta-analysis of neuroimaging studies shows functional overlap for brain systems involved in music production and emotions. Musical emotion communication, however, differs from other common expressive channels (faces and bodies, utterances and semantics) in its timescale, context, spontaneity and interactivity. Musical emotional communication can be modulated by affective information conveyed through verbal and other non-verbal channels. This makes music a flexible and culturally malleable means of context-dependent emotional communication that is particularly well suited for facilitating large-scale social affiliation and bonding. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601867/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032933","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Drew H Abney, Stephen J Sheinkopf, Samantha G Mitsven, Ronald Seifer, Barry M Lester, Ed Z Tronick, Daniel S Messinger
{"title":"Mutual positive engagement with mother is uniquely associated with increases in human infant heart rate variability.","authors":"Drew H Abney, Stephen J Sheinkopf, Samantha G Mitsven, Ronald Seifer, Barry M Lester, Ed Z Tronick, Daniel S Messinger","doi":"10.1098/rstb.2026.0004","DOIUrl":"10.1098/rstb.2026.0004","url":null,"abstract":"<p><p>The ability of human infants to dynamically adapt their physiological states to changing social demands is critical for the development of emotion regulation. We investigated infant parasympathetic dynamics related to emotion regulation-changes in respiratory sinus arrhythmia (RSA)-using a large sample of 974 at-risk infant-mother dyads (infant age: 4 months). Infant RSA was elevated during bouts of positive engagement matching with their mothers, characterized by mutual positive engagement. Specifically, infant RSA began to increase more than 5 seconds before the onset of bouts of positive engagement matching and remained elevated for at least 4 seconds after bout onsets. These periods were substantially extended when the bouts began with positive expressions from the mother. Strikingly, there were no increases in infant RSA before or during bouts of either infant or mother solo positive engagement. Findings from these exploratory analyses suggest the possibility of bi-directional associations between ongoing social interactions and infant parasympathetic reactivity in which positive mother expressions support increases in infant RSA that in turn support prolonged infant engagement in mutual positivity. Thus, early social interaction is not simply an elicitor of infant positive expressions. Instead, mutual positive expressivity may facilitate infant neurophysiological changes that underlie emotion regulation. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601893/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032974","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"On the nature of grief: uncovering the evolutionary roots of loss across the animal realm.","authors":"Sarv Dashti, Andre Goncalves","doi":"10.1098/rstb.2025.0147","DOIUrl":"10.1098/rstb.2025.0147","url":null,"abstract":"<p><p>Understanding grief in non-human animals poses conceptual and empirical challenges. Unlike humans, whose grief is readily identified through affective, cognitive and behavioural manifestations, recognizing it in other species is complicated by differences in expression and by anthropomorphic bias. Research on grief has taken several approaches. Observational studies in the ethological tradition have documented grief-like responses (withdrawal, vocalizations and changes in social interaction) across diverse taxa. Separation studies have shown that disrupting attachment produces depressive-like behaviour both behaviourally and physiologically, underlining how early bonds shape later affect. Questionnaire studies have explored perceptions of grief in companion animals, revealing both cross-species continuities and potential interpretive biases. Meanwhile, evolutionary hypotheses for grief range from non-functional accounts, through functional accounts (either cognitive or socially bound), to integrative/mechanistic models. Across these lines of evidence, we suggest that grief is the response to the rupture of a social bond built by a deeply conserved attachment system and its homologous nonapeptide toolkit. We propose an inclusive definition of grief as a response to the disruption of significant social bonds, encompassing affective, cognitive and behavioural components across species where the body is either present or absent. This definition is intended to ground systematic, comparative study of grief across species. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601878/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149033041","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"The expression of awe: evolutionary origins and cultural elaborations of a basic emotion.","authors":"Dacher Keltner, Maria Monroy","doi":"10.1098/rstb.2025.0148","DOIUrl":"10.1098/rstb.2025.0148","url":null,"abstract":"<p><p>Awe is the emotion people experience when encountering stimuli that are vast and beyond their current knowledge structures. In this article, we take a basic emotion approach to the study of the expression of awe, reviewing studies of awe-related movements of facial muscles, patterns of vocalization and bodily and postural movements. We first present evidence suggesting that awe indeed is a distinct emotional state. Guided by advances in the study of multi-modal emotional expression, we then review studies of how people express awe in a pattern of mouth and gaze activity, specific vocal bursts and head orientation upward, unfolding in coordinated patterns of expressive behaviour that have some degree of universality. Building upon recent theorizing in cultural evolution, we detail how the expression of awe is elaborated into cultural forms, including folklore, kinds of music (choral music), dance and portrayals of the sublime in visual arts. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13601885/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149032991","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"From seeing to showing: towards a neuroscience of emotion production.","authors":"Seohyeon Lee, Solha Hwang, M Justin Kim","doi":"10.1098/rstb.2025.0136","DOIUrl":"10.1098/rstb.2025.0136","url":null,"abstract":"<p><p>Facial expressions are considered a critical feature in affective neuroscience. Consequently, extensive research has focused on how emotional facial expressions are perceived, including investigations into which brain regions are activated during their observation and how the brain processes their social significance. To advance a more comprehensive understanding of emotion, it is essential to examine all three aspects-perception, experience and production-in a balanced manner. From an evolutionary perspective, the production of facial expressions during emotional experiences is particularly significant, as it reflects both adaptive communication and survival functions. Nevertheless, a review of existing databases reveals that research on the production of facial expressions remains scarce compared with studies on perception. This gap may stem from several methodological constraints, including the difficulty of reliably inducing emotional experiences in laboratory settings, the challenge of recording participants' full facial expressions in a magnetic resonance imaging scanner and limitations in current analytical approaches for facial expressions. In this review, we argue for the importance of studying the production of facial expressions and outline potential strategies to address these challenges. This article is part of the theme issue 'Mechanisms, development, phylogeny and functions of emotional expressions'.</p>","PeriodicalId":520748,"journal":{"name":"Philosophical transactions of the Royal Society of London. Series B, Biological sciences","volume":"381 1959","pages":""},"PeriodicalIF":4.7,"publicationDate":"2026-09-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13644780/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"149033007","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":0,"RegionCategory":"","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}