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Communicating emotional intensity plays a vital ecological role because it provides valuable information about the nature and likelihood of the sender's behavior. 1 , 2 Yet how this complex signaling task is achieved remains unknown. We addressed this question using a perception-based, data-driven method to mathematically model the specific facial movements that receivers use to classify the six basic emotions-"happy," "surprise," "fear," "disgust," "anger," and "sad"-and judge their intensity in two distinct cultures (East Asian, Western European; total n = 120). In both cultures, receivers expected facial expressions to dynamically represent emotion category and intensity information over time, using a multi-component compositional signaling structure. Specifically, emotion intensifiers peaked earlier or later than emotion classifiers and represented intensity using amplitude variations. Emotion intensifiers are also more similar across emotions than classifiers are, suggesting a latent broad-plus-specific signaling structure. Cross-cultural analysis further revealed similarities and differences in expectations that could impact cross-cultural communication. Specifically, East Asian and Western European receivers have similar expectations about which facial movements represent high intensity for threat-related emotions, such as "anger," "disgust," and "fear," but differ on those that represent low threat emotions, such as happiness and sadness. Together, our results provide new insights into the intricate processes by which facial expressions can achieve complex dynamic signaling tasks by revealing the rich information embedded in facial expressions., 3 For example, attack often follows signals of intense aggression if receivers fail to retreat. 4 , 5 Humans regularly use facial expressions to communicate such information. 6 , 7 , 8 , 9 , 10 , 11 Yet how this complex signaling task is achieved remains unknown. We addressed this question using a perception-based, data-driven method to mathematically model the specific facial movements that receivers use to classify the six basic emotions-"happy," "surprise," "fear," "disgust," "anger," and "sad"-and judge their intensity in two distinct cultures (East Asian, Western European; total n = 120). In both cultures, receivers expected facial expressions to dynamically represent emotion category and intensity information over time, using a multi-component compositional signaling structure. Specifically, emotion intensifiers peaked earlier or later than emotion classifiers and represented intensity using amplitude variations. Emotion intensifiers are also more similar across emotions than classifiers are, suggesting a latent broad-plus-specific signaling structure. Cross-cultural analysis further revealed similarities and differences in expectations that could impact cross-cultural communication. Specifically, East Asian and Western European receivers have similar expectations about which facial movements represent high intensity for threat-related emotions, such as "anger," "disgust," and "fear," but differ on those that represent low threat emotions, such as happiness and sadness. Together, our results provide new insights into the intricate processes by which facial expressions can achieve complex dynamic signaling tasks by revealing the rich information embedded in facial expressions., Competing Interests: Declaration of interests P.G.S. is on the advisory board of this Current Biology., (Copyright © 2023 The Author(s). Published by Elsevier Inc. All rights reserved.)

Models are the hallmark of mature scientific inquiry. In psychology, this maturity has been reached in a pervasive question-what models best represent facial expressions of emotion? Several hypotheses propose different combinations of facial movements [action units (AUs)] as best representing the six basic emotions and four conversational signals across cultures. We developed a new framework to formalize such hypotheses as predictive models, compare their ability to predict human emotion categorizations in Western and East Asian cultures, explain the causal role of individual AUs, and explore updated, culture-accented models that improve performance by reducing a prevalent Western bias. Our predictive models also provide a noise ceiling to inform the explanatory power and limitations of different factors (e.g., AUs and individual differences). Thus, our framework provides a new approach to test models of social signals, explain their predictive power, and explore their optimization, with direct implications for theory development.

Human facial expressions are complex, multi-component signals that can communicate rich information about emotions, 1-5 including specific categories, such as "anger," and broader dimensions, such as "negative valence, high arousal." 6-8 An enduring question is how this complex signaling is achieved. Communication theory predicts that multi-component signals could transmit each type of emotion information-i.e., specific categories and broader dimensions-via the same or different facial signal components, with implications for elucidating the system and ontology of facial expression communication. 9 We addressed this question using a communication-systems-based method that agnostically generates facial expressions and uses the receiver's perceptions to model the specific facial signal components that represent emotion category and dimensional information to them. 10-12 First, we derived the facial expressions that elicit the perception of emotion categories (i.e., the six classic emotions 13 plus 19 complex emotions 3 ) and dimensions (i.e., valence and arousal) separately, in 60 individual participants. Comparison of these facial signals showed that they share subsets of components, suggesting that specific latent signals jointly represent-i.e., multiplex-categorical and dimensional information. Further examination revealed these specific latent signals and the joint information they represent. Our results-based on white Western participants, same-ethnicity face stimuli, and commonly used English emotion terms-show that facial expressions can jointly represent specific emotion categories and broad dimensions to perceivers via multiplexed facial signal components. Our results provide insights into the ontology and system of facial expression communication and a new information-theoretic framework that can characterize its complexities., Competing Interests: Declaration of interests The authors declare no competing interests., (Crown Copyright © 2021. Published by Elsevier Inc. All rights reserved.)

Real-world studies show that the facial expressions produced during pain and orgasm-two different and intense affective experiences-are virtually indistinguishable. However, this finding is counterintuitive, because facial expressions are widely considered to be a powerful tool for social interaction. Consequently, debate continues as to whether the facial expressions of these extreme positive and negative affective states serve a communicative function. Here, we address this debate from a novel angle by modeling the mental representations of dynamic facial expressions of pain and orgasm in 40 observers in each of two cultures (Western, East Asian) using a data-driven method. Using a complementary approach of machine learning, an information-theoretic analysis, and a human perceptual discrimination task, we show that mental representations of pain and orgasm are physically and perceptually distinct in each culture. Cross-cultural comparisons also revealed that pain is represented by similar face movements across cultures, whereas orgasm showed distinct cultural accents. Together, our data show that mental representations of the facial expressions of pain and orgasm are distinct, which questions their nondiagnosticity and instead suggests they could be used for communicative purposes. Our results also highlight the potential role of cultural and perceptual factors in shaping the mental representation of these facial expressions. We discuss new research directions to further explore their relationship to the production of facial expressions., Competing Interests: The authors declare no conflict of interest., (Copyright © 2018 the Author(s). Published by PNAS.)

As a highly social species, humans generate complex facial expressions to communicate a diverse range of emotions. Since Darwin's work, identifying among these complex patterns which are common across cultures and which are culture-specific has remained a central question in psychology, anthropology, philosophy, and more recently machine vision and social robotics. Classic approaches to addressing this question typically tested the cross-cultural recognition of theoretically motivated facial expressions representing 6 emotions, and reported universality. Yet, variable recognition accuracy across cultures suggests a narrower cross-cultural communication supported by sets of simpler expressive patterns embedded in more complex facial expressions. We explore this hypothesis by modeling the facial expressions of over 60 emotions across 2 cultures, and segregating out the latent expressive patterns. Using a multidisciplinary approach, we first map the conceptual organization of a broad spectrum of emotion words by building semantic networks in 2 cultures. For each emotion word in each culture, we then model and validate its corresponding dynamic facial expression, producing over 60 culturally valid facial expression models. We then apply to the pooled models a multivariate data reduction technique, revealing 4 latent and culturally common facial expression patterns that each communicates specific combinations of valence, arousal, and dominance. We then reveal the face movements that accentuate each latent expressive pattern to create complex facial expressions. Our data questions the widely held view that 6 facial expression patterns are universal, instead suggesting 4 latent expressive patterns with direct implications for emotion communication, social psychology, cognitive neuroscience, and social robotics. (PsycINFO Database Record, ((c) 2016 APA, all rights reserved).)

Visual categorization is the brain computation that reduces high-dimensional information in the visual environment into a smaller set of meaningful categories. An important problem in visual neuroscience is to identify the visual information that the brain must represent and then use to categorize visual inputs. Here we introduce a new mathematical formalism-termed space-by-time manifold decomposition-that describes this information as a low-dimensional manifold separable in space and time. We use this decomposition to characterize the representations used by observers to categorize the six classic facial expressions of emotion (happy, surprise, fear, disgust, anger, and sad). By means of a Generative Face Grammar, we presented random dynamic facial movements on each experimental trial and used subjective human perception to identify the facial movements that correlate with each emotion category. When the random movements projected onto the categorization manifold region corresponding to one of the emotion categories, observers categorized the stimulus accordingly; otherwise they selected "other." Using this information, we determined both the Action Unit and temporal components whose linear combinations lead to reliable categorization of each emotion. In a validation experiment, we confirmed the psychological validity of the resulting space-by-time manifold representation. Finally, we demonstrated the importance of temporal sequencing for accurate emotion categorization and identified the temporal dynamics of Action Unit components that cause typical confusions between specific emotions (e.g., fear and surprise) as well as those resolving these confusions.

The human face transmits a wealth of signals that readily provide crucial information for social interactions, such as facial identity and emotional expression. Yet, a fundamental question remains unresolved: does the face information for identity and emotional expression categorization tap into common or distinct representational systems? To address this question we tested PS, a pure case of acquired prosopagnosia with bilateral occipitotemporal lesions anatomically sparing the regions that are assumed to contribute to facial expression (de)coding (i.e., the amygdala, the insula and the posterior superior temporal sulcus--pSTS). We previously demonstrated that PS does not use information from the eye region to identify faces, but relies on the suboptimal mouth region. PS's abnormal information use for identity, coupled with her neural dissociation, provides a unique opportunity to probe the existence of a dichotomy in the face representational system. To reconstruct the mental models of the six basic facial expressions of emotion in PS and age-matched healthy observers, we used a novel reverse correlation technique tracking information use on dynamic faces. PS was comparable to controls, using all facial features to (de)code facial expressions with the exception of fear. PS's normal (de)coding of dynamic facial expressions suggests that the face system relies either on distinct representational systems for identity and expression, or dissociable cortical pathways to access them. Interestingly, PS showed a selective impairment for categorizing many static facial expressions, which could be accounted for by her lesion in the right inferior occipital gyrus. PS's advantage for dynamic facial expressions might instead relate to a functionally distinct and sufficient cortical pathway directly connecting the early visual cortex to the spared pSTS. Altogether, our data provide critical insights on the healthy and impaired face systems, question evidence of deficits obtained from patients by using static images of facial expressions, and offer novel routes for patient rehabilitation., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Animals use social camouflage as a tool of deceit to increase the likelihood of survival and reproduction. We tested whether humans can also strategically deploy transient facial movements to camouflage the default social traits conveyed by the phenotypic morphology of their faces. We used the responses of 12 observers to create models of the dynamic facial signals of dominance, trustworthiness, and attractiveness. We applied these dynamic models to facial morphologies differing on perceived dominance, trustworthiness, and attractiveness to create a set of dynamic faces; new observers rated each dynamic face according to the three social traits. We found that specific facial movements camouflage the social appearance of a face by modulating the features of phenotypic morphology. A comparison of these facial expressions with those similarly derived for facial emotions showed that social-trait expressions, rather than being simple one-to-one overgeneralizations of emotional expressions, are a distinct set of signals composed of movements from different emotions. Our generative face models represent novel psychophysical laws for social sciences; these laws predict the perception of social traits on the basis of dynamic face identities.

Designed by biological and social evolutionary pressures, facial expressions of emotion comprise specific facial movements to support a near-optimal system of signaling and decoding. Although highly dynamical, little is known about the form and function of facial expression temporal dynamics. Do facial expressions transmit diagnostic signals simultaneously to optimize categorization of the six classic emotions, or sequentially to support a more complex communication system of successive categorizations over time? Our data support the latter. Using a combination of perceptual expectation modeling, information theory, and Bayesian classifiers, we show that dynamic facial expressions of emotion transmit an evolving hierarchy of "biologically basic to socially specific" information over time. Early in the signaling dynamics, facial expressions systematically transmit few, biologically rooted face signals supporting the categorization of fewer elementary categories (e.g., approach/avoidance). Later transmissions comprise more complex signals that support categorization of a larger number of socially specific categories (i.e., the six classic emotions). Here, we show that dynamic facial expressions of emotion provide a sophisticated signaling system, questioning the widely accepted notion that emotion communication is comprised of six basic (i.e., psychologically irreducible) categories, and instead suggesting four., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Since Darwin's seminal works, the universality of facial expressions of emotion has remained one of the longest standing debates in the biological and social sciences. Briefly stated, the universality hypothesis claims that all humans communicate six basic internal emotional states (happy, surprise, fear, disgust, anger, and sad) using the same facial movements by virtue of their biological and evolutionary origins [Susskind JM, et al. (2008) Nat Neurosci 11:843-850]. Here, we refute this assumed universality. Using a unique computer graphics platform that combines generative grammars [Chomsky N (1965) MIT Press, Cambridge, MA] with visual perception, we accessed the mind's eye of 30 Western and Eastern culture individuals and reconstructed their mental representations of the six basic facial expressions of emotion. Cross-cultural comparisons of the mental representations challenge universality on two separate counts. First, whereas Westerners represent each of the six basic emotions with a distinct set of facial movements common to the group, Easterners do not. Second, Easterners represent emotional intensity with distinctive dynamic eye activity. By refuting the long-standing universality hypothesis, our data highlight the powerful influence of culture on shaping basic behaviors once considered biologically hardwired. Consequently, our data open a unique nature-nurture debate across broad fields from evolutionary psychology and social neuroscience to social networking via digital avatars.

Facial expressions have long been considered the "universal language of emotion." Yet consistent cultural differences in the recognition of facial expressions contradict such notions (e.g., R. E. Jack, C. Blais, C. Scheepers, P. G. Schyns, & R. Caldara, 2009). Rather, culture--as an intricate system of social concepts and beliefs--could generate different expectations (i.e., internal representations) of facial expression signals. To investigate, they used a powerful psychophysical technique (reverse correlation) to estimate the observer-specific internal representations of the 6 basic facial expressions of emotion (i.e., happy, surprise, fear, disgust, anger, and sad) in two culturally distinct groups (i.e., Western Caucasian [WC] and East Asian [EA]). Using complementary statistical image analyses, cultural specificity was directly revealed in these representations. Specifically, whereas WC internal representations predominantly featured the eyebrows and mouth, EA internal representations showed a preference for expressive information in the eye region. Closer inspection of the EA observer preference revealed a surprising feature: changes of gaze direction, shown primarily among the EA group. For the first time, it is revealed directly that culture can finely shape the internal representations of common facial expressions of emotion, challenging notions of a biologically hardwired "universal language of emotion."

Bubbles is a classification image technique that randomly samples visual information from input stimuli to derive the diagnostic features that observers use in visual categorization tasks. To reach statistical significance, Bubbles performs an exhaustive and repetitive search in the stimulus space. To reduce the search trials, we developed an adaptive method that uses reinforcement learning techniques to optimize sampling by exploiting the observer's history of categorization. We compared the performance of the original and the adaptive Bubbles algorithms in a model observer and eight human adults who all resolved the same visual categorization task (i.e., five facial expressions of emotion). We demonstrate the feasibility of a substantial reduction (by a factor of ∼2) in the number of search trials required to locate the same diagnostic features with the adaptive method, but only when the observer reaches a performance threshold of 50% correct for each expression category. When this threshold is not reached, both the original and adaptive algorithms converge in the same number of trials., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

It is well established that animal communication signals have adapted to the evolutionary pressures of their environment. For example, the low-frequency vocalizations of the elephant are tailored to long-range communications, whereas the high-frequency trills of birds are adapted to their more localized acoustic niche. Like the voice, the human face transmits social signals about the internal emotional state of the transmitter. Here, we address two main issues: First, we characterized the spectral composition of the facial features signaling each of the six universal expressions of emotion (happiness, sadness, fear, disgust, anger, and surprise). From these analyses, we then predicted and tested the effectiveness of the transmission of emotion signals over different viewing distances. We reveal a gradient of recognition over viewing distances constraining the relative adaptive usefulness of facial expressions of emotion (distal expressions are good signals over a wide range of viewing distances; proximal expressions are suited to closer-range communication).

Competent social organisms will read the social signals of their peers. In primates, the face has evolved to transmit the organism's internal emotional state. Adaptive action suggests that the brain of the receiver has co-evolved to efficiently decode expression signals. Here, we review and integrate the evidence for this hypothesis. With a computational approach, we co-examined facial expressions as signals for data transmission and the brain as receiver and decoder of these signals. First, we show in a model observer that facial expressions form a lowly correlated signal set. Second, using time-resolved EEG data, we show how the brain uses spatial frequency information impinging on the retina to decorrelate expression categories. Between 140 to 200 ms following stimulus onset, independently in the left and right hemispheres, an information processing mechanism starts locally with encoding the eye, irrespective of expression, followed by a zooming out to processing the entire face, followed by a zooming back in to diagnostic features (e.g. the opened eyes in "fear", the mouth in "happy"). A model categorizer demonstrates that at 200 ms, the left and right brain have represented enough information to predict behavioral categorization performance.

This article examines the human face as a transmitter of expression signals and the brain as a decoder of these expression signals. If the face has evolved to optimize transmission of such signals, the basic facial expressions should have minimal overlap in their information. If the brain has evolved to optimize categorization of expressions, it should be efficient with the information available from the transmitter for the task. In this article, we characterize the information underlying the recognition of the six basic facial expression signals and evaluate how efficiently each expression is decoded by the underlying brain structures.