Perceptual and conceptual organization of facial emotions: Hemispheric differences

Patients with lesions to either the right or left hemisphere and control subjects were asked to judge the similarity of pairs of photographs of a person displaying different emotions, and of pairs of emotion words. The results were submitted to a multidimensional scaling analysis. Right-hemisphere-damaged subjects were found to be more impaired at perceiving facial emotions than were left-hemisphere-damaged subjects or controls, and this impairment was not confined to the perception of a subset of facial emotions nor to judging emotional valence (Pleasantness versus Unpleasantness). Rather, subtle impairments in perceiving a wide range of facial emotions were found, mostly concerning differentiation of the Positive-Negative and Attention-Rejection dimensions, and concerning the strategies the subjects used to make their judgments. The right-hemisphere-damaged subjects performed comparably to controls in their ratings of emotion words, suggesting that their ability to conceptualize emotional states was intact and that their impairment was strictly in the perception of emotion.     

The influence of emotional faces on the spatial allocation of attention

Background objectives: Studies suggest that the right hemisphere is dominant for emotional facial recognition. In addition, whereas some studies suggest the right hemisphere mediates the processing of all emotions (dominance hypothesis), other studies suggest that the left hemisphere mediates positive emotions the right mediates negative emotions (valence hypothesis). Since each hemisphere primarily attends to contralateral space, the goals of this study was to learn if emotional faces would induce a leftward deviation of attention and if the valence of facial emotional stimuli can influence the normal viewer’s spatial direction of attention. Methods: Seventeen normal right handed participants were asked to bisect horizontal lines that had all combinations of sad, happy or neutral faces at ends of these lines. During this task the subjects were never requested to look at these faces and there were no task demands that depended on viewing these faces. Results: Presentation of emotional faces induced a greater leftward deviation compared to neutral faces, independent of where (spatial position) these faces were presented. However, faces portraying negative emotions tended to induce a greater leftward bias than positive emotions. Conclusions: Independent of location, the presence of emotional faces influenced the spatial allocation of attention, such that normal subjects shift the direction of their attention toward left hemispace and this attentional shift appears to be greater for negative (sad) than positive faces (happy).     

Facial expressions are more easily produced on the upper-lower compared to the right-left hemiface

Clinical research has concentrated on differences in intensity of expression between the right and left hemiface as a means to assess hemispheric differences in motor control. However, observations by social psychologists suggest that control of facial expression may be organized predominantly across the upper-lower hemiface because during social interactions individuals may produce brief facial blends of emotions, in which the upper and lower face display a different emotion. Full facial versus upper/lower and right/left facial blends of emotion were posed by 20 subjects, 10 men and 10 women ranging in age from 20 to 37 years. The subjects rated the difficulty of each pose on a 5-point Likert scale. Digital photographs of the poses were taken and the full and half-facial poses were shown in random order to four judges who indicated what pose was being performed. The results were very robust and confirmed that facial blends of emotion are more easily and accurately posed on the upper-lower than on the right-left hemiface. Our results are consistent with recent anatomical studies showing separate cortical areas for motor control of the upper versus lower face in primates. Based on recent research exploring hemispheric differences in perceiving facial blends of emotion, the left hemisphere may be more involved with modulating lower facial expressions and the right hemisphere more involved with modulating upper facial expressions.     

Recognition of emotional facial expressions in depressed children and adolescents

Of the neurobiological models of children's and adolescents' depression, the neuropsychological one is considered here. Experimental and clinical evidence has allowed us to identify a lateralization of emotional functions from the very beginning of development, and a right hemisphere dominance for emotions is by now well-known. Many studies have also correlated depression with a right hemisphere dysfunction in patients of different ages. The aim of our study was to analyze recognition of different facial emotions by a group of depressed children and adolescents. Patients affected by Major Depressive Disorder recognized less fear in six fundamental emotions than a group of healthy controls, and Dysthymic subjects recognized less anger. The group of patients' failure to recognize negative-aroused facial expressions could indicate a subtle right hemisphere dysfunction in depressed children and adolescents.     

Decoding facial blends of emotion: Visual field,attentional and hemispheric biases

Most clinical research assumes that modulation of facial expressions is lateralized predominantly across the right-left hemiface. However, social psychological research suggests that facial expressions are organized predominantly across the upper-lower face. Because humans learn to cognitively control facial expression for social purposes, the lower face may display a false emotion, typically a smile, to enable approach behavior. In contrast, the upper face may leak a person’s true feeling state by producing a brief facial blend of emotion, i.e. a different emotion on the upper versus lower face. Previous studies from our laboratory have shown that upper facial emotions are processed preferentially by the right hemisphere under conditions of directed attention if facial blends of emotion are presented tachistoscopically to the mid left and right visual fields. This paper explores how facial blends are processed within the four visual quadrants. The results, combined with our previous research, demonstrate that lower more so than upper facial emotions are perceived best when presented to the viewer’s left and right visual fields just above the horizontal axis. Upper facial emotions are perceived best when presented to the viewer’s left visual field just above the horizontal axis under conditions of directed attention. Thus, by gazing at a person’s left ear, which also avoids the social stigma of eye-to-eye contact, one’s ability to decode facial expressions should be enhanced.     

Decoding facial blends of emotion: Visual field,attentional and hemispheric biases

Most clinical research assumes that modulation of facial expressions is lateralized predominantly across the right-left hemiface. However, social psychological research suggests that facial expressions are organized predominantly across the upper-lower face. Because humans learn to cognitively control facial expression for social purposes, the lower face may display a false emotion, typically a smile, to enable approach behavior. In contrast, the upper face may leak a person’s true feeling state by producing a brief facial blend of emotion, i.e. a different emotion on the upper versus lower face. Previous studies from our laboratory have shown that upper facial emotions are processed preferentially by the right hemisphere under conditions of directed attention if facial blends of emotion are presented tachistoscopically to the mid left and right visual fields. This paper explores how facial blends are processed within the four visual quadrants. The results, combined with our previous research, demonstrate that lower more so than upper facial emotions are perceived best when presented to the viewer’s left and right visual fields just above the horizontal axis. Upper facial emotions are perceived best when presented to the viewer’s left visual field just above the horizontal axis under conditions of directed attention. Thus, by gazing at a person’s left ear, which also avoids the social stigma of eye-to-eye contact, one’s ability to decode facial expressions should be enhanced.     

A valence-specific lateral bias for discriminating emotional facial expressions in free field

Findings from subjects with unilateral brain damage, as well as from normal subjects studied with tachistoscopic paradigms, argue that emotion is processed differently by each brain hemisphere. An open question concerns the extent to which such lateralised processing might occur under natural, freeviewing conditions. To explore this issue, we asked 28 normal subjects to discriminate emotions expressed by pairs of faces shown side-by-side, with no time or viewing constraints. Images of neutral expressions were shown paired with morphed images of very faint emotional expressions (happiness, surprise, disgust, fear, anger, or sadness). We found a surprising and robust laterality effect: When discriminating negative emotional expressions, subjects performed significantly better when the emotional face was to the left of the neutral face; conversely, when discriminating positive expressions, subjects performed better when the emotional face was to the right. We interpret this valence-specific laterality effect as consistent with the idea that the right hemisphere is specialised to process negative emotions, whereas the left is specialised to process positive emotions. The findings have important implications for how humans perceive facial emotion under natural conditions.     

Laterality effects in processing tonal and atonal melodies with affective and nonaffective task instructions

Right-handed university subjects were presented with monaural melodies that either conformed to the rules of the Western tonal system (tonal melodies) or that systematically deviated from it (atonal melodies) while containing similar contours and pitch skips. Subjects were tested under two different task instructions. One group was requested to judge whether each melody sounded correct or not (the nonaffective task); the other group had to judge whether each melody sounded pleasant or not (the affective task). The nonaffective task was found to elicit essentially no ear difference. In contrast, the affective instruction induced opposite and reliable laterality effects, depending on the valence of the response. The pleasant responses were indicative of a left hemisphere predominance and the unpleasant responses of a right hemisphere predominance. The results are consistent with the claim that the left hemisphere is biased toward positive emotions and the right to negative emotions. Moreover, the results suggest that affective appreciation of melodies is dissociable from their nonaffective judgment.     

Recognition of facial expressions of emotions in criminal and non-criminal psychopaths: Valence-specific hypothesis

IntroductionPsychopaths with the dominant reduced interpersonal and affective ability are characterized by the hypofunction of the right hemisphere, while psychopaths with the dominant impulsivity and antisocial behavior are characterized by the hyperfunction of the left hemisphere. The assumption is that this interhemispheric imbalance in a psychopath will also be reflected in the recognition of facial emotional expressions.ObjectiveThe objective is to examine the lateralization of facial expressions of positive and negative emotions as well as processing of facial expressions of emotions in criminal and non-criminal psychopaths.Participants48 male participants age 24–40 were voluntarily recruited from the psychiatric hospital in Nis, Serbia.Stimuli48 black-and-white photographs in two separate tasks were used for the stimulation with central and lateral exposition.ResultsCriminality is related to the reduced recognition of facial expression of surprise and not necessarily to psychopathy, whereas reduced recognition of facial expression of fear is related to psychopathy, but not criminality. Valence-specific hypothesis has not been confirmed for positive and negative emotions in criminal and non-criminal psychopaths and non-psychopaths, but it was shown that positive emotions are equally well processed in both hemispheres, whereas negative emotions are more successfully processed in the left hemisphere.     

The valence-specific laterality effect in free viewing conditions: The influence of sex, handedness, and response bias

The right hemisphere has often been viewed as having a dominant role in the processing of emotional information. Other evidence indicates that both hemispheres process emotional information but their involvement is valence specific, with the right hemisphere dealing with negative emotions and the left hemisphere preferentially processing positive emotions. This has been found under both restricted (Reuter-Lorenz & Davidson, 1981) and free viewing conditions (Jansari, Tranel, & Adophs, 2000). It remains unclear whether the valence-specific laterality effect is also sex specific or is influenced by the handedness of participants. To explore this issue we repeated Jansari et al.'s free-viewing laterality task with 78 participants. We found a valence-specific laterality effect in women but not men, with women discriminating negative emotional expressions more accurately when the face was presented on the left-hand side and discriminating positive emotions more accurately when those faces were presented on the right-hand side. These results indicate that under free viewing conditions women are more lateralised for the processing of facial emotion than are men. Handedness did not affect the lateralised processing of facial emotion. Finally, participants demonstrated a response bias on control trials, where facial emotion did not differ between the faces. Participants selected the left-hand side more frequently when they believed the expression was negative and the right-hand side more frequently when they believed the expression was positive. This response bias can cause a spurious valence-specific laterality effect which might have contributed to the conflicting findings within the literature.     

The relationship of hemispheric preference,as measured by conjugate lateral eye movements,to accuracy of emotional facial expression

This study investigated the relationship between hemispheric preference and communication accuracy of facial affect. Hemispheric preference was determined by rating conjugate lateral eye movements. Forty right-handed females were classified: 10 right-movers (left hemisphere preference), 15 left-movers (right hemisphere preference), and 15 bidirectionals (no hemispheric preference). Subjects were shown emotionally loaded slides portraying happiness, fear, and disgust and their videotaped facial expressions were independently rated for accuracy. It was predicted that negative emotions involved right hemisphere activity and negative affect involved left hemisphere activity. As expected, it was found that left-movers were significantly better than right-movers at nonverbally communicating disgust and fear; hemispheric preference was unrelated to expression of happiness. The accuracy scores of the bidirectionals generally fell in between those of the left-movers and those of the right-movers. The results were discussed in reference to the differential lateralization of positive and negative emotions.This study was conducted as a senior research project by the first author under the supervision of the second author.     

Identifying facial emotions: valence specific effects and an exploration of the effects of viewer gender

The valence hypothesis suggests that the right hemisphere is specialised for negative emotions and the left hemisphere is specialised for positive emotions (Silberman & Weingartner, 1986). It is unclear to what extent valence-specific effects in facial emotion perception depend upon the gender of the perceiver. To explore this question 46 participants completed a free view lateralised emotion perception task which involved judging which of two faces expressed a particular emotion. Eye fixations of 24 of the participants were recorded using an eye tracker. A significant valence-specific laterality effect was obtained, with positive emotions more accurately identified when presented to the right of centre, and negative emotions more accurately identified when presented to the left of centre. The valence-specific laterality effect did not depend on the gender of the perceiver. Analysis of the eye tracking data showed that males made more fixations while recognising the emotions and that the left-eye was fixated substantially more than the right-eye during emotion perception. Finally, in a control condition where both faces were identical, but expressed a faint emotion, the participants were significantly more likely to select the right side when the emotion label was positive. This finding adds to evidence suggesting that valence effects in facial emotion perception are not only caused by the perception of the emotion but by other processes.     

A Voxel-based lesion study on facial emotion recognition after circumscribed prefrontal cortex damage

Previous studies have shown inconsistent findings regarding the contribution of the different prefrontal regions in emotion recognition. Moreover, the hemispheric lateralization hypothesis posits that the right hemisphere is dominant for processing all emotions regardless of affective valence, whereas the valence specificity hypothesis posits that the left hemisphere is specialized for processing positive emotions while the right hemisphere is specialized for negative emotions. However, recent findings suggest that the evidence for such lateralization has been less consistent. In this study, we investigated emotion recognition of fear, surprise, happiness, sadness, disgust, and anger in 30 patients with focal prefrontal cortex lesions and 30 control subjects. We also examined the impact of lesion laterality on recognition of the six basic emotions. The results showed that compared to control subjects, the frontal subgroups were impaired in recognition of three negative basic emotions of fear, sadness, and anger – regardless of the lesion laterality. Therefore, our findings did not establish that each hemisphere is specialized for processing specific emotions. Moreover, the voxel-based lesion symptom mapping analysis showed that recognition of fear, sadness, and anger draws on a partially common bilaterally distributed prefrontal network.     

LATERALIZATION FOR FACIAL EMOTIONAL BEHAVIOR: A METHODOLOGICAL PERSPECTIVE*

Recent attention has focused upon the neuropsychological mechanisms underlying facial emotional expression. The greater intensity of expression and more extensive muscular involvement of the left hemiface have suggested right hemisphere dominance in the expression of emotion. This interpretation is based on the fact that the lower portion of the face is predominantly innervated by the contralateral cerebral hemisphere. The current paper reviews a series of studies on (a) facial asymmetry in normal subjects, (b) facial emotional expression in brain-damaged subjects with unilateral lesions, and (c) peripheral factors mediating facial expression findings. Neuropsychological mechanisms underlying emotional processing are also discussed. Our finding of left-sided facial asymmetries in normals is underscored by our finding that right brain-damaged patients are impaired in the expression of facial emotion. Finally, our studies have demonstrated that the asymmetries in normals and the deficits in patient populations are not artifacts of hemiface size and mobility or facial paralysis and apraxia. Taken together, these studies provide evidence that the right hemisphere is specialized for the expression of facial emotion.     

Laterality of facial expressions of emotion: Universal and culture-specific influences

Recent research indicates that (a) the perception and expression of facial emotion are lateralized to a great extent in the right hemisphere, and, (b) whereas facial expressions of emotion embody universal signals, culture-specific learning moderates the expression and interpretation of these emotions. In the present article, we review the literature on laterality and universality, and propose that, although some components of facial expressions of emotion are governed biologically, others are culturally influenced. We suggest that the left side of the face is more expressive of emotions, is more uninhibited, and displays culture-specific emotional norms. The right side of face, on the other hand, is less susceptible to cultural display norms and exhibits more universal emotional signals.     

Consequences of brain tumour resection on emotion recognition

Emotion processing impairments are common in patients undergoing brain surgery for fronto-temporal tumour resection, with potential consequences on social interactions. However, evidence is controversial concerning side and site of lesions causing such deficits. This study investigates visual and auditory emotion recognition in brain tumour patients with the aim of clarifying which lesion sites are related to impairments in emotion processing from different modalities. Thirty-four patients were evaluated, before and after surgery, on facial expression and emotional prosody recognition; voxel-based lesion–symptom mapping (VLSM) analyses were performed on patients’ post-surgery MRI images. Results showed that patients’ performance decreased after surgery in both visual and auditory modalities, but, in general, recovered 3 months after surgery. In facial expression recognition, left brain-damaged patients showed greater post-surgery deterioration than right brain-damaged ones, whose performance specifically decreased for sadness and fear. VLSM analysis revealed two segregated areas in the left hemisphere accounting for post-surgery scores for happy (fronto-temporo-insular region) and surprised (middle frontal gyrus and inferior fronto-occipital fasciculus) facial expressions. Our findings demonstrate that surgical removal of tumours in the fronto-temporal region produces impairment in facial emotion recognition with an overall recovery at 3 months, suggesting a partially different representation of positive and negative emotions in the left and right hemispheres for visually – but not auditory – presented emotions; moreover, we show that deficits in specific expression recognition are associated with discrete lesion locations.     

Visual-field bias in the judgment of facial expression of emotion

The left and right hemispheres of the brain are differentially related to the processing of emotions. Although there is little doubt that the right hemisphere is relatively superior for processing negative emotions, controversy exists over the hemispheric role in the processing of positive emotions. Eighty right-handed normal male participants were examined for visual-field (left-right) differences in the perception of facial expressions of emotion. Facial composite (RR, LL) and hemifacial (R, L) sets depicting emotion expressions of happiness and sadness were prepared. Pairs of such photographs were presented bilaterally for 150 ms, and participants were asked to select the photographs that looked more expressive. A left visual-field superiority (a right-hemisphere function) was found for sad facial emotion. A hemispheric advantage in the perception of happy expression was not found.     

Visual-Field Bias in the Judgment of Facial Expression of Emotion

The left and right hemispheres of the brain are differentially related to the processing of emotions. Although there is little doubt that the right hemisphere is relatively superior for processing negative emotions, controversy exists over the hemispheric role in the processing of positive emotions. Eighty right-handed normal male participants were examined for visual-field (left-right) differences in the perception of facial expressions of emotion. Facial composite (RR, LL) and hemifacial (R, L) sets depicting emotion expressions of happiness and sadness were prepared. Pairs of such photographs were presented bilaterally for 150 ms, and participants were asked to select the photographs that looked more expressive. A left visual-field superiority (a right-hemisphere function) was found for sad facial emotion. A hemispheric advantage in the perception of happy expression was not found.     

Unequal impairment in the recognition of positive and negative emotions after right hemisphere lesions: a left hemisphere bias for happy faces

The processing of several important aspects of a human face was investigated in a single patient (LZ), who had a large infarct of the right hemisphere involving the parietal, and temporal lobes with extensions into the frontal region. LZ showed selective problems with recognizing emotional expressions, whereas she was flawless in recognizing gender, familiarity, and identity. She was very poor in recognizing negative facial expressions (fear, disgust, anger, sadness), but scored as well as the controls on the positive facial expression of happiness. However, in two experiments using both static and dynamic face stimuli, we showed that LZ also did not have a proper notion of what a facial expression of happiness looks like, and could not adequately apply this label. We conclude that the proper recognition of both negative and positive facial expressions relies on the right hemisphere, and that the left hemisphere produces a default state resulting in a bias towards evaluating expressions as happy. We discuss the implications of the current findings for the main models that aim to explain hemispheric specializations for processing of positive and negative emotions.