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Posterior thalamic pulvinar nuclei have been implicated in different aspects of spatial attention, but their exact role in humans remain unclear. Most neuropsychological studies of attention deficits after pulvinar lesion have concerned single patients or small samples. Here we examined a group of 13 patients with focal damage to posterior thalamus on a visual search task with faces, allowing us to test several hypotheses concerning pulvinar function in controlling attention to visually salient or emotionally significant stimuli. Our results identified two subgroups of thalamic patients with distinct patterns of attentional responsiveness to emotional and colour features in face targets. One group with lesions located in anterior and ventral portions of thalamus showed intact performance, with a normal facilitation of visual search for faces with emotional (fearful or happy) expressions on both side of space, similar to healthy controls. By contrast, a second group showed a slower and poorer detection of face targets, most severe for neutral faces, but with a paradoxically enhanced facilitation by both colour and emotional features. This second group had lesions centred on the pulvinar, involving mainly the dorso-medial sectors in patients showing enhanced effects of colour features, but extending to more dorso-lateral sectors in those with enhanced effects of emotional features. These findings reveal that pulvinar nuclei are not critical for orienting attention to emotionally or visually salient features, but instead provide new evidence in support of previous hypotheses suggesting an important role in controlling attention in visual scenes with distracting information., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The superior temporal sulcus (STS) is a major component of the human face perception network, implicated in processing dynamic changeable aspects of faces. However, it remains unknown whether STS holds functionally segregated subdivisions for different categories of facial movements. We used high-resolution functional magnetic resonance imaging (fMRI) at 7T in 16 volunteers to compare STS activation with faces displaying angry or happy expressions, eye-gaze shifts and lip-speech movements. Combining univariate and multivariate analyses, we show a systematic topological organization within STS, with gaze-related activity predominating in the most posterior and superior sector, speech-related activity in the anterior sector and emotional expressions represented in the intermediate middle STS. Right STS appeared to hold a finer functional segregation between all four types of facial movements, and best discriminative abilities within the face-selective posterior STS (pSTS). Conversely, left STS showed greater overlap between conditions, with a lack of distinction between mouth movements associated to speech or happy expression and better discriminative abilities (for gaze and speech vs emotion conditions) outside pSTS. Differential sensitivity to upper (eye) or lower (mouth) facial features may contribute to, but does not appear to fully account for, these response patterns., (© The Author (2017). Published by Oxford University Press.)

Attention and perception are potentiated for emotionally significant stimuli, promoting efficient reactivity and survival. But does such enhancement extend to stimuli simultaneously presented across different sensory modalities? We used functional magnetic resonance imaging in humans to examine the effects of visual emotional signals on concomitant sensory inputs in auditory, somatosensory, and visual modalities. First, we identified sensory areas responsive to task-irrelevant tones, touches, or flickers, presented bilaterally while participants attended to either a neutral or a fearful face. Then, we measured whether these responses were modulated by the emotional content of the face. Sensory responses in primary cortices were enhanced for auditory and tactile stimuli when these appeared with fearful faces, compared with neutral, but striate cortex responses to the visual stimuli were reduced in the left hemisphere, plausibly as a consequence of sensory competition. Finally, conjunction and functional connectivity analyses identified 2 distinct networks presumably responsible for these emotional modulatory processes, involving cingulate, insular, and orbitofrontal cortices for the increased sensory responses, and ventrolateral prefrontal cortex for the decreased sensory responses. These results suggest that emotion tunes the excitability of sensory systems across multiple modalities simultaneously, allowing the individual to adaptively process incoming inputs in a potentially threatening environment., (© The Author 2016. Published by Oxford University Press.)

The ability to decode facial emotions is of primary importance for human social interactions; yet, it is still debated how we analyze faces to determine their expression. Here we compared the processing of emotional face expressions through holistic integration and/or local analysis of visual features, and determined which brain systems mediate these distinct processes. Behavioral, physiological, and brain responses to happy and angry faces were assessed by presenting congruent global configurations of expressions (e.g., happy top+happy bottom), incongruent composite configurations (e.g., angry top+happy bottom), and isolated features (e.g. happy top only). Top and bottom parts were always from the same individual. Twenty-six healthy volunteers were scanned using fMRI while they classified the expression in either the top or the bottom face part but ignored information in the other non-target part. Results indicate that the recognition of happy and anger expressions is neither strictly holistic nor analytic Both routes were involved, but with a different role for analytic and holistic information depending on the emotion type, and different weights of local features between happy and anger expressions. Dissociable neural pathways were engaged depending on emotional face configurations. In particular, regions within the face processing network differed in their sensitivity to holistic expression information, which predominantly activated fusiform, inferior occipital areas and amygdala when internal features were congruent (i.e. template matching), whereas more local analysis of independent features preferentially engaged STS and prefrontal areas (IFG/OFC) in the context of full face configurations, but early visual areas and pulvinar when seen in isolated parts. Collectively, these findings suggest that facial emotion recognition recruits separate, but interactive dorsal and ventral routes within the face processing networks, whose engagement may be shaped by reciprocal interactions and modulated by task demands., (Copyright © 2016 Elsevier Inc. All rights reserved.)

The neuropeptide oxytocin (OT) has many potential social benefits. For example, intranasal administration of OT appears to trigger caregiving behavior and to improve the recognition of emotional facial expressions. But the mechanism for these effects is not yet clear. Recent findings relating OT to action imitation and to the visual processing of the eye region of faces point to mimicry as a mechanism through which OT improves processing of emotional expression. To test the hypothesis that increased levels of OT in the brain enhance facial mimicry, 60 healthy male participants were administered, in a double-blind between-subjects design, 24 international units (IUs) of OT or placebo (PLA) through nasal spray. Facial mimicry and emotion judgments were recorded in response to movie clips depicting changing facial expressions. As expected, facial mimicry was increased in the OT group, but effects were strongest for angry infant faces. These findings provide further evidence for the importance of OT in social cognitive skills, and suggest that facial mimicry mediates the effects of OT on improved emotion recognition., (Copyright © 2016 Elsevier Inc. All rights reserved.)

A fast, subcortical pathway to the amygdala is thought to have evolved to enable rapid detection of threat. This pathway's existence is fundamental for understanding nonconscious emotional responses, but has been challenged as a result of a lack of evidence for short-latency fear-related responses in primate amygdala, including humans. We recorded human intracranial electrophysiological data and found fast amygdala responses, beginning 74-ms post-stimulus onset, to fearful, but not neutral or happy, facial expressions. These responses had considerably shorter latency than fear responses that we observed in visual cortex. Notably, fast amygdala responses were limited to low spatial frequency components of fearful faces, as predicted by magnocellular inputs to amygdala. Furthermore, fast amygdala responses were not evoked by photographs of arousing scenes, which is indicative of selective early reactivity to socially relevant visual information conveyed by fearful faces. These data therefore support the existence of a phylogenetically old subcortical pathway providing fast, but coarse, threat-related signals to human amygdala.

We frequently need to change our current occupation, an operation requiring additional effortful cognitive demands. Switching from one task to another may involve two distinct processes: inhibition of the previously relevant task-set, and initiation of a new one. Here we tested whether these two processes are underpinned by separate neural substrates, and whether they differ depending on the nature of the task and the emotional content of stimuli. We used functional magnetic resonance imaging in healthy human volunteers who categorize emotional faces according to three different judgment rules (color, gender, or emotional expression). Our paradigm allowed us to separate neural activity associated with inhibition and switching based on the sequence of the tasks required on successive trials. We found that the bilateral medial superior parietal lobule and left intraparietal sulcus showed consistent activation during switching regardless of the task. On the other hand, no common region was activated (or suppressed) as a consequence of inhibition across all tasks. Rather, task-specific effects were observed in brain regions that were more activated when switching to a particular task but less activated after inhibition of the same task. In addition, compared to other conditions, the emotional task elicited a similar switching cost but lower inhibition cost, accompanied by selective decrease in the anterior cingulate cortex when returning to this task shortly after inhibiting it. These results demonstrate that switching relies on domain-general processes mediated by postero-medial parietal areas, engaged across all tasks, but also provide novel evidence that task inhibition produces domain-specific decreases as a function of particular task demands, with only the latter inhibition component being modulated by emotional information., (Copyright © 2013 Elsevier Inc. All rights reserved.)

We recorded intracranial local field potentials from structurally intact human visual cortex during several face processing tasks in a patient before brain surgery. Intracranial local field potentials were measured from subdural electrodes implanted in a right fusiform region with face-sensitive activity and a more medial location in posterior parahippocampal gyrus with house-selective activity. This electrode implantation allowed us to compare neural responses with different facial properties within two adjacent but functionally distinct cortical regions. Five experiments were conducted to determine the temporal dynamics of perceptual (Experiments 1 and 5), emotional (Experiments 2 and 3), and social (Experiment 4) effects on face-specific responses in the right fusiform. Our findings showed an early negative deflection (N200) that primarily reflected category-selective perceptual encoding of facial information, whereas higher order effects of face individuation, emotional expression, and gaze direction produced selective modulations in the same face-specific region during a later time period (from 200 to 1000 msec after onset). These results shed new lights on the time course of face recognition mechanisms in human visual cortex and reveal for the first time anatomically overlapping but temporally distinct influences of identity or emotional/social factors on face processing in right fusiform gyrus, which presumably reflect top-down feedback effects from distant brain areas.

Objective: The 22q11.2 deletion syndrome (22q11DS) is a neurogenetic syndrome with high risk for the development of psychiatric disorder. There is interest in identifying reliable markers for measuring and monitoring socio-emotional impairments in 22q11DS during development. The current study investigated eye gaze as a potential marker during a face-processing task in children and young adolescents with 22q11DS., Method: Eye gaze and behavioral correlates were investigated in 26 subjects (aged 8 to 15 years) with 22q11DS during the Jane Task, which targets featural and configural face processing. Individuals with 22q11DS were compared with chronologically age-matched healthy controls and individuals with idiopathic developmental delay (DD)., Results: Few differences in accuracy were observed between patients with 22q11DS and DD controls; however individuals with 22q11DS spent less time on the eyes and more time on the mouths than both comparison groups. IQ predicted time on the eyes in subjects with 22q11DS, and anxiety predicted time on the eyes in DD and 22q11DS subjects., Conclusions: These results provide evidence for abnormal exploration of faces in the syndrome and suggest that time spent on the eyes may contribute to face processing difficulties and interact with anxiety levels to exacerbate socio-emotional dysfunction in affected individuals., (2010 American Academy of Child and Adolescent Psychiatry. Published by Elsevier Inc. All rights reserved.)

The amygdala is known to influence processing of threat-related stimuli in distant brain regions, including visual cortex. The time-course of these distant influences is unknown, although this information is important for resolving debates over likely pathways mediating an apparent rapidity in emotional processing. To address this, we recorded event-related potentials (ERPs) to seen fearful face expressions, in preoperative patients with medial temporal lobe epilepsy who had varying degrees of amygdala pathology, plus healthy volunteers. We found that amygdala damage diminished ERPs for fearful versus neutral faces within the P1 time-range, approximately 100-150 ms, and for a later component at approximately 500-600 ms. Individual severity of amygdala damage determined the magnitude of both these effects, consistent with a causal amygdala role. By contrast, amygdala damage did not affect explicit perception of fearful expressions nor a distinct emotional ERP effect at 150-250 ms. These results demonstrate two distinct time-points at which the amygdala influences fear processing. The data also demonstrate that while not all aspects of expression processing are disrupted by amygdala damage, there is a crucial impact on an early P1 component. These findings are consistent with the existence of multiple processing stages or routes for fearful faces that vary in their dependence on amygdala function., ((c) 2009 Wiley-Liss, Inc.)

Affective and social processes play a major role in everyday life, but appropriate methods to assess disturbances in these processes after brain lesions are still lacking. Past studies have shown that amygdala damage can impair recognition of facial expressions, particularly fear, as well as processing of gaze direction; but the mechanisms responsible for these deficits remain debated. Recent accounts of human amygdala function suggest that it is a critical structure involved in self-relevance appraisal. According to such accounts, responses to a given facial expression may vary depending on concomitant gaze direction and perceived social meaning. Here we investigated facial emotion recognition and its interaction with gaze in patients with unilateral amygdala damage (n = 19), compared to healthy controls (n = 10), using computer-generated dynamic face stimuli expressing variable intensities of fear, anger or joy, with different gaze directions (direct versus averted). If emotion perception is influenced by the self-relevance of expression based on gaze direction, a fearful face with averted gaze should be more relevant than the same expression with direct gaze because it signals danger near the observer; whereas anger with direct gaze should be more relevant than with averted gaze because it directly threatens the observer. Our results confirm a critical role for the amygdala in self-relevance appraisal, showing an interaction between gaze and emotion in healthy controls, a trend for such interaction in left-damaged patients but not in right-damaged patients. Impaired expression recognition was generally more severe for fear, but with a greater deficit for right versus left damage. These findings do not only provide new insights on human amygdala function, but may also help design novel neuropsychological tests sensitive to amygdala dysfunction in various patient populations.

How the processing of emotional expression is influenced by perceived gaze remains a debated issue. Discrepancies between previous results may stem from differences in the nature of stimuli and task characteristics. Here we used a highly controlled set of computer-generated animated faces combining dynamic emotional expressions with varying intensity, and gaze shifts either directed at or averted from the observer. We predicted that perceived self-relevance of fearful faces would be higher with averted gaze-signaling a nearby danger; whereas conversely, direct gaze would be more relevant for angry faces-signaling aggressiveness. This interaction pattern was observed behaviorally for emotion intensity ratings, and neurally for functional magnetic resonance imaging activation in amygdala, as well as fusiform and medial prefrontal cortices, but only for mild- and not high-intensity expressions. These results support an involvement of human amygdala in the appraisal of self-relevance and reveal a crucial role of expression intensity in emotion and gaze interactions.

To determine how emotional information modulates subsequent traces for repeated stimuli, we combined simultaneous electro-encephalography (EEG) and magneto-encephalography (MEG) measures during long-lag incidental repetition of fearful, happy, and neutral faces. Repetition effects were modulated by facial expression in three different time windows, starting as early as 40-50 ms in both EEG and MEG, then arising at the time of the N170/M170, and finally between 280-320 ms in MEG only. The very early repetition effect, observed at 40-50 ms over occipito-temporo-parietal regions, showed a different MEG topography according to the facial expression. This differential response to fearful, happy and neutral faces suggests the existence of very early discriminative visual processing of expressive faces, possibly based on the low-level physical features typical of different emotions. The N170 and M170 face-selective components both showed repetition enhancement selective to neutral faces, with greater amplitude for emotional than neutral faces on the first but not the second presentation. These differential repetition effects may reflect valence acquisition for the neutral faces due to repetition, and suggest a combined influence of emotion- and experience-related factors on the early stage of face encoding. Finally, later repetition effects consisted in enhanced M300 (MEG) between 280 and 320 ms for fearful relative to happy and neutral faces that occurred on the first presentation, but levelled out on the second presentation. This effect may correspond to the higher arousing value of fearful stimuli that might habituate with repetition. Our results reveal that multiple stages of face processing are affected by the repetition of emotional information.

Every day we encounter new people, interact with them, and form person impressions based on quick and automatic inferences from minimal contextual information. Previous studies have identified an extensive network of brain areas involved in familiar face recognition, but there is little evidence to date concerning the neural bases of negative vs. positive person impressions. In the present study, participants were repeatedly exposed to 16 unfamiliar face identities within a pseudo-interactive game context to generate a perception of either "friends" or "foes". Functional magnetic resonance imaging (fMRI) was then performed during an old/new memory task to assess any difference in brain responses to these now familiar face identities, relative to unfamiliar faces. Importantly, whereas facial expressions were always emotional (either smiling or angry) during the encoding phase, they were always neutral during the memory task. Our results reveal that several brain regions involved in familiar face recognition, including fusiform cortex, posterior cingulate gyrus, and amygdala, plus additional areas involved in motivational control such as caudate and anterior cingulate cortex, were differentially modulated as a function of a previous encounter, and generally more activated when faces were perceived as "foes" rather than "friends". These findings underscore that a key dimension of social judgments, based on past impressions of who may be supportive or hostile, may lead to long-lasting effects on memory for faces and thus influence affective reactions to people during a subsequent encounter even in a different (neutral) context.

Adult attachment style refers to individual personality traits that strongly influence emotional bonds and reactions to social partners. Behavioral research has shown that adult attachment style reflects profound differences in sensitivity to social signals of support or conflict, but the neural substrates underlying such differences remain unsettled. Using functional magnetic resonance imaging (fMRI), we examined how the three classic prototypes of attachment style (secure, avoidant, anxious) modulate brain responses to facial expressions conveying either positive or negative feedback about task performance (either supportive or hostile) in a social game context. Activation of striatum and ventral tegmental area was enhanced to positive feedback signaled by a smiling face, but this was reduced in participants with avoidant attachment, indicating relative impassiveness to social reward. Conversely, a left amygdala response was evoked by angry faces associated with negative feedback, and correlated positively with anxious attachment, suggesting an increased sensitivity to social punishment. Secure attachment showed mirror effects in striatum and amygdala, but no other specific correlate. These results reveal a critical role for brain systems implicated in reward and threat processing in the biological underpinnings of adult attachment style, and provide new support to psychological models that have postulated two separate affective dimensions to explain these individual differences, centered on the ventral striatum and amygdala circuits, respectively. These findings also demonstrate that brain responses to face expressions are not driven by facial features alone but determined by the personal significance of expressions in current social context. By linking fundamental psychosocial dimensions of adult attachment with brain function, our results do not only corroborate their biological bases but also help understand their impact on behavior.

The perception of emotional facial expressions induces covert imitation in emotion-specific muscles of the perceiver's face. Neural processes involved in these spontaneous facial reactions remain largely unknown. Here we concurrently recorded EEG and facial EMG in 15 participants watching short movie clips displaying either happy or angry facial expressions. EMG activity was recorded for the zygomaticus major (ZM) that elevates the lips during a smile, and the corrugator supercillii (CS) that knits the eyebrows during a frown. We found increased EMG activity of CS in response to angry expressions, and enhanced EMG activity of ZM for happy expressions, replicating earlier EMG studies. More importantly, we found that the amplitude of an early visual evoked potential (right P1) was larger when ZM activity to happy faces was high, and when CS activity to angry faces was high, as compared to when muscle reactions were low. Conversely, the amplitude of right N170 component was smaller when the intensity of facial imitation was high. These combined EEG-EMG results suggest that early visual processing of face expression may determine the magnitude of subsequent facial imitation, with dissociable effects for P1 and N170. These findings are discussed against the classical dual-route model of face recognition.

Background: 22q11.2 deletion syndrome (22q11DS) is a neurogenetic syndrome associated with a high rate of psychiatric disorders. Previous research has revealed distinctive cognitive deficits, including impaired face processing. However, the neuro-functional substrates underlying these deficits have not been explored. Our aim was to investigate facial and emotional processing in 22q11DS., Methods: During event-related functional magnetic resonance imaging, 15 individuals with 22q11DS were compared with age- and gender-matched healthy control subjects on a simple visual categorization task (faces or houses). Each stimulus was presented twice, and faces had either neutral or emotional (fearful) expressions., Results: Abnormal responses to faces were observed in 22q11DS, including a lack of normal face-selectivity in fusiform gyrus. By contrast, responses to houses were comparable across groups, with preserved selectivity in parahippocampal gyrus. Results also revealed a repetition-suppression effect for fearful faces in the right amygdala, which arose in healthy control subjects only, suggesting a lack of amygdala modulation by fear expression in 22q11DS., Conclusions: Our results demonstrate selective anomalies in several brain regions critically implicated in visual and social function in 22q11DS. These findings suggest important new avenues for studying emotional processing and social deficits frequently observed in psychotic patients and establishing their relation to specific phenotypic manifestations in 22q11DS.

Emotionally expressive faces have been shown to modulate activation in visual cortex, including face-selective regions in ventral temporal lobe. Here, we tested whether emotionally expressive bodies similarly modulate activation in body-selective regions. We show that dynamic displays of bodies with various emotional expressions vs neutral bodies, produce significant activation in two distinct body-selective visual areas, the extrastriate body area and the fusiform body area. Multi-voxel pattern analysis showed that the strength of this emotional modulation was related, on a voxel-by-voxel basis, to the degree of body selectivity, while there was no relation with the degree of selectivity for faces. Across subjects, amygdala responses to emotional bodies positively correlated with the modulation of body-selective areas. Together, these results suggest that emotional cues from body movements produce topographically selective influences on category-specific populations of neurons in visual cortex, and these increases may implicate discrete modulatory projections from the amygdala.

We compared electrical brain responses to fearful vs. neutral facial expressions in healthy volunteers while they performed an orthogonal gender decision task. Face stimuli either had a broadband spatial-frequency content, or were filtered to create either low spatial-frequency (LSF) or high spatial-frequency (HSF) faces, always overlapped with their complementary SF content in upside-down orientation to preserve the total stimulus energy. We tested the hypothesis that the coarse LSF content of faces might be responsible for an early modulation of event-related potentials (ERPs) to fearful expressions. Consistent with previous findings, we show that broadband images of fearful faces, relative to neutral faces, elicit a higher global field power of approximately 130 ms poststimulus onset, corresponding to an increased P1 component over lateral occipital electrodes, with neural sources located within the extrastriate visual cortex. Bandpass filtering of faces strongly affected the latency and amplitude of ERPs, with a suppression of the normal N170 response for both LSF and HSF faces, irrespective of expression. Critically, we found that LSF information from fearful faces, unlike HSF information, produced a right-lateralized enhancement of the lateral occipital P1, without any change in the scalp topography, relative to unfiltered (broadband) fearful faces. These results demonstrate that an early P1 response to fear expression depends on a visual pathway preferentially tuned to coarse-magnocellular inputs, and can persist unchanged even when the N170 generators are disrupted by SF filtering.

Visuo-spatial attention tends to be prioritized towards emotionally negative stimuli such as fearful faces, as opposed to neutral or positive stimuli. Using a covert orienting task, we previously showed that a lateral occipital P1 component, with extrastriate neural sources, was selectively enhanced to lateralized visual targets replacing a fearful face (fear-valid trial) than the same targets replacing a neutral face (fear-invalid trial), providing evidence for exogenous spatial orienting of attention towards threat cues. Here, we describe a new analysis of these data, using topographic evoked potentials mapping methods combined with a distributed source localization technique. We show that an early field topography (40-80 ms post-target onset) with a centro-parietal negativity and a left posterior parietal source distinguished fear-valid from fear-invalid trials, whereas a distinct activity with anterior cingulate sources was selectively evoked during fear-invalid trials. At the same latency, or later, no difference in field topography was found for valid compared to invalid trials with happy faces. The early parietal map preceded a modulation in amplitude of the field strength (approximately 130 ms), corresponding to the enhanced lateral occipital P1 during valid trials in the fear condition. Furthermore, this early topography at 40-80 ms was positively correlated with the subsequent amplitude modulation of P1 at 130-160 ms in the fear condition, suggesting a possible functional coupling between these two successive events. These data have important implications for models of spatial attention and interactions with emotion. They suggest two successive stages of neural activity during exogenous orienting of attention towards visual targets following fearful faces, including an early posterior parietal negativity, followed by gain control mechanisms enhancing visual responses in extrastriate occipital cortex.

Faces are multi-dimensional stimuli bearing important social signals, such as gaze direction and emotion expression. To test whether perception of these two facial attributes recruits distinct cortical areas within the right hemisphere, we used single-pulse transcranial magnetic stimulation (TMS) in healthy volunteers while they performed two different tasks on the same face stimuli. In each task, two successive faces were presented with varying eye-gaze directions and emotional expressions, separated by a short interval of random duration. TMS was applied over either the right somatosensory cortex or the right superior lateral temporal cortex, 100 or 200 ms after presentation of the second face stimulus. Participants performed a speeded matching task on the second face during one of two possible conditions, requiring judgements about either gaze direction or emotion expression (same/different as the first face). Our results reveal a significant task-stimulation site interaction, indicating a selective TMS-related interference following stimulations of somatosensory cortex during the emotional expression task. Conversely, TMS of the superior lateral temporal cortex selectively interfered with the gaze direction task. We also found that the interference effect was specific to the stimulus content in each condition, affecting judgements of gaze shifts (not static eye positions) with TMS over the right superior temporal cortex, and judgements of fearful expressions (not happy expressions) with TMS over the right somatosensory cortex. These results provide for the first time a double dissociation in normal subjects during social face recognition, due to transient disruption of non-overlapping brain regions. The present study supports a critical role of the somatosensory and superior lateral temporal regions in the perception of fear expression and gaze shift in seen faces, respectively.

Emotional visual stimuli evoke enhanced responses in the visual cortex. To test whether this reflects modulatory influences from the amygdala on sensory processing, we used event-related functional magnetic resonance imaging (fMRI) in human patients with medial temporal lobe sclerosis. Twenty-six patients with lesions in the amygdala, the hippocampus or both, plus 13 matched healthy controls, were shown pictures of fearful or neutral faces in task-releant or task-irrelevant positions on the display. All subjects showed increased fusiform cortex activation when the faces were in task-relevant positions. Both healthy individuals and those with hippocampal damage showed increased activation in the fusiform and occipital cortex when they were shown fearful faces, but this was not the case for individuals with damage to the amygdala, even though visual areas were structurally intact. The distant influence of the amygdala was also evidenced by the parametric relationship between amygdala damage and the level of emotional activation in the fusiform cortex. Our data show that combining the fMRI and lesion approaches can help reveal the source of functional modulatory influences between distant but interconnected brain regions.

We investigated the spatio-temporal dynamic of attentional bias towards fearful faces. Twelve participants performed a covert spatial orienting task while recording visual event-related brain potentials (VEPs). Each trial consisted of a pair of faces (one emotional and one neutral) briefly presented in the upper visual field, followed by a unilateral bar presented at the location of one of the faces. Participants had to judge the orientation of the bar. Comparing VEPs to bars shown at the location of an emotional (valid) versus neutral (invalid) face revealed an early effect of spatial validity: the lateral occipital P1 component (approximately 130 ms post-stimulus) was selectively increased when a bar replaced a fearful face compared to when the same bar replaced a neutral face. This effect was not found with upright happy faces or inverted fearful faces. A similar amplification of P1 has previously been observed in electrophysiological studies of spatial attention using non-emotional cues. In a behavioural control experiment, participants were also better at discriminating the orientation of the bar when it replaced a fearful rather than a neutral face. In addition, VEPs time-locked to the face-pair onset revealed a C1 component (approximately 90 ms) that was greater for fearful than happy faces. Source localization (LORETA) confirmed an extrastriate origin of the P1 response showing a spatial validity effect, and a striate origin of the C1 response showing an emotional valence effect. These data suggest that activity in primary visual cortex might be enhanced by fear cues as early as 90 ms post-stimulus, and that such effects might result in a subsequent facilitation of sensory processing for a stimulus appearing at the same location. These results provide evidence for neural mechanisms allowing rapid, exogenous spatial orienting of attention towards fear stimuli.

To clarify the time course of neural responses to faces with different emotional expressions, we used event-related potential (ERP) and reaction time measures. Faces expressing four different emotions (happy, neutral, fearful, disgusted) and houses were shown in both upright and inverted orientations while subjects performed an immediate-repeats task. Results indicated that upright fearful expressions enhanced the frontocentral P200. However, emotional effects on the N170 and late positive component interacted with face orientation and were not selective for any specific expression. A unique negative component for upright disgust faces was observed at approximately 300 ms at occipital regions. These results provide evidence for emotion-specific ERPs associated with fear and disgust, distinct from other non-specific configurational and attentional effects.

To investigate whether the processing of faces and emotional facial expression can be modulated by spatial attention, ERPs were recorded in response to stimulus arrays containing two faces and two non-face stimuli (houses). In separate trials, attention was focused on the face pair or on the house pair, and facial expression was either fearful or neutral. When faces were attended, a greater frontal positivity in response to arrays containing fearful faces was obtained, starting about 100 ms after stimulus onset. In contrast, with faces located outside the attentional focus, this emotional expression effect was completely eliminated. This differential result demonstrates for the first time a strong attentional gating of brain processes involved in the analysis of emotional facial expression. It is argued that while an initial detection of emotionally relevant events mediated by the amygdala may occur pre-attentively, subsequent stages of emotional processing require focal spatial attention. The face-sensitive N170 component was unaffected by emotional facial expression, but N170 amplitudes were enhanced when faces were attended, suggesting that spatial attention can modulate the structural encoding of faces.

Affective and social processes play a major role in everyday life, but appropriate methods to assess disturbances in these processes after brain lesions are still lacking. Past studies have shown that amygdala damage can impair recognition of facial expressions, particularly fear, as well as processing of gaze direction; but the mechanisms responsible for these deficits remain debated. Recent accounts of human amygdala function suggest that it is a critical structure involved in self-relevance appraisal. According to such accounts, responses to a given facial expression may vary depending on concomitant gaze direction and perceived social meaning. Here we investigated facial emotion recognition and its interaction with gaze in patients with unilateral amygdala damage (n = 19), compared to healthy controls (n = 10), using computer-generated dynamic face stimuli expressing variable intensities of fear, anger or joy, with different gaze directions (direct versus averted). If emotion perception is influenced by the self-relevance of expression based on gaze direction, a fearful face with averted gaze should be more relevant than the same expression with direct gaze because it signals danger near the observer; whereas anger with direct gaze should be more relevant than with averted gaze because it directly threatens the observer. Our results confirm a critical role for the amygdala in self-relevance appraisal, showing an interaction between gaze and emotion in healthy controls, a trend for such interaction in left-damaged patients but not in right-damaged patients. Impaired expression recognition was generally more severe for fear, but with a greater deficit for right versus left damage. These findings do not only provide new insights on human amygdala function, but may also help design novel neuropsychological tests sensitive to amygdala dysfunction in various patient populations. [ABSTRACT FROM PUBLISHER]

The present study investigated whether emotionally expressive faces guide attention and modulate fMRI activity in fusiform gyrus in acquired prosopagnosia. Patient PS, a pure case of acquired prosopagnosia with intact right middle fusiform gyrus, performed two behavioral experiments and a functional imaging experiment to address these questions. In a visual search task involving face stimuli, PS was faster to select the target face when it was expressing fear or happiness as compared to when it was emotionally neutral. In a change detection task, PS detected significantly more changes when the changed face was fearful as compared to when it was neutral. Finally, an fMRI experiment showed enhanced activation to emotionally expressive faces and bodies in right fusiform gyrus. In addition, PS showed normal body-selective activation in right fusiform gyrus, partially overlapping the fusiform face area. Together these behavioral and neuroimaging results show that attention was preferentially allocated to emotional faces in patient PS, as observed in healthy subjects. We conclude that systems involved in the emotional guidance of attention by facial expression can function normally in acquired prosopagnosia, and can thus be dissociated from systems involved in face identification. [ABSTRACT FROM PUBLISHER]

A recent brain imaging study (Vuilleumier, Armony, Driver and Dolan 2003, Nature Neuroscience, 6, 624-631) has shown that amygdala responses to fearful expressions are preferentially driven by intact or low spatial frequency (LSF) images of faces, rather than by high spatial frequency (HSF) images. These results suggest that LSF components processed rapidly via magnocellular pathways within the visual system might be very efficiently conveyed to the amygdala for the rapid recognition of fearful expressions, perhaps via a subcortical pathway that activates the pulvinar and superior colliculus, but which bypasses any finer visual analysis of HSF cues in the striate and temporal extrastriate cortex. The purpose of this paper is to analyse the statistical properties of LSF compared with HSF and intact faces. The statistical analysis shows that the LSF components in faces, which are typically extracted rapidly by the visual system, provide a better source of information than HSF components for the correct categorisation of fearful expressions in faces. These results support the idea that visual pathways from the magnocellular visual neurons might be optimal, at a computational level, for the rapid classification of fearful emotional expressions in human faces. [ABSTRACT FROM AUTHOR]

This paper reports five experiments demonstrating that the low spatial frequency components of faces are critical to the production of rapid attentional responses towards fearful facial expressions. In our main experiments, low spatial frequency (LSF) or high spatial frequency (HSF) face pairs, consisting of one fearful and one neutral expression, were presented on a computer screen for a brief period. Participants were required to identify as quickly as possible the orientation of a bar target that immediately replaced one of the faces. Responses were faster when targets replaced the location of LSF fearful faces, compared with LSF neutral faces. By contrast, there were no differences between responses to targets replacing HSF fearful versus HSF neutral faces. This facilitation in spatial orienting occurred specifically with short time intervals between faces and target, and is consistent with a rapid processing of fear cues from LSF inputs that can serve to guide attention towards threat events. [ABSTRACT FROM AUTHOR]

Emotive faces elicit neural responses even when they are not consciously perceived . We used faces hybridized from spatial frequency-filtered individual stimuli to study processing of facial emotion. Employing event-related functional magnetic resonance imaging (fMRI), we show enhanced fusiform cortex responses to hybrid faces containing fearful expressions when such emotional cues are present in the low-spatial frequency (LSF) range. Critically, this effect is independent of whether subjects use LSF or high-spatial frequency (HSF) information to make gender judgments on the hybridized faces. The magnitude of this fusiform enhancement predicts behavioral slowing in response times when participants report HSF information of the hybrid stimulus in the presence of fear in the unreported LSF components. Thus, emotional modulation of a face-responsive region of fusiform is driven by the low-frequency components of the stimulus, an effect independent of subjects'' reported perception but evident in an incidental measure of behavioral performance. [Copyright &y& Elsevier]

Facial expressions can be systematically coded using the Facial Action Coding System (FACS) that describes the specific action unit (AU) or combination of AUs elicited during different kinds of expressions. This study investigated the thermal patterns concomitant to specific action units performance. As thermal imaging can track dynamic patterns in facial temperature at any distance (>0.4 m), with high temporal (<20 m) and thermal (<20 mK@300 K) resolutions, this noninvasive technique was tested as a method to assess fluctuations of facial heat patterns induced by facial muscles contractions. Four FACS-trained coders produced nine different AUs or combination of AUs at various speeds and intensities. Using a spatial pattern approach based on PCA decomposition of the thermal signal, we showed that thermal fluctuations are specific to the activated AUs and are sensitive to the kinetics and intensities of AU production. These results open new avenues for studying patterns of facial muscle activity related to emotion or other cognitively induced activities, in a noninvasive manner, avoiding potential lighting issues. [ABSTRACT FROM PUBLISHER]