Hierarchical status is rapidly assessed from behaviourally dominant faces.

Recognition of social hierarchy is a key feature that helps us navigate through our complex social environment. Neuroimaging studies have identified brain structures involved in the processing of hierarchical stimuli, but the precise temporal dynamics of brain activity associated with such processing remains largely unknown. In this investigation, we used event-related potentials (ERPs) to examine the effect of social hierarchy on the neural responses elicited by dominant and nondominant faces. Participants played a game where they were led to believe that they were middle-rank players, responding alongside other alleged players, whom they perceived as higher or lower-ranking. ERPs were examined in response to dominant and nondominant faces, and low-resolution electromagnetic tomography (LORETA) was used to identify the implicated brain areas. The results revealed that the amplitude of the N170 component was enhanced for faces of dominant individuals, showing that hierarchy influences the early stages of face processing. A later component, the late positive potential (LPP) appearing between 350-700 ms, also was enhanced for faces of higher-ranking players. Source localisation suggested that the early modulation was due to an enhanced response in limbic regions. These findings provide electrophysiological evidence for enhanced early visual processing of socially dominant faces.

Event-Related Potentials Index Prediction Error Signalling During Perceptual Processing of Emotional Facial Expressions.

Humans use socially relevant stimuli to guide perceptual processing of the surrounding environment, with emotional stimuli receiving preferential attention due to their social importance. Predictive coding theory asserts this cognitive process occurs efficiently by combining predictions about what is to be perceived with incoming sensory information, generating prediction errors that are then used to update future predictions. Recent evidence has identified differing neural activity that demonstrates how spatial and feature-based attention may interact with prediction, yet how emotion-guided attention may influence this relationship remains unknown. In the present study, participants viewed a display of two faces in which attention, prediction, and emotion were manipulated, and responded to a face expressing a specific emotion (anger or happiness). The N170 was found to be enhanced by unpredictable as opposed to predictable stimuli, indicating that it indexes general prediction error signalling processes. The N300 amplitudes were also enhanced by unpredictable stimuli, but they were also affected by the attentional status of angry but not happy faces, suggesting that there are differences in prediction error processes indexed by the N170 and N300. Overall, the findings suggest that the N170 and N300 both index violations of expectation for spatial manipulations of stimuli in accordance with prediction error responding processes.

Neural processing of lateralised task-irrelevant fearful faces under different awareness conditions.

The neural fate of task-irrelevant emotional faces under different awareness conditions is poorly understood. Here, we examined the electrophysiological activity during an experiment where the location of target information (contrast-induced line) was manipulated orthogonally to the location of task-irrelevant fearful faces, under subliminal or supraliminal viewing conditions. We found that only target lines elicited an N2-posterior-contralateral (N2pc), indexing spatial attention shifting, in the supraliminal condition. No N2pc was found for the targets in the subliminal condition or for task-irrelevant fearful faces in either conditions. However, the mere presence of a fearful face enhanced early neural activity between 200 and 300 ms only in the subliminal condition. Additionally, the presence of a target line, but not a fearful face, enhanced the P3. Our results suggest that the N2pc is dependent on visual awareness and task-relevancy of the information and that laterally-presented task-irrelevant fearful expressions can be processed without awareness during early visual processing.

Turning the Face Inversion Effect on Its Head: Violated Expectations of Orientation, Lighting, and Gravity Enhance N170 Amplitudes.

Face inversion effects occur for both behavioral and electrophysiological responses when people view faces. In EEG, inverted faces are often reported to evoke an enhanced amplitude and delayed latency of the N170 ERP. This response has been attributed to the indexing of specialized face processing mechanisms within the brain. However, inspection of the literature revealed that, although N170 is consistently delayed to a variety of face representations, only photographed faces invoke enhanced N170 amplitudes upon inversion. This suggests that the increased N170 amplitudes to inverted faces may have other origins than the inversion of the face's structure. We hypothesize that the unique N170 amplitude response to inverted photographed faces stems from multiple expectation violations, over and above structural inversion. For instance, rotating an image of a face upside-down not only violates the expectation that faces appear upright but also lifelong priors about illumination and gravity. We recorded EEG while participants viewed face stimuli (upright vs. inverted), where the faces were illuminated from above versus below, and where the models were photographed upright versus hanging upside-down. The N170 amplitudes were found to be modulated by a complex interaction between orientation, lighting, and gravity factors, with the amplitudes largest when faces consistently violated all three expectations. These results confirm our hypothesis that face inversion effects on N170 amplitudes are driven by a violation of the viewer's expectations across several parameters that characterize faces, rather than a disruption in the configurational disposition of its features.

Waist-to-hip ratio affects female body attractiveness and modulates early brain responses.

This investigation examined the electrophysiological response underlying the visual processing of waist-to-hip ratio (WHR) in female bodies, a characteristic known to affect perceived attractiveness. WHRs of female bodies were artificially adjusted to values of 0.6, 0.7, 0.8 or 0.9. Behavioural ratings of attractiveness of the bodies revealed a preference for WHRs of 0.7 in the overall group of participants, which included both male and female heterosexual individuals. Event-related potentials (ERPs) were then recorded while participants performed a selective attention task involving photographs of female models and scrambled images. Results showed that the P1 (80-120 ms) and N1 (130-170 ms) components situated over posterior brain regions were the earliest components to be modulated by attention and bodies. Interestingly, the vertex-positive potential, occurring between 120-180 ms, produced a greater positivity for WHRs of 0.7 compared to the other ratios. However, this increase was only observed when the body stimuli were attended, while no effect was observed for unattended bodies. These findings provide evidence of an early brain sensitivity to visual attributes that constitute secondary sexual characteristics. Although they are relatively discrete from the point of view of their physical quality, these signs possess strong behavioural significance, producing greater reported attractiveness, likely by conveying the biological meaning that signals good health and greater reproductive success. Our results therefore reveal that attributes associated with sexual attractiveness in female bodies are processed rapidly in the stream of visual processing.

Early sensitivity of evoked potentials to surface and volumetric structure during the visual perception of three-dimensional object shape.

This study used event-related potentials (ERPs) to elucidate how the human visual system processes three-dimensional (3-D) object shape structure. In particular, we examined whether the perceptual mechanisms that support the analysis of 3-D shape are differentially sensitive to higher order surface and volumetric part structure. Observers performed a whole-part novel object matching task in which part stimuli comprised sub-regions of closed edge contour, surfaces or volumetric parts. Behavioural response latency data showed an advantage in matching surfaces and volumetric parts to whole objects over contours, but no difference between surfaces and volumes. ERPs were analysed using a convergence of approaches based on stimulus dependent amplitude modulations of evoked potentials, topographic segmentation, and spatial frequency oscillations. The results showed early differential perceptual processing of contours, surfaces, and volumetric part stimuli. This was first reliably observed over occipitoparietal electrodes during the N1 (140-200 ms) with a mean peak latency of 170 ms, and continued on subsequent P2 (220-260 ms) and N2 (260-320 ms) components. The differential sensitivity in perceptual processing during the N1 was accompanied by distinct microstate patterns that distinguished among contours, surfaces and volumes, and predominant theta band activity around 4-7 Hz over right occipitoparietal and orbitofrontal sites. These results provide the first evidence of early differential perceptual processing of higher order surface and volumetric shape structure within the first 200 ms of stimulus processing. The findings challenge theoretical models of object recognition that do not attribute functional significance to surface and volumetric object structure during visual perception.

Postoperative memory prognosis in temporal lobe epilepsy surgery: The contribution of postictal memory.

OBJECTIVE: The prediction of verbal memory decline after temporal lobe epilepsy (TLE) surgery remains difficult at an individual level. We evaluated the prognostic value of postictal memory testing in predicting the postoperative verbal memory function. METHODS: Sixty-three consecutive patients were included in the analysis who underwent TLE surgery at our center with preoperative interictal/postictal and postoperative memory testing. Verbal memory was evaluated using the Rey Auditory Verbal Learning Test (RAVLT). We used reliable change indices with 90% confidence interval (90% RCIs) to evaluate a significant postoperative memory decline. The sensitivity (Sn), specificity (Sp), positive predictive value (PPV), negative predictive value (NPV), area under the curve (AUC), and accuracy (ACC) were calculated. The analysis was performed for all TLE patients and for the subgroup with hippocampal sclerosis (HS). RESULTS: Left-TLE patients (n = 31) had lower verbal memory scores on RAVLT than right-TLE at 3 months (57% vs 78%) and 12 months (53% vs 78%) after surgery. The 90% RCI was estimated to be a loss of 4 out of 15 items. The predictive value was Sn = 42%, Sp = 84%, PPV = 39%, NPV = 86%, AUC = 0.630, and ACC = 76% to predict a verbal memory decline in the whole group (n = 63). In HS patients (n = 41), the postictal verbal memory test had Sn = 50%, Sp = 88%, PPV = 50%, NPV = 88%, AUC = 0.689, and ACC = 81% to predict a significant postoperative decline. SIGNIFICANCE: Postictal memory is a noninvasive bedside memory test that can help predict the postoperative verbal memory decline in patients with HS with an overall accuracy of 81%.

Early and late cortical responses to directly gazing faces are task dependent.

Gender categorisation of human faces is facilitated when gaze is directed toward the observer (i.e., a direct gaze), compared with situations where gaze is averted or the eyes are closed (Macrae, Hood, Milne, Rowe, & Mason, Psychological Science, 13(5), 460-464, 2002). However, the temporal dynamics underlying this phenomenon remain to some extent unknown. Here, we used electroencephalography (EEG) to assess the neural correlates of this effect, focusing on the event-related potential (ERP) components known to be sensitive to gaze perception (i.e., P1, N170, and P3b). We first replicated the seminal findings of Macrae et al. (2002, Experiment 1) regarding facilitated gender discrimination, and subsequently measured the underlying neural responses. Our data revealed an early preferential processing of direct gaze as compared with averted gaze and closed eyes at the P1, which reverberated at the P3b (Experiment 2). Critically, using the same material, we failed to reproduce these effects when gender categorisation was not required (Experiment 3). Taken together, our data confirm that direct gaze enhances the early P1, as well as later cortical responses to face processing, although the effect appears to be task dependent.

Electrophysiological evidence of perceived sexual attractiveness for human female bodies varying in waist-to-hip ratio.

The dynamics of brain activation reflecting attractiveness in humans are unclear. Among the different features affecting attractiveness of the female body, the waist-to-hip ratio (WHR) is considered to be crucial. To date, however, no event-related potential (ERP) study has addressed the question of its associated pattern of brain activation. We carried out two different experiments: (a) a behavioural study, to judge the level of attractiveness of female realistic models depicting 4 different WHRs (0.6, 0.7, 0.8, 0.9) with and without clothes; (b) an EEG paradigm, to record brain activity while participants (heterosexual men and women) viewed these same models. Behavioural results showed that WHRs of 0.7 were considered more attractive than the others. ERP analyses revealed a different pattern of activation for male and female viewers. The 0.7 ratio elicited greater positivity at the P1 level in male viewers but not females. Naked bodies increased the N190 in both groups and peaked earlier for the 0.7 ratio in the male viewers. Finally, the late positive component (LPC) was found to be greater in male than in female viewers and was globally more marked for naked bodies as well as WHRs of 0.7 in both groups of viewers. These results provide the first electrophysiological evidence of specific time periods linked to the processing of a body feature denoting attractiveness and therefore playing a role in mate choice.

Opposite ERP effects for conscious and unconscious semantic processing under continuous flash suppression.

We examined whether semantic processing occurs without awareness using continuous flash suppression (CFS). In two priming tasks, participants were required to judge whether a target was a word or a non-word, and to report whether the masked prime was visible. Experiment 1 manipulated the lexical congruency between the prime-target pairs and Experiment 2 manipulated their semantic relatedness. Despite the absence of behavioral priming effects (Experiment 1), the ERP results revealed that an N4 component was sensitive to the prime-target lexical congruency (Experiment 1) and semantic relatedness (Experiment 2) when the prime was rendered invisible under CFS. However, these results were reversed with respect to those that emerged when the stimuli were perceived consciously. Our findings suggest that some form of lexical and semantic processing can occur during CFS-induced unawareness, but are associated with different electrophysiological outcomes.

Amygdala activation for eye contact despite complete cortical blindness.

Cortical blindness refers to the loss of vision that occurs after destruction of the primary visual cortex. Although there is no sensory cortex and hence no conscious vision, some cortically blind patients show amygdala activation in response to facial or bodily expressions of emotion. Here we investigated whether direction of gaze could also be processed in the absence of any functional visual cortex. A well-known patient with bilateral destruction of his visual cortex and subsequent cortical blindness was investigated in an fMRI paradigm during which blocks of faces were presented either with their gaze directed toward or away from the viewer. Increased right amygdala activation was found in response to directed compared with averted gaze. Activity in this region was further found to be functionally connected to a larger network associated with face and gaze processing. The present study demonstrates that, in human subjects, the amygdala response to eye contact does not require an intact primary visual cortex.

Looming Angry Faces: Preliminary Evidence of Differential Electrophysiological Dynamics for Filtered Stimuli via Low and High Spatial Frequencies.

Looming motion interacts with threatening emotional cues in the initial stages of visual processing. However, the underlying neural networks are unclear. The current study investigated if the interactive effect of threat elicited by angry and looming faces is favoured by rapid, magnocellular neural pathways and if exogenous or endogenous attention influences such processing. Here, EEG/ERP techniques were used to explore the early ERP responses to moving emotional faces filtered for high spatial frequencies (HSF) and low spatial frequencies (LSF). Experiment 1 applied a passive-viewing paradigm, presenting filtered angry and neutral faces in static, approaching, or receding motions on a depth-cued background. In the second experiment, broadband faces (BSF) were included, and endogenous attention was directed to the expression of faces. Our main results showed that regardless of attentional control, P1 was enhanced by BSF angry faces, but neither HSF nor LSF faces drove the effect of facial expressions. Such findings indicate that looming motion and threatening expressions are integrated rapidly at the P1 level but that this processing relies neither on LSF nor on HSF information in isolation. The N170 was enhanced for BSF angry faces regardless of attention but was enhanced for LSF angry faces during passive viewing. These results suggest the involvement of a neural pathway reliant on LSF information at the N170 level. Taken together with previous reports from the literature, this may indicate the involvement of multiple parallel neural pathways during early visual processing of approaching emotional faces.

Oxytocin differentially modulates the early neural responses to faces and non-social stimuli.

Oxytocin (OT) alters social cognition partly through effects on the processing and appraisal of faces. It is debated whether the hormone also impacts the processing of other, non-social, visual stimuli. To this end, we conducted a randomized, counter-balanced, double-blind, placebo (PL)-controlled within-subjects' electro-encephalography (EEG) study with cismale participants (to control for gender dimorphic hormonal effects; n = 37). Participants received intranasal OT (24IU) and completed a one-back task viewing emotional (fearful/ happy) and neutral faces, and threat (snakes/spiders) and non-threat (mushrooms/flowers) non-social stimuli. OT differentially impacted event-related potentials (ERP)s to faces and non-social stimuli. For faces regardless of emotion, OT evoked greater occipital N1 and anterior P1 amplitudes at ∼155 ms than after PL, and lead to sustained differences over anterior, bilateral parietal and occipital sites from 205 ms onwards. For all non-social stimuli, OT evoked greater right parietal N1 amplitudes, and later only impacted threat stimuli over right parietal and occipital sites. None of these OT-induced modulations was related to individual anxiety levels. This pattern of results indicates that OT differentially modulates the processing of faces and non-social stimuli, and that the hormone's effect on visual processing and cognition does not occur as a function of non-clinical levels of anxiety.

Dissociation between goal-directed and discrete response localization in a patient with bilateral cortical blindness.

We investigated localization performance of simple targets in patient TN, who suffered bilateral damage of his primary visual cortex and shows complete cortical blindness. Using a two-alternative forced-choice paradigm, TN was asked to guess the position of left-right targets with goal-directed and discrete manual responses. The results indicate a clear dissociation between goal-directed and discrete responses. TN pointed toward the correct target location in approximately 75% of the trials but was at chance level with discrete responses. This indicates that the residual ability to localize an unseen stimulus depends critically on the possibility to translate a visual signal into a goal-directed motor output at least in certain forms of blindsight.

The effect of social comparison on effort: When similar and slightly better peers increase effort-related cardiovascular responses.

Social comparison theory states that comparison with others should influence an individual's behavior (Festinger, 1954; Munkes & Diehl, 2003). This is primarily due to an upward pressure: the pressure to be better than others, which according to some theories should motivate individuals to increase their level of performance (Munkes & Diehl, 2003; Rijsman, 1974). The effect of upward pressure on individual performance has been tested, but never on effort. To address this gap, we conducted a within-subject design study with N = 40 participants engaged in five-minute video games while presented with scores of a similar, slightly better, or weaker peer, with a control condition omitting the peer's score. Effort-related cardiovascular responses were recorded with initial systolic time interval (ISTI) during the game and baseline conditions. The effect of social comparison on effort was tested with a 4 (social comparison) x 5 (minutes of the tasks) repeated-measures ANOVA on ISTI reactivity. Results showed higher ISTI reactivity, interpreted as increased effort, when participants competed with similar and slightly better peers compared to a weaker peer and the control condition in the last minute of the task, confirming our expectations (Pegna et al., 2019). These results illustrate that social comparison - through its effect on upward pressure - is sufficient to elicit changes in effort-related cardiovascular response.

Up close and emotional: Electrophysiological dynamics of approaching angry faces.

Recent evidence suggests that looming emotional faces are processed rapidly by the neural system, and that this apparent approach further interacts with emotion, causing an enhanced neural response for angry expressions. However, previous research has not demonstrated unequivocally if these effects are due to low-level visual features, or if they are indeed due to the emotional content of the stimuli. To address this question, the current study presented upright and inverted angry and neutral faces, which either expanded or contracted in size on a constant depth-cued background, such that they appeared to approach or retreat from the viewer. EEG/ERP measures were used to identify the time course of brain activity for these stimuli. The results showed that when faces were upright, both the P1 and N170 were enhanced for angry expressions, with the P1 being further increased with looming angry faces. The inversion of the faces caused an increase in both the P1 and N170 amplitudes, but no modulation was found for emotions. These findings show an early modulation of brain activity for upright looming angry faces and rule out the influence of low-level visual features as a contributing factor.

Decoding neural patterns for the processing of fearful faces under different visual awareness conditions: A multivariate pattern analysis.

Previous studies have provided mixed findings regarding the nonconscious processing of fearful faces. Here, we used multivariate pattern analysis on electroencephalography data from three backward masking experiments to examine the processing of fearful faces under different visual awareness conditions. Three groups of participants were shown pairs of face images presented very briefly (for 16 ms) or for sufficiently long (for 266 ms), and completed tasks where the faces were either relevant to the experimental task (Experiment 1) or not (Experiments 2 and 3). Three main decoding analyses were performed. First, in the visual awareness decoding, the visibility of the faces, and hence participants' awareness of them, was maximally decodable in three time windows: 158-168 ms, 235-260 ms and 400-600 ms where the earlier neural patterns were generalized to the later stage activity. Second, we found that the spatial location of a fearful face in the face pairs was decodable, however only when the faces were consciously seen and task-relevant. Finally, we successfully decoded distinct neural patterns associated with the fearful-face-present conditions, compared to the fearful-face-absent conditions, and these patterns were decodable during both short and long presentations of the faces. Together, our results suggest that, while the processing of the spatial location of fearful faces requires awareness and task-relevancy, the mere presence of fearful faces can be processed even when visual awareness is highly restricted.

Faces capture spatial attention only when we want them to: An inattentional blindness EEG study.

Previous research on emotional face processing has shown that emotional faces such as fearful faces may be processed without visual awareness. However, evidence for nonconscious attention capture by fearful faces is limited. In fact, studies using sensory manipulation of awareness (e.g., backward masking paradigms) have shown that fearful faces do not attract attention during subliminal viewings nor when they were task-irrelevant. Here, we used a three-phase inattentional blindness paradigm and electroencephalography to examine whether faces (fearful and neutral) capture attention under different conditions of awareness and task-relevancy. We found that the electrophysiological marker for attention capture, the N2-posterior-contralateral (N2pc), was elicited by face stimuli only when participants were aware of the faces and when they were task-relevant (phase 3). When participants were unaware of the presence of faces (phase 1) or when the faces were irrelevant to the task (phase 2), no N2pc was observed. Together with our previous work, we concluded that fearful faces, or faces in general, do not attract attention unless we want them to.

Attentional capture by fearful faces requires consciousness and is modulated by task-relevancy: A dot-probe EEG study.

In the current EEG study, we used a dot-probe task in conjunction with backward masking to examine the neural activity underlying awareness and spatial processing of fearful faces and the neural processes for subsequent cued spatial targets. We presented face images under different viewing conditions (subliminal and supraliminal) and manipulated the relation between a fearful face in the pair and a subsequent target. Our mass univariate analysis showed that fearful faces elicit the N2-posterior-contralateral, indexing spatial attention capture, only when they are presented supraliminally. Consistent with this, the multivariate pattern analysis revealed a successful decoding of the location of the fearful face only in the supraliminal viewing condition. Additionally, the spatial attention capture by fearful faces modulated the processing of subsequent lateralised targets that were spatially congruent with the fearful face, in both al and electrophysiological data. There was no evidence for nonconscious processing of the fearful faces in the current paradigm. We conclude that spatial attentional capture by fearful faces requires visual awareness and it is modulated by top-down task demands.

Fixation-related electrical potentials during a free visual search task reveal the timing of visual awareness.

It has been repeatedly claimed that emotional faces readily capture attention, and that they may be processed without awareness. Yet some observations cast doubt on these assertions. Part of the problem may lie in the experimental paradigms employed. Here, we used a free viewing visual search task during electroencephalographic recordings, where participants searched for either fearful or neutral facial expressions among distractor expressions. Fixation-related potentials were computed for fearful and neutral targets and the response compared for stimuli consciously reported or not. We showed that awareness was associated with an electrophysiological negativity starting at around 110 ms, while emotional expressions were distinguished on the N170 and early posterior negativity only when stimuli were consciously reported. These results suggest that during unconstrained visual search, the earliest electrical correlate of awareness may emerge as early as 110 ms, and fixating at an emotional face without reporting it may not produce any unconscious processing.