EEG correlates of categorical and graded face perception

Face perception is a critical social ability and identifying its neural correlates is important from both basic and applied perspectives. In EEG recordings, faces elicit a distinct electrophysiological signature, the N170, which has a larger amplitude and shorter latency in response to faces compared to other objects. However, determining the face specificity of any neural marker for face perception hinges on finding an appropriate control stimulus. We used a novel stimulus set consisting of 300 images that spanned a continuum between random patches of natural scenes and genuine faces, in order to explore the selectivity of face-sensitive ERP responses with a model-based parametric stimulus set. Critically, our database contained “false alarm” images that were misclassified as face by computational face-detection system and varied in their image-level similarity to real faces. High-density (128-channel) event-related potentials (ERPs) were recorded while 23 adult subjects viewed all 300 images in random order, and determined whether each image was a face or non-face. The goal of our analyses was to determine the extent to which a gradient of sensitivity to face-like structure was evident in the ERP signal. Traditional waveform analyses revealed that the N170 component over occipitotemporal electrodes was larger in amplitude for faces compared to all non-faces, even those that were high in image similarity to faces, suggesting strict selectivity for veridical face stimuli. By contrast, single-trial classification of the entire waveform measured at the same sensors revealed that misclassifications of non-face patterns as faces increased with image-level similarity to faces. These results suggest that individual components may exhibit steep selectivity, but integration of multiple waveform features may afford graded information regarding stimulus appearance.     

Autistic traits modulate cortical responses to affective but not discriminative touch

The sense of touch is primarily considered a discriminative and exteroceptive sense, facilitating the detection, manipulation and exploration of objects, via an array of low‐threshold mechanoreceptors and fast conducting A‐beta (Aβ) afferents. However, a class of unmyelinated, low‐threshold mechanoreceptors identified in the hairy skin of mammals have been proposed to constitute a second, anatomically distinct system coding the affective qualities of touch. Unlike Aβs, which increase their firing rate linearly with the velocity of a stimulus moving across their receptive field, the response of these C‐tactile afferents (CTs) is described by an inverted ‘U’ curve fit, responding optimally to a skin temperature stimulus moving at between 1 and 10 cm/s. Given the distinct velocity tuning of these fast and slow touch fibres, here we used event‐related potentials to compare the time course of neural responses to 1st (fast) and 2nd (slow) touch systems. We identified a higher amplitude P300 in response to fast, Aβ‐targeted, versus slow CT‐targeted, stroking touch. In contrast, we identified a previously described, C‐fibre specific, ultra‐late potential (ULP) associated with CT‐targeted input. Of special note as regards the function of CTs is that the amplitude of the ULP was negatively correlated with self‐reported levels of autistic traits, which is consistent with the hypothesized affective and social significance of this response. Taken together, these findings provide further support for distinct discriminative and affective touch systems and suggests the temporal resolution of EEG provides an as yet underutilized tool for exploring individual differences in response sensitivity to CT‐targeted touch.     

Effects of personal familiarity on early neuromagnetic correlates of face perception

This study investigated effects of familiarity and orientation on face processing by means of magnetoencephalography. Participants were presented with photographs of personally familiar, famous and unfamiliar faces in both upright and inverted orientation. They had to decide whether faces were familiar by means of manual yes/no responses. Independent of orientation, we observed a clear modulation of the M170 by familiarity, with personally familiar faces evoking larger amplitudes than unknown faces. The M170 was also sensitive to orientation, with larger amplitudes for inverted than upright faces. Moreover, the M170 exhibited larger amplitudes over the right than over the left hemisphere, but this asymmetry was present for upright faces only. The present data suggest that at least for personally familiar faces, neural correlates of identification start no later than approximately 170 ms, and underline a special role of the right hemisphere for faces in their typical upright orientation.     

Being BOLD: The neural dynamics of face perception

According to a non‐hierarchical view of human cortical face processing, selective responses to faces may emerge in a higher‐order area of the hierarchy, in the lateral part of the middle fusiform gyrus (fusiform face area [FFA]) independently from face‐selective responses in the lateral inferior occipital gyrus (occipital face area [OFA]), a lower order area. Here we provide a stringent test of this hypothesis by gradually revealing segmented face stimuli throughout strict linear descrambling of phase information [Ales et al., 2012]. Using a short sampling rate (500 ms) of fMRI acquisition and single subject statistical analysis, we show a face‐selective responses emerging earlier, that is, at a lower level of structural (i.e., phase) information, in the FFA compared with the OFA. In both regions, a face detection response emerging at a lower level of structural information for upright than inverted faces, both in the FFA and OFA, in line with behavioral responses and with previous findings of delayed responses to inverted faces with direct recordings of neural activity were also reported. Overall, these results support the non‐hierarchical view of human cortical face processing and open new perspectives for time‐resolved analysis at the single subject level of fMRI data obtained during continuously evolving visual stimulation. Hum Brain Mapp 38:120–139, 2017. © 2016 Wiley Periodicals, Inc.     

The perception of musical phrase structure: a cross-cultural ERP study

Electroencephalography (EEG) was used in a cross-cultural music study investigating phrase boundary perception. Chinese and German musicians performed a cultural categorization task under Chinese and Western music listening conditions. Western music was the major subject for both groups of musicians, while Chinese music was familiar to Chinese subjects only. By manipulating the presence of pauses between two phrases in the biphrasal melodies, EEG correlates for the perception of phrase boundaries were found in both groups under both music listening conditions. Between 450 and 600 ms, the music CPS (closure positive shift), which had been found in earlier studies with a false tone detection task, was replicated for the more global categorization task and for all combinations of subject group and musical style. At short latencies (100 and 450 ms post phrase boundary offset), EEG correlates varied as a function of musical styles and subject group. Both bottom-up (style properties of the music) and top-down (acculturation of the subjects) information interacted during this early processing stage.     

Mapping the time course of nonconscious and conscious perception of fear: an integration of central and peripheral measures

Neuroimaging studies using backward masking suggest that conscious and nonconscious responses to complex signals of fear (facial expressions) occur via parallel cortical and subcortical circuits. Little is known, however, about the temporal differentiation of these responses. Psychophysics procedures were first used to determine objective thresholds for both nonconscious detection (face vs. blank screen) and discrimination (fear vs. neutral face) in a backward masking paradigm. Event-related potentials (ERPs) were then recorded (n = 20) using these thresholds. Ten blocks of masked fear and neutral faces were presented under each threshold condition. Simultaneously recorded skin conductance responses (SCRs) provided an independent index of stimulus perception. It was found that Fear stimuli evoked faster SCR rise times than did neutral stimuli across all conditions, indicating that emotional content influenced responses, regardless of awareness. In the first 400 msec of processing, ERPs dissociated the time course of conscious (enhanced N4 component) from nonconscious (enhanced N2 component) perception of fear, relative to neutral. Nonconscious detection of fear also elicited relatively faster P1 responses within 100 msec post-stimulus. The N2 may provide a temporal correlate of the initial sensory processing of salient facial configurations, which is enhanced when top-down cortical feedback is precluded. By contrast, the N4 may index the conscious integration of emotion stimuli in working memory, subserved by greater cortical engagement. Hum. Brain Mapping 21:64-74, 2004.     

Stereotypes bias face perception via orbitofrontal–fusiform cortical interaction

Previous research has shown that social-conceptual associations, such as stereotypes, can influence the visual representation of faces and neural pattern responses in ventral temporal cortex (VTC) regions, such as the fusiform gyrus (FG). Current models suggest that this social-conceptual impact requires medial orbitofrontal cortex (mOFC) feedback signals during perception. Backward masking can disrupt such signals, as it is a technique known to reduce functional connectivity between VTC regions and regions outside VTC. During functional magnetic resonance imaging (fMRI), subjects passively viewed masked and unmasked faces, and following the scan, perceptual biases and stereotypical associations were assessed. Multi-voxel representations of faces across the VTC, and in the FG and mOFC, reflected stereotypically biased perceptions when faces were unmasked, but this effect was abolished when faces were masked. However, the VTC still retained the ability to process masked faces and was sensitive to their categorical distinctions. Functional connectivity analyses confirmed that masking disrupted mOFC–FG connectivity, which predicted a reduced impact of stereotypical associations in the FG. Taken together, our findings suggest that the biasing of face representations in line with stereotypical associations does not arise from intrinsic processing within the VTC and FG alone, but instead it depends in part on top-down feedback from the mOFC during perception.     

TMS-EEG approach unveils brain mechanisms underlying conscious and unconscious face perception

BackgroundConscious perception of external stimuli has been related to recurrent activity in distributed cortical networks, although brain mechanisms controlling unconscious processing and stimuli access to conscious report need to be clarified.ObjectiveThis study aims at investigating modulations in cortical excitability related to conscious perception and unconscious processing of face stimuli with different visibility levels.MethodsWe used TMS-EEG over the right occipital face area (rOFA), or the right premotor cortex (rPMC) as control site, to measure cortical excitability during a backward masking paradigm with individually defined stimuli visibility.ResultsEvent related potentials showed significant differences for faces compared to houses, and detected faces compared to missed ones, 200 ms post target onset. TMS over rOFA, but not over rPMC, triggered a relative positivity starting 150 ms post target when faces with high visibility were consciously reported. Moreover, rOFA TMS evoked differential responses for high versus low visible faces in conscious and unconscious processing at 290–390 and 180–240 ms, respectively.ConclusionResults unveiled a causal link between rOFA excitability and late responses related to access to conscious perception, suggesting a critical role of recurrent activity, but distinct components, for consciously perceived stimuli and unconscious face processing.     

Autistic traits are linked to reduced adaptive coding of face identity and selectively poorer face recognition in men but not women

Our ability to discriminate and recognize thousands of faces despite their similarity as visual patterns relies on adaptive, norm-based, coding mechanisms that are continuously updated by experience. Reduced adaptive coding of face identity has been proposed as a neurocognitive endophenotype for autism, because it is found in autism and in relatives of individuals with autism. Autistic traits can also extend continuously into the general population, raising the possibility that reduced adaptive coding of face identity may be more generally associated with autistic traits. In the present study, we investigated whether adaptive coding of face identity decreases as autistic traits increase in an undergraduate population. Adaptive coding was measured using face identity aftereffects, and autistic traits were measured using the Autism-Spectrum Quotient (AQ) and its subscales. We also measured face and car recognition ability to determine whether autistic traits are selectively related to face recognition difficulties. We found that men who scored higher on levels of autistic traits related to social interaction had reduced adaptive coding of face identity. This result is consistent with the idea that atypical adaptive face-coding mechanisms are an endophenotype for autism. Autistic traits were also linked with face-selective recognition difficulties in men. However, there were some unexpected sex differences. In women, autistic traits were linked positively, rather than negatively, with adaptive coding of identity, and were unrelated to face-selective recognition difficulties. These sex differences indicate that autistic traits can have different neurocognitive correlates in men and women and raise the intriguing possibility that endophenotypes of autism can differ in males and females.     

Electrophysiological measures of attention during speech perception predict metalinguistic skills in children

Event-related potential (ERP) evidence demonstrates that preschool-aged children selectively attend to informative moments such as word onsets during speech perception. Although this observation indicates a role for attention in language processing, it is unclear whether this type of attention is part of basic speech perception mechanisms, higher-level language skills, or general cognitive abilities. The current study examined these possibilities by measuring ERPs from 5-year-old children listening to a narrative containing attention probes presented before, during, and after word onsets as well as at random control times. Children also completed behavioral tests assessing verbal and nonverbal skills. Probes presented after word onsets elicited a more negative ERP response beginning around 100 ms after probe onset than control probes, indicating increased attention to word-initial segments. Crucially, the magnitude of this difference was correlated with performance on verbal tasks, but showed no relationship to nonverbal measures. More specifically, ERP attention effects were most strongly correlated with performance on a complex metalinguistic task involving grammaticality judgments. These results demonstrate that effective allocation of attention during speech perception supports higher-level, controlled language processing in children by allowing them to focus on relevant information at individual word and complex sentence levels.     

The other-race effect for face perception: an event-related potential study

Summary It is well known that a recognition bias can be observed whenever subjects have to decide whether they have seen a person before that belongs to a different ethical group. Although this “other-race effect” is well documented on a behavioural level, its underlying mechanisms remain unclear. One plausible explanation might be that cortical areas involved in face processing are not as effective for other-race faces due to a missing experience with individuals from other ethnical groups. This interpretation is strongly supported by a functional magnetic resonance imaging study showing decreased brain activity on other-race faces. Furthermore, two event-related potential studies revealed differences in brain activity in the first 250 ms after face presentation, but with inconsistent results. Therefore, we investigated 12 Caucasian subjects, showing them faces of Asian and Caucasian subjects in a perceptual priming paradigm and measured the event-related brain potentials. On a behavioural level we found slower reaction times to Asian faces compared to Caucasian faces in the unprimed condition, reflecting a deficit for Caucasian subjects to process other-race faces. In accordance with these behavioural data we see a significantly reduced late N250r amplitude in the unprimed condition to the Asian faces compared to the Caucasian faces. These results clearly indicate that the other-race effect was present in our sample and very specific only in the unprimed condition around 350–450 ms after stimulus onset.     

Autistic traits modulate frontostriatal connectivity during processing of rewarding faces

Deficits in facial mimicry have been widely reported in autism. Some studies have suggested that these deficits are restricted to spontaneous mimicry and do not extend to volitional mimicry. We bridge these apparently inconsistent observations by testing the impact of reward value on neural indices of mimicry and how autistic traits modulate this impact. Neutral faces were conditioned with high and low reward. Subsequently, functional connectivity between the ventral striatum (VS) and inferior frontal gyrus (IFG) was measured while neurotypical adults (n = 30) watched happy expressions made by these conditioned faces. We found greater VS–IFG connectivity in response to high reward vs low reward happy faces. This difference was negatively proportional to autistic traits, suggesting that reduced spontaneous mimicry of social stimuli seen in autism, may be related to a failure in the modulation of the mirror system by the reward system rather than a circumscribed deficit in the mirror system.     

The role of face shape and pigmentation in other-race face perception: An electrophysiological study

Adult observers generally find it difficult to recognize and distinguish faces that belong to categories with which they have limited visual experience. One aspect of this phenomenon is commonly known as the “Other-Race Effect” (ORE) since this behavior is typically highly evident in the perception of faces belonging to ethnic or racial groups other than that of the observer. This acquired disadvantage in face recognition likely results from highly specific “tuning” of the underlying representation of facial appearance, leading to efficient processing of commonly seen faces at the expense of poor generalization to other face categories. In the current study we used electrophysiological (event-related potentials or ERPs) and behavioral measures of performance to characterize face processing in racial categories defined by dissociable shape and pigmentation information. Our goal was to examine the specificity of the representation of facial appearance in more detail by investigating how race-specific face shape and pigmentation separately modulated neural responses previously implicated in face processing, the N170 and N250 components. We found that both components were modulated by skin color, independent of face shape, but that only the N250 exhibited sensitivity to face shape. Moreover, the N250 appears to only respond differentially to the skin color of upright faces, showing a lack of color sensitivity for inverted faces.     

The fusiform face area and occipital face area show sensitivity to spatial relations in faces

Behavioral research indicates that successful face individuation is associated with sensitivity to subtle spatial relations between facial features, as well as to the features themselves. We used a blocked functional magnetic resonance adaptation paradigm to examine the sensitivity of the core face network to spatial relations in faces. The fusiform face area (FFA) was sensitive to spatial relations, responding more strongly to a single face presented with various feature spacings than to repeated presentations of an identical face. This response to spacing variations was as strong as the response to a series of distinct identities. There were no hemisphere effects in sensitivity to spatial relations, although FFAs were larger on the right. The right occipital face area (OFA) was also sensitive to spatial relations in faces. Few participants showed left OFAs. The superior temporal sulcus (STS), which does not code identity, showed little sensitivity to either relational changes or changes in identity. We suggest that the sensitivity of the FFA and right OFA to spatial relations in faces may contribute to our impressive ability to individuate faces despite their similarity as visual patterns.     

Traumatic life events and its impact on fibromyalgia symptoms through serotonin activity on pain perception and personality traits

Fibromyalgia is a chronic syndrome characterized by widespread muscular pain and tenderness with no evidence of soft tissue inflammation. Early-life stressors and traumatic events have been described to increase the risk of developing a number of psychiatric conditions, and conditions related to chronic pain in later life such as fibromyalgia on the other hand. In this overview of the literature on the topic, we present the impact that traumatic life events can have on fibromyalgia symptoms through serotonin activity on pain perception and personality traits.     

Neural responses to faces reflect social personality traits

Faces are a developmentally primary and critically important source of social information, and they are processed differently from most other visual percepts. Studies of brain electrophysiology reveal a face-sensitive component, the N170, which is typically enhanced to faces relative to other stimuli. Research in social disabilities suggests that an atypical N170 response in this population may stem from decreased developmental exposure to faces secondary to reduced social interest. Here we examined the relationship between neural responses to faces and social personality characteristics in a normative sample. Participants were pre-screened to identify individuals scoring high on extraversion or introversion. Both groups were presented with upright and inverted face stimuli. An inversion effect, a marker of expertise for faces, was observed in people with high extraversion but not in those with high introversion. These findings suggest that, within typically developing populations, social attitudes are reflected in the neural correlates of face perception.     

Colour constancy and conscious perception of changes of illuminant

A sudden change in illuminant (e.g., the outcome of turning on a tungsten light in a room illuminated with dim, natural daylight) causes a "global" change in perceived colour which subjects often recognise as a change of illuminant. In spite of this distinct, global change in the perceptual appearance of the scene caused by significant changes in the wavelength composition of the light reflected from different objects under the new illuminant, the perceived colour of the objects remains largely unchanged and this cornerstone property of human vision is often described as instantaneous colour constancy (ICC). ICC mechanisms are often difficult to study. The generation of appropriate stimuli to isolate ICC mechanisms remains a difficult task since the extraction of colour signals is also confounded in the processing of spatial chromatic context that leads to ICC. The extraction of differences in chromaticity that describe spatial changes in the wavelength composition of the light on the retina is a necessary operation that must precede colour constancy computations. A change of illuminant or changes in the spectral reflectance of the elements that make up the scene under a constant illuminant cause spatial changes in chromatic context and are likely to drive colour constancy mechanisms, but not exclusively. The same stimulus changes also cause differences in local luminance contrast and overall light flux changes, stimulus attributes that can activate different areas of the visual cortex. In order to address this problem we carried out a series of dichoptic experiments designed to investigate how the colour signals from the two eyes are combined in dichoptically viewed Mondrians and the extent to which the processing of chromatic context in monocularly driven neurons contributes to ICC. The psychophysical findings show that normal levels of ICC can be achieved in dichoptic experiments, even when the subject remains unaware of any changes of illuminant. Functional MRI (fMRI) experiments using new stimuli that produce stimulation of colour constancy mechanisms only in one condition with little or no difference in the activity generated in colour processing mechanisms in both test and reference conditions were also carried out. The results show that the processing of ICC signals generates strong activation in V1 and the fusiform colour area (V4, V4A). Significant activation was also observed in areas V2 and V3.     

Abnormal activation of the social brain during face perception in autism

ASD involves a fundamental impairment in processing social-communicative information from faces. Several recent studies have challenged earlier findings that individuals with autism spectrum disorder (ASD) have no activation of the fusiform gyrus (fusiform face area, FFA) when viewing faces. In this study, we examined activation to faces in the broader network of face-processing modules that comprise what is known as the social brain. Using 3T functional resonance imaging, we measured BOLD signal changes in 10 ASD subjects and 7 healthy controls passively viewing nonemotional faces. We replicated our original findings of significant activation of face identity-processing areas (FFA and inferior occipital gyrus, IOG) in ASD. However, in addition, we identified hypoactivation in a more widely distributed network of brain areas involved in face processing [including the right amygdala, inferior frontal cortex (IFC), superior temporal sulcus (STS), and face-related somatosensory and premotor cortex]. In ASD, we found functional correlations between a subgroup of areas in the social brain that belong to the mirror neuron system (IFC, STS) and other face-processing areas. The severity of the social symptoms measured by the Autism Diagnostic Observation Schedule was correlated with the right IFC cortical thickness and with functional activation in that area. When viewing faces, adults with ASD show atypical patterns of activation in regions forming the broader face-processing network and social brain, outside the core FFA and IOG regions. These patterns suggest that areas belonging to the mirror neuron system are involved in the face-processing disturbances in ASD.     

Understanding early communication signals in autism: a study of the perception of infants' cry

Background Previous studies have highlighted that episodes of crying of children with autistic disorder (AD) were perceived as inexplicable from their parents who could not identify causative factors. These results supported the view of AD as related to a problem of expressing and sharing emotions. Moreover, no evidence has been presented on which characteristics of a cry episode influence the adult perception. Aim of our research is to investigate how acoustical features of crying episodes modulate their perception of infants with ASD compared with infants with typical development (TD) and infants with developmental delay (DD). Methods Two studies were employed. In study 1, we artificially modified structural parameters (fundamental frequency, duration of the pauses, waveform modulation) of a cry episode, and then 50 adults (parents and non‐parents) were asked to judge the level of distress elicited. In study 2, acoustic analysis was applied to episodes of crying selected from retrospective home videos of 42 children with AD, TD and DD at 18 months. Results The results showed that (1) differences in the fundamental frequency and in other structural parameters of the cry lead parents and non‐parents to perceive an episode of crying as more aversive and (2) at 18 months of age, AD episodes of crying have higher fundamental frequency (f0). Conclusion Our findings offer support for the hypothesis that acoustic characteristics of episodes of crying of children with autism, especially higher fundamental frequencies, may account for mental states of uneasiness in the listener.