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The face inversion effect is an important indicator of holistic face perception and reflects the developmental level of face processing. This study examined the face inversion effect in deaf or hard of hearing (DHH) children aged 7-17 using the face dimensions task. This task uses photographic images of a face, in which configural and featural information in the eye and mouth regions have been parametrically and independently manipulated. The study aimed to discuss the effect of face inversion on facial processing in DHH children, including two aspects of information processing types (configural versus featural) and processing regions (eyes versus mouth) and compared the results with hearing children. The results revealed that DHH children aged 7-17 years exhibit significant face inversion effect, with disruptions observed in both the featural and configural processing of eyes and mouths when faces were inverted. Configural processing was more affected by inversion than featural processing in all children, with larger differences observed in DHH children than in hearing children. This supports the dual-mode hypothesis of holistic face processing. Age correlations were observed in the sensitivity of DHH children to face inversion effect but not among hearing children. The inversion effect of configural mouth processing decreases with age in DHH children.

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We report a large study (n = 72) using combined transcranial direct current stimulation-electroencephalography (tDCS-EEG) to investigate the modulation of perceptual learning indexed by the face inversion effect. Participants were engaged with an old/new recognition task involving intermixed upright and inverted, normal and Thatcherized faces. The accuracy results showed anodal tDCS delivered at the Fp3 scalp area (cathode/reference electrode placed at Fp2) increased the behavioural inversion effect for normal faces versus sham/control and this covaried with a modulation of the N170 event-related potential component. A reduced inversion effect for normal faces was found on the N170 latency and amplitude versus sham/control, extending recent work that combined tDCS and EEG in circumstances where the behavioural face inversion effect was reduced. Our results advance understanding of the neural mechanisms responsible for perceptual learning by revealing a dissociation between the N170 amplitude and latency in response to the tDCS-induced modulation of the face inversion effect. The behavioural modulation of the inversion effect tracks the modulation of the N170 amplitudes, albeit it is negatively correlated (i.e., reduced inversion effect-larger N170 amplitude inversion effect, increased inversion effect-reduced N170 amplitude inversion effect). For the N170 latencies, the inversion effect is reduced by the tDCS protocol we use irrespective of any modulation of the behavioural inversion effect.

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It is widely held that upright faces are processed more holistically than inverted faces and that this difference is reflected in the face inversion effect. It is not clear, however, how the inversion effect can best be measured, whether it is task specific, or even whether it specifically correlates with processing of upright faces. We examined these questions in a large sample (N = 420) who provided data on processing of upright and inverted stimuli in two different tasks with faces and one with objects. We find that the inversion effects are task dependent, and that they do not correlate better among face processing tasks than they do across face and object processing tasks. These findings were obtained regardless of whether inversion effects were measured by means of difference scores or regression. In comparison, only inversion effects based on regression predicted performance with upright faces in tasks other than those the inversion effects were derived from. Critically, however, inversion effects based on regression also predicted performance with inverted faces to a similar degree as they predicted performance with upright faces. Consequently, and contrary to what is commonly assumed, inversion effects do not seem to capture effects specific to holistic processing of upright faces. While the present findings do not bring us closer to an understanding of which changes in cognitive processing are induced by inversion, they do suggest that inversion effects do not reflect a unitary construct; an implicit assumption that seems to characterize much of the research regarding face processing.

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Humans pay special attention to faces and speech from birth, but the interplay of developmental processes leading to specialization is poorly understood. We investigated the effects of face orientation on audiovisual (AV) speech perception in two age groups of infants (younger: 5- to 6.5-month-olds; older: 9- to 10.5-month-olds) and adults. We recorded event-related potentials (ERP) in response to videos of upright and inverted faces producing /ba/ articulation dubbed with auditory syllables that were either matching /ba/ or mismatching /ga/ the mouth movement. We observed an increase in the amplitude of audiovisual mismatch response (AVMMR) to incongruent visual /ba/-auditory /ga/ syllable in comparison to other stimuli in younger infants, while the older group of infants did not show a similar response. AV mismatch response to inverted visual /ba/-auditory /ga/ stimulus relative to congruent stimuli was also detected in the right frontal areas in the younger group and the left and right frontal areas in adults. We show that face configuration affects the neural response to AV mismatch differently across all age groups. The novel finding of the AVMMR in response to inverted incongruent AV speech may potentially imply the featural face processing in younger infants and adults when processing inverted faces articulating incongruent speech. The lack of visible differential responses to upright and inverted incongruent stimuli obtained in the older group of infants suggests a likely functional cortical reorganization in the processing of AV speech.

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Research has demonstrated that attractiveness evaluations of adult faces were less accurate when faces were inverted than upright. It remains unknown, however, whether a similar effect applies to perceived cuteness of infants, which is assumed to be based on elemental facial features called the "baby schema." In this research, we studied the face inversion effect on perceived cuteness of infant faces in a rating task and a two-alternative forced-choice (2AFC) task. We also examined beauty as a control dimension. Although the rating task revealed no inversion effect, the 2AFC task showed poorer discrimination performance with inverted faces than with upright faces in both evaluations. These results indicate that infant cuteness and beauty dimensions are correlated well with each other, and their perception not only relies on elemental features that are not strongly affected by inversion but is also affected by holistic facial configurations when a detailed comparison is required.

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In a spatial Stroop task, the eye-gaze target produces the reversed congruency effect-responses become shorter when the gaze direction and its location are incongruent than when they are congruent. The present study examined the face inversion effect on the gaze spatial Stroop task to clarify whether the holistic face processing or part-based processing of the eyes is responsible for the reversed congruency effect. In Experiment 1, participants judged the gaze direction of the upright or inverted face with a neutral expression presented either in the left or right visual field. In Experiment 2, we examined whether face inversion interacted with facial expressions (i.e., angry, happy, neutral, and sad). Face inversion disrupted holistic face processing, slowing down the overall performance relative to the performance with upright faces. However, face inversion did not affect the reversed congruency effect. These results further support the parts-based processing account and suggest that while faces are processed holistically, the reversed congruency effect, relying on the extracted local features (i.e., eyes), may be processed in a part-based manner.

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The face inversion effect (FIE) refers to the observation that presenting stimuli upside-down impairs the processing of faces disproportionally more than other mono-oriented objects. This has been taken as evidence that processing of faces and objects differ qualitatively. However, nearly all FIE studies are based on comparing individuation of upright faces, which most people are rather good at, with individuation of objects most people are much less familiar with individuating (e.g., radios and airplanes). Consequently, the FIE may mainly reflect differences between categories in how they are processed prior to inversion, with within-category discrimination of upright faces being a much more familiar task than within-category discrimination among members belonging to other object classes. We tested this hypothesis by comparing inversion effects for faces and objects using object recognition tasks that do not require within-category discrimination (object decision and old/new recognition memory tasks). In all tasks (seven with objects and two with faces) we find credible inversion effects, but in no instance were these effects significantly larger for faces than for objects. This suggests that the FIE can be a product of familiarity with the type of identification process required in the upright conditions rather than some process that is selectively affected for faces when stimuli are inverted.

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To explore the number of latent variables underlying recognition of own- and other-race faces for Chinese observers, we conducted a study-recognition task where orientation, stimuli type, and duration were manipulated in the study phase and applied state trace analysis as a statistic method. Results showed that each state trace plot on each pair of stimuli types matched a single monotonic curve when stimuli type was set to state factor, but separate curves between face and non-face showed up when the state factor was orientation. The results implied that at least two latent variables affected recognition performance in the inversion paradigm. Besides, the unidimensional structure between own- and other-race faces regardless of the state factor suggested that Chinese participants used the same recognition mechanism for unfamiliar own- and other-race faces in the inversion paradigm.

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BACKGROUND: Studying the neural processing of faces can illuminate the mechanisms of compromised social expertise in autism. To resolve a longstanding debate, we examined whether differences in configural face processing in autism are underpinned by quantitative differences in the activation of typical face processing pathways, or the recruitment of non-typical neural systems. METHODS: We investigated spatial and temporal characteristics of event-related EEG responses to upright and inverted faces in a large sample of children, adolescents, and adults with and without autism. We examined topographic analyses of variance and global field power to identify group differences in the spatial and temporal response to face inversion. We then examined how quasi-stable spatiotemporal profiles - microstates - are modulated by face orientation and diagnostic group. RESULTS: Upright and inverted faces produced distinct profiles of topography and strength in the topographical analyses. These topographical profiles differed between diagnostic groups in adolescents, but not in children or adults. In the microstate analysis, the autistic group showed differences in the activation strength of normative microstates during early-stage processing at all ages, suggesting consistent quantitative differences in the operation of typical processing pathways; qualitative differences in microstate topographies during late-stage processing became prominent in adults, suggesting the increasing involvement of non-typical neural systems with processing time and over development. CONCLUSIONS: These findings suggest that early difficulties with configural face processing may trigger later compensatory processes in autism that emerge in later development.

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The face inversion effect (FIE) is a behavioral marker of face-specific processing that the recognition of inverted faces is disproportionately disrupted than that of inverted non-face objects. One hypothesis is that while upright faces are represented by face-specific mechanism, inverted faces are processed as objects. However, evidence from neuroimaging studies is inconclusive, possibly because the face system, such as the fusiform face area, is interacted with the object system, and therefore the observation from the face system may indirectly reflect influences from the object system. Here we examined the FIE in an artificial face system, visual geometry group network-face (VGG-Face), a deep convolutional neural network (DCNN) specialized for identifying faces. In line with neuroimaging studies on humans, a stronger FIE was found in VGG-Face than that in DCNN pretrained for processing objects. Critically, further classification error analysis revealed that in VGG-Face, inverted faces were miscategorized as objects behaviorally, and the analysis on internal representations revealed that VGG-Face represented inverted faces in a similar fashion as objects. In short, our study supported the hypothesis that inverted faces are represented as objects in a pure face system.

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This study investigates whether stimulus inversion influences neural responses of Chinese character recognition similarly to its effect on face recognition in category-selective and object-related brain areas using functional magnetic resonance imaging. Participants performed a one-back matching task for simple (one radical) and compound (two radicals) Chinese characters and faces with upright and inverted orientations. Inverted stimuli produced slower response times with stronger activity within the fusiform gyrus (FG) than upright stimuli for faces and Chinese characters. While common inversion-related activation was identified in the left FG among stimulus types, we observed a significant inter-regional correlation between the left FG and the intraparietal sulcus for face inversion. Importantly, analyses of region-of-interest (ROI) multivariate pattern classification showed that classifiers trained on face inversion can decode the representations of character inversion in the character-selective ROI. However, this was not true for face inversion in face-selective ROIs when the classifiers were trained on characters. Similar activity patterns for character and face inversion were observed in the object-related ROIs. We also showed higher decoding accuracy for upright stimuli in the face-selective ROI than in the character-selective ROI but this was not true for inverted ones or when patterns were examined in the object-related ROIs. Together, our results support shared and distinct configural representations for character and face recognition in category-selective and object-related brain areas.

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Facial disfigurements can influence how observers attend to and interact with the person, leading to disease-avoidance behaviour and emotions (disgust, threat, fear for contagion). However, it is unclear whether this behaviour is reflected in the effect of the facial stigma on attention and perceptual encoding of facial information. We addressed this question by measuring, in a mixed antisaccade task, observers' speed and accuracy of orienting of visual attention towards or away from peripherally presented upright and inverted unfamiliar faces that had either a realistic looking disease-signalling feature (a skin discolouration), a non-disease-signalling control feature, or no added feature. The presence of a disfiguring or control feature did not influence the orienting of attention (in terms of saccadic latency) towards upright faces, suggesting that avoidance responses towards facial stigma do not occur during covert attention. However, disfiguring and control features significantly reduced the effect of face inversion on saccadic latency, thus suggesting an impact on the holistic processing of facial information. The implications of these findings for the encoding and appraisal of facial disfigurements are discussed.

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OBJECTIVE: Impaired face perception is considered as a hallmark of social disability in schizophrenia. It is widely believed that inverted faces and upright faces are processed by distinct mechanisms. Previous studies have identified that individuals with schizophrenia display poorer face processing than controls. However, the mechanisms underlying the face inversion effect (FIE) in patients with first-episode schizophrenia (FSZ) remain unclear. METHODS: We designed an fMRI task to investigate the FIE mechanism in patients with schizophrenia. Thirty-four patients with FSZ and thirty-five healthy controls (CON) underwent task-related fMRI scanning, clinical assessment, anhedonia experience examination, and social function and cognitive function evaluation. RESULTS: The patients with FSZ exhibited distinct functional activity regarding upright and inverted face processing within the cortical face and non-face network. These results suggest that the differences in quantitative processing might mediate the FIE in schizophrenia. Compared with controls, affected patients showed impairments in processing both upright and inverted faces; and for these patients with FSZ, upright face processing was associated with more severe and broader impairment than inverted face processing. Reduced response in the left middle occipital gyrus for upright face processing was related to poorer performance of social function outcomes evaluated using the Personal and Social Performance Scale. CONCLUSION: Our data suggested that patients with FSZ exhibited similar performance in processing inverted faces and upright faces, but were less efficient than controls; and for these patients, inverted faces are processed less efficiently than upright faces. We also provided a clue that the mechanism under abnormal FIE might be related to an aberrant activation of non-face-selective areas instead of abnormal activation of face-specific areas in patients with schizophrenia. Finally, our study indicated that the neural pathway for upright recognition might be relevant in determining the functional outcomes of this devastating disorder.

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Eye movement studies show that humans can make very fast saccades towards faces in natural scenes, but the visual mechanisms behind this process remain unclear. Here we investigate whether fast saccades towards faces rely on mechanisms that are sensitive to the orientation or contrast of the face image. We present participants pairs of images each containing a face and a car in the left and right visual field or the reverse, and we ask them to saccade to faces or cars as targets in different blocks. We assign participants to one of three image conditions: normal images, orientation-inverted images, or contrast-negated images. We report three main results that hold regardless of image conditions. First, reliable saccades towards faces are fast - they can occur at 120-130 ms. Second, fast saccades towards faces are selective - they are more accurate and faster by about 60-70 ms than saccades towards cars. Third, saccades towards faces are reflexive - early saccades in the interval of 120-160 ms tend to go to faces, even when cars are the target. These findings suggest that the speed, selectivity, and reflexivity of saccades towards faces do not depend on the orientation or contrast of the face image. Our results accord with studies suggesting that fast saccades towards faces are mainly driven by low-level image properties, such as amplitude spectrum and spatial frequency.

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Feed-forward deep neural networks have better performance in object categorization tasks than other models of computer vision. To understand the relationship between feed-forward deep networks and the primate brain, we investigated representations of upright and inverted faces in a convolutional deep neural network model and compared them with representations by neurons in the monkey anterior inferior-temporal cortex, area TE. We applied principal component analysis to feature vectors in each model layer to visualize the relationship between the vectors of the upright and inverted faces. The vectors of the upright and inverted monkey faces were more separated through the convolution layers. In the fully-connected layers, the separation among human individuals for upright faces was larger than for inverted faces. The Spearman correlation between each model layer and TE neurons reached a maximum at the fully-connected layers. These results indicate that the processing of faces in the fully-connected layers might resemble the asymmetric representation of upright and inverted faces by the TE neurons. The separation of upright and inverted faces might take place by feed-forward processing in the visual cortex, and separations among human individuals for upright faces, which were larger than those for inverted faces, might occur in area TE.

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This investigation examined whether impairment in configural processing could explain deficits in face emotion recognition in people with Parkinson's disease (PD). Stimuli from the Radboud Faces Database were used to compare recognition of four negative emotion expressions by older adults with PD (n = 16) and matched controls (n = 17). Participants were tasked with categorizing emotional expressions from upright and inverted whole faces and facial composites; it is difficult to derive configural information from these two types of stimuli so featural processing should play a larger than usual role in accurate recognition of emotional expressions. We found that the PD group were impaired relative to controls in recognizing anger, disgust and fearful expressions in upright faces. Then, consistent with a configural processing deficit, participants with PD showed no composite effect when attempting to identify facial expressions of anger, disgust and fear. A face inversion effect, however, was observed in the performance of all participants in both the whole faces and facial composites tasks. These findings can be explained in terms of a configural processing deficit if it is assumed that the disruption caused by facial composites was specific to configural processing, whereas inversion reduced performance by making it difficult to derive both featural and configural information from faces.

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The face inversion effect reflects the special nature of facial processing and appears not only in recognizing facial identity or expression but also in subjective evaluation, such as facial attractiveness. Previous studies have revealed that the way in which we perceive attractiveness (beauty versus cuteness) differs our perceptual behavior. Therefore, the face inversion effect on attractiveness might differ based on the viewpoint of attractiveness. In this study, we measured pupillary response when judging the cuteness of facial stimuli and focused on the mechanism of perceiving attractiveness in terms of the effect of involuntary physical reaction. We investigated whether perceived cuteness - a kind of attractiveness - was affected by face inversion and whether the face inversion effect appeared in pupillary responses. We then conducted experiments in which participants observed inverted faces and rated the subjective cuteness of the faces, and we measured the participants' pupil size while they observed the facial stimuli. The results revealed a negative correlation between pupil changes and the perceived cuteness of inverted faces, which is consistent with the previous result of upright faces. Thus, we found that the perception of facial cuteness is little affected by face inversion, suggesting that the judgment of cuteness is processed differently from other types of attractiveness such as beauty. We also found that pupillary response is related to perceiving cuteness, which could lead to consistency in the perception of cuteness.

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Introduction: Individuals tend to explore the left side of a face first and for a longer time in comparison to the right side. This left visual field (LVF) bias is suggested to reflect right hemispheric dominance for face processing. Social anxiety disorder (SAD) is associated with maladaptive interpretations of facial expressions, but it is not known whether this is linked to an atypical LVF bias. Previous studies have reported a reduced LVF bias in autism, a condition overlapping with SAD. This pre-registered study examined the LVF bias in adolescents with SAD. Methods: Eye-tracking was used to investigate the ratio of first fixations to the left on upright and inverted face stimuli in 26 adolescents (13-17 years) with SAD and 23 healthy controls primed to look either between the eyes or at the mouth. Results: The SAD group showed a smaller LVF bias and an atypical face inversion effect when primed to look at the eyes. Autistic traits predicted a smaller LVF bias, independently of social anxiety level. Conclusions: Results suggest that SAD is associated with impaired processing of faces at an early stage of visual scanning. The findings contribute to a better understanding of SAD and its overlap with autism.

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BACKGROUND: Body dysmorphic disorder (BDD) is characterized by an excessive preoccupation with one or more perceived flaws in one's own appearance. Previous studies provided evidence for deficits in configural and holistic processing in BDD. Preliminary evidence suggests abnormalities at an early stage of visual processing. The present study is the first examining early neurocognitive perception of the own face in BDD by using electroencephalography (EEG). We investigated the face inversion effect, in which inverted (upside-down) faces are disproportionately poorly processed compared to upright faces. This effect reflects a disruption of configural and holistic processing, and in consequence a preponderance of featural face processing. METHODS: We recorded face-sensitive event-related potentials (ERPs) in 16 BDD patients and 16 healthy controls, all unmedicated. Participants viewed upright and inverted (upside-down) images of their own face and an unfamiliar other face, each in two facial emotional expressions (neutral vs. smiling). We calculated the early ERP components P100, N170, P200, N250, and the late positive component (LPC), and compared amplitudes among both groups. RESULTS: In the early P100, no face inversion effects were found in both groups. In the N170, both groups exhibited the common face inversion effects, with significantly larger N170 amplitudes for inverted than upright faces. In the P200, both groups exhibited larger inversion effects to other (relative to own) faces, with larger P200 amplitudes for other upright than inverted faces. In the N250, no significant group differences were found in face processing. In the LPC, both groups exhibited larger inversion effects to other (relative to own) faces, with larger LPC amplitudes for other inverted than upright faces. These overall patterns appeared to be comparable for both groups. Smaller inversion effects to own (relative to other) faces were observed in none of these components in BDD, relative to controls. CONCLUSIONS: The findings suggest no evidence for abnormalities at all levels of early face processing in our observed sample of BDD patients. Further research should investigate the neural substrates underlying BDD symptomatology.

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Face and body perception is mediated by configural mechanisms, which allow the perception of these stimuli as a whole, rather than the sum of individual parts. Indirect measures of configural processing in visual cognition are the face and body inversion effects (FIE and BIE), which refer to the drop in performance when these stimuli are perceived upside-down. Albeit FIE and BIE have been well characterized at the behavioral level, much still needs to be understood in terms of the neurophysiological correlates of these effects. Thus, in the current study, the brain's electrical activity has been recorded by a 128 channel electroencephalogram (EEG) in 24 healthy participants while perceiving (upright and inverted) faces, bodies and houses. EEG data were analyzed in both the time domain (i.e., event-related potentials-ERPs) and the frequency domain [i.e., induced theta (5-7 Hz) and gamma (28-45 Hz) oscillations]. ERPs amplitude results showed increased N170 amplitude for inverted faces and bodies (compared to the same stimuli presented in canonical position) but not for houses. ERPs latency results showed delayed N170 components for inverted (vs. upright) faces, houses, but not bodies. Spectral analysis of induced oscillations indicated physiological FIE and BIE; that is decreased gamma-band synchronization over right occipito-temporal electrodes for inverted (vs. upright) faces, and increased bilateral frontoparietal theta-band synchronization for inverted (vs. upright) faces. Furthermore, increased left occipito-temporal and right frontal theta-band synchronization for upright (vs. inverted) bodies was found. Our findings, thus, demonstrate clear differences in the neurophysiological correlates of face and body perception. The neurophysiological FIE suggests disruption of feature binding processes (decrease in occipital gamma oscillations for inverted faces), together with enhanced feature-based attention (increase in frontoparietal theta oscillations for inverted faces). In contrast, the BIE may suggest that structural encoding for bodies is mediated by the first stages of configural processing (decrease in occipital theta oscillations for inverted bodies).