Linguistic and attentional factors - Not statistical regularities - Contribute to word-selective neural responses with FPVS-oddball paradigms.

Studies using frequency-tagging in electroencephalography (EEG) have dramatically increased in the past 10 years, in a variety of domains and populations. Here we used Fast Periodic Visual Stimulation (FPVS) combined with an oddball design to explore visual word recognition. Given the paradigm's high sensitivity, it is crucial for future basic research and clinical application to prove its robustness across variations of designs, stimulus types and tasks. This paradigm uses periodicity of brain responses to measure discrimination between two experimentally defined categories of stimuli presented periodically. EEG was recorded in 22 adults who viewed words inserted every 5 stimuli (at 2 Hz) within base stimuli presented at 10 Hz. Using two discrimination levels (deviant words among nonwords or pseudowords), we assessed the impact of relative frequency of item repetition (set size or item repetition controlled for deviant versus base stimuli), and of the orthogonal task (focused or deployed spatial attention). Word-selective occipito-temporal responses were robust at the individual level (significant in 95% of participants), left-lateralized, larger for the prelexical (nonwords) than lexical (pseudowords) contrast, and stronger with a deployed spatial attention task as compared to the typically used focused task. Importantly, amplitudes were not affected by item repetition. These results help understanding the factors influencing word-selective EEG responses and support the validity of FPVS-EEG oddball paradigms, as they confirm that word-selective responses are linguistic. Second, they show its robustness against design-related factors that could induce statistical (ir)regularities in item rate. They also confirm its high individual sensitivity and demonstrate how it can be optimized, using a deployed rather than focused attention task, to measure implicit word recognition processes in typical and atypical populations.

The neuropsychological evaluation of face identity recognition.

Facial identity recognition (FIR) is arguably the ultimate form of recognition for the adult human brain. Even if the term prosopagnosia is reserved for exceptionally rare brain-damaged cases with a category-specific abrupt loss of FIR at adulthood, subjective and objective impairments or difficulties of FIR are common in the neuropsychological population. Here we provide a critical overview of the evaluation of FIR both for clinicians and researchers in neuropsychology. FIR impairments occur following many causes that should be identified objectively by both general and specific, behavioral and neural examinations. We refute the commonly used dissociation between perceptual and memory deficits/tests for FIR, since even a task involving the discrimination of unfamiliar face images presented side-by-side relies on cortical memories of faces in the right-lateralized ventral occipito-temporal cortex. Another frequently encountered confusion is between specific deficits of the FIR function and a more general impairment of semantic memory (of people), the latter being most often encountered following anterior temporal lobe damage. Many computerized tests aimed at evaluating FIR have appeared over the last two decades, as reviewed here. However, despite undeniable strengths, they often suffer from ecological limitations, difficulties of instruction, as well as a lack of consideration for processing speed and qualitative information. Taking into account these issues, a recently developed behavioral test with natural images manipulating face familiarity, stimulus inversion, and correct response times as a key variable appears promising. The measurement of electroencephalographic (EEG) activity in the frequency domain from fast periodic visual stimulation also appears as a particularly promising tool to complete and enhance the neuropsychological assessment of FIR.

Extensive Visual Training in Adulthood Reduces an Implicit Neural Marker of the Face Inversion Effect.

Face identity recognition (FIR) in humans is supported by specialized neural processes whose function is spectacularly impaired when simply turning a face upside-down: the face inversion effect (FIE). While the FIE appears to have a slow developmental course, little is known about the plasticity of the neural processes involved in this effect-and in FIR in general-at adulthood. Here, we investigate whether extensive training (2 weeks, ~16 h) in young human adults discriminating a large set of unfamiliar inverted faces can reduce an implicit neural marker of the FIE for a set of entirely novel faces. In all, 28 adult observers were trained to individuate 30 inverted face identities presented under different depth-rotated views. Following training, we replicate previous behavioral reports of a significant reduction (56% relative accuracy rate) in the behavioral FIE as measured with a challenging four-alternative delayed-match-to-sample task for individual faces across depth-rotated views. Most importantly, using EEG together with a validated frequency tagging approach to isolate a neural index of FIR, we observe the same substantial (56%) reduction in the neural FIE at the expected occipito-temporal channels. The reduction in the neural FIE correlates with the reduction in the behavioral FIE at the individual participant level. Overall, we provide novel evidence suggesting a substantial degree of plasticity in processes that are key for face identity recognition in the adult human brain.

The anterior fusiform gyrus: The ghost in the cortical face machine.

Face-selective regions in the human ventral occipito-temporal cortex (VOTC) have been defined for decades mainly with functional magnetic resonance imaging. This face-selective VOTC network is traditionally divided in a posterior 'core' system thought to subtend face perception, and regions of the anterior temporal lobe as a semantic memory component of an extended general system. In between these two putative systems lies the anterior fusiform gyrus and surrounding sulci, affected by magnetic susceptibility artifacts. Here we suggest that this methodological gap overlaps with and contributes to a conceptual gap between (visual) perception and semantic memory for faces. Filling this gap with intracerebral recordings and direct electrical stimulation reveals robust face-selectivity in the anterior fusiform gyrus and a crucial role of this region, especially in the right hemisphere, in identity recognition for both familiar and unfamiliar faces. Based on these observations, we propose an integrated theoretical framework for human face (identity) recognition according to which face-selective regions in the anterior fusiform gyrus join the dots between posterior and anterior cortical face memories.

Stability of face recognition abilities after left or right anterior temporal lobectomy.

Patients with anterior temporal lobe (ATL) resection due to mesial temporal lobe epilepsy (MTLE) have difficulties at identifying familiar faces and explicitly remembering newly learned faces but their ability to individuate unfamiliar faces remains largely unknown. Moreover, the extent to which their difficulties with familiar face identity recognition and learning is truly due to the ATL resection remains unknown. Here, we report a study of 24 MTLE patients and matched healthy controls tested with an extensive set of seven face and visual object recognition tasks (including three tasks evaluating unfamiliar face individuation) before and about 6 months after unilateral (nine left, 15 right) ATL resection. We found that ATL resection has little or no effect on the patients' preserved pre-surgical ability to perform unfamiliar face individuation, both at the group and individual levels. More surprisingly, ATL resection also has little effect on the patients' performance at recognizing and naming famous faces as well as at learning new faces. A substantial proportion of right MTLE patients (33%) even improved their response times on several tasks, which may indicate a functional release of visuo-spatial processing after resection in the right ATL. Altogether this study shows that face recognition abilities are mainly unaffected by ATL resection in MTLE, either because the critical regions for face recognition are spared or because performance at some tasks is already lower than normal preoperatively. Overall, these findings urge caution when interpreting the causal effect of brain lesions on face recognition ability in patients with ATL resection due to MTLE. They also illustrate the complexity of predicting cognitive outcomes after epilepsy surgery because of the influence of many different intertwined factors.

Intracerebral electrical stimulation of the face-selective right lateral fusiform gyrus transiently impairs face identity recognition.

Neuroimaging and intracranial electrophysiological studies have consistently shown the largest and most consistent face-selective neural activity in the middle portion of the human right lateral fusiform gyrus ('fusiform face area(s)', FFA). Yet, direct evidence for the critical role of this region in face identity recognition (FIR) is still lacking. Here we report the first evidence of transient behavioral impairment of FIR during focal electrical stimulation of the right FFA. Upon stimulation of an electrode contact within this region, subject CJ, who shows typical FIR ability outside of stimulation, was transiently unable to point to pictures of famous faces among strangers and to match pictures of famous or unfamiliar faces presented simultaneously for their identity. Her performance at comparable tasks with other visual materials (written names, pictures of buildings) remained unaffected by stimulation at the same location. During right FFA stimulation, CJ consistently reported that simultaneously presented faces appeared as being the same identity, with little or no distortion of the spatial face configuration. Independent electrophysiological recordings showed the largest neural face-selective and face identity activity at the critical electrode contacts. Altogether, this extensive multimodal case report supports the causal role of the right FFA in FIR.

A neural marker of the human face identity familiarity effect.

Human adults associate different views of an identity much better for familiar than for unfamiliar faces. However, a robust and consistent neural index of this behavioral face identity familiarity effect (FIFE)-not found in non-human primate species-is lacking. Here we provide such a neural FIFE index, measured implicitly and with one fixation per face. Fourteen participants viewed 70 s stimulation sequences of a large set (n = 40) of widely variable natural images of a face identity at a rate of 6 images/second (6 Hz). Different face identities appeared every 5th image (1.2 Hz). In a sequence, face images were either familiar (i.e., famous) or unfamiliar, participants performing a non-periodic task unrelated to face recognition. The face identity recognition response identified at 1.2 Hz over occipital-temporal regions in the frequency-domain electroencephalogram was 3.4 times larger for familiar than unfamiliar faces. The neural response to familiar faces-which emerged at about 180 ms following face onset-was significant in each individual but a case of prosopdysgnosia. Besides potential clinical and forensic applications to implicitly measure one's knowledge of a face identity, these findings open new perspectives to clarify the neurofunctional source of the FIFE and understand the nature of human face identity recognition.

Single neuron responses underlying face recognition in the human midfusiform face-selective cortex.

Faces are critical for social interactions and their recognition constitutes one of the most important and challenging functions of the human brain. While neurons responding selectively to faces have been recorded for decades in the monkey brain, face-selective neural activations have been reported with neuroimaging primarily in the human midfusiform gyrus. Yet, the cellular mechanisms producing selective responses to faces in this hominoid neuroanatomical structure remain unknown. Here we report single neuron recordings performed in 5 human subjects (1 male, 4 females) implanted with intracerebral microelectrodes in the face-selective midfusiform gyrus, while they viewed pictures of familiar and unknown faces and places. We observed similar responses to faces and places at the single cell level, but a significantly higher number of neurons responding to faces, thus offering a mechanistic account for the face-selective activations observed in this region. Although individual neurons did not respond preferentially to familiar faces, a population level analysis could consistently determine whether or not the faces (but not the places) were familiar, only about 50 ms after the initial recognition of the stimuli as faces. These results provide insights into the neural mechanisms of face processing in the human brain.

Automatic, Early Color-Specific Neural Responses to Object Color Knowledge.

Some familiar objects are associated with specific colors, e.g., rubber ducks with yellow. Whether and at what stage neural responses occur to these color associations remain open questions. We recorded frequency-tagged electroencephalogram (EEG) responses to periodic presentations of yellow-associated objects, shown among sequences of non-periodic blue-, red-, and green-associated objects. Both color and grayscale versions of the objects elicited yellow-specific responses, indicating an automatic activation of color knowledge from object shape. Follow-up experiments replicated these effects with green-specific responses, and demonstrated modulated responses for incongruent color/object associations. Importantly, the onset of color-specific responses was as early to grayscale as actually colored stimuli (before 100 ms), the latter additionally eliciting a conventional later response (approximately 140-230 ms) to actual stimulus color. This suggests that the neural representation of familiar objects includes both diagnostic shape and color properties, such that shape can elicit associated color-specific responses before actual color-specific responses occur.

Spatial Resolution Evaluation Based on Experienced Visual Categories With Sweep Evoked Periodic EEG Activity.

Purpose: Visual function is typically evaluated in clinical settings with visual acuity (VA), a test requiring to behaviorally match or name optotypes such as tumbling E or Snellen letters. The ability to recognize these symbols has little in common with the automatic and rapid visual recognition of socially important stimuli in real life. Here we use sweep visual evoked potentials to assess spatial resolution objectively based on the recognition of human faces and written words. Methods: To this end, we tested unfamiliar face individuation1 and visual word recognition2 in 15 normally sighted adult volunteers with a 68-electrode electroencephalogram system. Results: Unlike previous measures of low-level visual function including VA, the most sensitive electrode was found at an electrode different from Oz in a majority of participants. Thresholds until which faces and words could be recognized were evaluated at the most sensitive electrode defined individually for each participant. Word recognition thresholds corresponded with the VA level expected from normally sighted participants, and even a VA significantly higher than expected from normally sighted individuals for a few participants. Conclusions: Spatial resolution can be evaluated based on high-level stimuli encountered in day-to-day life, such as faces or written words with sweep visual evoked potentials.

Intracerebral Electrophysiological Recordings to Understand the Neural Basis of Human Face Recognition.

Understanding how the human brain recognizes faces is a primary scientific goal in cognitive neuroscience. Given the limitations of the monkey model of human face recognition, a key approach in this endeavor is the recording of electrophysiological activity with electrodes implanted inside the brain of human epileptic patients. However, this approach faces a number of challenges that must be overcome for meaningful scientific knowledge to emerge. Here we synthesize a 10 year research program combining the recording of intracerebral activity (StereoElectroEncephaloGraphy, SEEG) in the ventral occipito-temporal cortex (VOTC) of large samples of participants and fast periodic visual stimulation (FPVS), to objectively define, quantify, and characterize the neural basis of human face recognition. These large-scale studies reconcile the wide distribution of neural face recognition activity with its (right) hemispheric and regional specialization and extend face-selectivity to anterior regions of the VOTC, including the ventral anterior temporal lobe (VATL) typically affected by magnetic susceptibility artifacts in functional magnetic resonance imaging (fMRI). Clear spatial dissociations in category-selectivity between faces and other meaningful stimuli such as landmarks (houses, medial VOTC regions) or written words (left lateralized VOTC) are found, confirming and extending neuroimaging observations while supporting the validity of the clinical population tested to inform about normal brain function. The recognition of face identity - arguably the ultimate form of recognition for the human brain - beyond mere differences in physical features is essentially supported by selective populations of neurons in the right inferior occipital gyrus and the lateral portion of the middle and anterior fusiform gyrus. In addition, low-frequency and high-frequency broadband iEEG signals of face recognition appear to be largely concordant in the human association cortex. We conclude by outlining the challenges of this research program to understand the neural basis of human face recognition in the next 10 years.

Towards an optimization of functional localizers in non-human primate neuroimaging with (fMRI) frequency-tagging.

Non-human primate (NHP) neuroimaging can provide essential insights into the neural basis of human cognitive functions. While functional magnetic resonance imaging (fMRI) localizers can play an essential role in reaching this objective (Russ et al., 2021), they often differ substantially across species in terms of paradigms, measured signals, and data analysis, biasing the comparisons. Here we introduce a functional frequency-tagging face localizer for NHP imaging, successfully developed in humans and outperforming standard face localizers (Gao et al., 2018). FMRI recordings were performed in two awake macaques. Within a rapid 6 Hz stream of natural non-face objects images, human or monkey face stimuli were presented in bursts every 9 s. We also included control conditions with phase-scrambled versions of all images. As in humans, face-selective activity was objectively identified and quantified at the peak of the face-stimulation frequency (0.111 Hz) and its second harmonic (0.222 Hz) in the Fourier domain. Focal activations with a high signal-to-noise ratio were observed in regions previously described as face-selective, mainly in the STS (clusters PL, ML, MF; also, AL, AF), both for human and monkey faces. Robust face-selective activations were also found in the prefrontal cortex of one monkey (PVL and PO clusters). Face-selective neural activity was highly reliable and excluded all contributions from low-level visual cues contained in the amplitude spectrum of the stimuli. These observations indicate that fMRI frequency-tagging provides a highly valuable approach to objectively compare human and monkey visual recognition systems within the same framework.

A neural marker of rapid discrimination of facial expression in 3.5- and 7-month-old infants.

Infants' ability to discriminate facial expressions has been widely explored, but little is known about the rapid and automatic ability to discriminate a given expression against many others in a single experiment. Here we investigated the development of facial expression discrimination in infancy with fast periodic visual stimulation coupled with scalp electroencephalography (EEG). EEG was recorded in eighteen 3.5- and eighteen 7-month-old infants presented with a female face expressing disgust, happiness, or a neutral emotion (in different stimulation sequences) at a base stimulation frequency of 6 Hz. Pictures of the same individual expressing other emotions (either anger, disgust, fear, happiness, sadness, or neutrality, randomly and excluding the expression presented at the base frequency) were introduced every six stimuli (at 1 Hz). Frequency-domain analysis revealed an objective (i.e., at the predefined 1-Hz frequency and harmonics) expression-change brain response in both 3.5- and 7-month-olds, indicating the visual discrimination of various expressions from disgust, happiness and neutrality from these early ages. At 3.5 months, the responses to the discrimination from disgust and happiness expressions were located mainly on medial occipital sites, whereas a more lateral topography was found for the response to the discrimination from neutrality, suggesting that expression discrimination from an emotionally neutral face relies on distinct visual cues than discrimination from a disgust or happy face. Finally, expression discrimination from happiness was associated with a reduced activity over posterior areas and an additional response over central frontal scalp regions at 7 months as compared to 3.5 months. This result suggests developmental changes in the processing of happiness expressions as compared to negative/neutral ones within this age range.

Low and high frequency intracranial neural signals match in the human associative cortex.

In vivo intracranial recordings of neural activity offer a unique opportunity to understand human brain function. Intracranial electrophysiological (iEEG) activity related to sensory, cognitive or motor events manifests mostly in two types of signals: event-related local field potentials in lower frequency bands (<30 Hz, LF) and broadband activity in the higher end of the frequency spectrum (>30 Hz, High frequency, HF). While most current studies rely exclusively on HF, thought to be more focal and closely related to spiking activity, the relationship between HF and LF signals is unclear, especially in human associative cortex. Here, we provide a large-scale in-depth investigation of the spatial and functional relationship between these 2 signals based on intracranial recordings from 121 individual brains (8000 recording sites). We measure category-selective responses to complex ecologically salient visual stimuli - human faces - across a wide cortical territory in the ventral occipito-temporal cortex (VOTC), with a frequency-tagging method providing high signal-to-noise ratio (SNR) and the same objective quantification of signal and noise for the two frequency ranges. While LF face-selective activity has higher SNR across the VOTC, leading to a larger number of significant electrode contacts especially in the anterior temporal lobe, LF and HF display highly similar spatial, functional, and timing properties. Specifically, and contrary to a widespread assumption, our results point to nearly identical spatial distribution and local spatial extent of LF and HF activity at equal SNR. These observations go a long way towards clarifying the relationship between the two main iEEG signals and reestablish the informative value of LF iEEG to understand human brain function.

Natural Contrast Statistics Facilitate Human Face Categorization.

The ability to detect faces in the environment is of utmost ecological importance for human social adaptation. While face categorization is efficient, fast and robust to sensory degradation, it is massively impaired when the facial stimulus does not match the natural contrast statistics of this visual category, i.e., the typically experienced ordered alternation of relatively darker and lighter regions of the face. To clarify this phenomenon, we characterized the contribution of natural contrast statistics to face categorization. Specifically, 31 human adults viewed various natural images of nonface categories at a rate of 12 Hz, with highly variable images of faces occurring every eight stimuli (1.5 Hz). As in previous studies, neural responses at 1.5 Hz as measured with high-density electroencephalography (EEG) provided an objective neural index of face categorization. Here, when face images were shown in their naturally experienced contrast statistics, the 1.5-Hz face categorization response emerged over occipito-temporal electrodes at very low contrast [5.1%, or 0.009 root-mean-square (RMS) contrast], quickly reaching optimal amplitude at 22.6% of contrast (i.e., RMS contrast of 0.041). Despite contrast negation preserving an image's spectral and geometrical properties, negative contrast images required twice as much contrast to trigger a face categorization response, and three times as much to reach optimum. These observations characterize how the internally stored natural contrast statistics of the face category facilitate visual processing for the sake of fast and efficient face categorization.

Superior neural individuation of mother's than stranger's faces by five months of age.

Human adults are better at recognizing different views of a given face as belonging to the same person when that person is familiar rather than unfamiliar. To clarify the developmental origin of this well-established phenomenon, one group of five-month-olds (N = 22) was presented with pictures of four different unfamiliar female faces at a fixed rate (6 Hz, 166 msec stimulus onset asynchrony), interrupted every 5th stimulus (1.2 Hz) by either their mother's face (mother oddball condition) or, in different stimulation sequences, a stranger's face (stranger oddball condition). In another group of five-month-olds (N = 17), stimulation sequences were reversed such that their mothers' or a given stranger's face were repeated at 6 Hz and interrupted every 5 stimuli by pictures of different female faces (mother standard, stranger standard conditions, respectively). Twelve variable images of each identity served as stimulus material. Besides clear frequency-tagged EEG responses at the 6 Hz stimulation rate over the medial occipital region in all conditions, significant activity at 1.2 Hz and harmonics (2.4 Hz, etc.) was observed in this region, reflecting selective responses to facial identity across changes of views. This effect was strongest when the mother's face was immediately repeated at every stimulation cycle (mother standard). Overall, these observations point to an early developmental advantage of identifying a familiar face presented from different views during immediate stimulus repetition.

Twenty years of investigation with the case of prosopagnosia PS to understand human face identity recognition.Part II: Neural basis.

Patient PS sustained her dramatic brain injury thirty years ago, in 1992, the same year as the first report of a neuroimaging study of human face recognition.The present paper complements the review on the functional nature of PS's prosopagnosia (part I), illustrating how her case study directly, i.e., through neuroimaging investigations of her brain structure and activity, but also indirectly, through neural studies performed on other clinical cases and neurotypical individuals, inspired and constrained neural models of human face recognition.In the dominant right hemisphere for face recognition in humans, PS's main lesion concerns (inputs to) the inferior occipital gyrus (IOG), in a region where face-selective activity is typically found in normal individuals ('Occipital Face Area', OFA).Her case study initially supported the criticality of this region for face identity recognition (FIR) and provided the impetus for transcranial magnetic stimulation (TMS), intracerebral electrical stimulation, and cortical surgery studies that have generally supported this view.Despite PS's right IOG lesion, typical face-selectivity is found anteriorly in the middle portion of the fusiform gyrus, a hominoid structure.This face-selective right 'Fusiform Face Area' (FFA) has been widely considered as the most important region for human face recognition.This finding led to the original proposal of direct anatomico-functional connections from early visual cortices to the FFA, bypassing the IOG/OFA , a hypothesis supported by further neuroimaging studies of PS, other neurological cases and neuro-typical individuals with original visual stimulation paradigms, data recordings and analyses.The proposal of a lack of sensitivity to face identity in PS's right FFA due to defective reentrant inputs from the IOG/FFA has also been supported by other cases, functional connectivity and cortical surgery studies.Overall, neural studies of, and based on, the case of prosopagnosia PS strongly question the hierarchical organization of the human neural face recognition system, supporting a more flexible and dynamic view of this key social brain function in our species.

Twenty years of investigation with the case of prosopagnosia PS to understand human face identity recognition. Part I: Function.

Following traumatic brain injury in adulthood, Pierrette Sapey (PS) became suddenly unable to recognize the identity of people from their faces. Thanks to her remarkable recovery of general brain function, liveliness, and willingness to be tested, PS's case of prosopagnosia has been extensively studied for more than 20 years. This investigation includes hundreds of hours of behavioral data collection that provide information about the nature of human face identity recognition (FIR). Here a theory-driven extensive review of behavioral and eye movement recording studies performed with PS is presented (part I). The specificity of PS's recognition disorder to the category of faces, i.e., with preserved visual object (identity) recognition, is emphasized, arguing that isolating this impairment is necessary to define prosopagnosia, offering a unique window to understand the nature of human FIR. Studies performed with both unfamiliar and experimentally or naturally familiar faces show that PS, while being able to perceive both detailed diagnostic facial parts and a coarse global facial shape, can no longer build a relatively fine-grained holistic visual representation of a face, preventing its efficient individuation. Her mandatory part-by-part analytic behavior during FIR causes increased difficulties at extracting diagnostic cues from the crowded eye region of the face, but also from relative distances between facial parts and from 3D shape more than from surface cues. PS's impairment is interpreted here for the first time in terms of defective (access to) cortical memories of faces following brain damage, causing her impaired holistic perception of face individuality. Implications for revising standard neurofunctional models of human face recognition and evaluation of this function in neurotypical individuals are derived.

Developmental experiences alter the temporal processing characteristics of the visual cortex: Evidence from deaf and hearing native signers.

To date, the extent to which early experience shapes the functional characteristics of neural circuits is still a matter of debate. In the present study, we tested whether congenital deafness and/or the acquisition of a sign language alter the temporal processing characteristics of the visual system. Moreover, we investigated whether, assuming cross-modal plasticity in deaf individuals, the temporal processing characteristics of possibly reorganised auditory areas resemble those of the visual cortex. Steady-state visual evoked potentials (SSVEPs) were recorded in congenitally deaf native signers, hearing native signers, and hearing nonsigners. The luminance of the visual stimuli was periodically modulated at 12, 21, and 40 Hz. For hearing nonsigners, the optimal driving rate was 12 Hz. By contrast, for the group of hearing signers, the optimal driving rate was 12 and 21 Hz, whereas for the group of deaf signers, the optimal driving rate was 21 Hz. We did not observe evidence for cross-modal recruitment of auditory cortex in the group of deaf signers. These results suggest a higher preferred neural processing rate as a consequence of the acquisition of a sign language.

Rapid neural categorization of facelike objects predicts the perceptual awareness of a face (face pareidolia).

The human brain rapidly and automatically categorizes faces vs. other visual objects. However, whether face-selective neural activity predicts the subjective experience of a face - perceptual awareness - is debated. To clarify this issue, here we use face pareidolia, i.e., the illusory perception of a face, as a proxy to relate the neural categorization of a variety of facelike objects to conscious face perception. In Experiment 1, scalp electroencephalogram (EEG) is recorded while pictures of human faces or facelike objects - in different stimulation sequences - are interleaved every second (i.e., at 1 Hz) in a rapid 6-Hz train of natural images of nonface objects. Participants do not perform any explicit face categorization task during stimulation, and report whether they perceived illusory faces post-stimulation. A robust categorization response to facelike objects is identified at 1 Hz and harmonics in the EEG frequency spectrum with a facelike occipito-temporal topography. Across all individuals, the facelike categorization response is of about 20% of the response to human faces, but more strongly right-lateralized. Critically, its amplitude is much larger in participants who report having perceived illusory faces. In Experiment 2, facelike or matched nonface objects from the same categories appear at 1 Hz in sequences of nonface objects presented at variable stimulation rates (60 Hz to 12 Hz) and participants explicitly report after each sequence whether they perceived illusory faces. The facelike categorization response already emerges at the shortest stimulus duration (i.e., 17 ms at 60 Hz) and predicts the behavioral report of conscious perception. Strikingly, neural facelike-selectivity emerges exclusively when participants report illusory faces. Collectively, these experiments characterize a neural signature of face pareidolia in the context of rapid categorization, supporting the view that face-selective brain activity reliably predicts the subjective experience of a face from a single glance at a variety of stimuli.