Cortical Face Research Articles

Face pareidolia minimally engages macaque face selective neurons.

The macaque cerebral cortex contains concentrations of neurons that prefer faces over inanimate objects. Although these so-called face patches are thought to be specialized for the analysis of facial signals, their exact tuning properties remain unclear. For example, what happens when an object by chance resembles a face? Everyday objects can sometimes, through the accidental positioning of their internal components, appear as faces. This phenomenon is known as face pareidolia. Behavioral experiments have suggested that macaques, like humans, perceive illusory faces in such objects. However, it is an open question whether such stimuli would naturally stimulate neurons residing in cortical face patches. To address this question, we recorded single unit activity from four fMRI-defined face-selective regions: the anterior medial (AM), anterior fundus (AF), prefrontal orbital (PO), and perirhinal cortex (PRh) face patches. We compared neural responses elicited by images of real macaque faces, pareidolia-evoking objects, and matched control objects. Contrary to expectations, we found no evidence of a general preference for pareidolia-evoking objects over control objects. Although a subset of neurons exhibited stronger responses to pareidolia-evoking objects, the population responses to both categories of objects were similar, and collectively much less than to real macaque faces. These results suggest that neural responses in the four regions we tested are principally concerned with the analysis of realistic facial characteristics, whereas the special attention afforded to face-like pareidolia stimuli is supported by activity elsewhere in the brain.

Effective connectivity of the human cortical face network through concurrent intracerebral electrical stimulation and frequency-tagged visual presentation

Effective connectivity of the human cortical face network through concurrent intracerebral electrical stimulation and frequency-tagged visual presentation

Impact of emotional valence on mismatch negativity in the course of cortical face processing.

Various aspects of cortical face processing have been studied by assessing event related potentials (ERP). It has been described in the literature that mismatch negativity (MMN), a well-studied ERP, is not only modulated by sensory features but also emotional valence. However, the exact impact of emotion on the temporo-spatial profile of visual MMN during face processing remains inconsistent. By employing a sequential oddball paradigm using both neutral and emotional deviants, we were able to differentiate two distinct vMMN subcomponents. While an early subcomponent at 150-250ms is elicited by emotional salient facial stimuli, the later subcomponent at 250-400ms seems to reflect the detection of regularity violations in facial recognition per se, unaffected by emotional salience. Our results suggest that emotional valence is encoded in vMMN signal strength at an early stage of facial processing. Furthermore, we assume that of facial processing consists of temporo-spatially distinct, partially overlapping levels concerning different facial aspects.

임실하가 구석기유적의 돌날제작기법 연구

The blade is a representative artifact of the upper Paleolithic of Korea. The study of the manufacturing techniques of the blade and restoration of it play an important role in understanding of survival skills of upper Paleolithic human. This paper researches into the technology and process of the blade production through the blade production technology we learned from ‘Refitting artifact’ of Imsil Haga site, and analogizes the kind of the hammer which they used for the blade production in comparison with the existing research result of experimental archaeology on the blade production. Restoring the blade production process as a result of the analysis of ‘refitting artifacts’, 1. Shaping, removed cortical face and unnecessary parts of the raw material are made into sizes and shapes suitable for blade production using stone hammer direct percussion (refitting artifacts 1, 3). 2. creation of a striking platform, 3. blade debitage using the Antler hammer direct percussion. (refitting artifacts 1, 3) 4. Change the striking platform according to the production process, and blade debitage by making the most of the stones several times (refitting artifacts 1, 2). 5. When knapping accidents (step, hinge, and flange) appeared on the blade core, or when the size of the core became smaller and it became difficult to make a blade of the desired size, the blade was discarded without production more blades. The reason for the abandonment of the blade was 1. If it is no longer possible to make the blade due to the Knapping accidents (step, plunging, hinged), 2. If the size of the blade cannot be made, The ‘Refitting artifact 1’ includes two tools (tanged point), which analyzed the length and production space of the tool that the tool maker wanted. First of all, considering that the length of the blade identified in ‘Refitting artifact 1’ is about 40 to 88mm, and the tanged point is about 51, 54mm, it is thought that the stone maker made a blade with a length of about 50 to 70mm or made a tool. The excavation location of tanged point is about 2m away from the central part of the bladecore, and it is thought that the tool was made by moving the blade after manufacturing it.

Top-down modulation and cortical-AMG/HPC interaction in familiar face processing.

Humans can accurately recognize familiar faces in only a few hundred milliseconds, but the underlying neural mechanism remains unclear. Here, we recorded intracranial electrophysiological signals from ventral temporal cortex (VTC), superior/middle temporal cortex (STC/MTC), medial parietal cortex (MPC), and amygdala/hippocampus (AMG/HPC) in 20 epilepsy patients while they viewed faces of famous people and strangers as well as common objects. In posterior VTC and MPC, familiarity-sensitive responses emerged significantly later than initial face-selective responses, suggesting that familiarity enhances face representations after they are first being extracted. Moreover, viewing famous faces increased the coupling between cortical areas and AMG/HPC in multiple frequency bands. These findings advance our understanding of the neural basis of familiar face perception by identifying the top-down modulation in local face-selective response and interactions between cortical face areas and AMG/HPC.

Brain-wide functional connectivity of face patch neurons during rest

The brain is a highly organized, dynamic system whose network architecture is often assessed through resting functional magnetic resonance imaging (fMRI) functional connectivity. The functional interactions between brain areas, including those observed during rest, are assumed to stem from the collective influence of action potentials carried by long-range neural projections. However, the contribution of individual neurons to brain-wide functional connectivity has not been systematically assessed. Here we developed a method to concurrently measure and compare the spiking activity of local neurons with fMRI signals measured across the brain during rest. We recorded spontaneous activity from neural populations in cortical face patches in the macaque during fMRI scanning sessions. Individual cells exhibited prominent, bilateral coupling with fMRI fluctuations in a restricted set of cortical areas inside and outside the face patch network, partially matching the pattern of known anatomical projections. Within each face patch population, a subset of neurons was positively coupled with the face patch network and another was negatively coupled. The same cells showed inverse correlations with distinct subcortical structures, most notably the lateral geniculate nucleus and brainstem neuromodulatory centers. Corresponding connectivity maps derived from fMRI seeds and local field potentials differed from the single unit maps, particularly in subcortical areas. Together, the results demonstrate that the spiking fluctuations of neurons are selectively coupled with discrete brain regions, with the coupling governed in part by anatomical network connections and in part by indirect neuromodulatory pathways.

Intracerebral electrical stimulation of the right anterior fusiform gyrus impairs human face identity recognition

Brain regions located between the right fusiform face area (FFA) in the middle fusiform gyrus and the temporal pole may play a critical role in human face identity recognition but their investigation is limited by a large signal drop-out in functional magnetic resonance imaging (fMRI). Here we report an original case who is suddenly unable to recognize the identity of faces when electrically stimulated on a focal location inside this intermediate region of the right anterior fusiform gyrus. The reliable transient identity recognition deficit occurs without any change of percept, even during nonverbal face tasks (i.e., pointing out the famous face picture among three options; matching pictures of unfamiliar or familiar faces for their identities), and without difficulty at recognizing visual objects or famous written names. The effective contact is associated with the largest frequency-tagged electrophysiological signals of face-selectivity and of familiar and unfamiliar face identity recognition. This extensive multimodal investigation points to the right anterior fusiform gyrus as a critical hub of the human cortical face network, between posterior ventral occipito-temporal face-selective regions directly connected to low-level visual cortex, the medial temporal lobe involved in generic memory encoding, and ventral anterior temporal lobe regions holding semantic associations to people's identity.

Cortical Pathways or Mechanism in the Face Inversion Effect in Patients with First-Episode Schizophrenia.

ObjectiveImpaired 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.MethodsWe 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.ResultsThe 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.ConclusionOur 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.

Complementary Phenomena: Phantom Hand and Phantom Face.

After tissue or limb loss, the development of sensation and perception of the lost or deafferent tissue is defined as a phantom phenomenon. We investigated the presence of phantom phenomena in individuals who underwent a full face transplant as well as those who had a hand transplant. Specifically, we investigated sensory perception of the face on the fingers and sensory perception of the fingers on the face in three full face and four hand transplant patients. In all seven individuals, we used a brush to separately stimulate the right and left sides of the face or the palmar and dorsal faces of the hand. We then asked the individuals if they felt a sensation of touch on any other part of their body and, if so, to describe their perceptions. Changes in the regions of the primary sensory cortex representing the hand and face were defined using fMRI obtained via tactile sensory stimulation of the clinical examination areas. Two of the full face transplant patients reported sensory perceptions such as a prominent sensation of touch on their faces during sensory stimulation of their fingers. Three of the hand transplant patients reported sensory perceptions, which we referred to as finger patches, during sensory stimulation of the face area. In fMRI, overlaps were observed in the cortical hand and face representation areas. We consider the phantom hand and phantom face phenomena we observed to be complementary due to the neighborhood of the representations of the hand and face in the somatosensory cortex.

Audiovisual integration in macaque face patch neurons

SummaryPrimate social communication depends on the perceptual integration of visual and auditory cues, reflected in the multimodal mixing of sensory signals in certain cortical areas. The macaque cortical face patch network, identified through visual, face-selective responses measured with fMRI, is assumed to contribute to visual social interactions. However, whether face patch neurons are also influenced by acoustic information, such as the auditory component of a natural vocalization, remains unknown. Here, we recorded single-unit activity in the anterior fundus (AF) face patch, in the superior temporal sulcus, and anterior medial (AM) face patch, on the undersurface of the temporal lobe, in macaques presented with audiovisual, visual-only, and auditory-only renditions of natural movies of macaques vocalizing. The results revealed that 76% of neurons in face patch AF were significantly influenced by the auditory component of the movie, most often through enhancement of visual responses but sometimes in response to the auditory stimulus alone. By contrast, few neurons in face patch AM exhibited significant auditory responses or modulation. Control experiments in AF used an animated macaque avatar to demonstrate, first, that the structural elements of the face were often essential for audiovisual modulation and, second, that the temporal modulation of the acoustic stimulus was more important than its frequency spectrum. Together, these results identify a striking contrast between two face patches and specifically identify AF as playing a potential role in the integration of audiovisual cues during natural modes of social communication.

Connectivity at the origins of domain specificity in the cortical face and place networks

It is well established that the adult brain contains a mosaic of domain-specific networks. But how do these domain-specific networks develop? Here we tested the hypothesis that the brain comes prewired with connections that precede the development of domain-specific function. Using resting-state fMRI in the youngest sample of newborn humans tested to date, we indeed found that cortical networks that will later develop strong face selectivity (including the "proto" occipital face area and fusiform face area) and scene selectivity (including the "proto" parahippocampal place area and retrosplenial complex) by adulthood, already show domain-specific patterns of functional connectivity as early as 27 d of age (beginning as early as 6 d of age). Furthermore, we asked how these networks are functionally connected to early visual cortex and found that the proto face network shows biased functional connectivity with foveal V1, while the proto scene network shows biased functional connectivity with peripheral V1. Given that faces are almost always experienced at the fovea, while scenes always extend across the entire periphery, these differential inputs may serve to facilitate domain-specific processing in each network after that function develops, or even guide the development of domain-specific function in each network in the first place. Taken together, these findings reveal domain-specific and eccentricity-biased connectivity in the earliest days of life, placing new constraints on our understanding of the origins of domain-specific cortical networks.

Connectivity at the origins of domain specificity: the case of the cortical face network

Connectivity at the origins of domain specificity: the case of the cortical face network

P11-F The cortical sources of face selective N170: A simultaneous multi-scale EEG study

The sudden onset of a face image leads to a prominent face-selective response in human scalp electroencephalographic (EEG) recordings, peaking 170 ms after stimulus onset at occipito-temporal (OT) scalp sites: the N170 (or M170 in magnetoencephalography). According to a widely held view, the main cortical source of the N170 lies in the fusiform gyrus (FG), whereas the posteriorly located inferior occipital gyrus (IOG) would rather generate earlier face-selective responses. Here, we report neural responses to upright and inverted faces recorded in a unique patient using multicontact intracerebral electrodes implanted in the right IOG and in the OT sulcus above the right lateral FG (LFG). Simultaneous EEG recordings on the scalp identified the N170 over the right OT scalp region. The latency and amplitude of this scalp N170 were correlated at the single-trial level with the N170 recorded in the lateral IOG, close to the scalp lateral occipital surface. In addition, positive component maximal around the latency of the N170 (a P170) was prominent above the internal LFG, whereas this region typically generates an N170 (or “N200”) over its external/ventral surface. This suggests that electrophysiological responses in the LFG manifest as an equivalent dipole oriented mostly along the vertical axis with likely minimal projection to the lateral OT scalp region. Altogether, these observations provide evidence that the IOG is a major cortical generator of the face-selective scalp N170, qualifying the potential contribution of the FG and questioning a strict serial spatiotemporal organization of the human cortical face network.

Threat-conditioned contexts modulate the late positive potential to faces-A mobile EEG/virtual reality study.

In everyday life, the motivational value of faces is bound to the contexts in which faces are perceived. Electrophysiological studies have demonstrated that inherent negatively valent contexts modulate cortical face processing as assessed with ERP components. However, it is not well understood whether learned (rather than inherent) and three-dimensional aversive contexts similarly modulate the neural processing of faces. Using full immersive virtual reality (VR) and mobile EEG techniques, 25 participants underwent a differential fear conditioning paradigm, in which one virtual room was paired with an aversive noise burst (threat context) and another with a nonaversive noise burst (safe context). Subsequently, avatars with neutral or angry facial expressions were presented in the threat and safe contexts while EEG was recorded. Analysis of the late positive potential (LPP), which presumably indicates motivational salience, revealed a significant interaction of context (threat vs. safe) and face type (neutral vs. angry). Neutral faces evoked increased LPP amplitudes in threat versus safe contexts, while angry faces evoked increased early LPP amplitudes regardless of context. In addition to indicating that threat-conditioned contexts alter the processing of ambiguous faces, the present study demonstrates the successful integration of EEG and VR with particular relevance for affective neuroscience research.

What can we learn about human individual face recognition from experimental studies in monkeys?

Typical human adults recognize numerous individuals from their faces accurately, rapidly and automatically, reaching a level of expertise at individual face recognition that is important for the quality of their social interactions. A non-human species of primates, the rhesus monkey, has been used for decades as a model of human face processing, in particular for understanding the neural basis of individual face recognition. However, despite responding specifically to faces behaviourally and neurally, this species, as well as other Old World and New World monkeys, is remarkably poor at individuating faces of conspecifics. Following extensive conditioning, monkeys only achieve moderate performance at individual face matching tasks where image-based cues are available. Contrary to humans, monkeys do not show a systematic inversion effect in such tasks, or an advantage for matching face pictures of familiar versus unfamiliar individuals, indicating that they do not rely on qualitatively similar individual face recognition processes as humans. These observations concur with the characteristics of the rhesus monkey cortical face processing system, which lacks two critical aspects for human expertise at individual face recognition: a distinct ventral face-selective pathway and a right hemispheric specialization. While the rhesus monkey brain is undoubtedly an informative non-human model for studying the neural basis of social behaviour and visual cognition, it does not provide an adequate model of human individual face recognition. More generally, this review urges for caution when drawing direct inferences across species without sufficient homologies in behaviour and anatomico-functional landmarks.

A face is more than just the eyes, nose, and mouth: fMRI evidence that face-selective cortex represents external features

What is a face? Intuition, along with abundant behavioral and neural evidence, indicates that internal features (e.g., eyes, nose, mouth) are critical for face recognition, yet some behavioral and neural findings suggest that external features (e.g., hair, head outline, neck and shoulders) may likewise be processed as a face. Here we directly test this hypothesis by investigating how external (and internal) features are represented in the brain. Using fMRI, we found highly selective responses to external features (relative to objects and scenes) within the face processing system in particular, rivaling that observed for internal features. We then further asked how external and internal features are represented in regions of the cortical face processing system, and found a similar division of labor for both kinds of features, with the occipital face area and posterior superior temporal sulcus representing the parts of both internal and external features, and the fusiform face area representing the coherent arrangement of both internal and external features. Taken together, these results provide strong neural evidence that a “face” is composed of both internal and external features.

The Fusiform Face Area Plays a Greater Role in Holistic Processing for Own-Race Faces Than Other-Race Faces.

Own-race faces are recognized more effectively than other-race faces. This phenomenon is referred to as other-race effect (ORE). Existing behavioral evidence suggests that one of the possible causes of ORE is that own-race faces are processed more holistically than other-race faces. However, little is known about whether such differences in processing also produce distinctive neural responses in the cortical face processing network. To bridge this gap, the present study used fMRI methodology and the composite face paradigm to examine the response patterns of the traditional face-preferential cortical areas (i.e., the bilateral fusiform face areas [FFA] and the bilateral occipital face areas [OFA]) elicited by own-race faces and other-race faces. We found that the right FFA exhibited a neural composite face effect only for own-race faces but not for other-race faces, even with the absence of the race-related difference in behavior composite face effect. These findings suggest that the right FFA plays a greater role in holistic processing of individual own-race faces than other-race faces. They also suggest that the neural composite effect observed in the right FFA is not the exact neural counterpart of the behavioral face composite effect. The findings of the present study revealed that, along the pathway of the bottom-up face processing, own-race faces and other-race faces presented the holistic processing difference as early as when they were processed in the right FFA.

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.

Mesial temporal lobe epilepsy diminishes functional connectivity during emotion perception

ObjectivesUnilateral mesial temporal lobe epilepsy (MTLE) has been associated with impaired recognition of emotional facial expressions. Correspondingly, imaging studies showed decreased activity of the amygdala and cortical face processing regions in response to emotional faces. However, functional connectivity among regions involved in emotion perception has not been studied so far. MethodsTo address this, we examined intrinsic functional connectivity (FC) modulated by the perception of dynamic fearful faces among the amygdala and limbic, frontal, temporal and brainstem regions. Regions of interest were identified in an activation analysis by presenting a block-design with dynamic fearful faces and dynamic landscapes to 15 healthy individuals. This led to 10 predominately right-hemispheric regions. Functional connectivity between these regions during the perception of fearful faces was examined in drug-refractory patients with left- (n=16) or right-sided (n=17) MTLE, epilepsy patients with extratemporal seizure onset (n=15) and a second group of 15 healthy controls. ResultsHealthy controls showed a widespread functional network modulated by the perception of fearful faces that encompassed bilateral amygdalae, limbic, cortical, subcortical and brainstem regions. In patients with left MTLE, a downsized network of frontal and temporal regions centered on the right amygdala was present. Patients with right MTLE showed almost no significant functional connectivity. A maintained network in the epilepsy control group indicates that findings in mesial temporal lobe epilepsy could not be explained by clinical factors such as seizures and antiepileptic medication. ConclusionFunctional networks underlying facial emotion perception are considerably changed in left and right MTLE. Alterations are present for both hemispheres in either MTLE group, but are more pronounced in right MTLE. Disruption of the functional network architecture possibly contributes to deficits in facial emotion recognition frequently reported in MTLE.

The cognitive and neural basis of developmental prosopagnosia.

Developmental prosopagnosia (DP) is a severe impairment of visual face recognition in the absence of any apparent brain damage. The factors responsible for DP have not yet been fully identified. This article provides a selective review of recent studies investigating cognitive and neural processes that may contribute to the face recognition deficits in DP, focusing primarily on event-related brain potential (ERP) measures of face perception and recognition. Studies that measured the face-sensitive N170 component as a marker of perceptual face processing have shown that the perceptual discrimination between faces and non-face objects is intact in DP. Other N170 studies suggest that faces are not represented in the typical fashion in DP. Individuals with DP appear to have specific difficulties in processing spatial and contrast deviations from canonical upright visual-perceptual face templates. The rapid detection of emotional facial expressions appears to be unaffected in DP. ERP studies of the activation of visual memory for individual faces and of the explicit identification of particular individuals have revealed differences between DPs and controls in the timing of these processes and in the links between visual face memory and explicit face recognition. These observations suggest that the speed and efficiency of information propagation through the cortical face network is altered in DP. The nature of the perceptual impairments in DP suggests that atypical visual experience with the eye region of faces over development may be an important contributing factor to DP.