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Evoked potential evidence for human brain mechanisms that respond to single, fixated faces.

The influence of visual fixation position and stimulus size on the scalp-recorded "vertex positive peak" (VPP) evoked by images of faces was studied in three subjects. Responses were recorded, in turn, for line-drawn, frontal-view faces of approximately 8, 4, 2, and 1 deg length, fixated at the centre (bridge of the nose), and at points 1, 2, 3, and 4 deg to the left and right, and above and below, centre. The results showed that central fixation produced VPPs of similar, maximal amplitude for all face sizes. By comparison, "on-face" eccentric viewing yielded attenuated and delayed responses, and the degree of response attenuation as a function of eccentricity was directly related to the face size, with similar amplitude responses being evoked for corresponding fixation locations on each face. Very small or no VPPs were recorded for most "off-face" fixations. Similar results were observed for profile faces, except that the maximal VPP was recorded for fixations near the eyes and not in the centre of the head, and almost identical VPPs were evoked by a centrally fixated face presented with and without an adjacent face or object. These response properties, which correspond to the subjective perception of the facial stimuli, suggest that the VPP reflects brain mechanisms optimized to respond to single, fixated faces, irrespective both of facial image size and of the presence of neighbouring figures.

Adult↗

The role of long-term and short-term familiarity in visual and haptic face recognition.

Recent studies have suggested that the familiarity of a face leads to more robust recognition, at least within the visual domain. The aim of our study was to investigate whether face familiarity resulted in a representation of faces that was easily shared across the sensory modalities. In Experiment 1, we tested whether haptic recognition of a highly familiar face (one's own face) was as efficient as visual recognition. Our observers were unable to recognise their own face models from tactile memory alone but were able to recognise their faces visually. However, haptic recognition improved when participants were primed by their own live face. In Experiment 2, we found that short-term familiarisation with a set of previously unfamiliar face stimuli improved crossmodal recognition relative to the recognition of unfamiliar faces. Our findings suggest that familiarisation provides a strong representation of faces but that the nature of the information encoded during learning is critical for efficient crossmodal recognition.

Adult↗

Cortical responses to invisible faces: dissociating subsystems for facial-information processing.

Perceiving faces is critical for social interaction. Evidence suggests that different neural pathways may be responsible for processing face identity and expression information. By using functional magnetic resonance imaging (fMRI), we measured brain responses when observers viewed neutral, fearful, and scrambled faces, either visible or rendered invisible through interocular suppression. The right fusiform face area (FFA), the right superior temporal sulcus (STS), and the amygdala responded strongly to visible faces. However, when face images became invisible, activity in FFA to both neutral and fearful faces was much reduced, although still measurable; activity in the STS was robust only to invisible fearful faces but not to neutral faces. Activity in the amygdala was equally strong in both the visible and invisible conditions to fearful faces but much weaker in the invisible condition for the neutral faces. In the invisible condition, amygdala activity was highly correlated with that of the STS but not with FFA. The results in the invisible condition support the existence of dissociable neural systems specialized for processing facial identity and expression information. When images are invisible, cortical responses may reflect primarily feed-forward visual-information processing and thus allow us to reveal the distinct functions of FFA and STS.

Amygdala↗

Activation of the fusiform gyrus when individuals with autism spectrum disorder view faces.

Prior imaging studies have failed to show activation of the fusiform gyrus in response to emotionally neutral faces in individuals with autism spectrum disorder (ASD) [Critchley et al., Brain 124 (2001) 2059; Schultz et al., Arch. Gen. Psychiatry 57 (2000) 331]. However, individuals with ASD do not typically exhibit the striking behavioral deficits that might be expected to result from fusiform gyrus damage, such as those seen in prosopagnosia, and their deficits appear to extend well beyond face identification to include a wide range of impairments in social perceptual processing. In this study, our goal was to further assess the question of whether individuals with ASD have abnormal fusiform gyrus activation to faces. We used high-field (3 T) functional magnetic resonance imaging to study face perception in 11 adult individuals with autism spectrum disorder (ASD) and 10 normal controls. We used face stimuli, object stimuli, and sensory control stimuli (Fourier scrambled versions of the face and object stimuli) containing a fixation point in the center to ensure that participants were looking at and attending to the images as they were presented. We found that individuals with ASD activated the fusiform face area and other brain areas normally involved in face processing when they viewed faces as compared to non-face stimuli. These data indicate that the face-processing deficits encountered in ASD are not due to a simple dysfunction of the fusiform area, but to more complex anomalies in the distributed network of brain areas involved in social perception and cognition.

Adult↗

The fusiform face area is tuned for curvilinear patterns with more high-contrasted elements in the upper part.

The ability to identify conspecifics from the face is of primary interest for human social behavior. Newborns' visual preference for schematic face-like stimuli has been recently related to a sensitivity for visual patterns with a greater number of elements in the upper compared to the lower part. At the adult level, neuroimaging studies have identified a network of cortical areas devoted to the detection and identification of faces. However, whether and how low-level structural properties of face stimuli contribute to the preferential response to faces in these areas remain to be clarified. Using functional magnetic resonance imaging (fMRI), here we investigated whether the adults' face-sensitive cortical areas show a preference for top-heavy patterns, similarly to newborns' preference. Twelve participants were presented with head-shaped and square patterns with either more elements in the upper or the lower vertical part. In the right fusiform gyrus ('fusiform face area', FFA), an area showing a preference for faces over other visual object categories, there was a larger activation for curvilinear patterns with more high-contrast elements in the upper part, even though these patterns were not perceived as face stimuli. These findings provide direct evidence that the FFA is tuned for geometrical properties fitting best with the structure of faces, a computational mechanism that might drive the automatic detection of faces in the visual world.

Adult↗

Faces are represented holistically in the human occipito-temporal cortex.

Two identical top parts of a face photograph look different if their bottom parts differ. This perceptual illusion, the "face composite effect", is taken as strong evidence that faces are processed as a whole rather than as a collection of independent features. To test the hypothesis that areas responding preferentially to faces in the human brain represent faces holistically, we recorded functional magnetic resonance imaging (fMRI) during an adaptation paradigm with the composite face illusion. In both the middle fusiform gyrus (MFG) and the inferior occipital gyrus (IOG), we observed a significantly larger response to the same top face when it was aligned with different bottom parts than with the same bottom part, with a most robust effect in the right middle fusiform gyrus. This difference was not found when the top and the bottom face parts were spatially misaligned or when the faces were presented upside-down. These findings indicate that facial features are integrated into holistic face representations in areas of the human visual cortex responding preferentially to faces.

Adult↗

Cortical correlates of face and scene inversion: a comparison.

Face recognition is more strongly impaired by stimulus inversion than nonface object recognition. This phenomenon, known as the face inversion effect (FIE), suggests that the visual system contains specialized processing mechanisms that are more engaged by upright faces than by inverted faces or nonface objects. Neuroimaging and neuropsychological studies indicate that environmental scenes may also recruit specialized-purpose processing machinery but a comparable inversion effect for scenes has not been established. Here we demonstrate that both face and scene inversion lead to behavioral penalties during performance of a continuous visual matching task; however, the scene inversion effect was less robust and declined in magnitude over the course of the experiment. Scene inversion led to greater neural response in the functionally defined lateral occipital (LO) object area for inverted versus upright scenes and reduced response in the parahippocampal place area (PPA), while face inversion lead to greater response in LO and the right middle fusiform (MF) object area for inverted versus upright faces but no change in the fusiform face area (FFA). A whole-brain analysis revealed several regions that responded more strongly to either upright versus inverted faces or upright versus inverted scenes, some of which may be involved in post-recognition processing. These results demonstrate that both face and scene inversion cause a shift from specialized processing streams towards generic object-processing mechanisms; however, this shift only leads to a reliable behavioral penalty in the case of face inversion.

Adolescent↗

Occipitotemporal activation evoked by the perception of human bodies is modulated by the presence or absence of the face.

We investigated the degree to which activation in regions of the brain known to participate in social perception is influenced by the presence or absence of the face and other body parts. Subjects continuously viewed a static image of a lecture hall in which actors appeared briefly in various poses. There were three conditions: Body-Face, in which the actor appeared with limbs, torso, and face clearly visible; Body-Only, in which the actor appeared with his or her face occluded by a book; and Face-Only, in which the actor appeared behind a podium with only face and shoulders visible. Using event-related functional MRI, we obtained strong activation in those regions previously identified as important for face and body perception. These included portions of the fusiform (FFG) and lingual gyri within ventral occipitotemporal cortex (VOTC), and portions of the middle occipital gyrus (corresponding to the previously defined extrastriate body area, or EBA) and posterior superior temporal sulcus (pSTS) within lateral occipitotemporal cortex (LOTC). Activation of the EBA was strongest for the Body-Only condition; indeed, exposing the face decreased EBA activation evoked by the body. In marked contrast, activation in the pSTS was largest when the face was visible, regardless of whether the body was also visible. Activity within the lateral lingual gyrus and adjacent medial FFG was strongest for the Body-Only condition, while activation in the lateral FFG was greatest when both the face and body were visible. These results provide new information regarding the importance of a visible face in both the relative activation and deactivation of brain structures engaged in social perception.

Adult↗

At first sight: a high-level pop out effect for faces.

To determine the nature of face perception, several studies used the visual search paradigm, whereby subjects detect an odd target among distractors. When detection reaction time is set-size independent, the odd element is said to "pop out", reflecting a basic mechanism or map for the relevant feature. A number of previous studies suggested that schematic faces do not pop out. We show that natural face stimuli do pop out among assorted non-face objects. Animal faces, on the other hand, do not pop out from among the same assorted non-face objects. In addition, search for a face among distractors of another object category is easier than the reverse search, and face search is mediated by holistic face characteristics, rather than by face parts. Our results indicate that the association of pop out with elementary features and lower cortical areas may be incorrect. Instead, face search, and indeed all feature search, may reflect high-level activity with generalization over spatial and other property details.

Animals↗

On second glance: still no high-level pop-out effect for faces.

A recent article in this journal (Hershler, O., & Hochstein, S. (2005). At first sight: A high-level pop out effect for faces. Vision Research, 45(13), 1707-1724) reported, in contradiction to several earlier studies, that photographs of human faces can be searched for efficiently (i.e., "pop out") among photographs of other objects (as long as these objects are not "too similar" to faces). An apparent search asymmetry between faces and other categories (houses, cars) pointed to the existence of a specialized "face map". Findings of impaired performance for scrambled images were presented as evidence that this face pop out is a high-level, "holistic" effect. While the main pop-out effect cannot be disputed, several choices made in that study in terms of experiment design, analysis and interpretation are questionable. After discussing these issues, I report novel experiments which show that (i) the face pop-out effect can be replicated, but under controlled conditions there is no asymmetry between faces and other objects (cars); (ii) inverting pictures and hence disrupting holistic face processing has only a minor effect on search performance; (iii) finally, search becomes inefficient when Fourier amplitude information (which carries global low-level statistical properties of images) is made irrelevant, and only phase information (carrying contour localization) can be used to detect faces. These results imply, contrary to the target article, that the face pop-out effect is mostly based on low-level factors.

Adult↗

Beyond localisation: a dynamical dual route account of face recognition.

After decades of research the notion that faces are special is still at the heart of heated debates. New techniques like brain imaging have advanced some of the arguments but empirical data from brain-damaged patients like paradoxical recognition effects have required more complex explanations aside from localisation of the face area in normal adults. In this paper we focus on configural face processes and discuss configural processes in prosopagnosics in the light of findings obtained in brain-imaging studies. In order to account for data like paradoxical face recognition effects we propose a dual route model of face recognition. The model is based on the distinction between two separate aspects of face recognition, detection and identification, considered as dynamical and interrelated. In this perspective the face detection system appears as the stronger candidate for face-specific processes. The face identification system on the other hand is part of the object recognition system but derives its specificity in part from interaction with the face-specific detection system. The fact that face detection appears intact in some patients provides us with a possible explanation for the interference of configural processes on feature-based identification.

Brain↗

Impaired discrimination of familiar from unfamiliar faces.

We report investigations of the face processing abilities of J.T., a man who had suffered a right hemisphere stroke. J.T. showed a marked problem in discriminating between familiar and unfamiliar faces, with no corresponding problem in discriminating familiar from unfamiliar names. The problem with faces was still found 2 years after the stroke, but had resolved at long-term follow-up (9 years post stroke). When given unlimited time to respond, J.T. did not show any problem in recognising familiar faces, but tended to think that he recognised unfamiliar faces. When under time pressure, however, J.T. also failed to recognise a number of familiar faces which he could readily identify when the time pressure was removed. J.T.'s ability to remember a face and to think of other people who might be similar in appearance was affected by whether or not the face seemed familiar or unfamiliar to him. Hence, whatever underlay the spurious sense of familiarity was sufficient to produce real differences between the way in which genuinely unfamiliar and spuriously familiar faces were seen, leading us to suggest that his impaired discrimination of unfamiliar from familiar faces reflected a malfunction of face recognition units.

Aged↗

Recognition and identification of famous faces in patients with unilateral temporal lobe epilepsy.

We examined the performance of 21 patients with unilateral temporal lobe epilepsy (TLE) and hippocampal damage (10 lefts, and 11 rights) and 10 age-matched controls on the recognition and identification (name and occupation) of well-known faces. Famous face stimuli were selected from four time periods; 1970s, 1980s, 1990-1994, and 1995-1996. Differential patterns of performance were observed for the left and right TLE group across distinct face processing components. The left TLE group showed a selective impairment in naming famous faces while they performed similar to the controls in face recognition and semantic identification (i.e. occupation). In contrast, the right TLE group was impaired across all components of face memory; face recognition, semantic identification, and face naming. Face naming impairment in the left TLE group was characterized by a temporal gradient with better naming performance for famous faces from more distant time periods. Findings are discussed in terms of the role of the temporal lobe system for the acquisition, retention, and retrieval of face semantic networks, and the differential effects of lateralized temporal lobe lesions in this process.

Adult↗

Face processing in 8-month-old infants: evidence for configural and analytical processing.

Two experiments examined whether 8-month-old infants process faces (photos in Experiment 1, schematic faces in Experiment 2) analytically by processing facial features independently of the facial context or configurally by processing the features in conjunction with the facial context. Infants were habituated to two faces and looking time was measured. After habituation they were tested with a habituation face, a switch face, or a novel face. In the switch faces, single features of the habituation faces were switched. The results showed that the infants processed facial features of photographs of faces configurally whereas they processed features of schematic faces (eyes, nose, facial contour) analytically. Thus, although infants have access to both processing modes, for real looking faces they use the configural mode.

Discrimination, Psychological↗

Visual scanpaths in schizophrenia: is there a deficit in face recognition?

There is substantial evidence that schizophrenics have deficits in face processing. We hypothesised that this difficulty is due to abnormalities in the visual scanning of faces. The specificity of these abnormalities to recognisable faces, the effect of task difficulty and their relationships to three primary symptom dimensions were examined. An infrared corneal reflection technique was used to record the visual scanpaths of 63 schizophrenics and 60 non-psychiatric controls while viewing non-degraded ('recognisable') and degraded ('not recognisable') faces. In the concurrent recognition task, subjects were asked to select the previously viewed face from among seven (exposure 1) or three (exposure 2) options. Both groups were unable to accurately recognise degraded faces, but schizophrenics were less accurate than controls for non-degraded faces in the more difficult task condition. Schizophrenics maintained a relatively 'restricted' scanpath style across both faces, but scanpath disturbances were most apparent for non-degraded faces. Analysis of fixation distribution to non-degraded faces showed that, unlike controls, schizophrenics did not concentrate their fixations on salient features. Scanpath aberrations showed only minimal associations with symptom dimensions. These results suggest that schizophrenic individuals have a specific deficit in the visual scanning of faces, that is a distinct case of a fundamental problem in complex object processing.

Adult↗

Repetition priming for familiar and unfamiliar faces in a sex-judgment task: evidence for a common route for the processing of sex and identity.

Repetition priming for faces was examined in a sex-judgment task given at test. Priming was found for edited, hair-removed photos of unfamiliar and familiar faces after a single presentation at study. Priming was also observed for the edited photos when study and test faces were different exemplars. Priming was not observed, however, when sex judgments were made at test to photos of complete, hair-included faces. These findings were interpreted by assuming that, for edited faces, internal features are attended, thereby activating face-recognition units that support performance. With complete faces, however, participants provided speeded judgments based primarily on the hairstyle. It is suggested that, for both familiar and unfamiliar faces, a common locus exists for the processing of the identity of a face and its sex. A single face-recognition model for the processing of familiar and unfamiliar faces is advocated.

Adult↗

When memory shifts toward more typical category exemplars: accentuation effects in the recollection of ethnically ambiguous faces.

In 4 studies, the authors examined the impact of categorization on the recollection of ethnically ambiguous faces. Participants were presented with faces lying at various locations on mixed-race continua (i.e., Caucasian-North African and Caucasian-Asian faces were used as source images in a morphing program). In all studies, the prevalence of exclusive ethnic features in a face distorted participants' recollections of the face toward faces more typical of the category. Specifically, the recollection of 30% North African (or 30% Asian) faces shifted toward Caucasian source faces, whereas the recollection of 70% North African (or 70% Asian) faces shifted toward North African (Asian) source faces. Memory distortions did not emerge for extremely ambiguous (50%) faces and proved larger on mixed-race than same-race continua (Studies 3 and 4). Memory distortions also emerged with high levels of confidence. The authors elaborate on the theoretical and practical implications of these findings.

Adult↗

Microstimulation of inferotemporal cortex influences face categorization.

The inferior temporal cortex (IT) of primates is thought to be the final visual area in the ventral stream of cortical areas responsible for object recognition. Consistent with this hypothesis, single IT neurons respond selectively to highly complex visual stimuli such as faces. However, a direct causal link between the activity of face-selective neurons and face perception has not been demonstrated. In the present study of macaque monkeys, we artificially activated small clusters of IT neurons by means of electrical microstimulation while the monkeys performed a categorization task, judging whether noisy visual images belonged to 'face' or 'non-face' categories. Here we show that microstimulation of face-selective sites, but not other sites, strongly biased the monkeys' decisions towards the face category. The magnitude of the effect depended upon the degree of face selectivity of the stimulation site, the size of the stimulated cluster of face-selective neurons, and the exact timing of microstimulation. Our results establish a causal relationship between the activity of face-selective neurons and face perception.

Aging↗