Marc Crommelinck

The N170 occipito-temporal component is delayed and enhanced to inverted faces but not to inverted objects: an electrophysiological account of face-specific processes in the human brain.

Behavioral studies have shown that picture-plane inversion impacts face and object recognition differently, thereby suggesting face-specific processing mechanisms in the human brain. Here we used event-related potentials to investigate the time course of this behavioral inversion effect in both faces and novel objects. ERPs were recorded for 14 subjects presented with upright and inverted visual categories, including human faces and novel objects (Greebles). A N170 was obtained for all categories of stimuli, including Greebles. However, only inverted faces delayed and enhanced N170 (bilaterally). These observations indicate that the N170 is not specific to faces, as has been previously claimed. In addition, the amplitude difference between faces and objects does not reflect face-specific mechanisms since it can be smaller than between non-face object categories. There do exist some early differences in the time-course of categorization for faces and non-faces across inversion. This may be attributed either to stimulus category per se (e.g. face-specific mechanisms) or to differences in the level of expertise between these categories.

Spatio-temporal localization of the face inversion effect : an event-related potentials study

Event-related potentials (ERPs) from 58 electrodes at standard EEG sites were recorded while 14 subjects performed a delayed-matching task on normal and inverted faces. A large and single difference between normal and inverted face processing was observed at occipito-temporal sites about 160 ms following stimulus onset, mainly in the right hemisphere (RH). Although the topographies indicate that similar areas are involved at this latency in processing the two types of stimuli, the electrophysiological activity, which corresponds to the previously described N170, was larger and delayed for inverted as compared to normal face processing. These results complement and specify, at a neural level, previous behavioral and divided visual field studies which have suggested that the loss of configural face information by inversion may slow down and increase the difficulty of face processing, particularly in the RH.

Hemispheric Asymmetries for Whole-Based and Part-Based Face Processing in the Human Fusiform Gyrus

Behavioral studies indicate a right hemisphere advantage for processing a face as a whole and a left hemisphere superiority for processing based on face features. The present PET study identifies the anatomical localization of these effects in well-defined regions of the middle fusiform gyri of both hemispheres. The right middle fusiform gyrus, previously described as a face-specific region, was found to be more activated when matching whole faces than face parts whereas this pattern of activity was reversed in the left homologous region. These lateralized differences appeared to be specific to faces since control objects processed either as wholes or parts did not induce any change of activity within these regions. This double dissociation between two modes of face processing brings new evidence regarding the lateralized localization of face individualization mechanisms in the human brain.

Expertise Training with Novel Objects Leads to Left-Lateralized Facelike Electrophysiological Responses

Scalp event-related potentials (ERPs) in humans indicate that face and object processing differ approximately 170 ms following stimulus presentation, at the point of the N170 occipitotemporal component. The N170 is delayed and enhanced to inverted faces but not to inverted objects. We tested whether this inversion effect reflects early mechanisms exclusive to faces or whether it generalizes to other stimuli as a function of visual expertise. ERPs to upright and inverted faces and novel objects (Greebles) were recorded in 10 participants before and after 2 weeks of expertise training with Greebles. The N170 component was observed for both faces and Greebles. The results are consistent with previous reports in that the N170 was delayed and enhanced for inverted faces at recording sites in both hemispheres. For Greebles, the same effect of inversion was observed only for experts, primarily in the left hemisphere. These results suggest that the mechanisms underlying the electrophysiological face-inversion effect extend to visually homogeneous nonface object categories, at least in the left hemisphere, but only when such mechanisms are recruited by expertise.

Effect of familiarity on the processing of human faces

Most brain imaging studies on face perception have investigated the processing of unknown faces and addressed mainly the question of specific face processing in the human brain. The goal of this study was to highlight the effects of familiarity on the visual processing of faces. Using [15O]water 3D Positron Emission Tomography, regional cerebral blood flow distribution was measured in 11 human subjects performing an identical task (gender categorization) on both unknown and known faces. Subjects also performed two control tasks (a face recognition task and a visual pattern discrimination task). They were scanned after a training phase using videotapes during which they had been familiarized with and learned to recognize a set of faces. Two major results were obtained. On the one hand, we found bilateral activations of the fusiform gyri in the three face conditions, including the so-called fusiform-face area, a region in the right fusiform gyrus specifically devoted to face processing. This common activation suggests that different cognitive tasks performed on known and unknown faces require the involvement of this fusiform region. On the other hand, specific regional cerebral blood flow changes were related to the processing of known and unknown faces. The left amygdala, a structure involved in implicit learning of visual representations, was activated by the categorization task on unknown faces. The same task on known faces induced a relative decrease of activity in early visual areas. These differences between the two categorization tasks reveal that the human brain processes known and unknown faces differently.

Stimulation of the superior colliculus in the alert cat

SummaryElectrical stimulation of the cat superior colliculus (SC), in conjunction with the accurate measurement of elicited eye movements and histologically verified electrode positions, has revealed a striking antero-posterior variation in collicular organization. Three zones could be defined in the SC on the basis of eye movement patterns and associated neck muscle EMG activity evoked from the deeper layers. The Anterior zone was coextensive with the central 25 ° of the visual retinotopically coded map contained in the superficial layers. Saccades evoked from this zone were also retinotopically coded, and the latency of EMG activity depended on the position of the eye in the orbit. A similar observation applies to the entire monkey SC. The Intermediate zone was coextensive with the 25 °–70 ° of visual projections. Saccades evoked from this region were “goal-directed” and were associated with invariant, short latency EMG responses. The Posterior zone was found in the extreme caudo-lateral portion of the SC. Eye movements evoked from this zone were centering saccades associated with constant latency EMG activity. The present results in conjunction with previously demonstrated antero-posterior variations in projections to the SC, suggest that the motor strategies controlling gaze shifts toward visual targets vary depending on the location of the target in the visual field.

Task modulation of brain activity related to familiar and unfamiliar face processing : An ERP study

In order to investigate stimulus-related and task-related electrophysiological activity relevant for face processing, event-related potentials (ERPs) from 58 electrodes at standard EEG sites were recorded while subjects performed a simple visual discrimination (control) task, in addition to various face processing tasks: recognition of previously learned faces and gender decision on familiar and unfamiliar faces. Three electrophysiological components or dipolar complex were recorded in all subjects: an occipital early component (P1, around 110 ms); a vertex positive potential (VPP; around 158 ms) which appeared to be specific to faces; and a negative central component, N2 (around 230 ms). Parametric analysis and source localization were applied to these components by means of a single-subject analysis methodology. No effect of familiarity was observed on any of these early component. While the VPP appears to be independent of the kind of processing performed, face task modulations of the early P1 and the N2 were observed, with a higher amplitude for the recognition than for the gender discrimination task. An attentional modulation of early visual areas is proposed for the first effect (P1 modulation), while the N2 seems to be related to general visual memory processing. This study strongly suggests that the VPP reflects an early visual stage of face processing in the fusiform gyrus that is strictly stimulus-related and independent of familiarity. It also shows that source localization algorithms may give reliable solutions on single subject averages for early visual components despite high inter-subject variability of the surface characteristics of ERPs.

Categorical Perception of Happiness and Fear Facial Expressions: An ERP Study

Behavioral studies have shown that two different morphed faces perceived as reflecting the same emotional expression are harder to discriminate than two faces considered as two different ones. This advantage of between-categorical differences compared with within-categorical ones is classically referred as the categorical perception effect. The temporal course of this effect on fear and happiness facial expressions has been explored through event-related potentials (ERPs). Three kinds of pairs were presented in a delayed samedifferent matching task: (1) two different morphed faces perceived as the same emotional expression (within-categorical differences), (2) two other ones reflecting two different emotions (between-categorical differences), and (3) two identical morphed faces (same faces for methodological purpose). Following the second face onset in the pair, the amplitude of the bilateral occipito-temporal negativities (N170) and of the vertex positive potential (P150 or VPP) was reduced for within and same pairs relative to between pairs. This suggests a repetition priming effect. We also observed a modulation of the P3b wave, as the amplitude of the responses for the between pairs was higher than for the within and same pairs. These results indicate that the categorical perception of human facial emotional expressions has a perceptual origin in the bilateral occipito-temporal regions, while typical prior studies found emotion-modulated ERP components considerably later.

The time-course of intermodal binding between seeing and hearing affective information.

Intermodal binding between affective information that is seen as well as heard triggers a mandatory process of audiovisual integration. In order to track the time course of this audiovisual binding, event related brain potentials were recorded while subjects saw facial expression and concurrently heard auditory fragment. The results suggest that the combination of the two inputs is early in time (110 ms post-stimulus) and translates as a specific enhancement in amplitude of the auditory N1 component. These findings are compatible with previous functional neuroimaging results of audiovisual speech showing strong audiovisual interactions in auditory cortex in the form of magnetic response amplifications, as well as with electrophysiological studies demonstrating early audiovisual interactions (before 200 ms post-stimulus). Moreover, our results show that the informational content present in the two modalities plays a crucial role in triggering the intermodal binding process.

Neuronal Mechanisms of Perceptual Learning: Changes in Human Brain Activity with Training in Orientation Discrimination

Using 15O-water 3D positron emission tomography, regional cerebral blood flow was measured twice in six human subjects: before and after extensive training in orientation discrimination. In each session subjects performed two orientation discrimination tasks, during which they discriminated the orientation of a grating at either the trained or untrained reference orientation, and a control task, during which they detected a randomly textured pattern. By comparing the discrimination to the detection tasks, we observed a main effect of task bilaterally in the posterior occipital cortex, extending into the left posterior fusiform gyrus and the right inferior occipital gyrus, bilaterally in the intraparietal sulcus, as well as in the cerebellum, thalamus, and brainstem. When we compared the activation pattern before and after the training period, all the changes observed were activity decreases. The nonspecific changes, which were not related to the orientation used during the training, were situated in the cerebellum and bilaterally in the extrastriate visual cortex. The orientation-specific changes, on the other hand, were restricted to the striate and extrastriate visual cortex, more precisely the right calcarine sulcus, the left lingual gyrus, the left middle occipital, and the right inferior occipital gyrus. These findings confirm our hypothesis concerning the existence of learning related changes at early levels of visual processing in human adults and suggest that mechanisms resulting in neuronal activity decreases might be involved in the present kind of learning.