Familiarity facilitates the corticocortical processing of face perception

The visual evoked potentials elicited by mosaic pictures were used to elucidate the initial step of face perception. Three different mosaic levels (subthreshold, threshold, suprathreshold) for familiar and unfamiliar faces and objects were randomly presented for 250 ms. The latencies of occipital N1 and posterior-temporal N2 were shortened by decreasing the mosaic levels of faces but not for object. The N2 amplitude significantly increased at threshold and suprathreshold levels for familiar and unfamiliar faces. The latency difference between N1 and N2 at threshold level for a familiar face was significantly shortened compared with that for an unfamiliar face. Our findings suggest the initial step of face perception is already set in the primary visual cortex, and familiarity can facilitate the corticocortical processing of face information.     

Hemispheric asymmetries in image-specific and abstractive priming of famous faces: evidence from reaction times and event-related brain potentials

We investigated hemispheric differences in image-specific and abstractive immediate repetition priming of famous faces. Participants performed speeded familiarity decisions for centrally presented famous and unfamiliar target faces. Target faces were preceded by lateralized primes (150 ms), presented either in the left or right visual field (LVF or RVF). Primes were either an identical photograph of the famous target face (image-specific priming), a different image of the famous target face (abstractive priming) or a different familiar face (unprimed condition). Reaction times (RTs) revealed significant effects of priming for both image-specific and abstractive priming overall. In addition, image-specific priming was more than twice the magnitude for targets following LVF primes as compared to RVF primes. By contrast, no hemispheric differences emerged for abstractive face priming across different images. Whereas ERPs revealed no evidence that priming affected the N170 component, both image-specific and abstractive priming significantly modulated the amplitudes of a right temporal N250r and a parietal N400 component. Behavioural and electrophysiological evidence for hemispheric differences in image-specific and abstractive face priming are discussed with respect to current theories of how the human left and right ventral temporal cortices represent abstractive and form-specific visual information.     

Early visual ERP sensitivity to the species and animacy of faces

Assessing the agency of potential actors in the visual world is a critically important aspect of social cognition. Adult observers are generally capable of distinguishing real faces from artificial faces (even allowing for recent advances in graphics technology and motion capture); even small deviations from real facial appearance can lead to profound effects on face recognition. Presently, we examined how early components of visual event-related potentials (ERPs) are affected by the “life” in human faces and animal faces. We presented participants with real and artificial faces of humans and dogs, and analyzed the response properties of the P100 and the N170 as a function of stimulus appearance and task (species categorization vs. animacy categorization). The P100 exhibited sensitivity to face species and animacy. We found that the N170’s differential responses to human faces vs. dog faces depended on the task participants’ performed. Also, the effect of species was only evident for real faces of humans and dogs, failing to obtain with artificial faces. These results suggest that face animacy does modulate early components of visual ERPs—the N170 is not merely a crude face detector, but reflects the tuning of the visual system to natural face appearance.     

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.     

Electrophysiological correlates of improved short-term memory for emotional faces

Long-term memory (LTM) is enhanced for emotional information, but the influence of stimulus emotionality on short-term memory (STM) is less clear. We examined the electrophysiological correlates of improved visual STM for emotional face identity, focusing on the P1, N170, P3b and N250r event-related potential (ERP) components. These correlates are taken to indicate which memory processing stages and cognitive processes contribute to the improved STM for emotional face identity. In the encoding phase, one or three angry, happy or neutral faces were presented for 2 s, resulting in a memory load of one or three. The subsequent 1-s retention phase was followed by a 2-s retrieval phase, in which participants indicated whether a probe face had been present or not during encoding. Memory performance was superior for angry and happy faces over neutral faces at load three. None of the ERP components during encoding were affected by facial expression. During retrieval, the early P3b was decreased for emotional compared to neutral faces, which presumably reflects greater resource allocation to the maintenance of the emotional faces. Furthermore, the N250r during retrieval was increased for emotional compared to neutral faces, reflecting an enhanced repetition effect for emotional faces. These findings suggest that enhanced visual STM for emotional faces arises from improved maintenance and from improved detection of face repetition at retrieval.     

Different responses to same stimuli

Identical comparison stimuli were induced to be recognized as schematic faces or the figures with three English letters 'O' (EngO) by verbal bias and two different contextual priming conditions. Our findings confirmed that there might be a coarse visual categorization around 40-100 ms at mid-parietal site. Larger P1 responses to schematic faces compared with EngO definitely revealed that P1 at occipito-temporal sites should be thought to represent an earlier face-specific processing stage, as early as around 100-120 ms after stimulus onset rather than simply reflect the low-level visual difference. A significant difference of N170/vertex positive potential between the identical comparison stimuli reinforced the evidence that they were early markers of face processing and corroborated the unequivocal influence of contextual effect.     

Event-related potentials reveal temporal staging of dynamic facial expression and gaze shift effects on attentional orienting

Abstract

Multiple sources of information from the face guide attention during social interaction. The present study modified the Posner cueing paradigm to investigate how dynamic changes in emotional expression and eye gaze in faces affect the neural processing of subsequent target stimuli. Event-related potentials (ERPs) were recorded while participants viewed centrally presented face displays in which gaze direction (left, direct, right) and facial expression (fearful, neutral) covaried in a fully crossed design. Gaze direction was not predictive of peripheral target location. ERP analysis revealed several sequential effects, including: (1) an early enhancement of target processing following fearful faces (P1); (2) an interaction between expression and gaze (N1), with enhanced target processing following fearful faces with rightward gaze; and (3) an interaction between gaze and target location (P3), with enhanced processing for invalidly cued left visual field targets. Behaviorally, participants responded faster to targets following fearful faces and targets presented in the right visual field, in concordance with the P1 and N1 effects, respectively. The findings indicate that two nonverbal social cues—facial expression and gaze direction—modulate attentional orienting across different temporal stages of processing. Results have implications for understanding the mental chronometry of shared attention and social referencing.     

Structural encoding of human and schematic faces: holistic and part-based processes

The range of specificity and the response properties of the extrastriate face area were investigated by comparing the N170 event-related potential (ERP) component elicited by photographs of natural faces, realistically painted portraits, sketches of faces, schematic faces, and by nonface meaningful and meaningless visual stimuli. Results showed that the N170 distinguished between faces and nonface stimuli when the concept of a face was clearly rendered by the visual stimulus, but it did not distinguish among different face types: Even a schematic face made from simple line fragments triggered the N170. However, in a second experiment, inversion seemed to have a different effect on natural faces in which face components were available and on the pure gestalt-based schematic faces: The N170 amplitude was enhanced when natural faces were presented upside down but reduced when schematic faces were inverted. Inversion delayed the N170 peak latency for both natural and schematic faces. Together, these results suggest that early face processing in the human brain is subserved by a multiple-component neural system in which both whole-face configurations and face parts are processed. The relative involvement of the two perceptual processes is probably determined by whether the physiognomic value of the stimuli depends upon holistic configuration, or whether the individual components can be associated with faces even when presented outside the face context.     

N250 ERP correlates of the acquisition of face representations across different images

We used ERPs to investigate neural correlates of face learning. At learning, participants viewed video clips of unfamiliar people, which were presented either with or without voices providing semantic information. In a subsequent face-recognition task (four trial blocks), learned faces were repeated once per block and presented interspersed with novel faces. To disentangle face from image learning, we used different images for face repetitions. Block effects demonstrated that engaging in the face-recognition task modulated ERPs between 170 and 900 msec poststimulus onset for learned and novel faces. In addition, multiple repetitions of different exemplars of learned faces elicited an increased bilateral N250. Source localizations of this N250 for learned faces suggested activity in fusiform gyrus, similar to that found previously for N250r in repetition priming paradigms [Schweinberger, S. R., Pickering, E. C., Jentzsch, I., Burton, A. M., & Kaufmann, J. M. Event-related brain potential evidence for a response of inferior temporal cortex to familiar face repetitions. Cognitive Brain Research, 14, 398-409, 2002]. Multiple repetitions of learned faces also elicited increased central-parietal positivity between 400 and 600 msec and caused a bilateral increase of inferior-temporal negativity (>300 msec) compared with novel faces. Semantic information at learning enhanced recognition rates. Faces that had been learned with semantic information elicited somewhat less negative amplitudes between 700 and 900 msec over left inferior-temporal sites. Overall, the findings demonstrate a role of the temporal N250 ERP in the acquisition of new face representations across different images. They also suggest that, compared with visual presentation alone, additional semantic information at learning facilitates postperceptual processing in recognition but does not facilitate perceptual analysis of learned faces.     

Spatial attention modulates early face processing

It is widely reported that inverting a face dramatically affects its recognition. Previous studies have shown that face inversion increases the amplitude and delays the latency of the face-specific N170 component of the event-related potential (ERP) and also enhances the amplitude of the occipital P1 component (latency 100–132 ms). The present study investigates whether these effects of face inversion can be modulated by visual spatial attention. Participants viewed two streams of visual stimuli, one to the left and one to the right of fixation. One stream consisted of a sequence of alphanumeric characters at 6.67 Hz, and the other stream consisted of a series of upright and inverted images of faces and houses presented in randomized order. The participants’ task was to attend selectively to one or the other of the streams (during different blocks) in order to detect infrequent target stimuli. ERPs elicited by inverted faces showed larger P1 amplitudes compared to upright faces, but only when the faces were attended. In contrast, the N170 amplitude was larger to inverted than to upright faces only when the faces were not attended. The N170 peak latency was delayed to inverted faces regardless of attention condition. These inversion effects were face specific, as similar effects were absent for houses. These results suggest that early stages of face-specific processing can be enhanced by attention, but when faces are not attended the onset of face-specific processing is delayed until the latency range of the N170.     

Working memory load for faces modulates P300, N170, and N250r

We used event-related potential (ERP) methodology to examine neural activity associated with visual working memory (WM) for faces. There were two main goals. First, to extend previous findings of P300 load modulation to WM for faces. Second, to examine whether N170 and N250r are also influenced by WM load. Between one and four unfamiliar faces were simultaneously presented for memory encoding. After a 1-sec delay, a target face appeared, and participants had to judge whether this face was part of the previous face array. P300 amplitude decreased as WM load increased, and this P300 suppression was observed at both encoding and retrieval. WM load was also found to modulate other ERPs. The amplitude of the N170 elicited by the target face decreased with load, and this N170 decrease leveled off at load 2, reflecting the behavioral WM capacity of around two faces. In addition, the N250r, observed as an ERP difference for target faces that were present in the encoding array relative to target faces that were absent, was also reduced for higher WM loads. These findings extend previous work by showing that P300 modulation by WM load also occurs for faces. Furthermore, we show, for the first time, that WM load affects the N250r and the early visual N170 component. This suggests that higher visual areas play an important role in WM for faces.     

Reversal of the face-inversion effect in N170 under unconscious visual processing

Many studies using electroencephalography consistently reported a larger N170 (N1) response in the visual cortices to inverted than upright face images (the face inversion effect in N1, FIE-N1). Here we report this robust effect is diminished and even reversed when face stimuli are processed unconsciously. We measured visual-evoked potentials to neutral faces either visible or rendered invisible by an inter-ocular suppression. In visible condition, we observed a larger N1 to inverted than upright faces, which replicated the traditional FIE-N1. When those faces became invisible, however, neural responses to the inverted faces were greatly reduced compared to visible condition, whereas those to the invisible upright faces were relatively preserved. Consequently, N1 amplitudes were found to be larger in upright, rather than inverted, faces in invisible condition, which was opposite to the traditional FIE-N1 (upright<inverted) in visible condition. Those results highlighted a special mechanism in the brain for the processing of the upright, but not inverted, face (e.g. fusiform face area) that retains vigorous responses even when the face becomes invisible.     

An event-related brain potential study of explicit face recognition

To determine the time course of face recognition and its links to face-sensitive event-related potential (ERP) components, ERPs elicited by faces of famous individuals and ERPs to non-famous control faces were compared in a task that required explicit judgements of facial identity. As expected, the face-selective N170 component was unaffected by the difference between famous and non-famous faces. In contrast, the occipito-temporal N250 component was linked to face recognition, as it was selectively triggered by famous faces. Importantly, this component was present for famous faces that were judged to be definitely known relative to famous faces that just appeared familiar, demonstrating that it is associated with the explicit identification of a particular face. The N250 is likely to reflect early perceptual stages of face recognition where long-term memory traces of familiar faces in ventral visual cortex are activated by matching on-line face representations. Famous faces also triggered a broadly distributed longer-latency positivity (P600f) that showed a left-hemisphere bias and was larger for definitely known faces, suggesting links between this component and name generation. These results show that successful face recognition is predicted by ERP components over face-specific visual areas that emerge within 230 ms after stimulus onset.     

Altered automatic face processing in individuals with high-functioning autism spectrum disorders: Evidence from visual evoked potentials

Individuals with autism spectrum disorders (ASDs) have different automatic responses to faces than typically developing (TD) individuals. We recorded visual evoked potentials (VEPs) in 10 individuals with high-functioning ASD (HFASD) and 10 TD individuals. Visual stimuli consisted of upright and inverted faces (fearful and neutral) and objects presented subliminally in a backward-masking paradigm. In all participants, the occipital N1 (about 100 ms) and P1 (about 120 ms) peaks were major components of the evoked response. We calculated “subliminal face effect (SFE)” scores by subtracting the N1/P1 amplitudes and latencies of the object stimuli from those of the face stimuli. In the TD group, the SFE score for the N1 amplitude was significantly higher for upright fearful faces but not neutral faces, and this score was insignificant when the stimuli were inverted. In contrast, the N1 amplitude of the HFASD subjects did not show this SFE in the upright orientation. There were no significant group differences in SFE scores for P1 amplitude, latency, or N1 latency. Our findings suggest that individuals with HFASD have altered automatic visual processing for emotional faces within the lower level of the visual cortex. This impairment could be a neural component of the disrupted social cognition observed in individuals with HFASD.     

Event‐related potential and eye tracking evidence of the developmental dynamics of face processing

Although the wide neural network and specific processes related to faces have been revealed, the process by which face‐processing ability develops remains unclear. An interest in faces appears early in infancy, and developmental findings to date have suggested a long maturation process of the mechanisms involved in face processing. These developmental changes may be supported by the acquisition of more efficient strategies to process faces (theory of expertise) and by the maturation of the face neural network identified in adults. This study aimed to clarify the link between event‐related potential (ERP) development in response to faces and the behavioral changes in the way faces are scanned throughout childhood. Twenty‐six young children (4–10 years of age) were included in two experimental paradigms, the first exploring ERPs during face processing, the second investigating the visual exploration of faces using an eye‐tracking system. The results confirmed significant age‐related changes in visual ERPs (P1, N170 and P2). Moreover, an increased interest in the eye region and an attentional shift from the mouth to the eyes were also revealed. The proportion of early fixations on the eye region was correlated with N170 and P2 characteristics, highlighting a link between the development of ERPs and gaze behavior. We suggest that these overall developmental dynamics may be sustained by a gradual, experience‐dependent specialization in face processing (i.e. acquisition of face expertise), which produces a more automatic and efficient network associated with effortless identification of faces, and allows the emergence of human‐specific social and communication skills.     

Time course of spatial and feature selective attention for partly-occluded objects

Attention selects objects/groups as the most fundamental units, and this may be achieved by an attention-spreading mechanism. Previous event-related potential (ERP) studies have found that attention-spreading is reflected by a decrease in the N1 spatial attention effect. The present study tested whether the electrophysiological attention effect is associated with the perception of object unity or amodal completion through the use of partly-occluded objects. ERPs were recorded in 14 participants who were required to pay attention to their left or right visual field and to press a button for a target shape in the attended field. Bilateral stimuli were presented rapidly, and were separated, connected, or connected behind an occluder. Behavioral performance in the connected and occluded conditions was worse than that in the separated condition, indicating that attention spread over perceptual object representations after amodal completion. Consistently, the late N1 spatial attention effect (180-220 ms post-stimulus) and the early phase (230-280 ms) of feature selection effects (target N2) at contralateral sites decreased, equally for the occluded and connected conditions, while the attention effect in the early N1 latency (140-180 ms) shifted most positively for the occluded condition. These results suggest that perceptual organization processes for object recognition transiently modulate spatial and feature selection processes in the visual cortex.     

Spatiotemporal analyses of the N170 for human faces, animal faces and objects in natural scenes

We assessed the specificity to human faces of the N170 ERP component in the context of natural scenes. Subjects categorized photographs containing human faces, animal faces and various objects. Spatiotemporal topography analyses were performed on the individual ERP data. ERPs elicited by animal faces were similar to human faces ERPs but with a delayed face activity. In the N170 time window, ERPs to human and animal faces had a different topography compared with object ERPs. Such data suggest that N170 generators might process various stimuli with a coarse facial organization and show the care that must be taken in comparing scalp signal to faces and other objects as they are probably generated, at least partially, by different cortical sources.     

Effects of face inversion on the structural encoding and recognition of faces. Evidence from event-related brain potentials

It was investigated how face inversion affects face-specific components of event-related brain potentials (ERPs) which are assumed to reflect the structural encoding and the recognition of faces. ERPs were recorded to upright and inverted photographs of familiar faces, unfamiliar faces, and houses. In Part I, participants had to detect infrequently presented targets (hands), in Part II, attention was either directed towards or away from the pictorial stimuli. When compared with upright unfamiliar faces, upright familiar faces elicited an enhanced negativity between 300 ms and 450 ms ('N400f') and an enhanced positivity between 450 and 650 ms post-stimulus ('P600f'). It is suggested that these ERP modulations are generated by processes involved in the recognition of faces. Face inversion is known to disrupt face recognition processes. Accordingly, 'N400f' and 'P600f' were generally absent in response to inverted familiar and unfamiliar faces. The face-specific N170 component at lateral posterior electrodes was not affected by face familiarity, indicating that it reflects processing stages prior to face identification. N170 was delayed and enhanced for inverted relative to upright faces. While N170 enhancements were also observed for inverted relative to upright houses, the N170 latency shift caused by stimulus inversion was face-specific. Directing attention away from the faces towards a demanding primary visual task resulted in an N170 delay for inverted as well as for upright faces, suggesting that the time course of structural encoding of faces is affected by attentional factors. These results demonstrate that ERPs can be used as electrophysiological markers of specialised brain processes underlying the structural encoding and subsequent recognition of faces.     

Long-latency ERPs and recognition of facial identity

N400 brain event-related potential (ERP) is a mismatch negativity originally found in response to semantic incongruences of a linguistic nature and is used paradigmatically to investigate memory organization in various domains of information, including that of faces. In the present study, we analyzed different mismatch negativities evoked in N400-like paradigms related to recognition of newly learned faces with or without associated verbal information. ERPs were compared in the following conditions: (1) mismatching features (eyes-eyebrows) using a facial context corresponding to the faces learned without associated verbal information ("pure" intradomain facial processing); (2) mismatching features using a facial context corresponding to the faces learned with associated occupations and proper names ("nonpure" intradomain facial processing); (3) mismatching occupations using a facial context (cross-domain processing); and (4) mismatching names using an occupation context (intradomain verbal processing). Results revealed that mismatching stimuli in the four conditions elicited a mismatch negativity analogous to N400 but with different timing and topographical patterns. The onset of the mismatch negativity occurred earliest in Conditions 1 and 2, followed by Condition 4, and latest in Condition 3. The negativity had the shortest duration in Task 1 and the longest duration in Task 3. Bilateral parietal activity was confirmed in all conditions, in addition to a predominant right posterior temporal localization in Condition 1, a predominant right frontal localization in Condition 2, an occipital localization in Condition 3, and a more widely distributed (although with posterior predominance) localization in Condition 4. These results support the existence of multiple N400, and particularly of a nonlinguistic N400 related to purely visual information, which can be evoked by facial structure processing in the absence of verbal-semantic information.     

Form completion across a hemianopic boundary: behindsight?

Patients with homonymous hemianopia may report the completion of forms that overlap the vertical meridian of their field defects. While previous investigations of "hemianopic completion" have variously attributed to the disorder to inattention, residual vision or unstable fixation, we believe that our investigation has controlled for such potentially confounding factors. We report patient P.O.V. who experienced hemianopic completion in everyday life following a surgical lesion of his left occipital lobe. He showed normal spatial attention and normal spatial orienting: hemianopic completion can therefore occur in the absence of inattention. His completion was retinotopic and affected partial as well as complete forms: his hemianopic completion cannot be attributed to residual visual input or poor fixation. P.O.V.'s completion was also systematically affected by varying stimulus contrast and pattern masking. We argue that while other explanations may be appropriate for different cases, P.O.V.'s hemianopic completion reflects normal "constructive" visual processes and can be attributed to the unconstrained operation of visual routines that are normally involved in the perception of partially occluded forms. As such, this disorder has the potential to shed light on some of the most basic aspects of visual perception.