Shlomo Bentin

Electrophysiological studies of face perception in humans

Event-related potentials (ERPs) associated with face perception were recorded with scalp electrodes from normal volunteers. Subjects performed a visual target detection task in which they mentally counted the number of occurrences of pictorial stimuli from a designated category such as butterflies. In separate experiments, target stimuli were embedded within a series of other stimuli including unfamiliar human faces and isolated face components, inverted faces, distorted faces, animal faces, and other nonface stimuli. Human faces evoked a negative potential at 172 msec (N170), which was absent from the ERPs elicited by other animate and inanimate nonface stimuli. N170 was largest over the posterior temporal scalp and was larger over the right than the left hemisphere. N170 was delayed when faces were presented upside-down, but its amplitude did not change. When presented in isolation, eyes elicited an N170 that was significantly larger than that elicited by whole faces, while noses and lips elicited small negative ERPs about 50 msec later than N170. Distorted human faces, in which the locations of inner face components were altered, elicited an N170 similar in amplitude to that elicited by normal faces. However, faces of animals, human hands, cars, and items of furniture did not evoke N170. N170 may reflect the operation of a neural mechanism tuned to detect (as opposed to identify) human faces, similar to the structural encoder suggested by Bruce and Young (1986). A similar function has been proposed for the face-selective N200 ERP recorded from the middle fusiform and posterior inferior temporal gyri using subdural electrodes in humans (Allison, McCarthy, Nobre, Puce, & Belger, 1994c). However, the differential sensitivity of N170 to eyes in isolation suggests that N170 may reflect the activation of an eye-sensitive region of cortex. The voltage distribution of N170 over the scalp is consistent with a neural generator located in the occipitotemporal sulcus lateral to the fusiform/inferior temporal region that generates N200.

Guidelines for using human event-related potentials to study cognition: recording standards and publication criteria.

Event-related potentials (ERPs) recorded from the human scalp can provide important information about how the human brain normally processes information and about how this processing may go awry in neurological or psychiatric disorders. Scientists using or studying ERPs must strive to overcome the many technical problems that can occur in the recording and analysis of these potentials. The methods and the results of these ERP studies must be published in a way that allows other scientists to understand exactly what was done so that they can, if necessary, replicate the experiments. The data must then be analyzed and presented in a way that allows different studies to be compared readily. This paper presents guidelines for recording ERPs and criteria for publishing the results.

Event-related potentials, lexical decision and semantic priming☆

ERPs were recorded during a lexical decision task in order to investigate electrophysiological concomitants of semantic priming. The stimuli were 240 words and 240 nonwords presented one per trial at a fixed intertrial interval. Subjects were required to classify each stimulus as a word or nonword by pressing one of two response buttons. ERPs were recorded from 14 scalp locations, the right suborbital ridge, and the left earlobe, all referred to a balanced non-cephalic reference. RT and error data confirmed that semantic priming occurred under the conditions employed: primed words (those preceded by a semantically related word) were identified as words faster and more accurately than were unprimed words (those preceded by semantically unrelated words or nonwords). ERPs for all stimulus types were characterized by a large positivity peaking between 550 and 650 msec, preceded by a negative-going deflection peaking at approximately 400 msec. ERPs for primed and unprimed words were shown to differ significantly, diverging 200-250 msec following stimulus onset, reaching a maximum near the peak of the negative-going deflection at 400 msec. These differences were observed at locations over both hemispheres and were maximal in the centroparietal region. Although P300 latency differences between primed and unprimed words were also obtained, the priming effect on ERPs at shorter latencies could not be explained solely by P300 latency effects. Possible relationships between these ERP concomitants of semantic priming and P300, N200, and N400 were discussed.

Structural encoding and identification in face processing: ERP evidence for separate mechanisms

The present study had two aims. The first aim was to explore the possible top-down effect of face-recognition and/or face-identification processes on the formation of structural representation of faces, as indexed by the N170 ERP component. The second aim was to examine possible ERP manifestations of face identification processes as an initial step for assessing their time course and functional neuroanatomy. Identical N170 potentials were elicited by famous and unfamiliar faces in Experiment 1, when both were irrelevant to the task, suggesting that face familiarity does not affect structural encoding processes. Small but significant differences were observed, however, during later-occurring epochs of the ERPs. In Experiment 2 the participants were instructed to count occasionally occurring portraits of famous politicians while rejecting faces of famous people who were not politicians and faces of unfamiliar people. Although an attempt to identify each face was required, no differences were found in the N170 elicited by faces of unfamiliar people and faces of familiar non-politicians. Famous faces, however, elicited a negative potential that was significantly larger than that elicited by unfamiliar faces between about 250 and 500msec from stimulus onset. This negative component was tentatively identified as an N400 analogue elicited by faces. Both the absence of an effect of familiarity on the N170 and the familiarity face-N400 effect were replicated in Experiment 3, in which the participants made speeded button-press responses in each trial, distinguishing among faces of politicians and faces of famous and unfamiliar non-politicians. In addition, ERP components later than the N400 were found to be associated with the speed of the response but not with face familiarity. We concluded that (1) although reflected by the N170, the structural encoding mechanism is not influenced by the face recognition and identification processes, and (2) the negative component modulated by face familiarity is associated with the semantic activity involved in the identification of familiar faces.

Strategies for Visual Word Recognition and Orthographical Depth: A Multilingual Comparison

We investigated the psychological reality ofthe concept of orthographical depth and its influence on visual word recognition by examining naming performance in Hebrew, English, and Serbo-Croatian. We ran three sr of experiments in which we used native speakers and identical experimental methods in each language. Experiment 1 revealed that the lexical status ofthe stimulus (high-frequency words, low-frequency words, and nonwords) significantly affected naming in Hebrew (the deepest of the three orthographies). This effect was only moderate in English and nonsignificant in SerboCroatian (the shallowest of the three orthographies). Moreover, only in Hebrew did lexical status have similar effects on naming and lexical decision performance. Experiment 2 revealed that semantic priming effects in naming were larger in Hebrew than in English and completely absent in SerboCroatian. Experiment 3 revealed that a large proportion of nonlexical tokens (nonwords) in the stimulus list affects naming words in Hebrew and in English, but not in Serbo-Croatian. These resuits were interpreted as strong support for the orthographical depth hypothesis and suggest, in general, that in shallow orthographies phonology is generated dirr from print, whereas in deep orthographies phonology is derived from the internal lexicon.

Language-related field potentials in the anterior-medial temporal lobe: I. Intracranial distribution and neural generators.

Field potentials were recorded from intracranial electrodes in humans to study language-related processing. Subjects viewed sentences in which each word was presented successively in the center of a video monitor. Half of the sentences ended normally, while the other half ended with a semantically anomalous word. The anomalous sentence-ending words elicited a large negative field potential with a peak latency near 400 msec, which was focally distributed bilaterally in the anterior medial temporal lobe (AMTL), anterior to the hippocampus and near the amygdala. Subdural electrodes positioned near the collateral sulcus just inferior and lateral to the amygdala recorded a positive field potential at the same latency. This spatial distribution of voltage suggested that this language-sensitive field potential was generated in the neocortex near the collateral sulcus and anterior fusiform gyrus. Additional task-related field potentials were recorded in the hippocampus. The AMTL field potential at 400 msec shares characteristics with the N400 potential recorded from scalp electrodes that has been associated with semantic processing.

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 reduded 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.

Domain specificity versus expertise: factors influencing distinct processing of faces

To explore face specificity in visual processing, we compared the role of task-associated strategies and expertise on the N170 event-related potential (ERP) component elicited by human faces with the ERPs elicited by cars, birds, items of furniture, and ape faces. In Experiment 1, participants performed a car monitoring task and an animacy decision task. In Experiment 2, participants monitored human faces while faces of apes were the distracters. Faces elicited an equally conspicuous N170, significantly larger than the ERPs elicited by non-face categories regardless of whether they were ignored or had an equal status with other categories (Experiment 1), or were the targets (in Experiment 2). In contrast, the negative component elicited by cars during the same time range was larger if they were targets than if they were not. Furthermore, unlike the posterior-temporal distribution of the N170, the negative component elicited by cars and its modulation by task were more conspicuous at occipital sites. Faces of apes elicited an N170 that was similar in amplitude to that elicited by the human face targets, albeit peaking 10 ms later. As our participants were not ape experts, this pattern indicates that the N170 is face-specific, but not specie-specific, i.e. it is elicited by particular face features regardless of expertise. Overall, these results demonstrate the domain specificity of the visual mechanism implicated in processing faces, a mechanism which is not influenced by either task or expertise. The processing of other objects is probably accomplished by a more general visual processor, which is sensitive to strategic manipulations and attention.

Angry, Disgusted, or Afraid? Studies on the Malleability of Emotion Perception

Current theories of emotion perception posit that basic facial expressions signal categorically discrete emotions or affective dimensions of valence and arousal. In both cases, the information is thought to be directly “read out” from the face in a way that is largely immune to context. In contrast, the three studies reported here demonstrated that identical facial configurations convey strikingly different emotions and dimensional values depending on the affective context in which they are embedded. This effect is modulated by the similarity between the target facial expression and the facial expression typically associated with the context. Moreover, by monitoring eye movements, we demonstrated that characteristic fixation patterns previously thought to be determined solely by the facial expression are systematically modulated by emotional context already at very early stages of visual processing, even by the first time the face is fixated. Our results indicate that the perception of basic facial expressions is not context invariant and can be categorically altered by context at early perceptual levels.

Selective visual streaming in face recognition: evidence from developmental prosopagnosia.

Computational considerations suggest that efficient face identification requires the categorization and exclusive streaming of previously encoded face visual primitives into a dedicated face recognition system. Unique evidence supporting this claim is provided by a rare case of developmental pure prosopagnosia with otherwise normal visual and cognitive functions. Despite his normal visual memory and ability to describe faces, he is extremely impaired in face recognition. An early event related brain potential (N170) that is normally elicited exclusively by human faces, showed no specificity in this person. MRI revealed a smaller then normal right temporal lobe. These data emphasize the indispensability of the early streaming process for face recognition.