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Inversion leads to quantitative, not qualitative, changes in face processing.

Humans are remarkably adept at recognizing objects across a wide range of views. A notable exception to this general rule is that turning a face upside down makes it particularly difficult to recognize. This striking effect has prompted speculation that inversion qualitatively changes the way faces are processed. Researchers commonly assume that configural cues strongly influence the recognition of upright, but not inverted, faces. Indeed, the assumption is so well accepted that the inversion effect itself has been taken as a hallmark of qualitative processing differences. Here, we took a novel approach to understand the inversion effect. We used response classification to obtain a direct view of the perceptual strategies underlying face discrimination and to determine whether orientation effects can be explained by differential contributions of nonlinear processes. Inversion significantly impaired performance in our face discrimination task. However, surprisingly, observers utilized similar, local regions of faces for discrimination in both upright and inverted face conditions, and the relative contributions of nonlinear mechanisms to performance were similar across orientations. Our results suggest that upright and inverted face processing differ quantitatively, not qualitatively; information is extracted more efficiently from upright faces, perhaps as a by-product of orientation-dependent expertise.

Adult↗

The effects of inversion and eye displacements of familiar and unknown faces on early and late-stage ERPs.

OBJECTIVES: The objective of this study is to examine whether configural alterations of faces affect early or late processing stages as a function of their familiarity and their level of representation in memory. We then sought to verify whether the structural encoding stage is susceptible to top-down influences. METHODS: Electrophysiologic and behavioral studies were undertaken, during which unknown and familiar faces were presented upright or upside-down with or without feature alterations. The subjects were asked to determine whether the faces were familiar or not. RESULTS: N170 and N360 amplitudes were larger for familiar faces as well as altered ones. A higher degree of familiarity decreased reaction times (RTs) and N360 latencies, but increased N170 latencies, whereas face alterations increased RTs and latencies of both components examined. However, familiarity interacted with altered face configurations only for RTs and the N170. SIGNIFICANCE: In the perceptual stage, familiar faces seem to develop a more elaborate type of processing because of top-down influences linked to the robust nature of their representations in memory. The more elaborate type of processing for familiar faces has advantageous consequences for the following steps of information processing, by facilitating access to structural representations in memory (N360) as well as the final step reflected by RTs. The fact that configural alterations cause different effects for familiar as opposed to unfamiliar faces indicate that these stimuli are processed in a qualitatively different manner and solicit different representations in memory.

Adult↗

Naso-temporal asymmetry of the N170 for processing faces in normal viewers but not in developmental prosopagnosia.

Some elementary aspects of faces can be processed before cortical maturation or after lesion of primary visual cortex. Recent findings suggesting a role of an evolutionary ancient visual system in face processing have exploited the relative advantage of the temporal hemifield (nasal hemiretina). Here, we investigated whether under some circumstances face processing also shows a temporal hemifield advantage. We measured the face sensitive N170 to laterally presented faces viewed passively under monocular conditions and compared face recognition in the temporal and nasal hemiretina. A N170 response for upright faces was observed which was larger for projections to the nasal hemiretina/temporal hemifields. This pattern was not observed in a developmental prosopagnosic. These results point to the importance of the early stages of face processing for normal face recognition abilities and suggest a potentially important factor in the origins of developmental prosopagnosia.

Adult↗

How chemical information processing interferes with face processing: a magnetoencephalographic study.

Magnetic field changes related to face encoding were recorded in 20 healthy young participants. Faces had to be deeply encoded under four kinds of simultaneous nasal chemical stimulation. Neutral room air, phenyl ethyl alcohol (PEA, rose flavor), carbon dioxide (CO2, pain), and hydrogen sulfide (H2S, rotten eggs flavor) were used as chemical stimuli. PEA and H2S represented odor stimuli, whereas CO2 was used for trigeminal stimulation (pain sensation). After the encoding of faces, the respective recognition performances were tested focusing on recognition effects related to specific chemical stimulation during encoding. The number of correctly recognized faces (hits) varied between chemical conditions. PEA stimulation during face encoding significantly increased the number of hits compared to the control condition. H2S also led to an increased mean number of hits, whereas simultaneous CO2 administration during face encoding resulted in a reduction. Analysis of the physiological data revealed two latency regions of interest. Compared to the control condition, both olfactory stimulus conditions resulted in reduced activity components peaking at about 260 ms after stimulus onset, whereas CO2 produced a strongly pronounced enhanced activity component peaking at about 700 ms after stimulus onset. Both olfactory conditions elicited only weak enhanced activities at about 700 ms, and CO2 did not show any difference activity at 260 ms after stimulus onset compared to the control condition. It is concluded that the early activity differences represent subconscious olfactory information processing leading to enhanced memory performances irrespective of the hedonic value, at least if they are only subconsciously processed. The later activity is suggested to reflect conscious CO2 perception negatively affecting face encoding and therefore leading to reduced subsequent face recognition. We interpret that conscious processing of nasal chemical stimulation competes with deep face encoding with respect to cortical resources, whereas subconscious processing of nasal chemical stimulation does not.

Adult↗

Familiarity enhances invariance of face representations in human ventral visual cortex: fMRI evidence.

Face recognition across different viewing conditions is strongly improved by familiarity. In the present study, we tested the hypothesis that the neural basis of this effect is a less view-dependent representation of familiar faces in ventral visual cortex by assessing priming-related fMRI repetition effects. 15 healthy volunteers made male/female judgements on familiar (famous) and unfamiliar (novel) faces preceded by the same image, a different image of the same face, or another (unprimed) face. Reaction times revealed priming by same and different images independent of familiarity and more pronounced for same than different images. In the imaging data, a main effect of prime condition was found in bilateral fusiform and orbitofrontal regions. A right anterior fusiform region expressed stronger response decreases to repetition of familiar than unfamiliar faces. Bilateral mid-fusiform areas showed stronger response decreases to repetition of same than different images. A regions-of-interest analysis focussing specifically on face responsive regions suggested differences in the degree of image dependency across fusiform cortex. Collapsing across familiarity, there was greater image dependency of repetition effects in right than left anterior fusiform, replicating previous imaging findings obtained with common objects. For familiar faces alone, there was greater generalisation of repetition effects over different images in anterior than middle fusiform. This suggests a role of anterior fusiform cortex in coding image-independent representations of familiar faces.

Adult↗

Face perception: domain specific, not process specific.

Evidence that face perception is mediated by special cognitive and neural mechanisms comes from fMRI studies of the fusiform face area (FFA) and behavioral studies of the face inversion effect. Here, we used these two methods to ask whether face perception mechanisms are stimulus specific, process specific, or both. Subjects discriminated pairs of upright or inverted faces or house stimuli that differed in either the spatial distance among parts (configuration) or the shape of the parts. The FFA showed a much higher response to faces than to houses, but no preference for the configuration task over the part task. Similarly, the behavioral inversion effect was as large in the part task as the configuration task for faces, but absent in both part and configuration tasks for houses. These findings indicate that face perception mechanisms are not process specific for parts or configuration but are domain specific for face stimuli per se.

Brain↗

Evaluation of a shape-based model of human face discrimination using FMRI and behavioral techniques.

Understanding the neural mechanisms underlying object recognition is one of the fundamental challenges of visual neuroscience. While neurophysiology experiments have provided evidence for a "simple-to-complex" processing model based on a hierarchy of increasingly complex image features, behavioral and fMRI studies of face processing have been interpreted as incompatible with this account. We present a neurophysiologically plausible, feature-based model that quantitatively accounts for face discrimination characteristics, including face inversion and "configural" effects. The model predicts that face discrimination is based on a sparse representation of units selective for face shapes, without the need to postulate additional, "face-specific" mechanisms. We derive and test predictions that quantitatively link model FFA face neuron tuning, neural adaptation measured in an fMRI rapid adaptation paradigm, and face discrimination performance. The experimental data are in excellent agreement with the model prediction that discrimination performance should asymptote as faces become dissimilar enough to activate different neuronal populations.

Adolescent↗

Are Greebles like faces? Using the neuropsychological exception to test the rule.

Which image geometries count as face-like and which do not? Across multiple experiments, novel objects called Greebles have been used to argue that face-specific effects can be obtained with non-face stimuli under certain situations, in particular with expert observers. However, this claim depends on the argument that these non-face stimuli are not a priori treated by the face processing system. To address this question, CK, a neuropsychological patient well-known for exhibiting severe visual object agnosia and dyslexia but intact face processing, was tested with Greebles. CK performed poorly on Greebles, indicating that his intact face-specific abilities do not extend to include Greebles. These results suggest that insofar as CK is relying on face-specific visual processes, these processes do not a priori treat Greebles as faces.

Adolescent↗

A behavioural and ERP investigation of 3-month-olds' face preferences.

Recent behavioural work suggests that newborns' face preferences are derived from a general, non-specific attentional bias toward patterns with more features in the upper versus lower half. In the current study, we predicted that selectivity for the specific geometry of the face may emerge during the first 3 months of life as a product of perceptual narrowing, leading to the construction of the first broadly defined face category segregating faces from other visual objects which may share with faces one or more visual properties. This was investigated behaviourally, using a standard preferential looking paradigm, and electrophysiologically, using high-density ERPs. Behavioural results indicated that, at 3 months, the top-heavy property is no longer a crucial factor in determining face preferences. ERP results showed evidence of differentiation between the two stimuli only for the N700. No differentiation was found for earlier components that are thought to reflect the adult-like structural encoding stage of face processing in infants (N290 and P400). Together, ERP and behavioural results suggest that, by 3 months, the perceptual narrowing process has led to a behavioural response specific to the geometry of the human face, but that this response is not purely perceptual in nature. Rather, it seems related to the acquired salience of this stimulus category, which may reflect the high degree of familiarity and/or the social value faces have gained over the infants' first 3 months of life.

Electroencephalography↗

Accuracy of naming laterally presented known faces by children and adults.

Two experiments investigating asymmetries in the naming of laterally presented known faces are reported. In Experiment I, upright and inverted faces of classmates or colleagues were presented bilaterally and unilaterally to seven-year-old children, eleven-year-old children, and adults. A LVF (left visual hemifield) superiority was found for naming upright faces, unrelated to age. For inverted faces there was no difference between the visual hemifields. In Experiment II faces of famous people were used as stimuli. When adult subjects were given little indication as to which faces to expect (Condition A) there was no difference between the visual hemifields. When they were given a list of the faces to be used (Condition B) a LVF naming superiority was found. The results are interpreted as indicating superiority of the right cerebral hemisphere for a component of the processing of known faces that is of importance when the subject knows which faces he can expect to see.

Adult↗

Perceived gaze direction in faces and spatial attention: a study in patients with parietal damage and unilateral neglect.

Perceived gaze in faces is an important social signal that may influence orienting of attention in normal observers. Would such effects of gaze still occur in patients with right parietal damage and left neglect who usually fail to attend to contralesional space? Two experiments tested for effects of perceived gaze on visual extinction. Face or shape stimuli were presented in the right, left, or both hemifields, with faces looking either straight ahead or toward the opposite field. On bilateral trials, patients extinguished a left shape much less often when a concurrent right face looked leftward rather than straight ahead. This occurred, even though gaze was not relevant to the task and processing of facial signals implied attention to a competing ipsilesional stimulus. By contrast, rightward gaze in faces presented on the left side had no effect on extinction, suggesting that gaze cues are not extracted without attention. Two other experiments examined effects of perceived gaze on the detection of peripheral targets. Targets appeared at one of four possible locations to the right or left of a central face looking either toward the target location, another location on the same side, the opposite side, or straight ahead. Face and gaze were not relevant to the task and not predictive of target location. Patients responded faster when the face looked toward the target on both the contralesional and ipsilesional sides. In contralesional space, gaze allowed shifting of attention in a specific quadrant direction, but only to the first target along the scan path when there were different possible locations on the same side. By contrast, in intact ipsilesional space, attention was selectively directed to one among different eccentric locations. Control experiments showed that symbolic arrow cues did not produce similar effects. These results indicate that even though parietal damage causes spatial neglect and impairs the representation of location on the contralesional side, perceived gaze in faces can still trigger automatic shifts of attention in the contralesional direction, suggesting the existence of specific and anatomically distinct attentional mechanisms.

Aged↗

Understanding memory for faces in Parkinson's disease: the role of configural processing.

It has previously been reported that unfamiliar face recognition memory is impaired in Parkinson's disease (PD) [(Dewick, H. C., Hanley, J. R., Davies, A. D. M., Playfer, J. R. & Turnbull, C. J., Perception and memory for faces in Parkinson's disease. Neuropsychologia, 1991, 29, 785-802), (Haeske-Dewick, H. C., Are perception and memory for faces influenced by a specific age at onset factor in Parkinson's disease? Neuropsychologia, 1996, 34, 315-320), (Levin, B. E., Llabre, M. M. & Weiner, W. J., Cognitive impairments associated with early Parkinson's disease. Neurology, 1989, 39, 557-561)]. In the work reported here, we consider the possible mechanisms that might underlie this impairment. 28 PD patients and 28 controls were given a two-part test of recognition memory for words and faces, and two perceptual tests to measure their configural and componential processing ability. We found that PD patients were significantly worse than controls on the recognition memory test for faces, but not when the stimuli were words. In addition, PD patients were significantly impaired relative to controls on the closure test (FCT) used to measure configural processing, but there was no difference between the two groups on a test of componential processing ability. Multiple regression analyses revealed that even after accounting for the influence of age, intelligence and level of depression, configural processing ability was the important predictor of unfamiliar face recognition memory in Parkinson's disease. There was no effect of Parkinson's disease specific variables on either face recognition or FCT performance. In addition, some recently diagnosed patients were poor at face recognition. It is suggested that face configuration skills may be affected very early in the course of Parkinson's disease, and that this may be connected to the fact that considerable nigrostriatal degeneration and alteration in brain neurotransmitter levels occur before the clinical symptoms of PD appear.

Aged↗

Event-related potentials in macaque monkey during passive and attentional processing of faces in a priming paradigm.

Epidural event-related potentials (ERPs) were recorded in monkeys using a priming (S1-S2) paradigm. One juvenile and 2 young adult macaque monkeys (Macaca fascicularis) viewed tachistoscopically presented pairs of faces in four possible combinations: human-human, monkey-monkey, monkey-human, human-monkey. Faces were presented upright to untrained subjects. Subsequently, one adult monkey was trained to associate the occurrence of a monkey-monkey pair of faces with juice reward. In a separate control experiment, the same stimuli were presented upside down. In both the upright and inverted conditions, adult monkey ERPs exhibited a P1-N1-P2 complex of peaks in the 400 ms following stimulus presentation. However, only upright faces elicited a prominent, widely distributed negativity (N4) in the 400-800 ms interval and a long-duration negativity (LDN) in the last 400 ms of the 1,400-ms epoch. N4 was earlier in latency, larger in amplitude, and larger over left hemisphere in response to monkey faces compared to human faces. It was also sensitive to the preceding or priming stimulus, exhibiting larger amplitudes when S1 matched S2 than when it did not. Trained monkey waveforms differed from untrained monkey responses in terms of the appearance of an N2-P3 complex. P3 was bilaterally symmetrical, larger in magnitude, and delayed in latency in response to monkey faces. The differences in components in the two conditions suggest that monkey ERPs can distinguish between passive and attentional processing of faces. N4 appears to reflect the recognition of conspecifics, is sensitive to priming, and occurs primarily over left hemisphere. Rewarded presentation of faces results in P3-like components that may reflect the novelty, meaningfulness, or active nature of the response. The differences in distribution between P3 and N4 suggest that they do not share the same neural sources.

Aging↗

Scopolamine but not lorazepam modulates face repetition priming: a psychopharmacological fMRI study.

Repetition priming is a basic form of learning associated with decreased neuronal responses following stimulus repetition. In this experiment, we address cholinergic and GABAergic modulation of repetition priming in a face recognition paradigm. In experiment 1, we used event-related functional magnetic resonance imaging (fMRI) in combination with pharmacological challenge where participants were given placebo, lorazepam (2mg po), or scopolamine (0.4 mg IV) prior to study. Behavioral data showed intact priming for famous faces in the placebo and lorazepam group but impaired priming following scopolamine. In within-group analyses, a right fusiform region showed a fame by repetition interaction characterized by a response decrease to repetition of famous faces and a response enhancement to repetition of unfamous faces in the placebo group. In subjects treated with lorazepam, a main effect of repetition, driven by response decreases to repetition of famous faces, was seen in this right fusiform region. No significant repetition effects were found after scopolamine. In experiment 2, we further investigated behaviorally the cholinergic impairment of repetition priming. Participants were given either placebo or scopolamine (0.4 mg IV) after study. Behavioral data showed intact priming for famous faces in the placebo and scopolamine group. The results suggest that scopolamine but not lorazepam impair repetition priming for famous faces in a face recognition paradigm. These cholinergic impairments are likely to reflect interference with acquisition processes during study that may co-occur with a modulation of right fusiform decreases to repetition of famous faces.

Acetylcholine↗

Perceptual dominance of oriented faces mirrors the distribution of orientation tunings in inferotemporal neurons.

In three experiments participants viewed pairs of overlapping transparent faces, with one face upright and the other oriented, and they reported which face was dominant. In each trial, an upright face was presented with a face at 45, 90, 135 or 180 degrees, with transparency set using a linear weighted algorithm, so that relative contrast across faces was biased in favour of oriented faces. Exposure duration was restricted in experiment 1 to 250, 500 or 1000 ms, but was unlimited in experiments 2 and 3. Adults were tested in experiments 1 and 2 and children aged 6-9 years of age were tested in experiment 3. Irrespective of exposure duration, the results showed the probability of dominance being ceded by oriented faces to upright faces was a function of orientation. In comparable conditions, the function found with young children was flatter than with adults. These patterns, and those of earlier perceptual studies, can be explained by the distribution of different orientation tunings found in physiological studies of inferotemporal cortex in macaques.

Adult↗

Searching for face-specific long latency ERPs: a topographic study of effects associated with mismatching features.

In a previous study [E. Olivares, M.A. Bobes, E. Aubert, M. Valdés-Sosa, Associative ERPs effects with memories of artificial faces, Cogn. Brain Res. 2 (1994) 39-48] we reported the presence of a negativity associated with mismatching features when subjects carried out a face-feature matching task whilst their evoked potentials were recorded. Since the stimuli used were learned faces (realistic drawings), for which the subjects possessed no semantic information or associated verbal labels, the mismatch negativity obtained was considered a face-specific N400. In this work we present a new experiment to study the topographic distribution of these mismatch effects. As in the above-mentioned study, in each trial the subjects observed previously an incomplete (without the eyes/eyebrows fragment) familiar face, which served as a structural context for the face recognition. The face was then completed by grafting either matching (learned) features or mismatching features (from another face). In line with neuropsychological studies on prosopagnosia and electrophysiological findings in humans and non-human primates, we found as one of the most relevant items of data that the most-posterior (principally, left occipital) cortices appear to be a region in which are located the possible neural generators of the negativity associated with the detection of incongruencies in the structure of familiar faces. We also reported a late positivity, distributed in more anterior regions, which follows the mismatch negativity. This complex N-P is interpreted as reflecting a dual process of retrieval and integration of information in memory.

Adult↗

Effects of perceptual quality on the processing of human faces presented to the left and right cerebral hemispheres.

Three experiments examined the effects of stimulus duration, retinal eccentricity, and visual noise on the processing of human faces presented to the left visual field/right hemisphere (LVF-RH) and right visual field/left hemisphere (RVF-LH). In Experiment 1 observers identified which of 10 similar male faces was presented on a screen. The single face was presented for 10, 55, or 100 ms at 1 degree, 4 degrees, or 9 degrees of visual angle to the left or right of fixation. Decreasing stimulus duration and increasing retinal eccentricity lowered face recognition. The effect of duration was the same for LVF-RH and RVF-LH trials, but the detrimental effect of increasing retinal eccentricity was larger on LVF-RH trials than on RVF-LH trials. In Experiment 2 observers indicated whether a single face from this same set was a member of a memorized set of five positive faces. The probe face on each trial was presented alone or embedded in visual noise. Visual noise increased the error rate more on LVF-RH trials than on RVF-LH trials. This effect was replicated in Experiment 3, which also required observers to make a much easier discrimination between male and female faces. In the male/female task visual noise tended to impair performance more on RVF-LH trials than on LVF-RH trials, opposite the effect for the male/male task. These results are discussed in terms of hemispheric asymmetry for global versus local features of faces, the level of feature analysis demanded by a task, and the level of feature analysis most disrupted by perceptual degradation.

Attention↗

About face: left-hemisphere involvement in processing physiognomies.

Three experiments were concluded to investigate the involvement of the two cerebral hemispheres in processing faces. Perceptual discrimination of pairs of faces was equally speedy overall when the stimuli were presented in the right visual field (RVF) or left visual field (LVF). For faces differing in one or two features, however, a qualitatively different pattern of results was obtained for the two visual fields, and an RVF advantage emerged when the difference lay in the upper part of the faces (Experiment 1). An examination of the discriminability of the facial features from which the faces were constructed (Experiment 2) showed that the processes involved in RVF comparisons of faces were not dependent on the saliency of the features but, rather, followed a top-to-bottom serial analysis of the stimuli; the speed of the processing involved in LVF presentations was a function of the degree of similarity of the different comparison faces. Evidence for a serial type of comparison faces were used (Experiment 3). It was concluded that even though comparisons were equally speedy overall in LVF and RVF presentations, qualitatively different processes take place in the two hemispheres, which prove competent at processing faces, each in its own way. Some methodological problems inherent in tachistoscopic studies are discussed, and it is proposed that the quality of the stimulus representation achieved or required for cognitive processing may be determinant in the emergence of functional hemispheric asymmetries.

Adult↗