The horizontal and vertical relations in upright faces are transmitted by different spatial frequency ranges

Faces convey distinct types of information: features and their spatial relations, which are differentially vulnerable to inversion. While inversion largely disrupts the processing of vertical spatial relations (e.g. eyes’ height), its effect is moderate for horizontal relations (e.g. interocular distance) and local feature properties. The SF ranges optimally transmitting horizontal and vertical face relations were here investigated to further address their functional role in face perception. Participants matched upright and inverted pairs of faces that differed at the level of local featural properties, horizontal relations in vertical relations. Irrespective of SF, the inversion effect was larger for vertical than horizontal and featural cues. Most interestingly, SF differentially influenced the processing of vertical, horizontal and featural cues in upright faces. Vertical relations were optimally processed in intermediate SF, which are known to carry useful information for face individuation. In contrast, horizontal relations were best conveyed by high SF, which are involved in the processing of local face properties. These findings not only confirm that horizontal and vertical relations play distinct functional roles in face perception, but they also further suggest a unique role of vertical relations in face individuation.     

Face inversion disproportionately impairs the perception of vertical but not horizontal relations between features

Upside-down inversion disrupts the processing of spatial relations between the features of a face, while largely preserving local feature analysis. However, recent studies on face inversion failed to observe a clear dissociation between relational and featural processing. To resolve these discrepancies and clarify how inversion affects face perception, the authors monitored inversion effects separately for vertical and horizontal distances between features. Inversion dramatically declined performance in the vertical-relational condition, but it impaired featural and horizontal-relational performance only moderately. Identical observations were made whether upright and inverted trials were blocked or randomly interleaved. The largest performance decrement was found for vertical relations even when faces were rotated by 90 degrees. Evidence that inversion dramatically disrupts the ability to extract vertical but not horizontal feature relations supports the view that inversion qualitatively changes face perception by rendering some of the processes activated by upright faces largely ineffective.     

The time course of face matching for featural and relational image manipulations

It was found recently that horizontal and vertical relationships of facial features are differently vulnerable to inversion (Goffaux & Rossion, 2007). When faces are upside down manipulations of vertical relations are difficult to detect, while only moderate performance deficits are found for manipulations of horizontal relations, or when features differ. We replicate the findings of Goffaux and Rossion, and record the temporal courses of face matching performance and the effects of inversion. For vertical relations and featural changes inversion effects arise immediately, starting with the first 50 ms of processing. For horizontal relations inversion effects are absent at brief timings, but arise later, after about 200 ms. The final magnitudes of inversion effects are different in all three conditions, reaching 23.4% for vertical displacement, 10.7% for feature replacement, and 5.8% for horizontal displacement. Our results underline the special status of horizontal in contrast to vertical feature relations, and indicate that local and global configural information is handled in parallel by specific routines, as proposed recently (Senunkova & Barton, 2008).     

The effects of face inversion on the perception of long-range and local spatial relations in eye and mouth configuration

A recent study hypothesized a configurational anisotropy in the face inversion effect, with vertical relations more difficult to process. However, another difference in the stimuli of that report was that the vertical but not horizontal shifts lacked local spatial references. Difficulty processing long-range spatial relations might also be predicted from a relevance-interaction explanation, which proposes that in inverted faces, spatial relations are processed efficiently only within high-relevance local regions. The authors performed 2 experiments to distinguish between these hypotheses. Experiment 1 showed that the inversion effect for vertical shifts of the eyes alone was more similar to that for horizontal eye shifts than for vertical shifts of the eyes and eyebrows. In Experiment 2, focused attention reduced the inversion effect for vertical mouth position more than that for vertical shifts of the eyes and brows. The authors concluded that face inversion impairs the perception of both local spatial relations in low-relevance regions and long-range spatial relations extending across multiple facial regions, consistent with a loss of efficient whole-face processing of the spatial relations between features.     

Structure and function in acquired prosopagnosia: Lessons from a series of 10 patients with brain damage

Acquired prosopagnosia varies in both behavioural manifestations and the location and extent of underlying lesions. We studied 10 patients with adult‐onset lesions on a battery of face‐processing tests. Using signal detection methods, we found that discriminative power for the familiarity of famous faces was most reduced by bilateral occipitotemporal lesions that involved the fusiform gyri, and better preserved with unilateral right‐sided lesions. Tests of perception of facial structural configuration showed severe deficits with lesions that included the right fusiform gyrus, whether unilateral or bilateral. This deficit was most consistent for eye configuration, with some patients performing normally for mouth configuration. Patients with anterior temporal lesions had better configuration perception, though at least one patient showed a more subtle failure to integrate configural data from different facial regions. Facial imagery, an index of facial memories, was severely impaired by bilateral lesions that included the right anterior temporal lobe and marginally impaired by fusiform lesions alone; unilateral right fusiform lesions tended to spare imagery for facial features. These findings suggest that (1) prosopagnosia is more severe with bilateral than unilateral lesions, indicating a minor contribution of the left hemisphere to face recognition, (2) perception of facial configuration critically involves the right fusiform gyrus and (3) access to facial memories is most disrupted by bilateral lesions that also include the right anterior temporal lobe. This supports assertions that more apperceptive variants of prosopagnosia are linked to fusiform damage, whereas more associative variants are linked to anterior temporal damage. Next, we found that behavioural indices of covert recognition correlated with measures of overt familiarity, consistent with theories that covert behaviour emerges from the output of damaged neural networks, rather than alternative pathways. Finally, to probe the face specificity of the prosopagnosic defect, we tested recognition of fruits and vegetables: While face specificity was not found in most of our patients, the data of one patient suggested that this may be possible with more focal lesions of the right fusiform gyrus.     

Early development of letter specialization in left fusiform is associated with better word reading and smaller fusiform face area

A functional region of left fusiform gyrus termed “the visual word form area” (VWFA) develops during reading acquisition to respond more strongly to printed words than to other visual stimuli. Here, we examined responses to letters among 5‐ and 6‐year‐old early kindergarten children (N = 48) with little or no school‐based reading instruction who varied in their reading ability. We used functional magnetic resonance imaging (fMRI) to measure responses to individual letters, false fonts, and faces in left and right fusiform gyri. We then evaluated whether signal change and size (spatial extent) of letter‐sensitive cortex (greater activation for letters versus faces) and letter‐specific cortex (greater activation for letters versus false fonts) in these regions related to (a) standardized measures of word‐reading ability and (b) signal change and size of face‐sensitive cortex (fusiform face area or FFA; greater activation for faces versus letters). Greater letter specificity, but not letter sensitivity, in left fusiform gyrus correlated positively with word reading scores. Across children, in the left fusiform gyrus, greater size of letter‐sensitive cortex correlated with lesser size of FFA. These findings are the first to suggest that in beginning readers, development of letter responsivity in left fusiform cortex is associated with both better reading ability and also a reduction of the size of left FFA that may result in right‐hemisphere dominance for face perception.     

Effects of geometric distortions on face-recognition performance

The importance of 'configural' processing for face recognition is now well established, but it remains unclear precisely what it entails. Through four experiments we attempted to clarify the nature of configural processing by investigating the effects of various affine transformations on the recognition of familiar faces. Experiment 1 showed that recognition was markedly impaired by inversion of faces, somewhat impaired by shearing or horizontally stretching them, but unaffected by vertical stretching of faces to twice their normal height. In experiment 2 we investigated vertical and horizontal stretching in more detail, and found no effects of either transformation. Two further experiments were performed to determine whether participants were recognising stretched faces by using configural information. Experiment 3 showed that nonglobal vertical stretching of faces (stretching either the top or the bottom half while leaving the remainder undistorted) impaired recognition, implying that configural information from the stretched part of the face was influencing the process of recognition--ie that configural processing involves global facial properties. In experiment 4 we examined the effects of Gaussian blurring on recognition of undistorted and vertically stretched faces. Faces remained recognisable even when they were both stretched and blurred, implying that participants were basing their judgments on configural information from these stimuli, rather than resorting to some strategy based on local featural details. The tolerance of spatial distortions in human face recognition suggests that the configural information used as a basis for face recognition is unlikely to involve information about the absolute position of facial features relative to each other, at least not in any simple way.     

Dividing the self: distinct neural substrates of task-based and automatic self-prioritization after brain damage

Facial self-awareness is a basic human ability dependent on a distributed bilateral neural network and revealed through prioritized processing of our own over other faces. Using non-prosopagnosic patients we show, for the first time, that facial self-awareness can be fractionated into different component processes. Patients performed two face perception tasks. In a face orientation task, they judged whether their own or others' faces were oriented to the left or right. In the 'cross' experiment, they judged which horizontal or vertical element in a cross was relatively longer while ignoring a task-irrelevant face presented as background. The data indicate that impairments to a distinct task-based prioritization process (when faces had to be attended) were present after brain damage to right superior frontal gyrus, bilateral precuneus, and left middle temporal gyrus. In contrast, impairments to automatic prioritization processes (when faces had to be ignored) were associated only with left hemisphere damage (the cingulate gyrus, superior parietal lobe, and superior temporal gyrus). In addition, both automatic and task-based self-prioritizations were affected by damage to left supramarginal and angular gyrus. The results for the gray matter analyses also extended to the adjacent white matter fiber tracts including the inferior occipital-frontal fasciculus, cingulum, and optic radiation. The data provide the first empirical evidence for separate functional roles of the left and right hemispheres in different aspects of self-face perception and suggest distinct functional processes respectively for paying attention to and for ignoring self-related information.     

Picture-plane inversion leads to qualitative changes of face perception

Presenting a face stimulus upside-down generally causes a larger deficit in perceiving metric distances between facial features ("configuration") than local properties of these features. This effect supports a qualitative account of face inversion: the same transformation affects the processing of different kinds of information differently. However, this view has been recently challenged by studies reporting equal inversion costs of performance for discriminating featural and configural manipulations on faces. In this paper I argue that these studies did not replicate previous results due to methodological factors rather than largely irrelevant parameters such as having equal performance for configural and featural conditions at upright orientation, or randomizing trials across conditions. I also argue that identifying similar diagnostic features (eyes and eyebrows) for discriminating individual faces at upright and inverted orientations by means of response classification methods does not dismiss at all the qualitative view of face inversion. Considering these elements as well as both behavioral and neuropsychological evidence, I propose that the generally larger effect of inversion for processing configural than featural cues is a mere consequence of the disruption of holistic face perception. That is, configural relations necessarily involve two or more distant features on the face, such that their perception is most dependent on the ability to perceive simultaneously multiple features of a face as a whole.     

Neural developmental changes in processing inverted faces

We explored developmental changes in neural substrates for face processing, using fMRI. Children and adults performed a perceptual-matching task with upright and inverted face and animal stimuli. Behaviorally, inversion disrupted face processing more than animal processing for adults and older children. In line with this behavioral pattern, the left middle occipital gyrus showed a stronger face than animal inversion effect in adults. Moreover, a superior aspect of this region showed a greater face inversion effect in older than in younger children, indicating a developmental change in the processing of inverted faces. The visual regions recruited for inverted face processing in adults also overlapped more with brain regions involved in the viewing of upright objects than with regions involved in the viewing of upright faces in an independent localizer task. Hence, when faces are inverted, adults recruit regions normally engaged for recognizing objects, possibly pointing to a role for the featural processing of inverted faces.     

Explaining the face-inversion effect: the face–scheme incompatibility (FSI) model

The face-inversion effect (FIE) can be viewed as being based on two kinds of findings. According to the face(UI) effect, perception and recognition are better for faces presented upright (U) than for faces presented inverted (I). According to the face/object(UI) effect, inversion impairs the processing of faces more than the processing of nonfacial objects (e.g., buildings or cars). Part I of this article focuses on the face(UI) effect and the configural-processing hypothesis, which is considered the most popular explanatory hypothesis of the FIE. In this hypothesis, it is proposed that inversion impairs the processing of configural information (the spatial relations between features) but hardly (if at all) impairs the processing of featural information (e.g., eyes, nose, and mouth). Part II of the article starts from the conclusion reached in part I, that the configural-processing hypothesis has not succeeded in explaining a substantial number of the findings and in resolving certain theoretical problems. The part then goes on to outline a new alternative model, the face–scheme incompatibility (FSI) model, which contends with these theoretical problems, accounts for the configural-processing hypothesis, succeeds in explaining a considerable portion of the empirical findings related to the face(UI) effect, and proposes a relatively new research program on the concept of the face scheme. The basic assumption of the FSI model is that schemes and prototypes are involved in processing a visual stimulus of a face and in transforming it to a “meaning-bearing” face, and that different schemes are involved if the face is presented upright or inverted.     

Older adults' configural processing of faces: role of second-order information

Problems with face recognition are frequent in older adults. However, the mechanisms involved have only been partially discovered. In particular, it is unknown to what extent these problems may be related to changes in configural face processing. Here, we investigated the face inversion effect (FIE) together with the ability to detect modifications in the vertical or horizontal second-order relations between facial features. We used a same/different unfamiliar face discrimination task with 33 young and 33 older adults. The results showed dissociations in the performances of older versus younger adults. There was a lack of inversion effect during the recognition of original faces by older adults. However, for modified faces, older adults showed a pattern of performance similar to that of young participants, with preserved FIE for vertically modified faces and no detectable FIE for horizontally modified faces. Most importantly, the detection of vertical modifications was preserved in older relative to young adults whereas the detection of horizontal modifications was markedly diminished. We conclude that age has dissociable effects on configural face-encoding processes, with a relative preservation of vertical compared to horizontal second-order relations processing. These results help to understand some divergent results in the literature and may explain the spared familiar face identification abilities in the daily lives of older adults.     

眼睛区域构型信息与特征信息的跨维共变增益效应及其加工特异性

面孔知觉可能在区域尺度上发生多维信息整合, 但迄今无特异性实验证据。本研究在两个实验中操纵面孔眼睛区域或嘴巴区域的单维构型或特征信息, 测量人们觉察单维变化或跨维共变的敏感度, 以此检测面孔区域尺度上的多维信息整合有何现象与规律, 进而揭示面孔知觉的多维信息整合机制。实验获得3个发现：(1)正立面孔眼睛区域的信息变化觉察呈现出“跨维共变增益效应”, 跨维信息共变觉察的敏感度显著高于任意一种单维信息变化觉察的敏感度; (2)“跨维共变增益效应”只在正立面孔的眼睛区域出现, 在倒置面孔的眼睛区域、正立面孔的嘴巴区域或倒置面孔的嘴巴区域都没有出现, 因此具有面孔区域特异性和面孔朝向特异性; (3)就单维信息变化觉察而言, 眼睛区域的敏感度不会受到面孔倒置的损伤, 但是嘴巴区域的敏感度会受到面孔倒置的显著损伤。综合可知, 面孔知觉确实会发生区域尺度上的信息整合, 它不是普遍性的信息量效应, 而是特异性的眼睛效应(只发生在正立面孔的眼睛区域)。这是面孔整体加工(face holistic processing)在单维信息分辨和多维信息整合之间建立联系的必经环节。这提示我们对全脸多维信息知觉整合的理解需要从传统的面孔整体加工假设升级到以眼睛为中心的层级化多维信息整合算法(a hierarchical algorithm for multi-dimensional information integration)。     

Animal,but not human,faces engage the distributed face network in adolescents with autism

Multiple hypotheses have been offered to explain the impaired face‐processing behavior and the accompanying underlying disruptions in neural circuitry among individuals with autism. We explored the specificity of atypical face‐processing activation and potential alterations to fusiform gyrus (FG) morphology as potential underlying mechanisms. Adolescents with high functioning autism (HFA) and age‐matched typically developing (TD) adolescents were scanned with sMRI and fMRI as they observed human and animal faces. In spite of exhibiting comparable face recognition behavior, the HFA adolescents evinced hypo‐activation throughout the face‐processing system in response to unfamiliar human, but not animal, faces. They also exhibited greater activation in affective regions of the face‐processing network in response to animal, but not human, faces. Importantly, this atypical pattern of activation in response to human faces was not related to atypical structural properties of the FG. This atypical neural response to human faces in autism may stem from abnormalities in the ability to represent the reward value of social (i.e. conspecific) stimuli.     

Haptic face identification activates ventral occipital and temporal areas: An fMRI study

Many studies in visual face recognition have supported a special role for the right fusiform gyrus. Despite the fact that faces can also be recognized haptically, little is known about the neural correlates of haptic face recognition. In the current fMRI study, neurologically intact participants were intensively trained to identify specific facemasks (molded from live faces) and specific control objects. When these stimuli were presented in the scanner, facemasks activated left fusiform and right hippocampal/parahippocampal areas (and other regions) more than control objects, whereas the latter produced no activity greater than the facemasks. We conclude that these ventral occipital and temporal areas may play an important role in the haptic identification of faces at the subordinate level. We further speculate that left fusiform gyrus may be recruited more for facemasks than for control objects because of the increased need for sequential processing by the haptic system.     

Haptic face identification activates ventral occipital and temporal areas: an fMRI study

Many studies in visual face recognition have supported a special role for the right fusiform gyrus. Despite the fact that faces can also be recognized haptically, little is known about the neural correlates of haptic face recognition. In the current fMRI study, neurologically intact participants were intensively trained to identify specific facemasks (molded from live faces) and specific control objects. When these stimuli were presented in the scanner, facemasks activated left fusiform and right hippocampal/parahippocampal areas (and other regions) more than control objects, whereas the latter produced no activity greater than the facemasks. We conclude that these ventral occipital and temporal areas may play an important role in the haptic identification of faces at the subordinate level. We further speculate that left fusiform gyrus may be recruited more for facemasks than for control objects because of the increased need for sequential processing by the haptic system.     

Influence of subjective difficulty on the degree of configural and featural processing in face recognition1

This study aimed to assess the effect of subjective difficulty on the degree of featural or configural processing in face recognition. It could be assumed that featural processing is analytic processing ( Peterson & Rhodes, 2003 ), while configural processing is automatic processing ( Dunning & Stern, 1994 ). It has been suggested that task difficulty affected the automaticity of processing. Task difficulty was manipulated using a number of alternatives or time pressure. Subjective difficulty could also affect the automaticity of processing. If so, then subjective difficulty may in turn affect the degree of featural or configural processing. Participants in a difficult condition were given instructions mentioning that the face‐recognition task was difficult, while participants in the control condition were given no such instruction. The amount of retrieved featural information in the former condition was less than that in the latter. These results suggested that the degree of featural processing decreased when the participants found the task difficult.     

Regional variation in the inversion effect for faces: differential effects for feature shape, feature configuration, and external contour

Faces are perceived via an orientation-dependent expert mechanism. We previously showed that inversion impaired perception of the spatial relations of features more in the lower face than in the (more salient) upper face, suggests a failure to rapidly process this type of structural data from the entire face. In this study we wished to determine if this interaction between inversion and regional salience, which we consider a marker for efficient whole-face processing, was specific to second-order (coordinate) spatial relations or also affected other types of structural information in faces. We used an oddity paradigm to test the ability of seventeen subjects to discriminate changes in feature size, feature spatial relations, and external contour in both the upper and lower face. We also tested fourteen subjects on perception of two different types of spatial relations: second-order changes that create plausible alternative faces, and illegal spatial changes that transgress normal rules of facial geometry. In both experiments we examined for asymmetries between upper-face and lower-face perceptual accuracy with brief stimulus presentations. While all structural changes were less easily discerned in inverted faces, only changes to spatial relations showed a marked asymmetry between the upper and lower face, with far worse performance in the mouth region. Furthermore, this asymmetry was found only for second-order spatial relations and not illegal spatial changes. These results suggest that the orientation-dependent face mechanism has a rapid whole-face processing capacity specific to the internal second-order (coordinate) spatial relations of facial features.     

The respective role of low and high spatial frequencies in supporting configural and featural processing of faces

One distinctive feature of processing faces, as compared to other categories, is thought to be the large dependence on configural cues such as the metric relations among features. To test the role of low spatial frequencies (LSFs) and high spatial frequencies (HSFs) in configural and featural processing, subjects were presented with triplets of faces that were filtered to preserve either LSFs (below 8 cycles per face width), HSFs (above 32 cycles per face width), or the full frequency spectrum. They were asked to match one of two probe faces to a target face. The distractor probe face differed from the target either configurally, featurally, or both featurally and configurally. When the difference was at the configural level, performance was better with LSF faces than with HSF faces. In contrast, with a featural difference, a strong performance advantage was found for HSF faces as compared to LSF faces. These results support the dominant role that LSFs play in the configural processing of faces, whereas featural processing is largely dependent on HSFs.     

Features are also important: contributions of featural and configural processing to face recognition

It has been suggested that face recognition is primarily based on configural information, with featural information playing little or no role. We investigated this idea by comparing the prototype effect for face prototypes that emphasized either featural or configural processing. In Experiment 1, participants showed a tendency to commit false alarms in response to nonstudied prototypes, and this tendency was equivalent for featural and configural prototypes. Experiment 2 replicated this finding, and provided support for the assumption that the two types of prototypes differed in terms of featural and configural processing: Face inversion eliminated the prototype effect for configural prototypes but not for featural prototypes. These results suggest that both featural and configural processing make important contributions to face recognition, and that their effects are dissociable.