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.     

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.     

Expertise training with novel objects leads to left-lateralized facelike electrophysiological responses

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

Combined effects of inversion and feature removal on N170 responses elicited by faces and car fronts

The face-sensitive N170 is typically enhanced for inverted compared to upright faces. Itier, Alain, Sedore, and McIntosh (2007) recently suggested that this N170 inversion effect is mainly driven by the eye region which becomes salient when the face configuration is disrupted. Here we tested whether similar effects could be observed with non-face objects that are structurally similar to faces in terms of possessing a homogeneous within-class first-order feature configuration. We presented upright and inverted pictures of intact car fronts, car fronts without lights, and isolated lights, in addition to analogous face conditions. Upright cars elicited substantial N170 responses of similar amplitude to those evoked by upright faces. In strong contrast to face conditions however, the car-elicited N170 was mainly driven by the global shape rather than the presence or absence of lights, and was dramatically reduced for isolated lights. Overall, our data confirm a differential influence of the eye region in upright and inverted faces. Results for car fronts do not suggest similar interactive encoding of eye-like features and configuration for non-face objects, even when these objects possess a similar feature configuration as faces.     

Effect of inversion on the recognition of external and internal facial features

The purpose of the present study was to find out whether inversion affects recognition of external and internal facial features. 24 participants matched, under two experimental conditions (pair and multiple-choice matchings), upright target faces with three categories of facial test stimuli: full faces, external features and internal features, which were presented in either upright or inverted orientations. Data analysis showed that matching of facial stimuli was faster, more accurate and more consistent under upright than under inverted orientations for all stimulus categories; mostly for full faces, and least for internal features. As a rule, there were no speed-accuracy trade-offs. Implications of the data for accounts of the inversion effect in face recognition in terms of a shift from configurational to componential processing were discussed.     

面孔倒置效应的研究与理论述评

面孔倒置效应是指, 个体对倒置面孔的再认成绩显著低于对正立面孔的再认成绩, 而且, 与普通物体的倒置效应相比, 面孔的倒置效应更大。研究者们针对面孔倒置效应提出了多种理论假说, 两种主要的理论解释是：第一, 倒置面孔破坏了面孔的结构特征, 影响了面孔的整体加工, 所以, 个体对倒置面孔的再认成绩较差; 第二, 眼睛在面孔加工中起重要作用, 尤其在倒置面孔中的作用更明显, 使得倒置面孔N170波幅较高。ERP和fMRI研究提示, 未来研究需要进一步明确眼睛、注意等在面孔倒置效应中的作用。     

Beyond U-Shaped Development in Infants' Processing of Faces: An Information-Processing Account

The development of the "inversion" effect in face processing was examined in infants 3 to 6 months of age by testing their integration of the internal and external features of upright and inverted faces using a variation of the "switch" visual habituation paradigm. When combined with previous findings showing that 7-month-olds use integrative processing of an upright face, but featural processing of an inverted face (Cohen & Cashon, 2001a), the present findings suggest that from 3 to 7 months, infants' ability to integrate facial features follows an N-shaped developmental pattern for upright faces and an inverted U-shaped pattern for inverted faces. We discuss these results in terms of a set of domain-general information-processing principles.     

An inversion effect modified by expertise in capuchin monkeys

The face inversion effect may be defined as the general impairment in recognition that occurs when faces are rotated 180°. This phenomenon seems particularly strong for faces as opposed to other objects and is often used as a marker of a specialized face-processing mechanism. Four brown capuchin monkeys (Cebus apella) were tested on their ability to discriminate several classes of facial and non-facial stimuli presented in both their upright and inverted orientations in an oddity task. Results revealed significantly better performance on upright than inverted presentations of capuchin and human face stimuli, but not on chimpanzee faces or automobiles. These data support previous studies in humans and other primates suggesting that the inversion effect occurs for stimuli for which subjects have developed an expertise.     

The body-inversion effect

Researchers argue that faces are recognized via the configuration of their parts. An important behavioral finding supporting this claim is the face-inversion effect, in which inversion impairs recognition of faces more than nonface objects. Until recently, faces were the only class of objects producing the inversion effect for untrained individuals. This study investigated whether the inversion effect extends to human body positions, a class of objects whose exemplars are structurally similar to each other. Three experiments compared the recognition of upright and inverted faces, houses, and body positions using a forced-choice, same/different paradigm. For both reaction time and error data, the recognition of possible human body postures was more affected by inversion than the recognition of houses. Further, the recognition of possible human body postures and recognition of faces showed similar effects of inversion. The inversion effect was diminished for impossible body positions that violated the biomechanical constraints of human bodies. These data suggest that human body positions, like faces, may be processed configurally by untrained viewers.     

Physical and perceptual accuracy of upright and inverted face drawings

This study considers the conception that drawing or copying a face that is vertically inverted will improve the accuracy of the drawing by preventing holistic interference. We used a novel parameterized face space both for generating face stimuli and for measuring the physical accuracy of drawings. One group of participants (the artists) were asked to draw 16 parameterized faces (eight upright and eight inverted). We computed two physical measures of accuracy by comparing the face-space representation of each drawing to the original face. A second and third group of participants (the raters) compared the similarity between each original face and each pair of drawings of that face (one upright and one inverted per artist). For the second group, all faces were presented upright; for the third group, all faces were presented inverted. Our results showed that upright drawings were more accurate than inverted drawings, both in terms of the physical face-space measure and in terms of the perceptual judgments for both orientations. Our data suggest that holistic processing may aid rather than hinder face drawing accuracy.     

Privileged detection of conspecifics: Evidence from inversion effects during continuous flash suppression

The rapid visual detection of other people in our environment is an important first step in social cognition. Here we provide evidence for selective sensitivity of the human visual system to upright depictions of conspecifics. In a series of seven experiments, we assessed the impact of stimulus inversion on the detection of person silhouettes, headless bodies, faces and other objects from a wide range of animate and inanimate control categories. We used continuous flash suppression (CFS), a variant of binocular rivalry, to render stimuli invisible at the beginning of each trial and measured the time upright and inverted stimuli needed to overcome such interocular suppression. Inversion strongly interfered with access to awareness for human faces, headless human bodies, person silhouettes, and even highly variable body postures, while suppression durations for control objects were not (inanimate objects) or only mildly (animal faces and bodies) affected by inversion. Furthermore, inversion effects were eliminated when the normal body configuration was distorted. The absence of strong inversion effects in a binocular control condition not involving interocular suppression suggests that non-conscious mechanisms mediated the effect of inversion on body and face detection during CFS. These results indicate that perceptual mechanisms that govern access to visual awareness are highly sensitive to the presence of conspecifics.     

When feature information comes first! Early processing of inverted faces

We investigated the early stages of face recognition and the role of featural and holistic face information. We exploited the fact that, on inversion, the alienating disorientation of the eyes and mouth in thatcherised faces is hardly detectable. This effect allows featural and holistic information to be dissociated and was used to test specific face-processing hypotheses. In inverted thatcherised faces, the cardinal features are already correctly oriented, whereas in undistorted faces, the whole Gestalt is coherent but all information is disoriented. Experiment 1 and experiment 3 revealed that, for inverted faces, featural information processing precedes holistic information. Moreover, the processing of contextual information is necessary to process local featural information within a short presentation time (26 ms). Furthermore, for upright faces, holistic information seems to be available faster than for inverted faces (experiment 2). These differences in processing inverted and upright faces presumably cause the differential importance of featural and holistic information for inverted and upright faces.     

Impaired configural body processing in anorexia nervosa: Evidence from the body inversion effect

Patients with anorexia nervosa (AN) suffer from severe disturbances of body perception. It is unclear, however, whether such disturbances are linked to specific alterations in the processing of body configurations with respect to the local processing of body part details. Here, we compared a consecutive sample of 12 AN patients with a group of 12 age‐, gender‐ and education‐matched controls using an inversion effect paradigm requiring the visual discrimination of upright and inverted pictures of whole bodies, faces and objects. The AN patients presented selective deficits in the discrimination of upright body stimuli, which requires configural processing. Conversely, patients and controls showed comparable abilities in the discrimination of inverted bodies, which involves only detail‐based processing, and in the discrimination of both upright and inverted faces and objects. Importantly, the body inversion effect negatively correlated with the persistence scores at the Temperament and Character Inventory, which evaluates increased tendency to convert a signal of punishment into a signal of reinforcement. These results suggest that the deficits of configural processing in AN patients may be associated with their obsessive worries about body appearance and to the excessive attention to details that characterizes their general perceptual style.     

The influence of divided attention on holistic face perception.

There is evidence that upright, but not inverted, faces are encoded holistically. The holistic coding of faces was examined in four experiments by manipulating the attention allocated to target faces. In Experiment 1, participants in a divided attention condition were asked to match two upright flanker faces while encoding a centrally presented upright target face. Although holistic coding was evident in the full attention conditions, dividing attention disrupted holistic coding of target faces. In Experiment 2, we found that while matching upright flanker faces disrupted holistic coding, matching inverted flanker faces did not. Experiment 3 demonstrated that the differential effects of flanker orientation were not due to participants taking longer to match upright, than inverted, flanker faces. In Experiment 4, we found that matching fractured faces had an intermediate effect to that of matching upright and inverted flankers, on the holistic coding of the target faces. The findings emphasize the differences in processing of upright, fractured and inverted faces and suggest that there are limitations in the number of faces that can be holistically coded in a brief time.     

Specificity of face processing without awareness

The recognition memory for inverted faces is especially difficult when compared with that for non-face stimuli. This face inversion effect has often been used as a marker of face-specific holistic processing. However, whether face processing without awareness is still specific remains unknown. The present study addressed this issue by examining the face inversion effect with the technique of binocular rivalry. Results showed that invisible upright faces could break suppression faster than invisible inverted faces. Nevertheless, no difference was found for invisible upright houses and invisible inverted houses. This suggested that face processing without awareness is still specific. Some face-specific information can be processed by high-level brain areas even when that information is invisible.     

The effect of inversion on the encoding of normal and “thatcherized” faces

In the “Thatcher illusion” a face, in which the eyes and mouth are inverted relative to the rest of the face, looks grotesque when shown upright but not when inverted. In four experiments we investigated the contribution of local and global processing to this illusion in normal observers. We examined inversion effects (i.e., better performance for upright than for inverted faces) in a task requiring discrimination of whether faces were or were not “thatcherized”. Observers made same/different judgements to isolated face parts (Experiments 1-2) and to whole faces (Experiments 3-4). Face pairs had the same or different identity, allowing for different process- ing strategies using feature-based or configural information, respectively. In Experiment 1, feature-based matching of same-person face parts yielded only a small inversion effect for normal face parts. However, when feature-based matching was prevented by using the face parts of different people on all trials (Experiment 2) an inversion effect occurred for normal but not for thatcherized parts. In Experiments 3 and 4, inversion effects occurred with normal but not with thatcherized whole faces, on both same- and different-person matching tasks. This suggests that a common configural strategy was used with whole (normal) faces. Face context facilitated attention to misoriented parts in same-person but not in different-person matching. The results indicate that (1) face inversion disrupts local configural processing, but not the processing of image features, and (2) thatcherization disrupts local configural processing in upright faces.     

Remembering parts and wholes: Configural processing in face recollection

In two experiments, we examined the role of configural processing on states of awareness in face recognition. Configural processing was manipulated by presenting upright and inverted faces during study and test. After studying facial photographs of humans (Experiment 1) and horses (Experiment 2), participants provided a remember/know judgement for each recognised test face. In both experiments, disrupted configural processing had selective effects on states of awareness, so that inversion reduced remember, but not know, responses. Experiment 2 revealed disproportionate inversion effects in that the difference in remember responses between upright and inverted items was significant for human faces but not horses. These findings suggest that configural processing facilitates explicit recollection by providing a distinctive combination of nonsalient event attributes.     

Recognition of emotion from inverted schematic drawings of faces

The present study investigated whether facial expressions of emotion are recognized holistically, i.e., all at once as an entire unit, as faces are or featurally as other nonface stimuli. Evidence for holistic processing of faces comes from a reliable decrement in recognition performance when faces are presented inverted rather than upright. If emotion is recognized holistically, then recognition of facial expressions of emotion should be impaired by inversion. To test this, participants were shown schematic drawings of faces showing one of six emotions (surprise, sadness, anger, happiness, disgust, and fear) in either an upright or inverted orientation and were asked to indicate the emotion depicted. Participants were more accurate in the upright than in the inverted orientation, providing evidence in support of holistic recognition of facial emotion. Because recognition of facial expressions of emotion is important in social relationships, this research may have implications for treatment of some social disorders.     

Does Thompson's Thatcher Effect reflect a face-specific mechanism?

The Thatcher Illusion or Thatcher Effect (TE--Thompson 1980, Perception 9 483-484) reflects the difficulty in perceiving the local inversion of parts when the whole object, generally a face, is globally inverted. We tested the generality of the TE with a range of faces and nonface objects, and observed the TE with many non-face categories including cars, buildings, bikes, and letter strings. In terms of magnitude, the face TE is not exceptionally large compared to other object categories, and the magnitude of the TE can be predicted by performance on this task for upright stimuli, regardless of whether the object is a face or not. We did not observe evidence for a unique mechanism contributing to the TE for faces. We discuss factors that influence the magnitude of the TE, some common across domains and others more specific to a particular category.