TMS to the “occipital face area” affects recognition but not categorization of faces

The human cortical system for face perception is comprised of a network of connected regions including the middle fusiform gyrus (“fusiform face area” or FFA), the inferior occipital cortex (“occipital face area” or OFA), and the superior temporal sulcus. The traditional hierarchical feedforward model of visual processing suggests information flows from early visual cortex to the OFA for initial face feature analysis to higher order regions including the FFA for identity recognition. However, patient data suggest an alternative model. Patients with acquired prosopagnosia, an inability to visually recognize faces, have been documented with lesions to the OFA but who nevertheless show face-selective activation in the FFA. Moreover, their ability to categorize faces remains intact. This suggests that the FFA is not solely responsible for face recognition and the network is not strictly hierarchical, but may be organized in a reverse hierarchical fashion. We used transcranial magnetic stimulation (TMS) to temporarily disrupt processing in the OFA in neurologically-intact individuals and found participants’ ability to categorize intact versus scrambled faces was unaffected, however face identity discrimination was significantly impaired. This suggests that face categorization but not recognition can occur without the “earlier” OFA being online and indicates that “lower level” face category processing may be assumed by other intact face network regions such as the FFA. These results are consistent with the patient data and support a non-hierarchical, global-to-local model with re-entrant connections between the OFA and other face processing areas.     

TMS to the “occipital face area” affects recognition but not categorization of faces

The human cortical system for face perception is comprised of a network of connected regions including the middle fusiform gyrus (“fusiform face area” or FFA), the inferior occipital cortex (“occipital face area” or OFA), and the superior temporal sulcus. The traditional hierarchical feedforward model of visual processing suggests information flows from early visual cortex to the OFA for initial face feature analysis to higher order regions including the FFA for identity recognition. However, patient data suggest an alternative model. Patients with acquired prosopagnosia, an inability to visually recognize faces, have been documented with lesions to the OFA but who nevertheless show face-selective activation in the FFA. Moreover, their ability to categorize faces remains intact. This suggests that the FFA is not solely responsible for face recognition and the network is not strictly hierarchical, but may be organized in a reverse hierarchical fashion. We used transcranial magnetic stimulation (TMS) to temporarily disrupt processing in the OFA in neurologically-intact individuals and found participants’ ability to categorize intact versus scrambled faces was unaffected, however face identity discrimination was significantly impaired. This suggests that face categorization but not recognition can occur without the “earlier” OFA being online and indicates that “lower level” face category processing may be assumed by other intact face network regions such as the FFA. These results are consistent with the patient data and support a non-hierarchical, global-to-local model with re-entrant connections between the OFA and other face processing areas.     

面孔认知的四个层面及FFA在其中的作用

面孔具有明显不同于其他刺激物的特点,面孔认知的目的也因此与其他物体认知的目的大相径庭。依据面孔认知的目的,本文将面孔认知划分为四个层面：将面孔区别于一般物体的第一层面,对面孔物理属性进行识别的第二层面,对面孔的生物属性进行识别的第三层面和对面孔的社会属性进行识别的第四层面。FFA是面孔加工的一个重要脑区,通过论述它对面孔认知各层面的作用,FFA在面孔加工中的作用被进一步明确。     

双侧梭状回面孔区在面孔加工中的功能分工与协作

梭状回面孔区(fusiform face area,FFA)是视觉皮层上专门加工面孔的区域。然而,双侧FFA在面孔加工中的功能分工与协作还存在争议。在特异性刺激的加工上,右侧FFA主要负责人类面孔类别的知觉,而左侧FFA的功能与面孔精细特征的感知有关;在皮层可塑性上,右侧FFA主要参与青少年的社会适应学习,而左侧FFA负责成年人的知觉学习;在面孔网络中,二者与不同区域的连接用以适应不同的认知需求;他们之间的有向协作具有任务特定性。未来研究需要回答三个问题:左侧FFA的可塑性程度及这一可塑性是否是认知特定的、左侧FFA及其形成的网络连接的认知意义,双侧FFA在面孔网络中的连接有向性等问题。     

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.     

Neural perspectives on the other-race effect

Psychological studies have long shown that human memory is superior for faces of our own-race than for faces of other-races. In this paper, we review neural studies of own- versus other-race face processing. These studies divide naturally into those focused on socioaffective aspects of the other-race effect and those directed at high-level visual processing differences. The socioaffective studies consider how subconscious bias and emotional responses affect brain areas such as the amygdala, anterior cingulate cortex, and parahippocampal gyrus. The visual studies focus on face-selective areas in the ventral stream, such as the fusiform face area (FFA). In both cases, factors such as experience, familiarity, social/emotional responses, cultural learning, and bias modulate the patterns of neural activity elicited in response to own- and other-race faces.     

Perceiving age and gender in unfamiliar faces: an fMRI study on face categorization

Efficient processing of unfamiliar faces typically involves their categorization (e.g., into old vs. young or male vs. female). However, age and gender categorization may pose different perceptual demands. In the present study, we employed functional magnetic resonance imaging (fMRI) to compare the activity evoked during age vs. gender categorization of unfamiliar faces. In different blocks, participants performed age and gender classifications for old or young unfamiliar faces (50% female respectively). Both tasks elicited activations in the bilateral fusiform gyri (fusiform face area, FFA) and bilateral inferior occipital gyri (occipital face area, OFA). Importantly, the same stimuli elicited enhanced activation during gender as compared to age categorization. This enhancement was significant in the right FFA and the left OFA, and may be related to increased configural processing. Our findings replicate and extend recent work, and shows that the activation of core components of the face processing network is strongly dependent on task demands.     

Visual category-selectivity for faces, places and objects emerges along different developmental trajectories

The organization of category-selective regions in ventral visual cortex is well characterized in human adults. We investigated a crucial, previously unaddressed, question about how this organization emerges developmentally. We contrasted the developmental trajectories for face-, object-, and place-selective activation in the ventral visual cortex in children, adolescents, and adults. Although children demonstrated adult-like organization in object- and place-related cortex, as a group they failed to show consistent face-selective activation in classical face regions. The lack of a consistent neural signature for faces was attributable to (1) reduced face-selectivity and extent of activation within the regions that will become the FFA, OFA, and STS in adults, and (2) smaller volumes and considerable variability in the locus of face-selective activation in individual children. In contrast, adolescents showed an adult-like pattern of face-selective activation, although it was more right-lateralized. These findings reveal critical age-related differences in the emergence of category-specific functional organization in the visual cortex and support a model of brain development in which specialization emerges from interactions between experience-dependent learning and the maturing brain.     

A defense of the subordinate-level expertise account for the N170 component

A recent paper in this journal reports two event-related potential (ERP) experiments interpreted as supporting the domain specificity of the visual mechanisms implicated in processing faces (Cognition 83 (2002) 1). The authors argue that because a large neurophysiological response to faces (N170) is less influenced by the task than the response to objects, and because the response for human faces extends to ape faces (for which we are not expert), we should reject the hypothesis that the face-sensitivity reflected by the N170 can be accounted for by the subordinate-level expertise model of object recognition (Nature Neuroscience 3 (2000) 764). In this commentary, we question this conclusion based on some of our own ERP work on expert object recognition as well as the work of others.     

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.     

Impaired face recognition does not preclude intact whole face perception

We studied intact and impaired processes in a prosopagnosic patient (RP). In Experiment 1, RP showed an inversion superiority effect with both faces and objects, with better performance when stimuli were presented upside down than in normal upright orientation. In Experiment 2, we studied the effect of face configuration directly by comparing matching performance with normal vs. scrambled faces. RP was worse with normal than with scrambled faces, whereas normal controls showed an advantage of a good face context. In Experiment 3, RP showed interference from external face features on the evaluation of internal face features. These results indicate, first, that although RP is impaired in face recognition and face matching, he does still encode the whole face rather than relying completely on parts-based procedures. Second, RP has a deficit at the level of the configural processes involved in finding subtle differences between individual faces, as his performance is worse when presented with a normal face configuration than with scrambled or inverted faces.     

Domain‐specific development of face memory but not face perception

How does the remarkable human ability for face recognition arise over development? Competing theories have proposed either late maturity (beyond 10 years) or early maturity (before 5 years), but have not distinguished between perceptual and memory aspects of face recognition. Here, we demonstrate a perception–memory dissociation. We compare rate of development for (adult, human) faces versus other social stimuli (bodies), other discrete objects (cars), and other categories processed in discrete brain regions (scenes, bodies), from 5 years to adulthood. For perceptual discrimination, performance improved with age at the same rate for faces and all other categories, indicating no domain‐specific development. In contrast, face memory increased more strongly than non‐face memory, indicating domain‐specific development. The results imply that each theory is partly true: the late maturity theory holds for face memory, and the early maturity theory for face perception.     

Expertise increases the functional overlap between face and object perception

Recent studies indicate that expertise with objects can interfere with face processing. Although competition occurs between faces and objects of expertise, it remains unclear whether this reflects an expertise-specific bottleneck or the fact that objects of expertise grab attention and thereby consume more central resources. We investigated the perceptual costs of expertise by measuring visual thresholds for identifying targets embedded within RSVP sequences presented at varying temporal rates. Car experts and novices searched for face targets among face and car distractors, or watch targets among watch and car distractors. Remarkably, car experts were slower than novices at identifying faces among task-irrelevant cars, yet faster than novices at identifying watches among cars. This suggests that car expertise leads to greater functional overlap between cars and faces while reducing the functional overlap between cars and objects, a result incompatible with the notion of an encapsulated module for exclusive processing of faces.     

Voice recognition and the posterior cingulate: An fMRI study of prosopagnosia

Voices, in addition to faces, enable person identification. Voice recognition has been shown to evoke a distributed network of brain regions that includes, in addition to the superior temporal sulcus (STS), the anterior temporal pole, fusiform face area (FFA), and posterior cingulate gyrus (pCG). Here we report an individual (MS) with acquired prosopagnosia who, despite bilateral damage to much of this network, demonstrates the ability to distinguish voices of several well‐known acquaintances from voices of people that he has never heard before. Functional magnetic resonance imaging (fMRI) revealed that, relative to speech‐modulated noise, voices rated as familiar and unfamiliar by MS elicited enhanced haemodynamic activity in the left angular gyrus, left posterior STS, and posterior midline brain regions, including the retrosplenial cortex and the dorsal pCG. More interestingly, relative to noise and unfamiliar voices, the familiar voices elicited greater haemodynamic activity in the left angular gyrus and medial parietal regions including the dorsal pCG and precuneus. The findings are consistent with theories implicating the pCG in recognizing people who are personally familiar, and furthermore suggest that the pCG region of the voice identification network is able to make functional contributions to voice recognition even though other areas of the network, namely the anterior temporal poles, FFA, and the right parietal lobe, may be compromised.     

Individual differences in FFA activity suggest independent processing at different spatial scales

The brain processes images at different spatial scales, but it is unclear how far into the visual stream different scales remain segregated. Using functional magnetic resonance imaging, we found evidence that BOLD activity in the fusiform face area (FFA) reflects computations based on separate spatial frequency inputs. When subjects perform different tasks (attend location vs. identity; attend whole vs. parts) or the same task with different stimuli (upright or inverted) with high- and low-pass images of cars and faces, individual differences in the FFA in one condition are correlated with those in the other condition. However, FFA activity in response to low-pass stimuli is independent of its response to highpass stimuli. These results suggest that spatial scales are not integrated before the FFA and that processing in this area could support the flexible use of different sources of information present in broad-pass images.     

The improbable simplicity of the fusiform face area

The fusiform face area (FFA) is described as an easily identifiable module on the fusiform gyrus. However, the organization of face-selective regions in ventral temporal cortex (VTC) is more complex than this prevailing view. We highlight methodological factors contributing to these complexities and the extensive variability in how the FFA is identified. We suggest a series of constraints to aid researchers when defining any functionally specialized region with a pleasing realization: anatomy matters.     

Brain correlates of aesthetic expertise: a parametric fMRI study

Several studies have demonstrated that acquired expertise influences aesthetic judgments. In this paradigm we used functional magnetic resonance imaging (fMRI) to study aesthetic judgments of visually presented architectural stimuli and control-stimuli (faces) for a group of architects and a group of non-architects. This design allowed us to test whether level of expertise modulates neural activity in brain areas associated with either perceptual processing, memory, or reward processing. We show that experts and non-experts recruit bilateral medial orbitofrontal cortex (OFC) and subcallosal cingulate gyrus differentially during aesthetic judgment, even in the absence of behavioural aesthetic rating differences between experts and non-experts. By contrast, activity in nucleus accumbens (NAcc) exhibits a differential response profile compared to OFC and subcallosal cingulate gyrus, suggesting a dissociable role between these regions in the reward processing of expertise. Finally, categorical responses (irrespective of aesthetic ratings) resulted in expertise effects in memory-related areas such as hippocampus and precuneus. These results highlight the fact that expertise not only modulates cognitive processing, but also modulates the response in reward related brain areas.     

Differential development of selectivity for faces and bodies in the fusiform gyrus

Viewing faces or bodies activates category‐selective areas of visual cortex, including the fusiform face area (FFA), fusiform body area (FBA), and extrastriate body area (EBA). Here, using fMRI, we investigate the development of these areas, focusing on the right FFA and FBA. Despite the overlap of functionally defined FFA and FBA (54%–75% overlap), we found that these regions developed along different trajectories. With age (7–32 years old), the FFA gradually increased in size and selectivity, and was significantly larger and more face‐selective in adults than children. By contrast, the size and selectivity of the FBA did not correlate with age, and were equivalent in children and adults. Whereas in adults the FFA and FBA were comparable in size, in children the FBA was on average 70% larger than the FFA. These findings suggest that, in children, the fusiform gyrus is predominantly selective for bodies, with commensurate face‐selective responses apparent later in development. Moreover, differences in the development of the FFA and FBA indicate that overlapping functional brain areas, supported by the same anatomical structure, can develop along different trajectories.     

Development of preferences for the human body shape in infancy

Two studies investigated the development of infants' visual preferences for the human body shape. In Study 1, infants of 12, 15 and 18 months were tested in a standard preferential looking experiment, in which they were shown paired line drawings of typical and scrambled bodies. Results indicated that the 18-month-olds had a reliable preference for the scrambled body shapes over typical body shapes, while the younger infants did not show differential responding. In Study 2, 12- and 18-month-olds were tested with the same procedure, except that the typical and scrambled body stimuli were photographic images. The results of Study 2 again indicated that only the 18-month-olds had a reliable preference for the scrambled body shapes. This finding contrasts sharply with infants' precocious preferences for human faces, suggesting that infants' learning about human faces and human bodies follow different developmental trajectories.