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.     

Developmental trajectory of neural specialization for letter and number visual processing

Adult neuroimaging studies have demonstrated dissociable neural activation patterns in the visual cortex in response to letters (Latin alphabet) and numbers (Arabic numerals), which suggest a strong experiential influence of reading and mathematics on the human visual system. Here, developmental trajectories in the event‐related potential (ERP) patterns evoked by visual processing of letters, numbers, and false fonts were examined in four different age groups (7‐, 10‐, 15‐year‐olds, and young adults). The 15‐year‐olds and adults showed greater neural sensitivity to letters over numbers in the left visual cortex and the reverse pattern in the right visual cortex, extending previous findings in adults to teenagers. In marked contrast, 7‐ and 10‐year‐olds did not show this dissociable neural pattern. Furthermore, the contrast of familiar stimuli (letters or numbers) versus unfamiliar ones (false fonts) showed stark ERP differences between the younger (7‐ and 10‐year‐olds) and the older (15‐year‐olds and adults) participants. These results suggest that both coarse (familiar versus unfamiliar) and fine (letters versus numbers) tuning for letters and numbers continue throughout childhood and early adolescence, demonstrating a profound impact of uniquely human cultural inventions on visual cognition and its development.     

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。     

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.     

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.     

Letter processing in the visual system: different activation patterns for single letters and strings

One would expect that a lifetime of experience recognizing letters would have an important influence on the visual system. Surprisingly, there is limited evidence of a specific neural response to letters over visual control stimuli. We measured brain activation during a sequential matching task using isolated characters (Roman letters, digits, and Chinese characters) and strings of characters. We localized the visual word form area (VWFA) by contrasting the response to pseudowords against that for letter strings, but this region did not show any other sign of visual specialization for letters. In addition, a left fusiform area posterior to the VWFA was selective for letter strings, whereas a more anterior left fusiform region showed selectivity for single letters. The results of different analyses using both large regions of interest and inspections of individual patterns of response reveal a dissociation between selectivity for letter strings and selectivity for single letters. The results suggest that reading experience fine-tunes visual representations at different levels of processing. An important conclusion is that the processing of nonpronounceable letter strings cannot be assumed to be equivalent to single-letter perception.     

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.     

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.     

Visual experiences during letter production contribute to the development of the neural systems supporting letter perception

Letter production through handwriting creates visual experiences that may be important for the development of visual letter perception. We sought to better understand the neural responses to different visual percepts created during handwriting at different levels of experience. Three groups of participants, younger children, older children, and adults, ranging in age from 4.5 to 22 years old, were presented with dynamic and static presentations of their own handwritten letters, static presentations of an age‐matched control's handwritten letters, and typeface letters during fMRI. First, data from each group were analyzed through a series of contrasts designed to highlight neural systems that were most sensitive to each visual experience in each age group. We found that younger children recruited ventral‐temporal cortex during perception and this response was associated with the variability present in handwritten forms. Older children and adults also recruited ventral‐temporal cortex; this response, however, was significant for typed letter forms but not variability. The adult response to typed letters was more distributed than in the children, including ventral‐temporal, parietal, and frontal motor cortices. The adult response was also significant for one's own handwritten letters in left parietal cortex. Second, we compared responses among age groups. Compared to older children, younger children demonstrated a greater fusiform response associated with handwritten form variability. When compared to adults, younger children demonstrated a greater response to this variability in left parietal cortex. Our results suggest that the visual perception of the variability present in handwritten forms that occurs during handwriting may contribute to developmental changes in the neural systems that support letter perception.     

Neural adaptation across viewpoint and exemplar in fusiform cortex

The visual system has the remarkable ability to generalize across different viewpoints and exemplars to recognize abstract categories of objects, and to discriminate between different viewpoints and exemplars to recognize specific instances of particular objects. Behavioral experiments indicate the critical role of the right hemisphere in specific-viewpoint and -exemplar visual form processing and the left hemisphere in abstract-viewpoint and -exemplar visual form processing. Neuroimaging studies indicate the role of fusiform cortex in these processes, however results conflict in their support of the behavioral findings. We investigated this inconsistency in the present study by examining adaptation across viewpoint and exemplar changes in the functionally defined fusiform face area (FFA) and in fusiform regions exhibiting adaptation. Subjects were adapted to particular views of common objects and then tested with objects appearing in four critical conditions: same-exemplar, same-viewpoint adapted, same-exemplar, different-viewpoint adapted, different-exemplar adapted, and not adapted. In line with previous results, the FFA demonstrated a release from neural adaptation for repeated different viewpoints and exemplars of an object. In contrast to previous work, a (non-FFA) right medial fusiform area also demonstrated a release from neural adaptation for repeated different viewpoints and exemplars of an object. Finally, a left lateral fusiform area demonstrated neural adaptation for repeated different viewpoints, but not exemplars, of an object. Test-phase task demands did not affect adaptation in these regions. Together, results suggest that dissociable neural subsystems in fusiform cortex support the specific identification of a particular object and the abstract recognition of that object observed from a different viewpoint. In addition, results suggest that areas within fusiform cortex do not support abstract recognition of different exemplars of objects within a category.     

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.     

Neural correlates of subliminal and supraliminal letter processing--an event-related fMRI study

One problem of interpreting research on subconscious processing is the possibility that participants are weakly conscious of the stimuli. Here, we compared the fMRI BOLD response in healthy adults to clearly visible single letters (supraliminal presentation) with the response to letters presented in the absence of any behavioural evidence of visibility (subliminal presentation). No letter catch trials served as a control condition. Forced-choice responses did not differ from chance when letter-to-background contrast was low, whereas they were almost 100% correct when contrast was high. A comparison of fMRI BOLD signals for supraliminal and subliminal letters with the control trials revealed a signal increase in left BA 37 (fusiform gyrus). Comparison of supraliminal with subliminal letters showed a significant increase in the right inferior frontal gyrus (BA 44, partly extending to BA 9 and BA 45, as well as BA 46). Finally, a comparison of subliminal with supraliminal letters showed increases in the left middle temporal gyrus (BA 21) and the right extrastriate cortex (BA 19).     

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.     

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.     

识别名人与陌生人面孔的功能磁共振成像

探讨名人与陌生人身份判断的脑功能。6名被试参加实验,运用功能磁共振技术采集数据,AFNI软件进行统计处理。实验结果显示,名人面孔身份识别主要激活了额颞区和梭状回等,陌生人面孔识别主要激活了脑岛和梭状回。实验表明,熟悉名人面孔与陌生人面孔识别有其不同的区域;都有梭状回的视觉加工,但是激活的程度不同。     

Lexical processes and eye movements in neglect dyslexia

Neglect dyslexia is a disturbance in the allocation of spatial attention over a letter string following unilateral brain damage. Patients with this condition may fail to read letters on the contralesional side of an orthographic string. In some of these cases, reading is better with words than with non-words. This word superiority effect has received a variety of explanations that differ, among other things, with regard to the spatial distribution of attention across the letter string during reading. The primary goal of the present study was to explore the interaction between attention and lexical processes by recording eye movements in a patient (F.C.) with severe left neglect dyslexia who was required to read isolated word and non-word stimuli of various length. F.C.'s ocular exploration of orthographic stimuli was highly sensitive to the lexical status of the letter string. We found that: (1) the location to which F.C. directed his initial saccade (obtained approximately 230 ms post-stimulus onset) differed between word and non-word stimuli; (2) the patient spent a greater amount of time fixating the contralesional side of word than non-word strings. Moreover, we also found that F.C. failed to identify the left letters of a string despite having fixated them, thus showing a clear dissociation between eye movement responses and conscious access to orthographic stimuli. Our data suggest the existence of multiple interactions between lexical, attentional and eye movement systems that occur from very initial stages of visual word recognition.     

Not all visual features are created equal: early processing in letter and word recognition

In four experiments, we investigated the effect of deleting specific features of letters on letter and word recognition in the context of reading aloud. Experiments 1 and 2 assessed the relative importance of vertices versus midsegments in letter recognition. Experiments 3 and 4 tested the relative importance of vertices versus midsegments in word recognition. The results demonstrate that deleting vertices is more detrimental to letter and word identification than is deleting midsegments of letters. These results converge with those of previous research on the role of vertices in object identification. Theoretical implications for early processing in reading are noted.     

Behavioral and fMRI evidence that cognitive ability modulates the effect of semantic context on speech intelligibility

Text cues facilitate the perception of spoken sentences to which they are semantically related (Zekveld, Rudner, et al., 2011). In this study, semantically related and unrelated cues preceding sentences evoked more activation in middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) than nonword cues, regardless of acoustic quality (speech in noise or speech in quiet). Larger verbal working memory (WM) capacity (reading span) was associated with greater intelligibility benefit obtained from related cues, with less speech-related activation in the left superior temporal gyrus and left anterior IFG, and with more activation in right medial frontal cortex for related versus unrelated cues. Better ability to comprehend masked text was associated with greater ability to disregard unrelated cues, and with more activation in left angular gyrus (AG). We conclude that individual differences in cognitive abilities are related to activation in a speech-sensitive network including left MTG, IFG and AG during cued speech perception.