Language-specific tuning of visual cortex? Functional properties of the Visual Word Form Area (original) (raw)
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Brain, 2000
A standard model of word reading postulates that visual information is initially processed by occipitotemporal areas contralateral to the stimulated hemifield, from whence it is subsequently transferred to the visual word form (VWF) system, a left inferior temporal region specifically devoted to the processing of letter strings. For stimuli displayed in the left visual field, this transfer proceeds from the right to the left hemisphere through the posterior portion of the corpus callosum. In order to characterize the spatial and temporal organization of these processes, reading tasks with split-field presentation were performed by five control subjects and by two patients suffering from left hemialexia following posterior callosal lesions. The subjects' responses were studied using behavioural measures and functional brain imaging techniques, providing both high spatial resolution (functional MRI, fMRI) and high temporal resolution (high-density event-related potentials, ERPs). Early visual processing was revealed as activations contralateral to stimulation, located by fMRI in the inferior occipitotemporal region and presumably coincident with Abbreviations: BOLD ϭ blood oxygenation level-dependent; ERPs ϭ event-related potentials; fMRI ϭ functional MRI; LVF ϭ left visual field; RVF ϭ right visual field; SPM ϭ statistical parametric mapping; VWF ϭ visual word form
Brain activations during letter-by-letter reading: A follow-up study
Neuropsychologia, 2005
Lesions affecting the ventral cortex of the left temporal lobe commonly yield a selective reading impairment known as pure alexia. It is thought to result from the disruption or deafferentation of the Visual Word Form Area (VWFA), a region in the left lateral occipitotemporal sulcus activated whenever normal subjects are viewing alphabetic strings. Most pure alexic patients retain the ability to identify single letters, and develop a strategy of letter-by-letter (LBL) reading. We recently studied fMRI activations in LBL readers and clarified the underlying mechanisms. However, LBL reading is a dynamic process which may improve over months or years of practice, although the cerebral bases of this continuing improvement are currently unknown. We had the opportunity to run the same behavioural testing and fMRI experiment a second time in an alexic patient, 8 months after collecting the data reported by Cohen et al. ]. We analyze the changes that occurred over this period in the pattern of reading-related activations, while the patient's LBL reading improved. The activation level decreased in most of the overall network between the two sessions. This general trend contrasted with a focal increase restricted to specific left frontal and parietal areas. When studying the contrast between words and consonant strings, which may be taken as a correlate of LBL reading, we also found a general decrease, except for similar left frontal and parietal regions, which showed a significant increase. We suggest that the pattern of evolution fits with the minimal hypothesis of normal strategic abilities and skill learning, associated with perceptual tuning in right-hemispheric structures able to substitute the disrupted VWFA.
Brain and Language, 2011
Right visual field advantage Magnetoencephalography MEG Visual word form area a b s t r a c t Right-handed participants respond more quickly and more accurately to written words presented in the right visual field (RVF) than in the left visual field (LVF). Previous attempts to identify the neural basis of the RVF advantage have had limited success. Experiment 1 was a behavioral study of lateralized word naming which established that the words later used in Experiment 2 showed a reliable RVF advantage which persisted over multiple repetitions. In Experiment 2, the same words were interleaved with scrambled words and presented in the LVF and RVF to right-handed participants seated in an MEG scanner. Participants read the real words silently and responded ''pattern" covertly to the scrambled words. A beamformer analysis created statistical maps of changes in oscillatory power within the brain. Those whole-brain maps revealed activation of the reading network by both LVF and RVF words. Virtual electrode analyses used the same beamforming method to reconstruct the responses to real and scrambled words in three regions of interest in both hemispheres. The middle occipital gyri showed faster and stronger responses to contralateral than to ipsilateral stimuli, with evidence of asymmetric channeling of information into the left hemisphere. The left mid fusiform gyrus at the site of the 'visual word form area' responded more strongly to RVF than to LVF words. Activity in speech-motor cortex was lateralized to the left hemisphere, and stronger to RVF than LVF words, which is interpreted as representing the proximal cause of the RVF advantage for naming written words.
Word and non-word reading: What role for the Visual Word Form Area
Neuroimage, 2005
The putative role of the so-called Visual Word Form Area (VWFA) during reading remains under debate. For some authors, this region is specifically involved in a pre-lexical processing of words and pseudowords, whereas such specificity is challenged by others given the VWFA involvement during both non-word reading and word listening. Here, we further investigated this issue, measuring BOLD variations and their lateralization with fMRI during word and nonword reading, in order to evaluate the lexicality effect, and during reading and listening of words, in order to evaluate the impact of stimulus delivery modality on word processing networks. Region of interest (ROI) analysis was first performed in three target areas: 1-VWFA as defined by a meta-analysis of the word reading literature, 2-a middle temporal area (T2) found co-activated by both word reading and listening, 3-an inferior occipital area (OI) belonging to the unimodal visual cortex of the inferior occipital gyrus. VWFA activity was found not different between word and non-word reading but was more leftward lateralized during word reading due to a reduction of activity in the VWFA right counterpart. A similar larger leftward lateralization during word reading was also uncovered in the T2 ROI but was related to a larger left side activity. Such a lexicality effect was not observed in the OI ROI. By contrast, BOLD increases during listening were restricted to the left VWFA and T2 ROIs. Voxel-based analysis (SPM99) showed that semantic areas were more active during word than non-word reading and co-activated by both reading and listening, exhibiting a left lateralized activity in all tasks. These results indicate that the left VWFA would be the place where visual and verbal representations bind under the control of left semantic areas. D
Attention to single letters activates left extrastriate cortex
Neuroimage, 2004
Brain imaging studies examining the component processes of reading using words, non-words, and letter strings frequently report task-related activity in the left extrastriate cortex. Processing of these linguistic materials involves varying degrees of semantic, phonological, and orthographic analysis that are sensitive to individual differences in reading skill and history. In contrast, single letter processing becomes automatized early in life and is not modulated by later linguistic experience to the same degree as are words. In this study, skilled readers attended to different aspects (single letters, symbols, and colors) of an identical stimulus set during separate sessions of functional magnetic resonance imaging (fMRI). Whereas activation in some portions of ventral extrastriate cortex was shared by attention to both alphabetic and non-alphabetic features, a letter-specific area was identified in a portion of left extrastriate cortex (Brodmann's Area 37), lateral to the visual word form area. Our results demonstrate that while minimizing activity related to word-level lexical properties, cortical responses to letter recognition can be isolated from figural and color characteristics of simple stimuli. The practical utility of this finding is discussed in terms of early identification of reading disability.
The left fusiform area is affected by written frequency of words
Neuropsychologia, 2008
The recent neuroimaging literature gives conflicting evidence about whether the left fusiform gyrus (FG) might recognize words as unitary visual objects. The sensitivity of the left FG to word frequency might provide a neural basis for the orthographic input lexicon theorized by reading models . Surface dyslexia: Cognitive and neuropsychological studies of phonological reading. London: Lawrence Erlbaum]. The goal of this study was to investigate the time course and neural correlates of word processing in right-handed readers engaged in an orthographic decision task. Three hundred and twenty Italian words of high and low written frequency and 320 non-derived legal pseudowords were presented for 250 ms in the central visual field. ERPs were recorded from 128 scalp sites in 10 Italian University students. Behavioural data showed a word superiority effect, with faster RTs to words than pseudo-words. Left occipito/temporal N2 (240 ms) was greater to high-frequency than lowfrequency words and pseudo-words. According to the swLORETA inverse solution, the underlying neural source of this effect was located in the left fusiform gyrus of the occipital lobe (X = −29, Y = −66, Z = −10, BA19) and the right superior temporal gyrus (X = 51, Y = 6, Z = −5, BA22), which are probably involved in word recognition and semantic representation, respectively. Later frontal ERP components, LPN (300-350) and P3 (400-500), also showed strong lexical sensitivity, thus suggesting implicit semantic processes. The results shed some light on the possible neural substrate of visual reading disabilities such as developmental surface dyslexia or pure alexia.
2012
Although the left posterior occipitotemporal sulcus (pOTS) has been called a visual word form area, debate persists over the selectivity of this region for reading relative to general nonorthographic visual object processing. We used high-resolution functional magnetic resonance imaging to study left pOTS responses to combinatorial orthographic and object shape information. Participants performed naming and visual discrimination tasks designed to encourage or suppress phonological encoding. During the naming task, all participants showed subregions within left pOTS that were more sensitive to combinatorial orthographic information than to object information. This difference disappeared, however, when phonological processing demands were removed. Responses were stronger to pseudowords than to words, but this effect also disappeared when phonological processing demands were removed. Subregions within the left pOTS are preferentially activated when visual input must be mapped to a phonological representation (i.e., a name) and particularly when component parts of the visual input must be mapped to corresponding phonological elements (consonant or vowel phonemes). Results indicate a specialized role for subregions within the left pOTS in the isomorphic mapping of familiar combinatorial visual patterns to phonological forms. This process distinguishes reading from picture naming and accounts for a wide range of previously reported stimulus and task effects in left pOTS.
Models of the ''visual word form system'' postulate that a left occipitotemporal region implements the automatic visual word recognition required for efficient reading. This theory was assessed in a patient in whom reading was explored with behavioral measures, fMRI, and intracranial local field potentials. Prior to surgery, when reading was normal, fMRI revealed a normal mosaic of ventral visual selectivity for words, faces, houses, and tools. Intracranial recordings demonstrated that the left occipitotemporal cortex responded with a short latency to conscious but also to subliminal words. Surgery removed a small portion of wordresponsive occipitotemporal cortex overlapping with the word-specific fMRI activation. The patient developed a marked reading deficit, while recognition of other visual categories remained intact. Furthermore, in the post-surgery fMRI map of visual cortex, only word-specific activations disappeared. Altogether, these results provide direct evidence for the causal role of the left occipitotemporal cortex in the recognition of visual words. Neuron 192
Neuron, 2006
Models of the ''visual word form system'' postulate that a left occipitotemporal region implements the automatic visual word recognition required for efficient reading. This theory was assessed in a patient in whom reading was explored with behavioral measures, fMRI, and intracranial local field potentials. Prior to surgery, when reading was normal, fMRI revealed a normal mosaic of ventral visual selectivity for words, faces, houses, and tools. Intracranial recordings demonstrated that the left occipitotemporal cortex responded with a short latency to conscious but also to subliminal words. Surgery removed a small portion of wordresponsive occipitotemporal cortex overlapping with the word-specific fMRI activation. The patient developed a marked reading deficit, while recognition of other visual categories remained intact. Furthermore, in the post-surgery fMRI map of visual cortex, only word-specific activations disappeared. Altogether, these results provide direct evidence for the causal role of the left occipitotemporal cortex in the recognition of visual words. Neuron 192
The Neural Basis of Visual Word Form Processing: A Multivariate Investigation
Cerebral Cortex, 2013
Current research on the neurobiological bases of reading points to the privileged role of a ventral cortical network in visual word processing. However, the properties of this network and, in particular, its selectivity for orthographic stimuli such as words and pseudowords remain topics of significant debate. Here, we approached this issue from a novel perspective by applying pattern-based analyses to functional magnetic resonance imaging data. Specifically, we examined whether, where and how, orthographic stimuli elicit distinct patterns of activation in the human cortex. First, at the category level, multivariate mapping found extensive sensitivity throughout the ventral cortex for words relative to false-font strings. Secondly, at the identity level, the multi-voxel pattern classification provided direct evidence that different pseudowords are encoded by distinct neural patterns. Thirdly, a comparison of pseudoword and face identification revealed that both stimulus types exploit common neural resources within the ventral cortical network. These results provide novel evidence regarding the involvement of the left ventral cortex in orthographic stimulus processing and shed light on its selectivity and discriminability profile. In particular, our findings support the existence of sublexical orthographic representations within the left ventral cortex while arguing for the continuity of reading with other visual recognition skills.