Attention to single letters activates left extrastriate cortex (original) (raw)
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Extrastriate representation of letter discrimination
Neuroimage, 2001
Brain imaging studies examining the component processes of reading using words, non-words, and letter strings frequently report taskrelated 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. D
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.
fMRI correlates of cortical specialization and generalization for letter processing
NeuroImage, 2006
The present study used functional magnetic resonance imaging to examine cortical specialization for letter processing. We assessed whether brain regions that were involved in letter processing exhibited domain-specific and/or mandatory responses, following Fodor's definition of properties of modular systems . The Modularity of Mind. The MIT Press, Cambridge, MA.). Domainspecificity was operationalized as selective, or exclusive, activation for letters relative to object and visual noise processing and a baseline fixation task. Mandatory processing was operationalized as selective activation for letters during both a silent naming and a perceptual matching task. In addition to these operational definitions, other operational definitions of selectivity for letter processing discussed by [Pernet, C., Celsis, P., Demonet, J., 2005. Selective response to letter categorization within the left fusiform gyrus. NeuroImage 28, 738 -744] were applied to the data. Although the left fusiform gyrus showed a specialized response to letters using the definition of selectivity put forth by [Pernet, C., Celsis, P., Demonet, J., 2005. Selective response to letter categorization within the left fusiform gyrus. NeuroImage 28, 738 -744], this region did not exhibit specialization for letters according to our more conservative definition of selectivity. Instead, this region showed equivalent activation by letters and objects in both the naming and matching tasks. Hence, the left fusiform gyrus does not exhibit domain-specific or mandatory processing but may reflect a shared input system for both stimulus types. The left insula and some portions of the left inferior parietal lobule, however, did show a domain-specific response for letter naming but not for letter matching. These regions likely subserve some linguistically oriented cognitive process that is unique to letters, such as grapheme-to-phoneme translation or retrieval of phonological codes for letter names. Hence, cortical specialization for letters emerged in the naming task in some peri-sylvian language related cortices, but not in occipito-temporal cortex. Given that the domain-specific response for letters in left peri-sylvian regions was only present in the naming task, these regions do not process letters in a mandatory fashion, but are instead modulated by the linguistic nature of the task. D
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
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.
Neuroimaging studies of word reading
Proceedings of the National Academy of Sciences, 1998
This review discusses how neuroimaging can contribute to our understanding of a fundamental aspect of skilled reading: the ability to pronounce a visually presented word. One contribution of neuroimaging is that it provides a tool for localizing brain regions that are active during word reading. To assess the extent to which similar results are obtained across studies, a quantitative review of nine neuroimaging investigations of word reading was conducted. Across these studies, the results converge to reveal a set of areas active during word reading, including left-lateralized regions in occipital and occipitotemporal cortex, the left frontal operculum, bilateral regions within the cerebellum, primary motor cortex, and the superior and middle temporal cortex, and medial regions in the supplementary motor area and anterior cingulate. Beyond localization, the challenge is to use neuroimaging as a tool for understanding how reading is accomplished. Central to this challenge will be the integration of neuroimaging results with information from other methodologies. To illustrate this point, this review will highlight the importance of spelling-to-sound consistency in the transformation from orthographic (word form) to phonological (word sound) representations, and then explore results from three neuroimaging studies in which the spelling-to-sound consistency of the stimuli was deliberately varied. Emphasis is placed on the pattern of activation observed within the left frontal cortex, because the results provide an example of the issues and benefits involved in relating neuroimaging results to behavioral results in normal and brain damaged subjects, and to theoretical models of reading.
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
Specialization for written words over objects in the visual cortex
NeuroImage, 2011
The Visual Word Form Area (VWFA) is part of the left ventral visual stream that underlies the invariant identification of visual words. It remains debated whether this region is truly selective for words relative to common objects; why this particular part of the visual system is reproducibly engaged in reading; and whether reading expertise also relies on perceptual learning within earlier visual areas. In this fMRI study we matched written words and line-drawings of objects in luminance, contour length and number of features. We then compared them to control images made by scrambling procedures that kept local features intact. Greater responses to written words than to objects were found not only in the VWFA, but also in areas V1/V2 and V3v/V4. Furthermore, by contrasting stimuli reduced either to line junctions (vertices) or to line midsegments, we showed that the VWFA partially overlaps with regions of ventral visual cortex particularly sensitive to the presence of line junctions that are useful for object recognition. Our results indicate that preferential processing of written words can be observed at multiple levels of the visual system. It is possible that responses in early visual areas might be due to some remaining differences between words and controls not eliminated in the present stimuli. However, our results concur with recent comparisons of literates and illiterates and suggest that these early visual activations reflect the effects of perceptual learning under pressure for fast, parallel processing that is more prominent in reading than other visual cognitive processes.
Lexical processing in deaf readers: an FMRI investigation of reading proficiency
PloS one, 2013
Individuals with significant hearing loss often fail to attain competency in reading orthographic scripts which encode the sound properties of spoken language. Nevertheless, some profoundly deaf individuals do learn to read at age-appropriate levels. The question of what differentiates proficient deaf readers from less-proficient readers is poorly understood but topical, as efforts to develop appropriate and effective interventions are needed. This study uses functional magnetic resonance imaging (fMRI) to examine brain activation in deaf readers (N = 21), comparing proficient (N = 11) and less proficient (N = 10) readers' performance in a widely used test of implicit reading. Proficient deaf readers activated left inferior frontal gyrus and left middle and superior temporal gyrus in a pattern that is consistent with regions reported in hearing readers. In contrast, the less-proficient readers exhibited a pattern of response characterized by inferior and middle frontal lobe acti...