Word selectivity in high-level visual cortex and reading skill.
ABSTRACT: Word-selective neural responses in human ventral occipito-temporal cortex (VOTC) emerge as children learn to read, creating a visual word form area (VWFA) in the literate brain. It has been suggested that the VWFA arises through competition between pre-existing selectivity for other stimulus categories, changing the topography of VOTC to support rapid word recognition. Here, we hypothesized that competition between words and objects would be resolved as children acquire reading skill. Using functional magnetic resonance imaging (fMRI), we examined the relationship between responses to words and objects in VOTC in two ways. First, we defined the VWFA using a words?>?objects contrast and found that only skilled readers had a region that responded more to words than objects. Second, we defined the VWFA using a words?>?faces contrast and examined selectivity for words over objects in this region. We found that word selectivity strongly correlated with reading skill, suggesting reading skill-dependent tuning for words. Furthermore, we found that low word selectivity in struggling readers was not due to a lack of response to words, but to a high response to objects. Our results suggest that the fine-tuning of word-selective responses in VOTC is a critical component of skilled reading.
Project description:The visual word form area (VWFA) in the left ventral occipito-temporal (vOT) cortex is key to fluent reading in children and adults. Diminished VWFA activation during print processing tasks is a common finding in subjects with severe reading problems. Here, we report fMRI data from a multicentre study with 140 children in primary school (7.9-12.2 years; 55 children with dyslexia, 73 typical readers, 12 intermediate readers). All performed a semantic task on visually presented words and a matched control task on symbol strings. With this large group of children, including the entire spectrum from severely impaired to highly fluent readers, we aimed to clarify the association of reading fluency and left vOT activation during visual word processing. The results of this study confirm reduced word-sensitive activation within the left vOT in children with dyslexia. Interestingly, the association of reading skills and left vOT activation was especially strong and spatially extended in children with dyslexia. Thus, deficits in basic visual word form processing increase with the severity of reading disability but seem only weakly associated with fluency within the typical reading range suggesting a linear dependence of reading scores with VFWA activation only in the poorest readers.
Project description:Seeing words involves the activity of neural circuitry within a small region in human ventral temporal cortex known as the visual word form area (VWFA). It is widely asserted that VWFA responses, which are essential for skilled reading, do not depend on the visual field position of the writing (position invariant). Such position invariance supports the hypothesis that the VWFA analyzes word forms at an abstract level, far removed from specific stimulus features. Using functional MRI pattern-classification techniques, we show that position information is encoded in the spatial pattern of VWFA responses. A right-hemisphere homolog (rVWFA) shows similarly position-sensitive responses. Furthermore, electrophysiological recordings in the human brain show position-sensitive VWFA response latencies. These findings show that position-sensitive information is present in the neural circuitry that conveys visual word form information to language areas. The presence of position sensitivity in the VWFA has implications for how word forms might be learned and stored within the reading circuitry.
Project description:In human occipitotemporal cortex, category-specific processing for visual objects seems to involve pairs of cortical regions, often with one located in the occipital cortex and another more anteriorly. We investigated whether such an arrangement might be the case for visual word processing. In addition to the Visual Word Form Area (VWFA) located in the occipitotemporal sulcus, we observed that another region in occipital lobe with robust responses to written words (Chinese characters). The current fMRI study investigated this area's precise location and its functional selectivity using Chinese characters and other categories of visual images (cars, chairs and insects). In all the 13 subjects we could identify a cluster of voxels near the inferior occipital gyrus or middle occipital gyrus with stronger responses to Chinese characters than scrambled objects. We tentatively label this area as the Occipital Word Form Sensitive Area (OWA). The OWA's response amplitudes showed similar preference to written words as the VWFA, with the VWFA showing a higher degree of word selectivity, which was confirmed by the result from spatial patterns of response. These results indicate that the OWA, together with the VWFA, are critical parts of the network for processing and representing the category information for word.
Project description:In most environments, the visual system is confronted with many relevant objects simultaneously. That is especially true during reading. However, behavioral data demonstrate that a serial bottleneck prevents recognition of more than one word at a time. We used fMRI to investigate how parallel spatial channels of visual processing converge into a serial bottleneck for word recognition. Participants viewed pairs of words presented simultaneously. We found that retinotopic cortex processed the two words in parallel spatial channels, one in each contralateral hemisphere. Responses were higher for attended than for ignored words but were not reduced when attention was divided. We then analyzed two word-selective regions along the occipitotemporal sulcus (OTS) of both hemispheres (subregions of the visual word form area, VWFA). Unlike retinotopic regions, each word-selective region responded to words on both sides of fixation. Nonetheless, a single region in the left hemisphere (posterior OTS) contained spatial channels for both hemifields that were independently modulated by selective attention. Thus, the left posterior VWFA supports parallel processing of multiple words. In contrast, activity in a more anterior word-selective region in the left hemisphere (mid OTS) was consistent with a single channel, showing (i) limited spatial selectivity, (ii) no effect of spatial attention on mean response amplitudes, and (iii) sensitivity to lexical properties of only one attended word. Therefore, the visual system can process two words in parallel up to a late stage in the ventral stream. The transition to a single channel is consistent with the observed bottleneck in behavior.
Project description:Word learning depends not only on efficient online binding of phonological, orthographic and lexical information, but also on consolidation of new word representations into permanent lexical memory. Work on word learning under a variety of contexts indicates that reading and language skill impact facility of word learning in both print and speech. In addition, recent research finds that individuals with language impairments show deficits in both initial word form learning and in maintaining newly learned representations over time, implicating mechanisms associated with maintenance that may be driven by deficits in overnight consolidation. Although several recent studies have explored the neural bases of overnight consolidation of newly learned words, no extant work has examined individual differences in overnight consolidation at the neural level. The current study addresses this gap in the literature by investigating how individual differences in reading and language skills modulate patterns of neural activation associated with newly learned words following a period of overnight consolidation. Specifically, a community sample of adolescents and young adults with significant variability in reading and oral language (vocabulary) ability were trained on two spoken artificial lexicons, one in the evening on the day before fMRI scanning and one in the morning just prior to scanning. Comparisons of activation between words that were trained and consolidated vs. those that were trained but not consolidated revealed increased cortical activation in a number of language associated and memory associated regions. In addition, individual differences in age, reading skill and vocabulary modulated learning rate in our artificial lexicon learning task and the size of the cortical consolidation effect in the precuneus/posterior cingulate, such that older readers and more skilled readers had larger cortical consolidation effects in this learning-critical region. These findings suggest that age (even into late adolescence) and reading and language skills are important individual differences that affect overnight consolidation of newly learned words. These findings have significant implications for understanding reading and language disorders and should inform pedagogical models.
Project description:Resilient readers are characterized by impaired phonological processing despite skilled text comprehension. We investigated orthographic and semantic processing in resilient readers to examine mechanisms of compensation for poor phonological decoding. Performance on phonological (phoneme deletion, pseudoword reading), orthographic (orthographic choice, orthographic analogy), and semantic (semantic priming, homograph resolution) tasks was compared between resilient, poor and proficient readers. Asymmetry of the planum temporale was investigated in order to determine whether atypical readers showed unusual morphology in this language-relevant region. Resilient readers showed deficits on phonological tasks similar to those shown by poor readers. We obtained no evidence that resilient readers compensate via superior orthographic processing, as they showed neither exceptional orthographic skill nor increased reliance on orthography to guide pronunciation. Resilient readers benefited more than poor or proficient readers from semantic relationships between words and experienced greater difficulty when such relationships were not present. We suggest, therefore, that resilient readers compensate for poor phonological decoding via greater reliance on word meaning relationships. The reading groups did not differ in mean asymmetry of the planum temporale. However, resilient readers showed greater variability in planar asymmetry than proficient readers. Poor readers also showed a trend towards greater variability in planar asymmetry, with more poor readers than proficient readers showing extreme asymmetry. Such increased variability suggests that university students with less reading skill display less well regulated brain anatomy than proficient readers.
Project description:How does education affect cortical organization? All literate adults possess a region specialized for letter strings, the visual word form area (VWFA), within the mosaic of ventral regions involved in processing other visual categories such as objects, places, faces, or body parts. Therefore, the acquisition of literacy may induce a reorientation of cortical maps towards letters at the expense of other categories such as faces. To test this cortical recycling hypothesis, we studied how the visual cortex of individual children changes during the first months of reading acquisition. Ten 6-year-old children were scanned longitudinally 6 or 7 times with functional magnetic resonance imaging (fMRI) before and throughout the first year of school. Subjects were exposed to a variety of pictures (words, numbers, tools, houses, faces, and bodies) while performing an unrelated target-detection task. Behavioral assessment indicated a sharp rise in grapheme-phoneme knowledge and reading speed in the first trimester of school. Concurrently, voxels specific to written words and digits emerged at the VWFA location. The responses to other categories remained largely stable, although right-hemispheric face-related activity increased in proportion to reading scores. Retrospective examination of the VWFA voxels prior to reading acquisition showed that reading encroaches on voxels that are initially weakly specialized for tools and close to but distinct from those responsive to faces. Remarkably, those voxels appear to keep their initial category selectivity while acquiring an additional and stronger responsivity to words. We propose a revised model of the neuronal recycling process in which new visual categories invade weakly specified cortex while leaving previously stabilized cortical responses unchanged.
Project description:A century of neurology and neuroscience shows that seeing words depends on ventral occipital-temporal (VOT) circuitry. Typically, reading is learned using high-contrast line-contour words. We explored whether a specific VOT region, the visual word form area (VWFA), learns to see only these words or recognizes words independent of the specific shape-defining visual features. Word forms were created using atypical features (motion-dots, luminance-dots) whose statistical properties control word-visibility. We measured fMRI responses as word form visibility varied, and we used TMS to interfere with neural processing in specific cortical circuits, while subjects performed a lexical decision task. For all features, VWFA responses increased with word-visibility and correlated with performance. TMS applied to motion-specialized area hMT+ disrupted reading performance for motion-dots, but not line-contours or luminance-dots. A quantitative model describes feature-convergence in the VWFA and relates VWFA responses to behavioral performance. These findings suggest how visual feature-tolerance in the reading network arises through signal convergence from feature-specialized cortical areas.
Project description:Selective attention to speech versus nonspeech signals in complex auditory input could produce top-down modulation of cortical regions previously linked to perception of spoken, and even visual, words. To isolate such top-down attentional effects, we contrasted 2 equally challenging active listening tasks, performed on the same complex auditory stimuli (words overlaid with a series of 3 tones). Instructions required selectively attending to either the speech signals (in service of rhyme judgment) or the melodic signals (tone-triplet matching). Selective attention to speech, relative to attention to melody, was associated with blood oxygenation level-dependent (BOLD) increases during functional magnetic resonance imaging (fMRI) in left inferior frontal gyrus, temporal regions, and the visual word form area (VWFA). Further investigation of the activity in visual regions revealed overall deactivation relative to baseline rest for both attention conditions. Topographic analysis demonstrated that while attending to melody drove deactivation equivalently across all fusiform regions of interest examined, attending to speech produced a regionally specific modulation: deactivation of all fusiform regions, except the VWFA. Results indicate that selective attention to speech can topographically tune extrastriate cortex, leading to increased activity in VWFA relative to surrounding regions, in line with the well-established connectivity between areas related to spoken and visual word perception in skilled readers.
Project description:In typical readers, orthographic knowledge has been shown to influence phonological decisions. In the present study, we used visual rhyme and spelling tasks to investigate the interaction of orthographic and phonological information in adults with varying reading skill. Word pairs that shared both orthography and phonology (e.g., throat/boat), differed in both orthography and phonology (e.g., snow/arm), shared only orthography (e.g., farm/warm), and shared only phonology (e.g., vote/boat) were visually presented to university students who varied in reading ability. For rhyme judgment, participants were slower and less accurate to accept rhyming pairs when words were spelled differently and to reject non-rhyming pairs when words were spelled similarly. Similarly, for spelling judgments, participants were slower and less accurate when indicating that word endings were spelled differently when words rhymed, and slower and less accurate when indicating that words were spelled similarly when words did not rhyme. Crucially, while these effects were clear at the group level, there were large individual differences in the extent to which participants were impacted by conflict. In two separate samples, reading skill was associated with the extent to which orthographic conflict impacted rhyme decisions such that individuals with better nonword reading performance were less impacted by orthographic conflict. Thus, university students with poorer reading skills may differ from their peers either in the reading strategies they use or in the degree to which they automatically access word form information. Understanding these relationships is important for understanding the roles that reading processes play in readers of different skill.