Dysfunctional visual word form processing in progressive alexia.
ABSTRACT: Progressive alexia is an acquired reading deficit caused by degeneration of brain regions that are essential for written word processing. Functional imaging studies have shown that early processing of the visual word form depends on a hierarchical posterior-to-anterior processing stream in occipito-temporal cortex, whereby successive areas code increasingly larger and more complex perceptual attributes of the letter string. A region located in the left lateral occipito-temporal sulcus and adjacent fusiform gyrus shows maximal selectivity for words and has been dubbed the 'visual word form area'. We studied two patients with progressive alexia in order to determine whether their reading deficits were associated with structural and/or functional abnormalities in this visual word form system. Voxel-based morphometry showed left-lateralized occipito-temporal atrophy in both patients, very mild in one, but moderate to severe in the other. The two patients, along with 10 control subjects, were scanned with functional magnetic resonance imaging as they viewed rapidly presented words, false font strings, or a fixation crosshair. This paradigm was optimized to reliably map brain regions involved in orthographic processing in individual subjects. All 10 control subjects showed a posterior-to-anterior gradient of selectivity for words, and all 10 showed a functionally defined visual word form area in the left hemisphere that was activated for words relative to false font strings. In contrast, neither of the two patients with progressive alexia showed any evidence for a selectivity gradient or for word-specific activation of the visual word form area. The patient with mild atrophy showed normal responses to both words and false font strings in the posterior part of the visual word form system, but a failure to develop selectivity for words in the more anterior part of the system. In contrast, the patient with moderate to severe atrophy showed minimal activation of any part of the visual word form system for either words or false font strings. Our results suggest that progressive alexia is associated with a dysfunctional visual word form system, with or without substantial cortical atrophy. Furthermore, these findings demonstrate that functional MRI has the potential to reveal the neural bases of cognitive deficits in neurodegenerative patients at very early stages, in some cases before the development of extensive atrophy.
Project description:Despite decades of cognitive, neuropsychological and neuroimaging studies, it is unclear if letters are identified before word-form encoding during reading, or if letters and their combinations are encoded simultaneously and interactively. Here using functional magnetic resonance imaging, we show that a 'letter-form' area (responding more to consonant strings than false fonts) can be distinguished from an immediately anterior 'visual word-form area' in ventral occipito-temporal cortex (responding more to words than consonant strings). Letter-selective magnetoencephalographic responses begin in the letter-form area ?60?ms earlier than word-selective responses in the word-form area. Local field potentials confirm the latency and location of letter-selective responses. This area shows increased high-gamma power for ?400?ms, and strong phase-locking with more anterior areas supporting lexico-semantic processing. These findings suggest that during reading, visual stimuli are first encoded as letters before their combinations are encoded as words. Activity then rapidly spreads anteriorly, and the entire network is engaged in sustained integrative processing.
Project description:While behavioral and educational data characterize a fourth grade shift in reading development, neuroscience evidence is relatively lacking. We used the N400 component of the event-related potential waveform to investigate the development of single word processing across the upper elementary years, in comparison to adult readers. We presented third graders, fourth graders, fifth graders, and college students with a well-controlled list of real words, pseudowords, letter strings, false font strings, and animal name targets. Words and pseudowords elicited similar N400s across groups. False font strings elicited N400s similar to words and letter strings in the three groups of children, but not in college students. The pattern of findings suggests relatively adult-like semantic and phonological processing by third grade, but a long developmental time course, beyond fifth grade, for orthographic processing in this context. Thus, the amplitude of the N400 elicited by various word-like stimuli does not reflect some sort of shift or discontinuity in word processing around the fourth grade. However, the results do suggest different developmental time courses for the processes that contribute to automatic single word reading and the integrative N400.
Project description:The visual word form area (VWFA) is a region of left inferior occipitotemporal cortex that is critically involved in visual word recognition. Previous studies have investigated whether and how experience shapes the functional characteristics of VWFA by comparing neural response magnitude in response to words and nonwords. Conflicting results have been obtained, however, perhaps because response magnitude can be influenced by other factors such as attention. In this study, we measured neural activity in monozygotic twins, using functional magnetic resonance imaging. This allowed us to quantify differences in unique environmental contributions to neural activation evoked by words, pseudowords, consonant strings, and false fonts in the VWFA and striate cortex. The results demonstrate significantly greater effects of unique environment in the word and pseudoword conditions compared to the consonant string and false font conditions both in VWFA and in left striate cortex. These findings provide direct evidence for environmental contributions to the neural architecture for reading, and suggest that learning phonology and/or orthographic patterns plays the biggest role in shaping that architecture.
Project description:Reading is a complex cognitive skill subserved by a distributed network of visual and language-related regions. Disruptions of connectivity within this network have been associated with developmental dyslexia but their relation to individual differences in the severity of reading problems remains unclear. Here we investigate whether dysfunctional connectivity scales with the level of reading dysfluency by examining EEG recordings during visual word and false font processing in 9-year-old typically reading children (TR) and two groups of dyslexic children: severely dysfluent (SDD) and moderately dysfluent (MDD) dyslexics. Results indicated weaker occipital to inferior-temporal connectivity for words in both dyslexic groups relative to TRs. Furthermore, SDDs exhibited stronger connectivity from left central to right inferior-temporal and occipital sites for words relative to TRs, and for false fonts relative to both MDDs and TRs. Importantly, reading fluency was positively related with forward and negatively with backward connectivity. Our results suggest disrupted visual processing of words in both dyslexic groups, together with a compensatory recruitment of right posterior brain regions especially in the SDDs during word and false font processing. Functional connectivity in the brain's reading network may thus depend on the level of reading dysfluency beyond group differences between dyslexic and typical readers.
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:The functional role of the left ventral occipito-temporal cortex (vOT) in visual word processing has been studied extensively. A prominent observation is higher activation for unfamiliar but pronounceable letter strings compared to regular words in this region. Some functional accounts have interpreted this finding as driven by top-down influences (e.g., Dehaene and Cohen : Trends Cogn Sci 15:254-262; Price and Devlin : Trends Cogn Sci 15:246-253), while others have suggested a difference in bottom-up processing (e.g., Glezer et al. : Neuron 62:199-204; Kronbichler et al. : J Cogn Neurosci 19:1584-1594). We used dynamic causal modeling for fMRI data to test bottom-up and top-down influences on the left vOT during visual processing of regular words and unfamiliar letter strings. Regular words (e.g., taxi) and unfamiliar letter strings of pseudohomophones (e.g., taksi) were presented in the context of a phonological lexical decision task (i.e., "Does the item sound like a word?"). We found no differences in top-down signaling, but a strong increase in bottom-up signaling from the occipital cortex to the left vOT for pseudohomophones compared to words. This finding can be linked to functional accounts which assume that the left vOT contains neurons tuned to complex orthographic features such as morphemes or words [e.g., Dehaene and Cohen : Trends Cogn Sci 15:254-262; Kronbichler et al. : J Cogn Neurosci 19:1584-1594]: For words, bottom-up signals converge onto a matching orthographic representation in the left vOT. For pseudohomophones, the propagated signals do not converge, but (partially) activate multiple orthographic word representations, reflected in increased effective connectivity.
Project description:Visual tuning for words vs. symbol strings yields complementary increases of fast occipito-temporal activity (N1 or N170) in the event-related potential (ERP), and posterior-anterior gradients of increasing word-specific activity with functional magnetic resonance imaging (fMRI) in the visual word form system (VWFS). However, correlation of these coarse ERP and fMRI tuning responses seems limited to the most anterior part of the VWFS in adult and adolescent readers (Brem et al. [ 2006]: Neuroimage 29:822-837). We thus focused on fMRI tuning gradients of young readers with their more pronounced ERP print tuning, and compared developmental aspects of ERP and fMRI response tuning in the VWFS. Children (10.3 y, n = 19), adolescents (16.2 y, n = 13) and adults (25.2 y, n = 18) were tested with the same implicit reading paradigm using counterbalanced ERP and fMRI imaging. The word-specific occipito-temporal N1 specialization, its corresponding source activity, as well as the integrated source activity (0-700 ms) were most prominent in children and showed a marked decrease with age. The posterior-anterior fMRI gradient of word-specific activity instead which was fully established in children did not develop further, but exhibited a dependence on reading skills independent of age. To conclude, prominent developmental dissociation of the ERP and fMRI tuning patterns emerged despite convergent VWFS localization. The ERP response may selectively reflect fast visual aspects of print specialization, which become less important with age, while the fMRI response seems dominated by integrated task- and reading-related activations in the same regions.
Project description:We read jubmled wrods effortlessly, but the neural correlates of this remarkable ability remain poorly understood. We hypothesized that viewing a jumbled word activates a visual representation that is compared to known words. To test this hypothesis, we devised a purely visual model in which neurons tuned to letter shape respond to longer strings in a compositional manner by linearly summing letter responses. We found that dissimilarities between letter strings in this model can explain human performance on visual search, and responses to jumbled words in word reading tasks. Brain imaging revealed that viewing a string activates this letter-based code in the lateral occipital (LO) region and that subsequent comparisons to stored words are consistent with activations of the visual word form area (VWFA). Thus, a compositional neural code potentially contributes to efficient reading.
Project description:Prior lesion and functional imaging studies have highlighted the importance of the left ventral occipito-temporal (LvOT) cortex for visual word recognition. Within this area, there is a posterior-anterior hierarchy of subregions that are specialized for different stages of orthographic processing. The aim of the present fMRI study was to dissociate the effects of subword orthographic typicality (e.g., cider [high] vs. cynic [low]) from the effect of lexicality (e.g., pollen [word] vs. pillen [pseudoword]). We therefore orthogonally manipulated the orthographic typicality of written words and pseudowords (nonwords and pseudohomophones) in a visual lexical decision task. Consistent with previous studies, we identified greater activation for pseudowords than words (i.e., an effect of lexicality) in posterior LvOT cortex. In addition, we revealed higher activation for atypical than typical strings, irrespective of lexicality, in a left inferior occipital region that is posterior to LvOT cortex. When lexical decisions were made more difficult in the context of pseudohomophone foils, left anterior temporal activation also increased for atypical relative to typical strings. The latter finding agrees with the behavior of patients with progressive anterior temporal lobe degeneration, who have particular difficulty recognizing words with atypical orthography. The most novel outcome of this study is that, within a distributed network of regions supporting orthographic processing, we have identified a left inferior occipital region that is particularly sensitive to the typicality of subword orthographic patterns.
Project description:Parents and teachers worldwide believe that a visual environment rich with print can contribute to young children's literacy. Children seem to recognize words in familiar logos at an early age. However, most of previous studies were carried out with alphabetic scripts. Alphabetic letters regularly correspond to phonological segments in a word and provide strong cues about the identity of the whole word. Thus it was not clear whether children can learn to read words by extracting visual word form information from environmental prints. To exclude the phonological-cue confound, this study tested children's knowledge of Chinese words embedded in familiar logos. The four environmental logos were employed and transformed into four versions with the contextual cues (i.e., something apart from the presentation of the words themselves in logo format like the color, logo and font type cues) gradually minimized. Children aged from 3 to 5 were tested. We observed that children of different ages all performed better when words were presented in highly familiar logos compared to when they were presented in a plain fashion, devoid of context. This advantage for familiar logos was also present when the contextual information was only partial. However, the role of various cues in learning words changed with age. The color and logo cues had a larger effect in 3- and 4- year-olds than in 5-year-olds, while the font type cue played a greater role in 5-year-olds than in the other two groups. Our findings demonstrated that young children did not easily learn words by extracting their visual form information even from familiar environmental prints. However, children aged 5 begin to pay more attention to the visual form information of words in highly familiar logos than those aged 3 and 4.