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1.  Dissociating object familiarity from linguistic properties in mirror word reading 
It is known that the orthographic properties of linguistic stimuli are processed within the left occipitotemporal cortex at about 150–200 ms. We recorded event-related potentials (ERPs) to words in standard or mirror orientation to investigate the role of visual word form in reading. Word inversion was performed to determine whether rotated words lose their linguistic properties.
About 1300 Italian words and legal pseudo-words were presented to 18 right-handed Italian students engaged in a letter detection task. EEG was recorded from 128 scalp sites.
ERPs showed an early effect of word orientation at ~150 ms, with larger N1 amplitudes to rotated than to standard words. Low-resolution brain electromagnetic tomography (LORETA) revealed an increase in N1 to rotated words primarily in the right occipital lobe (BA 18), which may indicate an effect of stimulus familiarity. N1 was greater to target than to non-target letters at left lateral occipital sites, thus reflecting the first stage of orthographic processing. LORETA revealed a strong focus of activation for this effect in the left fusiform gyrus (BA 37), which is consistent with the so-called visual word form area (VWFA). Standard words (compared to pseudowords) elicited an enhancement of left occipito/temporal negativity at about 250–350 ms, followed by a larger anterior P3, a reduced frontal N400 and a huge late positivity. Lexical effects for rotated strings were delayed by about 100 ms at occipito/temporal sites, and were totally absent at later processing stages. This suggests the presence of implicit reading processes, which were pre-attentive and of perceptual nature for mirror strings.
The contrast between inverted and standard words did not lead to the identification of a purely linguistic brain region. This finding suggests some caveats in the interpretation of the inversion effect in subtractive paradigms.
PMCID: PMC1995209  PMID: 17708767
2.  Dysfunctional visual word form processing in progressive alexia 
Brain  2013;136(4):1260-1273.
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.
PMCID: PMC3613714  PMID: 23471694
progressive alexia; letter-by-letter reading; posterior cortical atrophy; logopenic primary progressive aphasia; visual word form system
3.  Orthographic familiarity, phonological legality and number of orthographic neighbours affect the onset of ERP lexical effects 
It has been suggested that the variability among studies in the onset of lexical effects may be due to a series of methodological differences. In this study we investigated the role of orthographic familiarity, phonological legality and number of orthographic neighbours of words in determining the onset of word/non-word discriminative responses.
ERPs were recorded from 128 sites in 16 Italian University students engaged in a lexical decision task. Stimuli were 100 words, 100 quasi-words (obtained by the replacement of a single letter), 100 pseudo-words (non-derived) and 100 illegal letter strings. All stimuli were balanced for length; words and quasi-words were also balanced for frequency of use, domain of semantic category and imageability. SwLORETA source reconstruction was performed on ERP difference waves of interest.
Overall, the data provided evidence that the latency of lexical effects (word/non-word discrimination) varied as a function of the number of a word's orthographic neighbours, being shorter to non-derived than to derived pseudo-words. This suggests some caveats about the use in lexical decision paradigms of quasi-words obtained by transposing or replacing only 1 or 2 letters. Our findings also showed that the left-occipito/temporal area, reflecting the activity of the left fusiform gyrus (BA37) of the temporal lobe, was affected by the visual familiarity of words, thus explaining its lexical sensitivity (word vs. non-word discrimination). The temporo-parietal area was markedly sensitive to phonological legality exhibiting a clear-cut discriminative response between illegal and legal strings as early as 250 ms of latency.
The onset of lexical effects in a lexical decision paradigm depends on a series of factors, including orthographic familiarity, degree of global lexical activity, and phonologic legality of non-words.
PMCID: PMC2491646  PMID: 18601726
4.  Experience-Dependent Hemispheric Specialization of Letters and Numbers is Revealed in Early Visual Processing 
Journal of cognitive neuroscience  2014;26(10):2239-2249.
Recent functional magnetic resonance imaging research has demonstrated that letters and numbers are preferentially processed in distinct regions and hemispheres in the visual cortex. In particular, the left visual cortex preferentially processes letters compared to numbers, while the right visual cortex preferentially processes numbers compared to letters. Because letters and numbers are cultural inventions and are otherwise physically arbitrary, such a double dissociation is strong evidence for experiential effects on neural architecture. Here, we use the high temporal resolution of event-related potentials (ERPs) to investigate the temporal dynamics of the neural dissociation between letters and numbers. We show that the divergence between ERP traces to letters and numbers emerges very early in processing. Letters evoked greater N1 waves (latencies 140–170 ms) than did numbers over left occipital channels, while numbers evoked greater N1s than letters over the right, suggesting letters and numbers are preferentially processed in opposite hemispheres early in visual encoding. Moreover, strings of letters, but not single letters, elicited greater P2 ERP waves, (starting around 250 ms) than numbers did over the left hemisphere, suggesting that the visual cortex is tuned to selectively process combinations of letters, but not numbers, further along in the visual processing stream. Additionally, the processing of both of these culturally defined stimulus types differentiated from similar but unfamiliar visual stimulus forms (false fonts) even earlier in the processing stream (the P1 at 100 ms). These findings imply major cortical specialization processes within the visual system driven by experience with reading and mathematics.
PMCID: PMC4261939  PMID: 24669789
Letter processing; number processing; ERP; hemispheric specialization
5.  Fast, visual specialization for reading in English revealed by the topography of the N170 ERP response 
N170 effects associated with visual words may be related to perceptual expertise effects that have been demonstrated for faces and other extensively studied classes of visual stimuli. Although face and other object expertise effects are typically bilateral or right-lateralized, the spatial topography of reading-related N170 effects are often left-lateralized, providing potential insights into the unique aspects of reading-related perceptual expertise.
Extending previous research in German [1], we use a high-density channel array to characterize the N170 topography for reading-related perceptual expertise in English, a language with inconsistent spelling-to-sound mapping. N170 effects related to overall reading-related expertise are defined by contrasting responses to visual words versus novel symbol strings. By contrasting each of these conditions to pseudowords, we examined how this reading-related N170 effect generalizes to well-ordered novel letter strings.
A sample-by-sample permutation test computed on word versus symbol ERP topographies revealed differences during two time windows corresponding to the N170 and P300 components. Topographic centroid analysis of the word and symbol N170 demonstrated significant differences in both left-right as well as inferior-superior dimensions. Words elicited larger N170 negativities than symbols at inferior occipito-temporal channels, with the maximal effect over left inferior regions often unsampled in conventional electrode montages. Further contrasts produced inferior-superior topographic effects for the pseudoword-symbol comparison and left-lateralized topographic effects for the word-pseudoword comparison.
Fast specialized perception related to reading experience produces an N170 modulation detectable across different EEG systems and different languages. Characterization of such effects may be improved by sampling with greater spatial frequency recordings that sample inferior regions. Unlike in German, reading-related expertise effects in English produced only partial generalization in N170 responses to novel pseudowords. The topographic inferior-superior N170 differences may reflect general perceptual expertise for orthographic strings, as it was found for words and pseudowords across both languages. The topographic left-right N170 difference between words and pseudowords was only found in English, and may suggest that ambiguity in pronunciating novel pseudowords due to inconsistency in spelling-to-sound mapping influences early stages of letter string processing.
PMCID: PMC1208852  PMID: 16091138
6.  N170 ERPs could represent a logographic processing strategy in visual word recognition 
Occipito-temporal N170 component represents the first step where face, object and word processing are discriminated along the ventral stream of the brain. N170 leftward asymmetry observed during reading has been often associated to prelexical orthographic visual word form activation. However, some studies reported a lexical frequency effect for this component particularly during word repetition that appears in contradiction with this prelexical orthographic step. Here, we tested the hypothesis that under word repetition condition, discrimination between words would be operated on visual rather than orthographic basis. In this case, N170 activity may correspond to a logographic processing where a word is processed as a whole.
To test such an assumption, frequent words, infrequent words and pseudowords were presented to the subjects that had to complete a visual lexical decision task. Different repetition conditions were defined 1 – weak repetition, 2 – massive repetition and 3 – massive repetition with font alternation. This last condition was designed to change visual word shape during repetition and therefore to interfere with a possible visual strategy during word recognition.
Behavioral data showed an important frequency effect for the weak repetition condition, a lower but significant frequency effect for massive repetition, and no frequency effect for the changing font repetition. Moreover alternating font repetitions slowed subject's responses in comparison to "simple" massive repetition.
ERPs results evidenced larger N170 amplitude in the left hemisphere for frequent than both infrequent words and pseudowords during massive repetition. Moreover, when words were repeated with different fonts this N170 effect was not present, suggesting a visual locus for such a N170 frequency effect.
N170 represents an important step in visual word recognition, consisting probably in a prelexical orthographic processing. But during the reading of very frequent words or after a massive repetition of a word, it could represent a more holistic process where words are processed as a global visual pattern.
PMCID: PMC1884163  PMID: 17451598
7.  Does the Reading of Different Orthographies Produce Distinct Brain Activity Patterns? An ERP Study 
PLoS ONE  2012;7(5):e36030.
Orthographies vary in the degree of transparency of spelling-sound correspondence. These range from shallow orthographies with transparent grapheme-phoneme relations, to deep orthographies, in which these relations are opaque. Only a few studies have examined whether orthographic depth is reflected in brain activity. In these studies a between-language design was applied, making it difficult to isolate the aspect of orthographic depth. In the present work this question was examined using a within-subject-and-language investigation. The participants were speakers of Hebrew, as they are skilled in reading two forms of script transcribing the same oral language. One form is the shallow pointed script (with diacritics), and the other is the deep unpointed script (without diacritics). Event-related potentials (ERPs) were recorded while skilled readers carried out a lexical decision task in the two forms of script. A visual non-orthographic task controlled for the visual difference between the scripts (resulting from the addition of diacritics to the pointed script only). At an early visual-perceptual stage of processing (∼165 ms after target onset), the pointed script evoked larger amplitudes with longer latencies than the unpointed script at occipital-temporal sites. However, these effects were not restricted to orthographic processing, and may therefore have reflected, at least in part, the visual load imposed by the diacritics. Nevertheless, the results implied that distinct orthographic processing may have also contributed to these effects. At later stages (∼340 ms after target onset) the unpointed script elicited larger amplitudes than the pointed one with earlier latencies. As this latency has been linked to orthographic-linguistic processing and to the classification of stimuli, it is suggested that these differences are associated with distinct lexical processing of a shallow and a deep orthography.
PMCID: PMC3352908  PMID: 22615746
8.  Activation of the Left Inferior Frontal Gyrus in the First 200 ms of Reading: Evidence from Magnetoencephalography (MEG) 
PLoS ONE  2009;4(4):e5359.
It is well established that the left inferior frontal gyrus plays a key role in the cerebral cortical network that supports reading and visual word recognition. Less clear is when in time this contribution begins. We used magnetoencephalography (MEG), which has both good spatial and excellent temporal resolution, to address this question.
Methodology/Principal Findings
MEG data were recorded during a passive viewing paradigm, chosen to emphasize the stimulus-driven component of the cortical response, in which right-handed participants were presented words, consonant strings, and unfamiliar faces to central vision. Time-frequency analyses showed a left-lateralized inferior frontal gyrus (pars opercularis) response to words between 100–250 ms in the beta frequency band that was significantly stronger than the response to consonant strings or faces. The left inferior frontal gyrus response to words peaked at ∼130 ms. This response was significantly later in time than the left middle occipital gyrus, which peaked at ∼115 ms, but not significantly different from the peak response in the left mid fusiform gyrus, which peaked at ∼140 ms, at a location coincident with the fMRI–defined visual word form area (VWFA). Significant responses were also detected to words in other parts of the reading network, including the anterior middle temporal gyrus, the left posterior middle temporal gyrus, the angular and supramarginal gyri, and the left superior temporal gyrus.
These findings suggest very early interactions between the vision and language domains during visual word recognition, with speech motor areas being activated at the same time as the orthographic word-form is being resolved within the fusiform gyrus. This challenges the conventional view of a temporally serial processing sequence for visual word recognition in which letter forms are initially decoded, interact with their phonological and semantic representations, and only then gain access to a speech code.
PMCID: PMC2671164  PMID: 19396362
9.  Cultural Constraints on Brain Development: Evidence from a Developmental Study of Visual Word Processing in Mandarin Chinese 
Cerebral Cortex (New York, NY)  2009;20(5):1223-1233.
Developmental differences in phonological and orthographic processing in Chinese were examined in 9 year olds, 11 year olds, and adults using functional magnetic resonance imaging. Rhyming and spelling judgments were made to 2-character words presented sequentially in the visual modality. The spelling task showed greater activation than the rhyming task in right superior parietal lobule and right inferior temporal gyrus, and there were developmental increases across tasks bilaterally in these regions in addition to bilateral occipital cortex, suggesting increased involvement over age on visuo-orthographic analysis. The rhyming task showed greater activation than the spelling task in left superior temporal gyrus and there were developmental decreases across tasks in this region, suggesting reduced involvement over age on phonological representations. The rhyming and spelling tasks included words with conflicting orthographic and phonological information (i.e., rhyming words spelled differently or nonrhyming words spelled similarly) or nonconflicting information. There was a developmental increase in the difference between conflicting and nonconflicting words in left inferior parietal lobule, suggesting greater engagement of systems for mapping between orthographic and phonological representations. Finally, there were developmental increases across tasks in an anterior (Broadman area [BA] 45, 46) and posterior (BA 9) left inferior frontal gyrus, suggesting greater reliance on controlled retrieval and selection of posterior lexical representations.
PMCID: PMC2852508  PMID: 19773547
Chinese; development; rhyming; spelling
10.  Sensitivity to Orthographic Familiarity in the Occipito-Temporal Region 
NeuroImage  2007;39(4):1988-2001.
The involvement of the left hemisphere occipito-temporal (OT) junction in reading has been established, yet there is current controversy over the region’s specificity for reading and the nature of its role in the reading process. Recent neuroimaging findings suggest that the region is sensitive to orthographic familiarity (Kronbichler et al., 2007), and the present study tested that hypothesis. Using fMRI, the OT region and other regions in the reading network were localized in 28 adult, right-handed participants. The BOLD signal in these regions was measured during a phonological judgment task (i.e., “Does it sound like a word?”). Stimuli included words, pseudohomophones (phonologically familiar yet orthographically unfamiliar), and pseudowords (phonologically and orthographically unfamiliar) that were matched on lexical properties including sublexical orthography. Relative to baseline, BOLD signal in the OT region was greater for pseudohomophones than for words, suggesting that the region is sensitive to orthographic familiarity at the whole-word level. Further contrasts of orthographic frequency within the word condition revealed increased BOLD signal for low- than high-frequency words. Specialization in the OT area for recognition of frequent letter strings may support the development of reading expertise. Additionally, BOLD signal in the OT region correlates positively with reading efficiency, supporting the idea that this region is a skill zone for reading printed words. BOLD signal in the IFG and STG correlate negatively with reading efficiency, indicating that processing effort in these classic phonological regions is inversely related to reading efficiency.
PMCID: PMC2267437  PMID: 18180168
11.  Paying attention to orthography: a visual evoked potential study 
In adult readers, letters, and words are rapidly identified within visual networks to allow for efficient reading abilities. Neuroimaging studies of orthography have mostly used words and letter strings that recruit many hierarchical levels in reading. Understanding how single letters are processed could provide further insight into orthographic processing. The present study investigated orthographic processing using single letters and pseudoletters when adults were encouraged to pay attention to or away from orthographic features. We measured evoked potentials (EPs) to single letters and pseudoletters from adults while they performed an orthographic-discrimination task (letters vs. pseudoletters), a color-discrimination task (red vs. blue), and a target-detection task (respond to #1 and #2). Larger and later peaking N1 responses (~170 ms) and larger P2 responses (~250 ms) occurred to pseudoletters as compared to letters. This reflected greater visual processing for pseudoletters. Dipole analyses localized this effect to bilateral fusiform and inferior temporal cortices. Moreover, this letter-pseudoletter difference was not modulated by task and thus indicates that directing attention to or away from orthographic features did not affect early visual processing of single letters or pseudoletters within extrastriate regions. Paying attention to orthography or color as compared to disregarding the stimuli (target-detection task) elicited selection negativities at about 175 ms, which were followed by a classical N2-P3 complex. This indicated that the tasks sufficiently drew participant's attention to and away from the stimuli. Together these findings revealed that visual processing of single letters and pseudoletters, in adults, appeared to be sensory-contingent and independent of paying attention to stimulus features (e.g., orthography or color).
PMCID: PMC3659343  PMID: 23734115
orthography; visual evoked potential (VEP); attention; reading; dipole modeling
12.  Lateralisation of language function in young adults born very preterm 
Objective: To explore, using functional magnetic resonance imaging (MRI), the functional organisation of phonological processing in young adults born very preterm.
Subjects: Six right handed male subjects with radiological evidence of thinning of the corpus callosum were selected from a cohort of very preterm subjects. Six normal right handed male volunteers acted as controls.
Method: Blood oxygenation level dependent contrast echoplanar images were acquired over five minutes at 1.5 T while subjects performed the tasks. During the ON condition, subjects were visually presented with pairs of non-words and asked to press a key when a pair of words rhymed (phonological processing). This task alternated with the OFF condition, which required subjects to make letter case judgments of visually presented pairs of consonant letter strings (orthographic processing). Generic brain activation maps were constructed from individual images by sinusoidal regression and non-parametric testing. Between group differences in the mean power of experimental response were identified on a voxel wise basis by analysis of variance.
Results: Compared with controls, the subjects with thinning of the corpus callosum showed significantly reduced power of response in the left hemisphere, including the peristriate cortex and the cerebellum, as well as in the right parietal association area. Significantly increased power of response was observed in the right precentral gyrus and the right supplementary motor area.
Conclusions: The data show evidence of increased frontal and decreased occipital activation in male subjects with neurodevelopmental thinning of the corpus callosum, which may be due to the operation of developmental compensatory mechanisms.
PMCID: PMC1756037  PMID: 14977893
13.  Developmental changes in brain regions involved in phonological and orthographic processing during spoken language processing 
NeuroImage  2008;41(2):623-635.
Developmental differences in brain activation of 9- to 15-year-old children were examined during an auditory rhyme decision task to spoken words using functional magnetic resonance imaging (fMRI). As a group, children showed activation in left superior/middle temporal gyri (BA 22, 21), right middle temporal gyrus (BA 21), dorsal (BA 45, pars opercularis) and ventral (BA 46, pars triangularis) aspects of left inferior frontal gyrus, and left fusiform gyrus (BA 37). There was a developmental increase in activation in left middle temporal gyrus (BA 22) across all lexical conditions, suggesting that automatic semantic processing increases with age regardless of task demands. Activation in left dorsal inferior frontal gyrus also showed developmental increases for the conflicting (e.g. PINT-MINT) compared to the non-conflicting (e.g. PRESS-LIST) non-rhyming conditions, indicating that this area becomes increasingly involved in strategic phonological processing in the face of conflicting orthographic and phonological representations. Left inferior temporal/fusiform gyrus (BA 37) activation was also greater for the conflicting (e.g. PINT-MINT) condition, and a developmental increase was found in the positive relationship between individuals' reaction time and activation in left lingual/fusiform gyrus (BA 18) in this condition, indicating an age-related increase in the association between longer reaction times and greater visual-orthographic processing in this conflicting condition. These results suggest that orthographic processing is automatically engaged by children in a task that does not require access to orthographic information for correct performance, especially when orthographic and phonological representations conflict, and especially for longer response latencies in older children.
PMCID: PMC2443702  PMID: 18413290
14.  Opposite effects of visual and auditory word-likeness on activity in the visual word form area 
The present fMRI study investigated the effects of word-likeness of visual and auditory stimuli on activity along the ventral visual stream. In the context of a one-back task, we presented visual and auditory words, pseudowords, and artificial stimuli (i.e., false-fonts and reversed-speech, respectively). Main findings were regionally specific effects of word-likeness on activation in a left ventral occipitotemporal region corresponding to the classic localization of the Visual Word Form Area (VWFA). Specifically, we found an inverse word-likeness effect for the visual stimuli in the form of decreased activation for words compared to pseudowords which, in turn, elicited decreased activation compared to the artificial stimuli. For the auditory stimuli, we found positive word-likeness effects as both words and pseudowords elicited more activation than the artificial stimuli. This resulted from a marked deactivation in response to the artificial stimuli and no such deactivation for words and pseudowords. We suggest that the opposite effects of visual and auditory word-likeness on VWFA activation can be explained by assuming the involvement of visual orthographic memory representations. For the visual stimuli, these representations reduce the coding effort as a function of word-likeness. This results in highest activation to the artificial stimuli and least activation to words for which corresponding representations exist. The positive auditory word-likeness effects may result from activation of orthographic information associated with the auditory words and pseudowords. The view that the VWFA has a primarily visual function is supported by our findings of high activation to the visual artificial stimuli (which have no phonological or semantic associations) and deactivation to the auditory artificial stimuli. According to the phenomenon of cross-modal sensory suppression such deactivations during demanding auditory processing are expected in visual regions.
PMCID: PMC3756304  PMID: 24009569
fMRI; neuroimaging; one-back task; word-likeness; word processing; VWFA; orthographic representations
15.  Word learning and the cerebral hemispheres: from serial to parallel processing of written words 
Reading familiar words differs from reading unfamiliar non-words in two ways. First, word reading is faster and more accurate than reading of unfamiliar non-words. Second, effects of letter length are reduced for words, particularly when they are presented in the right visual field in familiar formats. Two experiments are reported in which right-handed participants read aloud non-words presented briefly in their left and right visual fields before and after training on those items. The non-words were interleaved with familiar words in the naming tests. Before training, naming was slow and error prone, with marked effects of length in both visual fields. After training, fewer errors were made, naming was faster, and the effect of length was much reduced in the right visual field compared with the left. We propose that word learning creates orthographic word forms in the mid-fusiform gyrus of the left cerebral hemisphere. Those word forms allow words to access their phonological and semantic representations on a lexical basis. But orthographic word forms also interact with more posterior letter recognition systems in the middle/inferior occipital gyri, inducing more parallel processing of right visual field words than is possible for any left visual field stimulus, or for unfamiliar non-words presented in the right visual field.
PMCID: PMC2846318  PMID: 19933140
word learning; reading; hemispheres; visual fields; word length; case alternation
16.  Word and Letter String Processing Networks in Schizophrenia: Evidence for Anomalies and Compensation 
Brain and language  2008;107(2):158-166.
Imaging studies show that in normal language correlated activity between anterior and posterior brain regions increases as the linguistic and semantic content (i.e., from false fonts, letter strings, pseudo words, to words) of stimuli increase. In schizophrenia however, disrupted functional connectivity between frontal and posterior brain regions has been frequently reported and these disruptions may change the nature of language organization. We characterized basic linguistic operations in word and letter string processing in a region-of-interest network using structural equation modeling (SEM). Healthy volunteers and volunteers with schizophrenia performed an fMRI one-back matching task with real words and consonant letter strings. We hypothesized that left hemisphere network dysfunction in schizophrenia would be present during processes dealing with linguistic/semantic content. The modeling results suggest aberrant left hemisphere function in schizophrenia, even in tasks requiring minimal access to language. Alternative mechanisms included increases in right hemisphere involvement and increased top-down influence from frontal to posterior regions.
PMCID: PMC2599869  PMID: 18829095
Schizophrenia and language; Lateralization; Lexical-semantic processing; Imaging; Effective Connectivity; Modeling
17.  A Dual-Route Perspective on Poor Reading in a Regular Orthography: An fMRI Study 
This study examined functional brain abnormalities in dyslexic German readers who – due to the regularity of German in the reading direction – do not exhibit the reading accuracy problem of English dyslexic readers, but suffer primarily from a reading speed problem. The in-scanner task required phonological lexical decisions (i.e., Does xxx sound like an existing word?) and presented familiar and unfamiliar letter strings of existing phonological words (e.g., Taxi-Taksi) together with nonwords (e.g., Tazi). Dyslexic readers exhibited the same response latency pattern (words < pseudohomophones < nonwords) as nonimpaired readers, but latencies to all item types were much prolonged. The imaging results were suggestive for a different neural organization of reading processes in dyslexic readers. Specifically, dyslexic readers, in response to lexical route processes, exhibited underactivation in a left ventral occipitotemporal region which presumably is engaged by visual-orthographic whole word recognition. This region was also insensitive to the increased visual-orthographic processing demands of the sublexical route. Reduced engagement in response to sublexical route processes was also found in a left inferior parietal region, presumably engaged by attentional processes, and in a left inferior frontal region, presumably engaged by phonological processes. In contrast to this reduced engagement of the optimal left hemisphere reading network (ventral OT, inferior parietal, inferior frontal), our dyslexic readers exhibited increased engagement of visual occipital regions and of regions presumably engaged by silent articulatory processes (premotor/motor cortex and subcortical caudate and putamen).
PMCID: PMC3073233  PMID: 20650450
18.  A dual-route perspective on poor reading in a regular orthography: An fMRI study 
This study examined functional brain abnormalities in dyslexic German readers who – due to the regularity of German in the reading direction – do not exhibit the reading accuracy problem of English dyslexic readers, but suffer primarily from a reading speed problem. The in-scanner task required phonological lexical decisions (i.e., Does xxx sound like an existing word?) and presented familiar and unfamiliar letter strings of existing phonological words (e.g., Taxi-Taksi) together with nonwords (e.g., Tazi). Dyslexic readers exhibited the same response latency pattern (words < pseudohomophones < nonwords) as nonimpaired readers, but latencies to all item types were much prolonged. The imaging results were suggestive for a different neural organization of reading processes in dyslexic readers. Specifically, dyslexic readers, in response to lexical route processes, exhibited underactivation in a left ventral occipitotemporal (OT) region which presumably is engaged by visual-orthographic whole word recognition. This region was also insensitive to the increased visual-orthographic processing demands of the sublexical route. Reduced engagement in response to sublexical route processes was also found in a left inferior parietal region, presumably engaged by attentional processes, and in a left inferior frontal region, presumably engaged by phonological processes. In contrast to this reduced engagement of the optimal left hemisphere reading network (ventral OT, inferior parietal, inferior frontal), our dyslexic readers exhibited increased engagement of visual occipital regions and of regions presumably engaged by silent articulatory processes (premotor/motor cortex and subcortical caudate and putamen).
PMCID: PMC3073233  PMID: 20650450
Developmental dyslexia; fMRI; Reading; Phonological lexical decision; Dual-route
19.  Cortical Organization of Language Pathways in Children with Non-Localized Cryptogenic Epilepsy 
Children with a history of epilepsy are almost six times more likely than their unaffected siblings to be referred for speech or language therapy. However, the abnormalities in neural pathway that cause these delays are poorly understood. We recorded evoked fields using whole-head magnetoencephalography during real and non-word visual and auditory rhyme tasks in 15 children with non-localized cryptogenic epilepsy. Basic phonological and orthographic language skills were assessed using Woodcock–Johnson Test of Achievement subtests. Dynamic statistical parameter mapping was used with individual participant magnetic resonance images. Significant cortical activity was visualized on average and performance weighted maps. For the auditory rhyme tasks, bilateral primary and secondary auditory cortices, the superior temporal sulcus, and insular cortex were activated early with later increases in left hemisphere activity. Visual rhyme tasks evoked early bilateral primary and secondary occipital cortical and angular gyri activity followed by later activation of the planum temporale and supramarginal gyri and the left ventral occipitotemporal area. For the auditory rhyme tasks, performance weighted maps demonstrated that early right hemisphere activation was associated with poorer reading skills while later activity was associated with better reading skills; for the left hemisphere, greater early activation of the secondary auditory cortex, including the planum temporale, was related to better reading skills while relatively later activation of these areas was associated with poorer reading skills. For the visual rhyme tasks, greater activity in the bilateral ventral occipitotemporal and insular areas and angular and supramarginal gyri were associated with better performance. These data suggest that spatiotemporal cortical activation patterns are associated with variations in language performance in non-localized cryptogenic epilepsy.
PMCID: PMC4191195  PMID: 25346681
epilepsy; reading; language; magnetoencephalography; dynamic statistical parameter
20.  Evidence for Right Hemisphere Phonology in a Backward Masking Task 
Brain and language  2011;119(3):232-237.
The extent to which orthographic and phonological processes are available during the initial moments of word recognition within each hemisphere is under specified, particularly for the right hemisphere. Few studies have investigated whether each hemisphere uses orthography and phonology under constraints that restrict the viewing time of words and reduce overt phonological demands. The current study used backward masking in the divided visual field paradigm to explore hemisphere differences in the availability of orthographic and phonological word recognition processes. A 20 ms and 60 ms SOA were used to track the time course of how these processes develop during pre-lexical moments of word recognition. Nonword masks varied in similarity to the target words such that there were four types: orthographically and phonologically similar, orthographically but not phonologically similar, phonologically but not orthographically similar and unrelated. The results showed the left hemisphere has access to both orthography and phonology early in the word recognition process. With more time to process the stimulus, the left hemisphere is able to usephonology which benefits word recognition to a larger extent than orthography. The right hemisphere also demonstrates access to both orthography and phonology in the initial moments of word recognition, however, orthographic similarity improves word recognition to a greater extent than phonological similarity.
PMCID: PMC3191255  PMID: 21683434
Phonology; Orthography; Laterality; Word Recognition; Backward Masking; Visual Field; Right Hemisphere; Left Hemisphere
21.  A cross-linguistic evaluation of script-specific effects on fMRI lateralization in late second language readers 
Behavioral and neuroimaging studies have provided evidence that reading is strongly left lateralized, and the degree of this pattern of functional lateralization can be indicative of reading competence. However, it remains unclear whether functional lateralization differs between the first (L1) and second (L2) languages in bilingual L2 readers. This question is particularly important when the particular script, or orthography, learned by the L2 readers is markedly different from their L1 script. In this study, we quantified functional lateralization in brain regions involved in visual word recognition for participants' L1 and L2 scripts, with a particular focus on the effects of L1–L2 script differences in the visual complexity and orthographic depth of the script. Two different groups of late L2 learners participated in an fMRI experiment using a visual one-back matching task: L1 readers of Japanese who learnt to read alphabetic English and L1 readers of English who learnt to read both Japanese syllabic Kana and logographic Kanji. The results showed weaker leftward lateralization in the posterior lateral occipital complex (pLOC) for logographic Kanji compared with syllabic and alphabetic scripts in both L1 and L2 readers of Kanji. When both L1 and L2 scripts were non-logographic, where symbols are mapped onto sounds, functional lateralization did not significantly differ between L1 and L2 scripts in any region, in any group. Our findings indicate that weaker leftward lateralization for logographic reading reflects greater requirement of the right hemisphere for processing visually complex logographic Kanji symbols, irrespective of whether Kanji is the readers' L1 or L2, rather than characterizing additional cognitive efforts of L2 readers. Finally, brain-behavior analysis revealed that functional lateralization for L2 visual word processing predicted L2 reading competency.
PMCID: PMC4006067  PMID: 24795604
visual complexity; orthographic depth; second langauge reading; logogrpahic; functional lateralization
22.  Children with Reading Difficulties Show Differences in Brain Regions Associated with Orthographic Processing During Spoken Language Processing 
Brain research  2010;1356:73-84.
We explored the neural basis of spoken language deficits in children with reading difficulty, specifically focusing on the role of orthography during spoken language processing. We used functional magnetic resonance imaging (fMRI) to examine differences in brain activation between children with reading difficulties (aged 9-to-15 years) and age-matched children with typical achievement during an auditory rhyming task. Both groups showed activation in bilateral superior temporal gyri (BA 42, 22), a region associated with phonological processing, with no significant between-groups differences. Interestingly, typically achieving children, but not children with reading difficulties, showed activation of left fusiform cortex (BA 37), a region implicated in orthographic processing. Furthermore, this activation was significantly greater for typically achieving children compared to those with reading difficulties. These findings suggest that typical children automatically activate orthographic representations during spoken language processing, while those with reading difficulties do not. Follow-up analyses revealed that the intensity of the activation in the fusiform gyrus was associated with significantly stronger behavioral conflict effects in typically achieving children only (i.e., longer latencies to rhyming pairs with orthographically dissimilar endings than to those with identical orthographic endings; jazz-has vs. cat-hat). Finally, for reading disabled children, a positive correlation between left fusiform activation and non-word reading was observed, such that greater access to orthography was related to decoding ability. Taken together, the results suggest the integration of orthographic and phonological processing are directly related to reading ability.
PMCID: PMC2942963  PMID: 20691675
Phonology; Orthography; Dyslexia; fMRI; Language Deficits; Spoken Word Processing
23.  Taxi vs. Taksi: On orthographic word recognition in the left ventral occipitotemporal cortex 
Journal of cognitive neuroscience  2007;19(10):1584-1594.
The importance of the left occipitotemporal cortex for visual word processing is highlighted by numerous functional neuroimaging studies, but the precise function of the Visual Word Form Area (VWFA) in this brain region is still under debate. The present fMRI study varied orthographic familiarity independent from phonological-semantic familiarity by presenting orthographically familiar and orthographically unfamiliar forms (pseudohomophones) of the same words in a phonological lexical decision task. Consistent with orthographic word recognition in the VWFA, we found lower activation for familiar compared to unfamiliar forms, but no difference between pseudohomophones and pseudowords. This orthographic familiarity effect in the VWFA differed from the phonological familiarity effect in left frontal regions, where phonologically unfamiliar pseudowords led to higher activation than phonologically familiar pseudohomophones. We suggest that the VWFA not only computes letter string representations but also hosts word specific orthographic representations. These representations function as recognition units with the effect that letter strings, which readily match with stored representations lead to less activation than letter strings which do not.
PMCID: PMC2989180  PMID: 17933023
Functional MRI; orthographic word recognition; visual word processing; occipitotemporal cortex; reading
24.  What does the brain of children with developmental dyslexia tell us about reading improvement? ERP evidence from an intervention study 
Intervention is key to managing developmental dyslexia (DD), but not all children with DD benefit from treatment. Some children improve (improvers, IMP), whereas others do not improve (non-improvers, NIMP). Neurobiological differences between IMP and NIMP have been suggested, but studies comparing IMP and NIMP in childhood are missing. The present study examined whether ERP patterns change with treatment and differ between IMP and NIMP. We investigated the ERPs of 28 children with DD and 25 control children (CON) while performing a phonological lexical decision (PLD) task before and after a 6-month intervention. After intervention children with DD were divided into IMP (n = 11) and NIMP (n = 17). In the PLD–task children were visually presented with words, pseudohomophones, pseudowords, and false fonts and had to decide whether the presented stimulus sounded like an existing German word or not. Prior to intervention IMP showed higher N300 amplitudes over fronto-temporal electrodes compared to NIMP and CON and N400 amplitudes were attenuated in both IMP and NIMP compared to CON. After intervention N300 amplitudes of IMP were comparable to those of CON and NIMP. This suggests that the N300, which has been related to phonological access of orthographic stimuli and integration of orthographic and phonological representations, might index a compensatory mechanism or precursor that facilitates reading improvement. The N400, which is thought to reflect grapheme-phoneme conversion or the access to the orthographic lexicon increased in IMP from pre to post and was comparable to CON after intervention. Correlations between N300 amplitudes pre, growth in reading ability and N400 amplitudes post indicated that higher N300 amplitudes might be important for reading improvement and increase in N400 amplitudes. The results suggest that children with DD, showing the same cognitive profile might differ regarding their neuronal profile which could further influence reading improvement.
PMCID: PMC4071643  PMID: 25018723
developmental dyslexia; intervention; treatment; improvement; non-improvement; electrophysiology; N400; N300
25.  Effect of Orthographic Processes on Letter Identity and Letter-Position Encoding in Dyslexic Children 
The ability to identify letters and encode their position is a crucial step of the word recognition process. However and despite their word identification problem, the ability of dyslexic children to encode letter identity and letter-position within strings was not systematically investigated. This study aimed at filling this gap and further explored how letter identity and letter-position encoding is modulated by letter context in developmental dyslexia. For this purpose, a letter-string comparison task was administered to French dyslexic children and two chronological age (CA) and reading age (RA)-matched control groups. Children had to judge whether two successively and briefly presented four-letter strings were identical or different. Letter-position and letter identity were manipulated through the transposition (e.g., RTGM vs. RMGT) or substitution of two letters (e.g., TSHF vs. TGHD). Non-words, pseudo-words, and words were used as stimuli to investigate sub-lexical and lexical effects on letter encoding. Dyslexic children showed both substitution and transposition detection problems relative to CA-controls. A substitution advantage over transpositions was only found for words in dyslexic children whereas it extended to pseudo-words in RA-controls and to all type of items in CA-controls. Letters were better identified in the dyslexic group when belonging to orthographically familiar strings. Letter-position encoding was very impaired in dyslexic children who did not show any word context effect in contrast to CA-controls. Overall, the current findings point to a strong letter identity and letter-position encoding disorder in developmental dyslexia.
PMCID: PMC3356879  PMID: 22661961
letter-string processing; letter-position encoding; letter-identity encoding; letter transposition; letter substitution; reading acquisition; dyslexic children

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