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1.  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.
doi:10.1016/j.neuroimage.2007.10.044
PMCID: PMC2267437  PMID: 18180168
2.  Developmental dyslexia: dysfunction of a left hemisphere reading network 
This mini-review summarizes and integrates findings from recent meta-analyses and original neuroimaging studies on functional brain abnormalities in dyslexic readers. Surprisingly, there is little empirical support for the standard neuroanatomical model of developmental dyslexia, which localizes the primary phonological decoding deficit in left temporo-parietal (TP) regions. Rather, recent evidence points to a dysfunction of a left hemisphere reading network, which includes occipito-temporal (OT), inferior frontal, and inferior parietal regions.
doi:10.3389/fnhum.2012.00120
PMCID: PMC3340948  PMID: 22557962
brain; developmental dyslexia; fMRI; meta-analysis; neuroimaging; reading
3.  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.
doi:10.1093/brain/awt034
PMCID: PMC3613714  PMID: 23471694
progressive alexia; letter-by-letter reading; posterior cortical atrophy; logopenic primary progressive aphasia; visual word form system
4.  On the functional neuroanatomy of visual word processing: Effects of case and letter deviance 
Journal of cognitive neuroscience  2009;21(2):222-229.
This fMRI study contrasted case-deviant and letter-deviant forms with familiar forms of the same phonological words (e.g., TaXi and Taksi vs. Taxi) and found, that both types of deviance led to increased activation in a left occipitotemporal region corresponding to the Visual Word Form Area. Case-deviant items, in addition, led to increased activation in a right occipitotemporal region and in a left occipital and a left posterior occipitotemporal region, possibly reflecting the increased demands on letter form coding. For letter-deviant items, in addition to the increased left occipitotemporal activation, a main finding was increased activation primarily in extended left frontal regions, possibly reflecting sublexically mediated access to word phonology. These findings are consistent with general features of cognitive dual-route models of visual word processing. Furthermore, they add support to the main feature of Dehaene et al.’s (2005) neural model of early stages of visual word processing . However, the increased activation found for case-deviant items in the VWFA cannot be immediately reconciled with the assumption of completely abstract case-independent orthographic word codes in the VWFA.
doi:10.1162/jocn.2009.21002
PMCID: PMC2976854  PMID: 18476755
Functional MRI; visual word recognition; occipitotemporal cortex; visual word form area; orthographic processing
5.  Grey Matter Alterations Co-Localize with Functional Abnormalities in Developmental Dyslexia: An ALE Meta-Analysis 
PLoS ONE  2012;7(8):e43122.
The neural correlates of developmental dyslexia have been investigated intensively over the last two decades and reliable evidence for a dysfunction of left-hemispheric reading systems in dyslexic readers has been found in functional neuroimaging studies. In addition, structural imaging studies using voxel-based morphometry (VBM) demonstrated grey matter reductions in dyslexics in several brain regions. To objectively assess the consistency of these findings, we performed activation likelihood estimation (ALE) meta-analysis on nine published VBM studies reporting 62 foci of grey matter reduction in dyslexic readers. We found six significant clusters of convergence in bilateral temporo-parietal and left occipito-temporal cortical regions and in the cerebellum bilaterally. To identify possible overlaps between structural and functional deviations in dyslexic readers, we conducted additional ALE meta-analyses of imaging studies reporting functional underactivations (125 foci from 24 studies) or overactivations (95 foci from 11 studies ) in dyslexics. Subsequent conjunction analyses revealed overlaps between the results of the VBM meta-analysis and the meta-analysis of functional underactivations in the fusiform and supramarginal gyri of the left hemisphere. An overlap between VBM results and the meta-analysis of functional overactivations was found in the left cerebellum. The results of our study provide evidence for consistent grey matter variations bilaterally in the dyslexic brain and substantial overlap of these structural variations with functional abnormalities in left hemispheric regions.
doi:10.1371/journal.pone.0043122
PMCID: PMC3423424  PMID: 22916214
6.  Reading without the left ventral occipito-temporal cortex 
Neuropsychologia  2012;50-360(14):3621-3635.
The left ventral occipito-temporal cortex (LvOT) is thought to be essential for the rapid parallel letter processing that is required for skilled reading. Here we investigate whether rapid written word identification in skilled readers can be supported by neural pathways that do not involve LvOT. Hypotheses were derived from a stroke patient who acquired dyslexia following extensive LvOT damage. The patient followed a reading trajectory typical of that associated with pure alexia, re-gaining the ability to read aloud many words with declining performance as the length of words increased. Using functional MRI and dynamic causal modelling (DCM), we found that, when short (three to five letter) familiar words were read successfully, visual inputs to the patient’s occipital cortex were connected to left motor and premotor regions via activity in a central part of the left superior temporal sulcus (STS). The patient analysis therefore implied a left hemisphere “reading-without-LvOT” pathway that involved STS. We then investigated whether the same reading-without-LvOT pathway could be identified in 29 skilled readers and whether there was inter-subject variability in the degree to which skilled reading engaged LvOT. We found that functional connectivity in the reading-without-LvOT pathway was strongest in individuals who had the weakest functional connectivity in the LvOT pathway. This observation validates the findings of our patient’s case study. Our findings highlight the contribution of a left hemisphere reading pathway that is activated during the rapid identification of short familiar written words, particularly when LvOT is not involved. Preservation and use of this pathway may explain how patients are still able to read short words accurately when LvOT has been damaged.
Highlights
► Word reading can succeed after damage to the left occipito-temporal cortex. ► This is enabled by an alternative pathway via the left superior temporal sulcus. ► Connectivity analysis demonstrated the existence of this pathway in normal readers. ► We hypothesise that this pathway integrates semantics with phonology. ► Our work stresses the importance of mapping alternative degenerate reading pathways.
doi:10.1016/j.neuropsychologia.2012.09.030
PMCID: PMC3524457  PMID: 23017598
Effective connectivity; Alexia; Recovery; Degeneracy; Word reading pathways; Functional MRI
7.  ERP correlates of letter identity and letter position are modulated by lexical frequency 
Brain and language  2013;125(1):11-27.
The encoding of letter position is a key aspect in all recently proposed models of visual-word recognition. We analyzed the impact of lexical frequency on letter position assignment by examining the temporal dynamics of lexical activation induced by pseudowords extracted from words of different frequencies. For each word (e.g., BRIDGE), we created two pseudowords: A transposed-letter (TL: BRIGDE) and a replaced-letter pseudoword (RL: BRITGE). ERPs were recorded while participants read words and pseudowords in two tasks: Semantic categorization (Experiment 1) and lexical decision (Experiment 2). For high-frequency stimuli, similar ERPs were obtained for words and TL-pseudowords, but the N400 component to words was reduced relative to RL-pseudowords, indicating less lexical/semantic activation. In contrast, TL- and RL-pseudowords created from low-frequency stimuli elicited similar ERPs. Behavioral responses in the lexical decision task paralleled this asymmetry. The present findings impose constraints on computational and neural models of visual-word recognition.
doi:10.1016/j.bandl.2012.12.009
PMCID: PMC3612367  PMID: 23454070
visual-word recognition; position coding; ERPs; word-frequency; transposed-letter effects
8.  Inter-subject variability in the use of two different neuronal networks for reading aloud familiar words 
Neuroimage  2008;42(3-3):1226-1236.
Cognitive models of reading predict that high frequency regular words can be read in more than one way. We investigated this hypothesis using functional MRI and covariance analysis in 43 healthy skilled readers. Our results dissociated two sets of regions that were differentially engaged across subjects who were reading the same familiar words. Some subjects showed more activation in left inferior frontal and anterior occipito-temporal regions while other subjects showed more activation in right inferior parietal and left posterior occipito-temporal regions. To explore the behavioural correlates of these systems, we measured the difference between reading speed for irregularly spelled words relative to pseudowords outside the scanner in fifteen of our subjects and correlated this measure with fMRI activation for reading familiar words. The faster the lexical reading the greater the activation in left posterior occipito-temporal and right inferior parietal regions. Conversely, the slower the lexical reading the greater the activation in left anterior occipito-temporal and left ventral inferior frontal regions. Thus, the double dissociation in irregular and pseudoword reading behaviour predicted the double dissociation in neuronal activation for reading familiar words. We discuss the implications of these results which may be important for understanding how reading is learnt in childhood or re-learnt following brain damage in adulthood.
doi:10.1016/j.neuroimage.2008.05.029
PMCID: PMC2724104  PMID: 18639469
Functional MRI; Regional covariance; Inter-subject variability; Overt reading; Language; Connectivity; Network; Occipito-temporal sulcus
9.  Segregation of Lexical and Sub-Lexical Reading Processes in the Left Perisylvian Cortex 
PLoS ONE  2012;7(11):e50665.
A fundamental issue in cognitive neuroscience is the existence of two major, sub-lexical and lexical, reading processes and their possible segregation in the left posterior perisylvian cortex. Using cortical electrostimulation mapping, we identified the cortical areas involved on reading either orthographically irregular words (lexical, “direct” process) or pronounceable pseudowords (sublexical, “indirect” process) in 14 right-handed neurosurgical patients while video-recording behavioral effects. Intraoperative neuronavigation system and Montreal Neurological Institute (MNI) stereotactic coordinates were used to identify the localization of stimulation sites. Fifty-one reading interference areas were found that affected either words (14 areas), or pseudo-words (11 areas), or both (26 areas). Forty-one (80%) corresponded to the impairment of the phonological level of reading processes. Reading processes involved discrete, highly localized perisylvian cortical areas with individual variability. MNI coordinates throughout the group exhibited a clear segregation according to the tested reading route; specific pseudo-word reading interferences were concentrated in a restricted inferior and anterior subpart of the left supramarginal gyrus (barycentre x = −68.1; y = −25.9; z = 30.2; Brodmann’s area 40) while specific word reading areas were located almost exclusively alongside the left superior temporal gyrus. Although half of the reading interferences found were nonspecific, the finding of specific lexical or sublexical interferences is new evidence that lexical and sublexical processes of reading could be partially supported by distinct cortical sub-regions despite their anatomical proximity. These data are in line with many brain activation studies that showed that left superior temporal and inferior parietal regions had a crucial role respectively in word and pseudoword reading and were core regions for dyslexia.
doi:10.1371/journal.pone.0050665
PMCID: PMC3511309  PMID: 23226349
10.  Word Processing differences between dyslexic and control children 
BMC Psychiatry  2006;6:5.
Background
The aim of this study was to investigate brain responses triggered by different wordclasses in dyslexic and control children. The majority of dyslexic children have difficulties to phonologically assemble a word from sublexical parts following grapheme-to-phoneme correspondences. Therefore, we hypothesised that dyslexic children should mainly differ from controls processing low frequent words that are unfamiliar to the reader.
Methods
We presented different wordclasses (high and low frequent words, pseudowords) in a rapid serial visual word (RSVP) design and performed wavelet analysis on the evoked activity.
Results
Dyslexic children had lower evoked power amplitudes and a higher spectral frequency for low frequent words compared to control children. No group differences were found for high frequent words and pseudowords. Control children had higher evoked power amplitudes and a lower spectral frequency for low frequent words compared to high frequent words and pseudowords. This pattern was not present in the dyslexic group.
Conclusion
Dyslexic children differed from control children only in their brain responses to low frequent words while showing no modulated brain activity in response to the three word types. This might support the hypothesis that dyslexic children are selectively impaired reading words that require sublexical processing. However, the lacking differences between word types raise the question if dyslexic children were able to process the words presented in rapid serial fashion in an adequate way. Therefore the present results should only be interpreted as evidence for a specific sublexical processing deficit with caution.
doi:10.1186/1471-244X-6-5
PMCID: PMC1373620  PMID: 16441886
11.  A Dual-Route Perspective on Brain Activation in Response to Visual Words: Evidence for a Length by Lexicality Interaction in the Visual Word Form Area (VWFA) 
NeuroImage  2009;49(3):2649-2661.
Based on our previous work, we expected the Visual Word Form Area (VWFA) in the left ventral visual pathway to be engaged by both whole-word recognition and by serial sublexical coding of letter strings. To examine this double function, a phonological lexical decision task (i.e., “Does xxx sound like an existing word?”) presented short and long letter strings of words, pseudohomophones, and pseudowords (e.g., Taxi, Taksi and Tazi). Main findings were that the length effect for words was limited to occipital regions and absent in the VWFA. In contrast, a marked length effect for pseudowords was found throughout the ventral visual pathway including the VWFA, as well as in regions presumably engaged by visual attention and silent-articulatory processes. The length by lexicality interaction on brain activation corresponds to well-established behavioral findings of a length by lexicality interaction on naming latencies and speaks for the engagement of the VWFA by both lexical and sublexical processes.
doi:10.1016/j.neuroimage.2009.10.082
PMCID: PMC2989181  PMID: 19896538
12.  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.
doi:10.3389/fpsyg.2012.00154
PMCID: PMC3356879  PMID: 22661961
letter-string processing; letter-position encoding; letter-identity encoding; letter transposition; letter substitution; reading acquisition; dyslexic children
13.  A Treatment Sequence for Phonological Alexia/Agraphia 
Purpose
Damage to left perisylvian cortex often results in impaired phonological processing abilities with written language profiles consistent with phonological alexia and phonological agraphia. The purpose of this article was to examine a behavioral treatment sequence for such individuals intended to strengthen phonological processing and links between phonology and orthography, as well as train a means to maximize use of residual orthographic and phonological knowledge for spelling.
Method
Two women with persistent impairments of written language and phonological processing following damage to left perisylvian cortical regions participated in this study. Both exhibited characteristic features of phonological alexia and agraphia in that reading and spelling performance for real words was better preserved than nonwords (lexicality effect). A 2-stage treatment protocol was administered to strengthen sublexical skills (phonological treatment) and to train interactive use of lexical and sublexical information to maximize spelling performance (interactive treatment).
Results
Both participants improved phonological processing abilities and reading/spelling via the sublexical route. They also improved spelling of real words and were able to detect and correct most residual errors using an electronic spelling aid. Conclusions: Behavioral treatment served to strengthen phonological skills supporting reading and spelling, and provided a functional compensatory strategy to overcome residual weaknesses.
doi:10.1044/1092-4388(2009/08-0229)
PMCID: PMC3522177  PMID: 20360466
aphasia; dyslexia; dysgraphia; rehabilitation; stroke
14.  A dual-route perspective on eye movements of dyslexic readers 
Cognition  2010;115(3):367-379.
This study assessed eye movement abnormalities of adolescent dyslexic readers and interpreted the findings by linking the dual-route model of single word reading with the E-Z Reader model of eye movement control during silent sentence reading. A dysfunction of the lexical route was assumed to account for a reduced number of words which received only a single fixation or which were skipped and for the increased number of words with multiple fixations and a marked effect of word length on gaze duration. This pattern was interpreted as a frequent failure of orthographic whole-word recognition (based on orthographic lexicon entries) and on reliance on serial sublexical processing instead. Inefficiency of the lexical route was inferred from prolonged gaze durations for singly fixated words. These findings were related to the E-Z Reader model of eye movement control. Slow activation of word phonology accounted for the low skipping rate of dyslexic readers. Frequent reliance on sublexical decoding was inferred from a tendency to fixate word beginnings and from short forward saccades. Overall, the linkage of the dual-route model of single word reading and a model of eye movement control led to a useful framework for understanding eye movement abnormalities of dyslexic readers.
doi:10.1016/j.cognition.2009.11.004
PMCID: PMC2976468  PMID: 20227686
Developmental dyslexia; Dual-route model of reading; Eye movement control; E-Z Reader; Lexical route; Sublexical route
15.  Top-down modulation of ventral occipito-temporal responses during visual word recognition 
Neuroimage  2011;55(3-3):1242-1251.
Although interactivity is considered a fundamental principle of cognitive (and computational) models of reading, it has received far less attention in neural models of reading that instead focus on serial stages of feed-forward processing from visual input to orthographic processing to accessing the corresponding phonological and semantic information. In particular, the left ventral occipito-temporal (vOT) cortex is proposed to be the first stage where visual word recognition occurs prior to accessing nonvisual information such as semantics and phonology. We used functional magnetic resonance imaging (fMRI) to investigate whether there is evidence that activation in vOT is influenced top-down by the interaction of visual and nonvisual properties of the stimuli during visual word recognition tasks. Participants performed two different types of lexical decision tasks that focused on either visual or nonvisual properties of the word or word-like stimuli. The design allowed us to investigate how vOT activation during visual word recognition was influenced by a task change to the same stimuli and by a stimulus change during the same task. We found both stimulus- and task-driven modulation of vOT activation that can only be explained by top-down processing of nonvisual aspects of the task and stimuli. Our results are consistent with the hypothesis that vOT acts as an interface linking visual form with nonvisual processing in both bottom up and top down directions. Such interactive processing at the neural level is in agreement with cognitive and computational models of reading but challenges some of the assumptions made by current neuro-anatomical models of reading.
Research Highlights
►Activation in left vOT was modulated by task, even when stimuli were held constant. ►Activation in vOT was also modulated by stimulus differences. ►Activation in vOT could not be predicted by differences in response times. ►Top-down input to vOT is required to explain these effects.
doi:10.1016/j.neuroimage.2011.01.001
PMCID: PMC3221051  PMID: 21232615
Reading; Fusiform gyrus; fMRI; Lexical decision; Feedback
16.  N170 ERPs could represent a logographic processing strategy in visual word recognition 
Background
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.
Methods
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.
Results
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.
Conclusion
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.
doi:10.1186/1744-9081-3-21
PMCID: PMC1884163  PMID: 17451598
17.  Neural Correlates of Sublexical Processing in Phonological Working Memory 
Journal of cognitive neuroscience  2010;23(4):961-977.
This study investigated links between working memory and speech processing systems. We used delayed pseudoword repetition in fMRI to investigate the neural correlates of sublexical structure in phonological working memory (pWM). We orthogonally varied the number of syllables and consonant clusters in auditory pseudowords and measured the neural responses to these manipulations under conditions of covert rehearsal (Experiment 1). A left-dominant network of temporal and motor cortex showed increased activity for longer items, with motor cortex only showing greater activity concomitant with adding consonant clusters. An individual-differences analysis revealed a significant positive relationship between activity in the angular gyrus and the hippocampus, and accuracy on pseudoword repetition. As models of pWM stipulate that its neural correlates should be activated during both perception and production/rehearsal [Buchsbaum, B. R., & D’Esposito, M. The search for the phonological store: From loop to convolution. Journal of Cognitive Neuroscience, 20, 762–778, 2008; Jacquemot, C., & Scott, S. K. What is the relationship between phonological short-term memory and speech processing? Trends in Cognitive Sciences, 10, 480–486, 2006; Baddeley, A. D., & Hitch, G. Working memory. In G. H. Bower (Ed.), The psychology of learning and motivation: Advances in research and theory (Vol. 8, pp. 47–89). New York: Academic Press, 1974], we further assessed the effects of the two factors in a separate passive listening experiment (Experiment 2). In this experiment, the effect of the number of syllables was concentrated in posterior–medial regions of the supratemporal plane bilaterally, although there was no evidence of a significant response to added clusters. Taken together, the results identify the planum temporale as a key region in pWM; within this region, representations are likely to take the form of auditory or audiomotor “templates” or “chunks” at the level of the syllable [Papoutsi, M., de Zwart, J. A., Jansma, J. M., Pickering, M. J., Bednar, J. A., & Horwitz, B. From phonemes to articulatory codes: an fMRI study of the role of Broca’s area in speech production. Cerebral Cortex, 19, 2156–2165, 2009; Warren, J. E., Wise, R. J. S., & Warren, J. D. Sounds do-able: auditory–motor transformations and the posterior temporal plane. Trends in Neurosciences, 28, 636–643, 2005; Griffiths, T. D., & Warren, J. D. The planum temporale as a computational hub. Trends in Neurosciences, 25, 348–353, 2002], whereas more lateral structures on the STG may deal with phonetic analysis of the auditory input [Hickok, G. The functional neuroanatomy of language. Physics of Life Reviews, 6, 121–143, 2009].
doi:10.1162/jocn.2010.21491
PMCID: PMC3376447  PMID: 20350182
18.  Functional Brain Activation Differences in School-Age Children with Speech Sound Errors: Speech and Print Processing 
Purpose
To examine neural response to spoken and printed language in children with speech sound errors (SSE).
Method
Functional MRI was used to compare processing of auditorily and visually presented words and pseudowords in 17 children with SSE, ages 8;6–10;10, to 17 matched controls.
Results
When processing spoken words and pseudowords, the SSE group showed less activation than typically speaking controls in left middle temporal gyrus. They also showed greater activation than controls in several cortical and subcortical regions (e.g., left superior temporal gyrus, globus pallidus, insula, fusiform and bilateral parietal regions). In response to printed words and pseudowords, children with SSE had greater activation than controls in regions including bilateral fusiform and anterior cingulate. Some differences were found in both speech and print processing that that may be associated with children with SSE failing to show common patterns of task-induced deactivation and/or attentional resource allocation.
Conclusion
Compared with controls, children with SSE appear to rely more upon several dorsal speech perception regions and less on ventral speech perception regions. When processing print, numerous regions were observed to be activated more for the SSE group than for controls.
doi:10.1044/1092-4388(2011/11-0056)
PMCID: PMC3427927  PMID: 22232410
19.  Efficient Visual Object and Word Recognition Relies on High Spatial Frequency Coding in the Left Posterior Fusiform Gyrus: Evidence from a Case-Series of Patients with Ventral Occipito-Temporal Cortex Damage 
Cerebral Cortex (New York, NY)  2012;23(11):2568-2580.
Recent visual neuroscience investigations suggest that ventral occipito-temporal cortex is retinotopically organized, with high acuity foveal input projecting primarily to the posterior fusiform gyrus (pFG), making this region crucial for coding high spatial frequency information. Because high spatial frequencies are critical for fine-grained visual discrimination, we hypothesized that damage to the left pFG should have an adverse effect not only on efficient reading, as observed in pure alexia, but also on the processing of complex non-orthographic visual stimuli. Consistent with this hypothesis, we obtained evidence that a large case series (n = 20) of patients with lesions centered on left pFG: 1) Exhibited reduced sensitivity to high spatial frequencies; 2) demonstrated prolonged response latencies both in reading (pure alexia) and object naming; and 3) were especially sensitive to visual complexity and similarity when discriminating between novel visual patterns. These results suggest that the patients' dual reading and non-orthographic recognition impairments have a common underlying mechanism and reflect the loss of high spatial frequency visual information normally coded in the left pFG.
doi:10.1093/cercor/bhs224
PMCID: PMC3792736  PMID: 22923086
foveal/parafoveal vision; fusiform gyrus; letter-by-letter reading; pure alexia; spatial frequency; ventral occipito-temporal cortex; visual recognition
20.  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).
doi:10.1016/j.cortex.2010.06.004
PMCID: PMC3073233  PMID: 20650450
21.  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).
doi:10.1016/j.cortex.2010.06.004
PMCID: PMC3073233  PMID: 20650450
Developmental dyslexia; fMRI; Reading; Phonological lexical decision; Dual-route
22.  Neural correlates of sentence reading in children with reading difficulties 
Neuroreport  2011;22(14):674-678.
In contrast to word-level skills, the neural basis of sentence comprehension in children with reading difficulties is not well understood. Using magnetic source imaging, we investigated the spatiotemporal dynamics of regional activity associated with silent passage reading in non-impaired and students with reading difficulties. The latter exhibited underactivation of the temporo-parietal and visual cortices, bilaterally, as well as of the left posterior cingulate region. Late activity in left temporo-parietal and ventral occipito-temporal regions was found to be a significant predictor of individual reading ability in non-impaired, but not in students with reading difficulties. These findings support the notion that reduced temporo-parietal activation during word reading in context, is a hallmark of the functional deficit in reading disability.
doi:10.1097/WNR.0b013e328349ecf7
PMCID: PMC3164957  PMID: 21878791
Dyslexia; comprehension; temporoparietal cortex; magnetoencephalography
23.  Automatic Top-Down Processing Explains Common Left Occipito-Temporal Responses to Visual Words and Objects 
Cerebral Cortex (New York, NY)  2010;21(1):103-114.
Previous studies have demonstrated that a region in the left ventral occipito-temporal (LvOT) cortex is highly selective to the visual forms of written words and objects relative to closely matched visual stimuli. Here, we investigated why LvOT activation is not higher for reading than picture naming even though written words and pictures of objects have grossly different visual forms. To compare neuronal responses for words and pictures within the same LvOT area, we used functional magnetic resonance imaging adaptation and instructed participants to name target stimuli that followed briefly presented masked primes that were either presented in the same stimulus type as the target (word–word, picture–picture) or a different stimulus type (picture–word, word–picture). We found that activation throughout posterior and anterior parts of LvOT was reduced when the prime had the same name/response as the target irrespective of whether the prime-target relationship was within or between stimulus type. As posterior LvOT is a visual form processing area, and there was no visual form similarity between different stimulus types, we suggest that our results indicate automatic top-down influences from pictures to words and words to pictures. This novel perspective motivates further investigation of the functional properties of this intriguing region.
doi:10.1093/cercor/bhq063
PMCID: PMC3000565  PMID: 20413450
language; priming; repetition suppression; visual system; word–object recognition
24.  Spatial attention in written word perception 
The role of attention in visual word recognition and reading aloud is a long debated issue. Studies of both developmental and acquired reading disorders provide growing evidence that spatial attention is critically involved in word reading, in particular for the phonological decoding of unfamiliar letter strings. However, studies on healthy participants have produced contrasting results. The aim of this study was to investigate how the allocation of spatial attention may influence the perception of letter strings in skilled readers. High frequency words (HFWs), low frequency words and pseudowords were briefly and parafoveally presented either in the left or the right visual field. Attentional allocation was modulated by the presentation of a spatial cue before the target string. Accuracy in reporting the target string was modulated by the spatial cue but this effect varied with the type of string. For unfamiliar strings, processing was facilitated when attention was focused on the string location and hindered when it was diverted from the target. This finding is consistent the assumptions of the CDP+ model of reading aloud, as well as with familiarity sensitivity models that argue for a flexible use of attention according with the specific requirements of the string. Moreover, we found that processing of HFWs was facilitated by an extra-large focus of attention. The latter result is consistent with the hypothesis that a broad distribution of attention is the default mode during reading of familiar words because it might optimally engage the broad receptive fields of the highest detectors in the hierarchical system for visual word recognition.
doi:10.3389/fnhum.2014.00042
PMCID: PMC3918588  PMID: 24574990
visual word recognition; visuo-spatial attention; spatial cueing; reading aloud; phonological decoding; attentional focusing
25.  Does IQ affect the functional brain network involved in pseudoword reading in students with reading disability? A magnetoencephalography study 
The study examined whether individual differences in performance and verbal IQ affect the profiles of reading-related regional brain activation in 127 students experiencing reading difficulties and typical readers. Using magnetoencephalography in a pseudoword read-aloud task, we compared brain activation profiles of students experiencing word-level reading difficulties who did (n = 29) or did not (n = 36) meet the IQ-reading achievement discrepancy criterion. Typical readers assigned to a lower-IQ (n = 18) or a higher IQ (n = 44) subgroup served as controls. Minimum norm estimates of regional cortical activity revealed that the degree of hypoactivation in the left superior temporal and supramarginal gyri in both RD subgroups was not affected by IQ. Moreover, IQ did not moderate the positive association between degree of activation in the left fusiform gyrus and phonological decoding ability. We did find, however, that the hypoactivation of the left pars opercularis in RD was restricted to lower-IQ participants. In accordance with previous morphometric and fMRI studies, degree of activity in inferior frontal, and inferior parietal regions correlated with IQ across reading ability subgroups. Results are consistent with current views questioning the relevance of IQ-discrepancy criteria in the diagnosis of dyslexia.
doi:10.3389/fnhum.2013.00932
PMCID: PMC3884211  PMID: 24409136
magnetoencephalography; functional brain imaging; reading; dyslexia; intelligence

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