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1.  Does pronounceability modulate the letter string deficit of children with dyslexia? A study with the rate and amount model 
Frontiers in Psychology  2014;5:1353.
The locus of the deficit of children with dyslexia in dealing with strings of letters may be a deficit at a pre-lexical graphemic level or an inability to bind orthographic and phonological information. We evaluate these alternative hypotheses in two experiments by examining the role of stimulus pronounceability in a lexical decision task (LDT) and in a forced-choice letter discrimination task (Reicher–Wheeler paradigm). Seventeen fourth grade children with dyslexia and 24 peer control readers participated to two experiments. In the LDT children were presented with high-, low-frequency words, pronounceable pseudowords (such as DASU) and unpronounceable non-words (such as RNGM) of 4-, 5-, or 6- letters. No sign of group by pronounceability interaction was found when over-additivity was taken into account. Children with dyslexia were impaired when they had to process strings, not only of pronounceable stimuli but also of unpronounceable stimuli, a deficit well accounted for by a single global factor. Complementary results were obtained with the Reicher–Wheeler paradigm: both groups of children gained in accuracy in letter discrimination in the context of pronounceable primes (words and pseudowords) compared to unpronounceable primes (non-words). No global factor was detected in this task which requires the discrimination between a target letter and a competitor but does not involve simultaneous letter string processing. Overall, children with dyslexia show a selective difficulty in simultaneously processing a letter string as a whole, independent of its pronounceability; however, when the task involves isolated letter processing, also these children can make use of the ortho-phono-tactic information derived from a previously seen letter string. This pattern of findings is in keeping with the idea that an impairment in pre-lexical graphemic analysis may be a core deficit in developmental dyslexia.
PMCID: PMC4251298  PMID: 25520680
developmental dyslexia; lexical decision; Reicher–Wheeler paradigm; pronounceability; global factor; letter string
2.  Influence of context-sensitive rules on the formation of orthographic representations in Spanish dyslexic children 
Frontiers in Psychology  2014;5:1409.
Spanish-speaking developmental dyslexics are mainly characterized by poor reading fluency. One reason for this lack of fluency could be a difficulty in creating and accessing lexical representations, because, as the self-teaching theory suggest, it is necessary to develop orthographic representations to use direct reading (Share, 1995). It is possible that this difficulty to acquire orthographic representations can be specifically related to words that contain context-sensitive graphemes, since it has been demonstrated that reading is affected by this kind of graphemes (Barca et al., 2007). In order to test this possibility we compared a group of dyslexic children with a group of normal readers (9–13 years), in a task of repeated reading. Pseudo-words (half short and half long) with simple and contextual dependent rules were used. The length effect reduction on the reading speed, after repeated exposure, was considered an indicator of orthographic representation development, as the length effect is strong when reading unknown words, but absent when reading familiar words. The results show that dyslexic children have difficulties in developing orthographic representations, not only with context-sensitive graphemes, but also with simple graphemes. In contrast to the control children, in the dyslexic group differences between reading times for short and long stimuli remained without significant changes after six presentations. Besides, this happened with sensitive context rules and also with simple grapheme–phoneme conversion rules. On the other hand, response and articulation times were greatly affected by length in dyslexic children, indicating the use of serial reading. Results suggest that the problems related to storing orthographic representations could be caused by a learning deficit, independently of whether the word contained context-sensitive rules or not.
PMCID: PMC4255502  PMID: 25538659
dyslexia; orthographic representations; fluency; transparent orthography; context-sensitive rules
3.  The locus of impairment in English developmental letter position dyslexia 
Many children with reading difficulties display phonological deficits and struggle to acquire non-lexical reading skills. However, not all children with reading difficulties have these problems, such as children with selective letter position dyslexia (LPD), who make excessive migration errors (such as reading slime as “smile”). Previous research has explored three possible loci for the deficit – the phonological output buffer, the orthographic input lexicon, and the orthographic-visual analysis stage of reading. While there is compelling evidence against a phonological output buffer and orthographic input lexicon deficit account of English LPD, the evidence in support of an orthographic-visual analysis deficit is currently limited. In this multiple single-case study with three English-speaking children with developmental LPD, we aimed to both replicate and extend previous findings regarding the locus of impairment in English LPD. First, we ruled out a phonological output buffer and an orthographic input lexicon deficit by administering tasks that directly assess phonological processing and lexical guessing. We then went on to directly assess whether or not children with LPD have an orthographic-visual analysis deficit by modifying two tasks that have previously been used to localize processing at this level: a same-different decision task and a non-word reading task. The results from these tasks indicate that LPD is most likely caused by a deficit specific to the coding of letter positions at the orthographic-visual analysis stage of reading. These findings provide further evidence for the heterogeneity of dyslexia and its underlying causes.
PMCID: PMC4042363  PMID: 24917802
phonological output deficit; orthographic input lexicon deficit; orthographic-visual analysis deficit; migration errors; substitution errors; developmental dyslexia
4.  Modeling individual differences in text reading fluency: a different pattern of predictors for typically developing and dyslexic readers 
Frontiers in Psychology  2014;5:1374.
This study was aimed at predicting individual differences in text reading fluency. The basic proposal included two factors, i.e., the ability to decode letter strings (measured by discrete pseudo-word reading) and integration of the various sub-components involved in reading (measured by Rapid Automatized Naming, RAN). Subsequently, a third factor was added to the model, i.e., naming of discrete digits. In order to use homogeneous measures, all contributing variables considered the entire processing of the item, including pronunciation time. The model, which was based on commonality analysis, was applied to data from a group of 43 typically developing readers (11- to 13-year-olds) and a group of 25 chronologically matched dyslexic children. In typically developing readers, both orthographic decoding and integration of reading sub-components contributed significantly to the overall prediction of text reading fluency. The model prediction was higher (from ca. 37 to 52% of the explained variance) when we included the naming of discrete digits variable, which had a suppressive effect on pseudo-word reading. In the dyslexic readers, the variance explained by the two-factor model was high (69%) and did not change when the third factor was added. The lack of a suppression effect was likely due to the prominent individual differences in poor orthographic decoding of the dyslexic children. Analyses on data from both groups of children were replicated by using patches of colors as stimuli (both in the RAN task and in the discrete naming task) obtaining similar results. We conclude that it is possible to predict much of the variance in text-reading fluency using basic processes, such as orthographic decoding and integration of reading sub-components, even without taking into consideration higher-order linguistic factors such as lexical, semantic and contextual abilities. The approach validity of using proximal vs. distal causes to predict reading fluency is discussed.
PMCID: PMC4235379  PMID: 25477856
reading; individual differences; dyslexia; suppression effect; RAN; vocal reaction times
5.  Reduced Neural Integration of Letters and Speech Sounds in Dyslexic Children Scales with Individual Differences in Reading Fluency 
PLoS ONE  2014;9(10):e110337.
The acquisition of letter-speech sound associations is one of the basic requirements for fluent reading acquisition and its failure may contribute to reading difficulties in developmental dyslexia. Here we investigated event-related potential (ERP) measures of letter-speech sound integration in 9-year-old typical and dyslexic readers and specifically test their relation to individual differences in reading fluency. We employed an audiovisual oddball paradigm in typical readers (n = 20), dysfluent (n = 18) and severely dysfluent (n = 18) dyslexic children. In one auditory and two audiovisual conditions the Dutch spoken vowels/a/and/o/were presented as standard and deviant stimuli. In audiovisual blocks, the letter ‘a’ was presented either simultaneously (AV0), or 200 ms before (AV200) vowel sound onset. Across the three children groups, vowel deviancy in auditory blocks elicited comparable mismatch negativity (MMN) and late negativity (LN) responses. In typical readers, both audiovisual conditions (AV0 and AV200) led to enhanced MMN and LN amplitudes. In both dyslexic groups, the audiovisual LN effects were mildly reduced. Most interestingly, individual differences in reading fluency were correlated with MMN latency in the AV0 condition. A further analysis revealed that this effect was driven by a short-lived MMN effect encompassing only the N1 window in severely dysfluent dyslexics versus a longer MMN effect encompassing both the N1 and P2 windows in the other two groups. Our results confirm and extend previous findings in dyslexic children by demonstrating a deficient pattern of letter-speech sound integration depending on the level of reading dysfluency. These findings underscore the importance of considering individual differences across the entire spectrum of reading skills in addition to group differences between typical and dyslexic readers.
PMCID: PMC4199667  PMID: 25329388
6.  Age, dyslexia subtype and comorbidity modulate rapid auditory processing in developmental dyslexia 
The nature of Rapid Auditory Processing (RAP) deficits in dyslexia remains debated, together with the specificity of the problem to certain types of stimuli and/or restricted subgroups of individuals. Following the hypothesis that the heterogeneity of the dyslexic population may have led to contrasting results, the aim of the study was to define the effect of age, dyslexia subtype and comorbidity on the discrimination and reproduction of non-verbal tone sequences. Participants were 46 children aged 8–14 (26 with dyslexia, subdivided according to age, presence of a previous language delay, and type of dyslexia). Experimental tasks were a Temporal Order Judgment (TOJ) (manipulating tone length, ISI and sequence length), and a Pattern Discrimination Task. Dyslexic children showed general RAP deficits. Tone length and ISI influenced dyslexic and control children's performance in a similar way, but dyslexic children were more affected by an increase from 2 to 5 sounds. As to age, older dyslexic children's difficulty in reproducing sequences of 4 and 5 tones was similar to that of normally reading younger (but not older) children. In the analysis of subgroup profiles, the crucial variable appears to be the advantage, or lack thereof, in processing long vs. short sounds. Dyslexic children with a previous language delay obtained the lowest scores in RAP measures, but they performed worse with shorter stimuli, similar to control children, while dyslexic-only children showed no advantage for longer stimuli. As to dyslexia subtype, only surface dyslexics improved their performance with longer stimuli, while phonological dyslexics did not. Differential scores for short vs. long tones and for long vs. short ISIs predict non-word and word reading, respectively, and the former correlate with phonemic awareness. In conclusion, the relationship between non-verbal RAP, phonemic skills and reading abilities appears to be characterized by complex interactions with subgroup characteristics.
PMCID: PMC4032942  PMID: 24904356
developmental dyslexia; subgroups; rapid auditory processing; language impairment; dyslexia subtypes
7.  A Common Left Occipito-Temporal Dysfunction in Developmental Dyslexia and Acquired Letter-By-Letter Reading? 
PLoS ONE  2010;5(8):e12073.
We used fMRI to examine functional brain abnormalities of German-speaking dyslexics who suffer from slow effortful reading but not from a reading accuracy problem. Similar to acquired cases of letter-by-letter reading, the developmental cases exhibited an abnormal strong effect of length (i.e., number of letters) on response time for words and pseudowords.
Corresponding to lesions of left occipito-temporal (OT) regions in acquired cases, we found a dysfunction of this region in our developmental cases who failed to exhibit responsiveness of left OT regions to the length of words and pseudowords. This abnormality in the left OT cortex was accompanied by absent responsiveness to increased sublexical reading demands in phonological inferior frontal gyrus (IFG) regions. Interestingly, there was no abnormality in the left superior temporal cortex which—corresponding to the onological deficit explanation—is considered to be the prime locus of the reading difficulties of developmental dyslexia cases.
The present functional imaging results suggest that developmental dyslexia similar to acquired letter-by-letter reading is due to a primary dysfunction of left OT regions.
PMCID: PMC2920311  PMID: 20711448
8.  The Temporal Dynamics of Implicit Processing of Non-Letter, Letter, and Word-Forms in the Human Visual Cortex 
The decoding of visually presented line segments into letters, and letters into words, is critical to fluent reading abilities. Here we investigate the temporal dynamics of visual orthographic processes, focusing specifically on right hemisphere contributions and interactions between the hemispheres involved in the implicit processing of visually presented words, consonants, false fonts, and symbolic strings. High-density EEG was recorded while participants detected infrequent, simple, perceptual targets (dot strings) embedded amongst a of character strings. Beginning at 130 ms, orthographic and non-orthographic stimuli were distinguished by a sequence of ERP effects over occipital recording sites. These early latency occipital effects were dominated by enhanced right-sided negative-polarity activation for non-orthographic stimuli that peaked at around 180 ms. This right-sided effect was followed by bilateral positive occipital activity for false-fonts, but not symbol strings. Moreover the size of components of this later positive occipital wave was inversely correlated with the right-sided ROcc180 wave, suggesting that subjects who had larger early right-sided activation for non-orthographic stimuli had less need for more extended bilateral (e.g., interhemispheric) processing of those stimuli shortly later. Additional early (130–150 ms) negative-polarity activity over left occipital cortex and longer-latency centrally distributed responses (>300 ms) were present, likely reflecting implicit activation of the previously reported ‘visual-word-form’ area and N400-related responses, respectively. Collectively, these results provide a close look at some relatively unexplored portions of the temporal flow of information processing in the brain related to the implicit processing of potentially linguistic information and provide valuable information about the interactions between hemispheres supporting visual orthographic processing.
PMCID: PMC2796900  PMID: 20046826
word reading; ERPs; visual cortex; visual orthography
9.  Do Dyslexics Misread a ROWS for a ROSE? 
Reading and writing  2012;26(3):381-402.
Insufficient knowledge of the subtle relations between words’ spellings and their phonology is widely held to be the primary limitation in developmental dyslexia. In the present study the influence of phonology on a semantic-based reading task was compared for groups of readers with and without dyslexia. As many studies have shown, skilled readers make phonology-based false-positive errors to homophones and pseudohomophones in the semantic categorization task. The basic finding was extended to children, teens, and adults with dyslexia from familial and clinically-referred samples. Dyslexics showed the same overall pattern of phonology errors and the results were consistent across dyslexia samples, across age groups, and across experimental conditions using word and nonword homophone foils. The dyslexic groups differed from chronological-age matched controls by having elevated false-positive homophone error rates overall, and weaker effects of baseword frequency. Children with dyslexia also made more false-positive errors to spelling control foils. These findings suggest that individuals with dyslexia make use of phonology when making semantic decisions both to word homophone and non-word pseudohomophone foils and that dyslexics lack adequate knowledge of actual word spellings, compared to chronological-age and reading-level matched control participants.
PMCID: PMC4004072  PMID: 24791075
developmental dyslexia; homophone; pseudohomophone; semantic judgments; word frequency
10.  Investigating the role of visual and auditory search in reading and developmental dyslexia 
It has been suggested that auditory and visual sequential processing deficits contribute to phonological disorders in developmental dyslexia. As an alternative explanation to a phonological deficit as the proximal cause for reading disorders, the visual attention span hypothesis (VA Span) suggests that difficulties in processing visual elements simultaneously lead to dyslexia, regardless of the presence of a phonological disorder. In this study, we assessed whether deficits in processing simultaneously displayed visual or auditory elements is linked to dyslexia associated with a VA Span impairment. Sixteen children with developmental dyslexia and 16 age-matched skilled readers were assessed on visual and auditory search tasks. Participants were asked to detect a target presented simultaneously with 3, 9, or 15 distracters. In the visual modality, target detection was slower in the dyslexic children than in the control group on a “serial” search condition only: the intercepts (but not the slopes) of the search functions were higher in the dyslexic group than in the control group. In the auditory modality, although no group difference was observed, search performance was influenced by the number of distracters in the control group only. Within the dyslexic group, not only poor visual search (high reaction times and intercepts) but also low auditory search performance (d′) strongly correlated with poor irregular word reading accuracy. Moreover, both visual and auditory search performance was associated with the VA Span abilities of dyslexic participants but not with their phonological skills. The present data suggests that some visual mechanisms engaged in “serial” search contribute to reading and orthographic knowledge via VA Span skills regardless of phonological skills. The present results further open the question of the role of auditory simultaneous processing in reading as well as its link with VA Span skills.
PMCID: PMC3782690  PMID: 24093014
dyslexia; reading; visual search; auditory search; attention; temporal processing; visual attention span; phonology
11.  Letter-transposition effects are not universal: The impact of transposing letters in Hebrew 
Journal of memory and language  2009;61(3):285-302.
We examined the effects of letter transposition in Hebrew in three masked-priming experiments. Hebrew, like English has an alphabetic orthography where sequential and contiguous letter strings represent phonemes. However, being a Semitic language it has a non-concatenated morphology that is based on root derivations. Experiment 1 showed that transposed-letter (TL) root primes inhibited responses to targets derived from the non-transposed root letters, and that this inhibition was unrelated to relative root frequency. Experiment 2 replicated this result and showed that if the transposed letters of the root created a nonsense-root that had no lexical representation, then no inhibition and no facilitation were obtained. Finally, Experiment 3 demonstrated that in contrast to English, French, or Spanish, TL nonword primes did not facilitate recognition of targets, and when the root letters embedded in them consisted of a legal root morpheme, they produced inhibition. These results suggest that lexical space in alphabetic orthographies may be structured very differently in different languages if their morphological structure diverges qualitatively. In Hebrew, lexical space is organized according to root families rather than simple orthographic structure, so that all words derived from the same root are interconnected or clustered together, independent of overall orthographic similarity.
PMCID: PMC2748959  PMID: 20161017
Morphology; Letter Transposition; Hebrew; Masked-Priming
12.  Are All Letters Really Processed Equally and in Parallel? Further Evidence of a Robust First Letter Advantage 
Acta psychologica  2013;144(2):10.1016/j.actpsy.2013.07.018.
This present study examined accuracy and response latency of letter processing as a function of position within a horizontal array. In a series of 4 Experiments, target-strings were briefly (33 ms for Experiment 1 to 3, 83 ms for Experiment 4) displayed and both forward and backward masked. Participants then made a two alternative forced choice. The two alternative responses differed just in one element of the string, and position of mismatch was systematically manipulated. In Experiment 1, words of different lengths (from 3 to 6 letters) were presented in separate blocks. Across different lengths, there was a robust advantage in performance when the alternative response was different for the letter occurring at the first position, compared to when the difference occurred at any other position. Experiment 2 replicated this finding with the same materials used in Experiment 1, but with words of different lengths randomly intermixed within blocks. Experiment 3 provided evidence of the first position advantage with legal nonwords and strings of consonants, but did not provide any first position advantage for non-alphabetic symbols. The lack of a first position advantage for symbols was replicated in Experiment 4, where target-strings were displayed for a longer duration (83 ms). Taken together these results suggest that the first position advantage is a phenomenon that occurs specifically and selectively for letters, independent of lexical constraints. We argue that the results are consistent with models that assume a processing advantage for coding letters in the first position, and are inconsistent with the commonly held assumption in visual word recognition models that letters are equally processed in parallel independent of letter position.
PMCID: PMC3859888  PMID: 24012723
letter-processing; serial position; visual word recognition
13.  Reading networks in children with dyslexia compared to children with ocular motility disturbances revealed by fMRI 
Key Points Dyslexia is a neurological disorder with a genetic origin, but the underlying biological and cognitive causes are still being investigated.This study compares the brain activation pattern while reading in Spanish, a semitransparent language, in three groups of children: typically developing readers, dyslexic readers and readers with functional monocular vision.Based on our results Dyslexia would be a neurological disorder not related to vision impairments and would require a multidisciplinary treatment based on improving phonological awareness and language development.
Developmental dyslexia is a neurological disorder the underlying biological and cognitive causes of which are still being investigated, a key point, because the findings will determine the best therapeutic approach to use. Using functional magnetic resonance imaging, we studied the brain activation pattern while reading in the language-related cortical areas from the two reading routes, phonological and orthographic, and the strength of their association with reading scores in 66 Spanish-speaking children aged 9–12 years divided into three groups: typically developing readers (controls), dyslexic readers and readers with monocular vision due to ocular motility disorders but with normal reading development, to assess whether (or not) the neuronal network for reading in children with dyslexia has similarities with that in children with impaired binocular vision due to ocular motility disorders. We found that Spanish-speaking children with dyslexia have a brain circuit for reading that differs from that in children with monocular vision. Individuals with dyslexia tend to hypoactivate some of the language-related areas in the left hemisphere engaged by the phonological route, especially the visual word form area and left Wernicke's area, and try to compensate this deficit by activating language-related areas related to the orthographic route, such as the anterior part of the visual word form area and the posterior part of both middle temporal gyri. That is, they seem to compensate for impairment in the phonological route through orthographic routes of both hemispheres. Our results suggest that ocular motility disturbances do not play a causal role in dyslexia. Dyslexia seems to be a neurological disorder that is unrelated to vision impairments and requires early recognition and multidisciplinary treatment, based on improving phonological awareness and language development, to achieve the best possible outcome.
PMCID: PMC4237045  PMID: 25477808
developmental dyslexia; fMRI; ocular motility disorders; paradigm; pseudoword
14.  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.
PMCID: PMC3612367  PMID: 23454070
visual-word recognition; position coding; ERPs; word-frequency; transposed-letter effects
15.  Speed and accuracy of dyslexic versus typical word recognition: an eye-movement investigation 
Frontiers in Psychology  2014;5:1129.
Developmental dyslexia is often characterized by a dual deficit in both word recognition accuracy and general processing speed. While previous research into dyslexic word recognition may have suffered from speed-accuracy trade-off, the present study employed a novel eye-tracking task that is less prone to such confounds. Participants (10 dyslexics and 12 controls) were asked to look at real word stimuli, and to ignore simultaneously presented non-word stimuli, while their eye-movements were recorded. Improvements in word recognition accuracy over time were modeled in terms of a continuous non-linear function. The words' rhyme consistency and the non-words' lexicality (unpronounceable, pronounceable, pseudohomophone) were manipulated within-subjects. Speed-related measures derived from the model fits confirmed generally slower processing in dyslexics, and showed a rhyme consistency effect in both dyslexics and controls. In terms of overall error rate, dyslexics (but not controls) performed less accurately on rhyme-inconsistent words, suggesting a representational deficit for such words in dyslexics. Interestingly, neither group showed a pseudohomophone effect in speed or accuracy, which might call the task-independent pervasiveness of this effect into question. The present results illustrate the importance of distinguishing between speed- vs. accuracy-related effects for our understanding of dyslexic word recognition.
PMCID: PMC4191135  PMID: 25346708
visual word recognition; dyslexia; speed-accuracy trade-off; consistency; phonology
16.  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
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 (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
18.  The visual magnocellular deficit in Chinese-speaking children with developmental dyslexia 
Many alphabetic studies have evidenced that individuals with developmental dyslexia (DD) have deficits in visual magnocellular (M) pathway. However, there are few studies to investigate the M function of Chinese DD. Chinese is a logographic language, and Chinese characters are complicated in structure. Visual skills and orthographic processing abilities are particularly important for efficient reading in Chinese as compared to alphabetic languages. Therefore, it is necessary to investigate the visual M function of Chinese DD and whether the M function was associated with orthographic skills. In the present study, 26 dyslexic children (mean age: 10.03 years) and 27 age-matched normal children (mean age: 10.37 years) took part in a coherent motion (CM) detection task and an orthographic awareness test. The results showed that dyslexic children had a significantly higher threshold than age-matched children in CM detection task. Meanwhile, children with DD responded more slowly in orthographic awareness test, although the group difference was marginally significant. The results suggested that Chinese dyslexics had deficits both in visual M pathway processing and orthographic processing. In order to investigate the relationship between M function and orthographic skills, we made a correlation analysis between CM threshold and orthographic awareness by merging performance of dyslexic children and age-matched children. The results revealed that CM thresholds were positively correlated with reaction times in orthographic awareness test, suggesting that better M function was related to better orthographic processing skills.
PMCID: PMC4084406  PMID: 25071661
developmental dyslexia; magnocellular pathway; coherent motion detection; orthographic processing skills; Chinese reading
19.  Reading component skills in dyslexia: word recognition, comprehension and processing speed 
Frontiers in Psychology  2014;5:1339.
The cognitive model of reading comprehension (RC) posits that RC is a result of the interaction between decoding and linguistic comprehension. Recently, the notion of decoding skill was expanded to include word recognition. In addition, some studies suggest that other skills could be integrated into this model, like processing speed, and have consistently indicated that this skill influences and is an important predictor of the main components of the model, such as vocabulary for comprehension and phonological awareness of word recognition. The following study evaluated the components of the RC model and predictive skills in children and adolescents with dyslexia. 40 children and adolescents (8–13 years) were divided in a Dyslexic Group (DG; 18 children, MA = 10.78, SD = 1.66) and control group (CG 22 children, MA = 10.59, SD = 1.86). All were students from the 2nd to 8th grade of elementary school and groups were equivalent in school grade, age, gender, and IQ. Oral and RC, word recognition, processing speed, picture naming, receptive vocabulary, and phonological awareness were assessed. There were no group differences regarding the accuracy in oral and RC, phonological awareness, naming, and vocabulary scores. DG performed worse than the CG in word recognition (general score and orthographic confusion items) and were slower in naming. Results corroborated the literature regarding word recognition and processing speed deficits in dyslexia. However, dyslexics can achieve normal scores on RC test. Data supports the importance of delimitation of different reading strategies embedded in the word recognition component. The role of processing speed in reading problems remain unclear.
PMCID: PMC4246655  PMID: 25506331
phonological processing; reading disability; cognitive assessment; reading model; processing speed
20.  Visual processing of multiple elements in the dyslexic brain: evidence for a superior parietal dysfunction 
The visual attention (VA) span deficit hypothesis of developmental dyslexia posits that impaired multiple element processing can be responsible for poor reading outcomes. In VA span impaired dyslexic children, poor performance on letter report tasks is associated with reduced parietal activations for multiple letter processing. While this hints towards a non-specific, attention-based dysfunction, it is still unclear whether reduced parietal activity generalizes to other types of stimuli. Furthermore, putative links between reduced parietal activity and reduced ventral occipito-temporal (vOT) in dyslexia have yet to be explored. Using functional magnetic resonance imaging, we measured brain activity in 12 VA span impaired dyslexic adults and 12 adult skilled readers while they carried out a categorization task on single or multiple alphanumeric or non-alphanumeric characters. While healthy readers activated parietal areas more strongly for multiple than single element processing (right-sided for alphanumeric and bilateral for non-alphanumeric), similar stronger multiple element right parietal activations were absent for dyslexic participants. Contrasts between skilled and dyslexic readers revealed significantly reduced right superior parietal lobule (SPL) activity for dyslexic readers regardless of stimuli type. Using a priori anatomically defined regions of interest, we showed that neural activity was reduced for dyslexic participants in both SPL and vOT bilaterally. Finally, we used multiple regressions to test whether SPL activity was related to vOT activity in each group. In the left hemisphere, SPL activity covaried with vOT activity for both normal and dyslexic readers. In contrast, in the right hemisphere, SPL activity covaried with vOT activity only for dyslexic readers. These results bring critical support to the VA interpretation of the VA Span deficit. In addition, they offer a new insight on how deficits in automatic vOT based word recognition could arise in developmental dyslexia.
PMCID: PMC4083222  PMID: 25071509
developmental dyslexia; visual attention; reading; superior parietal lobes
21.  Brain-potential analysis of visual word recognition in dyslexics and typically reading children 
The specialization of visual brain areas for fast processing of printed words plays an important role in the acquisition of reading skills. Dysregulation of these areas may be among the deficits underlying developmental dyslexia. The present study examines the specificity of word activation in dyslexic children in 3rd grade by comparing early components of brain potentials elicited by visually presented words vs. strings of meaningless letter-like symbols. Results showed a more pronounced N1 component for words compared to symbols for both groups. The dyslexic group revealed larger left-lateralized, word-specific N1 responses than the typically reading group. Furthermore, positive correlations between N1 amplitudes and reading fluency were found in the dyslexic group. Our results support the notion of N1 as a sensitive index of visual word processing involved in reading fluency.
PMCID: PMC4075352  PMID: 25071507
reading fluency; developmental dyslexia; visual word recognition; event-related potentials; N1; visual attention
22.  Enhanced activation of the left inferior frontal gyrus in deaf and dyslexic adults during rhyming 
Brain  2009;132(7):1928-1940.
Hearing developmental dyslexics and profoundly deaf individuals both have difficulties processing the internal structure of words (phonological processing) and learning to read. In hearing non-impaired readers, the development of phonological representations depends on audition. In hearing dyslexics, many argue, auditory processes may be impaired. In congenitally profoundly deaf individuals, auditory speech processing is essentially absent. Two separate literatures have previously reported enhanced activation in the left inferior frontal gyrus in both deaf and dyslexic adults when contrasted with hearing non-dyslexics during reading or phonological tasks. Here, we used a rhyme judgement task to compare adults from these two special populations to a hearing non-dyslexic control group. All groups were matched on non-verbal intelligence quotient, reading age and rhyme performance. Picture stimuli were used since this requires participants to generate their own phonological representations, rather than have them partially provided via text. By testing well-matched groups of participants on the same task, we aimed to establish whether previous literatures reporting differences between individuals with and without phonological processing difficulties have identified the same regions of differential activation in these two distinct populations. The data indicate greater activation in the deaf and dyslexic groups than in the hearing non-dyslexic group across a large portion of the left inferior frontal gyrus. This includes the pars triangularis, extending superiorly into the middle frontal gyrus and posteriorly to include the pars opercularis, and the junction with the ventral precentral gyrus. Within the left inferior frontal gyrus, there was variability between the two groups with phonological processing difficulties. The superior posterior tip of the left pars opercularis, extending into the precentral gyrus, was activated to a greater extent by deaf than dyslexic participants, whereas the superior posterior portion of the pars triangularis extending into the ventral pars opercularis, was activated to a greater extent by dyslexic than deaf participants. Whether these regions play differing roles in compensating for poor phonological processing is not clear. However, we argue that our main finding of greater inferior frontal gyrus activation in both groups with phonological processing difficulties in contrast to controls suggests greater reliance on the articulatory component of speech during phonological processing when auditory processes are absent (deaf group) or impaired (dyslexic group). Thus, the brain appears to develop a similar solution to a processing problem that has different antecedents in these two populations.
PMCID: PMC2702837  PMID: 19467990
inferior frontal gyrus; deaf; dyslexia; rhyming; phonology
23.  Brain classification reveals the right cerebellum as the best biomarker of dyslexia 
BMC Neuroscience  2009;10:67.
Developmental dyslexia is a specific cognitive disorder in reading acquisition that has genetic and neurological origins. Despite histological evidence for brain differences in dyslexia, we recently demonstrated that in large cohort of subjects, no differences between control and dyslexic readers can be found at the macroscopic level (MRI voxel), because of large variances in brain local volumes. In the present study, we aimed at finding brain areas that most discriminate dyslexic from control normal readers despite the large variance across subjects. After segmenting brain grey matter, normalizing brain size and shape and modulating the voxels' content, normal readers' brains were used to build a 'typical' brain via bootstrapped confidence intervals. Each dyslexic reader's brain was then classified independently at each voxel as being within or outside the normal range. We used this simple strategy to build a brain map showing regional percentages of differences between groups. The significance of this map was then assessed using a randomization technique.
The right cerebellar declive and the right lentiform nucleus were the two areas that significantly differed the most between groups with 100% of the dyslexic subjects (N = 38) falling outside of the control group (N = 39) 95% confidence interval boundaries. The clinical relevance of this result was assessed by inquiring cognitive brain-based differences among dyslexic brain subgroups in comparison to normal readers' performances. The strongest difference between dyslexic subgroups was observed between subjects with lower cerebellar declive (LCD) grey matter volumes than controls and subjects with higher cerebellar declive (HCD) grey matter volumes than controls. Dyslexic subjects with LCD volumes performed worse than subjects with HCD volumes in phonologically and lexicon related tasks. Furthermore, cerebellar and lentiform grey matter volumes interacted in dyslexic subjects, so that lower and higher lentiform grey matter volumes compared to controls differently modulated the phonological and lexical performances. Best performances (observed in controls) corresponded to an optimal value of grey matter and they dropped for higher or lower volumes.
These results provide evidence for the existence of various subtypes of dyslexia characterized by different brain phenotypes. In addition, behavioural analyses suggest that these brain phenotypes relate to different deficits of automatization of language-based processes such as grapheme/phoneme correspondence and/or rapid access to lexicon entries.
PMCID: PMC2713247  PMID: 19555471
24.  Evidence for Letter-Specific Position Coding Mechanisms 
PLoS ONE  2013;8(7):e68460.
The perceptual matching (same-different judgment) paradigm was used to investigate precision in position coding for strings of letters, digits, and symbols. Reference and target stimuli were 6 characters long and could be identical or differ either by transposing two characters or substituting two characters. The distance separating the two characters was manipulated such that they could either be contiguous, separated by one intervening character, or separated by two intervening characters. Effects of type of character and distance were measured in terms of the difference between the transposition and substitution conditions (transposition cost). Error rates revealed that transposition costs were greater for letters than for digits, which in turn were greater than for symbols. Furthermore, letter stimuli showed a gradual decrease in transposition cost as the distance between the letters increased, whereas the only significant difference for digit and symbol stimuli arose between contiguous and non-contiguous changes, with no effect of distance on the non-contiguous changes. The results are taken as further evidence for letter-specific position coding mechanisms.
PMCID: PMC3699680  PMID: 23844204
25.  Devil in the details? Developmental dyslexia and visual long-term memory for details 
Cognitive theories on causes of developmental dyslexia can be divided into language-specific and general accounts. While the former assume that words are special in that associated processing problems are rooted in language-related cognition (e.g., phonology) deficits, the latter propose that dyslexia is rather rooted in a general impairment of cognitive (e.g., visual and/or auditory) processing streams. In the present study, we examined to what extent dyslexia (typically characterized by poor orthographic representations) may be associated with a general deficit in visual long-term memory (LTM) for details. We compared object- and detail-related visual LTM performance (and phonological skills) between dyslexic primary school children and IQ-, age-, and gender-matched controls. The results revealed that while the overall amount of LTM errors was comparable between groups, dyslexic children exhibited a greater portion of detail-related errors. The results suggest that not only phonological, but also general visual resolution deficits in LTM may play an important role in developmental dyslexia.
PMCID: PMC4078255  PMID: 25071656
picture processing; memory errors; orthographic representations; visual resolution deficit; phonology and reading; language and word processing

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