Search

This study utilized a neuroimaging task to assess working memory (WM) network recruitment during single word reading. Associations between WM and reading comprehension skills are well documented. Several converging models suggest WM may also contribute to foundational reading skills, but few studies have assessed this contribution directly. Two groups of children (77 developmental dyslexia (DD), 22 controls) completed a functional magnetic resonance imaging (fMRI) task to identify activation of a priori defined regions of the WM network. fMRI trials consisted of familiar word, pseudoword, and false font stimuli within a 1-back oddball task to assess how activation in the WM network differs in response to stimuli that can respectively be processed using word recognition, phonological decoding, or non-word strategies. Results showed children with DD recruited WM regions bilaterally in response to all stimulus types, whereas control children recruited left-lateralized WM regions during the pseudoword condition only. Group-level comparisons revealed activation differences in the defined WM network regions for false font and familiar word, but not pseudoword conditions. This effect was driven by increased activity in participants with DD in right hemisphere frontal, parietal, and motor regions despite poorer task performance. Findings suggest the WM network may contribute to inefficient decoding and word recognition strategies in children with DD., Competing Interests: Declaration of competing interest none., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Decades of research have led to several competing theories regarding the neural contributors to impaired reading. But how can we know which theory (or theories) identifies the types of markers that indeed differentiate between individuals with reading disabilities (RD) and their typically developing (TD) peers? To answer this question, we propose a new analytical tool for theory evaluation and comparison, grounded in the Bayesian latent-mixture modeling framework. We start by constructing a series of latent-mixture classification models, each reflecting one existing theoretical claim regarding the neurofunctional markers of RD (highlighting network-level differences in either mean activation, inter-subject heterogeneity, inter-region variability, or connectivity). Then, we run each model on fMRI data alone (i.e., while models are blind to participants' behavioral status), which enables us to interpret the fit between a model's classification of participants and their behavioral (known) RD/TD status as an estimate of its explanatory power. Results from n=127 adolescents and young adults (RD: n=59; TD: n=68) show that models based on network-level differences in mean activation and heterogeneity failed to differentiate between TD and RD individuals. In contrast, classifications based on variability and connectivity were significantly associated with participants' behavioral status. These findings suggest that differences in inter-region variability and connectivity may be better network-level markers of RD than mean activation or heterogeneity (at least in some populations and tasks). More broadly, the results demonstrate the promise of latent-mixture modeling as a theory-driven tool for evaluating different theoretical claims regarding neural contributors to language disorders and other cognitive traits., Competing Interests: Declaration of Competing Interest None., (Copyright © 2021. Published by Elsevier Inc.)

Understanding how pre-literate children's language abilities and neural function relate to future reading ability is important for identifying children who may be at-risk for reading problems. Pre-literate children are already proficient users of spoken language and their developing brain networks for language become highly overlapping with brain networks that emerge during literacy acquisition. In the present longitudinal study, we examined language abilities, and neural activation and connectivity within the language network in pre-literate children (mean age = 4.2 years). We tested how language abilities, brain activation, and connectivity predict children's reading abilities 1 year later (mean age = 5.2 years). At Time 1, children (n = 37) participated in a functional near infrared spectroscopy (fNIRS) experiment of speech processing (listening to words and pseudowords) and completed a standardized battery of language and cognitive assessments. At Time 2, children (n = 28) completed standardized reading assessments. Using psychophysiological interaction (PPI) analyses, we observed significant connectivity between the left IFG and right STG in pre-literate children, which was modulated by task (i.e., listening to words). Neural activation in left IFG and STG and increased task-modulated connectivity between the left IFG and right STG was predictive of multiple reading outcomes. Increased connectivity was associated later with increased reading ability., (© 2020 John Wiley & Sons Ltd.)

The BDNF gene is a prominent promoter of neuronal development, maturation and plasticity. Its Val 66 Met polymorphism affects brain morphology and function within several areas and is associated with several cognitive functions and neurodevelopmental disorder susceptibility. Recently, it has been associated with reading, reading-related traits and altered neural activation in reading-related brain regions. However, it remains unknown if the intermediate phenotypes (IPs, such as brain activation and phonological skills) mediate the pathway from gene to reading or reading disability. By conducting a serial multiple mediation model in a sample of 94 children (age 5-13), our findings revealed no direct effects of genotype on reading. Instead, we found that genotype is associated with brain activation in reading-related and more domain general regions which in turn is associated with phonological processing which is associated with reading. These findings suggest that the BDNF-Val 66 Met polymorphism is related to reading via phonological processing and functional activation. These results support brain imaging data and neurocognitive traits as viable IPs for complex behaviors., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Neuroimaging studies of basic achievement skills - reading and arithmetic - often control for the effect of IQ to identify unique neural correlates of each skill. This may underestimate possible effects of common factors between achievement and IQ measures on neuroimaging results. Here, we simultaneously examined achievement (reading and arithmetic) and IQ measures in young adults, aiming to identify MRI correlates of their common factors. Resting-state fMRI (rs-fMRI) data were analyzed using two metrics assessing local intrinsic functional properties; regional homogeneity (ReHo) and fractional amplitude low frequency fluctuation (fALFF), measuring local intrinsic functional connectivity and intrinsic functional activity, respectively. ReHo highlighted the thalamus/pulvinar (a subcortical region implied for selective attention) as a common locus for both achievement skills and IQ. More specifically, the higher the ReHo values, the lower the achievement and IQ scores. For fALFF, the left superior parietal lobule, part of the dorsal attention network, was positively associated with reading and IQ. Collectively, our results highlight attention-related regions, particularly the thalamus/pulvinar as a key region related to individual differences in performance on all the three measures. ReHo in the thalamus/pulvinar may serve as a tool to examine brain mechanisms underlying a comorbidity of reading and arithmetic difficulties, which could co-occur with weakness in general intellectual abilities., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020. Published by Elsevier Inc.)

Research using functional and structural magnetic resonance imaging has identified areas of reduced brain activation and gray matter volume in children and adults with reading disability, but associations between cortical structure and individual differences in reading in typically developing children remain underexplored. Furthermore, the majority of research linking gray matter structure to reading ability quantifies gray matter in terms of volume, and cannot specify unique contributions of cortical surface area and thickness to these relationships. Here, we applied a continuous analytic approach to investigate associations between distinct surface-based properties of cortical structure and individual differences in reading-related skills in a sample of typically developing young children. Correlations between cortical structure and reading-related skills were conducted using a surface-based vertex-wise approach. Cortical thickness in the left superior temporal cortex was positively correlated with word and pseudoword reading performance. The observed positive correlation between cortical thickness in the left superior temporal cortex and reading may have implications for the patterns of brain activation that support reading., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Objective: Decreased activation to print in the left ventral, dorsal, and anterior pathways has been implicated in readers with dyslexia (DRs) but also is characteristic for typical beginning readers. Because most studies have compared DRs with their age-matched peers, the observed results could represent a dyslexia phenotype or a developmental delay. This study aimed to disentangle reading and dyslexia effects using 2 control groups matched for age and skill and a longitudinal design., Method: Brain response for print was compared in DRs and typical readers (TRs) who, at the beginning of schooling (time point 1 [TP]; 6-7 years old), read on average 3 words per minute, as did DRs at TP1, but improved their reading to an average level, and advanced readers (ARs) who at TP1 read as well as DRs 2 years later (TP3; 8-9 years old). The TR and DR groups were tracked longitudinally to observe neurodevelopmental changes., Results: At TP1, DRs did not differ from TRs. Over time, only TRs developed a neural circuit for reading in the left inferior frontal and fusiform gyri. At TP3, DRs exhibited hypo-activation in these areas compared with age-matched (TRs at TP3) and reading-matched (ARs at TP1) controls. At TP3, TRs showed hypo-activation in the left frontal and bilateral ventral occipital regions compared with ARs, but these effects were nonoverlapping with DR hypo-activations and are partly explained by IQ., Conclusion: Decreased activation of the left fusiform and inferior frontal gyri to print in DRs results from an atypical developmental trajectory of reading and cannot be explained solely by lower reading skills., (Copyright © 2018 American Academy of Child and Adolescent Psychiatry. Published by Elsevier Inc. All rights reserved.)

Reading is a learned activity that engages multiple cognitive systems. In a cohort of typical and struggling adult readers we show evidence that successful oral reading of real words is related to gamma-amino-butyric acid (GABA) concentration in the higher-order language system, whereas reading of unfamiliar pseudo-words is not related to GABA in this system. We also demonstrate the capability of resting state functional connectivity (rsFC) combined with GABA measures to predict single real word compared to pseudo-word reading performance. Results show that the strength of rsFC between left fusiform gyrus (L-FG) and higher-order language systems predicts oral reading behavior of real words, irrespective of the local concentration of GABA. On the other hand, pseudo-words, which require grapheme-to-phoneme conversion, are not predicted by the connection between L-FG and higher-order language system. This suggests that L-FG may have a multi-functional role: lexical processing of real words and grapheme-to-phoneme processing of pseudo-words. Additionally, rsFC between L-FG, pre-motor, and putamen areas are positively related to the oral reading of both real and pseudo-words, suggesting that text may be converted into a phoneme sequence for speech initiation and production regardless of whether the stimulus is a real word or pseudo-word. In summary, from a systems neuroscience perspective, we show that: (i) strong rsFC between higher order visual, language, and pre-motor areas can predict and differentiate efficient oral reading of real and pseudo-words. (ii) GABA measures, along with rsFC, help to further differentiate the neural pathways for previously learned real words versus unfamiliar pseudo-words.

The aim of the current study was to develop an fMRI task capable of characterizing individual differences in reading and attentional domains. Forty-nine students with a range of reading and attentional control abilities completed an event-related fMRI oddball task consisting of printed word and false font stimuli. Reading network activation was assessed by contrasting printed words with false font stimuli. Left inferior frontal gyrus and superior/middle temporal gyrus showed a main effect of stimulus type. The magnitude of the difference in activation between words and false font was correlated with word reading for both regions and reading fluency for superior/middle temporal gyrus. Regions including bilateral middle cingulate, insula and right inferior frontal gyrus showed a main effect of trial type. The difference in activation between oddball and standard trials in the right superior/middle temporal gyrus and left cerebellum was correlated with attentional control measures. Results indicate the task tapped both reading and attentional control resources. Understanding the contribution of the neural networks supporting each of these domains may provide insight into the shared neural deficits underlying the co-morbidity between developmental dyslexia and attention deficit hyperactivity disorder., (Copyright © 2019 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Epidemiological population studies highlight the presence of substantial individual variability in reading skill, with approximately 5-10% of individuals characterized as having specific reading disability (SRD). Despite reported substantial heritability, typical for a complex trait, the specifics of the connections between reading and the genome are not understood. Recently, the SETBP1 gene has been implicated in several complex neurodevelopmental syndromes and disorders that impact language. Here, we examined the relationship between common polymorphisms in this gene, reading, and reading associated behaviors using data from an ongoing project on the genetic basis of SRD (n = 135). In addition, an exploratory analysis was conducted to examine the relationship between SETBP1 and brain activation using functional magnetic resonance imaging (fMRI; n = 73). Gene-based analyses revealed a significant association between SETBP1 and phonological working memory, with rs7230525 as the strongest associated single nucleotide polymorphism (SNP). fMRI analysis revealed that the rs7230525-T allele is associated with functional neural activation during reading and listening to words and pseudowords in the right inferior parietal lobule (IPL). These findings suggest that common genetic variation within SETBP1 is associated with reading behavior and reading-related brain activation patterns in the general population., (Copyright © 2018. Published by Elsevier Ltd.)

Recent work has suggested that variability in levels of neural activation may be related to behavioral and cognitive performance across a number of domains and may offer information that is not captured by more traditional measures that use the average level of brain activation. We examined the relationship between reading skill in school-aged children and neural activation variability during a functional MRI reading task after taking into account average levels of activity. The reading task involved matching printed and spoken words to pictures of items. Single trial activation estimates were used to calculate the mean and standard deviation of children's responses to print and speech stimuli; multiple regression analyses evaluated the relationship between reading skill and trial-by-trial activation variability. The reliability of observed findings from the discovery sample ( n = 44; ages 8-11; 18 female) was then confirmed in an independent sample of children ( n = 32; ages 8-11; 14 female). Across the two samples, reading skill was positively related to trial-by-trial variability in the activation response to print in the left inferior frontal gyrus pars triangularis. This relationship held even when accounting for mean levels of activation. This finding suggests that intrasubject variability in trial-by-trial fMRI activation responses to printed words accounts for individual differences in human reading ability that are not fully captured by traditional mean levels of brain activity. Furthermore, this positive relationship between trial-by-trial activation variability and reading skill may provide evidence that neural variability plays a beneficial role during early reading development. SIGNIFICANCE STATEMENT Recent work has suggested that neural activation variability, or moment-to-moment changes in the engagement of brain regions, is related to individual differences in behavioral and cognitive performance across multiple domains. However, differences in neural activation variability have not yet been evaluated in relation to reading skill. In the current study, we analyzed data from two independent groups of children who performed an fMRI task involving reading and listening to words. Across both samples, reading skill was positively related to trial-by-trial variability in activation to print stimuli in the left inferior frontal gyrus pars triangularis, even when accounting for the more conventional measure of mean levels of brain activity. This finding suggests that neural variability could be beneficial in developing readers., (Copyright © 2018 the authors 0270-6474/18/382982-09$15.00/0.)

Word learning depends not only on efficient online binding of phonological, orthographic and lexical information, but also on consolidation of new word representations into permanent lexical memory. Work on word learning under a variety of contexts indicates that reading and language skill impact facility of word learning in both print and speech. In addition, recent research finds that individuals with language impairments show deficits in both initial word form learning and in maintaining newly learned representations over time, implicating mechanisms associated with maintenance that may be driven by deficits in overnight consolidation. Although several recent studies have explored the neural bases of overnight consolidation of newly learned words, no extant work has examined individual differences in overnight consolidation at the neural level. The current study addresses this gap in the literature by investigating how individual differences in reading and language skills modulate patterns of neural activation associated with newly learned words following a period of overnight consolidation. Specifically, a community sample of adolescents and young adults with significant variability in reading and oral language (vocabulary) ability were trained on two spoken artificial lexicons, one in the evening on the day before fMRI scanning and one in the morning just prior to scanning. Comparisons of activation between words that were trained and consolidated vs. those that were trained but not consolidated revealed increased cortical activation in a number of language associated and memory associated regions. In addition, individual differences in age, reading skill and vocabulary modulated learning rate in our artificial lexicon learning task and the size of the cortical consolidation effect in the precuneus/posterior cingulate, such that older readers and more skilled readers had larger cortical consolidation effects in this learning-critical region. These findings suggest that age (even into late adolescence) and reading and language skills are important individual differences that affect overnight consolidation of newly learned words. These findings have significant implications for understanding reading and language disorders and should inform pedagogical models., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Background: Literacy acquisition is a demanding process that induces significant changes in the brain, especially in the spoken and written language networks. Nevertheless, large-scale paediatric fMRI studies are still limited., Methods: We analyzed fMRI data to show how individual differences in reading performance correlate with brain activation for speech and print in 111 children attending kindergarten or first grade and examined group differences between a matched subset of emergent-readers and prereaders., Results: Across the entire cohort, individual differences analysis revealed that reading skill was positively correlated with the magnitude of activation difference between words and symbol strings in left superior temporal, inferior frontal and fusiform gyri. Group comparisons of the matched subset of pre- and emergent-readers showed higher activity for emergent-readers in left inferior frontal, precentral, and postcentral gyri. Individual differences in activation for natural versus vocoded speech were also positively correlated with reading skill, primarily in the left temporal cortex. However, in contrast to studies on adult illiterates, group comparisons revealed higher activity in prereaders compared to readers in the frontal lobes. Print-speech coactivation was observed only in readers and individual differences analyses revealed a positive correlation between convergence and reading skill in the left superior temporal sulcus., Conclusions: These results emphasise that a child's brain undergoes several modifications to both visual and oral language systems in the process of learning to read. They also suggest that print-speech convergence is a hallmark of acquiring literacy., (© 2017 Association for Child and Adolescent Mental Health.)

Developmental dyslexia (decoding-based reading disorder; RD) is a complex trait with multifactorial origins at the genetic, neural, and cognitive levels. There is evidence that low-level sensory-processing deficits precede and underlie phonological problems, which are one of the best-documented aspects of RD. RD is also associated with impairments in integrating visual symbols with their corresponding speech sounds. Although causal relationships between sensory processing, print-speech integration, and fluent reading, and their neural bases are debated, these processes all require precise timing mechanisms across distributed brain networks. Neural excitability and neural noise are fundamental to these timing mechanisms. Here, we propose that neural noise stemming from increased neural excitability in cortical networks implicated in reading is one key distal contributor to RD., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

In the current study, we present a novel fMRI protocol in which words, pseudowords, and other word-like stimuli are passively presented in a rapid, sequential fashion. In this "fast" localizer paradigm, items are presented in groups of four; within sets, words are related in orthographic, phonological, and/or semantic properties. We tested this protocol with a group of skilled adult readers (N=18). Analyses uncovered key regions of the reading network that were sensitive to different component processes at the group level; namely, left fusiform gyrus as well as the pars opercularis subregion of inferior frontal gyrus were sensitive to lexicality; several regions including left precentral gyrus and left supramarginal gyrus were sensitive to spelling-sound consistency; the pars triangularis subregion of inferior frontal gyrus was sensitive to semantic similarity. Additionally, in a number of key brain regions, activation in response to semantically similar words was related to individual differences in reading comprehension outside the scanner. Importantly, these findings are in line with previous investigations of the reading network, yet data were obtained using much less imaging time than comparable paradigms currently available, especially relative to the number of indices of component processes obtained. This feature, combined with the relatively simple nature of the task, renders it appropriate for groups of subjects with a wide range of reading abilities, including children with impairments., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Understanding how genes impact the brain's functional activation for learning and cognition during development remains limited. We asked whether a common genetic variant in the BDNF gene (the Val66Met polymorphism) modulates neural activation in the young brain during a critical period for the emergence and maturation of the neural circuitry for reading. In animal models, the bdnf variation has been shown to be associated with the structure and function of the developing brain and in humans it has been associated with multiple aspects of cognition, particularly memory, which are relevant for the development of skilled reading. Yet, little is known about the impact of the Val66Met polymorphism on functional brain activation in development, either in animal models or in humans. Here, we examined whether the BDNF Val66Met polymorphism (dbSNP rs6265) is associated with children's (age 6-10) neural activation patterns during a reading task (n = 81) using functional magnetic resonance imaging (fMRI), genotyping, and standardized behavioral assessments of cognitive and reading development. Children homozygous for the Val allele at the SNP rs6265 of the BDNF gene outperformed Met allele carriers on reading comprehension and phonological memory, tasks that have a strong memory component. Consistent with these behavioral findings, Met allele carriers showed greater activation in reading-related brain regions including the fusiform gyrus, the left inferior frontal gyrus and left superior temporal gyrus as well as greater activation in the hippocampus during a word and pseudoword reading task. Increased engagement of memory and spoken language regions for Met allele carriers relative to Val/Val homozygotes during reading suggests that Met carriers have to exert greater effort required to retrieve phonological codes., Competing Interests: The authors have declared that no competing interests exist.

Phonological processing ability is a key factor in reading acquisition, predicting its later success or causing reading problems when it is weakened. Our aim here was to establish the neural correlates of auditory word rhyming (a standard phonological measure) in 102 young children with (FHD+) and without familial history of dyslexia (FHD-) in a shallow orthography (i.e. Polish). Secondly, in order to gain a deeper understanding on how schooling shapes brain activity to phonological awareness, a comparison was made of children who had had formal literacy instruction for several months (in first grade) and those who had not yet had any formal instruction in literacy (in kindergarten). FHD+ children compared to FHD- children in the first grade scored lower in an early print task and showed longer reaction times in the in-scanner rhyme task. No behavioral differences between FHD+ and FHD- were found in the kindergarten group. On the neuronal level, overall familial risk was associated with reduced activation in the bilateral temporal, tempo-parietal and inferior temporal-occipital regions, as well as the bilateral inferior and middle frontal gyri. Subcortically, hypoactivation was found in the bilateral thalami, caudate, and right putamen in FHD+. A main effect of the children's grade was present only in the left inferior frontal gyrus, where reduced activation for rhyming was shown in first-graders. Several regions in the ventral occipital cortex, including the fusiform gyrus, and in the right middle frontal and postcentral gyri, displayed an interaction between familial risk and grade. The present results show strong influence of familial risk that may actually increase with formal literacy instruction., (Copyright © 2016 Elsevier Inc. All rights reserved.)

We propose and test a theoretical perspective in which a universal hallmark of successful literacy acquisition is the convergence of the speech and orthographic processing systems onto a common network of neural structures, regardless of how spoken words are represented orthographically in a writing system. During functional MRI, skilled adult readers of four distinct and highly contrasting languages, Spanish, English, Hebrew, and Chinese, performed an identical semantic categorization task to spoken and written words. Results from three complementary analytic approaches demonstrate limited language variation, with speech-print convergence emerging as a common brain signature of reading proficiency across the wide spectrum of selected languages, whether their writing system is alphabetic or logographic, whether it is opaque or transparent, and regardless of the phonological and morphological structure it represents.

Despite anecdotal evidence of relative visuospatial processing strengths in individuals with reading disability (RD), only a few studies have assessed the presence or the extent of these putative strengths. The current study examined the cognitive and neural bases of visuospatial processing abilities in adolescents with RD relative to typically developing (TD) peers. Using both cognitive tasks and functional magnetic resonance imaging (fMRI) we contrasted printed word recognition with non-language visuospatial processing tasks. Behaviorally, lower reading skill was related to a visuospatial processing advantage (shorter latencies and equivalent accuracy) on a geometric figure processing task, similar to findings shown in two published studies. FMRI analyses revealed key group by task interactions in patterns of cortical and subcortical activation, particularly in frontostriatal networks, and in the distributions of right and left hemisphere activation on the two tasks. The results are discussed in terms of a possible neural tradeoff in visuospatial processing in RD., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Word reading in alphabetic language involves a cortical system with multiple components whose division of labor depends on the transparency of the writing system. To gain insight about the neural division of labor between phonology and semantics subserving word reading in Chinese, a deep non-alphabetic writing system, functional magnetic resonance imaging (fMRI) was used to investigate the effects of phonological and semantic training on the cortical circuitry for oral naming of Chinese characters. In a training study, we examined whether a training task that differentially focused readers' attention on the phonological or semantic properties of a Chinese character changes the patterns of cortical activation that was evoked by that character in a subsequent naming task. Our imaging results corroborate that the cortical regions underlying reading in Chinese largely overlap the left-hemisphere reading system responsible for reading in alphabetic languages, with some cortical regions in the left-hemisphere uniquely recruited for reading in Chinese. However, in contrast to findings from studies of English word naming, we observed considerable overlap in the neural activation patterns associated with phonological and semantic training on naming Chinese characters, which we suggest may reflect a balanced neural division of labor between phonology and semantics in Chinese character reading. The equitable division of labor for Chinese reading might be driven by the special statistical structure of the writing system, which includes equally systematic mappings in the correspondences between written forms and their pronunciations and meanings., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Reading disability is a brain-based difficulty in acquiring fluent reading skills that affects significant numbers of children. Although neuroanatomical and neurofunctional networks involved in typical and atypical reading are increasingly well characterized, the underlying neurochemical bases of individual differences in reading development are virtually unknown. The current study is the first to examine neurochemistry in children during the critical period in which the neurocircuits that support skilled reading are still developing. In a longitudinal pediatric sample of emergent readers whose reading indicators range on a continuum from impaired to superior, we examined the relationship between individual differences in reading and reading-related skills and concentrations of neurometabolites measured using magnetic resonance spectroscopy. Both continuous and group analyses revealed that choline and glutamate concentrations were negatively correlated with reading and related linguistic measures in phonology and vocabulary (such that higher concentrations were associated with poorer performance). Correlations with behavioral scores obtained 24 months later reveal stability for the relationship between glutamate and reading performance. Implications for neurodevelopmental models of reading and reading disability are discussed, including possible links of choline and glutamate to white matter anomalies and hyperexcitability. These findings point to new directions for research on gene-brain-behavior pathways in human studies of reading disability.

We employed brain-behavior analyses to explore the relationship between performance on tasks measuring phonological awareness, pseudoword decoding, and rapid auditory processing (all predictors of reading (dis)ability) and brain organization for print and speech in beginning readers. For print-related activation, we observed a shared set of skill-correlated regions, including left hemisphere temporoparietal and occipitotemporal sites, as well as inferior frontal, visual, visual attention, and subcortical components. For speech-related activation, shared variance among reading skill measures was most prominently correlated with activation in left hemisphere inferior frontal gyrus and precuneus. Implications for brain-based models of literacy acquisition are discussed., (Copyright © 2012 Elsevier Inc. All rights reserved.)

In both children and adults there is large variability in reading skill, with approximately 5-10% of individuals characterized as having reading disability; these individuals struggle to learn to read despite adequate intelligence and opportunity. Although it is well established that a substantial portion of this variability is attributed to the genetic differences between individuals, specifics of the connections between reading and the genome are not understood. This article presents data that suggest that variation in the COMT gene, which has previously been associated with variation in higher-order cognition, is associated with reading and reading-related skills, at the level of both brain and behavior. In particular, we found that the COMT Val/Met polymorphism at rs4680, which results in the substitution of the ancestral Valine (Val) by Methionine (Met), was associated with better performance on a number of critical reading measures and with patterns of functional neural activation that have been linked to better readers. We argue that this polymorphism, known for its broad effects on cognition, may modulate (likely through frontal lobe function) reading skill., (© 2012 Blackwell Publishing Ltd.)

Purpose: To examine neural response to spoken and printed language in children with speech sound errors (SSE)., Method: Functional magnetic resonance imaging was used to compare processing of auditorily and visually presented words and pseudowords in 17 children with SSE, ages 8;6[years;months] through 10;10, with 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 on 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.

Skilled reading depends upon successfully integrating orthographic, phonological, and semantic information; however, the process of becoming a skilled reader with efficient neural circuitry is not fully understood. Short-term learning paradigms can provide insight into learning mechanisms by revealing differential responses to training approaches. To date, neuroimaging studies have primarily focused on effects of teaching novel words either in isolation or in context, without directly comparing the two. The current study compared the behavioral and neurobiological effects of learning novel pseudowords (i.e., pronouncing and attaching meaning) trained either in isolation or in sentential context. Behavioral results showed generally comparable pseudoword learning for both conditions, but sentential context-trained pseudowords were spoken and comprehended slightly more quickly. Neurobiologically, fMRI activity for reading trained pseudowords was similar to real words; however, an interaction between training approach and reading proficiency was observed. Specifically, highly skilled readers showed similar levels of activity regardless of training approach. However, less skilled readers differentiated between training conditions, showing comparable activity to highly skilled readers only for isolation-trained pseudowords. Overall, behavioral and neurobiological findings suggest that training approach may affect rate of learning and neural circuitry, and that less skilled readers may need explicit training to develop optimal neural pathways., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Early language development sets the stage for a lifetime of competence in language and literacy. However, the neural mechanisms associated with the relative advantages of early communication success, or the disadvantages of having delayed language development, are not well explored. In this study, 174 elementary school-age children whose parents reported that they started forming sentences 'early', 'on-time' or 'late' were evaluated with standardized measures of language, reading and spelling. All oral and written language measures revealed consistent patterns for 'early' talkers to have the highest level of performance and 'late' talkers to have the lowest level of performance. We report functional magnetic resonance imaging data from a subset of early, on-time and late talkers matched for age, gender and performance intelligence quotient that allows evaluation of neural activation patterns produced while listening to and reading real words and pronounceable non-words. Activation in bilateral thalamus and putamen, and left insula and superior temporal gyrus during these tasks was significantly lower in late talkers, demonstrating that residual effects of being a late talker are found not only in behavioural tests of oral and written language, but also in distributed cortical-subcortical neural circuits underlying speech and print processing. Moreover, these findings suggest that the age of functional language acquisition can have long-reaching effects on reading and language behaviour, and on the corresponding neurocircuitry that supports linguistic function into the school-age years.

Using fMRI, we explored the relationship between phonological awareness (PA), a measure of metaphonological knowledge of the segmental structure of speech, and brain activation patterns during processing of print and speech in young readers from 6 to 10 years of age. Behavioral measures of PA were positively correlated with activation levels for print relative to speech tokens in superior temporal and occipito-temporal regions. Differences between print-elicited activation levels in superior temporal and inferior frontal sites were also correlated with PA measures with the direction of the correlation depending on stimulus type: positive for pronounceable pseudowords and negative for consonant strings. These results support and extend the many indications in the behavioral and neurocognitive literature that PA is a major component of skill in beginning readers and point to a developmental trajectory by which written language engages areas originally shaped by speech for learners on the path toward successful literacy acquisition.

Functional magnetic resonance imaging (fMRI) was used to investigate the impact of literacy skills in young adults on the distribution of cerebral activity during comprehension of sentences in spoken and printed form. The aim was to discover where speech and print streams merge, and whether their convergence is affected by the level of reading skill. The results from different analyses all point to the conclusion that neural integration of sentence processing across speech and print varies positively with the reader's skill. Further, they identify the inferior frontal region as the principal site of speech-print integration and a major focus of reading comprehension differences. The findings provide new evidence of the role of the inferior frontal region in supporting supramodal systems of linguistic representation.

Using functional magnetic resonance imaging, we explored the role of semantics in mediating orthographic-to-phonological processing in reading aloud, focusing on the interaction of imageability with spelling-to-sound consistency for low-frequency words. Behaviorally, high-imageable words attenuate the standard latency and accuracy disadvantage for low-frequency inconsistent words relative to their consistent counterparts. Neurobiologically, high-imageable words reduced consistency-related activation in the inferior frontal gyrus but increased posterior activation in the angular and middle temporal gyri, representing a possible neural signature of the tradeoff between semantics and phonology in reading aloud. We discuss implications for neurobiological models of reading in terms of understanding the interplay among areas associated with component processes and suggest that the results constitute an important step toward integrating neurobiological and computational models of reading.

Cortical regions engaged by sentence processing were mapped using functional MRI. The influence of input modality (spoken word vs. print input) and parsing difficulty (sentences containing subject-relative vs. object-relative clauses) was assessed. Auditory presentation was associated with pronounced activity at primary auditory cortex and across the superior temporal gyrus bilaterally. Printed sentences by contrast evoked major activity at several posterior sites in the left hemisphere, including the angular gyrus, supramarginal gyrus, and the fusiform gyrus in the occipitotemporal region. In addition, modality-independent regions were isolated, with greatest overlap seen in the inferior frontal gyrus (IFG). With respect to sentence complexity, object-relative sentences evoked heightened responses in comparison to subject-relative sentences at several left hemisphere sites, including IFG, the middle/superior temporal gyrus, and the angular gyrus. These sites showing modulation of activity as a function of sentence type, independent of input mode, arguably form the core of a cortical system essential to sentence parsing.

fMRI was used to investigate the separate influences of orthographic, phonological, and semantic processing on the ability to learn new words and the cortical circuitry recruited to subsequently read those words. In a behavioral session, subjects acquired familiarity for three sets of pseudowords, attending to orthographic, phonological, or (learned) semantic features. Transfer effects were measured in an event-related fMRI session as the subjects named trained pseudowords, untrained pseudowords, and real words. Behaviorally, phonological and semantic training resulted in better learning than did orthographic training. Neurobiologically, orthographic training did not modulate activation in the main reading regions. Phonological and semantic training yielded equivalent behavioral facilitation but distinct functional activation patterns, suggesting that the learning resulting from these two training conditions was driven by different underlying processes. The findings indicate that the putative ventral visual word form area is sensitive to the phonological structure of words, with phonologically analytic processing contributing to the specialization of this region.

Background: Converging evidence indicates a functional disruption in the neural systems for reading in adults with dyslexia. We examined brain activation patterns in dyslexic and nonimpaired children during pseudoword and real-word reading tasks that required phonologic analysis (i.e., tapped the problems experienced by dyslexic children in sounding out words)., Methods: We used functional magnetic resonance imaging (fMRI) to study 144 right-handed children, 70 dyslexic readers, and 74 nonimpaired readers as they read pseudowords and real words., Results: Children with dyslexia demonstrated a disruption in neural systems for reading involving posterior brain regions, including parietotemporal sites and sites in the occipitotemporal area. Reading skill was positively correlated with the magnitude of activation in the left occipitotemporal region. Activation in the left and right inferior frontal gyri was greater in older compared with younger dyslexic children., Conclusions: These findings provide neurobiological evidence of an underlying disruption in the neural systems for reading in children with dyslexia and indicate that it is evident at a young age. The locus of the disruption places childhood dyslexia within the same neurobiological framework as dyslexia, and acquired alexia, occurring in adults.

Becoming a skilled reader requires building a functional neurocircuitry for printed-language processing that integrates with spoken-language-processing networks. In this longitudinal study, functional MRI (fMRI) was used to examine convergent activation for printed and spoken language (print-speech coactivation) in selected regions implicated in printed-language processing (the reading network). We found that print-speech coactivation across the left-hemisphere reading network in beginning readers predicted reading achievement 2 years later beyond the effects of brain activity for either modality alone; moreover, coactivation effects accounted for variance in later reading after controlling for initial reading performance. Within the reading network, effects of coactivation were significant in bilateral inferior frontal gyrus (IFG) and left inferior parietal cortex and fusiform gyrus. The contribution of left and right IFG differed, with more coactivation in left IFG predicting better achievement but more coactivation in right IFG predicting poorer achievement. Findings point to the centrality of print-speech convergence in building an efficient reading circuitry in children.

Despite anecdotal evidence of relative visuospatial processing strengths in individuals with reading disability (RD), only a few studies have assessed the presence or the extent of these putative strengths. The current study examined the cognitive and neural bases of visuospatial processing abilities in adolescents with RD relative to typically developing (TD) peers. Using both cognitive tasks and functional magnetic resonance imaging (fMRI) we contrasted printed word recognition with non-language visuospatial processing tasks. Behaviorally, lower reading skill was related to a visuospatial processing advantage (shorter latencies and equivalent accuracy) on a geometric figure processing task, similar to findings shown in two published studies. FMRI analyses revealed key group by task interactions in patterns of cortical and subcortical activation, particularly in frontostriatal networks, and in the distributions of right and left hemisphere activation on the two tasks. The results are discussed in terms of a possible neural tradeoff in visuospatial processing in RD.

Statistical learning (SL) approaches to reading maintain that proficient reading requires assimilation of rich statistical regularities in the writing system. Reading skills in developing first-language readers are predicted by individual differences in sensitivity to regularities in mappings from orthography to phonology (O-P) and semantics (O-S), where good readers rely more on O-P consistency, and less on O-S associations. However, how these regularities are leveraged by second-language (L2) learners remains an open question. We utilize an individual-differences approach, measuring L2 English learners' sensitivity to O-P, O-S, and frequency during word-naming, across two years of immersion. We show that reliance on O-P is leveraged by better readers, while reliance on O-S is slower to develop, characterizing less proficient readers. All factors explain substantial individual variance in L2 reading skills. These findings show how SL plays a key role in L2 reading development through its role in assimilating sublexical regularities between print and speech. [ABSTRACT FROM AUTHOR]

In a quasiregular orthography like English, children inevitably encounter irregular words during reading. Previous research suggests successful reading of an irregular word depends at least partially on a child's ability to address the mismatch between decoded form and stored word pronunciation, referred to as a child's set for variability, and the word's relative transparency, measured here using a spelling to pronunciation transparency rating. Item-level analyses were used to explore the relationship between general child performance on the set for variability mispronunciation task, word specific set for variability (predicting reading of that word), spelling to pronunciation transparency rating, and irregular word reading. Significant predictors included general word reading, general set for variability performance, and item-specific set for variability performance; word frequency and spelling to pronunciation transparency rating; and an interaction between word reading and the transparency rating. Results underscore the importance of considering both general and item-specific factors affecting irregular word reading. [ABSTRACT FROM AUTHOR]

To investigate the neural basis of a common statistical learning mechanism involved in motor sequence learning and decoding, we recorded brain activation from participants during a serial reaction time (SRT) task and a word reading task using functional magnetic resonance imaging. In the SRT task, a manual response was made depending on the location of a visual cue, and the order of the locations was either fixed or random. In the word reading task, visual words were passively presented. In the inferior frontal gyrus pars triangularis (IFGpTr) and the insula, differences in activation between the ordered and random condition in the SRT task and activation to printed words in the word reading task were correlated with the participants' decoding ability. We speculate that extraction of statistically predictable patterns in the IFGpTr and insula contributes to both motor sequence learning and orthographic learning, and therefore predicts individual differences in decoding skill. [ABSTRACT FROM AUTHOR]

Recent work has suggested that variability in levels of neural activation may be related to behavioral and cognitive performance across a number of domains and may offer information that is not captured by more traditional measures that use the average level of brain activation. We examined the relationship between reading skill in school-aged children and neural activation variability during a functional MRI reading task after taking into account average levels of activity. The reading task involved matching printed and spoken words to pictures of items. Single trial activation estimates were used to calculate the mean and standard deviation of children's responses to print and speech stimuli; multiple regression analyses evaluated the relationship between reading skill and trial-by-trial activation variability. The reliability of observed findings from the discovery sample (n = 44; ages 8-11; 18 female) was then confirmed in an independent sample of children (n = 32; ages 8-11; 14 female). Across the two samples, reading skill was positively related to trial-by-trial variability in the activation response to print in the left inferior frontal gyrus pars triangularis. This relationship held even when accounting for mean levels of activation. This finding suggests that intrasubject variability in trial-by-trial fMRI activation responses to printed words accounts for individual differences in human reading ability that are not fully captured by traditional mean levels of brain activity. Furthermore, this positive relationship between trial-by-trial activation variability and reading skill may provide evidence that neural variability plays a beneficial role during early reading development. [ABSTRACT FROM AUTHOR]

Early language development sets the stage for a lifetime of competence in language and literacy. However, the neural mechanisms associated with the relative advantages of early communication success, or the disadvantages of having delayed language development, are not well explored. In this study, 174 elementary school-age children whose parents reported that they started forming sentences ‘early’, ‘on-time’ or ‘late’ were evaluated with standardized measures of language, reading and spelling. All oral and written language measures revealed consistent patterns for ‘early’ talkers to have the highest level of performance and ‘late’ talkers to have the lowest level of performance. We report functional magnetic resonance imaging data from a subset of early, on-time and late talkers matched for age, gender and performance intelligence quotient that allows evaluation of neural activation patterns produced while listening to and reading real words and pronounceable non-words. Activation in bilateral thalamus and putamen, and left insula and superior temporal gyrus during these tasks was significantly lower in late talkers, demonstrating that residual effects of being a late talker are found not only in behavioural tests of oral and written language, but also in distributed cortical-subcortical neural circuits underlying speech and print processing. Moreover, these findings suggest that the age of functional language acquisition can have long-reaching effects on reading and language behaviour, and on the corresponding neurocircuitry that supports linguistic function into the school-age years. [ABSTRACT FROM AUTHOR]

Converging evidence from a number of neuroimaging studies, including our own, suggest that fluent word identification in reading is related to the functional integrity of two consolidated left hemisphere (LH) posterior systems: a dorsal (temporo-parietal) circuit and a ventral (occipito-temporal) circuit. This posterior system is functionally disrupted in developmental dyslexia. Reading disabled readers, relative to nonimpaired readers, demonstrate heightened reliance on both inferior frontal and right hemisphere posterior regions, presumably in compensation for the LH posterior difficulties. We propose a neurobiological account suggesting that for normally developing readers the dorsal circuit predominates at first, and is associated with analytic processing necessary for learning to integrate orthographic features with phonological and lexical-semantic features of printed words. The ventral circuit constitutes a fast, late-developing, word identification system which underlies fluent word recognition in skilled readers. MRDD Research Reviews 6:207–213, 2000. © 2000 Wiley-Liss, Inc. [ABSTRACT FROM AUTHOR]

Provides experimental evidence for the psychological reality of the concept of neighborhood activation and tests the claim that letter positions yielding neighbors require greater processing than positions not yielding neighbors. Visual work recognition; Search models and activation models of word recognition.

There is now considerable evidence regarding the types of interventions that are effective at remediating reading disabilities on average. It is generally unclear, however, what predicts the magnitude of individual-level change following a given intervention. We examine new predictors of intervention gains that are theoretically grounded in computational models of reading and focus on individual differences in the functional organization of the reading system. Specifically, we estimate the extent to which children with reading disabilities (n = 118 3rd–4th graders) rely on two sources of information during an oral word reading task—print–speech correspondences and semantic imageability—before and after a phonologically-weighted intervention. We show that children who relied more on print–speech regularities and less on imageability preintervention had better intervention gains. In parallel, children who over the course of the intervention exhibited greater increases in their reliance on print–speech correspondences and greater decreases in their reliance on imageability, had better intervention outcomes. Importantly, these two factors were differentially related to specific reading task outcomes, with greater reliance on print–speech correspondences associated with pseudoword naming, while (lesser) reliance on imageability related to word reading and comprehension. We discuss the implications of these findings for theoretical models of reading acquisition and educational practice. Educational Impact and Implications Statement: Why do some children with reading disabilities show substantial gains when provided with an intervention program, whereas others show little to no improvement? Here we argue that such differences in improvement can be tied to the way in which different children read (reflecting the functional organization of their reading system). We measure how different children are impacted by two sources of information of words they read—print–speech correspondences (orthographic–phonological) and semantic imageability—before and after a phonologically-weighted intervention for children with reading disabilities. Results show that children who before the intervention relied more on orthographic–phonological knowledge, and less on semantic imageability, had greater reading skill gains over the intervention. Skill gains were also associated with changes children made in their use of these two sources of information over the intervention. These findings may help in detecting the children who may benefit more or less from a given intervention and advance us toward identifying an optimal intervention program for each individual based on the state of their reading system. [ABSTRACT FROM AUTHOR]

Despite reading being an essential and almost universal skill in the developed world, reading proficiency varies substantially from person to person. To study why, the fMRI field is beginning to turn from single-word or nonword reading tasks to naturalistic stimuli like connected text and listening to stories. To study reading development in children just beginning to read, listening to stories is an appropriate paradigm because speech perception and phonological processing are important for, and are predictors of, reading proficiency. Our study examined the relationship between behavioral reading-related skills and the neural response to listening to stories in the fMRI environment. Functional MRI were gathered in a 3T TIM-Trio scanner. During the fMRI scan, children aged approximately 7 years listened to professionally narrated common short stories and answered comprehension questions following the narration. Analyses of the data used inter-subject correlation (ISC), and representational similarity analysis (RSA). Our primary finding is that ISC reveals areas of increased synchrony in both high- and low-performing emergent readers previously implicated in reading ability/disability. Of particular interest are that several previously identified brain regions (medial temporal gyrus (MTG), inferior frontal gyrus (IFG), inferior temporal gyrus (ITG)) were found to "synchronize" across higher reading ability participants, while lower reading ability participants had idiosyncratic activation patterns in these regions. Additionally, two regions (superior frontal gyrus (SFG) and another portion of ITG) were recruited by all participants, but their specific timecourse of activation depended on reading performance. These analyses support the idea that different brain regions involved in reading follow different developmental trajectories that correlate with reading proficiency on a spectrum rather than the usual dichotomy of poor readers versus strong readers. [ABSTRACT FROM AUTHOR]

• 75 adolescents read and heard time-locked narratives during an fMRI scan. • Inter-subject correlation (ISC) was used to analyze reading ability's impact. • ISC was more sensitive to individual differences in reading ability than GLMs. • Reading ability affects brain activity in more regions than previously thought. • Worse readers had more idiosyncratic responses than better readers across the brain. Recent work using fMRI inter-subject correlation analysis has provided new information about the brain's response to video and audio narratives, particularly in frontal regions not typically activated by single words. This approach is very well suited to the study of reading, where narrative is central to natural experience. But since past reading paradigms have primarily presented single words or phrases, the influence of narrative on semantic processing in the brain – and how that influence might change with reading ability – remains largely unexplored. In this study, we presented coherent stories to adolescents and young adults with a wide range of reading abilities. The stories were presented in alternating visual and auditory blocks. We used a dimensional inter-subject correlation analysis to identify regions in which better and worse readers had varying levels of consistency with other readers. This analysis identified a widespread set of brain regions in which activity timecourses were more similar among better readers than among worse readers. These differences were not detected with standard block activation analyses. Worse readers had higher correlation with better readers than with other worse readers, suggesting that the worse readers had "idiosyncratic" responses rather than using a single compensatory mechanism. Close inspection confirmed that these differences were not explained by differences in IQ or motion. These results suggest an expansion of the current view of where and how reading ability is reflected in the brain, and in doing so, they establish inter-subject correlation as a sensitive tool for future studies of reading disorders. [ABSTRACT FROM AUTHOR]