Neural mechanism underlying preview effects and masked priming effects in visual word processing.

Two classic experimental paradigms - masked repetition priming and the boundary paradigm - have played a pivotal role in understanding the process of visual word recognition. Traditionally, these paradigms have been employed by different communities of researchers, with their own long-standing research traditions. Nevertheless, a review of the literature suggests that the brain-electric correlates of word processing established with both paradigms may show interesting similarities, in particular with regard to the location, timing, and direction of N1 and N250 effects. However, as of yet, no direct comparison has been undertaken between the two paradigms. In the current study, we used combined eye-tracking/EEG to perform such a within-subject comparison using the same materials (single Chinese characters) as stimuli. To facilitate direct comparisons, we used a simplified version of the boundary paradigm - the single word boundary paradigm. Our results show the typical early repetition effects of N1 and N250 for both paradigms. However, repetition effects in N250 (i.e., a reduced negativity following identical-word primes/previews as compared to different-word primes/previews) were larger with the single word boundary paradigm than with masked priming. For N1 effects, repetition effects were similar across the two paradigms, showing a larger N1 after repetitions as compared to alternations. Therefore, the results indicate that at the neural level, a briefly presented and masked foveal prime produces qualitatively similar facilitatory effects on visual word recognition as a parafoveal preview before a single saccade, although such effects appear to be stronger in the latter case.

Development of EEG alpha and theta oscillations in the maintenance stage of working memory.

Several studies have shown developmental changes in EEG oscillations during working memory tasks. Although the load-modulated theta and alpha activities in adults are well-documented, the findings are inconsistent if children possess the adult-like brain oscillations that are similarly modulated by memory load. The present study compares children's and adults' true theta and alpha EEG oscillations, separated from aperiodic components, in the maintenance stage of working memory. The EEG was recorded in 25 Chinese-speaking children (14 male, Mage = 9.4 yrs) and 31 adults (19 male, Mage = 20.8 yrs) in Hong Kong while they performed an n-back task that included four conditions differing in load (1- vs. 2-back) and stimulus type (Chinese character vs. visual pattern). The results show that aperiodic activities (i.e., broadband power and slope) during the maintenance stage in the n-back task were significantly higher in children than adults. The periodic theta and alpha oscillations also changed with age. More importantly, adults showed significant periodic theta increase with memory load, whereas such an effect was absent in children. Regardless of age, there was a significant alpha power decrease with load increase, and a significant theta power enhancement when maintaining visual patterns than Chinese characters. In adults, load-modulated alpha peak shift (towards higher frequency) was linked to higher behavioral efficiency in the n-back task. In children, higher load-modulated theta enhancement was linked to better behavioral efficiency. The findings suggest that the load-modulated theta power during working memory maintenance matures from childhood to adulthood.

A genome-wide association study of Chinese and English language phenotypes in Hong Kong Chinese children.

Dyslexia and developmental language disorders are important learning difficulties. However, their genetic basis remains poorly understood, and most genetic studies were performed on Europeans. There is a lack of genome-wide association studies (GWAS) on literacy phenotypes of Chinese as a native language and English as a second language (ESL) in a Chinese population. In this study, we conducted GWAS on 34 reading/language-related phenotypes in Hong Kong Chinese bilingual children (including both twins and singletons; total N = 1046). We performed association tests at the single-variant, gene, and pathway levels. In addition, we tested genetic overlap of these phenotypes with other neuropsychiatric disorders, as well as cognitive performance (CP) and educational attainment (EA) using polygenic risk score (PRS) analysis. Totally 5 independent loci (LD-clumped at r2 = 0.01; MAF > 0.05) reached genome-wide significance (p < 5e-08; filtered by imputation quality metric Rsq>0.3 and having at least 2 correlated SNPs (r2 > 0.5) with p < 1e-3). The loci were associated with a range of language/literacy traits such as Chinese vocabulary, character and word reading, and rapid digit naming, as well as English lexical decision. Several SNPs from these loci mapped to genes that were reported to be associated with EA and other neuropsychiatric phenotypes, such as MANEA and PLXNC1. In PRS analysis, EA and CP showed the most consistent and significant polygenic overlap with a variety of language traits, especially English literacy skills. To summarize, this study revealed the genetic basis of Chinese and English abilities in a group of Chinese bilingual children. Further studies are warranted to replicate the findings.

Expectation Modulates Repetition Suppression at Late But Not Early Stages during Visual Word Recognition: Evidence from Event-related Potentials.

Visual word recognition is commonly rapid and efficient, incorporating top-down predictive processing mechanisms. Neuroimaging studies with face stimuli suggest that repetition suppression (RS) reflects predictive processing at the neural level, as this effect is larger when repetitions are more frequent, that is, more expected. It remains unclear, however, at the temporal level whether and how RS and its modulation by expectation occur in visual word recognition. To address this gap, the present study aimed to investigate the presence and time course of these effects during visual word recognition using EEG. Thirty-six native Cantonese speakers were presented with pairs of Chinese written words and performed a nonlinguistic oddball task. The second word of a pair was either a repetition of the first or a different word (alternation). In repetition blocks, 75% of trials were repetitions and 25% were alternations, whereas the reverse was true in alternation blocks. Topographic analysis of variance of EEG at each time point showed robust RS effects in three time windows (141-227 msec, 242-445 msec, and 467-513 msec) reflecting facilitation of visual word recognition. Importantly, the modulation of RS by expectation was observed at the late rather than early intervals (334-387 msec, 465-550 msec, and 559-632 msec) and more than 100 msec after the first RS effects. In the predictive coding view of RS, only late repetition effects are modulated by expectation, whereas early RS effects may be mediated by lower-level predictions. Taken together, our findings provide the first EEG evidence revealing distinct temporal dynamics of RS effects and repetition probability on RS effects in visual processing of Chinese words.

Repetition Suppression for Familiar Visual Words Through Acceleration of Early Processing.

The visual N1 (N170) component with occipito-temporal negativity and fronto-central positivity is sensitive to visual expertise for print. Slightly later, an N200 component with an increase after stimulus repetition was reported to be specific for Chinese, but found at centro-parietal electrodes against a mastoid reference. Given the unusual location, temporal proximity to the N1, and atypical repetition behavior, we aimed at clarifying the relation between the two components. We collected 128-channel EEG data from 18 native Chinese readers during a script decision experiment. Familiar Chinese one- and two-character words were presented among unfamiliar Korean control stimuli with half of the stimuli immediately repeated. Stimulus repetition led to a focal increase in the N1 onset and to a wide-spread decrease in the N1 offset, especially for familiar Chinese and also prominently near the mastoids. A TANOVA analysis corroborated robust repetition effects in the N1 offset across ERP maps with a modulation by script familiarity around 300 ms. Microstate analyses revealed a shorter N1 microstate duration after repetitions, especially for Chinese. The results demonstrate that the previously reported centro-parietal N200 effects after repetitions reflect changes during the N1 offset at occipito-temporal electrodes including the mastoids. Although larger for Chinese, repetition effects could also be found for two-character Korean words, suggesting that they are not specific for Chinese. While the decrease of the N1 offset after repetition is in agreement with a repetition suppression effect, the microstate findings suggest that at least part of the facilitation is due to accelerated processing after repetition.

Dyslexia-related loci are significantly associated with language and literacy in Chinese-English bilingual Hong Kong Chinese twins.

A recent genome-wide association study on dyslexia in 51,800 affected European adults and 1,087,070 controls detected 42 genome-wide significant single nucleotide variants (SNPs). The association between rs2624839 in SEMA3F and reading fluency was replicated in a Chinese cohort. This study explores the genetic overlap between Chinese and English word reading, vocabulary knowledge and spelling, and aims at replicating the association in a unique cohort of bilingual (Chinese-English) Hong Kong Chinese twins. Our result showed an almost complete genetic overlap in vocabulary knowledge (r2 = 0.995), and some genetic overlaps in word reading and spelling (r2 = 0.846, 0.687) across the languages. To investigate the region near rs2624839, we tested proxy SNPs (rs1005678, rs12632110 and rs12494414) at the population level (n = 305-308) and the within-twin level (n = 342-344 [171-172 twin pairs]). All the three SNPs showed significant associations with quantitative Chinese and English vocabulary knowledge (p < 0.05). The strongest association after multiple testing correction was between rs12494414 and English vocabulary knowledge at the within-twin level (p = 0.004). There was a trend of associations with word reading and spelling in English but not in Chinese. Our result suggested that the region near rs2624839 is one of the common genetic factors across English and Chinese vocabulary knowledge and unique factors of English word reading and English spelling in bilingual Chinese twins. A larger sample size is required to validate our findings. Further studies on the relationship between variable expression of SEMA3F, which is important to neurodevelopment, and language and literacy are encouraged.

Neural specialization to English words in Chinese children: Joint contribution of age and English reading abilities.

N1 tuning to words, a neural marker of visual word recognition, develops by an interaction between age and ability. The development of N1 tuning to a second learnt print is unclear. The present study examined the joint contribution of age and English reading abilities to N1 amplitude and tuning to English print in Chinese children in Hong Kong. EEG signals were recorded from 179 children (six to nine years old) while they were performing a repetition detection task comprised of different print stimuli measuring three types of tuning, i.e., coarse tuning (real word versus false font), fine tuning (real versus nonword), and lexicality effect (real versus pseudo word). Children were assessed in English word reading accuracy (EWR) and English sub-lexical orthographic knowledge (EOK). Results indicated that coarse tuning decreased with age but increased with EWR and EOK. Fine tuning uniquely increased with EOK, and the lexicality effect increased with EWR. At last, higher EWR was linked to less right-lateralized coarse tuning in younger children. Taken together, the findings support the visual perceptual expertise account in the L2 context, in that N1 coarse tuning, fine tuning, and lexicality effect are driven by skill improvement.

Neural sensitivity to faces is increased by immersion into a novel ethnic environment: Evidence from ERPs.

Previous reports suggest that East-Asians may show larger face-elicited N170 components in the ERP as compared to Caucasian participants. Since the N170 can be modulated by perceptual expertise, such group differences may be accounted for by differential experience, for example, with logographic versus alphabetic scripts (script system hypothesis) or by exposure to abundant novel faces during the immersion into a new social and/or ethnic environment (social immersion hypothesis). We conducted experiments in Hong Kong and Berlin, recording ERPs in a series of one-back tasks, using same- and other-ethnicity face stimuli in upright and inverted orientation and doodle stimuli. In Hong Kong we tested local Chinese residents and foreign guest students who could not read the logographic script; in Berlin we tested German residents who could not read the logographic script and foreign Chinese visitors. In both experiments, we found significantly larger N170 amplitudes to faces, regardless of ethnicity, in the foreign than in the local groups. Moreover, this effect did not depend on stimulus orientation, suggesting that the N170 group differences do not reflect differences in configural visual processing. A group of short-term German residents in Berlin did not differ in N170 amplitude from long-term residents. Together, these findings indicate that the extensive confrontation with novel other-ethnicity faces during immersion in a foreign culture may enhance the neural response to faces, reflecting the short-term plasticity of the underlying neural system.

Spelling as a way to classify poor Chinese-English literacy skills in Hong Kong Chinese children.

Previous work has predominantly focused on word reading in studying literacy difficulties; very little work has focused on spelling difficulty instead. The present study adopted spelling (dictation) as the criterion to classify poor literacy skills in Hong Kong Chinese-English bilingual children. We examined the cognitive-linguistic skills profiles across four groups of children with different spelling abilities. Based on performances on Chinese and English dictation (criterion = below 25% in a larger sample), four groups were identified, 21 poor spellers of Chinese (PC), 18 poor spellers of English (PE), 27 poor spellers of both languages (PB), and 30 good spellers of both scripts (GB). Measures on language-specific tests of cognitive-linguistic skills (phonological awareness, lexical decision, morphological awareness, rapid naming, and delayed copying) were included to compare the degree of deficit exhibited by each group. With age, grade, and non-verbal intelligence controlled, one-way ANCOVA results revealed that, compared to GB, PC manifested significant deficits in Chinese-delayed copying but scored similarly on all English cognitive-linguistic skills. PE and PB showed significant deficits in Chinese and English phonological awareness compared to PC; they were significantly weaker in English-delayed copying, morphological awareness, and rapid naming (RAN). The PB group was significantly slower in both Chinese and English RAN compared to GB. Findings highlight the critical role of delayed copying in distinguishing poor spellers in both Chinese and English, the importance of phonological awareness for spelling in English but not in Chinese, and the role of automaticity in bilingual spelling difficulties.

Genome-wide analyses of individual differences in quantitatively assessed reading- and language-related skills in up to 34,000 people.

The use of spoken and written language is a fundamental human capacity. Individual differences in reading- and language-related skills are influenced by genetic variation, with twin-based heritability estimates of 30 to 80% depending on the trait. The genetic architecture is complex, heterogeneous, and multifactorial, but investigations of contributions of single-nucleotide polymorphisms (SNPs) were thus far underpowered. We present a multicohort genome-wide association study (GWAS) of five traits assessed individually using psychometric measures (word reading, nonword reading, spelling, phoneme awareness, and nonword repetition) in samples of 13,633 to 33,959 participants aged 5 to 26 y. We identified genome-wide significant association with word reading (rs11208009, P = 1.098 × 10-8) at a locus that has not been associated with intelligence or educational attainment. All five reading-/language-related traits showed robust SNP heritability, accounting for 13 to 26% of trait variability. Genomic structural equation modeling revealed a shared genetic factor explaining most of the variation in word/nonword reading, spelling, and phoneme awareness, which only partially overlapped with genetic variation contributing to nonword repetition, intelligence, and educational attainment. A multivariate GWAS of word/nonword reading, spelling, and phoneme awareness maximized power for follow-up investigation. Genetic correlation analysis with neuroimaging traits identified an association with the surface area of the banks of the left superior temporal sulcus, a brain region linked to the processing of spoken and written language. Heritability was enriched for genomic elements regulating gene expression in the fetal brain and in chromosomal regions that are depleted of Neanderthal variants. Together, these results provide avenues for deciphering the biological underpinnings of uniquely human traits.

Behavioral and neurophysiological aspects of working memory impairment in children with dyslexia.

The present study aimed to identify behavioral and neurophysiological correlates of dyslexia which could potentially predict reading difficulty. One hundred and three Chinese children with and without dyslexia (Grade 2 or 3, 7- to 11-year-old) completed both verbal and visual working memory (n-back) tasks with concurrent EEG recording. Data of 74 children with sufficient usable EEG data are reported here. Overall, the typically developing control group (N = 28) responded significantly faster and more accurately than the group with dyslexia (N = 46), in both types of tasks. Group differences were also found in EEG band power in the retention phase of the tasks. Moreover, forward stepwise logistic regression demonstrated that both behavioral and neurophysiological measures predicted reading difficulty uniquely. Dyslexia was associated with higher frontal midline theta activity and reduced upper-alpha power in the posterior region. This finding is discussed in relation to the neural efficiency hypothesis. Whether these behavioral and neurophysiological patterns can longitudinally predict later reading development among preliterate children requires further investigation.

Is there magnocellular facilitation of early neural processes underlying visual word recognition? Evidence from masked repetition priming with ERPs.

An influential theory in the field of visual object recognition proposes that it is the fast magnocellular (M) system that facilitates neural processing of spatially more fine-grained information rather the slower parvocellular (P) system. While written words can be considered as a special type of visual objects, it is unknown whether magnocellular facilitation also plays a role in reading. We used a masked priming paradigm that has been shown to result in neural facilitation in visual word processing and tested whether these facilitating effects are mediated by the magnocellular system. In two experiments, we manipulated the influence of magnocellular and parvocellular systems on visual processing of a contextually predictable target character by contrasting high versus low spatial frequency and luminance versus color contrast, respectively. In addition, unchanged (normal) primes were included in both experiments as a manipulation check. As expected, unchanged primes elicited typical repetition effects in the N1, N250 and P3 components of the ERP in both experiments. In the experiment manipulating spatial contrast, we obtained repetition effects only for the N1 component for both M- and P-biased primes. In the luminance versus color contrast experiment, repetition effects were found in N1 and N250 for both M- and P- biased primes. Furthermore, no interactions were found between M-vs. P-biased prime types and repetition. Together these results indicate that M- and P- information contributes jointly to early neural processes underlying visual word recognition.

Children With Chinese Dyslexia Acquiring English Literacy: Interaction Between Cognitive Subtypes of Dyslexia and Orthographies.

This study investigated the impact of Chinese dyslexia subtypes on English literacy skills (i.e., reading fluency and dictation) in Hong Kong children. Eighty-four Cantonese-speaking children officially diagnosed with dyslexia (Mage = 103 months) and 48 age-matched typically developing (TD) children were tested. Cluster analysis with performances on Chinese syllable awareness (CSA), Chinese phonemic awareness (CPA), Chinese phonological memory (CPM), Chinese orthographic awareness (COA), and matrix reasoning (MR) yielded three cognitive subtypes: the phonological deficit (PD) subtype, the orthographic deficit (OD) subtype, and the global deficit (GD) subtype. After controlling for English language experience, age, and gender, all three dyslexia subtype groups performed significantly worse in English word reading fluency and dictation than the TD children. In addition, the PD group performed worse in English PA; the OD group performed worse in English OA; and the GD group performed worse in all English skills except English PM. We compared the level of impairment in literacy between languages and dyslexia subtypes. In word reading fluency, all subtype groups experienced less impairment in English than Chinese, while the OD group showed the largest English advantage. In dictation, only the OD group showed a significant language effect favoring English. The findings suggest that different subtypes of Chinese dyslexia bear different risks for difficulties in English literacy.

Predicting Reading From Behavioral and Neural Measures - A Longitudinal Event-Related Potential Study.

Fluent reading is characterized by fast and effortless decoding of visual and phonological information. Here we used event-related potentials (ERPs) and neuropsychological testing to probe the neurocognitive basis of reading in a sample of children with a wide range of reading skills. We report data of 51 children who were measured at two time points, i.e., at the end of first grade (mean age 7.6 years) and at the end of fourth grade (mean age 10.5 years). The aim of this study was to clarify whether next to behavioral measures also basic unimodal and bimodal neural measures help explaining the variance in the later reading outcome. Specifically, we addressed the question of whether next to the so far investigated unimodal measures of N1 print tuning and mismatch negativity (MMN), a bimodal measure of audiovisual integration (AV) contributes and possibly enhances prediction of the later reading outcome. We found that the largest variance in reading was explained by the behavioral measures of rapid automatized naming (RAN), block design and vocabulary (46%). Furthermore, we demonstrated that both unimodal measures of N1 print tuning (16%) and filtered MMN (7%) predicted reading, suggesting that N1 print tuning at the early stage of reading acquisition is a particularly good predictor of the later reading outcome. Beyond the behavioral measures, the two unimodal neural measures explained 7.2% additional variance in reading, indicating that basic neural measures can improve prediction of the later reading outcome over behavioral measures alone. In this study, the AV congruency effect did not significantly predict reading. It is therefore possible that audiovisual congruency effects reflect higher levels of multisensory integration that may be less important for reading acquisition in the first year of learning to read, and that they may potentially gain on relevance later on.

Resting state EEG network modularity predicts literacy skills in L1 Chinese but not in L2 English.

EEG network modularity, as a proxy for cognitive plasticity, has been proposed to be a more reliable neural marker than power and coherence in predicting learning outcomes. The present study examined the associations between resting state EEG network modularity and both L1 Chinese and L2 English literacy skills among 90 Hong Kong first to fifth graders. The modularity indices of different frequency bands were highly correlated with one another. An exploratory factor analysis, performed to extract a general modularity index, explained 77.1% of the total variance. The modularity index was positively associated with Chinese word reading, Chinese phonological awareness, Chinese morphological awareness, and Chinese reading comprehension but was not significantly correlated with English word reading or English morphological awareness. Findings suggest that resting state EEG network modularity is likely to serve as a reasonable, reliable, and cost-effective neural marker of the development of first language but not second language literacy skills.

Remediation of a phonological representation deficit in Chinese children with dyslexia: A comparison between metalinguistic training and working memory training.

A form-preparation task in the language production field was adopted to examine output phonological representations in Chinese dyslexia and their susceptibility to training. Forty-one Chinese children with dyslexia (7-11 years old) and 36 chronological age controls completed this task. The controls demonstrated a marginally significant syllable facilitation effect (d = -0.13), indicating their use of syllable-sized phonological representations during speech production, while the group with dyslexia showed a significantly different pattern (d = 0.04), opposite to the direction of a facilitation effect. The children with dyslexia were then randomly assigned to either metalinguistic training (N = 22) or working memory training (N = 19). Only the metalinguistic training subgroup demonstrated a significant syllable facilitation effect afterward (metalinguistic: d = -0.13; working memory: d = -0.01). The results suggest the presence of a phonological representation deficit at the syllable level in Chinese dyslexia and its possible remediation by metalinguistic training. Such a phonological deficit in readers of a logographic script strongly supports the impaired phonological representation view of developmental dyslexia. A video abstract of this article can be viewed at https://youtu.be/zT2Be0xMkh0.

Electroencephalography decoding of Chinese characters in primary school children and its prediction for word reading performance and development.

Research on what neural mechanisms facilitate word reading development in non-alphabetic scripts is relatively rare. The present study was among the first to adopt a multivariate pattern classification analysis to decode electroencephalographic signals recorded for primary school children (N = 236) while performing a Chinese character decision task. Chinese is an ideal script for studying the relationship between neural discriminability (i.e., decodability) of the orthography and behavioral word reading skills since the mapping from orthography to phonology is relatively arbitrary in Chinese. This was also among the first empirical attempts to examine the extent to which decoding performance can predict current and subsequent word reading skills using a longitudinal design. Results showed that neural activation patterns of real characters can be distinguished from activation patterns for pseudo-characters, non-characters, and random stroke combinations in both younger and older children. Topography of the transformed classifier weights revealed two distinct cognitive sub-processes underlying single character recognition, but temporal generalization analysis suggested common neural mechanisms between the distinct cognitive sub-processes. Suggestive evidence from correlational and hierarchical regression analyses showed that decoding performance, assessed on average 2 months before the year 2 behavioral testing, predicted both year 1 word reading performance and the development of word reading fluency over the year. Results demonstrate that decoding performance, one indicator of how the neural system is functionally organized in processing characters and character-like stimuli, can serve as a useful neural marker in predicting current word reading skills and the capacity to learn to read.

Genome-wide association study reveals new insights into the heritability and genetic correlates of developmental dyslexia.

Developmental dyslexia (DD) is a learning disorder affecting the ability to read, with a heritability of 40-60%. A notable part of this heritability remains unexplained, and large genetic studies are warranted to identify new susceptibility genes and clarify the genetic bases of dyslexia. We carried out a genome-wide association study (GWAS) on 2274 dyslexia cases and 6272 controls, testing associations at the single variant, gene, and pathway level, and estimating heritability using single-nucleotide polymorphism (SNP) data. We also calculated polygenic scores (PGSs) based on large-scale GWAS data for different neuropsychiatric disorders and cortical brain measures, educational attainment, and fluid intelligence, testing them for association with dyslexia status in our sample. We observed statistically significant (p < 2.8 × 10-6) enrichment of associations at the gene level, for LOC388780 (20p13; uncharacterized gene), and for VEPH1 (3q25), a gene implicated in brain development. We estimated an SNP-based heritability of 20-25% for DD, and observed significant associations of dyslexia risk with PGSs for attention deficit hyperactivity disorder (at pT = 0.05 in the training GWAS: OR = 1.23[1.16; 1.30] per standard deviation increase; p = 8 × 10-13), bipolar disorder (1.53[1.44; 1.63]; p = 1 × 10-43), schizophrenia (1.36[1.28; 1.45]; p = 4 × 10-22), psychiatric cross-disorder susceptibility (1.23[1.16; 1.30]; p = 3 × 10-12), cortical thickness of the transverse temporal gyrus (0.90[0.86; 0.96]; p = 5 × 10-4), educational attainment (0.86[0.82; 0.91]; p = 2 × 10-7), and intelligence (0.72[0.68; 0.76]; p = 9 × 10-29). This study suggests an important contribution of common genetic variants to dyslexia risk, and novel genomic overlaps with psychiatric conditions like bipolar disorder, schizophrenia, and cross-disorder susceptibility. Moreover, it revealed the presence of shared genetic foundations with a neural correlate previously implicated in dyslexia by neuroimaging evidence.