Brain activation during phonological and semantic processing of Chinese characters in deaf signers

Previous studies found altered brain function in deaf individuals reading alphabetic orthographies. However, it is not known whether similar alterations of brain function are characteristic of non-alphabetic writing systems and whether alterations are specific to certain kinds of lexical tasks. Here we examined differences in brain activation between Chinese congenitally deaf individuals (CD) and hearing controls (HC) during character reading tasks requiring phonological and semantic judgments. For both tasks, we found that CD showed less activation than HC in left inferior frontal gyrus, but greater activation in several right hemisphere regions including inferior frontal gyrus, angular gyrus, and inferior temporal gyrus. Although many group differences were similar across tasks, greater activation in right middle frontal gyrus was more pronounced for the rhyming compared to the meaning task. Finally, within the deaf individuals better performance on the rhyming task was associated with less activation in right inferior parietal lobule and angular gyrus. Our results in Chinese CD are broadly consistent with previous studies in alphabetic languages suggesting greater engagement of inferior frontal gyrus and inferior parietal cortex for reading that is largely independent of task, with the exception of right middle frontal gyrus for phonological processing. The brain behavior correlations potentially indicate that CD that more efficiently use the right hemisphere are better readers.

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title: The Neural System Underlying Chinese Logograph Reading
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title: Functional Anatomy of Intra- and Cross-Modal Lexical Tasks
abstract: Functional magnetic resonance imaging (fMRI) was used to examine lexical processing in normal adults (20-35 years). Two tasks required only intramodal processing (spelling judgments with visual input and rhyming judgments with auditory input) and two tasks required cross-modal processing between phonologic and orthographic representations (spelling judgments with auditory input and rhyming judgments with visual input). Each task led to greater activation in the unimodal association area concordant with the modality of input, namely fusiform gyrus (BA 19, 37) for written words and superior temporal gyrus (BA 22, 42) for spoken words. Cross-modal tasks generated greater activation in posterior heteromodal regions including the supramarginal and angular gyri (BA 40, 39). Cross-modal tasks generated additional activation in unimodal areas representing the target of conversion, superior temporal gyrus for visual rhyming and fusiform gyrus for auditory spelling. Our findings suggest that the fusiform gyrus processes orthographic word forms, the superior temporal gyrus processes phonologic word forms, and posterior heteromodal regions are involved in the conversion between orthography and phonology.

title: Development of Brain Mechanisms for Processing Orthographic and Phonologic Representations
abstract: Developmental differences in the neurocognitive networks for lexical processing were examined in 15 adults and 15 children (9-to 12-year-olds) using functional magnetic resonance imaging (fMRI). The lexical tasks involved spelling and rhyming judgments in either the visual or auditory modality. These lexical tasks were compared with nonlinguistic control tasks involving judgments of line patterns or tone sequences. The first main finding was that adults showed greater activation than children during the cross-modal lexical tasks in a region proposed to be involved in mapping between orthographic and phonologic representations. The visual rhyming task, which required conversion from orthography to phonology, produced greater activation for adults in the angular gyrus. The auditory spelling task, which required the conversion from phonology to orthography, also produced greater activation for adults in the angular gyrus. The greater activation for adults suggests they may have a more elaborated posterior heteromodal system for mapping between representational systems. The second main finding was that adults showed greater activation than children during the intra-modal lexical tasks in the angular gyrus. The visual spelling and auditory rhyming did not require conversion between orthography and phonology for correct performance but the adults showed greater activation in a system implicated for this mapping. The greater activation for adults suggests that they have more interactive convergence between representational systems during lexical processing.

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abstract: Brain imaging studies have explored the neural mechanisms of recovery in adults following acquired disorders and, more recently, childhood developmental disorders. However, the neural systems underlying adult rehabilitation of neurobiologically based learning disabilities remain unexplored, despite their high incidence. Here we characterize the differences in brain activity during a phonological manipulation task before and after a behavioral intervention in adults with developmental dyslexia. Phonologically targeted training resulted in performance improvements in tutored compared to nontutored dyslexics, and these gains were associated with signal increases in bilateral parietal and right perisylvian cortices. Our findings demonstrate that behavioral changes in tutored dyslexic adults are associated with (1) increased activity in those left-hemisphere regions engaged by normal readers and (2) compensatory activity in the right perisylvian cortex. Hence, behavioral plasticity in adult developmental dyslexia involves two distinct neural mechanisms, each of which has previously been observed either for remediation of developmental or acquired reading disorders.

title: Modality independence of word comprehension
abstract: Functional magnetic resonance imaging (fMRI) was used to examine the functional anatomy of word comprehension in the auditory and visual modalities of presentation. We asked our subjects to determine if word pairs were semantically associated (e.g., table, chair) and compared this to a reference task where they were asked to judge whether word pairs rhymed (e.g., bank, tank). This comparison showed task‐specific and modality‐independent activation for semantic processing in the heteromodal cortices of the left inferior frontal gyrus (BA 46, 47) and left middle temporal gyrus (BA 21). There were also modality‐specific activations in the fusiform gyrus (BA 37) for written words and in the superior temporal gyrus (BA 22) for spoken words. Our findings are consistent with the hypothesis that word form recognition (lexical encoding) occurs in unimodal cortices and that heteromodal brain regions in the anterior as well as posterior components of the language network subserve word comprehension (semantic decoding).  Brain Mapping 16:251–261, 2002.

title: Effects of Lexicality, Frequency, and Spelling-to-Sound Consistency on the Functional Anatomy of Reading
abstract: Functional neuroimaging was used to investigate three factors that affect reading performance: first, whether a stimulus is a word or pronounceable non-word (lexicality), second, how often a word is encountered (frequency), and third, whether the pronunciation has a predictable spelling-to-sound correspondence (consistency). Comparisons between word naming (reading) and visual fixation scans revealed stimulus-related activation differences in seven regions. A left frontal region showed effects of consistency and lexicality, indicating a role in orthographic to phonological transformation. Motor cortex showed an effect of consistency bilaterally, suggesting that motoric processes beyond high-level representations of word phonology influence reading performance. Implications for the integration of these results into theoretical models of word reading are discussed.

title: Functional Specialization for Semantic and Phonological Processing in the Left Inferior Prefrontal Cortex
abstract: Neuroimaging and neuropsychological studies have implicated left inferior prefrontal cortex (LIPC) in both semantic and phonological processing. In this study, functional magnetic resonance imaging was used to examine whether separate LIPC regions participate in each of these types of processing. Performance of a semantic decision task resulted in extensive LIPC activation compared to a perceptual control task. Phonological processing of words and pseudowords in a syllable-counting task resulted in activation of the dorsal aspect of the left inferior frontal gyrus near the inferior frontal sulcus (BA 44/45) compared to a perceptual control task, with greater activation for nonwords compared to words. In a direct comparison of semantic and phonological tasks, semantic processing preferentially activated the ventral aspect of the left inferior frontal gyrus (BA 47/45). A review of the literature demonstrated a similar distinction between left prefrontal regions involved in semantic processing and phonological/lexical processing. The results suggest that a distinct region in the left inferior frontal cortex is involved in semantic processing, whereas other regions may subserve phonological processes engaged during both semantic and phonological tasks.

title: Cultural Constraints on Brain Development: Evidence from a Developmental Study of Visual Word Processing in Mandarin Chinese
abstract: Developmental differences in phonological and orthographic processing in Chinese were examined in 9 year olds, 11 year olds, and adults using functional magnetic resonance imaging. Rhyming and spelling judgments were made to 2-character words presented sequentially in the visual modality. The spelling task showed greater activation than the rhyming task in right superior parietal lobule and right inferior temporal gyrus, and there were developmental increases across tasks bilaterally in these regions in addition to bilateral occipital cortex, suggesting increased involvement over age on visuo-orthographic analysis. The rhyming task showed greater activation than the spelling task in left superior temporal gyrus and there were developmental decreases across tasks in this region, suggesting reduced involvement over age on phonological representations. The rhyming and spelling tasks included words with conflicting orthographic and phonological information (i.e., rhyming words spelled differently or nonrhyming words spelled similarly) or nonconflicting information. There was a developmental increase in the difference between conflicting and nonconflicting words in left inferior parietal lobule, suggesting greater engagement of systems for mapping between orthographic and phonological representations. Finally, there were developmental increases across tasks in an anterior (Broadman area [BA] 45, 46) and posterior (BA 9) left inferior frontal gyrus, suggesting greater reliance on controlled retrieval and selection of posterior lexical representations.

title: Segregating semantic and syntactic aspects of processing in the human brain: an fMRI investigation of different word types.
abstract: The processing of single words that varied in their semantic (concrete/abstract word) and syntactic (content/function word) status was investigated under different task demands (semantic/ syntactic task) in an event-related functional magnetic resonance imaging experiment. Task demands to a large degree determined which subparts of the neuronal network supporting word processing were activated. Semantic task demands selectively activated the left pars triangularis of the inferior frontal gyrus (BA 45) and the posterior part of the left middle/superior temporal gyrus (BA 21/22/37). In contrast, syntactic processing requirements led to an increased activation in the inferior tip of the left frontal operculum (BA 44) and the cortex lining the junction of the inferior frontal and inferior precentral sulcus (BA 44/6). Moreover, for these latter areas a word class by concreteness interaction was observed when a syntactic judgement was required. This interaction can be interpreted as a prototypicality effect: non-prototypical members of a word class, i.e. concrete function words and abstract content words, showed a larger activation than prototypical members, i.e. abstract function words and concrete content words. The combined data suggest that the activation pattern underlying word processing is predicted neither by syntactic class nor semantic concreteness but, rather, by task demands focusing either on semantic or syntactic aspects. Thus, our findings that semantic and syntactic aspects of processing are both functionally distinct and involve different subparts of the neuronal network underlying word processing support a domain-specific organization of the language system.

title: Specialization of phonological and semantic processing in Chinese word reading
abstract: The purpose of this study was to examine the neurocognitive network for processing visual word forms in native Chinese speakers using functional magnetic resonance imaging (fMRI). In order to compare the processing of phonological and semantic representations, we developed parallel rhyming and meaning association judgment tasks that required explicit access and manipulation of these representations. Subjects showed activation in left inferior/middle frontal gyri, bilateral medial frontal gyri, bilateral middle occipital/fusiform gyri, and bilateral cerebella for both the rhyming and meaning tasks. A direct comparison of the tasks revealed that the rhyming task showed more activation in the posterior dorsal region of the inferior/middle frontal gyrus (BA 9/44) and in the inferior parietal lobule (BA 40). The meaning task showed more activation in the anterior ventral region of the inferior/middle frontal gyrus (BA 47) and in the superior/middle temporal gyrus (BA 22,21). These findings are consistent with previous studies in English that suggest specialization of inferior frontal regions for the access and manipulation of phonological vs. semantic representations, but also suggest that this specialization extends to the middle frontal gyrus for Chinese. These findings are also consistent with the suggestion that the left middle temporal gyrus is involved in representing semantic information and the left inferior parietal lobule is involved in mapping between orthographic and phonological representations.

title: Cross‐cultural effect on the brain revisited: Universal structures plus writing system variation
abstract: Recognizing printed words requires the mapping of graphic forms, which vary with writing systems, to linguistic forms, which vary with languages. Using a newly developed meta‐analytic approach, aggregated Gaussian‐estimated sources (AGES; Chein et al. [ 2002 ]: Psychol Behav 77:635–639), we examined the neuroimaging results for word reading within and across writing systems and languages. To find commonalities, we compiled 25 studies in English and other Western European languages that use an alphabetic writing system, 9 studies of native Chinese reading, 5 studies of Japanese Kana (syllabic) reading, and 4 studies of Kanji (morpho‐syllabic) reading. Using the AGES approach, we created meta‐images within each writing system, isolated reliable foci of activation, and compared findings across writing systems and languages. The results suggest that these writing systems utilize a common network of regions in word processing. Writing systems engage largely the same systems in terms of gross cortical regions, but localization within those regions suggests differences across writing systems. In particular, the region known as the visual word form area (VWFA) shows strikingly consistent localization across tasks and across writing systems. This region in the left mid‐fusiform gyrus is critical to word recognition across writing systems and languages.  Brain Mapp 25:92–104, 2005.

title: Developmental differences of neurocognitive networks for phonological and semantic processing in Chinese word reading
abstract: Developmental differences in the neurocognitive networks for phonological and semantic processing in Chinese word reading were examined in 13 adults and 13 children using functional magnetic resonance imaging (fMRI). Rhyming and semantic association judgments were made to two‐character words that were presented sequentially in the visual modality. These lexical tasks were compared with a nonlinguistic control task involving judgment of line patterns. The first main finding was that adults showed greater activation than children in right middle occipital gyrus on both the meaning and rhyming task, suggesting adults more effectively engage right hemisphere brain regions involved in the visual‐spatial analysis of Chinese characters. The second main finding was that adults showed greater activation than children in left inferior parietal lobule for the rhyming as compared with the meaning task, suggesting greater specialization of phonological processing in adults. The third main finding was that children who had better performance in the rhyming task on characters with conflicting orthographic and phonological information relative to characters with nonconflicting information showed greater activation in left middle frontal gyrus, suggesting greater engagement of brain regions involved in the integration of orthography and phonology. Hum Brain Mapp, 2009.

title: Neural systems of second language reading are shaped by native language
abstract: Reading in a second language (L2) is a complex task that entails an interaction between L2 and the native language (L1). To study the underlying mechanisms, we used functional magnetic resonance imaging (fMRI) to visualize Chinese–English bilinguals' brain activity in phonological processing of logographic Chinese and alphabetic English, two written languages with a sharp contrast in phonology and orthography. In Experiment 1, we found that phonological processing of Chinese characters recruits a neural system involving left middle frontal and posterior parietal gyri, cortical regions that are known to contribute to spatial information representation, spatial working memory, and coordination of cognitive resources as a central executive system. We assume that the peak activation of this system is relevant to the unique feature of Chinese that a logographic character has a square configuration that maps onto a monosyllabic unit of speech. Equally important, when our bilingual subjects performed a phonological task on English words, this neural system was most active, whereas brain areas mediating English monolinguals' fine‐grained phonemic analysis, as demonstrated by Experiment 2, were only weakly activated. This suggests that our bilingual subjects were applying their L1 system to L2 reading and that the lack of letter‐to‐sound conversion rules in Chinese led Chinese readers to being less capable of processing English by recourse to an analytic reading system on which English monolinguals rely. Our brain imaging findings lend strongest support to the idea that language experience tunes the cortex.  2003.

title: Evidence for developmental changes in the visual word processing network beyond adolescence
abstract: Late development of specialization in the visual word processing system was examined using event-related potentials (ERP) and functional magnetic resonance imaging (fMRI) of word and symbol string processing in groups of adolescents (15.2-17.3 years) and adults (19.8-30.8 years). We focused our ERP analyses on fast visual activity: the occipital P1 (82-131 ms) modulated by physical stimulus characteristics and the occipito-temporal N1 (132-256 ms) reflecting visual tuning for print. Our fMRI analyses concentrated on basal occipito-temporal activations in the visual word form area VWFA. For words, the correlation of fMRI activation in the VWFA and N1 amplitude confirmed the close relationship of the electrophysiological N1 with metabolic activity in the VWFA. Further support for this relationship came from low resolution electromagnetic tomography localizing the word-specific N1 near the VWFA. Both imaging techniques revealed age-independent differences between words and symbol strings. Late development, however, was preferentially detected with ERPs. Decreases of P1 and N1 amplitudes with age were not limited to words and suggested further maturation of the underlying brain microstructure and function. Following adolescence, decreasing N1 latencies specific to words point to continued specialization of the visual word processing system. Both N1 and fMRI measures correlated with reading performance. In summary, the similarity of global fMRI activation patterns between groups suggests a fully established distribution of the reading network in adolescence, while the decreasing N1 latencies for words indicate protracted fine tuning after adolescence.

title: Neural correlates of mapping from phonology to orthography in children performing an auditory spelling task.
abstract: Age-related differences (9- to 15-year-olds) in the neural correlates of mapping from phonology to orthography were examined with functional magnetic resonance imaging (fMRI). Participants were asked to determine if two spoken words had the same spelling for the rime (corresponding letters after the first consonant or consonant cluster). Some of the word pairs had conflicting orthography and phonology (e.g. jazz-has, pint-mint) whereas other pairs had non-conflicting information (e.g. press-list, gate-hate) . There were age-related increases in activation for lexical processing (across conflicting and non-conflicting conditions) in left inferior parietal lobule, suggesting that older children have a more elaborated system for mapping between phonology and orthography that includes connections at different grain sizes (e.g. phonemes, onset-rimes, syllables). In addition, we found that the conflicting conditions had lower accuracy, slower reaction time and greater activation in left inferior frontal gyrus as compared to non-conflicting conditions. Higher accuracy was also correlated with greater activation in left inferior frontal gyrus for the most difficult conflicting condition (e.g. jazz-has). The finding of both a conflict effect and a correlation with accuracy in left inferior frontal gyrus suggests that this region may be involved in resolving the conflict between orthographic and phonological representations.

title: Development of neural mechanisms for reading
abstract: The complexities of pediatric brain imaging have precluded studies that trace the neural development of cognitive skills acquired during childhood. Using a task that isolates reading-related brain activity and minimizes confounding performance effects, we carried out a cross-sectional functional magnetic resonance imaging (fMRI) study using subjects whose ages ranged from 6 to 22 years. We found that learning to read is associated with two patterns of change in brain activity: increased activity in left-hemisphere middle temporal and inferior frontal gyri and decreased activity in right inferotemporal cortical areas. Activity in the left-posterior superior temporal sulcus of the youngest readers was associated with the maturation of their phonological processing abilities. These findings inform current reading models and provide strong support for Orton's 1925 theory of reading development.

title: Developmental changes in the neural correlates of semantic processing
abstract: Functional magnetic resonance imaging (fMRI) was used to explore the neural correlates of semantic judgments in the auditory modality in a group of 9- to 15-year-old children. Subjects were required to indicate if word pairs were related in meaning. Consistent with previous findings in adults, children showed activation in bilateral superior temporal gyri (BA 22) for recognizing spoken words as well as activations in bilateral inferior frontal gyri (BAs 47, 45) and left middle temporal gyrus (BA 21) for semantic processing. The neural substrates of semantic association and age differences were also investigated. Words with strong semantic association elicited significantly greater activation in the left inferior parietal lobule (BA 40), whereas words with weak semantic association elicited activation in left inferior frontal gyrus (BAs 47/45). Correlations with age were observed in the left middle temporal gyrus (BA 21) and the right inferior frontal gyrus (BA 47). The pattern of results for semantic association implies that the left inferior parietal lobule effectively integrates highly related semantic features and the left inferior frontal gyrus becomes more active for words that require a greater search for semantic associations. The developmental results suggest that older children recruit the right inferior frontal gyrus as they conduct a broader semantic search and the left middle temporal gyrus to provide more efficient access to semantic representations.

title: Developmental aspects of language processing: fMRI of verbal fluency in children and adults
abstract: We examined developmental differences, in location and extent of fMRI language activation maps, between adults and children while performing a semantic fluency task. We studied 29 adults and 16 children with echo planar imaging BOLD fMRI at 1.5 T using covert semantic verbal fluency (generation of words to categories compared to rest) using a block design. Post task testing was administered to assess performance. Individual data were analyzed with an a priori region of interest approach from t maps (t = 4) and asymmetry indices (AI). Group studies were analyzed using SPM 99 (Wellcome, UK; fixed effect, corrected P < 0.0001). We found no significant differences in location or laterality of activation between adults and children for a semantic verbal fluency task. Adults activated more pixels than children in left inferior frontal gyrus and left middle frontal gyrus, but AIs were the similar across ages (r2 < 0.09). Extent or laterality of activation was not affected by performance (r2 < 0.15). The brain areas that process semantic verbal fluency are similar in children and adults. The laterality of activation does not change appreciably with age and appears to be strongly lateralized by age 7 years.  2003.

title: A longitudinal functional magnetic resonance imaging study of language development in children 5 to 11 years old
abstract: Language skills continue to develop rapidly in children during the school‐age years, and the “snapshot” view of the neural substrates of language provided by current neuroimaging studies cannot capture the dynamic changes associated with brain development. The aim of this study was to conduct a 5‐year longitudinal investigation of language development using functional magnetic resonance imaging in healthy children.

title: Stanford Achievement Test—8th Edition: Reading Comprehension Subgroup Results
abstract: The following pages contain a set of five graphs depicting the median scaled scores and corresponding grade equivalents (GEs) on the 8th Edition Stanford Achievement Test (SAT-8) Reading Comprehension subtest for various groups of deaf and hard of hearing students. The median scores used in these graphs were extracted from a series of special decile norms computed for the population of deaf and hard of hearing students in the United States during a special achievement test project conducted by the Center for Assessment and Demographic Studies (CADS). Since this population is heterogeneous with respect to various demographic, audiological, and educational characteristics, misconceptions could occur when interpreting the scores of individual students. For example, if a particular student has handicaps in addition to hearing impairment, it may not be useful to compare that student to the entire population of deaf and hard of hearing students. An individual student may also belong to more than one relevant group. Thus, it is appropriate at times to use more than one set of group norms when interpreting a score (e.g., handicap status norms, ethnic group norms, and program type norms). A complete listing of the special group norms is available in:

title: The role of inferior frontal gyrus and inferior parietal lobule in semantic processing of Chinese characters
abstract: Functional magnetic resonance imaging was used to explore the neural correlates of semantic judgments to Chinese characters. Adult participants were asked to indicate if character pairs were related in meaning that were arranged in a continuous variable according to association strength. This parametric manipulation allowed for a more precise determination of the role of the left inferior parietal lobule in processing meaning, which has not been reported in previous Chinese studies. Consistent with previous findings in English, participants showed activation in left inferior frontal gyrus (BA 47, 45) and left posterior middle temporal gyrus (BA 21). Characters with stronger semantic association elicited greater activation in left inferior parietal lobule (BA 39), suggesting stronger integration of highly related semantic features. By contrast, characters with weaker semantic association elicited greater activation in both an anterior ventral region (BA 47) and a mid-ventral region of left inferior frontal gyrus (BA 45), suggesting a controlled retrieval process and a selection process. Our findings of association strength are discussed in a proposed neuro-anatomical model of semantic processing.

title: Sign-based short-term coding of American Sign Language signs and printed English words by congenitally deaf signers
abstract: A short-term serial recall task consisting of lists of five American Sign Language signs and lists of five printed English words was administered to eight congenitally deaf subjects. For both modes of presentation, the stimuli in a given list were (1) high in phonological relatedness, (2) high in cherological relatedness (i.e., they were similar in terms of the formational attributes of their sign-equivalents), or (3) low in both phonological and cherological similarity (control). For both modes of presentation, recall accuracy was significantly lower on cherologically related lists than on both phonologically related lists and control lists. Phonologically related lists did not differ significantly from control lists. These results suggest the possibility that ways of looking at human memory which rely on a preference for phonological coding in short-term memory processes may reflect the audiocentricity of the experimenter rather than the inherent nature of such processes.

title: When deaf signers read English: Do written words activate their sign translations?
abstract: Deaf bilinguals for whom American Sign Language (ASL) is the first language and English is the second language judged the semantic relatedness of word pairs in English. Critically, a subset of both the semantically related and unrelated word pairs were selected such that the translations of the two English words also had related forms in ASL. Word pairs that were semantically related were judged more quickly when the form of the ASL translation was also similar whereas word pairs that were semantically unrelated were judged more slowly when the form of the ASL translation was similar. A control group of hearing bilinguals without any knowledge of ASL produced an entirely different pattern of results. Taken together, these results constitute the first demonstration that deaf readers activate the ASL translations of written words under conditions in which the translation is neither present perceptually nor required to perform the task.

title: Signed and spoken language perception studied by positron emission tomography
abstract: Sign and spoken language seem to be localized in the same brain areas. They elicit similar regional cerebral blood flow (rCBF) patterns, even though sign language is dependent on spatial information. We investigated sign and spoken language perception in a group of healthy bilingual subjects. Four videotaped activation conditions were used during PET imaging: (1) sign language, (2) spoken language, (3) spoken language with mouth covered, and(4) spoken language on a sound track while showing a motionless face. Spoken language (condition 4) activated significantly the perisylvian cortex(Brodmann areas 22 and 43) bilaterally. Sign language activated the visual association areas (Brodmann areas 37 and 19) but did not selectively activate parietal regions. A reciprocal relationship was observed between the level of activation in visual language perception areas and that in auditory perception areas. We conclude that when healthy bilingual subjects use the visual route for sign language perception, the functional anatomy overlaps that of language processing containing both auditory and visual components.

title: Neural Systems Underlying Spatial Language in American Sign Language
abstract: A [(15)O]water PET experiment was conducted to investigate the neural regions engaged in processing constructions unique to signed languages: classifier predicates in which the position of the hands in signing space schematically represents spatial relations among objects. Ten deaf native signers viewed line drawings depicting a spatial relation between two objects (e.g., a cup on a table) and were asked either to produce a classifier construction or an American Sign Language (ASL) preposition that described the spatial relation or to name the figure object (colored red). Compared to naming objects, describing spatial relationships with classifier constructions engaged the supramarginal gyrus (SMG) within both hemispheres. Compared to naming objects, naming spatial relations with ASL prepositions engaged only the right SMG. Previous research indicates that retrieval of English prepositions engages both right and left SMG, but more inferiorly than for ASL classifier constructions. Compared to ASL prepositions, naming spatial relations with classifier constructions engaged left inferior temporal (IT) cortex, a region activated when naming concrete objects in either ASL or English. Left IT may be engaged because the handshapes in classifier constructions encode information about object type (e.g., flat surface). Overall, the results suggest more right hemisphere involvement when expressing spatial relations in ASL, perhaps because signing space is used to encode the spatial relationship between objects.

title: Hand and Mouth: Cortical Correlates of Lexical Processing in British Sign Language and Speechreading English
abstract: Spoken languages use one set of articulatorsthe vocal tract, whereas signed languages use multiple articulators, including both manual and facial actions. How sensitive are the cortical circuits for language processing to the particular articulators that are observed? This question can only be addressed with participants who use both speech and a signed language. In this study, we used functional magnetic resonance imaging to compare the processing of speechreading and sign processing in deaf native signers of British Sign Language (BSL) who were also proficient speechreaders. The following questions were addressed: To what extent do these different language types rely on a common brain network? To what extent do the patterns of activation differ? How are these networks affected by the articulators that languages use? Common peri-sylvian regions were activated both for speechreading English words and for BSL signs. Distinctive activation was also observed reflecting the language form. Speechreading elicited greater activation in the left mid-superior temporal cortex than BSL, whereas BSL processing generated greater activation at the temporo-parieto-occipital junction in both hemispheres. We probed this distinction further within BSL, where manual signs can be accompanied by different types of mouth action. BSL signs with speech-like mouth actions showed greater superior temporal activation, whereas signs made with non-speech-like mouth actions showed more activation in posterior and inferior temporal regions. Distinct regions within the temporal cortex are not only differentially sensitive to perception of the distinctive articulators for speech and for sign but also show sensitivity to the different articulators within the (signed) language.

title: Enhanced activation of the left inferior frontal gyrus in deaf and dyslexic adults during rhyming
abstract: Hearing developmental dyslexics and profoundly deaf individuals both have difficulties processing the internal structure of words (phonological processing) and learning to read. In hearing non-impaired readers, the development of phonological representations depends on audition. In hearing dyslexics, many argue, auditory processes may be impaired. In congenitally profoundly deaf individuals, auditory speech processing is essentially absent. Two separate literatures have previously reported enhanced activation in the left inferior frontal gyrus in both deaf and dyslexic adults when contrasted with hearing non-dyslexics during reading or phonological tasks. Here, we used a rhyme judgement task to compare adults from these two special populations to a hearing non-dyslexic control group. All groups were matched on non-verbal intelligence quotient, reading age and rhyme performance. Picture stimuli were used since this requires participants to generate their own phonological representations, rather than have them partially provided via text. By testing well-matched groups of participants on the same task, we aimed to establish whether previous literatures reporting differences between individuals with and without phonological processing difficulties have identified the same regions of differential activation in these two distinct populations. The data indicate greater activation in the deaf and dyslexic groups than in the hearing non-dyslexic group across a large portion of the left inferior frontal gyrus. This includes the pars triangularis, extending superiorly into the middle frontal gyrus and posteriorly to include the pars opercularis, and the junction with the ventral precentral gyrus. Within the left inferior frontal gyrus, there was variability between the two groups with phonological processing difficulties. The superior posterior tip of the left pars opercularis, extending into the precentral gyrus, was activated to a greater extent by deaf than dyslexic participants, whereas the superior posterior portion of the pars triangularis extending into the ventral pars opercularis, was activated to a greater extent by dyslexic than deaf participants. Whether these regions play differing roles in compensating for poor phonological processing is not clear. However, we argue that our main finding of greater inferior frontal gyrus activation in both groups with phonological processing difficulties in contrast to controls suggests greater reliance on the articulatory component of speech during phonological processing when auditory processes are absent (deaf group) or impaired (dyslexic group). Thus, the brain appears to develop a similar solution to a processing problem that has different antecedents in these two populations.

title: Cerebral asymmetry in children when reading Chinese characters.
abstract: This study examined cerebral asymmetry, especially in the hierarchical visual system, when reading Chinese characters. Twelve right-handed Chinese children (mean age = 11.6 years) were scanned while performing semantic and phonological tasks. Strong leftward asymmetry was found in the left inferior frontal cortex (BA44/45/47), the parietal lobule (BA40), and the cingulate cortex (BA24/32). In the visual system, we found significant left-hemispheric dominance in the fusiform cortex (BA19/37), but no asymmetry was found in the primary visual cortex (BA17/18). The differential results for the primary visual cortex versus high-order visual cortex (i.e., the fusiform cortex) are discussed in terms of the contribution of the logographic nature of Chinese characters to the asymmetry pattern in the hierarchical visual system.

title: Cerebral lateralization of deaf and hearing individuals for linguistic comparison judgments
abstract: A group of congenitally deaf adults and a group of hearing adults, both fluent in sign language, were tested to determine cerebral lateralization. In the most revealing task, subjects were given a series of trials in which they were fist presented with a videotaped sign and then with a word exposed tachistoscopically to the right visual field or left visual field, and were required to judge whether the word corresponded to the sign or not. The results suggested that the comparison processes involved in the decision were performed more efficiently by the left hemisphere for hearing subjects and by the right hemisphere for deaf subjects. However, the deaf subjects performed as well as the hearing subjects in the left hemisphere, suggesting that the deaf are not impeded by their auditory-speech handicap from developing the left hemisphere for at least some types of linguistic processing.

title: Lexical Processing in Deaf Readers: An fMRI Investigation of Reading Proficiency
abstract: Individuals with significant hearing loss often fail to attain competency in reading orthographic scripts which encode the sound properties of spoken language. Nevertheless, some profoundly deaf individuals do learn to read at age-appropriate levels. The question of what differentiates proficient deaf readers from less-proficient readers is poorly understood but topical, as efforts to develop appropriate and effective interventions are needed. This study uses functional magnetic resonance imaging (fMRI) to examine brain activation in deaf readers (N = 21), comparing proficient (N = 11) and less proficient (N = 10) readers’ performance in a widely used test of implicit reading. Proficient deaf readers activated left inferior frontal gyrus and left middle and superior temporal gyrus in a pattern that is consistent with regions reported in hearing readers. In contrast, the less-proficient readers exhibited a pattern of response characterized by inferior and middle frontal lobe activation (right>left) which bears some similarity to areas reported in studies of logographic reading, raising the possibility that these individuals are using a qualitatively different mode of orthographic processing than is traditionally observed in hearing individuals reading sound-based scripts. The evaluation of proficient and less-proficient readers points to different modes of processing printed English words. Importantly, these preliminary findings allow us to begin to establish the impact of linguistic and educational factors on the neural systems that underlie reading achievement in profoundly deaf individuals.

title: Functional dissociations within the inferior parietal cortex in verbal working memory
abstract: Neuroimaging studies of working memory have revealed two sites in the left supramarginal gyrus that may support the short-term storage of phonological information. Activation in the left dorsal aspect of the inferior parietal cortex (DIPC) has been observed in contrasts of working memory load, whereas activation in the ventral aspect of the inferior parietal cortex (VIPC) has been found primarily in contrast of information type (verbal vs. nonverbal). Our goal was to determine whether these two areas are functionally distinct or if instead they are part of a homogeneous region with large variations in the focus of peak activity. Toward this end, we used fMRI to assess the neural response in two working memory tasks (N-back and item recognition) in which we also manipulated memory load and the type of information to be recalled (verbal vs. nonverbal). We found both DIPC and VIPC activation in the same group of subjects and further demonstrated that they have differential sensitivity to our experimental factors. Only the DIPC showed robust load effects, whereas only the VIPC showed reliable effects of information type. These results help to account for the differences observed in between-subject comparisons, and they indicate that the two regions are functionally dissociable. In contrast to the DIPC, activity of the VIPC was also recruited in the fixation and low-load conditions, a surprising result that has not been fully explored in prior studies. Despite their distinctive patterns of performance, neither of these regions displayed a pattern of activity that entirely corresponds to common assumptions of a dedicated phonological short-term store (STS). Instead, we hypothesize that the DIPC may support domain-general executive processes, while the VIPC may support phonological encoding-recoding processes central to a variety of language tasks.

title: Left insula activation: a marker for language attainment in bilinguals.
abstract: Several lines of evidence suggest the importance of phonological working memory (PWM) in language acquisition. We investigated the neural correlates of PWM in young adults who were under compelling social pressure to be bilingual. Equal bilinguals had high proficiency in English and Chinese as measured by a standardized examination, whereas unequal bilinguals were proficient in English but not Chinese. Both groups were matched on several measures of nonverbal intelligence and working memory. In-scanner behavioral results did not show between-group differences. Of the regions showing load-dependent increments in activation, the left insula showed greater activation in equal bilinguals. Unequal bilinguals showed greater task-related deactivation in the anterior medial frontal region and greater anterior cingulate activation. Although unequal bilinguals kept apace with equal bilinguals in the simple PWM task, the differential cortical activations suggest that more optimal engagement of PWM in the latter may correlate with better second-language attainment.

title: Orthographic and phonological processing of Chinese characters: an fMRI study
abstract: The present study used functional magnetic resonance imaging (fMRI) to investigate the neural mechanisms underlying the orthographic and phonological processing of Chinese characters. Four tasks were devised, including one homophone judgment and three physical judgments of characters, pseudo-characters, and Korean-like nonsense figures. While the left occipitotemporal region, left dorsal processing stream, and right middle frontal gyrus constitute a network for orthographic processing, the left premotor gyrus, left middle/inferior frontal gyrus, supplementary motor area (SMA), and the left temporoparietal region work in concert for phonological processing. The ventral part of the left inferior frontal cortex responds specifically to the character stimuli, suggesting a general lexical processing role for this region for linguistic material. The stronger activation of the dorsal visual stream by Chinese homophone judgment pinpoints a tight coupling between phonological representation of Chinese characters and corresponding orthographic percepts. The concomitant engagement of sets of regions for different levels of Chinese orthographic and phonological processing is consistent with the notion of distributed parallel processing.

title: Biological abnormality of impaired reading is constrained by culture
abstract: Developmental dyslexia is characterized by a severe reading problem in people who have normal intelligence and schooling. Impaired reading of alphabetic scripts is associated with dysfunction of left temporoparietal brain regions. These regions perform phonemic analysis and conversion of written symbols to phonological units of speech (grapheme-to-phoneme conversion); two central cognitive processes that mediate reading acquisition. Furthermore, it has been assumed that, in contrast to cultural diversities, dyslexia in different languages has a universal biological origin. Here we show using functional magnetic resonance imaging with reading-impaired Chinese children and associated controls, that functional disruption of the left middle frontal gyrus is associated with impaired reading of the Chinese language (a logographic rather than alphabetic writing system). Reading impairment in Chinese is manifested by two deficits: one relating to the conversion of graphic form (orthography) to syllable, and the other concerning orthography-to-semantics mapping. Both of these processes are critically mediated by the left middle frontal gyrus, which functions as a centre for fluent Chinese reading that coordinates and integrates various information about written characters in verbal and spatial working memory. This finding provides an insight into the fundamental pathophysiology of dyslexia by suggesting that rather than having a universal origin, the biological abnormality of impaired reading is dependent on culture.