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      Neural Signatures of the Reading-Writing Connection: Greater Involvement of Writing in Chinese Reading than English Reading

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          Abstract

          Research on cross-linguistic comparisons of the neural correlates of reading has consistently found that the left middle frontal gyrus (MFG) is more involved in Chinese than in English. However, there is a lack of consensus on the interpretation of the language difference. Because this region has been found to be involved in writing, we hypothesize that reading Chinese characters involves this writing region to a greater degree because Chinese speakers learn to read by repeatedly writing the characters. To test this hypothesis, we recruited English L1 learners of Chinese, who performed a reading task and a writing task in each language. The English L1 sample had learned some Chinese characters through character-writing and others through phonological learning, allowing a test of writing-on-reading effect. We found that the left MFG was more activated in Chinese than English regardless of task, and more activated in writing than in reading regardless of language. Furthermore, we found that this region was more activated for reading Chinese characters learned by character-writing than those learned by phonological learning. A major conclusion is that writing regions are also activated in reading, and that this reading-writing connection is modulated by the learning experience. We replicated the main findings in a group of native Chinese speakers, which excluded the possibility that the language differences observed in the English L1 participants were due to different language proficiency level.

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          Most cited references53

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          Representation of manipulable man-made objects in the dorsal stream.

          We used fMRI to examine the neural response in frontal and parietal cortices associated with viewing and naming pictures of different categories of objects. Because tools are commonly associated with specific hand movements, we predicted that pictures of tools, but not other categories of objects, would elicit activity in regions of the brain that store information about motor-based properties. We found that viewing and naming pictures of tools selectively activated the left ventral premotor cortex (BA 6). Single-unit recording studies in monkeys have shown that neurons in the rostral part of the ventral premotor cortex (canonical F5 neurons) respond to the visual presentation of graspable objects, even in the absence of any subsequent motor activity. Thus, the left ventral premotor region that responded selectively to tools in the current study may be the human homolog of the monkey canonical F5 area. Viewing and naming tools also selectively activated the left posterior parietal cortex (BA 40). This response is similar to the firing of monkey anterior intraparietal neurons to the visual presentation of graspable objects. In humans and monkeys, there appears to be a close link between manipulable objects and information about the actions associated with their use. The selective activation of the left posterior parietal and left ventral premotor cortices by pictures of tools suggests that the ability to recognize and identify at least one category of objects (tools) may depend on activity in specific sites of the ventral and dorsal visual processing streams.
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            Functional anatomy of execution, mental simulation, observation, and verb generation of actions: a meta-analysis.

            There is a large body of psychological and neuroimaging experiments that have interpreted their findings in favor of a functional equivalence between action generation, action simulation, action verbalization, and perception of action. On the basis of these data, the concept of shared motor representations has been proposed. Indeed several authors have argued that our capacity to understand other people's behavior and to attribute intention or beliefs to others is rooted in a neural, most likely distributed, execution/observation mechanism. Recent neuroimaging studies have explored the neural network engaged during motor execution, simulation, verbalization, and observation. The focus of this metaanalysis is to evaluate in specific detail to what extent the activated foci elicited by these studies overlap.
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              Evaluation of the dual route theory of reading: a metanalysis of 35 neuroimaging studies.

              Numerous studies concerned with cerebral structures underlying word reading have been published during the last decade. A few controversies, however, together with methodological or theoretical discrepancies between laboratories, still contribute to blurring the overall view of advances effected in neuroimaging. Carried out within the dual route of reading framework, the aim of this metanalysis was to provide an objective picture of these advances. To achieve this, we used an automated analysis method based on the inventory of activation peaks issued from word or pseudoword reading contrasts of 35 published neuroimaging studies. A first result of this metanalysis was that no cluster of activations has been found more recruited by word than pseudoword reading, implying that the first steps of word access may be common to word and word-like stimuli and would take place within a left occipitotemporal region (previously referred to as the Visual Word Form Area-VWFA) situated in the ventral route, at the junction between inferior temporal and fusiform gyri. The results also indicated the existence of brain regions predominantly involved in one of the two routes to access word. The graphophonological conversion seems indeed to rely on left lateralized brain structures such as superior temporal areas, supramarginal gyrus, and the opercular part of the inferior frontal gyrus, these last two regions reflecting a greater load in working memory during such an access. The lexicosemantic route is thought to arise from the coactivation of the VWFA and semantic areas. These semantic areas would encompass a basal inferior temporal area, the posterior part of the middle temporal gyrus, and the triangular part of inferior frontal gyrus. These results confirm the suitability of the dual route framework to account for activations observed in nonpathological subjects while they read.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                16 December 2016
                2016
                : 11
                : 12
                : e0168414
                Affiliations
                [1 ]Department of Communicative Sciences and Disorders, Michigan State University, East Lansing, MI, United States of America
                [2 ]Learning Research and Development Center, University of Pittsburgh, Pittsburgh, PA, United States of America
                University of Akron, UNITED STATES
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                • Conceptualization: FC CAP.

                • Data curation: FC CAP.

                • Formal analysis: FC CAP.

                • Funding acquisition: FC CAP.

                • Investigation: FC CAP.

                • Methodology: FC CAP.

                • Project administration: FC CAP.

                • Resources: FC CAP.

                • Software: FC CAP.

                • Supervision: FC CAP.

                • Validation: FC CAP.

                • Visualization: FC CAP.

                • Writing – original draft: FC CAP.

                • Writing – review & editing: FC CAP.

                Article
                PONE-D-16-34959
                10.1371/journal.pone.0168414
                5161366
                27992505
                1b50e2dd-254d-47f7-bde9-dc74657c8a3e
                © 2016 Cao, Perfetti

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 31 August 2016
                : 30 November 2016
                Page count
                Figures: 6, Tables: 4, Pages: 23
                Funding
                This work was supported by faculty start-up grant awarded to Dr. Fan Cao by Michigan State University.
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                Data are available from the Michigan State University Institutional Data Access / Ethics Committee for public access after 7 years of publication of the original article. Confidential data regarding to the identity of participants is strictly limited to researchers approved by the IRB. Public access of the data, please contact IRB Office: 408 W. Circle Drive, Olds Hall, Rm 207, East Lansing, MI 48824, phone: 517-355-2180.

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