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      Mental Imagery of Speech and Movement Implicates the Dynamics of Internal Forward Models

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          Abstract

          The classical concept of efference copies in the context of internal forward models has stimulated productive research in cognitive science and neuroscience. There are compelling reasons to argue for such a mechanism, but finding direct evidence in the human brain remains difficult. Here we investigate the dynamics of internal forward models from an unconventional angle: mental imagery, assessed while recording high temporal resolution neuronal activity using magnetoencephalography. We compare two overt and covert tasks; our covert, mental imagery tasks are unconfounded by overt input/output demands – but in turn necessitate the development of appropriate multi-dimensional topographic analyses. Finger tapping (studies 1 and 2) and speech experiments (studies 3–5) provide temporally constrained results that implicate the estimation of an efference copy. We suggest that one internal forward model over parietal cortex subserves the kinesthetic feeling in motor imagery. Secondly, observed auditory neural activity ~170 ms after motor estimation in speech experiments (studies 3–5) demonstrates the anticipated auditory consequences of planned motor commands in a second internal forward model in imagery of speech production. Our results provide neurophysiological evidence from the human brain in favor of internal forward models deploying efference copies in somatosensory and auditory cortex, in finger tapping and speech production tasks, respectively, and also suggest the dynamics and sequential updating structure of internal forward models.

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          Most cited references74

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          Predicting human brain activity associated with the meanings of nouns.

          The question of how the human brain represents conceptual knowledge has been debated in many scientific fields. Brain imaging studies have shown that different spatial patterns of neural activation are associated with thinking about different semantic categories of pictures and words (for example, tools, buildings, and animals). We present a computational model that predicts the functional magnetic resonance imaging (fMRI) neural activation associated with words for which fMRI data are not yet available. This model is trained with a combination of data from a trillion-word text corpus and observed fMRI data associated with viewing several dozen concrete nouns. Once trained, the model predicts fMRI activation for thousands of other concrete nouns in the text corpus, with highly significant accuracies over the 60 nouns for which we currently have fMRI data.
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            Motor imagery activates primary sensorimotor area in humans.

            The spatiotemporal patterns of Rolandic mu and beta rhythms were studied during motor imagery with a dense array of EEG electrodes. The subjects were instructed to imagine movements of either the right or the left hand, corresponding to visual stimuli on a computer screen. It was found that unilateral motor imagery results in a short-lasting and localized EEG change over the primary sensorimotor area. The Rolandic rhythms displayed an event-related desynchronization (ERD) only over the contralateral hemisphere. In two of the three investigated subjects, an enhanced Rolandic rhythm was found over the ipsilateral side. The pattern of EEG desynchronization related to imagination of a movement was similar to the pattern during planning of a voluntary movement.
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              Intentional maps in posterior parietal cortex.

              The posterior parietal cortex (PPC), historically believed to be a sensory structure, is now viewed as an area important for sensory-motor integration. Among its functions is the forming of intentions, that is, high-level cognitive plans for movement. There is a map of intentions within the PPC, with different subregions dedicated to the planning of eye movements, reaching movements, and grasping movements. These areas appear to be specialized for the multisensory integration and coordinate transformations required to convert sensory input to motor output. In several subregions of the PPC, these operations are facilitated by the use of a common distributed space representation that is independent of both sensory input and motor output. Attention and learning effects are also evident in the PPC. However, these effects may be general to cortex and operate in the PPC in the context of sensory-motor transformations.
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                Author and article information

                Journal
                Front Psychol
                Front. Psychology
                Frontiers in Psychology
                Frontiers Research Foundation
                1664-1078
                21 October 2010
                2010
                : 1
                : 166
                Affiliations
                [1] 1simpleDepartment of Psychology, New York University New York, NY, USA
                Author notes

                Edited by: Barry Horwitz, National Institutes of Health, USA

                Reviewed by: Iiro P. Jääskeläinen, University of Helsinki, Finland; Heikki Lyytinen, University of Jyväskylä, Finland

                *Correspondence: Xing Tian, Department of Psychology, New York University, 6 Washington Pl. Suite 275, New York, NY 10003, USA. e-mail: xing.tian@ 123456nyu.edu

                This article was submitted to Frontiers in Auditory Cognitive Neuroscience, a specialty of Frontiers in Psychology.

                Article
                10.3389/fpsyg.2010.00166
                3158430
                21897822
                97a88331-cf27-4d52-b44c-4759d458dba4
                Copyright © 2010 Tian and Poeppel.

                This is an open-access article subject to an exclusive license agreement between the authors and the Frontiers Research Foundation, which permits unrestricted use, distribution, and reproduction in any medium, provided the original authors and source are credited.

                History
                : 05 August 2010
                : 20 September 2010
                Page count
                Figures: 10, Tables: 2, Equations: 1, References: 103, Pages: 23, Words: 13989
                Categories
                Psychology
                Original Research

                Clinical Psychology & Psychiatry
                parietal cortex,imagined speech,corollary discharge,articulation,efference copy,meg,motor,auditory cortex

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