Neural mechanisms underlying auditory feedback control of speech

The neural substrates underlying auditory feedback control of speech were investigated using a combination of functional magnetic resonance imaging (fMRI) and computational modeling. Neural responses were measured while subjects spoke monosyllabic words under two conditions: (i) normal auditory feedback of their speech and (ii) auditory feedback in which the first formant frequency of their speech was unexpectedly shifted in real time. Acoustic measurements showed compensation to the shift within approximately 136 ms of onset. Neuroimaging revealed increased activity in bilateral superior temporal cortex during shifted feedback, indicative of neurons coding mismatches between expected and actual auditory signals, as well as right prefrontal and Rolandic cortical activity. Structural equation modeling revealed increased influence of bilateral auditory cortical areas on right frontal areas during shifted speech, indicating that projections from auditory error cells in posterior superior temporal cortex to motor correction cells in right frontal cortex mediate auditory feedback control of speech.

[1]  Daniel M. Wolpert,et al.  Forward Models for Physiological Motor Control , 1996, Neural Networks.

[2]  Robert J Zatorre,et al.  Neural Substrates Governing Audiovocal Integration for Vocal Pitch Regulation in Singing , 2005, Annals of the New York Academy of Sciences.

[3]  Yihong Yang,et al.  Physiological Mapping of Human Auditory Cortices with a Silent Event-Related fMRI Technique , 2002, NeuroImage.

[4]  G. Rizzolatti,et al.  The Cortical Motor System , 2001, Neuron.

[5]  G E Alexander,et al.  Neural representations of the target (goal) of visually guided arm movements in three motor areas of the monkey. , 1990, Journal of neurophysiology.

[6]  S. Lehéricy,et al.  3-D diffusion tensor axonal tracking shows distinct SMA and pre-SMA projections to the human striatum. , 2004, Cerebral cortex.

[7]  M. Kawato,et al.  Reorganization of Brain Activity for Multiple Internal Models After Short But Intensive Training , 2007, Cortex.

[8]  Daniel S. O'Leary,et al.  An MRI-Based Parcellation Method for the Temporal Lobe , 2000, NeuroImage.

[9]  A. Nobre,et al.  The Cerebellum Predicts the Timing of Perceptual Events , 2008, The Journal of Neuroscience.

[10]  Reza Shadmehr,et al.  Evolution of motor memory during the seconds after observation of motor error. , 2007, Journal of neurophysiology.

[11]  Rachel E. Stark STAGES OF SPEECH DEVELOPMENT IN THE FIRST YEAR OF LIFE 1 1This work was supported by NINCDS Grant NS.09628 and by NICHD Grant HD. 11970. , 1980 .

[12]  Riitta Salmelin,et al.  Subject's own speech reduces reactivity of the human auditory cortex , 1999, Neuroscience Letters.

[13]  D. Louis Collins,et al.  Application of Information Technology: A Four-Dimensional Probabilistic Atlas of the Human Brain , 2001, J. Am. Medical Informatics Assoc..

[14]  H. Lüders,et al.  Functional connectivity in the human language system: a cortico-cortical evoked potential study. , 2004, Brain : a journal of neurology.

[15]  Dylan F. Cooke,et al.  The Cortical Control of Movement Revisited , 2002, Neuron.

[16]  J. Kalaska,et al.  A comparison of movement direction-related versus load direction- related activity in primate motor cortex, using a two-dimensional reaching task , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[17]  G. Curio,et al.  Speaking modifies voice‐evoked activity in the human auditory cortex , 2000, Human brain mapping.

[18]  G. Schaltenbrand,et al.  The effects on speech and language of stereotactical stimulation in thalamus and corpus callosum , 1975, Brain and Language.

[19]  Alan C. Evans,et al.  Morphology, morphometry and probability mapping of the pars opercularis of the inferior frontal gyrus: an in vivo MRI analysis , 1999, The European journal of neuroscience.

[20]  Srikantan S. Nagarajan,et al.  How is auditory feedback processed during speaking , 2007 .

[21]  G. Rizzolatti,et al.  Hearing Sounds, Understanding Actions: Action Representation in Mirror Neurons , 2002, Science.

[22]  Hermann Ackermann,et al.  Phonemic Vowel Length Contrasts in Cerebellar Disorders , 1999, Brain and Language.

[23]  D. Kennedy,et al.  The young adult human brain: an MRI-based morphometric analysis. , 1994, Cerebral cortex.

[24]  D. Wolpert,et al.  The cerebellum is involved in predicting the sensory consequences of action , 1999, Neuroreport.

[25]  D. Poeppel,et al.  Dorsal and ventral streams: a framework for understanding aspects of the functional anatomy of language , 2004, Cognition.

[26]  Michael I. Jordan,et al.  Sensorimotor adaptation in speech production. , 1998, Science.

[27]  Michael J. Martinez,et al.  Bias between MNI and Talairach coordinates analyzed using the ICBM‐152 brain template , 2007, Human brain mapping.

[28]  J. Kalinowski,et al.  Effect of delayed auditory feedback on normal speakers at two speech rates. , 2002, The Journal of the Acoustical Society of America.

[29]  Alan C. Evans,et al.  Modulation of cerebral blood-flow in the human auditory cortex during speech: role of motor-to-sensory discharges , 1996, NeuroImage.

[30]  J. Desmond,et al.  Lobular Patterns of Cerebellar Activation in Verbal Working-Memory and Finger-Tapping Tasks as Revealed by Functional MRI , 1997, The Journal of Neuroscience.

[31]  T. M. Nearey,et al.  Effects of consonant environment on vowel formant patterns. , 1997, The Journal of the Acoustical Society of America.

[32]  K. Hegedüs,et al.  Akinetic mutism associated with bicingular lesions: Clinicopathological and functional anatomical correlates , 2004, European archives of psychiatry and neurological sciences.

[33]  Yan Wang,et al.  Spatial distribution and density of prefrontal cortical cells projecting to three sectors of the premotor cortex , 2002, Neuroreport.

[34]  Ellen Douglas-Cowie,et al.  Speech production in profound postlingual deafness. , 1983 .

[35]  J. Karhu,et al.  Anticipatory cerebellar responses during somatosensory omission in man , 2000, Human brain mapping.

[36]  J. Penney,et al.  The functional anatomy of disorders of the basal ganglia , 1995, Trends in Neurosciences.

[37]  Gregory Hickok,et al.  Left posterior auditory-related cortices participate both in speech perception and speech production: Neural overlap revealed by fMRI , 2006, Brain and Language.

[38]  G. E. Alexander,et al.  Parallel organization of functionally segregated circuits linking basal ganglia and cortex. , 1986, Annual review of neuroscience.

[39]  Edward E. Smith,et al.  The Role of Parietal Cortex in Verbal Working Memory , 1998, The Journal of Neuroscience.

[40]  F. Chollet,et al.  Differential fMRI Responses in the Left Posterior Superior Temporal Gyrus and Left Supramarginal Gyrus to Habituation and Change Detection in Syllables and Tones , 1999, NeuroImage.

[41]  M. K. Wolf,et al.  Lacunar and Other Subcortical Infarctions , 1997, Neurology.

[42]  S. Petersen,et al.  Neuroimaging studies of word reading. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[43]  S Kuriki,et al.  Motor planning center for speech articulation in the normal human brain. , 1999, Neuroreport.

[44]  M. Merzenich,et al.  Modulation of the Auditory Cortex during Speech: An MEG Study , 2002, Journal of Cognitive Neuroscience.

[45]  P. Morosan,et al.  Human Primary Auditory Cortex: Cytoarchitectonic Subdivisions and Mapping into a Spatial Reference System , 2001, NeuroImage.

[46]  Terry M. Peters,et al.  3D statistical neuroanatomical models from 305 MRI volumes , 1993, 1993 IEEE Conference Record Nuclear Science Symposium and Medical Imaging Conference.

[47]  Yasmin L. Hashambhoy,et al.  Neural Correlates of Reach Errors , 2005, The Journal of Neuroscience.

[48]  John F. Houde,et al.  Compensatory responses to brief perturbations of speech amplitude , 2005 .

[49]  Michael I. Jordan,et al.  Sensorimotor adaptation of speech I: Compensation and adaptation. , 2002, Journal of speech, language, and hearing research : JSLHR.

[50]  D. Wolpert,et al.  Why can't you tickle yourself? , 2000, Neuroreport.

[51]  M. Arbib,et al.  Language within our grasp , 1998, Trends in Neurosciences.

[52]  EFFECTS OF CATEGORIZATION TRAINING ON AUDITORY PERCEPTION AND CORTICAL REPRESENTATIONS , 2001 .

[53]  Jeffery A. Jones,et al.  Remapping Auditory-Motor Representations in Voice Production , 2005, Current Biology.

[54]  David Silbersweig,et al.  Functional neuroanatomy of verbal self-monitoring , 1996 .

[55]  A. J. Yates Delayed Auditory Feedback , 1958, Psychological bulletin.

[56]  D. Poeppel,et al.  The cortical organization of speech processing , 2007, Nature Reviews Neuroscience.

[57]  G. Rizzolatti,et al.  Corticocortical connections of area F3 (SMA‐proper) and area F6 (pre‐SMA) in the macaque monkey , 1993, The Journal of comparative neurology.

[58]  P. Bentler,et al.  Cutoff criteria for fit indexes in covariance structure analysis : Conventional criteria versus new alternatives , 1999 .

[59]  Stephen Grossberg,et al.  How laminar frontal cortex and basal ganglia circuits interact to control planned and reactive saccades , 2004, Neural Networks.

[60]  R. Passingham The frontal lobes and voluntary action , 1993 .

[61]  J. A. Shafer,et al.  Understanding aphasia. , 1954, Archives of Physical Medicine and Rehabilitation.

[62]  Guinevere F. Eden,et al.  Meta-Analysis of the Functional Neuroanatomy of Single-Word Reading: Method and Validation , 2002, NeuroImage.

[63]  D. Poeppel,et al.  Towards a functional neuroanatomy of speech perception , 2000, Trends in Cognitive Sciences.

[64]  D. Wolpert,et al.  Sensorimotor attenuation by central motor command signals in the absence of movement , 2006, Nature Neuroscience.

[65]  Mitsuo Kawato,et al.  Internal models for motor control and trajectory planning , 1999, Current Opinion in Neurobiology.

[66]  J. Mazziotta,et al.  Cortical mechanisms of human imitation. , 1999, Science.

[67]  S. Clarke,et al.  Intrinsic connectivity of human auditory areas: a tracing study with DiI , 2001, The European journal of neuroscience.

[68]  J M VANBUREN,et al.  CONFUSION AND DISTURBANCE OF SPEECH FROM STIMULATION IN VICINITY OF THE HEAD OF THE CAUDATE NUCLEUS. , 1963, Journal of neurosurgery.

[69]  S. Clarke,et al.  Cytochrome Oxidase, Acetylcholinesterase, and NADPH-Diaphorase Staining in Human Supratemporal and Insular Cortex: Evidence for Multiple Auditory Areas , 1997, NeuroImage.

[70]  N. Makris,et al.  Cortical abnormalities in schizophrenia identified by structural magnetic resonance imaging. , 1999, Archives of general psychiatry.

[71]  Masao Ito Mechanisms of motor learning in the cerebellum 1 1 Published on the World Wide Web on 24 November 2000. , 2000, Brain Research.

[72]  R J Wise,et al.  Separate neural subsystems within 'Wernicke's area'. , 2001, Brain : a journal of neurology.

[73]  C Büchel,et al.  Brain regions involved in articulation , 1999, The Lancet.

[74]  Hiroaki Gomi,et al.  The motor cortex is involved in reflexive compensatory adjustment of speech articulation , 2005, Neuroreport.

[75]  S. Scott,et al.  Identification of a pathway for intelligible speech in the left temporal lobe. , 2000, Brain : a journal of neurology.

[76]  Dottie M. Clower,et al.  The Inferior Parietal Lobule Is the Target of Output from the Superior Colliculus, Hippocampus, and Cerebellum , 2001, The Journal of Neuroscience.

[77]  R. Shadmehr,et al.  Long-term adaptation to dynamics of reaching movements: a PET study , 2001, Experimental Brain Research.

[78]  G. Rizzolatti,et al.  Premotor cortex and the recognition of motor actions. , 1996, Brain research. Cognitive brain research.

[79]  Satrajit S. Ghosh,et al.  Effects of jaw perturbation on cortical activity during speech production , 2004 .

[80]  S. Koyama,et al.  Neural correlates of auditory feedback control in human , 2007, Neuroscience.

[81]  Xiaoqin Wang,et al.  Sensory-motor interaction in the primate auditory cortex during self-initiated vocalizations. , 2003, Journal of neurophysiology.

[82]  D. Kewley-Port,et al.  Vowel intelligibility in clear and conversational speech for normal-hearing and hearing-impaired listeners. , 2002, The Journal of the Acoustical Society of America.

[83]  Thomas M Donath,et al.  Control of voice fundamental frequency in speaking versus singing. , 2003, The Journal of the Acoustical Society of America.

[84]  Gabriel Curio,et al.  Differential effects of overt, covert and replayed speech on vowel-evoked responses of the human auditory cortex , 1999, Neuroscience Letters.

[85]  D. Kennedy,et al.  Anatomic segmentation and volumetric calculations in nuclear magnetic resonance imaging. , 1989, IEEE transactions on medical imaging.

[86]  E. Kokmen,et al.  Motor Speech Disorders , 1976 .

[87]  Jason A. Tourville,et al.  A cortical and cerebellar parcellation system for speech studies , 2010 .

[88]  Christine Preibisch,et al.  Cortical plasticity associated with stuttering therapy. , 2005, Journal of fluency disorders.

[89]  P. Strick,et al.  Cerebellar Projections to the Prefrontal Cortex of the Primate , 2001, The Journal of Neuroscience.

[90]  P. Strick,et al.  Cerebellar output channels. , 1997, International review of neurobiology.

[91]  Sylvain Houle,et al.  Functional neuroimaging of cerebellar activation during single word reading and verb generation in stuttering and nonstuttering adults , 2001, Neuroscience Letters.

[92]  Jun Tanji,et al.  Prefrontal cortical cells projecting to the supplementary eye field and presupplementary motor area in the monkey , 2005, Neuroscience Research.

[93]  Shinji Maeda,et al.  Compensatory Articulation During Speech: Evidence from the Analysis and Synthesis of Vocal-Tract Shapes Using an Articulatory Model , 1990 .

[94]  Trigeminal evoked potentials in somatosensory cortex of the Macaca mulatta. , 1995, Journal of neurosurgery.

[95]  Jeremy D W Greenlee,et al.  Functional connections within the human inferior frontal gyrus , 2007, The Journal of comparative neurology.

[96]  Dottie M. Clower,et al.  Basal ganglia and cerebellar inputs to 'AIP'. , 2005, Cerebral cortex.

[97]  F. Binkofski,et al.  Motor functions of the Broca’s region , 2004, Brain and Language.

[98]  T. Wiley,et al.  Recognition of speech produced in noise. , 2001, Journal of speech, language, and hearing research : JSLHR.

[99]  A R Damasio,et al.  The anatomical basis of conduction aphasia. , 1980, Brain : a journal of neurology.

[100]  Kevin G. Munhall,et al.  The role of auditory feedback during phonation: studies of Mandarin tone production , 2002, J. Phonetics.

[101]  D. Oller THE EMERGENCE OF THE SOUNDS OF SPEECH IN INFANCY , 1980 .

[102]  C. Larson,et al.  Effects of pitch-shift velocity on voice Fo responses. , 2000, The Journal of the Acoustical Society of America.

[103]  R. Shadmehr,et al.  Effects of Deep Brain Stimulation on Adaptive Control of Reaching , 2005, 2005 IEEE Engineering in Medicine and Biology 27th Annual Conference.

[104]  Frank H. Guenther The representation of prototypical and non-prototypical vowels in peri-sylvian cortical areas , 2000 .

[105]  A. Friederici,et al.  Auditory Language Comprehension: An Event-Related fMRI Study on the Processing of Syntactic and Lexical Information , 2000, Brain and Language.

[106]  G. Rizzolatti,et al.  Mirror neurons responding to the observation of ingestive and communicative mouth actions in the monkey ventral premotor cortex , 2003, The European journal of neuroscience.

[107]  Yi Xu,et al.  Compensation for pitch-shifted auditory feedback during the production of Mandarin tone sequences. , 2004, The Journal of the Acoustical Society of America.

[108]  Satrajit S. Ghosh,et al.  ROI-based analysis of fMRI data incorporating individual differences in brain anatomy , 2001, NeuroImage.

[109]  Alan C. Evans,et al.  MRI Atlas of the Human Cerebellum , 2000 .

[110]  Peter Stoeter,et al.  Cerebellar speech representation: lesion topography in dysarthria as derived from cerebellar ischemia and functional magnetic resonance imaging. , 2003, Archives of neurology.

[111]  Richard G. Lomax,et al.  A Beginner's Guide to Structural Equation Modeling , 2022 .

[112]  Bruce Crosson,et al.  Subcortical functions in language and memory , 1992 .

[113]  G. Fesl,et al.  Inferior central sulcus: variations of anatomy and function on the example of the motor tongue area , 2003, NeuroImage.

[114]  M. E. Lutman,et al.  Hearing science and hearing disorders , 1983 .

[115]  H. Lüders,et al.  Functional connectivity in human cortical motor system: a cortico-cortical evoked potential study. , 2006, Brain : a journal of neurology.

[116]  A M Galaburda,et al.  The planum temporale. , 1993, Archives of neurology.

[117]  Daniel M. Wolpert,et al.  The Cerebellum Contributes to Somatosensory Cortical Activity during Self-Produced Tactile Stimulation , 1999, NeuroImage.

[118]  Willem J. M. Levelt,et al.  A theory of lexical access in speech production , 1999, Behavioral and Brain Sciences.

[119]  Robert Sekuler,et al.  Corticolimbic Interactions Associated with Performance on a Short-Term Memory Task Are Modified by Age , 2000, The Journal of Neuroscience.

[120]  R. Ingham,et al.  Stuttered and fluent speech production: An ALE meta‐analysis of functional neuroimaging studies , 2005, Human brain mapping.

[121]  E. Formisano,et al.  Neural correlates of verbal feedback processing: An fMRI study employing overt speech , 2007, Human brain mapping.

[122]  N. Makris,et al.  MRI-Based Topographic Parcellation of Human Neocortex: An Anatomically Specified Method with Estimate of Reliability , 1996, Journal of Cognitive Neuroscience.

[123]  Charles R Larson,et al.  Early pitch-shift response is active in both steady and dynamic voice pitch control. , 2002, The Journal of the Acoustical Society of America.

[124]  F H Guenther,et al.  Speech sound acquisition, coarticulation, and rate effects in a neural network model of speech production. , 1995, Psychological review.

[125]  K. Amunts,et al.  Multimodal architectonic mapping of human superior temporal gyrus , 2005, Anatomy and Embryology.

[126]  R. Schumacker,et al.  A beginner's guide to structural equation modeling, 2nd ed. , 2004 .

[127]  U Klose,et al.  Dynamic pattern of brain activation during sequencing of word strings evaluated by fMRI. , 1999, Brain research. Cognitive brain research.

[128]  G. Rizzolatti,et al.  Parietal cortex: from sight to action , 1997, Current Opinion in Neurobiology.

[129]  Mark D. Johnson,et al.  The Role of the Human Thalamus in Language and Memory: Evidence from Electrophysiological Studies , 2000, Brain and Cognition.

[130]  Hermann Ackermann,et al.  Speech deficits in ischaemic cerebellar lesions , 1992, Journal of Neurology.

[131]  Frank H Guenther,et al.  The distinctness of speakers' /s/-/S/ contrast is related to their auditory discrimination and use of an articulatory saturation effect. , 2004, Journal of speech, language, and hearing research : JSLHR.

[132]  J Ashburner,et al.  Functional neuroimaging of speech perception in six normal and two aphasic subjects. , 1999, The Journal of the Acoustical Society of America.

[133]  K. Simonyan,et al.  Efferent subcortical projections of the laryngeal motorcortex in the rhesus monkey , 2003, Brain Research.

[134]  H. Scheich,et al.  Phonetic Perception and the Temporal Cortex , 2002, NeuroImage.

[135]  O. Foerster,et al.  THE MOTOR CORTEX IN MAN IN THE LIGHT OF HUGHLINGS JACKSON'S DOCTRINES , 1936 .

[136]  Anne-Lise Giraud,et al.  Distinct functional substrates along the right superior temporal sulcus for the processing of voices , 2004, NeuroImage.

[137]  Hiroshi Imamizu,et al.  Human cerebellar activity reflecting an acquired internal model of a new tool , 2000, Nature.

[138]  D. Wolpert,et al.  Internal models in the cerebellum , 1998, Trends in Cognitive Sciences.

[139]  Mitsuo Kawato,et al.  A computational model of four regions of the cerebellum based on feedback-error learning , 2004, Biological Cybernetics.

[140]  Kuansan Wang,et al.  Spectral shape analysis in the central auditory system , 1995, IEEE Trans. Speech Audio Process..

[141]  M Lotze,et al.  The representation of articulation in the primary sensorimotor cortex , 2000, Neuroreport.

[142]  R P Lesser,et al.  Cortical tongue area studied by chronically implanted subdural electrodes--with special reference to parietal motor and frontal sensory responses. , 1994, Brain : a journal of neurology.

[143]  J. Kropotov,et al.  Selection of actions in the basal ganglia-thalamocortical circuits: review and model. , 1999, International journal of psychophysiology : official journal of the International Organization of Psychophysiology.

[144]  C. Larson,et al.  Instructing subjects to make a voluntary response reveals the presence of two components to the audio-vocal reflex , 1999, Experimental Brain Research.

[145]  Michael I. Jordan,et al.  Sensorimotor Adaptation of Speech I , 2002 .

[146]  David Poeppel,et al.  The analysis of speech in different temporal integration windows: cerebral lateralization as 'asymmetric sampling in time' , 2003, Speech Commun..

[147]  T. Robbins,et al.  Inhibition and the right inferior frontal cortex , 2004, Trends in Cognitive Sciences.

[148]  D. Pandya,et al.  The cerebrocerebellar system. , 1997, International review of neurobiology.

[149]  A. Palmer,et al.  Histochemical identification of cortical areas in the auditory region of the human brain , 2002, Experimental Brain Research.

[150]  G Weismer,et al.  Interspeaker variation in habitual speaking rate: evidence for a neuromuscular component. , 1997, Journal of speech, language, and hearing research : JSLHR.

[151]  T. J. Grabowski Re-examining the Brain Regions Crucial for Orchestrating Speech Articulation , 2006 .

[152]  Stephen C. Strother,et al.  Mapping cerebral blood flow during speech production in hereditary ataxia , 2006, NeuroImage.

[153]  P. R. Davidson,et al.  Widespread access to predictive models in the motor system: a short review , 2005, Journal of neural engineering.

[154]  Wolfgang Grodd,et al.  Hemispheric Lateralization Effects of Rhythm Implementation during Syllable Repetitions: An fMRI Study , 2002, NeuroImage.

[155]  S. Grossberg,et al.  Cortical networks for control of voluntary arm movements under variable force conditions. , 1998, Cerebral cortex.

[156]  Benjamin Wallace,et al.  Generalization of prism adaptation. , 2006, Journal of experimental psychology. Human perception and performance.

[157]  C. Ludlow Central nervous system control of the laryngeal muscles in humans , 2005, Respiratory Physiology & Neurobiology.

[158]  David C Reutens,et al.  Functional topography of the low postcentral area. , 2002, Journal of neurosurgery.

[159]  Frank H Guenther,et al.  Effects of short- and long-term changes in auditory feedback on vowel and sibilant contrasts. , 2007, Journal of speech, language, and hearing research : JSLHR.

[160]  G. Bruyn Atlas of the Cerebral Sulci, M. Ono, S. Kubik, Chad D. Abernathey (Eds.). Georg Thieme Verlag, Stuttgart, New York (1990), 232, DM 298 , 1990 .

[161]  Satrajit S. Ghosh,et al.  Region of interest based analysis of functional imaging data , 2003, NeuroImage.

[162]  A. Murata,et al.  Largely segregated parietofrontal connections linking rostral intraparietal cortex (areas AIP and VIP) and the ventral premotor cortex (areas F5 and F4) , 1999, Experimental Brain Research.

[163]  H. Lane,et al.  Speech deterioration in postlingually deafened adults. , 1991, The Journal of the Acoustical Society of America.

[164]  R. A. Chase Effect of delayed auditory feedback on the repetition of speech sounds. , 1958, The Journal of speech and hearing disorders.

[165]  Cynthia G. Clopper,et al.  Acoustic characteristics of the vowel systems of six regional varieties of American English. , 2005, The Journal of the Acoustical Society of America.

[166]  R. Zatorre,et al.  Structure and function of auditory cortex: music and speech , 2002, Trends in Cognitive Sciences.

[167]  Christophe Habas,et al.  Cortical areas functionally linked with the cerebellar second homunculus during out-of-phase bimanual movements , 2005, Neuroradiology.

[168]  Y Yang,et al.  A silent event‐related functional MRI technique for brain activation studies without interference of scanner acoustic noise , 2000, Magnetic resonance in medicine.

[169]  E. T. Possing,et al.  Human temporal lobe activation by speech and nonspeech sounds. , 2000, Cerebral cortex.

[170]  Wolfgang Grodd,et al.  The cerebral control of speech tempo: Opposite relationship between speaking rate and BOLD signal changes at striatal and cerebellar structures , 2006, NeuroImage.

[171]  Pascal Perrier,et al.  A theory of speech motor control and supporting data from speakers with normal hearing and with profound hearing loss , 2000, J. Phonetics.

[172]  J. Rauschecker,et al.  A PET study of human auditory spatial processing , 1999, Neuroscience Letters.

[173]  U. Castiello,et al.  The Human Premotor Cortex Is 'Mirror' Only for Biological Actions , 2004, Current Biology.

[174]  T Allison,et al.  Localization of the face area of human sensorimotor cortex by intracranial recording of somatosensory evoked potentials. , 1993, Journal of neurosurgery.

[175]  W. Levelt,et al.  Do speakers have access to a mental syllabary? , 1994, Cognition.

[176]  J. Rauschecker,et al.  Mechanisms and streams for processing of "what" and "where" in auditory cortex. , 2000, Proceedings of the National Academy of Sciences of the United States of America.

[177]  Christoph E Schreiner,et al.  Order and disorder in auditory cortical maps , 1995, Current Opinion in Neurobiology.

[178]  Frank H. Guenther,et al.  A neural network model of speech acquisition and motor equivalent speech production , 2004, Biological Cybernetics.

[179]  S E Shaywitz,et al.  Neurobiological studies of reading and reading disability. , 2001, Journal of communication disorders.

[180]  D. Wolpert,et al.  Attenuation of Self-Generated Tactile Sensations Is Predictive, not Postdictive , 2006, PLoS biology.

[181]  W. Grodd,et al.  Cerebellum and Speech Perception: A Functional Magnetic Resonance Imaging Study , 2002, Journal of Cognitive Neuroscience.

[182]  Anthony R. McIntosh,et al.  Clustered functional MRI of overt speech production , 2006, NeuroImage.

[183]  F. Guenther,et al.  Overreliance on auditory feedback may lead to sound/syllable repetitions: simulations of stuttering and fluency-inducing conditions with a neural model of speech production. , 2010, Journal of fluency disorders.

[184]  P. Strick,et al.  Basal ganglia and cerebellar loops: motor and cognitive circuits , 2000, Brain Research Reviews.

[185]  R A Andersen,et al.  Multimodal integration for the representation of space in the posterior parietal cortex. , 1997, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[186]  Mark D. Humphries,et al.  A robot model of the basal ganglia: Behavior and intrinsic processing , 2006, Neural Networks.

[187]  F. Gonzalez-Lima,et al.  Structural equation modeling and its application to network analysis in functional brain imaging , 1994 .

[188]  K. Kalveram,et al.  Effects of frequency-shifted auditory feedback on fundamental frequency of long stressed and unstressed syllables. , 2001, Journal of speech, language, and hearing research : JSLHR.

[189]  R. Sperry Neural basis of the spontaneous optokinetic response produced by visual inversion. , 1950, Journal of comparative and physiological psychology.

[190]  Colin Humphries,et al.  Role of left posterior superior temporal gyrus in phonological processing for speech perception and production , 2001, Cogn. Sci..

[191]  J. Mink THE BASAL GANGLIA: FOCUSED SELECTION AND INHIBITION OF COMPETING MOTOR PROGRAMS , 1996, Progress in Neurobiology.

[192]  Alan C. Evans,et al.  Digital brain atlases , 1995, Trends in Neurosciences.

[193]  P. Strick,et al.  Cerebellar Loops with Motor Cortex and Prefrontal Cortex of a Nonhuman Primate , 2003, The Journal of Neuroscience.

[194]  G. Rizzolatti,et al.  Understanding motor events: a neurophysiological study , 2004, Experimental Brain Research.

[195]  Peter D. Neilson,et al.  Speech motor control and stuttering: A computational model of adaptive sensory-motor processing , 1987, Speech Commun..

[196]  J. Krakauer,et al.  Sensory prediction errors drive cerebellum-dependent adaptation of reaching. , 2007, Journal of neurophysiology.

[197]  S. Bouisset,et al.  [Voluntary movement]. , 1953, Journal de physiologie.

[198]  K. Sakai,et al.  Brain activations during conscious self‐monitoring of speech production with delayed auditory feedback: An fMRI study , 2003, Human brain mapping.

[199]  R. Turner,et al.  Characterizing Dynamic Brain Responses with fMRI: A Multivariate Approach , 1995, NeuroImage.

[200]  J. Voogd The human cerebellum , 2003, Journal of Chemical Neuroanatomy.

[201]  Kirrie J. Ballard,et al.  Understanding the nature of apraxia of speech: Theory, analysis, and treatment , 2000 .

[202]  L. Cohen,et al.  The role of the supplementary motor area (SMA) in word production , 2006, Brain Research.

[203]  C. Larson,et al.  Voice F0 responses to manipulations in pitch feedback. , 1998, The Journal of the Acoustical Society of America.

[204]  L S Illis,et al.  Neurotrauma , 1998, Spinal Cord.

[205]  Julien Doyon,et al.  Cerebellum and M1 interaction during early learning of timed motor sequences , 2005, NeuroImage.

[206]  R. Zatorre,et al.  Voice-selective areas in human auditory cortex , 2000, Nature.

[207]  G. Schaltenbrand,et al.  The effects of stereotactic electrical stimulation in the depth of the brain. , 1965, Brain : a journal of neurology.

[208]  L. Parsons,et al.  Location-Probability Profiles for the Mouth Region of Human Primary Motor–Sensory Cortex: Model and Validation , 2001, NeuroImage.

[209]  P. McGuire,et al.  An fMRI study of verbal self-monitoring: neural correlates of auditory verbal feedback. , 2006, Cerebral cortex.

[210]  S Jonas,et al.  The supplementary motor region and speech emission. , 1981, Journal of communication disorders.

[211]  D K Oller,et al.  The role of audition in infant babbling. , 1988, Child development.

[212]  André Brechmann,et al.  Sound-level-dependent representation of frequency modulations in human auditory cortex: a low-noise fMRI study. , 2002, Journal of neurophysiology.

[213]  A. Dale,et al.  Whole Brain Segmentation Automated Labeling of Neuroanatomical Structures in the Human Brain , 2002, Neuron.

[214]  Neural Bases of Speech, Hearing, and Language , 1990 .

[215]  S. Cobb Speech and Brain-Mechanisms. , 1960 .

[216]  Christopher K. Kovach,et al.  A functional connection between inferior frontal gyrus and orofacial motor cortex in human. , 2004, Journal of neurophysiology.

[217]  H. Fukuyama,et al.  Cortical processing mechanism for vocalization with auditory verbal feedback , 1997, Neuroreport.

[218]  Sylvain Houle,et al.  A positron emission tomography study of short- and long-term treatment effects on functional brain activation in adults who stutter. , 2003, Journal of fluency disorders.

[219]  M. Molinari,et al.  Verbal short-term store-rehearsal system and the cerebellum. Evidence from a patient with a right cerebellar lesion. , 1998, Brain : a journal of neurology.

[220]  W. Levelt,et al.  The spatial and temporal signatures of word production components , 2004, Cognition.

[221]  S. Nagarajan,et al.  Magnetoencephalographic evidence for a precise forward model in speech production , 2006, Neuroreport.

[222]  Bradley R. Buchsbaum,et al.  Reading, hearing, and the planum temporale , 2005, NeuroImage.

[223]  K. Cullen Sensory signals during active versus passive movement , 2004, Current Opinion in Neurobiology.

[224]  N. Geschwind Disconnexion syndromes in animals and man. I. , 1965, Brain : a journal of neurology.

[225]  W. T. Thach,et al.  Basal ganglia intrinsic circuits and their role in behavior , 1993, Current Opinion in Neurobiology.

[226]  F. Guenther,et al.  Neural mechanisms underlying sensory feedback control of speech , 2007 .

[227]  H. Saito,et al.  Mesial frontal lobe syndromes: correlations between neurological deficits and radiological localizations. , 1990, The Tohoku journal of experimental medicine.

[228]  K. Munhall,et al.  Compensation following real-time manipulation of formants in isolated vowels. , 2006, The Journal of the Acoustical Society of America.

[229]  T. H. Le,et al.  Functional MRI of human auditory cortex using block and event‐related designs , 2001, Magnetic resonance in medicine.

[230]  P. Fox Brain imaging in stuttering: where next? , 2003, Journal of fluency disorders.

[231]  R. Miall,et al.  Functional imaging of changes in cerebellar activity related to learning during a novel eye–hand tracking task , 2005, Experimental Brain Research.

[232]  F. Guenther,et al.  Modeling the representation of speech sounds in auditory cortical areas , 2003 .

[233]  A. Saykin,et al.  Functional MRI localisation of central nervous system regions associated with volitional inspiration in humans , 1999, The Journal of physiology.

[234]  A. Nakamura,et al.  Somatosensory Homunculus as Drawn by MEG , 1998, NeuroImage.

[235]  Philip Lieberman,et al.  Selective speech motor, syntax and cognitive deficits associated with bilateral damage to the putamen and the head of the caudate nucleus: a case study , 1998, Neuropsychologia.

[236]  H. Ackermann,et al.  Speech rate and rhythm in cerebellar dysarthria: an acoustic analysis of syllabic timing. , 1994, Folia phoniatrica et logopaedica : official organ of the International Association of Logopedics and Phoniatrics.

[237]  M. Merello,et al.  [Functional anatomy of the basal ganglia]. , 2000, Revista de neurologia.

[238]  J. Perkell,et al.  Sensorimotor adaptation to feedback perturbations of vowel acoustics and its relation to perception. , 2007, The Journal of the Acoustical Society of America.

[239]  D. Wolpert,et al.  Motor prediction , 2001, Current Biology.

[240]  Nikos Makris,et al.  Automatically parcellating the human cerebral cortex. , 2004, Cerebral cortex.

[241]  Jay J Bauer,et al.  Vocal responses to unanticipated perturbations in voice loudness feedback: an automatic mechanism for stabilizing voice amplitude. , 2006, The Journal of the Acoustical Society of America.

[242]  W. Grodd,et al.  Differential Contributions of Motor Cortex, Basal Ganglia, and Cerebellum to Speech Motor Control: Effects of Syllable Repetition Rate Evaluated by fMRI , 2001, NeuroImage.

[243]  M. Erb,et al.  fMRI Evaluation of Somatotopic Representation in Human Primary Motor Cortex , 2000, NeuroImage.

[244]  Xiaoqin Wang,et al.  Dynamics of auditory-vocal interaction in monkey auditory cortex. , 2005, Cerebral cortex.

[245]  R. Luján Fiber Pathways of the Brain, J.D. Schmahmann, D.N. Pandya (Eds.). Oxford University Press (2006), ISBN: 0-19-510423-4 , 2008 .

[246]  Iwona Stepniewska,et al.  Ipsilateral cortical connections of motor, premotor, frontal eye, and posterior parietal fields in a prosimian primate, Otolemur garnetti , 2005, The Journal of comparative neurology.

[247]  G. Ojemann Cortical organization of language , 1991, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[248]  J. Perkell,et al.  The distinctness of speakers' productions of vowel contrasts is related to their discrimination of the contrasts. , 2004, The Journal of the Acoustical Society of America.

[249]  Jefferson E. Roy,et al.  Dissociating Self-Generated from Passively Applied Head Motion: Neural Mechanisms in the Vestibular Nuclei , 2004, The Journal of Neuroscience.

[250]  A R McIntosh,et al.  Lateralization and behavioral correlation of changes in regional cerebral blood flow with classical conditioning of the human eyeblink response. , 1997, Journal of neurophysiology.

[251]  R. Turner,et al.  Dynamic magnetic resonance imaging of human brain activity during primary sensory stimulation. , 1992, Proceedings of the National Academy of Sciences of the United States of America.

[252]  C. Habas,et al.  Specific neocerebellar activation during out-of-phase bimanual movements , 2004, Neuroreport.

[253]  J. Kaas,et al.  Subdivisions of auditory cortex and processing streams in primates. , 2000, Proceedings of the National Academy of Sciences of the United States of America.

[254]  M. Kawato,et al.  Internal forward models in the cerebellum: fMRI study on grip force and load force coupling. , 2003, Progress in brain research.

[255]  Guy Marchal,et al.  Automated multi-modality image registration based on information theory , 1995 .

[256]  Sharlene D. Newman,et al.  Differences in auditory processing of words and pseudowords: An fMRI study , 2001, Human brain mapping.

[257]  Thomas M Donath,et al.  Effects of frequency-shifted auditory feedback on voice F0 contours in syllables. , 2002, The Journal of the Acoustical Society of America.

[259]  Satrajit S. Ghosh,et al.  Mindboggle: Automated brain labeling with multiple atlases , 2005, BMC Medical Imaging.

[260]  W Grodd,et al.  Opposite hemispheric lateralization effects during speaking and singing at motor cortex, insula and cerebellum , 2000, Neuroreport.

[261]  Alfonso Nieto-Castanon,et al.  A modeling investigation of articulatory variability and acoustic stability during American English /r/ production. , 2005, The Journal of the Acoustical Society of America.

[262]  F. Guenther,et al.  A theoretical investigation of reference frames for the planning of speech movements. , 1998, Psychological review.

[263]  Kevin G Munhall,et al.  Adaptive control of vowel formant frequency: evidence from real-time formant manipulation. , 2006, The Journal of the Acoustical Society of America.

[264]  J. Fiez,et al.  A Comment on the Functional Localization of the Phonological Storage Subsystem of Working Memory , 1999, Brain and Cognition.

[265]  W. Zemlin Speech and Hearing Science : Anatomy and Physiology , 1968 .

[266]  Alan C. Evans,et al.  Lateralization of phonetic and pitch discrimination in speech processing. , 1992, Science.

[267]  S. Sapir,et al.  Some effects of auditory stimulation on perioral motor unit discharge and their implications for speech production. , 1981, The Journal of the Acoustical Society of America.

[268]  G. Hickok,et al.  Auditory–Motor Interaction Revealed by fMRI: Speech, Music, and Working Memory in Area Spt , 2003 .

[269]  N Makris,et al.  Analysis of lesions by MRI in stroke patients with acoustic‐phonetic processing deficits , 1995, Neurology.

[270]  D. Wolpert,et al.  Central cancellation of self-produced tickle sensation , 1998, Nature Neuroscience.

[271]  G. Rizzolatti,et al.  Functional organization of inferior area 6 in the macaque monkey , 1988, Experimental Brain Research.

[272]  P. Skudlarski,et al.  Event-related fMRI of auditory and visual oddball tasks. , 2000, Magnetic resonance imaging.

[273]  S. Scott,et al.  The neuroanatomical and functional organization of speech perception , 2003, Trends in Neurosciences.

[274]  W PENFIELD,et al.  The supplementary motor area of the cerebral cortex; a clinical and experimental study. , 1951, A.M.A. archives of neurology and psychiatry.

[275]  A Olivier,et al.  Sensory cortical tongue representation in man. , 1983, Journal of neurosurgery.

[276]  Scott T. Grafton,et al.  Forward modeling allows feedback control for fast reaching movements , 2000, Trends in Cognitive Sciences.

[277]  H. Lane,et al.  The Lombard Sign and the Role of Hearing in Speech , 1971 .

[278]  Frank H. Guenther,et al.  Unstable or Insufficiently Activated Internal Models and Feedback-Biased Motor Control as Sources of Dysfluency: A Theoretical Model of Stuttering , 2004 .

[279]  M. Mauk,et al.  What the cerebellum computes , 2003, Trends in Neurosciences.

[280]  J. Ford,et al.  Fine-tuning of auditory cortex during speech production. , 2005, Psychophysiology.

[281]  Satrajit S. Ghosh,et al.  Neural modeling and imaging of the cortical interactions underlying syllable production , 2006, Brain and Language.

[282]  Kevin R. Smith,et al.  Word length modulates neural activity in auditory cortex during covert object naming , 2003, Neuroreport.

[283]  P. Tallal,et al.  Neurobiological Basis of Speech: A Case for the Preeminence of Temporal Processing , 1993, Annals of the New York Academy of Sciences.

[284]  Karl J. Friston,et al.  Role of the human rostral supplementary motor area and the basal ganglia in motor sequence control: investigations with H2 15O PET. , 1998, Journal of neurophysiology.

[285]  Joseph S Perkell,et al.  Effects of masking noise on vowel and sibilant contrasts in normal-hearing speakers and postlingually deafened cochlear implant users. , 2005, The Journal of the Acoustical Society of America.

[286]  Emily B. Myers,et al.  An event-related fMRI investigation of phonological–lexical competition , 2006, Neuropsychologia.

[287]  Wolfram Ziegler,et al.  The role of the left mesial frontal cortex in fluent speech: Evidence from a case of left supplementary motor area hemorrhage , 1997, Neuropsychologia.

[288]  C. Price,et al.  The Constraints Functional Neuroimaging Places on Classical Models of Auditory Word Processing , 2001, Journal of Cognitive Neuroscience.

[289]  T. Ebner,et al.  Functional magnetic resonance imaging of cerebellar activation during the learning of a visuomotor dissociation task , 1996, Human brain mapping.

[290]  D M Wolpert,et al.  Multiple paired forward and inverse models for motor control , 1998, Neural Networks.

[291]  N. Dronkers A new brain region for coordinating speech articulation , 1996, Nature.

[292]  W. Grodd,et al.  Articulatory/Phonetic Sequencing at the Level of the Anterior Perisylvian Cortex: A Functional Magnetic Resonance Imaging (fMRI) Study , 2000, Brain and Language.

[293]  Vincent A Magnotta,et al.  Cerebral cortex: a topographic segmentation method using magnetic resonance imaging , 2000, Psychiatry Research: Neuroimaging.

[294]  Karl J. Friston,et al.  Analysis of fMRI Time-Series Revisited , 1995, NeuroImage.

[295]  P. Redgrave,et al.  The basal ganglia: a vertebrate solution to the selection problem? , 1999, Neuroscience.

[296]  Richard S. J. Frackowiak,et al.  Human cerebral activity with increasing inspiratory force: a study using positron emission tomography. , 1996, Journal of applied physiology.

[297]  Frank H. Guenther,et al.  An fMRI investigation of syllable sequence production , 2006, NeuroImage.

[298]  Gottfried Schlaug,et al.  Shared and distinct neural correlates of singing and speaking , 2006, NeuroImage.

[299]  Guy Marchal,et al.  Automated multi-moda lity image registration based on information theory , 1995 .

[300]  Satrajit S. Ghosh,et al.  Representation of sound categories in auditory cortical maps. , 2004, Journal of speech, language, and hearing research : JSLHR.