The DIVA model: A neural theory of speech acquisition and production

“Similarly, comparisons of auditory responses during self-produced speech and while listening to recordings of one’s own speech indicate attenuation of auditory cortex responses during speech production (Curio, Neuloh, Numminen, Jousmaki, & Hari, 2000; Heinks-Maldonado, Mathalon, Gray, & Ford, 2005;…”

“1 Performance of these motor programs relies on contributions from bilateral premotor, motor, and subcortical regions (see below), areas that are active bilaterally during normal speech production.”

“Blood oxygenation level-dependent (BOLD) functional magnetic resonance imaging (fMRI; Belliveau et al., 1992; Kwong et al., 1992; Ogawa et al.,”

“Similarly, comparisons of auditory responses during self-produced speech and while listening to recordings of one’s own speech indicate attenuation of auditory cortex responses during speech production (Curio, Neuloh, Numminen, Jousmaki, & Hari, 2000; Heinks-Maldonado, Mathalon, Gray, & Ford, 2005;…”

“1 Performance of these motor programs relies on contributions from bilateral premotor, motor, and subcortical regions (see below), areas that are active bilaterally during normal speech production.”

“This mapping from desired sensory outcome to the appropriate motor action is an inverse kinematic transformation and is often referred to as an inverse model (e.g., Kawato, 1999; Wolpert & Kawato, 1998).”

“Evidence of mirror neurons in humans has implicated left precentral gyrus for grasping actions (Tai, Scherfler, Brooks, Sawamoto, & Castiello, 2004), and left opercular inferior frontal gyrus for finger movements (Iacoboni et al., 1999).”

“Von Holst and Mittelstaedt (1950) proposed the ‘principle of reafference’ in which a copy of the expected sensory consequences of a motor command, termed an efference copy, was subtracted from the realised sensory consequences.”

“For this purpose, the proposed locations of model components have been mapped to anatomical landmarks of the canonical brain provided with the SPM image analysis software package (Friston et al, 1995 ; <ext-link ext-link-type="uri" xmlns:xlink="http://www.”

“Mapping the model’s components onto this reference, then, provides a convenient means to compare the results of a large pool of neuroimaging experiments from a common theoretical framework, a framework that accounts for a wide range of data from diverse experimental modalities.”

“The goal of our research programme over the past 16 years has been to improve our understanding of the neural mechanisms that underlie speech motor control.”

“The somatosensory target and state maps lie in ventral supramarginal gyrus, a region Hickok and colleagues (e.g., Hickok & Poeppel, 2004) have argued supports the integration of speech motor commands and sensory feedback.”

“This mapping from desired sensory outcome to the appropriate motor action is an inverse kinematic transformation and is often referred to as an inverse model (e.g., Kawato, 1999; Wolpert & Kawato, 1998).”

“# 2010 Psychology Press, an imprint of the Taylor & Francis Group, an Informa business http://www.psypress.com/lcp DOI: 10.1080/01690960903498424 Indefrey & Levelt, 2004; Turkeltaub, Eden, Jones, & Zeffiro, 2002).”

“1 Performance of these motor programs relies on contributions from bilateral premotor, motor, and subcortical regions (see below), areas that are active bilaterally during normal speech production.”

“Predictions from the model have guided studies of the role of auditory feedback in normally hearing persons, deaf persons, and persons who have recently regained some hearing through the use of cochlear implants (Lane et al., 2007a; Perkell et al., 2000, 2004a, 2004b, 2007).”

“The DIVA model: A neural theory of speech acquisition and production”

“Local field potentials are thought to reflect local synaptic activity, i.e., a weighted sum of the inputs to a given region (Raichle & Mintun, 2006).”

“Projections such as these, which predict the sensorimotor state resulting from a movement, are typically described as representing a forward model of the movement (e.g., Davidson & Wolpert, 2005; Desmurget & Grafton, 2000; Kawato, 1999; Miall & Wolpert, 1996).”

“Browman & Goldstein, 1989 ) for that speech sound.”

“Broca’s early findings linking left inferior frontal damage with speech production deficits have been corroborated by a large body of lesion studies (Dronkers, 1996; Duffy, 2005; Hillis et al., 2004; Kent & Tjaden, 1997).”

“…velocity cells is analogous to the process of ‘phonetic encoding’ as conceptualised by Levelt and colleagues (e.g., Levelt, Roelofs, & Meyer, 1999; Levelt & Wheeldon, 1994); i.e., it transforms a phonological input from adjacent inferior frontal cortex into the set of feedforward motor…”

“Reciprocal pathways between inferior frontal and auditory and somatosensory cortex have been demonstrated in humans (Makris et al., 2005; Matsumoto et al., 2004) and non-human primates (Morel & Kaas, 1992; Ojemann, 1991; Romanski et al.”

“A dominant role for the left hemisphere has been a hallmark of language processing models for several decades (Geschwind, 1970).”

“…cortex have been demonstrated in humans (Makris et al., 2005; Matsumoto et al., 2004) and non-human primates (Morel & Kaas, 1992; Ojemann, 1991; Romanski et al., 1999; Schmahmann & Pandya, 2006); also see Duffau (2008) for a description of the putative fronto-parietal and frontotemporal…”

“Neuroimaging studies of motor learning have noted cerebellar activity that is associated with the size or frequency of sensory error (e.g., Blakemore, Frith, & Wolpert, 2001; Blakemore, Wolpert, & Frith, 1999; Diedrichsen, Hashambhoy, Rane, & Shadmehr, 2005; Flament, Ellermann, Kim, Ugurbil, & Ebner, 1996; Grafton et…”

“Model components were mapped to locations in the Montreal Neurological Institute (MNI; Mazziotta et al., 2001) standard reference frame based on relevant neuroanatomical and neurophysiological studies (Guenther et al., 2006).”

“Maeda, S. (1990). Compensatory articulation during speech: Evidence from the analysis and synthesis of vocal tract shapes using an articulatory model.”

“…inverse kinematics (e.g., Kawato, 1999; Wolpert & Kawato, 1998), a role for which it is anatomically well-suited; the cerebellum receives inputs from higher-order auditory and somatosensory areas (e.g., Schmahmann & Pandya, 1997), and projects heavily to the motor cortex (Middleton & Strick, 1997).”

“Support for these modifications and the impact they have on the model are discussed below.”

“…past decade and a half of imaging research has provided a tremendous amount of functional data regarding the brain regions involved in speech (e.g., Fiez & Petersen, 1998; Ghosh, Bohland, & Guenther, 2003; Indefrey & Levelt, 2004; Riecker et al., 2000b; Soros, Sokoloff, Bose, McIntosh, Graham, &…”

“The SMA is strongly interconnected with lateral motor and premotor cortex and the basal ganglia (Jurgens, 1984; Lehericy et al., 2004; Luppino, Matelli, Camarda, & Rizzolatti, 1993; Matsumoto et al., 2004, 2007).”

“Once trained, the model takes a speech sound as input, and generates a time varying sequence of articulator positions that command movements of the simulated vocal tract that produce the desired sound.”

“…1996; Ghacibeh & Heilman, 2003; Kotz et al., 2003; Mitchell, Elliott, Barry, Cruttenden, & Woodruff, 2003; Pihan, Altenmuller, & Ackermann, 1997; Ross & Mesulam, 1979; Williamson, Harrison, Shenal, Rhodes, & Demaree, 2003), though the view of affective prosody as a unitary, purely right…”

“It has been argued by many that the cerebellum uses sensory error to build forward models that generate sensory predictions (Blakemore et al., 2001; Imamizu et al., 2000; Kawato et al., 2003; O’Reilly, Mesulam, & Nobre, 2008), the role of projections from the speech sound map to sensory target maps…”

“…Arieli, Hasson, Fried, & Malach, 2005), consensus is building around the hypotheses that BOLD signal is correlated with local field potentials (Goense & Logothetis, 2008; Logothetis, Pauls, Augath, Trinath, & Oeltermann, 2001; Mathiesen, Caesar, Akgoren, & Lauritzen, 1998; Viswanathan &…”

“The synaptic weights are learned during a babbling phase meant to coarsely represent that typically experienced by a normally developing infant (e.g., Oller & Eilers, 1988).”

“The mapping from a cell in the speech sound map to the articulator velocity cells is analogous to the process of ‘phonetic encoding’ as conceptualised by Levelt and colleagues (eg, Levelt, Roelofs, & Meyer, 1999; Levelt & Wheeldon, 1994 ); ie, it transforms a phonological input from adjacent inferior frontal cortex in…”

“These data and recent imaging findings have led a number of investigators to conclude that the SMA plays a critical role in controlling the initiation of speech motor commands (e.g., Alario, Chainay, Lehericy, & Cohen, 2006; Bohland & Guenther, 2006; Jonas, 1987; Ziegler et al., 1997).”

“Cells in the articulator position maps are hypothesised to correspond to ‘tonic’ neurons that have been identified in monkey primary motor cortex (e.g., Kalaska et al., 1989).”

“Wise and colleagues, using positron emission tomography (PET) to indirectly assess neural activity, noted reduced superior temporal gyrus activation during speech production compared with a listening task (Wise, Greene, Buchel, & Scott, 1999 ).”

“This hypothesised role for ventral parietal cortex during speech production is analogous to the visual-motor integration role associated with more dorsal parietal regions during limb movements (Andersen, 1997 ; Rizzolatti, Fogassi, & Gallese, 1997 ).”

“…in the speech sound map to the articulator velocity cells is analogous to the process of ‘phonetic encoding’ as conceptualised by Levelt and colleagues (e.g., Levelt, Roelofs, & Meyer, 1999; Levelt & Wheeldon, 1994); i.e., it transforms a phonological input from adjacent inferior frontal cortex…”

“Broca’s early findings linking left inferior frontal damage with speech production deficits have been corroborated by a large body of lesion studies (Dronkers, 1996; Duffy, 2005; Hillis et al., 2004; Kent & Tjaden, 1997).”

“Thus, as DIVA has come to account for a wide range of speech production phenomena (e.g., Callan et al., 2000; Guenther, 1994, 1995; Guenther et al., 1998; Nieto-Castanon et al., 2005), it does so from a unified quantitative and neurobiologically grounded framework.”

“Experiments are designed to test these predictions, and the empirical findings are, in turn, used to further refine the model.”

“Neuroimaging studies of speech production in persons who stutter consistently demonstrate increased right hemisphere activation relative to normal speakers in the precentral and inferior frontal gyrus regions (see Brown, Ingham, Ingham, Laird, & Fox, 2005 for review), the same frontal regions identified as part of the…”

“…of affective prosody, which appear to rely more on the right cerebral hemisphere than the left hemisphere (Adolphs, Damasio, & Tranel, 2002; Buchanan et al., 2000; George et al., 1996; Ghacibeh & Heilman, 2003; Kotz et al., 2003; Mitchell, Elliott, Barry, Cruttenden, & Woodruff, 2003;…”

“Based on these findings, two sets of cells representing laryngeal parameters of the Maeda articulator model (Maeda, 1990) associated with laryngeal functions have been assigned to MNI locations: cells in ventrolateral precentral gyrus (labelled Larynx, Extrinsic in Table 1) represent larynx height, whereas cells in th…”

“…carryover coarticulation, velocity/ distance relationships, speaking rate effects, and speaking skill acquisition and retention throughout development (e.g., Callan, Kent, Guenther, & Vorperian, 2000; Guenther, 1994, 1995; Guenther et al., 2006; Guenther et al., 1998; Nieto-Castanon et al., 2005).”

“Modulation of pitch, loudness, duration, and rhythm convey meaningful linguistic and affective cues (Bolinger, 1961, 1989; Lehiste, 1970, 1976; Netsell, 1973; Shriberg & Kent, 1982).”

“…Hasson, Fried, & Malach, 2005), consensus is building around the hypotheses that BOLD signal is correlated with local field potentials (Goense & Logothetis, 2008; Logothetis, Pauls, Augath, Trinath, & Oeltermann, 2001; Mathiesen, Caesar, Akgoren, & Lauritzen, 1998; Viswanathan & Freeman, 2007).”

“…speculated that a representation of sensory errors in the cerebellum drives corrective motor commands (Grafton et al., 2008; Penhune & Doyon, 2005) and contributes to feedback-based motor learning (Ito, 2000; Tseng, Diedrichsen, Krakauer, Shadmehr, & Bastian, 2007; Wolpert, Miall, & Kawato, 1998).”

“These data and recent imaging findings have led a number of investigators to conclude that the SMA plays a critical role in controlling the initiation of speech motor commands (eg, Alario, Chainay, Lehericy, & Cohen, 2006 ; Bohland & Guenther, 2006 ; Jonas, 1987 ; Ziegler et al, 1997 ).”

“The SMA is reciprocally connected with the basal ganglia, another region widely believed to contribute to gating motor commands (eg, Albin, Young, & Penney, 1995 ; Pickett, Kuniholm, Protopapas, Friedman, & Lieberman, 1998 ; Van Buren, 1963 ).”

“A wide body of evidence suggests such a mechanism plays an important role in the motor control of eye and hand movements, as well as speech (e.g., Bays, Flanagan, & Wolpert, 2006; Cullen, 2004; Heinks-Maldonado & Houde, 2005; Reppas, Usrey, & Reid, 2002; Roy & Cullen, 2004; Voss, Ingram, Haggard, &…”

“…in humans (Makris et al., 2005; Matsumoto et al., 2004) and non-human primates (Morel & Kaas, 1992; Ojemann, 1991; Romanski et al., 1999; Schmahmann & Pandya, 2006); also see Duffau (2008) for a description of the putative fronto-parietal and frontotemporal pathways involved in language processing.”

“Evidence of mirror neurons in humans has implicated left precentral gyrus for grasping actions (Tai, Scherfler, Brooks, Sawamoto, & Castiello, 2004 ), and left opercular inferior frontal gyrus for finger movements (Iacoboni et al, 1999 ).”

“The SMA is strongly interconnected with lateral motor and premotor cortex and the basal ganglia (Jurgens, 1984; Lehericy et al., 2004; Luppino, Matelli, Camarda, & Rizzolatti, 1993; Matsumoto et al., 2004, 2007).”

“The SMA is strongly interconnected with lateral motor and premotor cortex and the basal ganglia (Jurgens, 1984; Lehericy et al., 2004; Luppino, Matelli, Camarda, & Rizzolatti, 1993; Matsumoto et al., 2004, 2007).”

“…data regarding the brain regions involved in speech (e.g., Fiez & Petersen, 1998; Ghosh, Bohland, & Guenther, 2003; Indefrey & Levelt, 2004; Riecker et al., 2000b; Soros, Sokoloff, Bose, McIntosh, Graham, & Stuss, 2006; Turkeltaub et al., 2002), including the identification of a ‘minimal…”

“The SMA is strongly interconnected with lateral motor and premotor cortex and the basal ganglia (Jurgens, 1984; Lehericy et al., 2004; Luppino, Matelli, Camarda, & Rizzolatti, 1993; Matsumoto et al., 2004, 2007).”

“The map consists of 10 pairs of antagonistic cells2 that correspond to parameters of the Maeda vocal tract that determine lip protrusion, upper and lower lip height, jaw height, tongue height, tongue shape, tongue body position, tongue tip location, larynx height, and glottal opening and pressure.”

“It has been hypothesised that stuttering involves excessive reliance upon auditory feedback control due to poor feedforward commands (Max et al., 2004).”

“Support for these modifications and the impact they have on the model are discussed below.”

“The model has proven to be a valuable tool for studying the mechanisms underlying normal (Callan et al, 2000 ; Lane et al, 2007b ; Perkell et al, 2000 , 2004a, 2004b, 2007 ; Villacorta, Perkell, & Guenther, 2007 ) and disordered speech (Max et al, 2004 ; Robin et al, 2008 ; Terband et al, 2008 ).”

“Based on recent imaging work (Brown, Ngan, & Liotti, 2008 ; Olthoff, Baudewig, Kruse, & Dechent, 2008 ), additional motor cortex cells have been added to the model that represent the intrinsic laryngeal muscles.”

“This area has been proposed to correspond to the caudal portion of ventral inferior frontal gyrus (Brodmann’s area 44) in the human (see Binkofski & Buccino, 2004; Rizzolatti & Arbib, 1998).”

“One of the more consistent findings in the literature concerns perception and production of affective prosody, which appear to rely more on the right cerebral hemisphere than the left hemisphere (Adolphs, Damasio, & Tranel, 2002; Buchanan et al., 2000; George et al., 1996; Ghacibeh & Heilman, 2003; Kotz et al., 2003;…”

“It implies that damage to the left inferior frontal cortex is more commonly associated with speech disruptions than damage to the same region in the right hemisphere because feedforward motor programs are disrupted.”

“Projections such as these, which predict the sensorimotor state resulting from a movement, are typically described as representing a forward model of the movement (e.g., Davidson & Wolpert, 2005; Desmurget & Grafton, 2000; Kawato, 1999; Miall & Wolpert, 1996).”

“Thus, as DIVA has come to account for a wide range of speech production phenomena (e.g., Callan et al., 2000; Guenther, 1994, 1995; Guenther et al., 1998; Nieto-Castanon et al., 2005), it does so from a unified quantitative and neurobiologically grounded framework.”

“The model has also been employed in investigations of the etiology of stuttering (Max, Guenther, Gracco, Ghosh, & Wallace, 2004), and acquired apraxia of speech (Robin et al., 2008; Terband, Maassen, Brumberg, & Guenther, 2008).”

“However, rather than being limited to a repository of motor programs for frequently produced syllables, as Levelt proposed, the speech sound map also represents common syllabic, sub-syllabic (phonemes), and multi-syllabic speech sounds (e.g., words, phrases).”

“It is widely held that the cerebellum is involved with the learning and maintenance of feedforward motor commands (though see Grafton et al, 2008 ; Kawato, 1999 ; Ohyama, Nores, Murphy, & Mauk, 2003 ).”

“…suggests such a mechanism plays an important role in the motor control of eye and hand movements, as well as speech (e.g., Bays, Flanagan, & Wolpert, 2006; Cullen, 2004; Heinks-Maldonado & Houde, 2005; Reppas, Usrey, & Reid, 2002; Roy & Cullen, 2004; Voss, Ingram, Haggard, & Wolpert, 2006).”

“…frontal and auditory and somatosensory cortex have been demonstrated in humans (Makris et al., 2005; Matsumoto et al., 2004) and non-human primates (Morel & Kaas, 1992; Ojemann, 1991; Romanski et al., 1999; Schmahmann & Pandya, 2006); also see Duffau (2008) for a description of the putative…”

“Studies have begun to explore putative sensory expectation projections from lateral frontal cortex to auditory cortex, establishing an inhibitory effect linked to ongoing articulator movements (eg, Heinks-Maldonado et al, 2006 ).”

“Additional activity is typically found in adjacent lateral cortex bilaterally (Bohland & Guenther, 2006 ; Ghosh et al, 2008 ; Riecker, Ackermann, Wildgruber, Dogil, & Grodd, 2000; Riecker, Wildgruber, Dogil, Grodd, & Ackermann, 2002 ; Tourville et al, 2008 ; Wildgruber, Ackermann, & Grodd, 2001 ), an area less commonl…”

“This work bridges the gap between computational models of the linguistic/phonological level of speech production (Dell, 1986; Hartley & Houghton, 1996; Levelt et al., 1999), and DIVA, which addresses production at the speech motor control level.”

“Predictions from the model have guided studies of the role of auditory feedback in normally hearing persons, deaf persons, and persons who have recently regained some hearing through the use of cochlear implants (Lane et al., 2007a; Perkell et al., 2000, 2004a, 2004b, 2007).”

“The model has also been employed in investigations of the etiology of stuttering (Max, Guenther, Gracco, Ghosh, & Wallace, 2004), and acquired apraxia of speech (Robin et al., 2008; Terband, Maassen, Brumberg, & Guenther, 2008).”

“, & Kawato, M. (2007). Reorganization of brain activity for”

“…a laryngeal representation in motor cortex has typically been associated with the ventrolateral extreme of the precentral gyrus (e.g., Duffy, 2005; Ludlow, 2005), an assumption based heavily upon findings from non-human primates (e.g. Simonyan & Jurgens, 2003) and supported by the intracortical…”

“Lesions to anterior vermal and paravermal cerebellum have been associated with disruptions of speech production (Ackermann, Vogel, Petersen, & Poremba, 1992 ; Urban et al, 2003 ), termed ataxic dysarthria, characterised by an impaired ability to produce speech with fluent timing and gestural coordination.”

“Predictions from the model have guided studies of the role of auditory feedback in normally hearing persons, deaf persons, and persons who have recently regained some hearing through the use of cochlear implants (Lane et al., 2007a; Perkell et al., 2000, 2004a, 2004b, 2007).”

“The model has been used to guide a number of behavioural and functional imaging studies of speech processing (e.g., Bohland & Guenther, 2006; Ghosh, Tourville & Guenther, 2008; Guenther et al., 1999; Lane et al., 2005, 2007a, 2007b; Nieto-Castanon, Guenther, Perkell, & Curtin, 2005; Perkell et al., 2004a, 2004b; Tourv…”

“Modulation of pitch, loudness, duration, and rhythm convey meaningful linguistic and affective cues (Bolinger, 1961, 1989; Lehiste, 1970, 1976; Netsell, 1973; Shriberg & Kent, 1982).”

“…of auditory response suppression during self-initiated vocalisations is provided by single unit recordings from non-human primates; for example, attenuation of auditory cortical responses prior to self-initiated vocalisations has been demonstrated in the marmoset (Eliades & Wang, 2003, 2005).”

“…Wolpert, 2001; Blakemore, Wolpert, & Frith, 1999; Diedrichsen, Hashambhoy, Rane, & Shadmehr, 2005; Flament, Ellermann, Kim, Ugurbil, & Ebner, 1996; Grafton et al., 2008; Imamizu et al., 2000; Imamizu, Higuchi, Toda, & Kawato, 2007; Miall & Jenkinson, 2005; Schreurs et al., 1997; Tesche & Karhu,…”

“This hypothesis is consistent with the effects of fluency-inducing therapy on BOLD responses; successful treatment has been associated with a shift toward more normal, left-lateralised frontal activation (De Nil, Kroll, Lafaille, & Houle, 2003 ; Neumann et al, 2005 ).”

“The model has proven to be a valuable tool for studying the mechanisms underlying normal (Callan et al., 2000; Lane et al., 2007b; Perkell et al., 2000, 2004a, 2004b, 2007; Villacorta, Perkell, & Guenther, 2007) and disordered speech (Max et al., 2004; Robin et al., 2008; Terband et al., 2008).”

“…debate over which and to what extent prosodic feature control is lateralised (e.g., Doherty, West, Dilley, Shattuck-Hufnagel, & Caplan, 2004; Emmorey, 1987; Gandour et al., 2003; Meyer, Alter, Friederici, Lohmann, & von Cramon, 2002; Stiller et al., 1997; Walker, Pelletier, & Reif, 2004).”

“This hypothesis is consistent with the effects of fluency-inducing therapy on BOLD responses; successful treatment has been associated with a shift toward more normal, leftlateralised frontal activation (De Nil, Kroll, Lafaille, & Houle, 2003; Neumann et al., 2005).”

“It has been speculated that a representation of sensory errors in the cerebellum drives corrective motor commands (Grafton et al., 2008; Penhune & Doyon, 2005) and contributes to feedback-based motor learning (Ito, 2000; Tseng, Diedrichsen, Krakauer, Shadmehr, & Bastian, 2007; Wolpert, Miall, &…”

“Neuroimaging studies of motor learning have noted cerebellar activity that is associated with the size or frequency of sensory error (e.g., Blakemore, Frith, & Wolpert, 2001; Blakemore, Wolpert, & Frith, 1999; Diedrichsen, Hashambhoy, Rane, & Shadmehr, 2005; Flament, Ellermann, Kim, Ugurbil, & Ebner, 1996; Grafton et…”

“…debate over which and to what extent prosodic feature control is lateralised (e.g., Doherty, West, Dilley, Shattuck-Hufnagel, & Caplan, 2004; Emmorey, 1987; Gandour et al., 2003; Meyer, Alter, Friederici, Lohmann, & von Cramon, 2002; Stiller et al., 1997; Walker, Pelletier, & Reif, 2004).”

“Our findings corroborate characterisations of AOS as a disruption of the use and development of speech motor programs (Ballard, Granier, & Robin, 2000 ; McNeil, Robin, & Schmidt, 2007 ) and suggest that rehabilitative treatments focused on restoring motor programs, eg, sound production treatment (Wambaugh, Duffy, McNe…”

“Blood oxygenation level-dependent (BOLD) functional magnetic resonance imaging (fMRI; Belliveau et al., 1992; Kwong et al., 1992; Ogawa et al.,”

“The SMA is reciprocally connected with the basal ganglia, another region widely believed to contribute to gating motor commands (eg, Albin, Young, & Penney, 1995 ; Pickett, Kuniholm, Protopapas, Friedman, & Lieberman, 1998 ; Van Buren, 1963 ).”

“…to anterior vermal and paravermal cerebellum have been associated with disruptions of speech production (Ackermann, Vogel, Petersen, & Poremba, 1992; Urban et al., 2003), termed ataxic dysarthria, characterised by an impaired ability to produce speech with fluent timing and gestural coordination.”

“…extreme of the precentral gyrus (e.g., Duffy, 2005; Ludlow, 2005), an assumption based heavily upon findings from non-human primates (e.g. Simonyan & Jurgens, 2003) and supported by the intracortical mapping studies performed by Penfield and colleagues in humans prior to epilepsy surgery…”

“…& Tanji, 2000), intervals between specific movements within in a sequence (Shima & Tanji, 2000), the ordinal position of movements within a sequence (Clower & Alexander, 1998), and the number of items remaining in a sequence (Sohn & Lee, 2007) have been noted in recordings of neurons in the SMA.”

“These data and recent imaging findings have led a number of investigators to conclude that the SMA plays a critical role in controlling the initiation of speech motor commands (e.g., Alario, Chainay, Lehericy, & Cohen, 2006; Bohland & Guenther, 2006; Jonas, 1987; Ziegler et al., 1997).”

“Additional activity is typically found in adjacent lateral cortex bilaterally (Bohland & Guenther, 2006 ; Ghosh et al, 2008 ; Riecker, Ackermann, Wildgruber, Dogil, & Grodd, 2000; Riecker, Wildgruber, Dogil, Grodd, & Ackermann, 2002 ; Tourville et al, 2008 ; Wildgruber, Ackermann, & Grodd, 2001 ), an area less commonl…”

“…data regarding the brain regions involved in speech (e.g., Fiez & Petersen, 1998; Ghosh, Bohland, & Guenther, 2003; Indefrey & Levelt, 2004; Riecker et al., 2000b; Soros, Sokoloff, Bose, McIntosh, Graham, & Stuss, 2006; Turkeltaub et al., 2002), including the identification of a ‘minimal…”

“Other imaging studies of speech production that have included a perturbed auditory feedback condition have demonstrated greater right hemisphere involvement in auditory feedback-based control of speech (eg, Fu et al, 2006 ; Toyomura et al, 2007 ).”

“E-mail: guenther@cns.bu.edu Supported by the National Institute on Deafness and other Communication Disorders (R01 DC02852, F. Guenther PI).”

“The relative weight of feedforward and feedback commands in the overall motor command is dependent upon the size of the error signal since this determines the size of the feedback control contribution.”

“Ipsilateral and contralateral premotor-to-motor projections that would underlie this hypothesised connectivity have been demonstrated in monkeys (e.g., Dancause et al., 2006, 2007; Fang, Stepniewska, & Kaas, 2005; Stepniewska, Preuss, & Kaas, 2006).”

“Ipsilateral and contralateral premotor-to-motor projections that would underlie this hypothesised connectivity have been demonstrated in monkeys (eg, Dancause et al, 2006 , 2007 ; Fang, Stepniewska, & Kaas, 2005 ; Stepniewska, Preuss, & Kaas, 2006 ).”

“The SMA is reciprocally connected with the basal ganglia, another region widely believed to contribute to gating motor commands (e.g., Albin, Young, & Penney, 1995; Pickett, Kuniholm, Protopapas, Friedman, & Lieberman, 1998; Van Buren, 1963).”

“…suggests such a mechanism plays an important role in the motor control of eye and hand movements, as well as speech (e.g., Bays, Flanagan, & Wolpert, 2006; Cullen, 2004; Heinks-Maldonado & Houde, 2005; Reppas, Usrey, & Reid, 2002; Roy & Cullen, 2004; Voss, Ingram, Haggard, & Wolpert, 2006).”

“…2001; Blakemore, Wolpert, & Frith, 1999; Diedrichsen, Hashambhoy, Rane, & Shadmehr, 2005; Flament, Ellermann, Kim, Ugurbil, & Ebner, 1996; Grafton et al., 2008; Imamizu et al., 2000; Imamizu, Higuchi, Toda, & Kawato, 2007; Miall & Jenkinson, 2005; Schreurs et al., 1997; Tesche & Karhu, 2000).”

“…& Tanji, 2000), intervals between specific movements within in a sequence (Shima & Tanji, 2000), the ordinal position of movements within a sequence (Clower & Alexander, 1998), and the number of items remaining in a sequence (Sohn & Lee, 2007) have been noted in recordings of neurons in the SMA.”

“…carryover coarticulation, velocity/ distance relationships, speaking rate effects, and speaking skill acquisition and retention throughout development (e.g., Callan, Kent, Guenther, & Vorperian, 2000; Guenther, 1994, 1995; Guenther et al., 2006; Guenther et al., 1998; Nieto-Castanon et al., 2005).”

“Ipsilateral and contralateral premotor-to-motor projections that would underlie this hypothesised connectivity have been demonstrated in monkeys (e.g., Dancause et al., 2006, 2007; Fang, Stepniewska, & Kaas, 2005; Stepniewska, Preuss, & Kaas, 2006).”

“Brown et al. (2008) and Olthoff et al. (2008) have since demonstrated a bilateral representation in a more dorsal region of motor cortex adjacent to the lip area and near a second ‘vocalisation’ region identified by Penfield and Roberts (1959, p. 200).”

“These projections encode the time-varying sensory expectations, or targets, associated with the active speech sound map cell.”

“The map consists of 10 pairs of antagonistic cells2 that correspond to parameters of the Maeda vocal tract that determine lip protrusion, upper and lower lip height, jaw height, tongue height, tongue shape, tongue body position, tongue tip location, larynx height, and glottal opening and pressure.”

“The model has been used to guide a number of behavioural and functional imaging studies of speech processing (eg, Bohland & Guenther, 2006 ; Ghosh, Tourville & Guenther, 2008; Guenther et al, 1999 ; Lane et al, 2005 , 2007a, 2007b; Nieto-Castanon, Guenther, Perkell, & Curtin, 2005; Perkell et al, 2004a , 2004b; Tourvi…”

“Support for these modifications and the impact they have on the model are discussed below.”

“…debate over which and to what extent prosodic feature control is lateralised (e.g., Doherty, West, Dilley, Shattuck-Hufnagel, & Caplan, 2004; Emmorey, 1987; Gandour et al., 2003; Meyer, Alter, Friederici, Lohmann, & von Cramon, 2002; Stiller et al., 1997; Walker, Pelletier, & Reif, 2004).”

“Based on recent imaging work (Brown, Ngan, & Liotti, 2008; Olthoff, Baudewig, Kruse, & Dechent, 2008), additional motor cortex cells have been added to the model that represent the intrinsic laryngeal muscles.”

“…of how speech and language are rooted in the brain as extensive research has shown how many other cortical and subcortical regions besides Broca’s and Wernicke’s areas are involved in processing speech and language (Hickok and Poeppel, 2007; Tourville and Guenther, 2011; Ueno et al., 2011).”

“As some models of speech production assemble articulatory targets from phone segments (e.g., Tourville and Guenther, 2011), the present implementation can be seen as a front-end for this type of model.”

“, [65]).”

“This interpretation is reflected in the most recent version of the DIVA model (e.g., Golfinopoulos et al., 2010; Tourville and Guenther, 2010).”

“…to forward models underlying articulation (as proposed in Jaeger & Ferreira, 2013; for specific models, see Hickok, 2012; Houde & Nagarajan, 2011; Tourville & Guenther, 2011) or selection between different forward models (similar to recent proposals for speech perception, Kleinschmidt &…”

“(Guenther et al., 2006; Tourville and Guenther, 2011) (see red arrow in Figure 2C ).”

“The speech production process has been widely studied (Tian and Poeppel, 2010; Tourville and Guenther, 2011; Hickok, 2012), and while it is not fully understood, a number of brain areas are known to be involved at the level of producing articulation.”

“Although the cerebellum appears in some models of feedback control in speech (Tourville and Guenther, 2011), our results are in accord with past work in other motor domains, where patients with cerebellar pathology were found to retain the ability to use sensory feedback to correct ongoing…”

“Other brain regions, such as the pars opercularis, that are known to be active during both speech perception and production have been proposed to play a key role in speech acquisition (Guenther and Vladusich, 2012; Hickok, 2009; Lotto et al., 2009).”

“…during feedback processing have also been related to error detection in perception-based models, as it is hypothesized to drive corrective motor commands to the motor cortex after receiving input from somatosensory and auditory areas (Ito, 2008; Tourville & Guenther, 2011; Hickok, 2012).”

“The DIVA model of speech production (Guenther, et al., 2006; Guenther et al., 1998; Tourville and Guenther, 2011) is another model that incorporates auditory feedback and feed-forward commands for voice and speech control (Guenther, et al.”

“Theories of speech production make such claims about syllables and phonemes (Hickok et al., 2011; Tourville and Guenther, 2011).”

“Importantly, our word recognition in noise task involved speech production to assess recognition, which means that cingulo-opercular activity could reflect the engagement of hypothesized speech control mechanisms (Tourville and Guenther, 2011; Hickok, 2012).”

“As an example of a more interactive model, we considered the hierarchical state feedback control (HSFC) model proposed by Hickok (2012; see also Tourville & Guenther, 2011).”

“…suggestions that after the acquisition of speech around age of three, the role of auditory feedback may gradually start to change in comparison to its role during speech acquisition (Callan, Kent, Guenther, & Vorperian, 2000; Guenther, 2006; Kuhl, 1994; Kuhl, 2004; Tourville & Guenther, 2011).”

“Modulation of these language-motor interactions may be especially useful for people with nonfluent aphasias and apraxia of speech, consistent with the proposed role of the cerebellum in both feed-forward and feedback control of speech acquisition and production (Tourville & Guenther, 2011).”

“…of atypical speech in participants with nfvPPA correlated with regions typically associated with motor speech planning and execution (Guenther, 2006; Tourville & Guenther, 2011), namely the left middle and inferior frontal gyrus, cingulate, premotor area, and insula (Grossman et al., 2013;…”

“According to an influential neuro-computational model of speech, three interactive components are critical for fluent speech production [Civier et al., 2013; Tourville and Guenther, 2011].”

“Sensorimotor integration (SMI) provides an interface for speech perception and production and is fundamental to efficient verbal communication (e.g., Perrier et al., 1996; Rogalsky et al., 2011; Tourville and Guenther, 2011; Guenther and Vladusich, 2012; Moulin-Frier and Arbib, 2013).”

“the DIVA model (Tourville and Guenther, 2011) and the WEAVER++ model (Levelt et al.”

“In speech, this combined feedforward/feedback architecture is used in the DIVA model (Guenther, 2016; Tourville & Guenther, 2011).”

“The activation peak in the right middle frontal gyrus (+52, +16, +38) for speaking (>signing) is near the coordinates for the right lateralized feedback control component for speech production proposed by the DIVA model (Tourville and Guenther, 2011).”

“The plant in DIVA has historically been Maeda’s model (Tourville and Guenther, 2011), but several models of the plant have been used throughout DIVA’s development [e.g., Callan et al. (2000)], and most recently a custom plant model has been used (Guenther, 2016).”

“Nonetheless, it is plausible that the speech-production system could produce such effects, given the tight coupling between production and perception (Tourville & Guenther, 2011).”

“Data from Bouchard et al. (2013). tion, semantic processing, memory encoding, preparation, and execution of vocal motor commands (e.g., Hickok and Poeppel, 2007; Houde and Nagarajan, 2011; Tourville and Guenther, 2011; Price, 2012).”

“…notions that PWS are overly reliant on sensory feedback (Max et al., 2004) and reports that corrective feedback signals are mediated by the right hemisphere (Tourville and Guenther, 2011), these findings suggest a proportionally larger contribution of reafference to speech motor control in PWS.”

“6), supporting the notion of top-down suppression of a “wrong” linguistic prediction after the same speaker used a term that mismatched with the expected precedent.”

“The Directions Into Velocities of Articulators (DIVA) is one such example of neuroanatomically specific computational account of speech production (Guenther, 2016; Guenther et al., 2006; Tourville & Guenther, 2011).”

“…– usually called burst generators (Scudder et al. 2002) – have been the inspiration for models of the command generating processes for skills like reaching, grasping and speaking (Bullock & Grossberg 1988, Guenther 1995, Bullock et al. 1998, Shadmehr & Wise 2005, Tourville & Guenther 2011).”

“Existing models of speech motor control provide a mechanism for such an association in the form of feedback control architectures (Hickok, 2012; Hickok, Houde, & Rong, 2011; Houde & Nagarajan, 2011; Tourville & Guenther, 2011).”

“…monitoring of speech output (Allen et al., 1997; Craig-McQuaide et al., 2014), which aligns with the repeated finding of hyperactive cerebellum in stuttering (Brown et al., 2005) and with the DIVA model of speech production (Guenther et al., 2006; Civier et al., 2010; Tourville and Guenther, 2011).”

“Output phonology supports speech production when prior knowledge of speech sounds drives and monitors Frontiers in Human Neuroscience www.frontiersin.org May 2014 | Volume 8 | Article 246 | 1 HUMAN NEUROSCIENCE articulatory activity (Tourville and Guenther, 2011; Guenther and Vladusich, 2012).”

“In the DIVA model (Tourville & Guenther, 2011), the cerebellum is hypothesized to use feedback signals from auditory and somatosensory areas for feedforward control of speech production.”

“Previous studies have shown that some adults who stutter may have a defect at the level of the auditory temporal cortex (Foundas, et al., 2004) that could infl uence the shift to a left-lateralized feedforward and a rightlateralized feedback control mechanism (Tourville & Guenther, 2011).”

“Speech execution has long been argued to be mediated by cortical connections (e.g., Jurgens, 2002; Simonyan and Horwitz, 2011; Tourville and Guenther, 2011), with fMRI studies confirming activation during speech production in such areas as the anterior cingulate gyrus, motor cortex, and…”

“It is rather uncontroversial that during the development of speech production, auditory-articulatory and orosensoryarticulatory relationships must be established and encoded into internal models (Callan et al., 2000b; Tourville and Guenther, 2011; Guenther and Vladusich, 2012).”

“See Houde and Nagarajan (2011) and Tourville and Guenther (2011) for speech and Todorov (2005) and Shadmehr andMussa-Ivaldi (2012) for general treatments of this interplay.”

“Several current theories of speech motor control hypothesize that speech perception contributes to speech production through an auditory feedback mechanism that informs speech motor control (Hickok et al., 2011; Houde and Nagarajan, 2011; Tourville and Guenther, 2011).”

“…of the areas included within these networks also feature prominently in existing models of speech production, such as the primary motor cortex, ventral premotor cortex, supplementary motor area, supramarginal gyrus, deep gray nuclei, and cerebellum in the DIVA model (Tourville and Guenther, 2011).”

“As mentioned earlier, current models of speech motor control emphasize that speech production relies on a combination of feedback and feedforward control (Houde and Nagarajan, 2011; Tourville and Guenther, 2011).”

“…or sequence of speech sounds, it accurately predicts the auditory consequences of the planned movements, and it uses this prediction to inform the auditory cortex about expected feedback input (E. F. Chang et al., 2013; Houde & Nagarajan, 2011; Niziolek et al., 2013; Tourville & Guenther, 2011).”

“…of speech production consistently report bilateral STC activation, due at least in part to the auditory feedback that accompanies speech, and some models propose that auditory targets for production are represented in middle and posterior STC (e.g., the DIVA model; Tourville & Guenther, 2011).”

“According to contemporary models of speech such as the Directions into Velocities of Articulators (DIVA; Tourville and Guenther, 2011; Guenther and Vladusich, 2012) and State Feedback Control (SFC; Hickok et al.”

“Current models of auditory-motor integration for music and speech comprise a feed-forward and a feed-back component (Brown et al., 2004; Zatorre et al., 2007; Tourville and Guenther, 2011; Kleber and Zarate, 2014).”

“Thus, although this alternative feedback-based speech production network is initially recruited to alleviate stuttering (Tourville & Guenther, 2011; Tourville et al., 2008), it turns out to be inadequate, and ends up worsening symptoms (Civier et al., 2010).”

“It is commonly recorded over anterior dorsal stream regions (Tamura et al., 2012; Jenson et al., 2014), including the premotor cortex, which serves as a computational hub for processing sensorimotor information (Hickok et al., 2011; Tourville and Guenther, 2011; Guenther and Vladusich, 2012).”

“…theories of implementation (e.g., Guenther, 1995; Guenther & Perkell, 2004; Keating & Shattuck-Hufnagel, 2002; Turk & Shattuck-Hufnagel, 2014), and by prediction-based models of speech motor control (e.g., Hickok, 2012; Houde & Nagarajan, 2011; Niziolek et al., 2013; Tourville & Guenther, 2011).”

“In a TD talker, sensory-motor neural traces are posited to strengthen over repeated productions as the child maps his/ her productions onto corresponding adult forms (Guenther, 2006; Tourville & Guenther, 2011).”