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Neural Representations and Mechanisms for the Performance of Simple Speech Sequences
Open accessAbstract
Speakers plan the phonological content of their utterances before their release as speech motor acts. Using a finite alphabet of learned phonemes and a relatively small number of syllable structures, speakers are able to rapidly plan and produce arbitrary syllable sequences that fall within the rules of their language. The class of computational models of sequence planning and performance termed competitive queuing models have followed K. S. Lashley [The problem of serial order in behavior. In L. A. Jeffress (Ed.), Cerebral mechanisms in behavior (pp. 112-136). New York: Wiley, 1951] in assuming that inherently parallel neural representations underlie serial action, and this idea is increasingly supported by experimental evidence. In this article, we developed a neural model that extends the existing DIVA model of speech production in two complementary ways. The new model includes paired structure and content subsystems [cf. MacNeilage, P. F. The frame/content theory of evolution of speech production. Behavioral and Brain Sciences, 21, 499-511, 1998 ] that provide parallel representations of a forthcoming speech plan as well as mechanisms for interfacing these phonological planning representations with learned sensorimotor programs to enable stepping through multisyllabic speech plans. On the basis of previous reports, the model's components are hypothesized to be localized to specific cortical and subcortical structures, including the left inferior frontal sulcus, the medial premotor cortex, the basal ganglia, and the thalamus. The new model, called gradient order DIVA, thus fills a void in current speech research by providing formal mechanistic hypotheses about both phonological and phonetic processes that are grounded by neuroanatomy and physiology. This framework also generates predictions that can be tested in future neuroimaging and clinical case studies.
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“Finally, the left putamen and thalamus were incorporated into a motor loop that passes activity from the SMA via the putamen to the thalamus and into the motor cortex (Bohland et al., 2010).”
From A review and synthesis of the first 20years of PET and fMRI studies of heard speech, spoken language and reading (2012) · cites this paper as Bohland et al., 2010
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“A blueprint for the organization of neural control in speech production is given by the DIVA model (Directions Into Velocities of Articulators, see Guenther 2006, Bohland et al., 2010, and Guenther and Vladusich 2012).”
BackgroundMethodHybbinette, H., Östberg, P., Schalling, E., Deboussard, C. N., Plantin, J., Borg, J., & Lindberg, P. G. (2022) · Frontiers in Neurology · doi:10.3389/fneur.2022.1013652 · Cited by 7
“The importance of the premotor cortex in speech production has also been highlighted in neurocomputational models of speech production, such as the Directions Into Velocities of Articulators (DIVA) model (20) and its expanded version, the Gradient Order DIVA (GODIVA) model (21, 22).”
BackgroundLinn, W., Barrios‐Martinez, J., Fernandes-Cabral, D. T., Jacquesson, T., Nunez, M. A., Gómez, R. J., Anania, Y., Fernandez-Miranda, J. C., & Yeh, F.-C. F. (2024) · Human Brain Mapping · doi:10.1002/hbm.26630 · Cited by 7
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“The GODIVA model of the neural mechanisms of speech production postulates that a cortical-basal ganglia loop circuit controls the release of planned speech sounds to the execution system so that a new motor program is initiated only upon completion of the previous program (22).”
Chrabaszcz, A. V., Wang, D., Lipski, W. J., Bush, A., Crammond, D. J., Shaiman, S., Dickey, M. W., Holt, L. L., Turner, R. S., Fiez, J. A., et al. (2021) · Journal of Neurolinguistics · doi:10.1016/j.jneuroling.2021.101019 · Cited by 7
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“38 shown to receive preparatory signals on the order of information from pre-SMA and to send driving signals to the motor regions that execute motor commands (Bohland et al., 2010).”
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Xu, D., Chung, J. E., Silva, A. B., Metzger, S. L., Greicius, Q., Zhang, Y., Leonard, M. K., Aghi, M. K., Hervey‐Jumper, S. L., & Chang, E. F. (2024) · SSRN Electronic Journal · doi:10.1101/2024.12.27.630563 · Cited by 9
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“We brie fl y address the gradient order DIVA (GODIVA) model here because it concerns speech planning mechanisms 96,98 , which is informative for the θ – α SPC differences we observed >1 s before speech onset in accurate versus error trials.”
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“Under this view, coordinated neural mechanisms must sequentially select the appropriate phonological units from a planning buffer, while concurrently activating lower-level sensory-motor programs that drive the production of individual syllables (Bohland et al., 2010).”
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“We used the DIVA/GODIVA model(4) (Bohland et al., 2010; Guenther, 2016; Guenther et al., 2006), a well-supported detailed model of speech motor planning and control, because it meets these criteria.”
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“This model proposes that the premotor elements of different movements to be executed in sequence are activated (or readied) in parallel and then selected sequentially through a mechanism where movements occurring earlier in the sequence suppress later ones (Bohland et al., 2010; Bullock, 2004; Houghton and Hartley, 19…”
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“While the pre-SMA is often included in models of the neural bases of speech (i.e., the motor planning and execution aspects of language production; Bohland et al., 2010; Dick et al., 2019; Price, 2010), it is absent from many of the dominant models of the broader language network (e.g., Dell et…”
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A Novel Investigation of Generalized Motor Program Theory: Syllable Stress as a Motor-Class Variable
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“The complexity of speech production would intuitively suggest that multiple GMPs are encoded and retrieved from memory during speech production (e.g., phonemes and syllable levels; Bohland et al., 2010; Roelofs & Meyer, 1998); therefore, the results of this study should not be surprising.”
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Phonological variation and lexical form
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“Despite decades of investigation, a clear account of the interaction between the neural processes underlying speech production and their dynamics is still needed (Bohland et al. 2010; Laga-naro 2019; Miller & Guenther 2021; Verwoert et al. 2022).”
Rowe, H. P., Miller, H. E., Belisle, R. M., Perrachione, T. K., & Guenther, F. H. (2025) · Neurobiology of Language · doi:10.1162/nol.a.18 · Cited by 2
“Both LpIFS and RCbm are thought to be involved in the learning of speech sequences, such as the sequences of phonemes making up the phonotactically illegal syllables used here (Bohland et al., 2010; Bohland & Guenther, 2006; Guenther, 2016).”
Kröger, B. J. (2023) · Journal of Integrative Neuroscience · doi:10.31083/j.jin2205124 · Cited by 2
“The sensorimotor neural production models DIVA [4] and GODIVA [5] represent further components of the production pathway from the phonological form via activation of motor programs down to motor execution and include auditory and somatosensory feedback to allow motor corrections.”
MethodInfluentialLateralization and Time-Course of Cortical Phonological Representations during Syllable Production
Meier, A. M., Kuzdeba, S., Jackson, L., Daliri, A., Tourville, J. A., Guenther, F. H., & Greenlee, J. D. W. (2023) · eNeuro · doi:10.1523/eneuro.0474-22.2023 · Cited by 3
Kröger, B. J. (2023) · Frontiers in Language Sciences · doi:10.3389/flang.2023.1100774 · Cited by 2
Cummine, J., Ngo, T., & Nisbet, K. (2023) · Brain Sciences · doi:10.3390/brainsci13121622 · Cited by 2
“In the speech production literature, the caudate is implicated in speech planning [2] and has been noted for playing a role in rhythm.”
BackgroundInfluentialDesjardins, M., Jomphe, V., Lagadec-Gaulin, L., Cohen, M. L., & Abbott, K. V. (2024) · Journal of Speech Language and Hearing Research · doi:10.1044/2024_jslhr-23-00673 · Cited by 2
Distinct neural processes link speech planning and execution
Duraivel, S., Rahimpour, S., Barth, K. J., Chiang, C., Wang, C., Harward, S. C., Lad, S. P., Sexton, D., Friedman, A. H., Sinha, S. R., et al. (2026) · Nature Human Behaviour · doi:10.1038/s41562-026-02546-w · Cited by 3
“Based on neuro-anatomical studies that investigated speech articulation 4,16,20,65,68 , we divided these ROIs into three distinct networks based on previous literature: planning (IFG, rMFG, and cMFG), articulation (PrCG & PoCG, and IPC), and monitoring (aSTG, pSTG, STS, and PAC).”
InfluentialTerband, H., Rodd, J., & Maas, E. (2015) · MPG.PuRe (Max Planck Society) · Cited by 2
Phoneme and Stress Programming Interact During Nonword Repetition Learning
Meigh, K. M., Cobun, E. R., & Yunusova, Y. (2020) · Journal of Speech Language and Hearing Research · doi:10.1044/2020_jslhr-19-00262 · Cited by 2
Ahmadi, Z. Z., DiBacco, M. L., & Pearl, P. L. (2021) · Journal of Child Neurology · doi:10.1177/08830738211001210 · Cited by 2
“Concerning the connectivity of subcortical areas and SMA, it is mostly cited that the anterior regions of the SMA are strongly connected to the caudate,(35) which is involved in cognitive processes(36,37) and phonological segmentation.(38) In contrast, the posterior regions of SMA are strongly connected to the thalamu…”
BackgroundJackson, E. S. (2015) · CUNY Academic Works (City University of New York) · Cited by 2
Children’s Use of Syntax In Word Learning
Lidz, J. L. (2022) · Oxford University Press eBooks · doi:10.1093/oxfordhb/9780198845003.013.10 · Cited by 2
Phonological Abstraction in The Mental Lexicon
Bakovic, E. J., Heinz, J. N., & Rawski, J. (2022) · Oxford University Press eBooks · doi:10.1093/oxfordhb/9780198845003.013.14 · Cited by 2
Nozari, N., & Pinet, S. (2019) · doi:10.31234/osf.io/x9wq7 · Cited by 2
Defining brain pathways for human spoken language and vocal learningPreprint
Theofanopoulou, C., & Jarvis, E. D. (2023) · doi:10.31234/osf.io/pa5fq · Cited by 2
Manes, J. L., & Guenther, F. H. (2024) · Elsevier eBooks · doi:10.1016/b978-0-12-820480-1.00120-0 · Cited by 2
Some properties of mental speech preparation as revealed by self-monitoring
Quené, H., & Nooteboom, S. G. (2024) · Speech Communication · doi:10.1016/j.specom.2024.103043 · Cited by 1
Speaking demands: concurrent visuomotor rotor pursuit tracking measures cognitive load for speech
Kriegel, Z., & Whitfield, J. A. (2024) · Speech Language and Hearing · doi:10.1080/2050571x.2024.2317027 · Cited by 2
“However, it was expected that sentence repetition and counting forward by one might slightly degrade percent time on target in the visuomotor rotor pursuit tracking task because of a more complex sequence of intended phonemes translated into speech motor commands (Bohland et al., 2010).”
InfluentialProduction training and contextual similarity hurt the comprehension of new vocabulary
Waller, M., Yurovsky, D., & Nozari, N. (2024) · Language Cognition and Neuroscience · doi:10.1080/23273798.2024.2429127 · Cited by 1
“This link is critical for both learning and monitoring language production (e.g. Bohland et al., 2010; Hickok, 2012; Levelt et al., 1999; Nozari, 2020).”
Right-Hemispheric White Matter Organization Is Associated With Speech Timing in Autistic Children
Davison, K. E., Liu, T., Belisle, R. M., Perrachione, T. K., Qi, Z., Gabrieli, J. D. E., Tager-Flusberg, H. B., & Zuk, J. (2025) · Journal of Speech Language and Hearing Research · doi:10.1044/2025_jslhr-24-00548 · Cited by 1
“…a region known to play a major role in auditory processing of spectrotemporal information important for speech perception (Hickok & Poeppel, 2007); and (c) the pre – supplementary motor area (preSMA), a speech motor region involved in the planning of articulatory movements (Bohland et al., 2010).”
Prem, P., Saggu, S., Boadu, A., Saju, S., Nisbet, K., & Cummine, J. (2025) · Neuroscience · doi:10.1016/j.neuroscience.2025.09.035 · Cited by 2
Höbler, F., Liu, Y., Beltz, A. M., Becker, H. C., Angstadt, M., Guenther, F. H., & Chang, S. (2025) · Neurobiology of Language · doi:10.1162/nol.a.26 · Cited by 1
“With most of the DIVA and GODIVA nodes implicated in developmental stuttering, moments of stuttering are hypothesized to arise from specific disruptions to the functioning of the basal ganglia proper (Alm, 2004, 2021; Civier et al., 2013), or in corticostriatal WM connectivity (Civier et al., 2013), thereby impacting…”
InfluentialLi, S., Chen, Z., Luo, X., Wang, J., Teng, P., Luan, G., Wang, Q., & Tian, X. (2025) · Communications Biology · doi:10.1038/s42003-025-08847-6 · Cited by 1
Testing theories of stuttering: cognitive heterogeneity but shared orosensory tactile weaknesses
Thorre, A. V., Patri, J.-F., & Vannuscorps, G. (2025) · Cognitive Neuropsychology · doi:10.1080/02643294.2025.2572357 · Cited by 1
“In the GODIVA model (Bohland et al., 2010), stut-tering arises from delays in the deactivation of the ongoing syllable (leading to prolongations) or in the activation of the subsequent syllable (leading to blocks).”
An Imaging‐Guided Neural Model Explains Lexical Stress Alteration in Acquired Apraxia of Speech
Civier, O., Ramage, A. E., Tourville, J. A., Robin, D. A., Guenther, F. H., & Ballard, K. J. (2025) · Human Brain Mapping · doi:10.1002/hbm.70412 · Cited by 1
“…explanation of the atypical neural dynamics that give rise to AOS, and to simulate them numerically using a neuro-computational model of speech production (i.e., a custom instantiation of the DIVA/ GODIVA family of speech production models, see Bohland et al. 2010; Guenther et al. 2006).”
InfluentialKasper, J., Kell, C. A., & Perrier, P. (2025) · Journal of Neurolinguistics · doi:10.1016/j.jneuroling.2025.101298 · Cited by 1
The neural correlates of mid- and end-clause silent pauses in L1 and L2 speech
Révész, A., Jeong, H., Suzuki, S., Cui, H., Matsuura, S., Saito, K., & Sugiura, M. (2026) · Bilingualism Language and Cognition · doi:10.1017/s1366728926101011 · Cited by 2
Erasmus clinical model of the onset and development of stuttering 2.0
Franken, M.-C. J. P., Oonk, L. C., Bast, B. J. E. G., Bouwen, J., & Nil, L. D. (2024) · Journal of Fluency Disorders · doi:10.1016/j.jfludis.2024.106040 · Cited by 1
Inhibitory modulation of speech trajectories: Evidence from a vowel-modified Stroop task
Beach, S. D., & Niziolek, C. A. (2024) · Cognitive Neuropsychology · doi:10.1080/02643294.2024.2315831 · Cited by 1
“As speaking is perhaps the most complex human motor behaviour, lateral inhibition underlies many theoretical and computational accounts of speech motor planning and selection (Bohland et al., 2010; Meyer & Gordon, 1985; Schade & Berg, 1992; Tilsen, 2013; Yaniv et al., 1990).”
BackgroundCai, X. F., & Zhang, Q. (2025) · Brain Research · doi:10.1016/j.brainres.2025.150010 · Cited by 1
Metadata
- Authors
- Bohland, J. W., Bullock, D., & Guenther, F. H.
- Affiliations
- Cold Spring Harbor Laboratory (US)
- Boston University (US)
- Venue
- Journal of Cognitive Neuroscience
- Publisher
- MIT Press
- Published
- 2010-07-01
- Volume
- 22
- Issue
- 7
- Pages
- 1504-1529
- ISSN
- 0898-929X (ISSN-L)
- 1530-8898
- PMC
- PMC2937837
- OpenAlex
- W2114774561
- Cited by
- 393
- References
- 158
- Citations by year
- 2026: 21 · 2025: 26 · 2024: 29 · 2023: 33 · 2022: 64 · 2021: 15 · 2020: 22 · 2019: 40 · 2018: 18 · 2017: 26 · 2016: 19 · 2015: 23 · 2014: 16 · 2013: 18 · 2012: 10
- FWCI
- 3.72
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Journal of Cognitive Neuroscience (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Cold Spring Harbor Laboratory Institutional Repository (Cold Spring Harbor Laboratory) (acceptedVersion)
- DSpace@MIT (Massachusetts Institute of Technology) (submittedVersion)
- Funders
- National Science Foundation (0354378)
- National Institutes of Health (NIH-R01, DC002852)
- National Institute on Deafness and Other Communication Disorders (R01DC002852, R01DC007683)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neurobiology of Language and Bilingualism
- Social Sciences › Psychology › Experimental and Cognitive Psychology › Phonetics and Phonology Research
- Social Sciences › Psychology › Developmental and Educational Psychology › Language Development and Disorders
- Keywords
- speech production, phonological planning, sequence planning, DIVA model, serial order, basal ganglia, thalamus
- MeSH terms
- Basal Ganglia, Computer Simulation, Frontal Lobe, Models, Neurological, Neural Pathways, Phonetics, Psychomotor Performance, Speech, Thalamus
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