From phonemes to articulatory codes: an fMRI study of the role of Broca's area in speech production
Open accessAbstract
We used event-related functional magnetic resonance imaging to investigate the neuroanatomical substrates of phonetic encoding and the generation of articulatory codes from phonological representations. Our focus was on the role of the left inferior frontal gyrus (LIFG) and in particular whether the LIFG plays a role in sublexical phonological processing such as syllabification or whether it is directly involved in phonetic encoding and the generation of articulatory codes. To answer this question, we contrasted the brain activation patterns elicited by pseudowords with high- or low-sublexical frequency components, which we expected would reveal areas related to the generation of articulatory codes but not areas related to phonological encoding. We found significant activation of a premotor network consisting of the dorsal precentral gyrus, the inferior frontal gyrus bilaterally, and the supplementary motor area for low- versus high-sublexical frequency pseudowords. Based on our hypothesis, we concluded that these areas and in particular the LIFG are involved in phonetic and not phonological encoding. We further discuss our findings with respect to the mechanisms of phonetic encoding and provide evidence in support of a functional segregation of the posterior part of Broca's area, the pars opercularis.
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Broca's area plays a role in normal speech production.
“We do not question the role that Broca’s area has been shown to play in normal speech production”
From Damage to Broca’s area does not contribute to long-term speech production outcome after stroke (2021) · cites this paper as Papoutsi et al., 2009
The ventral left pars opercularis has a role in articulatory recoding of speech.
“consistent with a role in articulatory recoding of speech (Papoutsi et al., 2009).”
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 Papoutsi et al., 2009
The left dorsal pars opercularis is associated with processing prior to articulatory recoding.
“The left dorsal pars opercularis was associated with processing prior to articulatory recoding (Papoutsi et al., 2009) and was found to be commonly activated by both the production and observation of nonsemantic, nonsyntactic, and nonsense syllables (Fridriksson et al., 2009).”
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 Papoutsi et al., 2009
The ventral left pars opercularis is activated during covert articulation.
“The ventral left pars opercularis was activated during covert articulation (Zheng et al., 2010; Papoutsi et al., 2009) with increased activation for phonological relative to semantic retrieval (Heim et al., 2009a), consistent with a role in articulatory recoding of speech (Papoutsi et al., 2009).”
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 Papoutsi et al., 2009
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Cites
Rizzolatti, G., & Craighero, L. (2004) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.27.070203.144230 · Cited by 7,433 · see citing works
The cortical organization of speech processing
Hickok, G., & Poeppel, D. (2007) · Nature Reviews Neuroscience · doi:10.1038/nrn2113 · Cited by 5,495 · see citing works
A theory of lexical access in speech production [target paper]
Levelt, W. J. M., Roelofs, A., & Meyer, A. S. (1999) · Radboud Repository (Radboud University) · doi:10.1017/s0140525x99001776 · Cited by 5,173 · see citing works
A new SPM toolbox for combining probabilistic cytoarchitectonic maps and functional imaging data
Eickhoff, S. B., Stephan, K. E., Mohlberg, H., Grefkes, C., Fink, G. R., Amunts, K., & Zilles, K. (2005) · NeuroImage · doi:10.1016/j.neuroimage.2004.12.034 · Cited by 4,033 · see citing works
Understanding motor events: a neurophysiological study
di Pellegrino, G., Fadiga, L., Fogassi, L., Gallese, V., & Rizzolatti, G. (1992) · Experimental Brain Research · doi:10.1007/bf00230027 · Cited by 3,539 · see citing works
Cattaneo, L., & Rizzolatti, G. (2009) · Archives of Neurology · doi:10.1001/archneurol.2009.41 · Cited by 3,137 · see citing works
The motor theory of speech perception revised
Liberman, A. M., & Mattingly, I. G. (1985) · Cognition · doi:10.1016/0010-0277(85)90021-6 · Cited by 2,865 · see citing works
Rizzolatti, G., & Arbib, M. A. (1998) · Trends in Neurosciences · doi:10.1016/s0166-2236(98)01260-0 · Cited by 2,858 · see citing works
Gazzaniga, M. S., & Bizzi, E. (2000) · Cited by 2,525 · see citing works
Working memory and language: an overview
Baddeley, A. (2003) · Journal of Communication Disorders · doi:10.1016/s0021-9924(03)00019-4 · Cited by 2,344 · see citing works
Region of interest analysis using an SPM toolbox
Brett, M. (2002) · NeuroImage · Cited by 2,273 · see citing works
Hickok, G., & Poeppel, D. (2004) · Cognition · doi:10.1016/j.cognition.2003.10.011 · Cited by 2,212 · see citing works
The spatial and temporal signatures of word production components
Indefrey, P., & Levelt, W. J. M. (2004) · Cognition · doi:10.1016/j.cognition.2002.06.001 · Cited by 2,144 · see citing works
Event-Related fMRI: Characterizing Differential Responses
Friston, K. J., Fletcher, P. C., Josephs, O., Holmes, A., Rugg, M. D., & Turner, R. (1998) · NeuroImage · doi:10.1006/nimg.1997.0306 · Cited by 2,017 · see citing works
Lateralization of Phonetic and Pitch Discrimination in Speech Processing
Zatorre, R. J., Evans, A. C., Meyer, E., & GJEDDE, A. H. (1992) · Science · doi:10.1126/science.1589767 · Cited by 1,507 · see citing works
Poldrack, R. A., Wagner, A. D., Prull, M. W., Desmond, J. E., Glover, G. H., & Gabrieli, J. D. E. (1999) · NeuroImage · doi:10.1006/nimg.1999.0441 · Cited by 1,448 · see citing works
Towards a functional neuroanatomy of speech perception
Hickok, G. S., & Poeppel, D. (2000) · Trends in Cognitive Sciences · doi:10.1016/s1364-6613(00)01463-7 · Cited by 1,136 · see citing works
Neural modeling and imaging of the cortical interactions underlying syllable production
Guenther, F. H., Ghosh, S. S., & Tourville, J. A. (2006) · Brain and Language · doi:10.1016/j.bandl.2005.06.001 · Cited by 879 · see citing works
Infants' sensitivity to phonotactic patterns in the native language.
Jusczyk, P., Luce, P. A., & Charles-Luce, J. (1994) · Journal of Memory and Language · doi:10.1006/jmla.1994.1030 · Cited by 858 · see citing works
Auditory-motor interaction revealed by fMRI: speech, music, and working memory in area Spt
Hickok, G., Buchsbaum, B., Humphries, C., & Muftuler, T. (2003) · Journal of Cognitive Neuroscience · doi:10.1162/089892903322307393 · Cited by 700 · see citing works
The motor theory of speech perception reviewed
Galantucci, B., Fowler, C. A., & Turvey, M. T. (2006) · Psychonomic Bulletin & Review · doi:10.3758/bf03193857 · Cited by 675 · see citing works
Levelt, W. J. M. (1999) · Trends in Cognitive Sciences · doi:10.1016/s1364-6613(99)01319-4 · Cited by 619 · see citing works
When Words Compete: Levels of Processing in Perception of Spoken Words
Vitevitch, M. S., & Luce, P. A. (1998) · Psychological Science · doi:10.1111/1467-9280.00064 · Cited by 603 · see citing works
Devlin, J. T., Matthews, P. M., & Rushworth, M. F. S. (2003) · Journal of Cognitive Neuroscience · doi:10.1162/089892903321107837 · Cited by 577 · see citing works
Do speakers have access to a mental syllabary?
Levelt, W. J. M., & Wheeldon, L. R. (1994) · Cognition · doi:10.1016/0010-0277(94)90030-2 · Cited by 559 · see citing works
A Web-based interface to calculate phonotactic probability for words and nonwords in English
Vitevitch, M. S., & Luce, P. A. (2004) · Behavior Research Methods, Instruments, & Computers · doi:10.3758/bf03195594 · Cited by 547 · see citing works
An fMRI investigation of syllable sequence production
Bohland, J. W., & Guenther, F. H. (2006) · NeuroImage · doi:10.1016/j.neuroimage.2006.04.173 · Cited by 545 · see citing works
Validating cluster size inference: random field and permutation methods
Hayasaka, S. (2003) · NeuroImage · doi:10.1016/j.neuroimage.2003.08.003 · Cited by 491 · see citing works
PET Studies of Phonetic Processing of Speech: Review, Replication, and Reanalysis
Zatorre, R. J., Meyer, E., GJEDDE, A. H., & Evans, A. C. (1996) · Cerebral Cortex · doi:10.1093/cercor/6.1.21 · Cited by 422 · see citing works
Some Notes on Syllable Structure in Articulatory Phonology
Browman, C. P., & Goldstein, L. (1988) · Phonetica · doi:10.1159/000261823 · Cited by 406 · see citing works
Perception of Wordlikeness: Effects of Segment Probability and Length on the Processing of Nonwords
Frisch, S. A., Large, N. R., & Pisoni, D. B. (2000) · Journal of Memory and Language · doi:10.1006/jmla.1999.2692 · Cited by 395 · see citing works
Fiez, J. A., Balota, D. A., Raichle, M. E., & Petersen, S. E. (1999) · Neuron · doi:10.1016/s0896-6273(00)80833-8 · Cited by 390 · see citing works
The role of segmentation in phonological processing: an fMRI investigation
Burton, M. W., Small, S. L., & Blumstein, S. E. (2000) · Journal of Cognitive Neuroscience · doi:10.1162/089892900562309 · Cited by 378 · see citing works
Processing Linguistic Complexity and Grammaticality in the Left Frontal Cortex
Friederici, A. D., Fiebach, C. J., Schlesewsky, M., Bornkessel‐Schlesewsky, I., & von Cramon, D. Y. (2005) · Cerebral Cortex · doi:10.1093/cercor/bhj106 · Cited by 366 · see citing works
fMRI reveals two distinct cerebral networks subserving speech motor control
Riecker, A., Mathiak, K., Wildgruber, D., Erb, M., Hertrich, I., Grodd, W., & Ackermann, H. (2005) · Neurology · doi:10.1212/01.wnl.0000152156.90779.89 · Cited by 327 · see citing works
Phonotactics and Syllable Stress: Implications for the Processing of Spoken Nonsense Words
Vitevitch, M. S., Luce, P. A., Charles-Luce, J., & Kemmerer, D. L. (1997) · Language and Speech · doi:10.1177/002383099704000103 · Cited by 322 · see citing works
Phonotactics, Neighborhood Activation, and Lexical Access for Spoken Words
Vitevitch, M. S., Luce, P. A., Pisoni, D. B., & Auer, E. T. (1999) · Brain and Language · doi:10.1006/brln.1999.2116 · Cited by 275 · see citing works
The neural correlates of language production
Indefrey, P., & Levelt, W. J. M. (2000) · MPG.PuRe (Max Planck Society) · Cited by 273 · see citing works
Efficient Design of Event-Related fMRI Experiments Using M-Sequences
Buračas, G. T., & Boynton, G. M. (2002) · NeuroImage · doi:10.1006/nimg.2002.1116 · Cited by 271 · see citing works
Molnar-Szakacs, I., Iacoboni, M., Koski, L., & Mazziotta, J. C. (2005) · Cerebral Cortex · doi:10.1093/cercor/bhh199 · Cited by 266 · see citing works
Effects of syllable frequency in speech production
Cholin, J., Levelt, W. J. M., & Schiller, N. O. (2005) · Cognition · doi:10.1016/j.cognition.2005.01.009 · Cited by 228 · see citing works
Speech perception, word recognition and the structure of the lexicon
Pisoni, D. B., Nusbaum, H. C., Luce, P. A., & Slowiaczek, L. M. (1985) · Speech Communication · doi:10.1016/0167-6393(85)90037-8 · Cited by 219 · see citing works
Bokde, A. L. W., Tagamets, M.-A., Friedman, R. B., & Horwitz, B. (2001) · Neuron · doi:10.1016/s0896-6273(01)00288-4 · Cited by 211 · see citing works
Self-citationComparison of fMRI motion correction software tools
Oakes, T., Johnstone, T., Walsh, K. S. O., Greischar, L., Alexander, A., Fox, A., & Davidson, R. (2005) · NeuroImage · doi:10.1016/j.neuroimage.2005.05.058 · Cited by 207 · see citing works
An fMRI investigation of covertly and overtly produced mono- and multisyllabic words
SHUSTER, L., & LEMIEUX, S. (2005) · Brain and Language · doi:10.1016/j.bandl.2004.07.007 · Cited by 205 · see citing works
Retinotopic organization of early visual spatial attention effects as revealed by PET and ERPs
Woldorff, M. G., Fox, P. T., Matzke, M., Lancaster, J. L., Veeraswamy, S., Zamarripa, F., Seabolt, M., Glass, T. G., Gao, J., Martin, C. C., et al. (1997) · Human Brain Mapping · doi:10.1002/(sici)1097-0193(1997)5:4 · Cited by 195 · see citing works
Lexical and post-lexical phonological representations in spoken production
Goldrick, M., & Rapp, B. C. (2006) · Cognition · doi:10.1016/j.cognition.2005.12.010 · Cited by 187 · see citing works
Barch, D. M., Sabb, F. W., Carter, C. S., Braver, T. S., Noll, D. C., & Cohen, J. D. (1999) · NeuroImage · doi:10.1006/nimg.1999.0500 · Cited by 186 · see citing works
Graves, W. W., Grabowski, T. J., Mehta, S., & Gordon, J. K. (2007) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2007.19.4.617 · Cited by 178 · see citing works
de Zwart, J. A., Ledden, P. J., van Gelderen, P., Bodurka, J., Chu, R., & Duyn, J. H. (2003) · Magnetic Resonance in Medicine · doi:10.1002/mrm.10678 · Cited by 176 · see citing works
Increases in phonotactic probability facilitate spoken nonword repetition
Vitevitch, M. S., & Luce, P. A. (2005) · Journal of Memory and Language · doi:10.1016/j.jml.2004.10.003 · Cited by 166 · see citing works
Dissociating Neural Mechanisms of Temporal Sequencing and Processing Phonemes
Gelfand, J. R., & Bookheimer, S. Y. (2003) · Neuron · doi:10.1016/s0896-6273(03)00285-x · Cited by 163 · see citing works
Experimental designs and processing strategies for fMRI studies involving overt verbal responses
Birn, R. M., Cox, R. W., & Bandettini, P. A. (2004) · NeuroImage · doi:10.1016/j.neuroimage.2004.07.039 · Cited by 154 · see citing works
Functional heterogeneity within Broca's area during verbal working memory
Chein, J. M., Fissell, K., Jacobs, S., & Fiez, J. A. (2002) · Physiology & Behavior · doi:10.1016/s0031-9384(02)00899-5 · Cited by 144 · see citing works
Heim, S., Eickhoff, S. B., Ischebeck, A. K., Friederici, A. D., Stephan, K. E., & Amunts, K. (2009) · Human Brain Mapping · doi:10.1002/hbm.20512 · Cited by 133 · see citing works
The Neural Correlate of Speech Rhythm as Evidenced by Metrical Speech Processing
Geiser, E., Zaehle, T., Jäncke, L., & Meyer, M. E. (2007) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2008.20029 · Cited by 118 · see citing works
PET Studies of Phonological Processing: A Critical Reply to Poeppel
Démonet, J., Fiez, J. A., Paulesu, E., Petersen, S. E., & Zatorre, R. J. (1996) · Brain and Language · doi:10.1006/brln.1996.0109 · Cited by 107 · see citing works
Okada, K., & Hickok, G. S. (2006) · Neuroreport · doi:10.1097/01.wnr.0000233091.82536.b2 · Cited by 95 · see citing works
Reduction of Gradient Acoustic Noise in MRI Using SENSE-EPI
de Zwart, J. A., Vangelderen, P., Kellman, P., & Duyn, J. H. (2002) · NeuroImage · doi:10.1006/nimg.2002.1119 · Cited by 65 · see citing works
Method for functional MRI mapping of nonlinear response
Kellman, P., van Gelderen, P., de Zwart, J. A., et al. (2003) · NeuroImage · doi:10.1016/s1053-8119(03)00056-9 · Cited by 30 · see citing works
Broca's region revisited: Cytoarchitecture and intersubject variability
Amunts, K., Schleicher, A., B�rgel, U., Mohlberg, H., Uylings, H. B. M., & Zilles, K. (1999) · The Journal of Comparative Neurology · doi:10.1002/(sici)1096-9861(19990920)412:2
Flagged first, then by citations
Research works printed but not linked: 3
- Bachoud-Lévi A Dupoux E Cohen L Mehler J 1998 Where Is the Length Effect? A Cross-Linguistic Study of Speech Production Journal of Memory and Language 39 3 331 346
- Iacoboni M 2005 Understanding others: Imitation, language, empathy Perspectives on imitation: from cognitive neuroscience to social science 1 Cambridge, MA MIT Press
- Mazaika P Whitfield-Gabrieli S Reiss A 2007 Artifact Repair for fMRI Data from High Motion Clinical Subjects Human Brain Mapping Conference
Cited by
Sorted by citations
Price, C. J. (2012) · NeuroImage · doi:10.1016/j.neuroimage.2012.04.062 · Cited by 2,159 · see citing works
The anatomy of language: a review of 100 fMRI studies published in 2009
Price, C. J. (2010) · Annals of the New York Academy of Sciences · doi:10.1111/j.1749-6632.2010.05444.x · Cited by 1,387 · see citing works
The Spatial and Temporal Signatures of Word Production Components: A Critical Update
Indefrey, P. (2011) · Frontiers in Psychology · doi:10.3389/fpsyg.2011.00255 · Cited by 767 · see citing works
“To date only one study (Papoutsi et al., 2009) used an experimental variable (biphone frequency) that convincingly disentangles syllabification from later phonetic and motor processing stages and confirmed a pattern predicted for syllabification in a subregion of Broca’s area.”
BackgroundMethodInfluentialNeural representations and mechanisms for the performance of simple speech sequences
Bohland, J. W., Bullock, D., & Guenther, F. H. (2010) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2009.21306 · Cited by 393 · see citing works
“A similar region in left dorsal inferior frontal gyrus has been suggested to be involved in sequencing discrete units, including phonemes (Gelfand&Bookheimer, 2003), and in phonological encoding tasks (Papoutsi et al., 2009; Chein & Fiez, 2001; Burton, Small, & Blumstein, 2000; Poldrack et al., 1999).”
Coupled neural systems underlie the production and comprehension of naturalistic narrative speech
Silbert, L. J., Honey, C. J., Simony, E., Poeppel, D., & Hasson, U. (2014) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1323812111 · Cited by 352 · see citing works
Laryngeal Motor Cortex and Control of Speech in Humans
Simonyan, K., & Horwitz, B. (2011) · The Neuroscientist · doi:10.1177/1073858410386727 · Cited by 310 · see citing works
Self-citationParker Jones, O., Green, D. W., Grogan, A., Pliatsikas, C., Filippopolitis, K., Ali, N., Lee, H. L., Ramsden, S., Gazarian, K., Prejawa, S., et al. (2012) · Cerebral Cortex · doi:10.1093/cercor/bhr161 · Cited by 221 · see citing works
Functional segregation of cortical regions underlying speech timing and articulation
Long, M. A., Katlowitz, K. A., Svirsky, M. A., Clary, R. C., Byun, T. M., Majaj, N., Oya, H., Howard, M. A., & Greenlee, J. D. W. (2016) · Neuron · doi:10.1016/j.neuron.2016.01.032 · Cited by 151 · see citing works
Damage to Broca’s area does not contribute to long-term speech production outcome after stroke
Gajardo-Vidal, A., Lorca-Puls, D. L., team, P., Warner, H., Pshdary, B., Crinion, J. T., Leff, A. P., Hope, T. M. H., Geva, S., Seghier, M. L., et al. (2021) · Brain · doi:10.1093/brain/awaa460 · Cited by 150 · see citing works
Distributed neural representations of phonological features during speech perception
Arsenault, J. S., & Buchsbaum, B. R. (2015) · The Journal of Neuroscience · doi:10.1523/jneurosci.2454-14.2015 · Cited by 111 · see citing works
Vaden, K. I., Piquado, T., & Hickok, G. (2011) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2011.21620 · Cited by 36 · see citing works
The effect of sublexical and lexical frequency on speech production: an fMRI investigation
Shuster, L. I. (2009) · Brain and Language · doi:10.1016/j.bandl.2009.06.003 · Cited by 32 · see citing works
- Authors
- Papoutsi, M., de Zwart, J. A., Jansma, J. M., Pickering, M. J., Bednar, J. A., & Horwitz, B.
- Affiliations
- National Institutes of Health (US)
- University of Edinburgh (GB)
- Venue
- Cerebral Cortex
- Publisher
- Oxford University Press (OUP)
- Published
- 2009-01-29
- Volume
- 19
- Issue
- 9
- Pages
- 2156-2165
- ISSN
- 1047-3211 (ISSN-L)
- 1460-2199
- PMC
- PMC2722428
- OpenAlex
- W2132092497
- Cited by
- 190
- References
- 62
- Citations by year
- 2026: 9 · 2025: 11 · 2024: 14 · 2023: 8 · 2022: 15 · 2021: 8 · 2020: 9 · 2019: 10 · 2018: 11 · 2017: 14 · 2016: 12 · 2015: 11 · 2014: 10 · 2013: 13 · 2012: 11
- FWCI
- 5.65
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Cerebral Cortex (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- UCL Discovery (University College London) (submittedVersion)
- Funders
- National Institutes of Health (Z99 DC999999)
- National Institute on Deafness and Other Communication Disorders (Z01DC000057)
- 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 › Reading and Literacy Development
- Keywords
- Broca's area, speech production, phonetic encoding, phonological representation, left inferior frontal gyrus, syllabification, supplementary motor area, pars opercularis, pseudowords, functional segregation
- MeSH terms
- Adult, Brain Mapping, Female, Frontal Lobe, Humans, Language, Magnetic Resonance Imaging, Male, Semantics, Speech
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review