The graphemic/motor frontal area Exner’s area revisited
Abstract
Objective: In 1881, Exner first described a “graphic motor image center” in the middle frontal gyrus. Current psycholinguistic models of handwriting involve the conversion of abstract, orthographic representations into motor representations before a sequence of appropriate hand movements is produced. Direct cortical stimulation and functional magnetic resonance imaging (fMRI) were used to study the human frontal areas involved in writing. Methods: Cortical electrical stimulation mapping was used intraoperatively in 12 patients during the removal of brain tumors to identify the areas involved in oral language (sentence reading and naming) and writing, and to spare them during surgery. The fMRI activation experiment involved 12 right‐handed and 12 left‐handed healthy volunteers using word dictation (without visual control) and 2 control tasks. Results: Direct cortical electrical stimulation of restricted areas rostral to the primary motor hand area (Brodmann area [BA] 6) impaired handwriting in 6 patients, without disturbing hand movements or oral language tasks. In 6 other patients, stimulation of lower frontal regions showed deficits combining handwriting with other language tasks. fMRI also revealed selective activation during word handwriting in left versus right BA6 depending on handedness. This area was anatomically matched to those areas that affected handwriting on electrical stimulation. Interpretation: An area in middle frontal gyrus (BA6) that we have termed the graphemic/motor frontal area supports bridging between orthography and motor programs specific to handwriting. Ann Neurol 2009;66:537–545
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Roux et al. found similar activation in the left superior parietal cortex during typing, suggesting that some aspects of orthographic encoding are shared between handwriting and typing.
“In contrast, Roux et al. [19] studied 24 participants via fMRI and found similar activation in the same area during typing, suggesting that some aspects of orthographic encoding are shared between both modalities.”
From The Neuroscience Behind Writing: Handwriting vs. Typing—Who Wins the Battle? (2025) · cites this paper as [19]
Cites
Different neural substrates for Kanji and Kana writing: a PET study
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A Neurosurgical Functional Dissection of the Middle Precentral Gyrus during Speech Production
Silva, A. B., Liu, J. R., Zhao, L., Levy, D. F., Scott, T. L., & Chang, E. F. (2022) · The Journal of Neuroscience · doi:10.1523/jneurosci.1614-22.2022 · Cited by 111 · see citing works
Functional architecture of the motor homunculus detected by electrostimulation
Roux, F., Niare, M., Charni, S., Giussani, C., & Durand, J. (2020) · The Journal of Physiology · doi:10.1113/jp280156 · Cited by 87 · see citing works
Self-citationHu, J., Small, H., Kean, H., Takahashi, A., Zekelman, L., Kleinman, D., Ryan, E., Nieto-Castañón, A., Ferreira, V., & Fedorenko, E. (2023) · Cerebral Cortex · doi:10.1093/cercor/bhac350 · Cited by 73 · see citing works
The Neuroscience Behind Writing: Handwriting vs. Typing—Who Wins the Battle?
Marano, G., Kotzalidis, G. D., Lisci, F. M., Anesini, M. B., Rossi, S., Barbonetti, S., Cangini, A., Ronsisvalle, A., Artuso, L., Falsini, C., et al. (2025) · Life · doi:10.3390/life15030345 · Cited by 32 · see citing works
Sierpowska, J., León-Cabrera, P., Camins, À., Juncadella, M., Gabarrós, A., & Rodríguez-Fornells, A. (2020) · Cortex · doi:10.1016/j.cortex.2020.06.009 · Cited by 10 · see citing works
Levy, D. F., Silva, A. B., Scott, T. L., Liu, J. R., Harper, S., Zhao, L., Hullett, P. W., Kurteff, G., Wilson, S. M., Leonard, M. K., et al. (2023) · Journal of Neurosurgery: Case Lessons · doi:10.3171/case22504 · Cited by 10 · see citing works
- Authors
- Roux, F., Dufor, O., Giussani, C., Wamain, Y., Draper, L., Longcamp, M., & Démonet, J.
- Affiliations
- Inserm (FR)
- Université Toulouse III - Paul Sabatier (FR)
- Centre Hospitalier Universitaire de Toulouse (FR)
- Corresponding author
- Roux, Franck‐Emmanuel
- ORCID
- Venue
- Annals of Neurology
- Publisher
- Wiley
- Published
- 2009-10
- Volume
- 66
- Issue
- 4
- Pages
- 537-545
- ISSN
- 0364-5134 (ISSN-L)
- 1531-8249
- OpenAlex
- W2114369124
- Cited by
- 173
- References
- 1
- Citations by year
- 2026: 6 · 2025: 10 · 2024: 8 · 2023: 11 · 2022: 15 · 2021: 15 · 2020: 9 · 2019: 14 · 2018: 13 · 2017: 7 · 2016: 8 · 2015: 11 · 2014: 7 · 2013: 13 · 2012: 10
- FWCI
- 3.94
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Annals of Neurology (publishedVersion)
- PubMed (publishedVersion)
- HAL (submittedVersion)
- BOA (University of Milano-Bicocca) (submittedVersion)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neurobiology of Language and Bilingualism
- Life Sciences › Neuroscience › Cognitive Neuroscience › Motor Control and Adaptation
- Life Sciences › Neuroscience › Cognitive Neuroscience › Hemispheric Asymmetry in Neuroscience
- Keywords
- handwriting, middle frontal gyrus, Brodmann area 6, orthographic representation, motor representations, handedness, word dictation
- MeSH terms
- Adult, Brain Mapping, Electric Stimulation, Female, Frontal Lobe, Handwriting, Humans, Magnetic Resonance Imaging, Male, Middle Aged, Motor Cortex, Young Adult
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