Words in the brain's language
Open accessAbstract
If the cortex is an associative memory, strongly connected cell assemblies will form when neurons in different cortical areas are frequently active at the same time. The cortical distributions of these assemblies must be a consequence of where in the cortex correlated neuronal activity occurred during learning. An assembly can be considered a functional unit exhibiting activity states such as full activation (“ignition”) after appropriate sensory stimulation (possibly related to perception) and continuous reverberation of excitation within the assembly (a putative memory process). This has implications for cortical topographies and activity dynamics of cell assemblies forming during language acquisition, in particular for those representing words. Cortical topographies of assemblies should be related to aspects of the meaning of the words they represent, and physiological signs of cell assembly ignition should be followed by possible indicators of reverberation. The following postulates are discussed in detail: (1) assemblies representing phonological word forms are strongly lateralized and distributed over perisylvian cortices; (2) assemblies representing highly abstract words such as grammatical function words are also strongly lateralized and restricted to these perisylvian regions; (3) assemblies representing concrete content words include additional neurons in both hemispheres; (4) assemblies representing words referring to visual stimuli include neurons in visual cortices; and (5) assemblies representing words referring to actions include neurons in motor cortices. Two main sources of evidence are used to evaluate these proposals: (a) imaging studies focusing on localizing word processing in the brain, based on stimulus-triggered event-related potentials (ERPs), positron emission tomography (PET), and functional magnetic resonance imaging (fMRI), and (b) studies of the temporal dynamics of fast activity changes in the brain, as revealed by high-frequency responses recorded in the electroencephalogram (EEG) and magnetoencephalogram (MEG). These data provide evidence for processing differences between words and matched meaningless pseudowords, and between word classes, such as concrete content and abstract function words, and words evoking visual or motor associations. There is evidence for early word class-specific spreading of neuronal activity and for equally specific high-frequency responses occurring later. These results support a neurobiological model of language in the Hebbian tradition. Competing large-scale neuronal theories of language are discussed in light of the data summarized. Neurobiological perspectives on the problem of serial order of words in syntactic strings are considered in closing.
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Recent studies have shown that the neural mechanisms of speech production and perception are connected and overlapping, even in motor areas.
“recent studies have shown that the neural mechanisms of speech production and perception are connected and overlapping, even in motor areas68–71.”
From Inner speech in motor cortex and implications for speech neuroprostheses (2025) · cites this paper as [69]
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Cites
Hebb, D. O. (2005) · doi:10.4324/9781410612403 · Cited by 9,825 · see citing works
Cognitive maps in rats and men
Tolman, E. C. (1948) · Psychological Review · doi:10.1037/h0061626 · Cited by 6,641 · see citing works
Basic objects in natural categories
Rosch, E., Mervis, C. B., Gray, W. D., Johnson, D. M., & Boyes-Braem, P. (1976) · Cognitive Psychology · doi:10.1016/0010-0285(76)90013-x · Cited by 5,643 · see citing works
Gray, C. M., König, P., Engel, A. K., & Singer, W. (1989) · Nature · doi:10.1038/338334a0 · Cited by 4,183 · see citing works
Bienenstock, E., Cooper, L., & Munro, P. (1982) · The Journal of Neuroscience · doi:10.1523/jneurosci.02-01-00032.1982 · Cited by 2,896 · see citing works
Unconscious cerebral initiative and the role of conscious will in voluntary action
Libet, B. (1985) · Behavioral and Brain Sciences · doi:10.1017/s0140525x00044903 · Cited by 2,623 · see citing works
Emotion, attention, and the startle reflex
Lang, P. J., Bradley, M. M., & Cuthbert, B. N. (1990) · Psychological Review · doi:10.1037/0033-295x.97.3.377 · Cited by 2,205 · see citing works
CORTICAL PLASTICITY: From Synapses to Maps
Buonomano, D. V., & Merzenich, M. M. (1998) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.21.1.149 · Cited by 2,083 · see citing works
A Bio‐Informational Theory of Emotional Imagery
Lang, P. J. (1979) · Psychophysiology · doi:10.1111/j.1469-8986.1979.tb01511.x · Cited by 1,750 · see citing works
On the time relations of mental processes: An examination of systems of processes in cascade.
McClelland, J. L. (1979) · Psychological Review · doi:10.1037/0033-295x.86.4.287 · Cited by 1,298 · see citing works
Cortex: Statistics and Geometry of Neuronal Connectivity
Braitenberg, V., & Schüz, A. (1998) · doi:10.1007/978-3-662-03733-1 · Cited by 1,178 · see citing works
Memory processes, brain oscillations and EEG synchronization
Klimesch, W. (1996) · International Journal of Psychophysiology · doi:10.1016/s0167-8760(96)00057-8 · Cited by 923 · see citing works
Crick, F., & Mitchison, G. (1983) · Nature · doi:10.1038/304111a0 · Cited by 865 · see citing works
Development and plasticity of cortical processing architectures
Singer, W. (1995) · Science · doi:10.1126/science.270.5237.758 · Cited by 669 · see citing works
Models of visual word recognition: Sampling the state of the art.
Jacobs, A. M., & Grainger, J. (1994) · Journal of Experimental Psychology: Human Perception and Performance · doi:10.1037/0096-1523.20.6.1311 · Cited by 319 · see citing works
Neurobiological mechanisms of language acquisition
Pulvermüller, F., & Schumann, J. H. (1994) · Language Learning · doi:10.1111/j.1467-1770.1994.tb00635.x · Cited by 189 · see citing works
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Barsalou, L. W. (1999) · Behavioral and Brain Sciences · doi:10.1017/s0140525x99002149 · Cited by 6,887 · see citing works
Temporal binding and the neural correlates of sensory awareness.
Engel, A. K., & Singer, W. (2001) · Trends in Cognitive Sciences · doi:10.1016/s1364-6613(00)01568-0 · Cited by 1,372 · see citing works
Rogers, T. T., Lambon Ralph, M. A., Garrard, P., Bozeat, S., McClelland, J. L., Hodges, J. R., & Patterson, K. (2004) · Psychological Review · doi:10.1037/0033-295x.111.1.205 · Cited by 948 · see citing works
Grounded Cognition: Past, Present, and Future: Topics in Cognitive Science
Barsalou, L. W. (2010) · Topics in Cognitive Science · doi:10.1111/j.1756-8765.2010.01115.x · Cited by 993 · see citing works
Situated simulation in the human conceptual system
Barsalou, L. (2003) · Language and Cognitive Processes · doi:10.1080/01690960344000026 · Cited by 979 · see citing works
AT THE INTERFACE: DYNAMIC INTERACTIONS OF EXPLICIT AND IMPLICIT LANGUAGE KNOWLEDGE
Ellis, N. C. (2005) · Studies in Second Language Acquisition · doi:10.1017/s027226310505014x · Cited by 893 · see citing works
How neurons make meaning: brain mechanisms for embodied and abstract-symbolic semantics
Pulvermüller, F. (2013) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2013.06.004 · Cited by 631 · see citing works
Perceptions of perceptual symbols
Barsalou, L. W. (1999) · Behavioral and Brain Sciences · doi:10.1017/s0140525x99532147 · Cited by 620 · see citing works
Intentional attunement: Mirror neurons and the neural underpinnings of interpersonal relations
Gallese, V., Eagle, M. N., & Migone, P. (2007) · Journal of the American Psychoanalytic Association · doi:10.1177/00030651070550010601 · Cited by 496 · see citing works
Convergence of biological and psychological perspectives on cognitive coordination in schizophrenia
Phillips, W. A., & Silverstein, S. M. (2003) · Behavioral and Brain Sciences · doi:10.1017/s0140525x03000025 · Cited by 445 · see citing works
Longcamp, M., Zerbato-Poudou, M.-T., & Velay, J.-L. (2005) · Acta Psychologica · doi:10.1016/j.actpsy.2004.10.019 · Cited by 434 · see citing works
Neural 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
Activity and Imagined Activity Can Enhance Young Children's Reading Comprehension.
Glenberg, A. M., Gutierrez, T., Levin, J. R., Japuntich, S., & Kaschak, M. P. (2004) · Journal of Educational Psychology · doi:10.1037/0022-0663.96.3.424 · Cited by 401 · see citing works
The emergence of competing modules in bilingualism
HERNANDEZ, A., LI, P., & MACWHINNEY, B. (2005) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2005.03.003 · Cited by 366 · see citing works
Emotional and semantic networks in visual word processing: insights from ERP studies
Kissler, J., Assadollahi, R., & Herbert, C. (2006) · Progress in Brain Research · doi:10.1016/s0079-6123(06)56008-x · Cited by 329 · see citing works
Modelling the N400 brain potential as change in a probabilistic representation of meaning
Rabovsky, M., Hansen, S. S., & McClelland, J. L. (2018) · Nature Human Behaviour · doi:10.1038/s41562-018-0406-4 · Cited by 290 · see citing works
Challenges and opportunities for grounding cognition.
Barsalou, L. W. (2020) · Journal of Cognition · doi:10.5334/joc.116 · Cited by 319 · see citing works
Imagined speech can be decoded from low- and cross-frequency intracranial EEG features
Proix, T., Delgado Saa, J., Christen, A., Martin, S., Pasley, B. N., Knight, R. T., Tian, X., Poeppel, D., Doyle, W. K., Devinsky, O., et al. (2022) · Nature Communications · doi:10.1038/s41467-021-27725-3 · Cited by 212 · see citing works
Grounding symbolic operations in the brain’s modal systems
Barsalou, L. W. (2008) · Embodied Grounding · doi:10.1017/cbo9780511805837.002 · Cited by 179 · see citing works
Birba, A., García-Cordero, I., Kozono, G., Legaz, A., Ibáñez, A., Sedeño, L., & García, A. M. (2017) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2017.07.011 · Cited by 128 · see citing works
Frequency effects of Chinese character processing in the brain: an event-related fMRI study
Kuo, W.-J., Yeh, T.-C., Lee, C.-Y., Wu, Y. U.-T. E., Chou, C.-C., Ho, L.-T., Hung, D. L., Tzeng, O. J. L., & Hsieh, J.-C. (2003) · NeuroImage · doi:10.1016/s1053-8119(03)00015-6 · Cited by 129 · see citing works
The Oxford Handbook of Event-Related Potential Components
(2011) · doi:10.1093/oxfordhb/9780195374148.001.0001 · Cited by 2,263 · see citing works
Boudelaa, S. (2014) · Literacy Studies · doi:10.1007/978-94-017-8545-7_2 · Cited by 104 · see citing works
What differs in visual recognition of handwritten vs. printed letters? An fMRI study
Longcamp, M., Hlushchuk, Y., & Hari, R. (2011) · Human Brain Mapping · doi:10.1002/hbm.21105 · Cited by 85 · see citing works
The Language Network Is Recruited but Not Required for Nonverbal Event Semantics
Ivanova, A. A., Mineroff, Z., Zimmerer, V., Kanwisher, N., Varley, R., & Fedorenko, E. (2021) · Neurobiology of Language · doi:10.1162/nol_a_00030 · Cited by 65 · see citing works
Inner speech in motor cortex and implications for speech neuroprostheses
Kunz, E. M., Abramovich Krasa, B., Kamdar, F., Avansino, D. T., Hahn, N., Yoon, S., Singh, A., Nason-Tomaszewski, S. R., Card, N. S., Jude, J. J., et al. (2025) · Cell · doi:10.1016/j.cell.2025.06.015 · Cited by 55 · see citing works
Is prediction nothing more than multi-scale pattern completion of the future?
Falandays, J. B., Nguyen, B., & Spivey, M. J. (2021) · Brain Research · doi:10.1016/j.brainres.2021.147578 · Cited by 36 · see citing works
Language and motor cortex response to comprehending accidental and intentional action sentences
Kana, R. K., Ammons, C. J., Doss, C. F., Waite, M. E., Kana, B., Herringshaw, A. J., & Ver Hoef, L. (2015) · Neuropsychologia · doi:10.1016/j.neuropsychologia.2015.08.020 · Cited by 15 · see citing works
- Authors
- Pulvermüller, F.
- Venue
- Behavioral and Brain Sciences
- Publisher
- Cambridge University Press (CUP)
- Published
- 1999-04
- Volume
- 22
- Issue
- 2
- Pages
- 253-279
- ISSN
- 0140-525X (ISSN-L)
- 1469-1825
- OpenAlex
- W2099535060
- Cited by
- 1,284
- References
- 16
- Citations by year
- 2026: 27 · 2025: 30 · 2024: 45 · 2023: 22 · 2022: 27 · 2021: 35 · 2020: 29 · 2019: 52 · 2018: 63 · 2017: 29 · 2016: 63 · 2015: 39 · 2014: 51 · 2013: 53 · 2012: 48
- FWCI
- 20.08
- Type
- journal article
- Language
- en
- Open access
- green
- nbn-resolving.de
- Licence
- cc-by-nc-nd
- www.cambridge.org/core/terms (unspecified)
- Locations
- Behavioral and Brain Sciences (publishedVersion)
- PubMed (publishedVersion)
- KOPS (University of Konstanz) (submittedVersion, cc-by-nc-nd)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neurobiology of Language and Bilingualism
- Social Sciences › Psychology › Developmental and Educational Psychology › Reading and Literacy Development
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Keywords
- word processing, associative memory, language acquisition, function words, hemispheric lateralization, perisylvian cortex, cortical topography, reverberation, Hebbian learning, visual word recognition, action words, high-frequency response, event-related potentials
- MeSH terms
- Association Learning, Brain Mapping, Cerebral Cortex, Cognition, Dominance, Cerebral, Electroencephalography, Evoked Potentials, Humans, Linguistics, Magnetic Resonance Imaging, Memory, Models, Neurological, Motor Cortex, Periaqueductal Gray, Tomography, Emission-Computed, Visual Cortex, Magnetoencephalography
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