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Interaction of language, auditory and memory brain networks in auditory verbal hallucinations
Open accessAbstract
Auditory verbal hallucinations (AVH) occur in psychotic disorders, but also as a symptom of other conditions and even in healthy people. Several current theories on the origin of AVH converge, with neuroimaging studies suggesting that the language, auditory and memory/limbic networks are of particular relevance. However, reconciliation of these theories with experimental evidence is missing. We review 50 studies investigating functional (EEG and fMRI) and anatomic (diffusion tensor imaging) connectivity in these networks, and explore the evidence supporting abnormal connectivity in these networks associated with AVH. We distinguish between functional connectivity during an actual hallucination experience (symptom capture) and functional connectivity during either the resting state or a task comparing individuals who hallucinate with those who do not (symptom association studies). Symptom capture studies clearly reveal a pattern of increased coupling among the auditory, language and striatal regions. Anatomical and symptom association functional studies suggest that the interhemispheric connectivity between posterior auditory regions may depend on the phase of illness, with increases in non-psychotic individuals and first episode patients and decreases in chronic patients. Leading hypotheses involving concepts as unstable memories, source monitoring, top-down attention, and hybrid models of hallucinations are supported in part by the published connectivity data, although several caveats and inconsistencies remain. Specifically, possible changes in fronto-temporal connectivity are still under debate. Precise hypotheses concerning the directionality of connections deduced from current theoretical approaches should be tested using experimental approaches that allow for discrimination of competing hypotheses.
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“In addition, evidence supports the hypothesis that hallucinations arise from release of those sensory signals by reduced top-down inhibition from higher-order centers such as frontal areas and their connections with the thalamus. 40-43”
From Towards an Integrative Account of Potential Mechanisms Mediating the Path From Sleep Dysfunction to Hallucinations (2025) · cites this paper as [41]
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“Many hypotheses of AVHs have been established in recent decades; each hypothesis explains AVHs from a different perspective (Jones, 2010; Liemburg et al., 2012; Cho and Wu, 2014; McCarthy-Jones et al., 2014; Northoff, 2014; Wilkinson, 2014; Alderson-Day et al., 2015, 2017; Hugdahl, 2015; Conde et al., 2016; Baumeister…”
BackgroundIs psychosis a dysmyelination-related information-processing disorder?
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Hallucination-Specific structure-function associations in schizophrenia
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Cognitive-Developmental Mechanisms in Hallucinations
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“For example, disconnections or over-connections in psychopathologies (Fornito, Zalesky, Pantelis, & Bullmore, 2012; Ebisch and Aleman, 2016; Ćurčić-Blake et al., 2017; Romme, de Reus, Ophoff, Kahn, & van den Heuvel, 2017) or F I G U R E 1 Global and local trajectories of brain evolution.”
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“…and different pathological features of different AH subtypes should be explored to provide insight into the mechanisms underlying AHs (Allen et al., 2008; Ćurčić-Blake et al., 2017; Diederen, Daalman, et al., 2012; Diederen, van Lutterveld, et al., 2012; Hugdahl, 2015; Upthegrove et al., 2016).”
BackgroundHistorical pursuits of the language pathway hypothesis of schizophrenia
DeLisi, L. E. (2021) · Schizophrenia · doi:10.1038/s41537-021-00182-z · Cited by 8
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Noninvasive direct current stimulation for schizophrenia: a review
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“AVH have primarily been associated with cortical language regions [16], in particular with the left temporo-parietal-occipital junction [17], a region covering the posterior left superior temporal gyrus (STG: Wernicke’s area), suggesting that these may be effective targets for”
BackgroundFilosa, M., Rossi, E. D., Carbone, G. A., Farina, B., Massullo, C., Panno, A., Adenzato, M., Ardito, R. B., & Imperatori, C. (2024) · Neuroscience Letters · doi:10.1016/j.neulet.2024.137686 · Cited by 6
“The discrepancy with our results may be due to the different approach to networks investigation (e.g., EEG vs. fMRI) and studies design (e.g., clinical vs. non-clinical samples).”
ResultBoisvert, M., Dugre, J. R., & Potvin, S. (2024) · Psychological Medicine · doi:10.1017/s003329172400165x · Cited by 9
Sun, J., Deng, H., Wang, S., Cui, Y.-P., Yang, X., Wang, C., Chen, Y., Yang, Q., Wang, H., & Qin, W. (2021) · Biological Psychiatry Cognitive Neuroscience and Neuroimaging · doi:10.1016/j.bpsc.2021.07.013 · Cited by 7
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“YMRS score 2.8 ± 1.2 2.7 ± 1.4 – .677 AHRS score 18.7 ± 5.5 – – – TA B L E 1 Demographic and clinical characteristics of the subjects by group processing areas may constitute the basis of AVH (Curcic-Blake et al., 2017).”
BackgroundInfluentialSheaves, B., Cropley, V. L., Moseley, P., Woodruff, P. W. R., Punton, G., Speth, C., Speth, J., Meerlo, P., & Brederoo, S. G. (2025) · Schizophrenia Bulletin · doi:10.1093/schbul/sbaf107 · Cited by 6
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Multi feature fusion network for schizophrenia classification and abnormal brain network recognition
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“Our finding of increased gFCD in the bilateral Broca and Wernicke regions and decreased gFCD in the bilateral hippocampus in patients with BD and AVH supports the “unstable memory” hypothesis, which holds that the failure to control the contents of memories may cause remembered experiences to rise into a person’s cons…”
BackgroundInfluentialTargeted neural network interventions for auditory hallucinations: Can TMS inform DBS?
Taylor, J. J., Krystal, J. H., D'Souza, D. C., Gerrard, J. L., & Corlett, P. R. (2018) · Schizophrenia Research · doi:10.1016/j.schres.2017.09.020 · Cited by 5
“…resonance imaging (fMRI) or electroencephalograph (EEG) signals signify the correlations between regions (functional connectivity) (Chang et al., 2017; Curcic-Blake et al., 2017) or the directional relationships between regions (effective connectivity) (Grana et al., 2017; Li et al., 2017).”
A Meta-Analysis of Functional Neuroimaging Studies of Ketamine Administration in Healthy Volunteers
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Mapping alterations in the local synchrony of the cerebral cortex in schizophrenia
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“Positive symptoms of schizophrenia (e.g., the experience of hallucinations) are associated with cortical hyperactivity [60, 61] and increased functional connectivity [62].”
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“Based on hypothesized models of hallucinations, such as the “two-hit bottom-up top-down” models (76), and their associated brain regions, a multi-site network analysis using stereoelectrocephalography (sEEG), for example, could enable simultaneous LFP recording from all regions of interest.”
MethodResultTime varying dynamics of hallucinations in clinical and non-clinical voice-hearers
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Variability in white matter structure relates to hallucination proneness
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“An explicit review can be found in Ćurčić-Blake et al.(9) Shortly, psychological models point out the importance of the subjective experience of the context and the emotion expressed in the AVHs.”
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“involved in aberrant gamma oscillations to endogenous inputs from pyramidal neurons (83) despite absence of actual external stimuli (81, 82, 85).”
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“Unstable memory is another main concept in cognitive mechanisms of AVH, which endorses that hallucinators mis-integrate intrusive memories into current context, and AVH is thus conceptualized as failure of accessing context cues (79-81).”
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“Multiple cognitive theories have been proposed delineating the emergence and phenomenology of AVH (Jones, 2010; Ćurčić-Blake et al., 2017; Rollins et al., 2019).”
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EEG Connectivity Pattern: A Window into the Schizophrenia Mind?
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Brain rhythms, neural synchrony, and networks in health and disease
Eggermont, J. J. (2021) · Elsevier eBooks · doi:10.1016/b978-0-12-819818-6.00009-1 · Cited by 2
Variability in white matter structure relates to hallucination pronenessPreprint
Johnson, J. F., Schwartze, M., Belyk, M., Pinheiro, A. P., & Kotz, S. A. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.06.10.447901 · Cited by 2
Editorial: Genetic Mechanisms of Biomarkers in Schizophrenia, Bipolar Disorder and Depression
Tang, Y., Gui, H., Tarbox, S. I., Hu, S., Ma, X., & Wang, Q. (2021) · Frontiers in Psychiatry · doi:10.3389/fpsyt.2021.736055 · Cited by 2
“Auditory verbal hallucinations (AVH), a core feature of schizophrenia, refer to hearing voices without external stimulation in the awake state (6, 7).”
BackgroundInfluentialHow voices change the brain. Two fMRI follow-up case reports
González-Vivas, C., Valdés-Florido, M. J., García‐Martí, G., Soldevila-Matías, P., Martí‐Bonmatí, L., Nácher, J., & Sanjuán, J. (2022) · Schizophrenia Research · doi:10.1016/j.schres.2022.09.016 · Cited by 2
Demler, V. F., Sterner, E., & Knolle, F. (2025) · medRxiv · doi:10.64898/2025.12.07.25341628 · Cited by 2
Sato, Y., Sakuma, A., Ohmuro, N., Katsura, M., Abe, K., Tomimoto, K., Iizuka, K., Ito, F., Tomita, H., & Matsumoto, K. (2021) · Psychiatry and Clinical Neurosciences · doi:10.1111/pcn.13284 · Cited by 1
ON THE ‘HALLUCINATIONS’ OF ARTIFICIAL INTELLIGENCE AND THE HALLUCINATION EXPERIENCE IN HUMAN
İnce, E., & Üçok, A. (2024) · Turkish Journal of Psychiatry · doi:10.5080/u27608 · Cited by 1
Guo, Z., Zhou, Z., Wang, S., Yang, L., Gao, X.-S., Xia, Y., Yang, Y., Shan, Z., Huang, H., & Tian, L. (2025) · Frontiers in Psychiatry · doi:10.3389/fpsyt.2025.1641190 · Cited by 1
Inner speech and the neurobiology of psychosis
Skipper, J. I., Lametti, D. R., & Green, D. W. (2026) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2026.106805 · Cited by 1
Epileptic aura in medicine and literature
Vasilenko, А. V., Novichkova, D. A., Ismailov, S. G., Николаевич, Ч. С., Zaripov, A. S., Евгеньевна, Д. В., Pulotova, T. I., & Bannikov, E. R. (2025) · Epilepsy and Paroxysmal Conditions · doi:10.17749/2077-8333/epi.par.con.2025.234 · Cited by 1
Alotaibi, A. M., Orfali, R. S., Alorainy, A. I., Alosaimi, M. H., Alshanawani, M., Alwtban, A. A., Bentall, R. P., & Meyer, G. (2026) · NeuroImage · doi:10.1016/j.neuroimage.2026.121733 · Cited by 1
Huang, Z., Ma, Y., Wang, J., Song, Y., Zhang, J., Ye, Z., Ouyang, D., Ke, D., Cao, D., & Dong, W. (2026) · Translational Psychiatry · doi:10.1038/s41398-026-03839-4 · Cited by 1
“Although various explanatory models have been proposed, including source-monitoring de fi cit, and predictive-coding error models [11], the underlying mechanism of AVH remains unclear.”
InfluentialAntipsychotic-related remodeling of regional brain activity in schizophrenia
Liang, J., Chen, W., & Zhang, C. (2026) · Asian Journal of Psychiatry · doi:10.1016/j.ajp.2026.105067 · Cited by 1
A life in academia: My career in brief
Hugdahl, K. (2018) · Scandinavian Journal of Psychology · doi:10.1111/sjop.12406 · Cited by 1
Orlov, N. D., Garrison, J. R., & Allen, P. (2018) · Springer eBooks · doi:10.1007/978-3-319-75124-5_11 · Cited by 1
Hubl, D., Moor, N., Kindler, J., Kottlow, M., Dierks, T., & Koenig, T. W. (2020) · Timing & Time Perception · doi:10.1163/22134468-20191163 · Cited by 1
Lin, X., Li, G., Ji, F., Cheng, L., Chen, G., Zhang, W., Su, Z., Zhang, Z., Wang, W., Zhou, C., et al. (2020) · International Journal of Neuroscience · doi:10.1080/00207454.2020.1750387 · Cited by 1
Multilayer network instability underlying persistent auditory verbal hallucinations in schizophrenia
Li, J., He, J., Ren, H., Li, Z., Ma, X., Yuan, L., Ouyang, L., Li, C., Hé, Y., Hé, Y., et al. (2024) · Psychiatry Research · doi:10.1016/j.psychres.2024.116351 · Cited by 3
Auditory verbal hallucinations as ictal phenomena in a patient with drug-resistant epilepsy
Toro-Perez, J. E., Burneo, J. G., MacDougall, K. W., McLachlan, R. S., Mirsattari, S. M., Diosy, D. C., Hayman-Abello, B. A., Aluwari, M., Herrera, M. L., Arevalo, M., et al. (2022) · Clinical Neurology and Neurosurgery · doi:10.1016/j.clineuro.2022.107223 · Cited by 1
The Impact of Technology on the Developing Visual and/or Auditory Memory in School-Aged Children
Mayer, C. (2021) · doi:10.32597/honors/254/ · Cited by 1
Lee, J. H. (2016) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/072504
Changes in the Brain Connectome With Age
Eggermont, J. J. (2019) · Elsevier eBooks · doi:10.1016/b978-0-12-815304-8.00008-6
Falkenberg, L. E., Westerhausen, R., Johnsen, E., Kroken, R., Løberg, E., Bereśniewicz, J., Kazimierczak, K., Kompus, K., Ersland, L., & Hugdahl, K. (2019) · bioRxiv · doi:10.1101/784942
Geng, H. (2020) · University of Groningen research database (University of Groningen / Centre for Information Technology) · doi:10.33612/diss.131330743
Brain Networks Detectable by fMRI during On-Line Self Report of Hallucinations in SchizophreniaPreprint
Gill, K., Percival, C. M., Roes, M., Arreaza, L., Chinchani, A. M., Sanford, N., Sena, W., Mohammadsadeghi, H., Menon, M., Hughes, M. E., et al. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.11.06.467564
A comprehensive study on the cognitive mechanisms and neural substrates of hallucination proneness
Johnson, J. F. (2022) · Research Publications (Maastricht University) · doi:10.26481/dis.20221101jj
Experimental Approaches to Understanding Thought Insertion
Brann, E., Walsh, E., Mehta, M. A., Oakley, D. A., & Deeley, Q. (2023) · doi:10.1093/med/9780192896162.003.0012
Metadata
- Authors
- Ćurčić-Blake, B., Ford, J. M., Hubl, D., Orlov, N. D., Sommer, I. E., Waters, F., Allen, P., Jardri, R., Woodruff, P. W., David, O., Mulert, C., Woodward, T. S., & Aleman, A.
- Affiliations
- San Francisco VA Medical Center (US)
- University of Bern (CH)
- King's College London (GB)
- University Medical Center Utrecht (NL)
- The University of Western Australia (AU)
- Centre National de la Recherche Scientifique (FR)
- University of Sheffield (GB)
- University Medical Center Hamburg-Eppendorf (DE)
- University of British Columbia (CA)
- University Medical Center Groningen (NL)
- ORCID
- Venue
- Progress in Neurobiology
- Publisher
- Elsevier BV
- Published
- 2017-01
- Volume
- 148
- Pages
- 1-20
- ISSN
- 0301-0082 (ISSN-L)
- PMC
- PMC5240789
- OpenAlex
- W2552561399
- Cited by
- 224
- References
- 200
- Citations by year
- 2026: 13 · 2025: 21 · 2024: 20 · 2023: 18 · 2022: 21 · 2021: 28 · 2020: 34 · 2019: 35 · 2018: 20 · 2017: 13
- FWCI
- 2.43
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Progress in Neurobiology (publishedVersion, cc-by-nc-nd)
- Europe PMC (PubMed Central) (publishedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by-nc-nd)
- PubMed (publishedVersion)
- University of Groningen research database (University of Groningen / Centre for Information Technology) (publishedVersion, cc-by-nc-nd)
- HAL (submittedVersion, cc-by-nc-nd)
- UWA Profiles and Research Repository (UWA) (submittedVersion, cc-by-nc-nd)
- BORIS (University Library Bern) (submittedVersion, cc-by-nc-nd)
- Utrecht University Repository (Utrecht University) (submittedVersion, other-oa)
- Research Portal (King's College London) (submittedVersion, cc-by-nc-nd)
- LillOA (Université de Lille (University Of Lille)) (submittedVersion, cc-by-nc-nd)
- University of Groningen research database (University of Groningen / Centre for Information Technology) (submittedVersion, cc-by-nc-nd)
- Open Access CRIS of the University of Bern
- Funders
- NWO VICI (453-11-004)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Hallucinations in medical conditions
- Health Sciences › Medicine › Psychiatry and Mental health › Schizophrenia research and treatment
- Life Sciences › Neuroscience › Cognitive Neuroscience › Functional Brain Connectivity Studies
- Keywords
- auditory verbal hallucinations, language network, auditory network, functional connectivity, structural connectivity, symptom association, interhemispheric connectivity, striatum, source monitoring, top-down attention, psychosis, first-episode psychosis, chronic psychosis, fronto-temporal connectivity
- MeSH terms
- Auditory Perception, Hallucinations, Humans, Language, Memory, Nerve Net
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.
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