Expectation Cues and False Percepts Generate Stimulus-Specific Activity in Distinct Layers of the Early Visual Cortex
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Abstract
Perception has been proposed to result from the integration of feedforward sensory signals with internally generated feedback signals. Feedback signals are believed to play an important role in driving false percepts, that is, seeing things that are not actually there. Feedforward and feedback influences on perception can be studied using layer-specific fMRI, which we used here to interrogate neural activity underlying high-confidence false percepts while healthy human participants ( N = 25, male and female) performed a perceptual orientation discrimination task. Auditory cues implicitly signaled the most likely upcoming orientation (referred to here as expectations). These expectations induced orientation-specific templates in the deep and superficial layers of V2, without affecting perception. In contrast, the orientation of falsely perceived stimuli with high confidence was reflected in the middle input layers of V2, suggesting a feedforward signal contributing to false percepts. The prevalence of high-confidence false percepts was related to everyday hallucination severity in a separate online sample ( N = 100), suggesting a possible link with abnormal perceptual experiences. These results reveal a potential feedforward mechanism underlying false percepts, reflected by spontaneous stimulus-like activity in the input layers of the visual cortex, independent of top-down signals reflecting cued orientations. SIGNIFICANCE STATEMENT False percepts have been suggested to arise through excessive feedback signals. However, feedforward contributions to false percepts have remained largely understudied. Laminar fMRI has been shown to be useful in distinguishing feedforward from feedback activity as it allows the imaging of different cortical layers. In the present study we demonstrate that although cued orientations are encoded in the feedback layers of the visual cortex, the content of the false percepts are encoded in the feedforward layers and did not rely on these cued orientations. This shows that false percepts can in principle emerge from random feedforward signals in the visual cortex, with possible implications for disorders hallmarked by hallucinations like schizophrenia and Parkinson's disease.
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People are more likely to hallucinate when sensory information is noisy, especially when combined with strong expectations.
“recent work has shown that people are more likely to hallucinate when sensory information is noisy, especially when combined with strong expectations41”
From Deviations in effective connectivity explain different hallucination subtypes in Parkinson's disease psychosis (2026) · cites this paper as [41]
The Discussion (page unavailable) reports that lower reliance on sensory evidence was associated with more everyday hallucinations, and it describes a model in which reduced sensory precision can increase the influence of expectations. But the paper did not show that strong expectations especially increased false percepts under noisy conditions; its Discussion says expectations did not affect perception in the present study. (LLM, conf=0.86)
Experimental — accuracy not yet measuredSpontaneous feedforward activity in the visual cortex can also lead to hallucinations, rather than feedback activity alone.
“suggesting that spontaneous feedforward activity in the visual cortex can also lead to hallucinations, rather than feedback activity alone41”
From Deviations in effective connectivity explain different hallucination subtypes in Parkinson's disease psychosis (2026) · cites this paper as [41]
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Roth, Z. N., Kay, K., & Merriam, E. P. (2022) · Nature Communications · doi:10.1038/s41467-022-34134-7 · Cited by 25 · see citing works
Perceptual pathways to hallucinogenesis
Sheldon, A. D., Kafadar, E., Fisher, V., Greenwald, M. S., Aitken, F., Negreira, A. M., Woods, S. W., & Powers, A. R. (2022) · Schizophrenia Research · doi:10.1016/j.schres.2022.02.002 · Cited by 18 · see citing works
Leptourgos, P., Bouttier, V., Denève, S., & Jardri, R. (2021) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2022.104593 · Cited by 13 · see citing works
Subjective visibility report is facilitated by conscious predictions only
Alilović, J., Slagter, H. A., & van Gaal, S. (2020) · Consciousness and Cognition · doi:10.1016/j.concog.2020.103048 · Cited by 4 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
Predictions and errors are distinctly represented across V1 layers
Thomas, E. R., Haarsma, J., Nicholson, J. L., Yon, D., Kok, P., & Press, C. (2024) · Current Biology · doi:10.1016/j.cub.2024.04.036 · Cited by 51 · see citing works
Self-citationGoodwin, I., Kugel, J., Hester, R., & Garrido, M. I. (2023) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1011670 · Cited by 21 · see citing works
Carricarte, T., Iamshchinina, P., Trampel, R., Chaimow, D., Weiskopf, N., & Cichy, R. M. (2024) · iScience · doi:10.1016/j.isci.2024.110229 · Cited by 15 · see citing works
Shenyan, O., Lisi, M., Greenwood, J. A., Skipper, J. I., & Dekker, T. M. (2024) · Scientific Reports · doi:10.1038/s41598-024-52372-1 · Cited by 14 · see citing works
Warrington, O., Graedel, N. N., Callaghan, M. F., & Kok, P. (2025) · Science Advances · doi:10.1126/sciadv.ads4970 · Cited by 13 · see citing works
Self-citationPre-stimulus alpha oscillations encode stimulus-specific visual predictionsPreprint
Hetenyi, D., Haarsma, J., & Kok, P. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.03.13.584593 · Cited by 12 · see citing works
Self-citationGuided by Expectations: Overweighted Semantic Priors in Schizotypy and Their Links to Glutamate
Knolle, F., Sterner, E., Demler, V. F., MacGregor, L. J., & Mathys, C. D. (2025) · Biological Psychiatry · doi:10.1016/j.biopsych.2025.06.025 · Cited by 10 · see citing works
Predictive processing accounts of psychosis: bottom-up or top-down disruptions
Goodwin, I., Diederen, K. M. J., Hird, E., Weilnhammer, V. A., Garrido, M. I., & Knolle, F. (2026) · Nature Mental Health · doi:10.1038/s44220-025-00558-5 · Cited by 9 · see citing works
Haarsma, J., Kaltenmaier, A., Fleming, S. M., & Kok, P. (2024) · Cognition · doi:10.1016/j.cognition.2024.105961 · Cited by 7 · see citing works
Self-citationParanoid and teleological thinking give rise to distinct social hallucinations in vision
Castiello, S., Ongchoco, J. D. K., van Buren, B., Scholl, B. J., & Corlett, P. R. (2024) · Communications Psychology · doi:10.1038/s44271-024-00163-9 · Cited by 6 · see citing works
Shared and Diverging Neural Dynamics Underlying False and Veridical Perception
Haarsma, J., Hetenyi, D., & Kok, P. (2025) · Journal of Neuroscience · doi:10.1523/jneurosci.1479-24.2025 · Cited by 5 · see citing works
Self-citationContents of Visual Predictions Oscillate at Alpha Frequencies
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Self-citationTesting circuit-level theories of consciousness in humans
Dykstra, A. R., Zhu, Y., Fernandez Pujol, C., Zhou, D. W., Jones, S. R., Marvan, T., & Bonaiuto, J. J. (2026) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2025.08.012 · Cited by 3 · see citing works
Stockart, F., Robin, A., Blumenfeld, H., Brázdil, M., Kahane, P., Mudrik, L., Thum, J. A., Pereira, M., & Faivre, N. (2026) · eLife · doi:10.7554/elife.109604.4 · Cited by 5 · see citing works
Mu, Y., Liu, Q., Chu, Y., Zhao, X., Ding, W., Wang, H., & Jiang, Y. (2025) · Nature Communications · doi:10.1038/s41467-025-67380-6 · Cited by 3 · see citing works
Learning alters salience and proactive attentional priority
Duncan, D. H., van Moorselaar, D., & Theeuwes, J. (2026) · Communications Psychology · doi:10.1038/s44271-026-00411-0 · Cited by 5 · see citing works
Haarsma, J., & Kok, P. (2025) · Current topics in behavioral neurosciences · doi:10.1007/7854_2025_595 · Cited by 2 · see citing works
Self-citationCastiello, S., Ongchoco, J. D. K., van Buren, B. F., Scholl, B. J., & Corlett, P. R. (2024) · doi:10.31234/osf.io/4wnfv · Cited by 2 · see citing works
Current status and perspectives of ultrahigh-field magnetic resonance imaging
Bolin, Q. I. N., & Gao, J. (2025) · Acta Physica Sinica · doi:10.7498/aps.74.20241759 · Cited by 1 · see citing works
Marsh, D. C., Sokoliuk, R., Aquino, K. M., Pakenham, D. O., Wilson, R., Panchuelo, R. S., Brookes, M. J., Hanslmayr, S., Mayhew, S. D., Francis, S., et al. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.09.09.674189 · Cited by 1 · see citing works
Vignando, M., Ffytche, D., Mazibuko, N., Palma, G., Bhat, A., Montagnese, M., Dave, S., Tai, Y. F., Batzu, L., Leta, V., et al. (2026) · Nature Mental Health · doi:10.1038/s44220-026-00669-7
Flicker and Ganzfeld induced visual hallucinations differ in frequency and contentPreprint
Shenyan, O., Lisi, M., Greenwood, J. A., Skipper, J. I., & Dekker, T. M. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.08.08.552408
The Future of Perceptual Dysregulation in Psychiatric Nosology
Powers, A. R., & Khalsa, S. S. (2025) · Current topics in behavioral neurosciences · doi:10.1007/7854_2025_596
Stockart, F., Robin, A., Blumenfeld, H., Brázdil, M., Kahane, P., Mudrik, L., Thum, J., Pereira, M., & Faivre, N. (2026) · eLife · doi:10.7554/elife.109604
Stockart, F. (2024) · theses.fr (ABES) · doi:10.70675/c4f62883zfadez4797z9c4ezc049226f96a4
False Percepts as a Window onto Visual Processing
Quinn, K. R., & Sandhaeger, F. (2026) · Journal of Neuroscience · doi:10.1523/jneurosci.1942-25.2026
- Authors
- Haarsma, J., Deveci, N., Corbin, N., Callaghan, M. F., & Kok, P.
- Affiliations
- Wellcome Centre for Human Neuroimaging (GB)
- Corresponding author
- Haarsma, Joost
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2023-09-22
- Volume
- 43
- Issue
- 47
- Pages
- 7946-7957
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC10669763
- OpenAlex
- W4386952214
- Cited by
- 26
- References
- 94
- Citations by year
- 2026: 6 · 2025: 10 · 2024: 8 · 2023: 2
- FWCI
- 3.88
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Locations
- Journal of Neuroscience (publishedVersion, cc-by)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- UCL Discovery (University College London) (submittedVersion, cc-by)
- HAL (submittedVersion)
- Oskar-Bordeaux (Universite de Bordeaux) (submittedVersion)
- Funders
- Wellcome Trust ([218535/Z/19/Z], [203147/Z/16/Z])
- European Research Council ([948548])
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Visual perception and processing mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Hallucinations in medical conditions
- Life Sciences › Neuroscience › Cognitive Neuroscience › Face Recognition and Perception
- Keywords
- visual hallucinations, feedback signals, early visual cortex, V2, cortical layers, orientation discrimination, auditory cues, spontaneous neuronal activity, schizophrenia, Parkinson's disease
- MeSH terms
- Cues, Feedback, Female, Humans, Magnetic Resonance Imaging, Male, Motivation, Visual Cortex, Visual Perception
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 ›
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