Perceptual Learning beyond Perception: Mnemonic Representation in Early Visual Cortex and Intraparietal Sulcus
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Abstract
The ability to discriminate between stimuli relies on a chain of neural operations associated with perception, memory and decision-making. Accumulating studies show learning-dependent plasticity in perception or decision-making, yet whether perceptual learning modifies mnemonic processing remains unclear. Here, we trained human participants of both sexes in an orientation discrimination task, while using functional magnetic resonance imaging (fMRI) and transcranial magnetic stimulation (TMS) to separately examine training-induced changes in working memory (WM) representation. fMRI decoding revealed orientation-specific neural patterns during the delay period in primary visual cortex (V1) before, but not after, training, whereas neurodisruption of V1 during the delay period led to behavioral deficits in both phases. In contrast, both fMRI decoding and disruptive effect of TMS showed that intraparietal sulcus (IPS) represented WM content after, but not before, training. These results suggest that training does not affect the necessity of sensory area in representing WM information, consistent with the sensory recruitment hypothesis in WM, but likely alters the coding format of the stored stimulus in this region. On the other hand, training can render WM content to be maintained in higher-order parietal areas, complementing sensory area to support more robust maintenance of information. SIGNIFICANCE STATEMENT There has been accumulating progresses regarding experience-dependent plasticity in perception or decision-making, yet how perceptual experience moulds mnemonic processing of visual information remains less explored. Here, we provide novel findings that learning-dependent improvement of discriminability accompanies altered WM representation at different cortical levels. Critically, we suggest a role of training in modulating cortical locus of WM representation, providing a plausible explanation to reconcile the discrepant findings between human and animal studies regarding the recruitment of sensory or higher-order areas in WM.
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Transfer has been observed from high-precision visual perception training to relatively low-precision WM tasks, but not in the reverse direction.
“Consistent with this interpretation, transfer has been observed from high-precision visual perception training to relatively low-precision WM tasks (Jia et al., 2021), but not in the reverse direction (Cai et al., 2022).”
From The asymmetry of working memory training transfer: A systematic review and meta-analysis (2026) · cites this paper as Jia et al., 2021
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From remembering to reconstruction: The transformative neural representation of episodic memory.
Xue, G. (2022) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2022.102351 · Cited by 49 · see citing works
Sensory Recruitment Revisited: Ipsilateral V1 Involved in Visual Working Memory
Zhao, Y.-J., Kay, K., Tian, Y., & Ku, Y. (2021) · Cerebral Cortex · doi:10.1093/cercor/bhab300 · Cited by 39 · see citing works
Ultra-High Field Imaging of Human Visual Cognition
Jia, K., Goebel, R., & Kourtzi, Z. (2023) · Annual Review of Vision Science · doi:10.1146/annurev-vision-111022-123830 · Cited by 28 · see citing works
Self-citationPhylactou, P., Traikapi, A., Papadatou-Pastou, M., & Konstantinou, N. (2022) · Psychonomic Bulletin & Review · doi:10.3758/s13423-022-02107-y · Cited by 15 · see citing works
Different Forms of Plasticity Interact in Adult Humans
Sarı, İ. D., & Lunghi, C. (2023) · eNeuro · doi:10.1523/eneuro.0204-22.2023 · Cited by 14 · see citing works
Causal evidence for the role of the sensory visual cortex in visual short-term memory maintenance
Phylactou, P., Shimi, A. G., & Konstantinou, N. (2023) · Royal Society Open Science · doi:10.1098/rsos.230321 · Cited by 9 · see citing works
Top-down modulation of DLPFC in visual search: a study based on fMRI and TMS
Tian, Y., Tan, C., Tan, J., Li, Y., & Tang, Y. (2024) · Cerebral Cortex · doi:10.1093/cercor/bhad540 · Cited by 8 · see citing works
Self-citationElectrophysiological hallmarks for event relations and event roles in working memory
Yu, X., Li, J., Zhu, H., Tian, X., & Lau, E. F. (2024) · Frontiers in Neuroscience · doi:10.3389/fnins.2023.1282869 · Cited by 6 · see citing works
Divergent roles of early visual cortex and inferior frontal junction in visual working memory
Zhao, Y., Zhang, X., & Ku, Y. (2024) · Brain stimulation · doi:10.1016/j.brs.2024.06.001 · Cited by 6 · see citing works
A Test of Three Models of Character Reversal in Typically Developing Children’s Writing
Fischer, J., & Luxembourger, C. (2021) · Frontiers in Communication · doi:10.3389/fcomm.2021.719652 · Cited by 4 · see citing works
Comparative analysis of working memory tasks in fMRI and MEG studies
Pechenkova, E., Korolkova, O. A., Panikratova, Y. R., Pchelintseva, M. E., & Sinitsyn, V. E. (2025) · Experimental Psychology (Russia) · doi:10.17759/exppsy.2025180112 · Cited by 3 · see citing works
Consorti, A., & Sale, A. (2025) · Frontiers in Neural Circuits · doi:10.3389/fncir.2025.1636023 · Cited by 2 · see citing works
Huang, H., Zhang, Y., & Li, S. (2022) · Attention Perception & Psychophysics · doi:10.3758/s13414-022-02520-4 · Cited by 2 · see citing works
Self-citationReactivation-induced memory integration prevents proactive interference in perceptual learning
Huang, Z., Niu, Z., & Li, S. (2023) · Journal of Vision · doi:10.1167/jov.23.5.1 · Cited by 2 · see citing works
Self-citationTask-irrelevant features in working memory alter current visual processing.
Shi, H., Zhang, Q., Zhou, J., Ding, Y., Wang, Y., & Li, Y. (2025) · Journal of Experimental Psychology Learning Memory and Cognition · doi:10.1037/xlm0001553 · Cited by 2 · see citing works
Self-citationTricking our brains to learn and remember; is all learning incidental?
Seitz, A. R. (2025) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2025.103020 · Cited by 1 · see citing works
The causal involvement of the visual cortex in visual working memory remains uncertain
Grassi, P. R., Bannert, M. M., & Bartels, A. (2024) · Royal Society Open Science · doi:10.1098/rsos.231884 · Cited by 1 · see citing works
The asymmetry of working memory training transfer: A systematic review and meta-analysis
Fu, J., Zhou, Y., Yuan, M., & Cai, Y. (2026) · Psychonomic Bulletin & Review · doi:10.3758/s13423-026-02994-5
“Consistent with this interpretation, transfer has been observed from high-precision visual perception training to relatively low-precision WM tasks (Jia et al., 2021), but not in the reverse direction (Cai et al., 2022).”
Self-medication for Cough: a Study of Smokers and Non-smokers in Surabaya Colleges
Lorensia, A., Suryadinata, R. V., Hayati, S. N., & Suud, S. (2024) · Global Medical & Health Communication (GMHC) · doi:10.29313/gmhc.v12i2.12367
Asymmetric transfer between the learning of the complex stimulus
Du, Y., Kou, H., Liu, H., & Bi, T. (2025) · Frontiers in Neuroscience · doi:10.3389/fnins.2025.1578862
Vectorial boundary representation of 3D layout in human visual cortexPreprint
Wu, Y., & Li, S. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.11.17.688791
Self-citationPerceptual learning reformats working memory codes along a reverse visual hierarchyPreprint
Cheng, S., Ge, Y., & Chen, N. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.03.20.712287
Wu, Y., & Li, S. (2026) · PLoS Biology · doi:10.1371/journal.pbio.3003541
Self-citationJin, Y., Zhen, Z., Hua, R., Ding, Y., Wang, Y., & Li, Y. W. (2026) · Frontiers in Neuroscience · doi:10.3389/fnins.2026.1794676
Self-citationEditorial: Advances in perceptual learning: new directions, techniques, and applications
He, Q., Huang, C., Chen, J., & Thompson, B. (2026) · Frontiers in Neuroscience · doi:10.3389/fnins.2026.1935934
- Authors
- Jia, K., Li, Y., Gong, M., Huang, H., Wang, Y., & Li, S.
- Affiliations
- Peking University (CN)
- Shaanxi Normal University (CN)
- Zhejiang University (CN)
- Corresponding author
- Li, Sheng
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2021-04-02
- Volume
- 41
- Issue
- 20
- Pages
- 4476-4486
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC8152602
- OpenAlex
- W3146276902
- Cited by
- 25
- References
- 105
- Citations by year
- 2026: 4 · 2025: 7 · 2024: 5 · 2023: 4 · 2022: 3 · 2021: 2
- FWCI
- 1.84
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Journal of Neuroscience (publishedVersion, cc-by-nc-sa)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Key R&D Program of China (2017YFB1002503)
- National Natural Science Foundation of China (31271081, 31230029, 31800910, 32000784)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Visual perception and processing mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural and Behavioral Psychology Studies
- Keywords
- perceptual learning, primary visual cortex, intraparietal sulcus, orientation discrimination, experience-dependent plasticity, neural decoding, transcranial magnetic stimulation, memory maintenance, visual working memory
- MeSH terms
- Adolescent, Adult, Female, Humans, Learning, Magnetic Resonance Imaging, Male, Memory, Short-Term, Parietal Lobe, Visual Cortex, Visual Perception, Transcranial Magnetic Stimulation, Young Adult
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.
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