The DOI matches its Crossref record; 222 of the 227 references it cites were found in a registry or an index, and 1 are not checked yet.
Pattern separation, completion, and categorisation in the hippocampus and neocortex
Abstract
The mechanisms for pattern completion and pattern separation are described in the context of a theory of hippocampal function in which the hippocampal CA3 system operates as a single attractor or autoassociation network to enable rapid, one-trial, associations between any spatial location (place in rodents, or spatial view in primates) and an object or reward, and to provide for completion of the whole memory during recall from any part. The factors important in the pattern completion in CA3 and also a large number of independent memories stored in CA3 include: a sparse distributed representation, representations that are independent due to the randomizing effect of the mossy fibres, heterosynaptic long-term depression as well as long-term potentiation in the recurrent collateral synapses, and diluted connectivity to minimize the number of multiple synapses between any pair of CA3 neurons which otherwise distort the basins of attraction. Recall of information from CA3 is implemented by the entorhinal cortex perforant path synapses to CA3 cells, which in acting as a pattern associator allow some pattern generalization. Pattern separation is performed in the dentate granule cells using competitive learning to convert grid-like entorhinal cortex firing to place-like fields, and in the dentate to CA3 connections that have diluted connectivity. Recall to the neocortex is achieved by a reverse hierarchical series of pattern association networks implemented by the hippocampo-cortical backprojections, each one of which performs some pattern generalization, to retrieve a complete pattern of cortical firing in higher-order cortical areas. New results on competitive networks show which factors contribute to their ability to perform pattern separation, pattern clustering, and pattern categorisation, and how these apply in different hippocampal and neocortical systems.
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“This feature seems to depend on a pattern completion mechanism that is able to reinstate complete memories after partial cueing, based on the operation of hippocampal autoassociative neural networks”
From Spatial Cognition in Teleost Fish: Strategies and Mechanisms (2021) · cites this paper as [81]
Citations
Cites
Neural networks and physical systems with emergent collective computational abilities.
Hopfield, J. J. (1982) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.79.8.2554 · Cited by 19,270
The magical number seven, plus or minus two: Some limits on our capacity for processing information.
Miller, G. A. (1956) · Psychological Review · doi:10.1037/h0043158 · Cited by 17,812
Plum, F. (1960) · Archives of Neurology · doi:10.1001/archneur.1960.03840090124020 · Cited by 10,388
Self-Organization and Associative Memory
Kohonen, T. (1989) · Springer Series in Information Sciences · doi:10.1007/978-3-642-88163-3 · Cited by 8,740
(1978) · UA Campus Repository (The University of Arizona) · Cited by 8,010
Introduction to the theory of neural computation
Hertz, J., Krogh, A., & Palmer, R. G. (1994) · Neural Networks · doi:10.1016/0893-6080(94)90014-0 · Cited by 6,410
McClelland, J. L., McNaughton, B. L., & O'Reilly, R. C. (1995) · Psychological Review · doi:10.1037/0033-295x.102.3.419 · Cited by 5,256
Microstructure of a spatial map in the entorhinal cortex
Hafting, T., Fyhn, M., Molden, S., Moser, M.-B., & Moser, E. I. (2005) · Nature · doi:10.1038/nature03721 · Cited by 4,068
Self-Organization and Associative Memory
Kohonen, T. (1988) · Springer series in information sciences · doi:10.1007/978-3-662-00784-6 · Cited by 3,843
Marr, D. (1969) · The Journal of Physiology · doi:10.1113/jphysiol.1969.sp008820 · Cited by 3,253
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4 of the 227 references couldn’t be matched to a known paper.
Research works printed but not linked: 3
- McNaughton, 1991, Associative pattern completion in hippocampal circuits: New evidence and new questions, Brain Research Reviews, 16, 193
- Rolls, 1995, A model of the operation of the hippocampus and entorhinal cortex in memory, International Journal of Neural Systems, 6, 51
- Witter, 2000, Cortico-hippocampal communication by way of parallel parahippocampal-subicular pathways, Hippocampus, 10, 398, 10.1002/1098-1063(2000)10:4<398::aid-hipo6>3.0.co;2-k · DOI
Cited by
Sorted by citations
Items 201–231 of 231
Development of memory generalization and specificity in childhood
Ngo, Z. T. (2023) · Brain and Cognition · doi:10.1016/j.bandc.2023.106019
Wahlheim, C. N., Garlitch, S. M., Mohamed, R. M., & Weidler, B. J. (2023) · doi:10.31234/osf.io/tfrh8
Effects of focal epilepsy and aging on discrimination and reactivation of memoriesPreprint
Tai, X. Y., Sen, A., Husain, M., & Manohar, S. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.05.13.593984
“Sparseness level of model inputs is an important characteris8c of an autoassocia8ve network and has implica8ons on the network storage capacity 44 .”
Addis, D. R. (2024) · Elsevier eBooks · doi:10.1016/b978-0-12-820480-1.00135-2
Nunes, A., Singh, S., Khayachi, A., Stern, S., Trappenberg, T. P., & Alda, M. (2025) · Brain and Behavior · doi:10.1002/brb3.70209
Field, T. A., & Ghoston, M. R. (2025) · doi:10.1002/9781394347230.ch9
A model for how grid cells contribute to complementary concept representations in the hippocampusPreprint
Houser, T. M. (2025) · Research Square · doi:10.21203/rs.3.rs-7708281/v1
Zhang, R., An, G., Hao, Y., & Wu, D. O. (2026) · IEEE Transactions on Image Processing · doi:10.1109/tip.2025.3650668
“The retrieval of a whole representation is initiated by the activation of some small part of the same representation (the recall cue) [23].”
Subregional activity in the dentate gyrus is amplified during elevated cognitive demandsPreprint
Castillon, C., Otsuka, S., Armstrong, J. N., & Contractor, A. (2026) · eLife · doi:10.7554/elife.109611.1
Sun, L., Liu, Q., Li, S., Ding, W., Li, Z., Li, K. X., Qu, W., & Liang, X. (2026) · Communications Biology · doi:10.1038/s42003-026-09554-6
“In contrast, CA3 and CA1 sub fi elds are more involved in pattern completion, which facilitates the retrieval of complete memories or contextual regularities from partial cues 22 – 24 .”
Age-related impairments in scene-based mnemonic pattern separation
Zhang, M., Yu, Q., Wu, T., Li, Y. J., Wen, W., & Deng, J. (2026) · Behavioural Brain Research · doi:10.1016/j.bbr.2026.116106
When and Where: A Model Hippocampal Network Unifies Formation of Time Cells and Place CellsPreprint
Yu, Q. S., Wang, Z., & Balasubramanian, V. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.03.22.713480
The neural code of perceptual inference
Shin, H. (2026) · Molecular Brain · doi:10.1186/s13041-026-01299-x
“More broadly, pattern separation and pattern completion can serve as the neural mechanism for discrimination and generalization, respectively [64].”
Subregional activity in the dentate gyrus is amplified during elevated cognitive demandsPreprint
Castillon, C., Otsuka, S., Armstrong, J. N., & Contractor, A. (2026) · eLife · doi:10.7554/elife.109611.2
Cupolillo, D. (2021) · theses.fr (ABES) · doi:10.70675/74419524zf753z4b53z850dz04482581bb2f · Cited by 2
Presynaptic plasticity and memory encoding in hippocampal circuits
Marneffe, C. (2022) · theses.fr (ABES) · doi:10.70675/2220ff87zc90dz425bza48fz15208b07595e
Identifying distinct ensembles recruited during memory acquisition to form the engram
Pouget, C. (2024) · theses.fr (ABES) · doi:10.70675/339fe468z82fbz4f38zbe9azb44f1082d215
Dynamic shifts between pattern separation and reinstatement in human CA3 and dentate gyrusPreprint
Gellersen, H. M., Naspi, L., Przygodda, X., Düzel, E., & Berron, D. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.04.24.720552 · Cited by 1
Computational mechanisms transforming visual codes into sparse representations
Cao, R., & Wang, S. (2026) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2026.106782
[Andreas M. Brandmaier and Markus Werkle-Bergner contributed equally to this work.]
Buchberger, E., Ngo, C., Peikert, A., Brandmaier, A., & Werkle‐Bergner, M. (2024) · MPG.PuRe (Max Planck Society)
Enhanced Contour‐Deviant Mismatch Negativity and Mnemonic Representations in Older Musicians
Bugos, J. A., Chow, R., Mo, S., Rosenbaum, R. S., & Alain, C. (2026) · European Journal of Neuroscience · doi:10.1111/ejn.70576
“…of MMN to abstract, complex irregularities, such as omission deviants within expected tone sequences (Chennu et al. 2016; Phillips et al. 2015, 2016; see Heilbron and Chait 2018), which reflect higher-order regularity violations, comparable with complex pitch contour violations used in the…”
Hippocampal prediction errors and claustral control at the perception–memory crossroads
Medina, C., Fernández, R. S., Krawczcyk, M. C., & Boccia, M. M. (2026) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2025.0253 · Cited by 1
Overlapping representations in the ventral hippocampus support rapid fear memory retrieval
Rozeske, R. R., Runtz, L., Lee, J. Q., Keinath, A. T., Sossin, A., & Brandon, M. P. (2026) · Nature Neuroscience · doi:10.1038/s41593-026-02435-5 · Cited by 4
Olfactory Emotion as a Model System for Emotion Measurement
Caricchio, F., Migone, C., Piras, A., Zambito, Y., Pietra, D., Borghini, A., & Fabiano, A. (2026) · Journal of Psychophysiology · doi:10.1027/0269-8803/a000364
Neuroplasticity under constraint: From sensory ecology to neuroinflammation
Diniz, D. G., Bento-Torres, N. V. O., Bento‐Torres, J., Jardim, N. Y. V., Sosthenes, M. C. K., Krejčová, L. V., dos Santos, Z. A., de Sousa, A. A., da Costa, E. R., Pereira, P. D. C., et al. (2026) · Brain Behavior and Immunity Integrative · doi:10.1016/j.bbii.2026.100199
Grigoryan, G. A., & Balaban, P. M. (2026) · Biosystems · doi:10.1016/j.biosystems.2026.105967
Mental Scene and Memory Construction and the Hippocampus
Rolls, E. T. (2026) · Hippocampus · doi:10.1002/hipo.70136
“…circuit in humans (Rogers Flattery et al. 2020) are shown in (b), and indicate very many dentate granule cells, consistent with expansion encoding and the production of sparse uncorrelated representations prior to CA3 (Rolls 2016, 2021a). of the episodic memory by pattern association retrieval.”
Self-citationLöffler, H., & Bahmer, A. (2026) · Research Square · doi:10.21203/rs.3.rs-11140391/v1
Metadata
- Authors
- Rolls, E. T.
- Affiliations
- University of Warwick (GB)
- Corresponding author
- Rolls, Edmund T.
- ORCID
- Rolls, Edmund T.
- Venue
- Neurobiology of Learning and Memory
- Publisher
- Elsevier BV
- Published
- 2016-03
- Volume
- 129
- Pages
- 4-28
- ISSN
- 1074-7427 (ISSN-L)
- OpenAlex
- W889908523
- Semantic Scholar
- dee8fed2414a8b21d9bf5724732f82ca2356190b
- Cited by
- 234
- References
- 227
- Citations by year
- 2026: 28 · 2025: 25 · 2024: 29 · 2023: 24 · 2022: 19 · 2021: 29 · 2020: 18 · 2019: 21 · 2018: 17 · 2017: 15 · 2016: 9
- FWCI
- 8.22
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Neurobiology of Learning and Memory (publishedVersion)
- PubMed (publishedVersion)
- Funders
- Medical Research Council
- Human Frontier Science Program
- EEC BRAIN
- MRC Oxford Interdisciplinary Research Centre in Cognitive Neuroscience
- Oxford McDonnell-Pew Centre in Cognitive Neuroscience
- Oxford Centre for Computational Neuroscience
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Keywords
- pattern separation, pattern completion, hippocampus, CA3, dentate gyrus, neocortex, attractor networks, sparse distributed representations, mossy fibers, long-term potentiation, perforant path, entorhinal cortex, place cells, grid cells, competitive learning, pattern generalization, memory retrieval
- MeSH terms
- Animals, Association Learning, Humans, Models, Neurological, Neural Pathways, Neurons, Mental Recall, Synapses, Neocortex, Pattern Recognition, Physiological, CA3 Region, Hippocampal
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