Evidence map›Paper›PMID 33181068›Full record

ArticleCell2020

The Tolman-Eichenbaum Machine: Unifying Space and Relational Memory through Generalization in the Hippocampal Formation.

James C R Whittington, Timothy H Muller, Shirley Mark, Guifen Chen, Caswell Barry, Neil Burgess, Timothy E J Behrens

Registry-linked trialAbstract read
In one paragraph

Article in Cell, 2020. The graph could read no effect estimate from its abstract, so it casts no vote on the map. It is linked to trial NCT05746299 (The Emergence of Abstract Structure Knowledge Across Learning and Sleep), which is not on this map. Cited by 257 papers.

0numbers the graph read from it
0cells of the map it votes in
257citing papers in PubMed
–field-weighted citation impact
1 · What the graph read from it

What it found

Each row is one number read from the abstract, on the scale the paper reported it, with its interval. Left of the dashed line favours the treatment, right favours the comparator. Under each row is the sentence it came from. New to these charts? A ten-minute tutorial.

The abstract states no effect estimate the extractor could read, or names no intervention and outcome on the map, so this paper lights no cell and moves no belief. It is still indexed, cited and linked below.

2 · The registry

The trial behind it

Trials whose registry record cites this paper, or whose number appears in the abstract. A trial that started after this paper was published is citing it as background, not reporting it.

NCT05746299 nacompletednot on this mapstarted 2023, after this paper: background citation

The Emergence of Abstract Structure Knowledge Across Learning and Sleep

TypeinterventionalSponsorUniversity of PennsylvaniaRan2023 to 2024Enrolled194ConditionsMemory ConsolidationArmsCongruent vs. Incongruent, Immediate vs. Awake vs. Asleep, Targeted memory reactivation (TMR)
3 · Its place in the literature

Who cites it

257 citing papers in PubMed.

  1. Article
  2. Article
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  4. Positive and Negative Retinotopic Codes in the Human Hippocampus.The Journal of neuroscience : the official journal of the Society for Neuroscience · 2026
    Article
  5. Review
  6. Article
  7. Article
  8. Article
  9. Article
  10. Article
  11. The hippocampus as a generative model.Philosophical transactions of the Royal Society of London. Series B, Biological sciences · 2026
    Review
  12. Article
  13. Article
  14. Article
  15. Planning in the Brain: It's Not What You Think It Is.Annual review of neuroscience · 2026
    Review
  16. Article
  17. Article
  18. Article
  19. Article
  20. Medial temporal lobe encodes cognitive maps of real-world social networks.Proceedings of the National Academy of Sciences of the United States of America · 2026
    Article

197 more citing papers are in PubMed but not listed here.

4 · The record

Corrections and comments

5 · Who and what money

Authors and funding

7 authors.

James C R WhittingtonWellcome Centre for Integrative Neuroimaging, University of Oxford, Oxford OX3 9DU, UK. Electronic address: jcrwhittington@gmail.com.
Timothy H MullerWellcome Centre for Integrative Neuroimaging, University of Oxford, Oxford OX3 9DU, UK; Institute of Neurology, UCL, London WC1N 3BG, UK.
Shirley MarkWellcome Centre for Human Neuroimaging, UCL, London WC1N 3AR, UK.
Guifen ChenInstitute of Cognitive Neuroscience, UCL, London WC1N 3AZ, UK; School of Biological and Chemical Sciences, QMUL, London E1 4NS, UK.
Caswell BarrySainsbury Wellcome Centre for Neural Circuits and Behaviour, UCL, London W1T 4JG, UK; Research department of Cell and Developmental Biology, UCL, London WC1E 6BT, UK.
Neil BurgessInstitute of Neurology, UCL, London WC1N 3BG, UK; Wellcome Centre for Human Neuroimaging, UCL, London WC1N 3AR, UK; Institute of Cognitive Neuroscience, UCL, London WC1N 3AZ, UK; Sainsbury Wellcome Centre for Neural Circuits and Behaviour, UCL, London W1T 4JG, UK.
Timothy E J BehrensWellcome Centre for Integrative Neuroimaging, University of Oxford, Oxford OX3 9DU, UK; Wellcome Centre for Human Neuroimaging, UCL, London WC1N 3AR, UK; Sainsbury Wellcome Centre for Neural Circuits and Behaviour, UCL, London W1T 4JG, UK.

Funding

Medical Research CouncilWellcome TrustWellcome Trust 104765/Z/14/ZWellcome Trust 202805/Z/16/ZWellcome Trust 203139/Z/16/ZWellcome Trust 203147/Z/16/ZWellcome Trust 212281/Z/18/ZWellcome Trust 214314/Z/18/ZWellcome Trust 219525/Z/19/Z
6 · The paper itself

Abstract

The hippocampal-entorhinal system is important for spatial and relational memory tasks. We formally link these domains, provide a mechanistic understanding of the hippocampal role in generalization, and offer unifying principles underlying many entorhinal and hippocampal cell types. We propose medial entorhinal cells form a basis describing structural knowledge, and hippocampal cells link this basis with sensory representations. Adopting these principles, we introduce the Tolman-Eichenbaum machine (TEM). After learning, TEM entorhinal cells display diverse properties resembling apparently bespoke spatial responses, such as grid, band, border, and object-vector cells. TEM hippocampal cells include place and landmark cells that remap between environments. Crucially, TEM also aligns with empirically recorded representations in complex non-spatial tasks. TEM also generates predictions that hippocampal remapping is not random as previously believed; rather, structural knowledge is preserved across environments. We confirm this structural transfer over remapping in simultaneously recorded place and grid cells.

Indexed as

Generalization, PsychologicalModels, NeurologicalAnimalsEntorhinal CortexHippocampusKnowledgeMemoryPlace CellsSensationTask Performance and Analysisentorhinal cortexgeneralizationgrid cellshippocampusneural networksnon-spatial reasoningplace cellsrepresentation learning

Identifiers

PMID33181068
PMCPMC7707106

What OpenQuestion holds

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Registered trials

Read under generation 80e0d062 · epoch 390. Bibliography from PubMed, PubMed Central and OpenAlex; grants from NIH RePORTER; trial links from ClinicalTrials.gov; estimates, votes and beliefs from the OpenQuestion graph.