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Memory engrams: Recalling the past and imagining the future
Open accessAbstract
The neural substrate of memory: The ability to form memory is an essential trait that allows learning and the accumulation of knowledge. But what is a memory? There has been a long history of searching for the neuronal substrate that forms memory in the brain, and the emerging view is that ensembles of engram cells explain how memories are formed and retrieved. In a Review, Josselyn and Tonegawa discuss the evidence for engram cells as a substrate of memory, particularly in rodents; what we have learned so far about the features of memory, including memory formation, retrieval over time, and loss; and future directions to understand how memory becomes knowledge. Science , this issue p. eaaw4325
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“These neurons encode the animal’s position in an environment1 and, more generally, events of episodic experience2–4 and have been causally linked to memory functions5–7.”
From Place and behavioral modulation of hippocampal neurons during immobility (2026) · cites this paper as [7]
Experimental — accuracy not yet measuredIn the citing paper
“For decades, small populations of neurons known as engram neurons have been identified as key mediators of memory storage and retrieval 1 – 3 .”
From Role of the hippocampus in systems consolidation of remote fear memory (2026) · cites this paper as [3]
In the citing paper
“Rodent studies have shown that hippocampal engram cells are reactivated during recent memory recall, whereas neocortical engram cells are reactivated during remote memory recall 3 , 5 , 6 .”
From Role of the hippocampus in systems consolidation of remote fear memory (2026) · cites this paper as [3]
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Metadata
- Authors
- Josselyn, S. A., & Tonegawa, S.
- Affiliations
- Canadian Institute for Advanced Research (CA)
- Howard Hughes Medical Institute (US)
- Corresponding author
- Josselyn, Sheena A.
- Tonegawa, Susumu
- Venue
- Science
- Published
- 2020-01-03
- Volume
- 367
- Issue
- 6473
- ISSN
- 0036-8075 (ISSN-L)
- 1095-9203
- PMC
- PMC7577560
- OpenAlex
- W2998470023
- Cited by
- 1,087
- References
- 240
- Citations by year
- 2026: 165 · 2025: 224 · 2024: 199 · 2023: 159 · 2022: 157 · 2021: 111 · 2020: 70
- FWCI
- 16.83
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Science (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- DSpace@MIT (Massachusetts Institute of Technology) (submittedVersion, cc-by-nc-sa)
- Funders
- Howard Hughes Medical Institute
- JPB Foundation
- Canadian Institute for Advanced Research
- RIKEN Brain Science Institute
- Canadian Institutes of Health Research (FDN-388455)
- NIMH (1 R01 MH119421-01)
- NSERC
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory Processes and Influences
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Keywords
- memory engrams, engram cells, memory storage, memory retrieval, intrinsic excitability, synaptic plasticity, single-unit recording, memory reconsolidation
- MeSH terms
- Animals, Brain, Brain Chemistry, Humans, Memory Disorders, Neurons, Mental Recall
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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