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Memory engrams: Recalling the past and imagining the future

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Josselyn, Sheena A., Tonegawa, Susumu·2020·Science

DOI 10.1126/science.aaw4325PMC PMC7577560

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Abstract

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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The Organization of Behavior
Hebb, D. O. (2005) · doi:10.4324/9781410612403 · Cited by 9,825
The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat
O'Keefe, J., & Dostrovsky, J. (1971) · Brain Research · doi:10.1016/0006-8993(71)90358-1 · Cited by 6,151
Millisecond-timescale, genetically targeted optical control of neural activity
Boyden, E. S., Zhang, F., Bamberg, E., Nagel, G., & Deisseroth, K. A. (2005) · Nature Neuroscience · doi:10.1038/nn1525 · Cited by 5,015
The memory function of sleep
Diekelmann, S., & Born, J. (2010) · Nature Reviews Neuroscience · doi:10.1038/nrn2762 · Cited by 3,760
Reactivation of hippocampal ensemble memories during sleep
Wilson, M. A., & McNaughton, B. L. (1994) · Science · doi:10.1126/science.8036517 · Cited by 3,118
Stimulation of 3T3 cells induces transcription of the c-fos proto-oncogene
Greenberg, M. E., & Ziff, E. B. (1984) · Nature · doi:10.1038/311433a0 · Cited by 2,992
Synaptic Plasticity and Memory: An Evaluation of the Hypothesis
Martin, S. J., Grimwood, P. D., & Morris, R. G. M. (2000) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.23.1.649 · Cited by 2,794
Simple memory: a theory for archicortex
Marr, D. (1971) · Philosophical Transactions of the Royal Society of London. B, Biological Sciences · doi:10.1098/rstb.1971.0078 · Cited by 2,771
Fear memories require protein synthesis in the amygdala for reconsolidation after retrieval
Nader, K., Schafe, G. E., & Le Doux, J. E. (2000) · Nature · doi:10.1038/35021052 · Cited by 2,706
Modality-Specific Retrograde Amnesia of Fear
Kim, J. J., & Fanselow, M. S. (1992) · Science · doi:10.1126/science.1585183 · Cited by 2,483

Flagged first, then by citations

Show all 240 references

Research works printed but not linked: 7

K. S. Lashley in Society of Experimental Biology Symposium No. 4: Psychological Mechanisms in Animal Behavior J. F. Danielli R. Brown Eds. (Academic Press 1950) pp. 454–482.
M. Moscovitch in Science of Memory: Concepts H. L. I. Roediger Y. Dudai S. M. Fitzpatrick Eds. (Oxford Univ. Press 2007) pp. 17–29.
M. S. Fanselow L. S. Lester in Evolution and Learning R. C. Bolles M. D. Beecher Eds. (Lawrence Erlbaum Associates Inc. 1988) pp. 185–212.
P. M. Milner, Cell assemblies: Whose idea? Psycholoquy 10, 1 (1999).
I. Pavlov Conditioned Reflexes (Oxford Univ. Press 1927).
10.1016/j.cub.2012.11.019 · DOI
H. B. Barlow in The Cognitive Neurosciences M. S. Gazzaniga Ed. (MIT Press 1995) pp. 415–435.

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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
ORCID: Josselyn, Sheena A.
Tonegawa, Susumu
Venue: Science
Publisher: American Association for the Advancement of Science (AAAS)
Published: 2020-01-03
Volume: 367
Issue: 6473
ISSN: 0036-8075 (ISSN-L)
1095-9203
DOI: 10.1126/science.aaw4325
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

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Memory engrams: Recalling the past and imagining the future

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