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Memory Consolidation
Open accessAbstract
Conscious memory for a new experience is initially dependent on information stored in both the hippocampus and neocortex. Systems consolidation is the process by which the hippocampus guides the reorganization of the information stored in the neocortex such that it eventually becomes independent of the hippocampus. Early evidence for systems consolidation was provided by studies of retrograde amnesia, which found that damage to the hippocampus-impaired memories formed in the recent past, but typically spared memories formed in the more remote past. Systems consolidation has been found to occur for both episodic and semantic memories and for both spatial and nonspatial memories, although empirical inconsistencies and theoretical disagreements remain about these issues. Recent work has begun to characterize the neural mechanisms that underlie the dialogue between the hippocampus and neocortex (e.g., "neural replay," which occurs during sharp wave ripple activity). New work has also identified variables, such as the amount of preexisting knowledge, that affect the rate of consolidation. The increasing use of molecular genetic tools (e.g., optogenetics) can be expected to further improve understanding of the neural mechanisms underlying consolidation.
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“Humans do not treat every new situation as entirely unfamiliar; instead, they gradually summarize common structure from past episodes and organize it into abstractions through memory consolidation (McClelland et al., 1995; Squire et al., 2015; Dudai et al., 2015; Moscovitch et al., 2016; Sun et al., 2023; Bartlett, 1932).”
From Useful Memories Become Faulty When Continuously Updated by LLMs (2026) · cites this paper as Squire et al., 2015
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“Cognitive theories of memory describe two complementary processes: a fast episodic system that stores individual experiences, and a slow process that extracts compact schemas -abstractions over many episodes that integrate with prior knowledge to support future generalization (McClelland et al., 1995; Squire et al., 2015; Dudai et al., 2015; Moscovitch et al., 2016).”
From Useful Memories Become Faulty When Continuously Updated by LLMs (2026) · cites this paper as Squire et al., 2015
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“We observe that humans' performance steadily improves because each experience is (i) encoded as an episodic trace (Pritzel et al., 2017), (ii) distilled into abstract rules during sleep-dependent consolidation (Squire et al., 2015), (iii) selectively reinforced by dopamine-driven credit assignment (Glimcher, 2011), and (iv) retrieved through case- or analogy-based reasoning when similar problems”
From Memento: Fine-tuning LLM Agents without Fine-tuning LLMs (2025) · cites this paper as Squire et al., 2015
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“Human memory relies on complementary learning systems (McClelland et al., 1995), where memory consolidation (Squire et al., 2015) progressively transforms rapid episodic encoding into more stable semantic structures (Cowan et al., 2021).”
From TiMem: Temporal-Hierarchical Memory Consolidation for Long-Horizon Conversational Agents (2026) · cites this paper as Squire et al., 2015
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“Human memory relies on complementary learning systems (McClelland et al., 1995), where memory consolidation (Squire et al., 2015) progressively transforms rapid episodic encoding into more stable semantic structures (Cowan et al., 2021).”
From TiMem: Temporal-Hierarchical Memory Consolidation for Long-Horizon Conversational Agents (2026) · cites this paper as Squire et al., 2015
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Computer Vision Based Two-stage Waste Recognition-Retrieval Algorithm for Waste Classification
Zhang, S., Chen, Y., Yang, Z., & Gong, H. (2021) · Resources Conservation and Recycling · doi:10.1016/j.resconrec.2021.105543 · Cited by 62
Mechanistic flexibility of the retrosplenial cortex enables its contribution to spatial cognition
Stacho, M., & Manahan‐Vaughan, D. (2022) · Trends in Neurosciences · doi:10.1016/j.tins.2022.01.007 · Cited by 67
“Moreover, reports of a gradual development of memory-related activity in the RC [13,16,26,31,32], as well as findings that show that RC inactivation after learning impairs long-term memory [16,33], suggest that the RC might be especially important in the consolidation of memory traces [34].”
BackgroundResultSemantic memory in developmental amnesia
Elward, R. L., & Vargha‐Khadem, F. (2018) · Neuroscience Letters · doi:10.1016/j.neulet.2018.04.040 · Cited by 59
The molecular memory code and synaptic plasticity: A synthesis
Gershman, S. J. (2023) · Biosystems · doi:10.1016/j.biosystems.2022.104825 · Cited by 63
“A standard answer appeals to systems consolidation, the process by which memories are gradually transferred from a temporary store in the hippocampus to a more durable representation in the neocortex (Squire et al., 2015).”
BackgroundBoccia, M., Teghil, A., & Guariglia, C. (2019) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2019.09.003 · Cited by 65
Contributions of post-learning REM and NREM sleep to memory retrieval
MacDonald, K. J., & Côté, K. A. (2021) · Sleep Medicine Reviews · doi:10.1016/j.smrv.2021.101453 · Cited by 66
Ali, M. A., Menze, E. T., Tadros, M., & Tolba, M. F. (2020) · Neuropharmacology · doi:10.1016/j.neuropharm.2020.108334 · Cited by 61
What can we teach Lymnaea and what can Lymnaea teach us?
Rivi, V., Benatti, C., Lukowiak, K., Colliva, C., Alboni, S., Tascedda, F., & Blom, J. M. C. (2021) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/brv.12716 · Cited by 57
“The process by which the transient STM is converted to a more stable LTM is generally referred to as consolidation and is dependent on altered gene activity and new protein synthesis (Squire et al., 2015).”
BackgroundPerez, D. M. (2021) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2021.652152 · Cited by 63
Evans, H. T., Bodea, L., & Götz, J. (2020) · eLife · doi:10.7554/elife.52990 · Cited by 55
Cassel, J., Ferraris, M., Quilichini, P., Cholvin, T., Boch, L., Stéphan, A., & de Vasconcelos, A. P. (2021) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2021.03.023 · Cited by 56
Hrybouski, S., MacGillivray, M., Huang, Y., Madan, C. R., Carter, R., Seres, P., & Malykhin, N. V. (2019) · NeuroImage · doi:10.1016/j.neuroimage.2019.01.061 · Cited by 56
“Performing odd/even judgments between the two task phases not only limits rote rehearsal, but also produces more accurate estimates of memory-related BOLD activation, particularly inside the HC formation (Stark & Squire, 2001).”
BackgroundCausal Contribution of Awake Post-encoding Processes to Episodic Memory Consolidation
Tambini, A., & D’Esposito, M. (2020) · Current Biology · doi:10.1016/j.cub.2020.06.063 · Cited by 54
Tortella, G., Seabra, A. B., Padrão, J., & Juan, R. D.-S. (2021) · Brain Sciences · doi:10.3390/brainsci11050552 · Cited by 66
“The term “memory consolidation” is closely related to the previous term, referring to the necessary process by which a labile or temporary memory can be transformed into long-term memory [27].”
BackgroundSmolen, P., Baxter, D. A., & Byrne, J. H. (2019) · Learning & Memory · doi:10.1101/lm.049395.119 · Cited by 56
“4 This process is referred to as systems consolidation (Frankland and Bontempi 2005; Haist et al. 2001; Kirwan et al. 2008; McGaugh 2000; Nadel and Moscovitch 1997; Squire et al. 2015) and can occur, in part, on a rapid time scale of ~1-2 h (Brodt et al.”
BackgroundA theory of hippocampal function: New developments
Rolls, E. T., & Treves, A. (2024) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2024.102636 · Cited by 54
“…memory systems (McClelland et al., 1995; McClelland et al., 2020), and evidence from for example the possible temporal gradient of retrograde amnesia following hippocampal damage (Scoville and Milner, 1957; Squire, 1992; Squire et al., 2015; Fernandez and Morris, 2018; Moscovitch and Gilboa, 2022).”
Network-wide abnormalities explain memory variability in hippocampal amnesia
Argyropoulos, G. P. D., Loane, C., Roca‐Fernández, A., Lage-Martinez, C., Gurau, O., Irani, S. R., & Butler, C. (2019) · eLife · doi:10.7554/elife.46156 · Cited by 50
“The ‘standard consolidation theory’ holds that newly-acquired memories initially depend upon the HPC but that, through a process of ‘consolidation’, memory traces gradually become HPCindependent and are stored in and retrieved by neocortical structures alone (Hardt and Nadel, 2018; Squire et al., 2015).”
BackgroundResultUgolini, F., Lana, D., Nardiello, P., Nosi, D., Pantano, D., Casamenti, F., & Giovannini, M. G. (2018) · Frontiers in Aging Neuroscience · doi:10.3389/fnagi.2018.00372 · Cited by 50
Emerging neurodevelopmental perspectives on mathematical learning
Menon, V., & Chang, H. (2021) · Developmental Review · doi:10.1016/j.dr.2021.100964 · Cited by 62
“…subdivision, in learning and memory consolidation for events in space and time is well known (Davachi, 2006; Diana, Yonelinas, & Ranganath, 2007; Eichenbaum, Yonelinas, & Ranganath, 2007; Squire, 1992; Squire, Stark, & Clark, 2004; Tulving, 1983; Davachi et al., 2003; Squire et al., 2015).”
BackgroundMemory Retrieval in Mice and MenPreprint
Ben-Yakov, A., Dudai, Y., & Mayford, M. R. (2015) · Cold Spring Harbor Perspectives in Biology · doi:10.1101/cshperspect.a021790 · Cited by 54
“What is the nature of this circuit-based consolidation of memory (“systems consolidation”) (Squire et al. 2015)? What is the anatomical locus of hippocampaldependent new memories and hippocampalindependent older, consolidated, memories? What is the role of the hippocampus and cortex in the encoding and retrieval of ne…”
Hypertension is associated with reduced hippocampal connectivity and impaired memory
Feng, R., Rolls, E. T., Cheng, W., & Feng, J. (2020) · EBioMedicine · doi:10.1016/j.ebiom.2020.103082 · Cited by 49
“Given this discovery, we next analysed whether hypertension was associated with memory and other cognitive problems, as the hippocampus is implicated in memory [32-35].”
BackgroundTask-dependent mixed selectivity in the subiculum
Ledergerber, D., Battistin, C., Blackstad, J. S., Gardner, R. J., Witter, M. P., Moser, M., Roudi, Y., & Moser, E. I. (2021) · Cell Reports · doi:10.1016/j.celrep.2021.109175 · Cited by 53
Hippocampal area CA1 and remote memory in ratsPreprint
Ocampo, A. C., Squire, L. R., & Clark, R. E. (2017) · Learning & Memory · doi:10.1101/lm.045781.117 · Cited by 51
“Rats with hippocampal lesions have consistently exhibited severe, ungraded retrograde memory impairment in this task, with remote memory as severely impaired as recent memory (Bolhuis et al. 1994; Mumby et al. 1999; Sutherland et al. 2001; Clark et al. 2005a).”
Self-citationA double dissociation between savings and long-term memory in motor learning
Hadjiosif, A. M., Morehead, J. R., & Smith, M. A. (2023) · PLoS Biology · doi:10.1371/journal.pbio.3001799 · Cited by 49
Understanding the dynamic and destiny of memories
Alvares, L. D. O., & Monte, F. H. D. (2021) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2021.03.009 · Cited by 51
Early- and Late-Onset Alzheimer’s Disease: Two Sides of the Same Coin?
Valdez-Gaxiola, C. A., Rosales-Leycegui, F., Gaxiola-Rubio, A., Moreno-Ortíz, J. M., & Figuera, L. E. (2024) · Diseases · doi:10.3390/diseases12060110 · Cited by 55
“This section provides a comprehensive comparison of the diagnostic and assessment strategies used in both EOAD and LOAD, taking into account various factors, including neuroimaging techniques, biomarkers, and cognitive assessments [73,74].”
MethodSugden, A. U., Zaremba, J. D., Sugden, L. A., McGuire, K. L., Lutas, A., Ramesh, R. N., Alturkistani, O., Lensjø, K. K., Burgess, C. R., & Andermann, M. L. (2020) · Nature Neuroscience · doi:10.1038/s41593-020-0651-5 · Cited by 43
Naïve to expert: Considering the role of previous knowledge in memory
Alonso, A., van der Meij, J., Tse, D., & Genzel, L. (2020) · Brain and Neuroscience Advances · doi:10.1177/2398212820948686 · Cited by 46
“But over time (weeks/months/years), cortical connections become reinforced in offline consolidation processes so that later on they are sufficient for retrieval (McClelland et al., 1995; Squire et al., 2015).”
BackgroundSelf-citationVirtual reality modulating dynamics of neuroplasticity: Innovations in neuro-motor rehabilitation
Wankhede, N. L., Koppula, S., Ballal, S., Doshi, H., Kumawat, R., Raju, S., Arora, I., Sammeta, S. S., Khalid, M., Zafar, A., et al. (2024) · Neuroscience · doi:10.1016/j.neuroscience.2024.12.040 · Cited by 52
Engram stability and maturation during systems consolidation
Refaeli, R., Kreisel, T., Groysman, M., Adamsky, A., & Goshen, I. (2023) · Current Biology · doi:10.1016/j.cub.2023.07.042 · Cited by 46
Metadata
- Authors
- Squire, L. R., Genzel, L., Wixted, J. T., & Morris, R. G.
- Affiliations
- University of California San Diego (US)
- University of Edinburgh (GB)
- Venue
- Cold Spring Harbor Perspectives in Biology
- Publisher
- Cold Spring Harbor Laboratory
- Published
- 2015-08
- Volume
- 7
- Issue
- 8
- Pages
- a021766
- ISSN
- 1943-0264 (ISSN-L)
- PMC
- PMC4526749
- OpenAlex
- W4247532959
- Cited by
- 700
- References
- 148
- Citations by year
- 2026: 70 · 2025: 97 · 2024: 100 · 2023: 78 · 2022: 83 · 2021: 83 · 2020: 68 · 2019: 45 · 2018: 26 · 2017: 27 · 2016: 14 · 2015: 6
- Type
- posted content
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Cold Spring Harbor Perspectives in Biology (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- U.S. Department of Veterans Affairs
- U.S. Public Health Service (24600)
- National Institute of Mental Health (24600, Grant 24600)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory Processes and Influences
- Life Sciences › Neuroscience › Cognitive Neuroscience › Sleep and Wakefulness Research
- Keywords
- memory consolidation, systems consolidation, hippocampus, neocortex, retrograde amnesia, episodic memory, semantic memory, spatial memory, neural replay, sharp-wave ripples, optogenetics
- MeSH terms
- Animals, Hippocampus, Humans, Memory, Neocortex
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