The hippocampus and related neocortical structures in memory transformation
Abstract
Episodic memories are multifaceted and malleable, capable of being transformed with time and experience at both the neural level and psychological level. At the neural level, episodic memories are transformed from being dependent on the hippocampus to becoming represented in neocortical structures, such as the medial prefrontal cortex (mPFC), and back again, while at the psychological level, detailed, perceptually rich memories, are transformed to ones retaining only the gist of an experience or a schema related to it. Trace Transformation Theory (TTT) initially proposed that neural and psychological transformations are linked and proceed in tandem. Building on recent studies on the neurobiology of memory transformation in rodents and on the organization of the hippocampus and its functional cortical connectivity in humans, we present an updated version of TTT that is more precise and detailed with respect to the dynamic processes and structures implicated in memory transformation. At the heart of the updated TTT lies the long axis of the hippocampus whose functional differentiation and connectivity to neocortex make it a hub for memory formation and transformation. The posterior hippocampus, connected to perceptual and spatial representational systems in posterior neocortex, supports fine, perceptually rich, local details of memories; the anterior hippocampus, connected to conceptual systems in anterior neocortex, supports coarse, global representations that constitute the gist of a memory. Notable among the anterior neocortical structures is the medial prefrontal cortex (mPFC) which supports representation of schemas that code for common aspects of memories across different episodes. Linking the aHPC with mPFC is the entorhinal cortex (EC) which conveys information needed for the interaction/translation between gist and schemas. Thus, the long axis of the hippocampus, mPFC and EC provide the representational gradient, from fine to coarse and from perceptual to conceptual, that can implement processes implicated in memory transformation. Each of these representations of an episodic memory can co-exist and be in dynamic flux as they interact with one another throughout the memory's lifetime, going from detailed to schematic and possibly back again, all mediated by corresponding changes in neural representation.
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In young adults, perception of the childhood odorant leads to increased functional connectivity in the memory system, including dorsal hippocampus, prefrontal cortex, orbitofrontal cortex, and olfactory bulb.
“the perception of the childhood odorant leads to an increased connectivity in the memory system, notably structures typically involved in autobiographical memory—the dorsal hippocampus, the prefrontal cortex—and olfactory memory—the orbitofrontal cortex and the olfactory bulb—both in humans and rodents [9,43,44].”
From Positive early-life olfactory memory is rooted in the olfactory bulb and triggers large-scale changes beyond the olfactory system (2026) · cites this paper as [44]
These findings challenge the Multiple Trace Theory, which posits that the dorsal hippocampus is constantly involved in episodic memory recall.
“These findings challenge the Multiple Trace Theory, which posits that the dorsal hippocampus is constantly involved in episodic memory recall [44,71].”
From Positive early-life olfactory memory is rooted in the olfactory bulb and triggers large-scale changes beyond the olfactory system (2026) · cites this paper as [44]
Some spatial memories retain their detailed HPC representations in perpetuity, along with their schematic representation, and these representations interact with each other, with one or the other dominating depending on conditions including time since acquisition, task demands and retrieval cues.
“However, some spatial memories retain their detailed HPC representations in perpetuity, along with their schematic representation, and these representations interact with each other, with one or the other dominating depending on various conditions, including time since acquisition, task demands and retrieval cues 92,93 .”
From From cognitive maps to spatial schemas (2023) · cites this paper as [93]
According to trace transformation theory, the hippocampus and distributed cortical regions encode distinct aspects of memory and engage in dynamic, bidirectional interactions throughout memory formation and retrieval.
“According to trace transformation theory, the hippocampus and distributed cortical regions encode distinct aspects of memory and engage in dynamic, bidirectional interactions throughout memory formation and retrieval 19 .”
From Role of the hippocampus in systems consolidation of remote fear memory (2026) · cites this paper as [19]
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Self-citationRole of the hippocampus in remembering the past and imagining the future
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Patterns of Autobiographical Memory Loss in Medial-Temporal Lobe Amnesic Patients
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Self-citationSchema representation in patients with ventromedial PFC lesions
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Hippocampal contributions to recollection in retrograde and anterograde amnesia
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Effects of Prior-Knowledge on Brain Activation and Connectivity During Associative Memory Encoding
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Investigating the functions of subregions within anterior hippocampus
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Self-citationThe Emergence of Knowledge and How it Supports the Memory for Novel Related Information
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Cognitive Neuroscience of Memory Consolidation
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Memory consolidation, retrograde amnesia, and the temporal lobe
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The perceptual richness of complex memory episodes is compromised by medial temporal lobe damage
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Mechanisms of Memory Consolidation and Transformation
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The retrieval of perceptual memory details depends on right hippocampal integrity and activation
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Self-citationOrbitofrontal Reality Filtering
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Familiar real-world spatial cues provide memory benefits in older and younger adults.
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Self-citationGeneralizable knowledge outweighs incidental details in prefrontal ensemble code over time
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Autobiographical memory, future imagining, and the medial temporal lobe
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Recollection versus strength as the primary determinant of hippocampal engagement at retrieval
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Self-citationMemory allocation and integration in rodents and humans
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A study of remote spatial memory in aged rats
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Self-citationMemory Distortion: From Misinformation to Rich False Memory.
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The hippocampal sharp wave-ripple in memory retrieval for immediate use and consolidation
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Self-citationMemory consolidation as an adaptive process
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From cognitive maps to spatial schemas
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From remembering to reconstruction: The transformative neural representation of episodic memory.
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Pathways for Contextual Memory: The Primate Hippocampal Pathway to Anterior Cingulate Cortex
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Tracking the relation between gist and item memory over the course of long-term memory consolidation
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Self-citationDistinct brain networks for remote episodic memory depending on content and emotional experience
Auguste, A., Fourcaud-Trocmé, N., Meunier, D., Gros, A., Garcia, S., Messaoudi, B., Thevenet, M., Ravel, N., & Veyrac, A. (2023) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2023.102422 · Cited by 14 · see citing works
Decoding episodic autobiographical memory in naturalistic virtual reality
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Ziegler, M. G., Liu, Z.-X., Arsenault, J., Dang, C., Grady, C., Rosenbaum, R. S., & Moscovitch, M. (2023) · Cerebral Cortex · doi:10.1093/cercor/bhad272 · Cited by 6 · see citing works
Self-citationDejou, J., Athanassi, A., Brunel, T., Thevenet, M., Didier, A., & Mandairon, N. (2026) · PLOS Biology · doi:10.1371/journal.pbio.3003845 · Cited by 1 · see citing works
“Interestingly, we observed a well-known subnetwork composed of structures typically involved in autobiographical memory (dHipp-mPFC-OFC) [9,43,44], also including the CPu and MOB which shows the involvement of olfactory modality and may explain part of the emotional potency of childhood olfactory…”
Role of the hippocampus in systems consolidation of remote fear memory
Park, H., & Kaang, B.-K. (2026) · Experimental & Molecular Medicine · doi:10.1038/s12276-026-01680-9 · Cited by 1 · see citing works
- Authors
- Sekeres, M. J., Winocur, G., & Moscovitch, M.
- Affiliations
- Baylor University (US)
- University of Toronto (CA)
- Corresponding author
- Moscovitch, Morris
- Venue
- Neuroscience Letters
- Publisher
- Elsevier BV
- Published
- 2018-07
- Volume
- 680
- Pages
- 39-53
- ISSN
- 0304-3940 (ISSN-L)
- OpenAlex
- W2802343044
- Cited by
- 370
- References
- 181
- Citations by year
- 2026: 37 · 2025: 53 · 2024: 59 · 2023: 47 · 2022: 52 · 2021: 41 · 2020: 39 · 2019: 30 · 2018: 11
- FWCI
- 4.05
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Neuroscience Letters (publishedVersion)
- PubMed (publishedVersion)
- Funders
- Canadian Institutes for Health Research (MGP 6694)
- Natural Sciences and Engineering Research Council and Canadian Institutes of Health Research (CIHR)
- 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 transformation, episodic memory, hippocampus, medial prefrontal cortex, entorhinal cortex, anterior hippocampus, posterior hippocampus, gastrointestinal stromal tumor, perceptual memory, cortical connectivity
- MeSH terms
- Memory Consolidation, Animals, Hippocampus, Humans, Models, Theoretical, Prefrontal Cortex, Neocortex, Memory, Episodic
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