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, James C R Whittington, Timothy H Muller, Shirley Mark, Guifen Chen, Caswell Barry, Neil Burgess, Timothy E J Behrens
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.
Keywords: entorhinal cortex; generalization; grid cells; hippocampus; neural networks; non-spatial reasoning; place cells; representation learning.
Conflict of interest statement
Declaration of Interests The authors declare no competing interests.
Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.
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References
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