The Tolman-Eichenbaum Machine: Unifying Space and Relational Memory through Generalization in the Hippocampal Formation

Cell. 2020 Nov 25;183(5):1249-1263.e23. doi: 10.1016/j.cell.2020.10.024. Epub 2020 Nov 11.


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.

Publication types

  • Research Support, Non-U.S. Gov't

MeSH terms

  • Animals
  • Entorhinal Cortex / physiology*
  • Generalization, Psychological*
  • Hippocampus / physiology*
  • Knowledge
  • Memory / physiology*
  • Models, Neurological*
  • Place Cells / cytology
  • Sensation
  • Task Performance and Analysis