1. Centre for the Biology of Memory, Norwegian University of Science and Technology, 7489 Trondheim, Norway
2. *These authors contributed equally to this work
3. †Present address: Department of Physiology, University of Oslo, PO Box 1103 Blindern, 0317 Oslo, Norway

Correspondence to: Edvard I. Moser¹ (Email: edvard.moser@ntnu.no).

Abstract

The ability to find one's way depends on neural algorithms that integrate information about place, distance and direction, but the implementation of these operations in cortical microcircuits is poorly understood. Here we show that the dorsocaudal medial entorhinal cortex (dMEC) contains a directionally oriented, topographically organized neural map of the spatial environment. Its key unit is the 'grid cell', which is activated whenever the animal's position coincides with any vertex of a regular grid of equilateral triangles spanning the surface of the environment. Grids of neighbouring cells share a common orientation and spacing, but their vertex locations (their phases) differ. The spacing and size of individual fields increase from dorsal to ventral dMEC. The map is anchored to external landmarks, but persists in their absence, suggesting that grid cells may be part of a generalized, path-integration-based map of the spatial environment.

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