The entorhinal grid map is discretized


The medial entorhinal cortex (MEC) is part of the brain’s circuit for dynamic representation of self-location. The metric of this representation is provided by grid cells, cells with spatial firing fields that tile environments in a periodic hexagonal pattern. Limited anatomical sampling has obscured whether the grid system operates as a unified system or a conglomerate of independent modules. Here we show with recordings from up to 186 grid cells in individual rats that grid cells cluster into a small number of layer-spanning anatomically overlapping modules with distinct scale, orientation, asymmetry and theta-frequency modulation. These modules can respond independently to changes in the geometry of the environment. The discrete topography of the grid-map, and the apparent autonomy of the modules, differ from the graded topography of maps for continuous variables in several sensory systems, raising the possibility that the modularity of the grid map is a product of local self-organizing network dynamics.

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Figure 1: Step-like increases in grid scale along the entorhinal dorsoventral axis.
Figure 2: Comodular organization of grid orientation and grid scale.
Figure 3: Distortions in grid shape are comodular with grid spacing.
Figure 4: Grid modules are organized as overlapping horizontal bands.
Figure 5: Scale relationship between grid modules.
Figure 6: Comodularity between grid spacing and theta frequency.
Figure 7: Grid modules are functionally independent.


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We thank A. M. Amundgård, K. Haugen, K. Jenssen, E. Kråkvik, R. Skjerpeng, and H. Waade for technical assistance, J. Whitlock for implanting an animal in the tangential group, and T. Bonhoeffer, B. Dunn, B. McNaughton, Y. Roudi, A. Treves and A. Witoelar for helpful discussion. The work was supported by an Advanced Investigator Grant from the European Research Council (‘CIRCUIT’, Grant Agreement no. 232608), the Kavli Foundation and the Centre of Excellence scheme of the Research Council of Norway.

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T.St., H.S., T.So., M.-B.M. and E.I.M. designed experiments and analyses; H.S., T.St. and T.So. implanted tetrodes; H.S. recorded multisite data; T.So. and K.F. tested animals with tangential implants; T.St. performed the majority of the analyses; and T.St. and E.I.M. wrote the paper with input from all authors.

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Correspondence to Hanne Stensola or Tor Stensola or Edvard I. Moser.

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The authors declare no competing financial interests.

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Stensola, H., Stensola, T., Solstad, T. et al. The entorhinal grid map is discretized. Nature 492, 72–78 (2012).

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