Publications and Research

Document Type


Publication Date



Grid cells in the brain respond when an animal occupies a periodic lattice of ‘grid fields’ during navigation. Grids are organized in modules with different periodicity. We propose that the grid system implements a hierarchical code for space that economizes the number of neurons required to encode location with a given resolution across a range equal to the largest period. This theory predicts that (i) grid fields should lie on a triangular lattice, (ii) grid scales should follow a geometric progression, (iii) the ratio between adjacent grid scales should be √e for idealized neurons, and lie between 1.4 and 1.7 for realistic neurons, (iv) the scale ratio should vary modestly within and between animals. These results explain the measured grid structure in rodents. We also predict optimal organization in one and three dimensions, the number of modules, and, with added assumptions, the ratio between grid periods and field widths.


This article was originally published in eLife on September 3, 2015 (4:e08362) under a Creative Commons Attribution License.




To view the content in your browser, please download Adobe Reader or, alternately,
you may Download the file to your hard drive.

NOTE: The latest versions of Adobe Reader do not support viewing PDF files within Firefox on Mac OS and if you are using a modern (Intel) Mac, there is no official plugin for viewing PDF files within the browser window.