ABSTRACT:Not all areas of neuronal systems investigation have matured to the stage where computation can be understood at the microcircuit level. In mammals, insights into cortical circuit functions have been obtained for the early stages of sensory systems, where signals can be followed through networks of increasing complexity from the receptors to the primary sensory cortices. These studies have suggested how neurons and neuronal networks extract features from the external world, but how the brain generates its own codes, in the higher-order nonsensory parts of the cortex, has remained deeply mysterious. In this terra incognita, a path was opened by the discovery of grid cells, place-modulated entorhinal neurons whose firing locations define a periodic triangular or hexagonal array covering the entirety of the animal's available environment. This array of firing is maintained in spite of ongoing changes in the animal's speed and direction, suggesting that grid cells are part of the brain's metric for representation of space. Because the crystal-like structure of the firing fields is created within the nervous system itself, grid cells may provide scientists with direct access to some of the most basic operational principles of cortical circuits. V