Modeling the grid cell activity on non-horizontal surfaces based on oscillatory interference modulated by gravity.

Internal representation of the space is a fundamental and crucial function of the animal's brain. Grid cells in the medial entorhinal cortex are thought to provide an environment-invariant metric system for the navigation of the animal. Most experimental and theoretical studies have focused on the horizontal planar codes of grid cell, while how this metric coordinate system is configured in the actual three-dimensional space remains unclear. Evidence has implied the spatial cognition may not be fully volumetric. We proposed an oscillatory interference model with a novel gravity and body plane modulation to simulate grid cell activity in complex space for rodents. The animal can perceive the rotation of its body plane along the local surface by sensing the gravity, causing the modulation to the dendritic oscillations. The results not only reproduce the firing patterns of the grid cell recorded from known experiments, but also predict the grid codes in novel environments. It further demonstrates that the gravity signal is indispensable for the animal's navigation, and supports the hypothesis that the periodic firing of the grid cell is intrinsically not a volumetric code in three-dimensional space. This will provide new insights to understand the spatial representation of the actual world in the brain.