Simultaneous representation of multiple time horizons by entorhinal grid cells and CA1 place cells

Grid cells and place cells constitute the basic building blocks of the medial entorhinal-hippocampal spatial cognitive map by representing the spatiotemporal continuum of an animal past, present and future locations. However, the spatiotemporal relationship between these different cell types is unclear. Here we co-recorded grid and place cells in freely foraging rats. We show that average time shifts in grid cells tend to be prospective and are proportional to their spatial scale, providing a nearly instantaneous readout of a spectrum of progressively increasing time horizons ranging hundreds of milliseconds. Average time shifts of place cells are generally larger compared to grid cells and also increase with place field sizes. Moreover, time shifts displayed nonlinear modulation by the animal trajectories in relation to the local boundaries and locomotion cues. Finally, long and short time shifts occurred at different parts of the theta cycle, which may facilitate their readout. Together, these findings suggest that progressively increasing time horizons of grid and place cells may provide a basis for calculating animal trajectories essential for goal-directed navigation and planning.