Grid cells encode reward distance during path integration in cue-rich environments

The medial entorhinal cortex supports both path integration and landmark anchoring, but how these computations interact during goal-directed navigation is unclear. We show that grid cells dissociate from landmarks and instead encode reward distance when mice perform a path integration task on a cue-rich treadmill. Grid cell population activity reset at rewards and shifted coherently across trials, consistent with continuous attractor dynamics realigned by rewards. Furthermore, grid cells exhibited reduced spatial scales, broadened theta frequency distributions, and altered temporal coordination. These phenomena were captured by a theta interference model incorporating cell competition and two sets of theta oscillating inputs whose frequencies shifted apart. Switching to cue-based navigation stabilized the firing fields and partially restored grid scale, theta frequencies and temporal structure. These results demonstrate that MEC circuits flexibly reset to encode goal-directed trajectories, and suggest that continuous attractor and interference mechanisms normally cooperate but can decouple under path integration demands.