Burst activity plays no role in the field-to-field variability and rate remapping of grid cells

Grid cells in rodent medial entorhinal cortex are thought to play a key role for spatial navigation. When the animal is freely moving in an open arena the firing fields of each grid cell tend to form a highly regular, hexagonal lattice spanning the environment. However, firing rates vary from field to field and change under contextual modifications, whereas the field locations shift at most by a small amount under such "rate remapping." The observed differences in firing rate could reflect overall activity changes or changes in the detailed spike-train statistics. As these two alternatives imply distinct neural coding schemes, we investigated whether temporal firing patterns vary from field to field and whether they change under rate remapping. Focusing on short time scales, we found that the proportion of bursts compared to all discharge events is similar in all firing fields of a given grid cell and does not change under rate remapping. For each cell, mean firing rates with bursts are proportional to mean firing rates without bursts. However, this ratio varies across cells. Additionally, we looked at how rate remapping relates to entorhinal theta-frequency oscillations. Theta-phase coding was preserved despite firing-rate changes from rate remapping but we did not observe differences between the first and second half of the theta cycle, as had been reported for CA1. Our results indicate that both, the heterogeneity between firing fields and rate remapping, are not due to altered firing patterns on short time scales but reflect location-specific changes at the firing-rate level.