Medial entorhinal cortex activates in a traveling wave in the rat

Traveling waves are hypothesized to support the long-range coordination of anatomically distributed circuits. Whether separate strongly interacting circuits exhibit traveling waves remains unknown. The hippocampus exhibits traveling 'theta' waves and interacts strongly with the medial entorhinal cortex (MEC). To determine whether the MEC also activates in a traveling wave, we performed extracellular recordings of local field potentials (LFP) and multi-unit activity along the MEC. These recordings revealed progressive phase shifts in activity, indicating that the MEC also activates in a traveling wave. Variation in theta waveform along the region, generated by gradients in local physiology, contributed to the observed phase shifts. Removing waveform-related phase shifts left significant residual phase shifts. The residual phase shifts covaried with theta frequency in a manner consistent with those generated by weakly coupled oscillators. These results show that the coordination of anatomically distributed circuits could be enabled by traveling waves but reveal heterogeneity in the mechanisms generating those waves.