Changes in hippocampal theta rhythm and their correlations with speed during different phases of voluntary wheel running in rats.

Hippocampal theta rhythm (4-12 Hz) can be observed during locomotor behavior, but findings on the relationship between locomotion speed and theta frequency are inconsistent if not contradictory. The inconsistency may be because of the difficulties that previous analyses and protocols have had excluding the effects of behavior training. We recorded the first or second voluntary wheel running each day, and assumed that theta frequency and activity are correlated with speed in different running phases. By simultaneously recording electroencephalography, physical activity, and wheel running speed, this experiment explored the theta oscillations during spontaneous running of the 12-h dark period. The recording was completely wireless and allowed the animal to run freely while being recorded in the wheel. Theta frequency and theta power of middle frequency were elevated before running and theta frequency, theta power of middle frequency, physical activity, and running speed maintained persistently high levels during running. The slopes of the theta frequency and theta activity (4-9.5 Hz) during the initial running were different compared to the same values during subsequent running. During the initial running, the running speed was positively correlated with theta frequency and with theta power of middle frequency. Over the 12-h dark period, the running speed did not positively correlate with theta frequency but was significantly correlated with theta power of middle frequency. Thus, theta frequency was associated with running speed only at the initiation of running. Furthermore, theta power of middle frequency was associated with speed and with physical activity during running when chronological order was not taken into consideration.