Older rats show slow modulation of hippocampal theta rhythm during voluntary running

Aging causes brain function degeneration and slows many motor and behavioural responses. The hippocampal theta rhythm (4-12 Hz) is related to cognition and locomotion. However, the findings on aging-related changes in the frequency and amplitude of hippocampal theta oscillations have been inconsistent. We hypothesized that older rats have slower responses in terms of hippocampal theta rhythm during voluntary wheel running than do young adult rats. By simultaneously recording electroencephalography and physical activity (PA), we evaluated theta oscillations in 8-week-old (young adult) and 60-week-old (middle-aged) rats before and during wheel running, which was conducted only during the rats' 12-h dark period. To test the alterations of hippocampal theta rhythm in voluntary wheel running, we analyzed the signals without (8-s) or with (2-s) chronological order. No significant difference was observed in total frequency (TP, 4-12 Hz), low-frequency (LT, 4-6.5 Hz), or high-frequency (9.5-12 Hz) theta activity between active waking and overall running in either group. The theta oscillations were slower in the middle-aged rats than in the young adult rats during wheel running but increased during running for both age groups. During wheel running, the middle-aged rats exhibited an increased LT, which was related to PA. On the basis of the chronological order of running, the young adult rats exhibited increased TP, and the middle-aged rats exhibited significant increases in middle-frequency (MT, 6.5-9.5 Hz) theta activity. The dominant modulations of MT in the middle-aged rats may have caused nonsignificant changes in total activity. These between-group differences in theta rhythm characteristics during voluntary running provide insights into age-related brain function decline.