Relation between activities of the cortex and vibrissae muscles during high-voltage rhythmic spike discharges in rats

Paroxysmal 5- to 12-Hz high-voltage rhythmic spike (HVRS) activities, which are accompanied by whisker twitching (WT), are found in Long Evans rats, but the function of these HVRS activities is still debated. In four major functional hypotheses of HVRS discharges, i.e., alpha tremor, attention/mu rhythm, idling/mu rhythm, and absence seizure, the first two hypotheses emphasize WT behavior in HVRS bouts. Whisker movement is primarily determined by activation of intrinsic and extrinsic muscles. To clarify the role of WT in HVRS activities, simultaneous recording of the activities from the cortex and intrinsic/extrinsic and neck muscles were performed. Most HVRS bouts (68.8%) revealed no time-locked WT behavior in a 2-h recording session. In addition, WT primarily arose from active protraction due to activation of intrinsic muscles followed by passive retraction. A small portion of WT resulted from activation of both vibrissae muscles with dynamic frequency-dependent phase shifts. Onset of the rhythmic vibrissae EMG significantly lagged behind HVRS onset, and the mean duration of vibrissae muscle activity was one-third to a one-half of a HVRS bout. Moreover, a greater number of HVRS bouts were associated with a longer HVRS duration and higher oscillation frequency. Oscillation frequencies of HVRS activities without WT behavior were significantly lower than those with WT. Under peripheral sensory/motor blockade by xylocaine injection, oscillation frequencies of HVRS bouts significantly decreased, but no remarkable changes in the number or duration of HVRS bouts were observed. Compared with vibrissa muscle activity during WT and exploratory whisking, the duration of muscular activity in each cycle was apparently longer during whisking bouts. Based on these results, overemphasis of the role of WT on HVRS activities might not be appropriate. Instead, HVRS discharges may be associated with absence seizure or idling state. In addition, peripheral inputs, including WT, may elevate the oscillation frequency of HVRS bouts. Moreover, different muscular controls may exist between WT and whisking.