Evidence that climbing fibers control an intrinsic spike generator in cerebellar Purkinje cells

It is well established that the climbing fiber (CF) input to a cerebellar Purkinje cell ( PC) can exert a controlling influence on the background simple spike (SS) activity of the cell, in that repetitive stimulation of CFs causes a decrease in SS activity, and removal or inactivation of CFs is followed by a rise in activity. In the present study, the effects of inactivation of CFs in the short term and longer term ( hours) were investigated in anesthetized rats to determine how the CFs control the PC SS activity. Inactivation of the CF input to a PC was accomplished by either reversibly inactivating with lignocaine or by microlesioning the inferior olive. Consistent with previous findings, CF removal caused a transformation of the PC firing pattern, with SSs discharging more regularly and rising to an exceptionally high level. In cases in which CF activity resumed, SS rate declined to control levels within a few seconds. However, with sustained CF inactivation ( 30 min to 5 hr), SS activity continues to rise progressively and develops an oscillating firing pattern, consisting of alternating bursts of high-frequency discharge at up to 100-150 Hz followed by 10-20 sec periods of electrical quiescence. No accompanying changes in the threshold for evoking SSs via the parallel fibers were seen to accompany the increases in tonic SS activity. We conclude that the increase in SS activity that follows CF inactivation could be caused by the removal of an inhibitory action that CFs exert on the intrinsic pacemaker present in PCs under normal conditions.