A mechanism for type III vestibular responses of frog cerebellar Purkinje cells

Type III Purkinje cells (P-cells), which are excited with both directions of horizontal rotation, are found in high numbers in the frog auricular lobe and adjacent cerebellar areas. To examine the mechanisms underlying these responses, recordings were made from P-cells in curarized animals during rotational stimulation of the horizontal canals. The horizontal canal input to these cells was then modified unilaterally by VIIth nerve section, intraperilymphatic injection of local anesthetic, or by caloric stimulation. Control recordings were also obtained from peripheral canal neurons. Type III responses were abolished by unilateral lesions or reversible blockage of the VIIIth nerve with local anesthetic. The remaining responses were attributable only to the unaffected horizontal canal, ie. only type II or type I responses were observed upon interruption of the ipsi-or contralateral nerve, respectively. The level of spontaneous activity of cerebellar input fibers was low and during rotation produced 'cell silencing' response waveform asymmetries (facilitation greater than disfacilitation). When the level of peripheral resting activity was increased (warm water irrigation), thereby increasing horizontal canal response symmetry, type III responses were reduced in magnitude or abolished. Conversely, cold water irrigation, which decreases the resting rate and response symmetry of input fibers, enhanced type III response magnitudes. On the basis of these results, it is suggested that type III responses result from the fact that single P-cells receive a facilitatory input from both horizontal canals. Since these inputs are 180 degrees phase-reversed and their response waveforms asymmetrical, their resulting postsynaptic effect is a net excitation during both portions of the stimulus cycle.