Control of cerebellar nuclear cells: a direct role for complex spikes?

The question of what modulates the firing of the cerebellar nuclei (CN) is one to which we presently have a surprisingly incomplete answer. Because most synaptic input to the CN originates from Purkinje cells (PCs), and simple spikes (SSs) are far more numerous than complex spikes (CSs), SSs are generally thought to be the dominant influence on the CN. However, evidence, reviewed here, suggests that this appears not to be the case in some physiologically important situations. As an alternative, we propose that CS activity may have at least as significant an effect on CN firing as do SSs. In particular, we suggest that CS activity has a role in controlling the bursts CN neurons show during various movements, during sleep states, and under ketamine-xylazine anesthesia. The ability to perform this role rests on the fact that CSs can be highly synchronized among PCs that project to the same CN neuron. Specifically, we suggest that synchronized CSs help determine the temporal course of the CN bursts, most often their offset, and that SSs and activity from cerebellar afferents may modulate the specific firing pattern within each burst. This joint control of CN activity may help explain anomalies present in the standard model for synaptic control of CN activity in which determination of CN firing patterns is attributed primarily to SSs.