Active contribution of dendrites to the tonic and trimodal patterns of activity in cerebellar Purkinje neurons.

The cerebellum is responsible for coordination of movement and maintenance of balance. Cerebellar architecture is based on repeats of an anatomically well defined circuit. At the center of these functional circuits are Purkinje neurons, which form the sole output of the cerebellar cortex. It is proposed that coordination of movement is achieved by encoding timing signals in the rate of firing and pattern of activity of Purkinje cells. An understanding of cerebellar timing requires an appreciation of the intrinsic firing behavior of Purkinje cells and the extent to which their activity is regulated within the functional circuits. We have examined the spontaneous firing of Purkinje neurons in isolation from the rest of the cerebellar circuitry by blocking fast synaptic transmission in acutely prepared cerebellar slices. We find that, intrinsically, mature Purkinje cells show a complex pattern of activity in which they continuously cycle among tonically firing, bursting, and silent modes. This trimodal pattern of activity emerges as the cerebellum matures anatomically and functionally. Concurrent with the transformation of the immature tonically firing cells to those with the trimodal pattern of activity, the dendrites assume a prominent role in regulating the excitability of Purkinje cells. Thus, alterations in the rate and pattern of activity of Purkinje neurons are not solely the result of synaptic input but also arise as a consequence of the intrinsic properties of the cells.