Variable timing of synaptic transmission in cerebellar unipolar brush cells.

The cerebellum ensures the smooth execution of movements, a task that requires accurate neural signaling on multiple time scales. Computational models of cerebellar timing mechanisms have suggested that temporal information in cerebellum-dependent behavioral tasks is in part computed locally in the cerebellar cortex. These models rely on the local generation of delayed signals spanning hundreds of milliseconds, yet the underlying neural mechanism remains elusive. Here we show that a granular layer interneuron, called the unipolar brush cell, is well suited to represent time intervals in a robust way in the cerebellar cortex. Unipolar brush cells exhibited delayed increases in excitatory synaptic input in response to presynaptic stimulation in mouse cerebellar slices. Depending on the frequency of stimulation, delays extended from zero up to hundreds of milliseconds. Such controllable protraction of delayed currents was the result of an unusual mode of synaptic integration, which was well described by a model of steady-state AMPA receptor activation. This functionality extends the capabilities of the cerebellum for adaptive control of behavior by facilitating appropriate output in a broad temporal window.