Timing and plasticity in the cerebellum: focus on the granular layer

Two of the most striking properties of the cerebellum are its control in timing of motor operations and its ability to adapt behavior to new sensorimotor associations. Here, we propose a 'time-window matching' hypothesis for granular layer processing. Our hypothesis states that mossy fiber inputs to the granular layer are transformed into well-timed spike bursts by intrinsic granule cell processing, that feedforward Golgi cell inhibition sets a limit to the duration of such bursts and that these activities are spread over particular fields in the granular layer so as to generate ongoing time-windows for proper control of interacting motor domains. The role of synaptic plasticity would be that of fine-tuning pre-wired circuits favoring activation of specific granule cell groups in relation to particular time windows. This concept has wide implications for processing in the olivo-cerebellar system as a whole.