Movement-related gating of climbing fibre input to cerebellar cortical zones

The inferior olive climbing fibre projection and associated spino-olivocerebellar paths (SOCPs) have been studied intensively over the last quarter of a century yet precisely what information they signal to the cerebellar cortex during movements remains unclear. A different approach is to consider the times during a movement when afferent signals are likely to be conveyed via these paths. Central regulation (gating) of afferent transmission during active movements is well documented in sensory pathways leading to the cerebral cortex and the present review examines the possibility that a similar phenomenon also occurs in SOCPs during movements such as locomotion and reaching. Several lines of evidence are considered which suggest that SOCPs are not always open for transmission. Instead, flow of sensory information to the cerebellum via climbing fibre paths is powerfully modulated during active movements. The findings are discussed in relation to the parasagittal zonal organization of the cerebellar cortex and, in particular, evidence is presented that different cerebellar zones are subject to similar patterns of gating during reaching but can differ appreciably in the pattern of modulation their SOCPs exhibit during locomotion. Furthermore, differences in gating can occur at different rostrocaudal loci within the same zone, suggesting that in the awake behaving animal, individual cerebellar zones are not functionally homogeneous. Finally, the data are interpreted in relation to the error detector hypothesis of climbing fibre function and the possibility explored that the gating serves as a task-dependent mechanism that operates to prevent self-generated 'irrelevant' sensory inputs from being relayed via the SOCPs to the cerebellar cortex, while behaviourally 'relevant' signals are selected for transmission.