Precise matching of olivo-cortical divergence and corticonuclear convergence between somatotopically corresponding areas in the medial C1 and medial C3 zones of the paravermal cerebellum

The paravermal cerebellar cortex contains three spatially separate zones (the C1, C3 and Y zones) which form a functionally coupled system involved in the control of voluntary limb movements. A series of 'modules' has been postulated, each defined by a set of olivary neurons with similar receptive fields, the cortical microzones innervated by these neurons and the group of deep cerebellar nuclear neurons upon which the microzones converge. A key feature of this modular organization is a correspondence between cortical input and output, irrespective of the zonal identity of the microzone. This was tested directly using a combined electrophysiological and bi-directional tracer technique in barbiturate-anaesthetized cats. During an initial operation, small injections of a mix of retrograde and anterograde tracer material (red beads combined with Fluoro-Ruby or green beads combined with biotinylated dextran amine or Fluoro-Emerald) were made into areas of the medial C1 and medial C3 zones in cerebellar lobule V characterized by olive-cerebellar input from the ventral forelimb. The inferior olive and the deep cerebellar nuclei were then scrutinized for retrogradely labelled cells and anterogradely labelled axon terminals, respectively. For individual experiments, the degree of C1-C3 zone terminal field overlap in the nucleus interpositus anterior was plotted as a function of either the regional overlap of single-labelled cells or the proportion of double-labelled cells in the dorsal accessory olive. The results were highly positively correlated, indicating that cortico-nuclear convergence between parts of the two zones is in close proportion to the corresponding olive-cerebellar divergence, entirely consistent with the modular hypothesis.