Salient anatomic features of the cortico-ponto-cerebellar pathway

Recent studies of the primate corticopontine projection show that the neocerebellum--in addition to connections from motor and sensory areas--receives connections from various association areas of the cerebral cortex, some of which are thought to be primarily engaged in cognitive tasks. The quantities of such connections in relation to those from more clearly motor-related parts of the cortex need to be more precisely determined, however. Furthermore, the anatomic data on origin of corticopontine fibers needs to be supplemented with physiological experiments to clarify their functional properties at the single-cell level. For example, nothing is known of the functional role of the large input from the cingulate gyrus, nor is the input from the posterior parietal cortex physiologically characterized. Finally, the scarcity of corticopontine connections from the prefrontal cortex in the monkey (and probably also in man) may not seem readily compatible with a prominent role of the neocerebellum in certain cognitive tasks. We discuss data--in particular from three-dimensional reconstructions--indicating that both corticopontine projects and pontocerebellar neurons are arranged in a lamellar pattern. Corticopontine and pontocerebellar lamellae have similar shapes and orientations but appear to differ in other respects. Corticopontine terminal fields are sharply delimited, apparently without gradual overlap between projections from different sites in the cortex, whereas pontocerebellar lamellae are more fuzzy and exhibit gradual overlap of neuronal populations projecting to different targets. In spite of the sharpness of the corticopontine projection, there may be many opportunities for convergence of inputs from different parts of the cortex. Thus, the wide divergence of corticopontine projections produces many sites of overlap, and extensive interfaces between different terminal fields enabling convergence of inputs onto each neuron. We suggest that the lamellar arrangement of corticopontine terminal fields and of pontocerebellar neurons serve to create diversity of pontocerebellar neuronal properties. Thus, each small part of the cerebellar cortex would receive a specific combination of messages from many different sites in the cerebral cortex. The spatial arrangement of cerebrocerebellar connections have to be understood both in terms of fairly simple large-scale, gradual topographic relationships and an apparently highly complex pattern of divergence and convergence. Developmental studies of corticopontine and of pontocerebellar projections together with three-dimensional reconstructions in adults suggest that the highly complex adult connectional pattern may be created by simple rules operating during development.