The cerebellum has a medial, cortico-nuclear zone consisting of the cerebellar vermis and the fastigial nucleus. Functionally, this zone is concerned with whole-body posture and locomotion. The vermis classically is thought to be included within the "spinocerebellum" and to receive somatic sensory input from ascending spinal pathways. In contrast, the lateral zone of the cerebellum is included in the "cerebro-cerebellum" because it is densely interconnected with the cerebral cortex. Here we report the surprising result that a portion of the vermis receives dense input from the cerebral cortex. We injected rabies virus into lobules VB-VIIIB of the vermis and used retrograde transneuronal transport of the virus to define disynaptic inputs to it. We found that large numbers of neurons in the primary motor cortex and in several motor areas on the medial wall of the hemisphere project to the vermis. Thus, our results challenge the classical view of the vermis and indicate that it no longer should be considered as entirely isolated from the cerebral cortex. Instead, lobules VB-VIIIB represent a site where the cortical motor areas can influence descending control systems involved in the regulation of whole-body posture and locomotion. We argue that the projection from the cerebral cortex to the vermis is part of the neural substrate for anticipatory postural adjustments and speculate that dysfunction of this system may underlie some forms of dystonia. motor system | virus tracing C lassically, the cerebellum is divided anatomically into three cortico-nuclear zones: a medial zone consisting of the cerebellar vermis and fastigial nucleus, an intermediate zone consisting of paravermal cortex and interposed nuclei, and a lateral zone consisting of the most lateral cerebellar cortex and the dentate nucleus (1, 2). Each of these zones is considered to be functionally distinct. For example, lesions of the vermis result in deficits in whole-body posture and locomotion, whereas lesions of the lateral zone result in deficits in limb movements (2, 3). Current texts include the vermis in the "spinocerebellum" and emphasize that it receives somatic sensory inputs related to the head and proximal parts of the body from ascending spinal pathways. In contrast, the lateral zone is included in the "cerebro-cerebellum" because it is densely interconnected with the cerebral cortex (4).We have begun to examine the inputs to the cerebellar cortex using retrograde transneuronal transport of rabies virus (RV) (5). In the course of these experiments we made the surprising observation that the primary motor cortex (M1) and several of the cortical motor areas on the medial wall of the hemisphere provide a major source of input to lobules VB-VIIIB of the vermis. Furthermore, a substantial portion of this input comes from cortical regions involved in the control of distal as well as proximal limb movements. Thus, our results challenge the classical view of the vermis and indicate that it no longer should be considered as entirely isolated from the...