The projections of the spinal cord upon the cerebellum of normal and Reeler mutant mice were compared by light and electron microscopic methods after hemicordotomy. In both genotypes this afferent system projects to the cerebellar cortex and to the roof nuclei. In the Reeler, there is an additional projection among the Purkinje cells and interneurons of the central cerebellar mass. In both normal and Reeler cerebellar cortex this mossy fiber system terminates as large glomeruli. In Reeler the spinal projection also gives rise to a smaller terminal which is distributed both to the cortex and the central cerebellar mass. In both genotypes the dendrites of granule cells and the somata and dendrites of Golgi cells are synaptic targets of the glomeruli of the cortical projection. In Reeler both the glomeruli and smaller terminals also form heterologous synaptic contacts with dendrite spines of heterotopic intracortical and subcortical Purkinje cells. In both genotypes the synapses are exclusively type I. A second class of heterologous synapse, a type I junction between axons of Golgi cells and Purkinje cell spines, is also recognized in electron micrographs. The present study is the first unequivocal demonstration by experimental hodologic method of heterologous synaptic junctions in the mammalian central nervous system. The existence of such junctions in the cytoarchitectonically anomalous cerebellum of this mutant emphasizes the critical role played by the cellular environment in shaping neural circuits in the developing nervous system.