6-OHDA-induced ectopia of external granule cells in the subarachnoid space covering the cerebellum. III. Morphology and synaptic organization of ectopic cerebellar neurons: a scanning and transmission electron microscopic study.

The present report describes the ultrastructure, surface morphology, and synaptic connectivity of ectopically placed cerebellar neurons after treatment of newborn rats with 100 μg 6-hydroxydopamine (6-OHDA), administered intracisternally. In addition to granule cells which form the majority of ectopic neurons, neurons exhibiting the ultrastructure of basket/stellate cells are found in the subarachnoid space over the cerebellum. The ectopic neurons present an almost complete spectrum of homologous efferent and afferent connections. Parallel fiber synapses are found on thorns of spiny branchlets of Purkinje cell dendrites which also have grown out into the subarachnoid space, and on the somata and dendrites of basket/stellate cells. Many ectopic parallel fibers are seen to pass into the molecular layer of the underlying cerebellar cortex through defects in the pial surface, presumably connecting with intracortical postsynaptic partners. Synapses between ectopic Purkinje cell dendrites and basket/stellate cell axons are also observed; however, the source of these axons remains uncertain. Granule cell dendrites are engaged in glomeruli with mossy fibers. Moreover, ectopic granule cell colonies are densely innervated by noradrenergic fibers. Our results show that the technique of generating ectopia of external granule cells provides an additional model for investigating influences of epigenetic factors on the development of nerve cells.