Axonal competition in the synaptic wiring of the cerebellar cortex during development and in the mature cerebellum

Purkinje cell (PC) dendrites are made by a proximal dendritic domain, which is provided with scattered clusters of spines innervated by a single climbing fiber (CF) and by a distal domain with a high density of spines innervated by parallel fibers (PFs). Following block of electrical activity a spine increase occurs in the proximal domain and the new spines are innervated by the PFs while the number of synaptic contacts formed by the CF is reduced. Also the GABAergic input expands its territory of innervation on the proximal domain, which undergoes a profound restructuring of the glutamate and GABA receptors. Excitatory-like postsynaptic assemblies appear not only on the new spines, but also on the smooth region of the dendrite and both of them may be innervated by GABAergic terminals. In this case GABA receptors coexist with the glutamate receptors leading to the formation of hybrid synapses. In contrast, PF synapses contain solely glutamate receptors. Thus, the expression of glutamate receptors appears to be an intrinsic property of the PC, while the expression of the GABA receptors is induced by the presence of GABAergic terminals. The data highlight an important feature of the CF input; its electrical activity, in addition to inducing a powerful phasic excitation and a tonic inhibition, controls the finer architecture of the cerebellar cortex.