Spontaneous electrical activity and structural plasticity in the mature cerebellar cortex.

The Purkinje cell of the cerebellar cortex presents two distinct dendritic domains: a distal one, with spiny branchlets and a high density of spines innervated by many parallel fibers, and a proximal one, with a few clusters of spines innervated by a single climbing fiber terminal arbor. In adult rats, after 7 days of blocked electrical activity by the administration of TTX into the cerebellar parenchyma, the proximal dendritic domain of the Purkinje cell shows a remarkable growth of new spines that are innervated by parallel fibers. At the same time, the climbing fiber terminal arbor tends to become atrophic. In contrast, in the branchlets, spine density remains unmodified. These changes are reversible when TTX is removed. TTX treatment also leads to a decrease in spine size both in the branchlets and in the new spines of the proximal dendritic compartment. Spontaneous electrical activity should therefore be regarded not simply as noise, but as a significant signal for maintaining the typical profile of afferent innervation of the Purkinje cell and for preventing spines from shrinking.

Spontaneous electrical activity and dendritic spine size in mature cerebellar Purkinje cells.

Previous experiments have shown that in the mature cerebellum both blocking of spontaneous electrical activity and destruction of the climbing fibres by a lesion of the inferior olive have a similar profound effect on the spine distribution on the proximal dendrites of the Purkinje cells. Many new spines develop that are largely innervated by parallel fibers. Here we show that blocking electrical activity leads to a significant decrease in size of the spines on the branchlets. We have also compared the size of the spines of the proximal dendritic domain that appear during activity block and after an inferior olive lesion. In this region also, the spines in the absence of activity are significantly smaller. In the proximal dendritic domain, the new spines that develop in the absence of activity are innervated by parallel fibers and are not significantly different in size from those of the branchlets, although they are shorter. Thus, the spontaneous activity of the cerebellar cortex is necessary not only to maintain the physiological spine distribution profile in the Purkinje cell dendritic tree, but also acts as a signal that prevents spines from shrinking.