Serotonin (5-HT) exerts a variety of effects on the excitability of motoneurons, interneurons, and ascending tract cells. Spinocerebellar-tract cells in the dorsal horn receive synaptic connections from serotoninergic axons, but little is known about the relationships between serotoninergic axons and dorsal spinocerebellar tract (DSCT) cells in Clarke's column. We studied these relationships by using a combination of immunohistochemical localization of 5-HT-immunoreactive boutons and intracellular staining with horseradish peroxidase (HRP) or neurobiotin of identified DSCT cells in vivo. In the adult cat, Clarke's column displayed a lower density of 5-HT-immunoreactive axons and boutons than adjacent regions of the spinal gray matter. Eleven intracellularly stained DSCT cells were analyzed with light microscopy, and six of these cells were entirely reconstructed in three dimensions. A total of 3,739 close appositions (340+/-101 per postsynaptic neuron: mean +/- SD) were observed on the labeled DSCT cells. The majority (97%) of the appositions were formed on dendrites, including proximal and distal branches, with an average density of about 1.4 appositions per 1,000 microm2 of dendritic membrane. These results indicate that the bulbo-spinal serotoninergic system(s) provide direct innervation of Clarke's column-DSCT cells in the upper lumbar spinal cord and that the inputs are spread widely over all regions of the target neurons' soma-dendritic membrane.
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