Gracile projection to the cat medial accessory olive: ultrastructural termination patterns and convergence with spino-olivary projection

Abstract

The caudal medial accessory subdivision of the inferior olive (cMAO) receives information from the hindlimb from both the gracile nucleus and the lumbosacral spinal cord. This study determined which elements in cMAO serve as the postsynaptic targets of the gracile projection and whether these elements also receive input from the lumbosacral spinal cord. Gracile axons were labeled in cats by anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP), visualized with tetramethylbenzidine. Convergence of gracile and lumbosacral axons was evaluated by labeling in the same animal, one pathway by WGA-HRP and the other by degeneration. In cMAO, gracile axons synapse with equal probability on dendritic spines and distal dendritic shafts. This termination pattern contrasts markedly with that of other somatosensory inputs to the inferior olive and may account for the greater heterogeneity in responses to somatosensory stimuli displayed by neurons in cMAO. The distal dendritic shafts receiving gracile input were more likely than dendritic spines to receive convergent input from putative inhibitory synapses. The most likely source of these inhibitory synapses is the parasolitary nucleus, a structure that has been shown by others to receive input from the cerebellum. Thus the parasolitary nucleus may serve as an inhibitory relay between the cerebellum and cMAO. The dendritic spines in cMAO that receive input from the gracile nucleus often receive additional input from the lumbosacral spinal cord. This convergence of somatosensory axons on dendritic spines may provide a mechanism through which the unusually complex receptive fields of neurons in cMAO are generated.