Nature of optokinetic response and zonal organization of climbing fiber afferents in the vestibulocerebellum of the pigmented rabbit

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Abstract
In pigmented rabbits anesthetized with N2O (70%) and halothane (2–4%), Purkinje cells were extracellularly recorded in the nodulus. Large field (60°×60°) optokinetic stimulation (OKS) with constant velocity was delivered to either the ipsi- or contralateral eye, and the direction and velocity selectivities of complex spike responses were examined. To ipsilateral OKS (n = 181), the preferred direction was forward (F, n = 72), upward (U, n = 38) or downward (D, n = 10), while the remaining cells (n = 61) showed no response (N). To contralateral OKS (n = 117), the preferred direction was backward (B, n = 22), upward (U, n = 7) or downward (D, n = 22), while the rest (n = 66) showed N. Cells tested with both eyes (n = 95) fell into 8 categories based on the preferred direction to ipsi- and contralateral OKS: (1) ipsi-F and contra-B (F/B type, n = 20), (2) ipsi-F but contra-N (F/N type, n = 12), (3) ipsi-U and contra-D (U/D type, n = 15), (4) ipsi-U but contra-N (U/N type, n = 13), (5) ipsi-N but contra-D (N/D type, n = 1), (6) ipsi-D but contra-N (D/N type, n = 5), (7) ipsi-N but contra-U (N/U type, n = 6), and (8) N to both eyes (N/N type, n = 23). The optimum velocity was in the range 0.1–2.0°/s for all cells responsive to OKS. In the ventral lamella, four medio-laterally aligned zones were demonstrated. In the most medial zone (0–0.5 mm from the midline), the majority of cells showed ipsi-N or contra-N responses. In the second zone (0.5–1.5 mm), most cells preferred ipsi-F or contra-B directions. In the third zone (1.5–2.5 mm), most cells preferred ipsi-U or contra-D directions. In the most lateral zone (2.5–3.5 mm), most cells preferred ipsi-F or contra-B directions. In the dorsal lamella, a longitudinal zone characterized with cells preferring ipsi-U or contra-D directions was found about 1.5–2.5 mm from the midline. This zone seemed to be the continuation of the third zone in the ventral lamella. Cells preferring ipsi-D or contra-U directions were scattered in the medial half of both the dorsal and ventral lamellae. Most cells responsive to electrical stimulation of the contralateral optic tract (OT) preferred the ipsi-F direction and were localized in the second and the most lateral zones of the ventral lamella. As for cells activated by a climbing fiber with a branching axon to the flocculus, no characteristic feature was found in terms of the preferred direction to OKS, except that none of the cells preferring ipsi-D or contra-U directions were activated by such branching climbing fibers. The results indicate that the nodulus consists of at least four functionally distinct zones in terms of direction selectivity of visual climbing fiber afferents.