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1. Complex and simple spike responses of Purkinje cells were recorded in the flocculus of anesthetized, paralyzed rabbits during rotating full-field visual stimuli produced by a three-axis planetarium projector. 2. On the basis of the spatial properties of their complex spike responses, floccular Purkinje cells could be placed into three distinct classes called Vertical Axis, Anterior (45 degrees) Axis and Posterior (135 degrees) Axis. The first two classes occurred in both monocular and binocular forms; the third class was encountered only in binocular form. For the binocular response forms, stimulation through one eye, called the dominant eye, elicited a stronger modulation of the complex spike firing rate than did stimulation of the other eye. The approximate orientation of that axis about which full-field rotation elicited the deepest modulation (the preferred axis) when presented to the dominant eye served as the class label. These classes are the same as those determined qualitatively for inferior olive neurons in the previous paper (47). The present study provides a quantitative description of their spatial tuning. 3. For Vertical Axis cells, the dominant eye was ipsilateral with respect to the flocculus recording site. The preferred axis was vertical and null (no-response) axes were in the horizontal plane. For the binocular response form of Vertical Axis cells (less than 10% of this class), the direction preferences for the two eyes were synergistic with respect to rotation about the vertical axis. 4. The dominant eye for the Anterior (45 degrees) Axis cells was contralateral, with the preferred axis oriented in the horizontal plane at approximately 45 degrees contralateral azimuth. The modulation depth showed a close to cosine relation with the angle between the preferred axis and the stimulus rotation axis. The average orientation (n = 10) for the dominant eye preferred axis, determined by the best-fit sinusoid, was 47 degrees contralateral azimuth. The preferred axis orientation for the ipsilateral (nondominant) eye in the binocular response forms was between 45 and 90 degrees azimuth in the horizontal plane. A null axis for each eye was at approximately 90 degrees to the preferred axis. 5. The Posterior (135 degrees) Axis cells were encountered only in binocular response forms. The dominant eye was ipsilateral, with the preferred axis oriented at approximately 135 degrees ipsilateral azimuth close to the horizontal plane. The modulation depth showed a close to cosine relation with the angle between the preferred axis and the stimulus rotation axis.(ABSTRACT TRUNCATED AT 400 WORDS)

Cerebellar cortical interneurons such as Golgi cells, basket cells, stellate cells, unipolar brush cells, and granule cells play an essential role in the operations of the cerebellum. However, detailed functional studies of the activity of these cells in both anesthetized and behaving animals have been hampered by problems in recognizing their physiological signatures. We have extracellularly recorded the spontaneous activity of vestibulocerebellar interneurons in ketamine/xylazine-anesthetized rats and subsequently labeled them with Neurobiotin using the juxtacellular technique. After recovery and morphological identification of these cells, they were related to statistical measures of their spontaneous activity. Golgi cells display a somewhat irregular firing pattern with relatively low average frequencies. Unipolar brush cells are characterized by more regular firing at higher rates. Basket and stellate cells are alike in their firing characteristics, which mainly stand out by their irregularity; some of them are set apart by their very slow average rate. The spontaneous activity of interneurons examined in the ketamine/xylazine rabbit fit within this general pattern. In the rabbit, granule cells were identified by the spontaneous occurrence of extremely high-frequency bursts of action potentials, which were also recognized in the rat. On the basis of these observations, we devised an algorithm that reliably determined the identity of 75% of the cells with only 2% incorrect classifications. The remaining cells were placed into border categories within which no classification was attempted. We propose that this algorithm can be used to help classify vestibulocerebellar interneurons recorded in awake, behaving animals.

The rabbit flocculus can be divided into five zones (zones 1, 2, 3, 4, and C2) with the use of acetylcholinesterase histochemistry. The projections of individual Purkinje cells in these zones to the vestibular and cerebellar nuclei were studied by using biocytin as an anterograde tracer. The zones were physiologically identified in terms of the Purkinje cell complex spike modulation occurring in response to optokinetic stimulation. In zones 1 and 3 neurons respond best to rotation about a horizontal axis that is close to perpendicular to the ipsilateral anterior semicircular canal, whereas in zones 2 and 4 neurons respond best to rotation about the vertical axis. Complex spike activity in zone C2 is unresponsive to optokinetic stimulation. Collectively, Purkinje cells of zone 1 projected to the ventral dentate nucleus, dorsal group y, and superior vestibular nucleus; Purkinje cells of zones 2 and 4 projected to the magnocellular and parvicellular parts of the medial vestibular nucleus; Purkinje cells of zone 3 projected to dorsal group y, ventral group y, and the superior vestibular nucleus; and Purkinje cells of zone C2 projected to the interposed posterior nucleus and dorsal group y. Some of the labeled Purkinje cell axons branched and innervated two nuclei. Branching axons from zone 1 either innervated both the ventral dentate nucleus and the superior vestibular nucleus or both dorsal group y and the superior vestibular nucleus. Branching axons from zones 2 and 4 innervated both the magnocellular and the parvicellular parts of the medial vestibular nucleus. Branching axons from zone 3 innervated both dorsal group y and the superior vestibular nucleus, or both ventral group y and the superior vestibular nucleus. Branching axons from zone C2 innervated both the interposed posterior nucleus and dorsal group y. Some of the target nuclei of the floccular Purkinje cell axons (e.g., dorsal group y and interposed posterior nucleus) project to the part of the inferior olive that, in turn, projects to the corresponding floccular zone, thus completing a closed pathway consisting of the inferior olive, the cerebellar cortex, and the cerebellar and vestibular nuclei. Other target nuclei (e.g., superior vestibular nucleus and medial vestibular nucleus) do not project back to the olivary subnuclei that innervate the flocculus and are part of an open olivofloccular pathway. An individual Purkinje cell thus can innervate a nucleus in the closed pathway as well as a nucleus in the open pathway.(ABSTRACT TRUNCATED AT 400 WORDS)

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