Eye movement related neurons in the cat pontine reticular formation: projection to the flocculus

This study examines projection to the cerebellar flocculus of eye movement-related neurons in the median and paramedian part of the cat pontine tegmentum between the trochlear and the abducens nucleus. They were identified by rhythmic activity related to horizontal vestibular nystagmus induced by sinusoidal rotation. These neurons were classified into several groups by their discharge patterns during nystagmus, using criteria of earlier studies on saccadic eye movements and vestibular nystagmus in the monkey. Electrical stimulation of the ipsilateral flocculus elicited antidromic spike responses in a number of burst-tonic neurons and long-lead and medium-lead burst neurons. These neurons were located in and around the medial longitudinal fasciculus, the nucleus raphe pontis and the nucleus reticularis tegmenti pontis. A few neurons tested were also activated antidromically by stimulation of the contralateral flocculus. In contrast, no pauser neurons were activated from the ipsi-lateral flocculus. It is concluded that eye movement-related neurons in the medial pontine tegmentum, except for pauser neurons, directly project to the flocculus and may convey information about eye movements of visual and vestibular origins to the flocculus.     

The effect of prolonged monocular occlusion on latent nystagmus in the treatment of amblyopia

We recorded eye movements in 5 patients with latent nystagmus (LN) before and after 2 days of occlusion of the better eye. The slow-phase speed of the nystagmus (SPS) was in general, before occlusion, lower when the better eye fixated but, after occlusion, lower when the worse eye fixated. However, the sum of SPS during right fixation and SPS during left fixation remained constant. Oscillopsia complaints gradually disappeared during the period of occlusion. These findings indicate that the difference between the SPS during fixation with the right eye and the SPS during fixation with the left eye in LN patients is caused by a compensatory drift that decreases LN during fixation with the better eye but increases LN during fixation with the worse eye. During occlusion, this compensatory drift changes its direction and magnitude slowly over days. Hence, occlusion of the better eye in children with amblyopia and LN should be prescribed only in days per week, not in hours per day.     

Vestibular primary afferent projection to the cerebellum of the rabbit

The vestibular primary afferent projection to the cerebellum of the rabbit was studied with retrograde and orthograde tracers. We injected individual lobules of the cerebellum with horseradish peroxidase (HRP) or wheat germ agglutinin-HRP (WGA-HRP). Following these injections the numbers of labeled and unlabeled cells in Scarpa's ganglion were counted. Approximately 64–89% of the cells in Scarpa's ganglion were labeled retrogradely following uvula-nodular injections. About 2% of the cells in the ipsilateral Scarpa's ganglion were labeled after injections of the flocculus. Virtually no cells were labeled following injections of the ventral paraflocculus. The vestibular primary afferent projection to the uvula-nodulus is so extensive that it must be part of a collateral system that also innervates the vestibular nuclei. This collateral projection pattern was confirmed by using fluorescent tracers injected into the uvula-nodulus and vestibular complex. Fluorogold was injected into the uvula-nodulus and peroxidaserhodamine isothiocyanate was injected into the vestibular complex. More than 50% of the neurons in Scarpa's ganglion were double labeled by these subtotal injections. The dense vestibular primary afferent projection to the uvula-nodulus was confirmed by using the C fragment of tetanus toxin (TTC) injected into the labyrinth as an orthograde tracer. With the TTC technique, the vestibular primary afferent projection to the uvula-nodulus terminated exclusively in the ipsilateral granule cell layer of lobules 9d and 10. Much sparser vestibular primary afferent projections were found in the banks of major cerebellar sulci. A barely detectable projection was found to the flocculus and ventral paraflocculus. © 1993 Wiley-Liss, Inc.     

Two groups of secondary vestibular neurons mediating horizontal canal signals, probably to the ipsilateral medial rectus muscle, under inhibitory influences from the cerebellar flocculus in rabbits

Vestibular nuclear neurons that mediate horizontal canal signals to the ipsilateral medial rectus motoneurons were explored in anesthetized and decerebrate rabbits. These neurons were identified by four criteria: (1) they were activated monosynaptically by ipsilateral vestibular nerve stimulation and (2) antidromically from the oculomotor nucleus region, while they were inhibited by (3) direct floccular stimulation and (4) ipsilateral retinal stimulation that activated floccular Purkinje cells via a climbing fiber afferent pathway. Neurons fulfilling these criteria were found in two anatomically different regions, i.e. the rostrolateral part of the medial vestibular nucleus and in the ventral part of the lateral vestibular nucleus. In decerebrate rabbits, neurons in both loci responded to horizontal rotation of the whole body with the type I pattern (excited by ipsilateral rotation). These results suggest that horizontal canal signals are conveyed to ipsilateral medial rectus motoneurons by two separate groups of vestibular nuclear neurons which may play different roles in the vestibulo-ocular reflex.     

Neuronal events correlated with long-term adaptation of the horizontal vestibulo-ocular reflex in the primate flocculus

The activity of flocculus Purkinje cells was examined in Japanese monkeys during sustained vestibular-visual stimulation which caused adaptation of the horizontal vestibulo-ocular reflex (H-VOR). In the floccular area related to the H-VOR by microstimulation. Purkinje cells consistently changed their simple spike responsiveness to head rotation in parallel with the adaptive H-VOR gain change. Similar changes occurred even after the H-VOR had been extinguished by lesioning of the vestibular nuclei. The complex spike discharge, on the other hand, modulated during vestibular-visual stimulation with a reciprocal pattern to the adaptive changes in the simple spike discharge. These results support the hypothesis that the flocculus adaptively modifies the H-VOR through their simple spike activity under the influence of visual climbing fiber signals.     

Functional localization in the three floccular zones related to eye movement control in the cat

Anatomically, the cat's cerebellar flocculus can be divided into 3 zones on the basis of differences in their efferent projection sites^13,14. The functional differences of these 3 zones in relation to eye movement control were investigated by observing the eye movements evoked by electric stimulation of each zone of the flocculus in ketamine-anesthetized cats. Stimulation of the flocculus elicited a slow eye movement. The direction of the slow eye movement was mapped. A downward eye movement was evoked by stimulation of the caudal zone. An ipsilateral horizontal eye movement was induced from the middle zone. An upward eye movement was elicited from the rostral zone. When prolonged stimulation was applied to the flocculus, the slow eye movement was followed by nystagmus in the opposite direction. This nystagmus persisted for many seconds after cessation of stimulation (afternystagmus). Nystagmus and afternystagmus could not be elicited in deeply anesthetized cats. Possibilities as to how the stimulation leads to various eye movements are discussed.     

Eye field in the cerebellar flocculus of pigmented rabbits determined with local electrical stimulation

The cerebellar flocculus was mapped with local stimulation techniques in alert pigmented rabbits. Triple-barrelled microelectrodes filled with solutions each containing one of three different dyes (Fast Green FCF, Pontamine Sky Blue and Nigrosine) were used for recording and stimulation. The H-zone from which local stimulation through the microelectrode induced abduction in the ipsilateral eye was visualized on the reconstructed model of the flocculus to span across major folia of the flocculus, forming a narrow strip 0.5-1.0 mm wide. The H-zone was flanked with two V-zones, rostral and caudal, from which downward eye movements were induced in the ipsilateral eye. The R-zone, from which clockwise rotation was induced in the contralateral eye, was extended rostrocaudally across the H- and V-zones. In addition to these eye movement-related zones, a restricted area specifically related to eye blinking was found in the rostroventral area of the flocculus.     

Optokinetic response of cells in the nucleus reticularis tegmenti pontis of the pigmented rabbit

Summary In immobilized pigmented rabbits anesthetized with N₂O (70%) and halothane (2–4%), extracellular spikes were recorded from neurons in the nucleus reticularis tegmenti pontis (NRTP) and their responses to optokinetic stimulation (OKS) were examined. OKS was delivered using constant-velocity (0.1–4.0°/s) movements of a random dot pattern (60° × 60°) at 0°, 45°, 90° or 135° to the horizon. With OKS delivered to the contralateral eye (n=43), the preferred directions of NRTP cells were forward (F, n = 10), backward (B, n = 7), downward (D, n = 5), and the remaining cells showed no response (N, n = 21). With OKS delivered to the ipsilateral eye (n = 43), the preferred directions were F (n = 8), B (n = 8), upward (U, n = 2), D (n= 1) and N (n = 24). The majority of cells which responded to OKS (17/22 for contralateral, and 16/19 for ipsilateral OKS) preferred the horizontal orientation. The optimum velocity ranged from 0.2 to l°/s. The results suggest that the NRTP cells mainly transfer horizontal optikinetic signals to the flocculus and control horizontal optokinetic eye movements.     

Role of the primate flocculus in adaptation of the vestibulo-ocular reflex

Neuronal events associated with adaptation of the horizontal vestibulo-ocular reflex (HVOR) induced by sustained vestibular-visual mismatching were investigated in the primate flocculus. The floccular area related to the HVOR (H-zone) was identified by electrical micro-stimulation which induced ipsilaterally directed horizontal eye movement. It was thus found that Purkinje cells in the H-zone consistently changed their simple spike responses to head rotation in parallel with the adaptive HVOR gain change. This was demonstrated by observing the change of simple spike firing of Purkinje cells during adaptation of HVOR either in a population study or an individual study. Since similar changes occurred even after bilateral lesioning of vestibular nuclei had extinguished the HVOR, these changes appear to represent vestibular, but not eye velocity, mossy fiber responsiveness. The complex spike discharge, on the other hand, modulated during vestibular-visual stimulation with a reciprocal pattern to the adaptive changes in the simple spike discharge. These results are consistent with the hypothesis that the flocculus Purkinje cells adaptively control the HVOR through their simple spike activity under influences of retinal error signals conveyed by visual climbing fiber pathways.