Comparison of vertical and horizontal optokinetic nystagmus in the squirrel monkey.

Horizontal and vertical optokinetic nystagmus (OKN) and optokinetic after-nystagmus (OKAN) of squirrel monkeys were compared with those of rabbits, cats and humans that were previously described. Squirrel monkeys showed similar findings to cats, in which vertical optokinetic nystagmus (VOKN) is not as well elicited as horizontal optokinetic nystagmus (HOKN) and down-pursuit OKN is poorer than up-pursuit OKN. As to the reasons that bring about different responses of OKN and OKAN (and vestibular nystagmus) in different planes, we speculated two possibilities: compensatory activation of horizontal eye movement for narrowed visual field accompanied by frontally positioned eyes, and the gravity that restricts and modifies posture and locomotion. Directional difference of VOKN may be caused by a physiological mechanism that makes visual fixation not susceptible to downward movement of the ground surface during forward locomotion.     

Effect of vestibulo-cerebellar lesions on asymmetry of vertical optokinetic functions in the squirrel monkey

The role of the cerebellar uvula and nodulus in vertical optokinetic after-nystagmus (OKAN) was studied in 4 squirrel monkeys. Aspiration ablation of the uvula and nodulus resulted in no significant change in the initial or peak gain of vertical optokinetic nystagmus (OKN) during the 24-week post-operative observation. However, the asymmetry of vertical OKAN was significantly altered. Using a protracted upward OK stimulus, slow phase-down OKAN-II, which was not seen pre-operatively, was significantly increased. In contrast, a downward OK stimulus produced little change in slow phase-up OKAN-II. Thus, the asymmetric degree of vertical OKAN-II was decreased after uvulonodulectomy. In addition, there was a post-operative reduction in the vertical oculomotor stability. When slow-phase eye velocity of OKAN was plotted along the time scale, the amplitude and frequency of the sinusoidal pattern was increased. OKAN-III and OKAN-IV were found in 50% of the monkeys after uvulonodulectomy. It is therefore thought that inhibition and directional control from the uvula and nodulus influence the stability and asymmetrical behaviour of the leaky integrator in the second order output system.     

Asymmetry of vertical optokinetic after-nystagmus in squirrel monkeys

Asymmetry of vertical optokinetic after-nystagmus (OKAN) was studied in 6 squirrel monkeys. The slow-phase eye velocity (SPEV) of upward OKAN first-phase (OKAN-I) increased with increasing stimulus velocity, whereas the SPEV of downward OKAN-I diminished. The time constant of OKAN-I was shortened with the increase in stimulus speed in both directions. With a downward stimulus, the short stimulus duration failed to produce OKAN second-phase (OKAN-II) (upward slow-phase); however, with an increase in stimulus duration, the percentage appearance increased. There was no change in percentage appearance, regardless of the duration of upward stimulus. The asymmetry of OKAN-I and that of OKAN-II differed to a certain degree.     

The effect of saccular ablation on vertical optokinetic after-nystagmus in squirrel monkeys

Summary The effect of bilateral saccular ablation on the asymmetry of vertical optokinetic after-nystagmus (OKAN) was studied in squirrel monkeys. No significant changes occurred in the initial slow-phase eye velocity (SPEV) or the time constant of the upward or downward OKAN first phase (OKAN-I) under various stimulus conditions. However, with a protracted downward stimulus, the maximum SPEV and the number of beats of the slow-phase-up OKAN second phase (OKAN-II) significantly increased. This increase should be the result from enhancement of the downward optokinetic input. In contrast, there was only minimal change in the slow-phase-down OKAN-II. Thus, the asymmetrical dominance of the vertical OKAN (dominance upward) remained the same after saccular deafferentation. Offprint request to: M. Igarashi     

Galvanically induced asymmetric optokinetic after-nystagmus

The effect of an asymmetric vestibular input on the symmetry of horizontal optokinetic after-nystagmus (OKAN) was studied in twenty healthy subjects. Optokinetic nystagmus (OKN) was elicited by a whole-field optokinetic drug, rotating at 90 degrees/s, and eye-movements were recorded by a DC electro-oculographic technique (EOG). The ratio of OKAN following right and left-beating OKN respectively was computed. An asymmetric vestibular input was generated by a continuous bi-polar, bi-aural galvanic stimulus (1 mA) to the vestibular nerves during the optokinetic stimulation and the recording of the OKAN. During galvanic stimulation the relation between left and right-beating OKAN was asymmetric, compared with the OKAN found after optokinetic stimulation only. The galvanic stimulus caused a preponderance for OKAN with the fast phase beating toward the cathode. Thus, the small vestibular asymmetry induced by the galvanic stimulus, which was not strong enough to produce nystagmus by itself, caused an asymmetric OKAN. These findings suggest that examination of OKAN may be of value to detect small vestibular asymmetries in peripheral vestibular disorders in man.     

Effect of otolith end organ ablation on horizontal optokinetic nystagmus, and optokinetic afternystagmus in the squirrel monkey.

Horizontal optokinetic nystagmus (OKN) and optokinetic afternystagmus (OKAN) (stimulus speed 0-200 degrees/sec with 1 degree/sec constant angular acceleration) were examined before and after utriculo-sacculectomy (bilateral, two-stage) in squirrel monkeys. OKN exhibited a slight decline only after bilateral otolith and organ ablations. OKAN showed a minimal decline after unilateral operation but no change after bilateral operations. Severe OKN reduction and disappearance of OKAN after bilateral labyrinthectomy in primates should basically reflect the elimination of inputs from the cristae ampullares, and not from the maculae.     

Effect of body positions in human optokinetic nystagmus and optokinetic after nystagmus

We determined whether whole body tilt would shift the axis of optokinetic nystagmus (OKN) and optokinetic-after nystagmus (OKAN) induced by full-field rotation at 35 degrees/sec. Fifteen normal people were positioned upright or tilted 30 degrees, 60 degrees or 90 degrees to both sides. Stripes of 5 degrees were projected on a 10-foot dome around the subject's yaw axis. Each trial lasted 45 sec. The lights were then extinguished, and the subject remained in darkness for 30 sec, while after nystagmus (OKAN) was recorded. Horizontal and vertical eye movements were recorded by video-oculography at 60 Hz. Eye position and velocity data were stored on optic disk cartridge by use of the data acquisition system. A. OKAN: For the subject in the upright position, the OKN velocity vector was aligned with both gravity and the subject's yaw axis with two minor exceptions. When the subject was tilted, a vertical OKN component (VOKN) appeared in a majority of subjects. For all 15 subjects, the mean angle of the OKN velocity vector regravity (Vectorg) was 22.6 +/- 7.2 degrees at 30 degrees tilted position. The Vectorg were 48.5 +/- 10.3 degrees at 60 degrees tilted position, and 76.4 +/- 12.6 degrees at 90 degrees tilted position. This represented shifts of the OKN velocity vector from the body axis of 7.4 degrees, 11.5 degrees and 13.6 degrees, respectively. The horizontal OKN (HOKN) gain remained unchanged in different positions. B. OKAN: The duration of HOKAN and initial slow phase velocity (SPV) of HOKAN decreased as the body position increased from upright to 30 degrees, 60 degrees and 90 degrees tilted position, respectively. The incidence and initial SPV of VOKAN and Re-Body did not change as the body position increased from upright to 30 degrees, 60 degrees and 90 degrees tilted position, respectively. Thus, VOKN was observed during HOKN as subjects were tilted and tended to vector to gravity, but VOKAN was not always observed during horizontal OKAN when subjects were tilted.     

Experimental parameter estimation of a visuo-vestibular interaction model in humans

Visuo-vestibular interactions in monkeys can be accurately modelled using the classical Raphan and Cohen's model. This model is composed of direct vestibular and visual contributions to the vestibulo-ocular reflex (VOR) and of a velocity storage. We applied this model to humans and estimated its parameters in a series of experiments: yaw rotations at moderate (60°/s) and high velocities (240°/s), suppression of the VOR by a head-fixed wide-field visual stimulus, and optokinetic stimulation with measurements of optokinetic nystagmus (OKN) and optokinetic afternystagmus (OKAN). We found the velocity storage time constant to be 13 s, which decreased to 8 s during visual suppression. OKAN initial velocity was 12% of the OKN stimulus velocity. The gain of the direct visual pathway was 0.75 during both visual suppression and OKN; however, the visual input to the velocity storage was higher during visual suppression than during OKN. We could not estimate the time constant of the semicircular canals accurately. Finally, we inferred from high-velocity rotations that the velocity storage saturates around 20-30°/s. Our results indicate that the dynamics of visuo-vestibular interactions in humans is similar as in monkeys. The central integration of visual cues, however, is weaker in humans.     

Horizontal and vertical optokinetic nystagmus in man

Horizontal and vertical optokinetic nystagmus (OKN) was studied in 20 healthy adults. Although the horizontal OKN showed no directional difference, a statistically significant difference was found between horizontal and vertical OKN. Upward optokinetic pursuit was better on average than downward pursuit, but more variable. The inferiority of vertical OKN seems to indicate a suppression of optokinetic pursuit due to a different direction of the rotational axis from that of gravity. Regarding the vertical OKN findings, it is speculated that manifest directionality found in quadrupeds is modified in men by the change of their visual field on forward locomotion accomplished by their upright walking posture.     

Effects on the optokinetic system of midline lesions in the pretectum of monkeys

The nucleus of the optic tract (NOT), an important visuo-motor relay between the retina and preoculomotor structures, is responsible for mediating horizontal optokinetic nystagmus (OKN) in monkeys, cats, rabbits and rats. In addition to its projection to the vestibular nuclei, the NOT has a prominent projection to the contralateral NOT via the posterior commissure. In order to evaluate the role of the commissural fibers between the NOTs in OKN, we cut the posterior commissure in three Macaca fuscata. The animals viewed the OKN stripes under three conditions: right eye viewing, left eye viewing, and both eyes viewing. OKN was recorded in response to counter-clockwise and clockwise stimulation at stimulus velocities of 30 degrees/s, 60 degrees/s and 90 degrees/s. After control data were gathered, the posterior commissure was transected with an operating knife. Before the animal was sacrificed, biocytin, an anterograde tracer, was injected into the left NOT in order to confirm that all of the commissural fibers had been cut. Although the midline lesions decreased the initial rapid rise and steady state OKN slow-phase velocity in all three animals, there were no directional differences observed during monocular clockwise or counter-clockwise visual stimulation to either eye. In two of the three animals, there were no significant differences in the time-constants of optokinetic after nystagmus (OKAN) after the lesion. In the remaining animal, the time-constants decreased at stimulus velocities of 30 degrees/s and 60 degrees/s. In conclusion, gain reduction in the rapid rise and steady state slow-phase velocity of OKN can be explained by removal of an excitatory signal mediated by commissural fibers to inhibitory interneurons in the contralateral NOT. However, interrupting the commissural fibers had no effect on the velocity storage mechanism, because the time-constants of OKAN mostly remained largely unchanged by the lesion.     

Human optokinetic afternystagmus. Slow-phase characteristics and analysis of the decay of slow-phase velocity

Events following the extinction of lights after 1-minute exposures of naive, normal subjects to an optokinetic stimulus at 40 deg/sec have been closely examined and quantified. Mean eye displacement in each slow phase decreased from 10.12 +/- 1.61 deg during optokinetic nystagmus (OKN) to 3.36 +/- 2.32 deg during optokinetic afternystagmus (OKAN). Slow-phase duration increased from 0.26 +/- 0.03 sec during OKN to 0.45 +/- 0.195 sec during OKAN. Eye displacement per slow phase remained fairly constant during OKAN, suggesting a spatial reference for the resetting of gaze. OKAN decay is a two-component process which can be closely approximated by a sum of two exponentials, one with a short time constant of 1.15 sec and the other with a long time constant of 48.8 sec. OKAN decay commenced at a time after lights out which depended upon the presence and timing of an intervening fast phase. When a fast phase intervened, OKAN decay commenced about 230 msec after it, and about 460 msec after lights out. When lights out occurred during the fast phase, OKAN decay commenced about 340 msec later.     

Computer analysis of optokinetic nystagmus in patients with spontaneous nystagmus of peripheral vestibular origin

The bias of slow phase velocity (SPV) of optokinetic nystagmus (OKN) caused by an acute labyrinthine lesion was examined in 8 patients using differnet optokinetic stimulus velocities. In all patients a directional preponderance of OKN-SPV was found corresponding to spontaneous nystagmus. This was due to enhancement of nystagmus SPV to the side of the lesion and depression of SPV in the opposite horizontal direction. The preponderance of OKN on the average increased with the intensity of spontaneous nystagmus and decreased along with recovery. These vestibularly induced differences in OKN-SPV range up to 70%. A differentiation is discussed between OKN preponderances caused by labyrinthine lesions and brain stem lesions.     

Cross-coupling between horizontal and vertical eye movements during optokinetic nystagmus and optokinetic afternystagmus elicited in microgravity

The occurrence of horizontal jerks with larger amplitudes than on Earth was observed during vertical optokinetic nystagmus in astronauts tested throughout a 7-day spaceflight. During early exposure to microgravity, a horizontal spontaneous-like nystagmus was recorded in darkness following both vertical and horizontal optokinetic stimulation. In addition, the time constant of vertical OKAN with slow phase up was larger than on the ground. These effects disappeared on flight day 2. Then, the horizontal and vertical OKAN time constants decreased, and gradually returned to the preflight values, as previously observed with the gain of the vestibulo-ocular reflex. The early changes in microgravity are in agreement with those obtained on Earth in monkeys and humans during static tilt relative to gravity. Our findings suggest that the absence of otolithic input in microgravity may have an effect on the optokinetic system which could be mediated by the velocity storage mechanism.     

Human optokinetic afternystagmus. Charging characteristics and stimulus exposure time dependence in the two-component model

The dependence of human optokinetic afternystagmus (OKAN) velocity storage (charging) and optokinetic nystagmus (OKN) characteristics on optokinetic (OK) stimulus exposure time was investigated, using the two-component double exponential model for OKAN decay. Results are compatible with our previously proposed concept of two velocity storage integrators, one responsible for the short time constant decay (pursuit-mediated) and the other for the long time constant decay (OK system-mediated). The dependence of the long time constant integrator of OKAN on stimulus exposure time was clearly demonstrated. The short time constant integrator appeared to be independent of stimulus exposure time within the range studied. We conclude that the charging time-course of each component is distinct from that of the other. The time constants of each component decay were found to be invariant. A left-right asymmetry observed in both OKN and OKAN responses suggests that the integrators are direction sensitive.     

Effect of alertness and visual attention on optokinetic nystagmus in humans

The effect of alertness and visual attention on optokinetic nystagmus (OKN) and optokinetic after-nystagmus (OKAN) was studied in 20 volunteers. Electroencephalographic (EEG) activity was recorded over the occipital lobe. Exposure to sound and vibration caused a significant increase in the mean slow-phase velocity of OKN, whereas its maximum slow-phase velocity remained unaffected. Vibration tended to increase the mean slow-phase velocity of OKN more than sound did, though the difference was not statistically significant. Vibration also significantly increased the OKAN. When alpha rhythm appeared in the occipital EEG during OKN, the velocity of concurrent slow phases was reduced. However, the periods of alpha rhythm did not differ between the different stimulus conditions. The findings suggest that sound and vibration activate the subcortical optokinetic mechanism, thus causing an increase in the mean velocity of OKN. Abatement of visual attention is reflected in temporary reduction of OKN in conjunction with the appearance of alpha waves and is to be interpreted as transient quiescence of the cortical optokinetic mechanism.     

Effect of macular ablation on vertical optokinetic nystagmus in the squirrel monkey.

After macular ablation in squirrel monkeys, a significant improvement of the slow-phase eye speed of vertical optokinetic nystagmus (VOKN) was found. This findings is in contrast to the result of our previous study in which horizontal optokinetic nystagmus (HOKN) was not improved following macular ablation. The different influence of macular ablation on HOKN and VOKN could be due to the directional characteristics of vestibular and visual interaction relative to the direction of gravity and to the difference in horizontal and vertical vestibulo-oculomotor neural pathways, including vestibular end-organs. The vertical directional dominance which was observed preoperatively was similarly observed in the postablative status; the upward slow-phase nystagmus was easier to provoke than the downward nystagmus by application of an identical stimulus.     

Optokinetic nystagmus and after-nystagmus during a 6 hour bedrest study

CONCLUSIONS: A lengthy alteration of gravity direction produced different effects on the intrinsic horizontal and vertical optokinetic oculomotor systems. OBJECTIVE: To examine both optokinetic nystagmus (OKN) and optokinetic after-nystagmus (OKAN) in a 6 h 6 degrees head-down bedrest study, in which the subjects were kept lying under simulated micro-gravity conditions. SUBJECTS AND METHODS: In six normal healthy adults, we repeatedly (five times) and comparatively studied OKN and OKAN evoked by horizontal and vertical stimuli. Stage 1 was an upright sitting position. During the 6 h bedrest condition, we studied OKN and OKAN in 90 degrees recumbent lateral positions (stages 2, 3, and 4). In stage 5 the subject returned to an upright position. RESULTS: We confirmed that the change in gravity direction had various effects on the condition of OKN and OKAN. Also, we found that it took more than 3 h to reach a desirable level of systemic adaptive modification to the unique environmental condition. We considered that the early change was basically due to the changes in sensory inputs through the otolith organs, and the latter changes represented the adaptive process of the spatial orientation system. During the tilt, the occurrence rates of both horizontal and vertical OKANs were decreased; however, the conditions of these changes were different.     

Suppression of optokinetic velocity storage in humans by static tilt in pitch.

The effects of static tilts about the pitch axis on human horizontal optokinetic after-nystagmus OKAN (HOKAN) were examined. Static tilts in pitch produced tilt-dependent HOKAN suppression. The slow decay (indirect pathway) component (coefficient C and long time constant 1/D) of the two-component model for OKAN was significantly reduced, while the short decay (direct pathway) component (coefficient A and short time constant 1/B) remained invariant as angle of tilt was increased. These results provide further evidence that otolith organ activity can couple to horizontal velocity storage in humans, in accordance with models proposed in the literature.     

Optokinetic nystagmus and after-nystagmus during a 6 hour bedrest study

Conclusions. A lengthy alteration of gravity direction produced different effects on the intrinsic horizontal and vertical optokinetic oculomotor systems. Objective. To examine both optokinetic nystagmus (OKN) and optokinetic after-nystagmus (OKAN) in a 6 h 6° head-down bedrest study, in which the subjects were kept lying under simulated micro-gravity conditions. Subjects and methods. In six normal healthy adults, we repeatedly (five times) and comparatively studied OKN and OKAN evoked by horizontal and vertical stimuli. Stage 1 was an upright sitting position. During the 6 h bedrest condition, we studied OKN and OKAN in 90° recumbent lateral positions (stages 2, 3, and 4). In stage 5 the subject returned to an upright position. Results. We confirmed that the change in gravity direction had various effects on the condition of OKN and OKAN. Also, we found that it took more than 3 h to reach a desirable level of systemic adaptive modification to the unique environmental condition. We considered that the early change was basically due to the changes in sensory inputs through the otolith organs, and the latter changes represented the adaptive process of the spatial orientation system. During the tilt, the occurrence rates of both horizontal and vertical OKANs were decreased; however, the conditions of these changes were different.     

Transfer of optokinetic activity to vestibular nystagmus

Transfer of activity generated by prior optokinetic (OK) stimuli of one minute's duration to nystagmus induced in darkness by a subsequent vestibular stimulus consisting of step velocities to and from 40 degrees/s-1 was studied in 10 normal subjects. Four types of OK stimuli were used: (a) full field 'passive'; (b) full field 'active'; (c) full field in the presence of optic fixation, and (d) small OK drum stimulation. Transfer (T) was evident under all conditions and resulted in an enhancement of the vestibulo-ocular (VO) response when activity from the two stimuli were in the same direction (S) and a suppression when in the opposite (O). Expressed by the equation: Formula See Text. the respective transfer values obtained for the above conditions were (a) 66%, (b) 58%, (c) 22%, and (d) 54%. In all tests, rightward OK drum movement was more effective than leftward. In respect of passive OKN the resultant response can be well represented as the algebraic summation of the expected optokinetic after-nystagmus (OKAN) and the VO response, though opposing OKAN is more effective than enhancing. Passive OKN is more effective than active and this can be accounted for by the small contribution made by retinal slip in the former (the indirect path). Surprisingly, the small drum proved almost as effective as active OKN in terms of transfer. Fixation in the presence of full-field OK stimuli induces a non-directionally specific depression of the subsequent VO response, implying that retinal slip could contribute to the mechanism of VO response suppression.