Torsional eye movements are facilitated during perception of self-motion

Visual motion in the roll plane elicits torsional optokinetic nystagmus (tOKN) with intermittent periods of illusory, contradirectional self-motion (circularvection, CV). The CV may also have a component of whole-body tilt if the axis of stimulus rotation is not aligned with the direction of gravity. We report how the characteristics of tOKN are affected by the presence of CV. Subjects had their eye movements recorded by VOG whilst viewing a full-field stimulus rotating at 30-60 degrees/s about their naso-occipital axis. They were tested in upright and supine posture and signalled the presence-absence of CV with a pushbutton. In both postures, during CV, tOKN slow-phase gain was found to be enhanced and average torsional eye position shifted in the direction opposite to stimulus rotation. When supine, slow-phase gain was greater than when upright both during the perception of object-motion and during CV. The effects may be explained in terms of a relegation of restraining vestibular input to the torsional oculomotor system during CV and illusory tilt.     

Spatial orientation of the vestibular system: dependence of optokinetic after-nystagmus on gravity.

1. Monkeys received optokinetic stimulation at 60 degrees/s about their yaw (animal vertical) and pitch (animal horizontal) axes, as well as about other head-centered axes in the coronal plane. The animals were upright or tilted in right-side-down positions with regard to gravity. The stimuli induced horizontal, vertical, and oblique optokinetic nystagmus (OKN). OKN was followed by optokinetic after-nystagmus (OKAN), which was recorded in darkness. 2. When monkeys were tilted, stimulation that generated horizontal or yaw axis eye velocity during OKN induced a vertical or pitch component of slow phase velocity during OKAN. This has been designated as "cross-coupling" of OKAN. Eigenvalues and eigenvectors associated with the system generating OKAN were found as a function of tilt. They were determined by use of the Levenberg-Marquardt algorithm to minimize the mean square error between the output of a model of OKAN and the data. 3. The eigenvector associated with yaw OKAN (yaw axis eigenvector) was maintained close to the spatial vertical regardless of the angle of tilt. The eigenvector associated with pitch OKAN (pitch axis eigenvector) was always aligned with the body axis. The data indicate that velocity storage can be modeled by a piecewise linear system, the structure of which is dependent on gravity and the yaw axis eigenvector, which tends to align with gravity. 4. Yaw axis eigenvectors were also determined by giving optokinetic stimulation about head-centered axes in the coronal plane with the animal in various angles of tilt. A technique using a spectral analysis of residuals was developed to estimate whether yaw and pitch OKAN slow phase velocities decayed concurrently at the same relative rate and over the same time course. The eigenvectors determined by this method were in agreement with those obtained by analyzing OKAN elicited by yaw OKN. 5. During yaw OKN with the animal in tilted positions, the mean vector of the ensuing nystagmus was closer to the body axis than to the spatial vertical. This suggests that there is suppression of the cross-coupled pitch component during OKN. The direction of the stimulus may be utilized to suppress components of velocity storage not coincident with the direction of stimulus motion. 6. There were similarities between the monkey eigenvectors and human perception of the spatial vertical, and the mean of eigenvectors for upward and downward eye velocities overlay human 1-g perceptual data.(ABSTRACT TRUNCATED AT 400 WORDS)     

Effect of side-down tilt on optokinetic nystagmus and optokinetic after-nystagmus in cats

Slow phase velocity (SPV) of optokinetic nystagmus (OKN) and after-nystagmus (OKAN) was examined in the cat in side-down tilts. In the upright position, the axis of SPV (direction of SPV vector) during OKN was always close to the stimulus axis. In side-down positions, yaw stimulation induced OKN with the vector deviating from the stimulus axis toward the pitch-axis, so as to make the SPV rotational plane be shifted toward the earth horizontal plane, while pitch-axis stimulation induced no change in vector direction. This yaw-to-pitch cross-coupling is qualitatively similar to that previously described in primates. SPV vector size during yaw stimulation is greatest when the stimulus axis is oriented vertically, and the vector size is smaller when the stimulus is in the gravity direction than when it is in the anti-gravity direction. The SPV vector for the rapid-rise response showed no clear change with head orientation, indicating that the direct optokinetic pathway has no contribution to the induction of cross-coupling. Cross-coupling was found also during OKAN. The SPV trajectories were fitted well using the velocity storage integrator model. SPV vector change of cat OKN/OKAN in head tilt could be fitted by changing gain elements in the model.     

Neural processing of gravito-inertial cues in humans. IV. Influence of visual rotational cues during roll optokinetic stimuli

Sensory systems often provide ambiguous information. For example, otolith organs measure gravito-inertial force (GIF), the sum of gravitational force and inertial force due to linear acceleration. However, according to Einstein's equivalence principle, a change in gravitational force due to tilt is indistinguishable from a change in inertial force due to translation. Therefore the central nervous system (CNS) must use other sensory cues to distinguish tilt from translation. For example, the CNS might use dynamic visual cues indicating rotation to help determine the orientation of gravity (tilt). This, in turn, might influence the neural processes that estimate linear acceleration, since the CNS might estimate gravity and linear acceleration such that the difference between these estimates matches the measured GIF. Depending on specific sensory information inflow, inaccurate estimates of gravity and linear acceleration can occur. Specifically, we predict that illusory tilt caused by roll optokinetic cues should lead to a horizontal vestibuloocular reflex compensatory for an interaural estimate of linear acceleration, even in the absence of actual linear acceleration. To investigate these predictions, we measured eye movements binocularly using infrared video methods in 17 subjects during and after optokinetic stimulation about the subject's nasooccipital (roll) axis (60 degrees /s, clockwise or counterclockwise). The optokinetic stimulation was applied for 60 s followed by 30 s in darkness. We simultaneously measured subjective roll tilt using a somatosensory bar. Each subject was tested in three different orientations: upright, pitched forward 10 degrees, and pitched backward 10 degrees. Five subjects reported significant subjective roll tilt (>10 degrees ) in directions consistent with the direction of the optokinetic stimulation. In addition to torsional optokinetic nystagmus and after nystagmus, we measured a horizontal nystagmus to the right during and following clockwise (CW) stimulation and to the left during and following counterclockwise (CCW) stimulation. These measurements match predictions that subjective tilt in the absence of real tilt should induce a nonzero estimate of interaural linear acceleration and, therefore, a horizontal eye response. Furthermore, as predicted, the horizontal response in the dark was larger for Tilters (n = 5) than for Non-Tilters (n = 12).     

The effects of head and trunk position on torsional vestibular and optokinetic eye movements in humans

We measured torsional vestibular and optokinetic eye movements in human subjects with the head and trunk erect, with the head supine and the trunk erect, and with the head and trunk supine, in order to quantify the effects of otolithic and proprioceptive modulation. During active head movements, the torsional vestibulo-ocular reflex (VOR) had significantly higher gain with the head upright than with the head supine, indicating that dynamic otolithic inputs can supplement the semicircular canal-ocular reflex. During passive earth-vertical axis rotation, torsional VOR gain was similar with the head and trunk supine and with the head supine and the trunk erect. This finding implies that static proprioceptive information from the neck and trunk has little effect upon the torsional VOR. VOR gain with the head supine was not increased by active, self-generated head movement compared with passive, whole body rotation, indicating that the torsional VOR is not augmented by dynamic proprioceptive inputs or by an efference copy of a command for head movement. Viewing earth-fixed surroundings enhanced the torsional VOR, while fixating a chair-fixed target suppressed the VOR, especially at low frequencies. Torsional optokinetic nystagmus (OKN) evoked by a full-field stimulus had a mean slow-phase gain of 0.22 for 10°/s drum rotation, but gain fell to 0.06 for 80°/s stimuli. Despite this fall in gain, mean OKN slow-phase velocities increased with drum speed, reaching maxima of 2.5°/s–8.0°/s in our subjects. Optokinetic afternystagmus (OKAN) was typically absent. Torsional OKN and OKAN were not modified by otolithic or proprioceptive changes caused by altering head and trunk position with respect to gravity. Torsional velocity storage is negligible in humans, regardless of head orientation.Presented in part at the Society for Neuroscience Annual Meeting, October 31, 1989, Phoenix, AZ     

The control and coordination of one-handed catching: the effect of temporal constraints

A transparent motion condition occurs when two different motion vectors appear at the same region of an image. Such transparency during self-motion has shown demonstrable effects on perception and on the underlying neurophysiology in the cortical and subcortical structures of primates. Presumably such stimulus conditions also influence oculomotor behavior. We investigated smooth-pursuit performance, using a transparent stimulus consisting of two oppositely-moving patterns. We found slight reduction in the mean eye velocity tracking a transparent pattern, compared with that when tracking a unidirectional pattern. Additionally, we investigated the behavior of the optokinetic system to transparency, demonstrating that it elicits antagonistic optokinetic nystagmus, with distinctly reduced gain of the slow phases. Furthermore, we observed, during optokinetic stabilization of transparent stimuli, directional dominances demonstrating that subjects preferably followed one direction. Presenting a transparent stimulus with oppositely moving patterns and different velocities we found a general velocity dominance demonstrating that patterns with a certain velocity are preferred. Performing all experiments under dichoptic conditions produced results comparable with those found under transparent stimulus conditions.     

Human ocular counterroll: assessment of static and dynamic properties from electromagnetic scleral coil recordings

Summary Static and dynamic components of ocular counterroll as well as cyclorotatory optokinetic nystagmus were measured with a scleral search coil technique. Static counterroll compensated for about 10% of head roll when the head was tilted to steady positions up to 20 deg from the upright position. The dynamic component of counterroll, which occurs only while the head is moving, is much larger. It consists of smooth compensatory cyclorotation opposite to the head rotation, interrupted frequently by saccades moving in the same direction as the head. During voluntary sinusoidal head roll, cyclorotation compensated from 40% to more than 70% of the head motion. In the range 0.16 to 1.33 Hz, gain increased with frequency and with the amount of visual information. The lowest values were found in darkness. The gain increased in the presence of a visual fixation point and a further rise was induced by a structured visual pattern. Resetting saccades were made more frequently in the dark than in the light. These saccades were somewhat slower than typical horizontal saccades. Cyclorotatory optokinetic nystagmus could be induced by a patterned disk rotating around the visual axis. It was highly variable even within a same subject and had in general a very low gain (mean value about 0.03 for stimulus velocities up to 30 deg/s). It is concluded that cyclorotational slip velocity on the retina is considerably reduced by counterroll during roll of the head, although the residual cyclorotation after the head has reached a steady position is very small.     

Difference between horizontal and vertical optokinetic nystagmus in cats at upright position

The slow-phase velocity (SPV) of optokinetic nystagmus (OKN) and optokinetic after nystagmus (OKAN) in response to a velocity step of surround rotation in the horizontal direction is composed of the rapid and slow components in the cat: a rapid rise, a slow rise to a steady state, a rapid fall, and a slow decline to 0 deg/s. The rapid and slow components are attributed to the direct pathway and velocity storage neuronal mechanisms, respectively. The difference between horizontal and vertical OKN has been reported in the monkey at the upright position, but the slow and rapid components have not been distinguished. The present study compared horizontal OKN-OKAN with vertical OKN-OKAN in the cat at the upright position, distinguishing the rapid and slow components. Constant velocity rotation of a random dot pattern at a velocity of 5 to 160 deg/s was used for optokinetic stimulation. The results: First, the amplitude of the rapid rise was relatively small in all SPV directions and all stimulus velocities investigated, with a slight upward-SPV preference to the downward-SPV (maximum 6.4, 6.0, and 3.4 deg/s in horizontal, upward, and downward SPV directions, respectively). Second, the steady state velocity was large during horizontal OKN (maximum 69.0 deg/s), small during upward-SPV OKN (12.9 deg/s), and missing (SPV is negligibly small and irregular) during downward-SPV OKN, indicating a large directional difference of OKN. Third, the acceleration of the slow rise decreased with the stimulus velocity at higher stimulus velocities >20 deg/s during both horizontal and upward-SPV OKN, suggesting strong nonlinearity in the velocity charge system. Fourth, the decay time course of the OKAN was described by the time constant of the exponential function, and the time constant was longer during horizontal (mean, 8.3 s at a stimulus velocity of 20 deg/s) than during upward-SPV (5.4 s) OKAN, suggesting that the velocity discharge system is relatively linear compared with the velocity charge system. It is concluded that horizontal OKN-OKAN is much larger than vertical OKN-OKAN in the cat at the upright position, and this directional difference is caused mainly by the directional difference in the velocity storage mechanism, but not in the direct pathway mechanism.     

Functional organization of the mechanisms subserving the optokinetic nystagmus in the cat

In the cat both crossed and uncrossed retinal fibres are able to mediate optokinetic nystagmus in both temporonasal and nasotemporal directions. There exists, however, a slight directional predominance of the nystagmus for the crossed fibre system in the temporonasal stimulus direction and for the uncrossed fibres in a nasotemporal direction.During the first 9 days following ablation of the visual cortex the optokinetic nystagmus elicited monocularly is greatly asymmetrical: the nystagmus elicited by a temporonasal stimulus is moderately affected particularly at higher stimulus velocities, whereas the nystagmus elicited by a nasotemporal stimulus is present only at stimulus velocities below 20–30 deg/s and has a low gain.Without the visual cortex, selective stimulation of the crossed retinal fibres of one eye may evoke a weak nystagmus on temporonasal stimulus motion only. In contrast, in absence of visual cortex, the uncrossed retinal fibres do not mediate any optokinetic nystagmus. The behaviour of the vestibulo-ocular reflex in light and of the visual fixation suppression of the postrotatory nystagmus in our lesioned animals provided another means to reach similar conclusions. Twenty-seven units recorded in the vestibular nuclei showed responses to optokinetic stimulations, which were in line with the behaviour of the optokinetic nystagmus.These data suggest that optokinetic nystagmus has two components: (i), a subcortical component in which the temporonasal direction of stimulation is predominant in eliciting the nystagmus and in which both the crossed and uncrossed retinal fibres are involved, although with a different weight, and (ii), a cortical component responsible for a symmetrical optomotor response, which also involves the crossed and uncrossed retinal fibres.     

Vestibular and optokinetic eye movements evoked in the cat by rotation about a tilted axis

Summary Horizontal and vertical eye movements were recorded from cats in response to either a) off-vertical axis rotation (OVAR) at a range of velocities (5–72 deg/s) and a range of tilts (0–60 deg) or b) horizontal (with respect to the cat) optokinetic stimulation (10–80 deg/s), also around a range of tilted axes (0–60 deg). The responses to stopping either of these stimuli were also measured: post-rotatory nystagmus (PRN) following actual rotation, and optokinetic after nystagmus (OKAN) following optokinetic stimulation. The response found during OVAR was a nystagmus with a bias slow-phase velocity that was sinusoidally modulated. The bias was dependent on the tilt and reached 50% of its maximum velocity (maximum was 73±23% of the table velocity) at a tilt of 16 deg. The phase of modulation in horizontal eye velocity bore no consistent relation to the angular rotation. The amplitude of this modulation was roughly correlated with the bias with a slope of 0.13 (deg/s) modulation/(deg/s) bias velocity. There was also a low-velocity vertical bias with the slow-phases upwardly directed. The vertical bias was also modulated and the amplitude depended on the bias velocity (0.27 (deg/s) modulation/ (deg/s) bias velocity). When separated from the canal dependent response, the build up of the OVAR response had a time constant of 5.0±0.8 s. Following OVAR there was no decline in the time constant of PRN which remained at the value measured during earth-vertical axis rotation (EVAR) (6.3±2 s). The peak amplitude of PRN was reduced, dependent on the tilt, reaching only 20% of its EVAR value for a tilt of 20 deg. When a measurable PRN was found, it was accompanied by a slowly-emerging vertical component (time constant 5.4±2s) the effect of which was to vector the PRN accurately onto the earth horizontal. OKN measured about a tilted axis showed no differences in magnitude or direction from EVAR OKN even for tilts as large as 60 deg. OKAN following optokinetic stimulation around a tilted axis appeared normal in the horizontal plane (with respect to the animal) but was accompanied by a slowly emerging (time constant 4.1±2 s) vertical component, the effect of which was to vector the overall OKAN response onto the earth horizontal for tilts less than 20 deg. These results are compared with data from monkey and man and discussed in terms of the involvement of the velocity storage mechanism.     

Orientation of human optokinetic nystagmus to gravity: a model-based approach

Optokinetic nystagmus (OKN) was induced by having subjects watch a moving display in a binocular, head-fixed apparatus. The display was composed of 3.3° stripes moving at 35°/s for 45 s. It subtended 88° horizontally by 72° vertically of the central visual field and could be oriented to rotate about axes that were upright or tilted 45° or 90°. The head was held upright or was tilted 45° left or right on the body during stimulation. Head-horizontal (yaw axis) and head-vertical (pitch axis) components of OKN were recorded with electro-oculography (EOG). Slow phase velocity vectors were determined and compared with the axis of stimulation and the spatial vertical (gravity axis). With the head upright, the axis of eye rotation during yaw axis OKN was coincident with the stimulus axis and the spatial vertical. With the head tilted, a significant vertical component of eye velocity appeared during yaw axis stimulation. As a result the axis of eye rotation shifted from the stimulus axis toward the spatial vertical. Vertical components developed within 1–2 s of stimulus onset and persisted until the end of stimulation. In the six subjects there was a mean shift of the axis of eye rotation during yaw axis stimulation of 18° with the head tilted 45° on the body. Oblique optokinetic stimulation with the head upright was associated with a mean shift of the axis of eye rotation toward the spatial vertical of 9.2°. When the head was tilted and the same oblique stimulation was given, the axis of eye rotation rotated to the other side of the spatial vertical by 5.4°. This counterrotation of the axis of eye rotation is similar to the Müller (E) effect, in which the perception of the upright is counterrotated to the opposite side of the spatial vertical when subjects are tilted in darkness. The data were simulated by a model of OKN with a direct and indirect pathway. It was assumed that the direct visual pathway is oriented in a body, not a spatial frame of reference. Despite the short optokinetic after-nystagmus time constants, strong horizontal to vertical cross-coupling could be produced if the horizontal and vertical time constants were in proper ratio and there were no suppression of nystagmus in directions orthogonal to the stimulus direction. The model demonstrates that the spatial orientation of OKN can be achieved by restructuring the system matrix of velocity storage. We conclude that an important function of velocity storage is to orient slow-phase velocity toward the spatial vertical during movement in a terrestrial environment.     

Effects of tilt of the gravito-inertial acceleration vector on the angular vestibuloocular reflex during centrifugation.

Effects of tilt of the gravito-inertial acceleration vector on the angular vestibuloocular reflex during centrifugation. Interaction of the horizontal linear and angular vestibuloocular reflexes (lVOR and aVOR) was studied in rhesus and cynomolgus monkeys during centered rotation and off-center rotation at a constant velocity (centrifugation). During centered rotation, the eye velocity vector was aligned with the axis of rotation, which was coincident with the direction of gravity. Facing and back to motion centrifugation tilted the resultant of gravity and linear acceleration, gravito-inertial acceleration (GIA), inducing cross-coupled vertical components of eye velocity. These components were upward when facing motion and downward when back to motion and caused the axis of eye velocity to reorient from alignment with the body yaw axis toward the tilted GIA. A major finding was that horizontal time constants were asymmetric in each monkey, generally being longer when associated with downward than upward cross coupling. Because of these asymmetries, accurate estimates of the contribution of the horizontal lVOR could not be obtained by simply subtracting horizontal eye velocity profiles during facing and back to motion centrifugation. Instead, it was necessary to consider the effects of GIA tilts on velocity storage before attempting to estimate the horizontal lVOR. In each monkey, the horizontal time constant of optokinetic after-nystagmus (OKAN) was reduced as a function of increasing head tilt with respect to gravity. When variations in horizontal time constant as a function of GIA tilt were included in the aVOR model, the rising and falling phases of horizontal eye velocity during facing and back to motion centrifugation were closely predicted, and the estimated contribution of the compensatory lVOR was negligible. Beating fields of horizontal eye position were unaffected by the presence or magnitude of linear acceleration during centrifugation. These conclusions were evaluated in animals in which the low-frequency aVOR was abolished by canal plugging, isolating the contribution of the lVOR. Postoperatively, the animals had normal ocular counterrolling and horizontal eye velocity modulation during off-vertical axis rotation (OVAR), suggesting that the otoliths were intact. No measurable horizontal eye velocity was elicited by centrifugation with angular accelerations 相似文献   

Eye movements induced by off-vertical axis rotation (OVAR) at small angles of tilt

Summary Off-vertical rotation (OVAR) in darkness induced continuous horizontal nystagmus in humans at small tilts of the rotation axis (5 to 30 degrees). The horizontal slow eye velocity had two components: a mean velocity in the direction opposite to head rotation and a sinusoidal modulation around the mean. Mean velocity generally did not exceed 10 deg/s, and was less than or equal to the maximum velocity of optokinetic after-nystagmus (OKAN). Both the mean and modulation components of horizontal nystagmus increased with tilt angle and rotational velocity. Vertical slow eye velocity was also modulated sinusoidally, generally around zero. The amplitude of the vertical modulation increased with tilt angle, but not with rotational velocity. In addition to modulations in eye velocity, there were also modulations in horizontal and vertical eye positions. These would partially compensate for head position changes in the yaw and pitch planes during each cycle of OVAR. Modulations in vertical eye position were regular, increased with increases in tilt angle and were separated from eye velocity by 90 deg. These results are compatible with the interpretation that, during OVAR, mean slow velocity of horizontal nystagmus is produced by the velocity storage mechanism in the vestibular system. In addition, they indicate that the otolith organs induce compensatory eye position changes with regard to gravity for tilts in the pitch, yaw and probably also the roll planes. Such compensatory changes could be utilized to study the function of the otolith organs. A functional interpretation of these results is that nystagmus attempts to stabilize the image on the retina of one point of the surrounding world. Mean horizontal velocity would then be opposite to the estimate of head rotational velocity provided by the output of the velocity storage mechanism, as charged by an otolithic input during OVAR. In spite of the lack of actual translation, an estimate of head translational velocity could, in this condition, be constructed from the otolithic signal. The modulation in horizontal eye position would then be compensatory for the perceived head translation. Modulation of vertical eye velocity would compensate for actual changes in head orientation with respect to gravity.     

Stimulation of the nodulus and uvula discharges velocity storage in the vestibulo-ocular reflex

The nodulus and sublobule d of the uvula of rhesus and cynomolgus monkeys were electrically stimulated with short trains of pulses to study changes in horizontal slow-phase eye velocity. Nodulus and uvula stimulation produced a rapid decline in horizontal slow phase velocity, one aspect of the spatial reorientation of the axis of eye rotation that occurs when the head is tilted with regard to gravity during per- and post-rotatory nystagmus and optokinetic after-nystagmus (OKAN). Nodulus and uvula stimulation also reproduced the reduction of the horizontal time constant of post-rotatory nystagmus and OKAN that occurs during visual suppression. The brief electric stimuli (4–5 s) induced little slow-phase velocity and had no effect on the initial jump in eye velocity at the onset or the end of angular rotation. Effects of stimulation were unilateral, suggesting specificity of the output pathways. Activation of more caudal sites in the uvula produced nystagmus with a rapid rise in eye velocity, but the effects did not outlast the stimulus and did not affect VOR or OKAN time constants. Thus, stimulation of caudal parts of the uvula did not affect eye velocity produced by velocity storage. We postulate that the nodulus and sublobule d of the uvula control the time constant of the yaw axis (horizontal) component of slow-phase eye velocity produced by velocity storage.     

Effect of semicircular canal stimulation on the perception of the visual vertical

The subjective visual vertical (SVV) is usually considered a measure of otolith function. Herewith we investigate the influence of semicircular canal (SCC) stimulation on the SVV by rotating normal subjects in yaw about an earth-vertical axis, with velocity steps of +/- 90 degrees /s, for 60 s. SVV was assessed by setting an illuminated line to perceived earth vertical in darkness, during a per- and postrotary period. Four head positions were tested: upright, 30 degrees backward (chin up) or forward, and approximately 40 degrees forward from upright. During head upright/backward conditions, a significant SVV tilt (P < 0.01) in the direction opposite to rotation was found that reversed during postrotary responses. The rotationally induced SVV tilt had a time constant of decay of approximately 30 s. Rotation with the head 30 degrees forward did not affect SVV, whereas the 40 degrees forward tilt caused a direction reversal of SVV responses compared with head upright/backward. Spearman correlation values (Rho) between individual SCC efficiencies in different head positions and mean SVV tilts were 0.79 for posterior, 0.34 for anterior, and - 0.80 for horizontal SCCs. Three-dimensional video-oculography showed that SVV and torsional eye position measurements were highly correlated (0.83) and in the direction opposite to the slow phase torsional vestibuloocular reflex. In conclusion: 1) during yaw axis rotation without reorientation of the head with respect to gravity, the SVV is influenced by SCC stimulation; 2) this effect is mediated by the vertical SCCs, particularly the posterior SCCs; 3) rotationally induced SVV changes are due to torsional ocular tilt; 4) SVV and ocular tilts occur in the "anticompensatory," fast phase direction of the torsional nystagmus; and 5) clinically, abnormal SVV tilts cannot be considered a specific indication of otolith system dysfunction.     

Pitch body orientation influences the perception of self-motion direction induced by optic flow

We studied the effect of static pitch body tilts on the perception of self-motion direction induced by a visual stimulus. Subjects were seated in front of a screen on which was projected a 3D cluster of moving dots visually simulating a forward motion of the observer with upward or downward directional biases (relative to a true earth horizontal direction). The subjects were tilted at various angles relative to gravity and were asked to estimate the direction of the perceived motion (nose-up, as during take-off or nose-down, as during landing). The data showed that body orientation proportionally affected the amount of error in the reported perceived direction (by 40% of body tilt magnitude in a range of ±20°) and these errors were systematically recorded in the direction of body tilt. As a consequence, a same visual stimulus was differently interpreted depending on body orientation. While the subjects were required to perform the task in a geocentric reference frame (i.e., relative to a gravity-related direction), they were obviously influenced by egocentric references. These results suggest that the perception of self-motion is not elaborated within an exclusive reference frame (either egocentric or geocentric) but rather results from the combined influence of both.     

Tilt and translation motion perception during off-vertical axis rotation

The effect of stimulus frequency on tilt and translation motion perception was studied during constant velocity off-vertical axis rotation (OVAR), and compared to the effect of stimulus frequency on eye movements. Fourteen healthy subjects were rotated in darkness about their longitudinal axis 10° and 20° off-vertical at 45°/s (0.125 Hz) and 20° off-vertical at 180°/s (0.5 Hz). Perceived motion was evaluated using verbal reports and a joystick capable of recording tilt and translation in both sagittal and lateral planes. Eye movements were also recorded using videography. At the lower frequency, subjects reported the perception of progressing along the edge of a cone, whereas at the higher frequency they had the sensation of progressing along the edge of an upright cylinder. Tilt perception and ocular torsion significantly increased as the tilt angle increased from 10° to 20° at the lower frequency, and then decreased at the higher frequency. The phase lag of ocular torsion increased as a function of frequency, while the phase lag of tilt perception did not change. Horizontal eye movements were small at the lower frequency and showed a phase lead relative to the linear acceleration stimulus. While the phase lead of horizontal eye movements decreased at 0.5 Hz, the phase of translation perception did not vary with stimulus frequency and was similar to the phase of tilt perception during all conditions. A second data set was obtained in 12 subjects to compare motion perception phase when using a simple push-button to indicate nose-up orientation, continuous setting of pitch tilt alone, or continuous setting of tilt and translation in both pitch and roll planes as in the first data set. This set of measurements indicated that in the frequency range studied subjects tend to lead the stimulus when using a push-button task while lagging the stimulus when using a continuous setting of tilt with a joystick. Both amplitude and phase of tilt perception using the joystick were not different whether concentrating on pitch tilt alone or attempting a more complex reporting of tilt and translation in both sagittal and lateral planes. During dynamic linear stimuli in the absence of canal and visual input, a change in stimulus frequency alone elicits similar changes in the amplitude of both self-motion perception and eye movements. However, in contrast to the eye movements, the phase of both perceived tilt and translation motion is not altered by stimulus frequency over this limited range. These results are consistent with the hypothesis that neural processing to distinguish tilt and translation stimuli differs between eye movements and motion perception.     

Nystagmus induced by off-vertical rotation axis in the cat

Summary 1) In the alert cat, nystagmus induced by off-vertical axis rotation (OVAR) was recorded following steps in head velocity or ramps of velocity at constant acceleration below canal threshold. Dependence of nystagmus characteristics on tilt angle of rotation axis and head velocity was studied. Similar results were obtained with both types of stimulation. 2) Mean and modulation amplitude of horizontal eye velocity increased with tilt angle in the range 0–30 degrees. 3) Both variables increased also with head velocity, but with different trends, probably because they are set by different mechanisms. When head rotational velocity was increased above 80°/s, mean eye velocity progressively decreased to zero. 4) In spite of variations from one animal to another, some regularity was observed in the phase of eye velocity modulation. In several cases, a reduction in phase lead of eye velocity with respect to conventional origin of phases (nose-down position) was observed when head velocity increased. 5) Time constant of post-OVAR nystagmus decreased with the tilt angle of the rotation axis from gravity, but not with the orientation of the head with respect to rotation axis. 6) The results could be accounted for by a general equation describing the vestibulo-ocular reflex, provided that estimates of kinematic variables of head movement (head rotational and translational velocities), and visual target distance could be computed by the Central Nervous System.     

Compensatory and orienting eye movements induced by off-vertical axis rotation (OVAR) in monkeys

Nystagmus induced by off-vertical axis rotation (OVAR) about a head yaw axis is composed of a yaw bias velocity and modulations in eye position and velocity as the head changes orientation relative to gravity. The bias velocity is dependent on the tilt of the rotational axis relative to gravity and angular head velocity. For axis tilts <15 degrees, bias velocities increased monotonically with increases in the magnitude of the projected gravity vector onto the horizontal plane of the head. For tilts of 15-90 degrees, bias velocity was independent of tilt angle, increasing linearly as a function of head velocity with gains of 0.7-0.8, up to the saturation level of velocity storage. Asymmetries in OVAR bias velocity and asymmetries in the dominant time constant of the angular vestibuloocular reflex (aVOR) covaried and both were reduced by administration of baclofen, a GABA(B) agonist. Modulations in pitch and roll eye positions were in phase with nose-down and side-down head positions, respectively. Changes in roll eye position were produced mainly by slow movements, whereas vertical eye position changes were characterized by slow eye movements and saccades. Oscillations in vertical and roll eye velocities led their respective position changes by approximately 90 degrees, close to an ideal differentiation, suggesting that these modulations were due to activation of the orienting component of the linear vestibuloocular reflex (lVOR). The beating field of the horizontal nystagmus shifted the eyes 6.3 degrees /g toward gravity in side down position, similar to the deviations observed during static roll tilt (7.0 degrees /g). This demonstrates that the eyes also orient to gravity in yaw. Phases of horizontal eye velocity clustered ~180 degrees relative to the modulation in beating field and were not simply differentiations of changes in eye position. Contributions of orientating and compensatory components of the lVOR to the modulation of eye position and velocity were modeled using three components: a novel direct otolith-oculomotor orientation, orientation-based velocity modulation, and changes in velocity storage time constants with head position re gravity. Time constants were obtained from optokinetic after-nystagmus, a direct representation of velocity storage. When the orienting lVOR was combined with models of the compensatory lVOR and velocity estimator from sequential otolith activation to generate the bias component, the model accurately predicted eye position and velocity in three dimensions. These data support the postulates that OVAR generates compensatory eye velocity through activation of velocity storage and that oscillatory components arise predominantly through lVOR orientation mechanisms.     

Arthrokinetic nystagmus and ego-motion sensation

Summary A compelling illusion of body rotation and nystagmus can be induced when the horizontally extended arm of a stationary subject is passively rotated about a vertical axis in the shoulder joint.Lateral nystagmus with the fast phase beating in the opposite direction to the arm movement was found consistently; the mean slow phase velocity increased with increasing actual arm velocity and reached about 15 °/sec; the mean position of the eyes was deviated towards the fast phase as in optokinetic nystagmus, and the nystagmus continued after the cessation of stimulation (arthrokinetic after-nystagmus).The existence of an arthrokinetic circularvection and nystagmus indicates a convergence of vestibular and somatosensory afferents from joint receptors. It is concluded that information about joint movements plays an important role within the multisensory processes of self-motion perception.Research was supported by Deutsche Forschungsgemeinschaft, Br. 639/1 Bewegungskrankheit