Eye movement responses to combined linear and angular head movement

Summary Lateral eye movements evoked by linear head motion were evaluated in human subjects by subtracting the eye movement responses to headcentred angular oscillation in the dark, about a vertical axis, from the responses evoked by similar oscillation with the head displaced 30 cm eccentrically from the axis. The centred oscillation gave a purely angular stimulus whereas the eccentric oscillation gave an additional tangential linear acceleration acting laterally to the head. The stimuli used were relatively unpredictable, enveloped sinewaves at 0.02 to 1.2 Hz, 60°/s peak angular velocity, 0.004 to 0.24 g peak tangential acceleration, and subjects were either given no instructions or were told to imagine fixating on targets at 60 cm or 5 m distance. Eye movements of significantly higher velocity were evoked in the eccentric position, particularly at the higher frequencies and when subjects imagined near targets. The increase in velocity of eye movement was attributed to the linear stimulus and probably derives from stimulation of the otolith organs. The frequency response of the gain (°/s/g) of these movements gave an approximate slope of –1, indicating that the eye velocity bears a constant proportionality to linear head velocity. The findings are in accord with the theoretical prediction that eye movements compensating for linear head motion should only be required for viewing near targets. These otolithic influences on eye movements could either the mediated by a direct otolith-ocular reflex which is subservient to viewing conditions or, alternatively, the otolith signals may modify the activity of other oculomotor mechanisms.A. M. Bronstein was supported by The Brain Research Trust     

The influence of target distance on eye movement responses during vertical linear motion

Summary Studies of the linear vestibulo-ocular reflex (LVOR) suggest that eye movement responses to linear head motion are rudimentary. This may be due to inadequate control of target distance (D). As D approaches infinity, eye movements are not required to maintain retinal image stability during linear head displacements, but must become increasingly large as D shortens. The LVOR in the presence of visual targets (VLVOR) was tested by recording human vertical eye and head movements during self-generated vertical linear oscillation (averaging 2.7 Hz at peak excursion of 3.2 cm) while subjects alternately fixated targets at D=36, 142, and 424 cm. Response sensitivity rose from 0.10 deg/cm (5.8 deg/s/g) for D=424 cm to 0.65 deg/cm (37.5 deg/s/g) for D=36 cm. Results employing optical manipulations, including spherical lenses to modify accommodation and accomodative convergence, and prisms to modify fusional vergence without altering accommodation, imply that the state of vergence is the most important variable underlying the effect. Trials in darkness (LVOR) and with head-fixed targets (visual suppression of the LVOR) suggest that, while visual following and perhaps mental set influences results, the major proportion of the VLVOR response is driven by vestibular (presumably otolith) inputs.     

Otolithic-acoustic interaction in the control of eye movement

Summary In order to examine otolithic contribution to eye movements ten subjects were asked to track either a moving acoustic target or a stationary target during subect linear motion on a cart. The relative displacement between the subject and the target was the same in the two situations. Recordings of eye movements during subject lateral acceleration in the dark without any task, or with the task of tracking an imagined stationary target were made as a control.The frequencies ranged between 0.15 and 0.3 Hz and peak acceleration between 0.55 and 1.2 m/s². No lateral eye movements (L-nystagmus) were recorded in the dark. Only saccadic eye movements were recorded during the tracking of a moving acoustic target.Slow eye movements interspersed by saccades were observed when the moving subject tracked an imagined or an acoustic stationary target. Contribution of the slow phase to tracking was more important in the presence of an acoustic target than in the presence of imagined target. The results are interpreted in terms of an otolithic contribution to the central reconstruction of the acoustic target velocity, or in terms of an adaptive control of the otolithic-ocular reflex gain. A conceptual model accounting for these interpretations is proposed.Work supported by CNRS (France) and CNR (Italy)     

Influence of eye and head position on the vestibulo-ocular reflex

Summary For the vestibulo-ocular reflex (VOR) to function properly, namely to ensure a stable retinal image under all circumstances, it should be able to take into account varying eye positions in the orbit and varying orientations of the head with respect to the axis about which it is rotating. We tested this capability by quantifying the gain and the time constant of the horizontal component of the VOR during rotation about an earth vertical axis when the line of sight (optical axis) was moved out of the plane of head rotation — either by rotating the eyes up or down in the orbit or by pitching the head up or down with respect to earth-horizontal. In either case the gain of the horizontal component of the VOR was attenuated precisely by the cosine of the angle made between the optical axis and the plane of head rotation. Furthermore, if the head was pitched up or down but the eye rotated oppositely in the orbit so as to keep the line of sight in the plane of head rotation the gain of the horizontal component of the VOR was the same value as with the head and eyes both straight ahead. In contrast, the time constant of the VOR varied only as a function of the orientation of the head and not as a function of eye position in the orbit. During rotation about an earth vertical axis, the time constant was longest (about 18 s) when the head was pitched forward to place the lateral canals near earth-horizontal and shortest (about 11 s) when the head was pitched backward to place the vertical canals near earth-horizontal. Finally, since during rotation in yaw the pattern of stimulation of the lateral and vertical semicircular canals varies with different head orientations one can use measurements of the horizontal component of the VOR, under varying degrees of pitch of the head, to calculate the relative ability of the lateral and vertical semicircular canals to transduce head velocity.Dr. Fetter is a visiting scientist from the Neurologische Universitätsklinik, Eberhard-Karls-Universität, Liebermeisterstr. 18-20, D-7400 Tübingen, Federal Republic of Germany     

The horizontal vestibulo-ocular reflex during linear acceleration in the frontal plane of the cat

Summary Horizontal and vertical eye movements were recorded in alert, restrained cats that were subjected to whole-body rotations with the horizontal semicircular canals in the plane of rotation and the body centered on the axis or 45 cm eccentric from the axis of rotation. Changes in the horizontal vestibulo-ocular reflex (HVOR) due to the resultant of the linear forces (i.e., gravity and linear acceleration) acting on the otolith organs were examined during off-axis rotation when there was a centripetal acceleration along the animal's interaural axis. The HVOR time constant was slightly shortened when the resultant otolith force was not parallel to the animal's vertical axis. This effect was independent of the direction of the otolith force relative to the direction of the slow phase eye velocity. No effect on the HVOR amplitude was observed. In addition to changes in the HVOR dynamics, a significant vertical component of eye velocity was observed during stimulation of the horizontal canals when the resultant otolith force was not parallel with the animal's vertical axis. The effect was greater for larger angles between the resultant otolith force and gravity. An upward or downward component was elicited, depending on the direction of the horizontal component of eye velocity and the direction of the resultant otolith force. The vertical component was always in the direction that would tend to align the eye velocity vector with the resultant otolith force and keep the eye movement in a plane that had been rotated by the angle between the resultant otolith force and gravity.     

Initiation of the human heave linear vestibulo-ocular reflex

The linear vestibulo-ocular reflex (LVOR) was studied in eight normal human subjects of average age 24±5 years. Subjects underwent a sudden heave (mediolateral) translation delivered by a pneumatic servo-driven chair with a peak acceleration of 0.5 g while viewing earth-fixed targets at 15, 25, 50, and 200 cm. Stimuli were provided both with targets continuously visible or extinguished just prior to motion. Cancellation was tested using chair-fixed targets at each viewing distance. Eye movements were recorded using binocular magnetic search coils. Head translation was measured using a linear accelerometer attached to the upper teeth, to which also was attached a magnetic search coil verifying absence of head rotation. Vergence angles achieved by all subjects were appropriate to interpupillary distance and target distance. Heave translations evoked horizontal ocular rotations in the opposite direction following a brief latency. Latency of the LVOR was determined by automated algorithms based on identification of times when eye position and head acceleration exceeded three standard deviations (SDs) of baseline noise, and was corrected for differing transducer delays. Mean LVOR latency was 30±16 ms (mean ± SD), range 12–53 ms. Slow phase LVOR amplitude was greater for near and less for more distant targets, although all observed responses were suboptimal. Measured 100 ms after head translation onset, mean response was 20% of ideal for the target at 15 cm, 22% at 25 cm, 31% at 50 cm, and 53% at 200 cm. Mean latency was significantly longer than the previously reported values for both the human angular VOR and the monkey LVOR, and had significant inverse correlation with response magnitude. The relatively longer latency of the human LVOR than angular VOR may be tailored to match human head movement dynamics. Electronic Publication     

Short-latency compensatory eye movements associated with a brief period of free fall

The vertical eye movements induced by a brief period of free fall were recorded from three monkeys (Macaca mulatta) using the electromagnetic search-coil technique. Free fall was initiated in total darkness immediately following binocular fixation of one of six target lights located at viewing distances ranging from 20 to 107 cm. Responses consisted of an initial transient downward eye movement (anticompensatory direction) with a latency of a few milliseconds at most followed by a sustained upward (compensatory) eye movement. The early transient was independent of viewing distance and attributed to an artifact, whereas the later component was a linear function of the inverse of the prior viewing distance and attributed to the translational vestibulo-ocular reflex (TVOR). Response latencies for the four nearer viewing distances were determined from the individual eye velocity traces using a computerized algorithm: after removing the initial transient by subtracting the mean response obtained with the most distant viewing, a regression line was fitted to the initial rising phase of the residual response and then extrapolated back to the baseline to determine the onset. When so determined, median latencies for the nearest viewing ranged from 16.4 to 18.5 ms, values appreciably shorter than any in the literature.     

Influence of otolithic stimulation by horizontal linear acceleration on optokinetic nystagmus and visual motion perception

Summary Several studies in the past have demonstrated the existence of an Otolith-Ocular Reflex (OOR) in man, although much less sensitive than canal ocular reflex. The present paper 1 confirms these previous results. Nystagmic eye movements (L-nystagmus) appear in the seated subject during horizontal acceleration along the interaural axis in the dark for an acceleration level (1 m/s²) about ten times the perception threshold with a sensitivity of about 0.035 rad/m.When sinusoidal linear acceleration is combined with optokinetic stimulation, the recorded nystagmus slow phase velocity exhibits strong periodic modulation related to subject motion. This marked effect of linear acceleration on the optokinetic nystagmus (OKN) appears at a level (0.1 m/s²) close to the acceleration perception threshold and has a 4-fold higher sensitivity than L-nystagmus. Modulation of OKN can reach a peak-to-peak amplitude as great as 20 °/s; for a given optokinetic field size it increases with the velocity of the optokinetic stimulus, i.e. with the slow phase eye velocity. In parallel with changes in OKN slow phase velocity, linear acceleration induces a motion related decrease in the perceived velocity of the visual scene and modifications in selfmotion perception.The results are interpreted in terms of a mathematical model of visual-vestibular interaction. They show that sensory interaction processes can magnify the contribution given to the control of eye movements by the otolithic system and provide a way of exploring its function at low levels of acceleration.The present work has been presented at III European Neurosciences Meeting, Rome, September 1979     

Torsional and horizontal vestibular ocular reflex adaptation: three-dimensional eye movement analysis

This study used visual-vestibular conflict to effect short-term torsional and horizontal adaptation of the vestibulo-ocular reflex (VOR). Seven normal subjects underwent sinusoidal whole-body rotation about the earth-vertical axis for 40 min (±37°/s, 0.3 Hz) while viewing a stationary radial pattern fixed to the chair (×0 viewing). During adaptation and testing in darkness, the head was pitched either up or down 35° to excite both the horizontal and torsional VOR. The eyes were kept close to zero orbital elevation. Eye movements were recorded with a dual search coil in a three-field magnetic system. VOR gain was determined by averaging peak eye velocity from ten cycles of chair oscillation in complete darkness. The gain of the angular horizontal VOR (response to rotation about the head rostral-caudal axis) was significantly reduced after training in both head orientations. Angular torsional VOR gain (head rotation about the naso-occipital axis) was reduced in both head orientations, but this reached statistical significance only in the head down position. These results suggest that torsional and horizontal VOR gain adaptation, even when elicited together, may be subject to different influences depending upon head orientation. Differences between head up and down could be due to the relatively greater contribution of the horizontal semicircular canals with nose-down pitch. Alternatively, different VOR-adaptation processes could depend on the usual association of the head down posture to near viewing, in which case the torsional VOR is relatively suppressed.     

Vestibular nerve and nuclei unit responses and eye movement responses to repetitive galvanic stimulation of the labyrinth in the rat

Summary Two-second cathodal current pulses were applied at one-minute intervals at a point external to the round window in the ear of each albino rat subject. Responses were recorded in the vestibular nerve ganglion, the vestibular nuclei (single units), or in the eye movements (search coil recording method) of anaesthetized, decerebrated, or alert rats. The unit responses to the galvanic stimuli were characterized and compared with responses to galvanic and rotational stimuli reported in the literature. The main focus of the study, however, was effects of stimulus repetition. In both the vestibular nerve and vestibular nuclei recordings, the responses of many units were substantially larger or smaller at the end of a 13-pulse stimulus train than at the beginning. In the vestibular nuclei, but not in the nerve, there was a slight bias towards a decrease in response magnitude, with 10/88 units showing decreases great enough to be considered as reflecting an habituation process. In contrast, the eye movement responses showed more consistent response decrements, especially in the alert condition, but also in the other conditions (none of the unit recordings were done in alert rats). It is concluded that some of the modifications underlying habituation of the vestibuloocular reflex probably occur in portions of the neuronal reflex pathways that are downstream from the vestibular nuclei.Prof. Precht died on March 12, 1985     

Vestibular catch-up saccades augmenting the human transient heave linear vestibulo-ocular reflex

Vestibular catch-up saccades (VCUS) cued by the semicircular canals can supplement the deficient angular vestibulo-ocular reflex during transient rotations to stabilize gaze in people with unilateral vestibular deafferentation (Tian et al. 2000). However, a possible analogous role for VCUS to augment a deficient linear vestibulo-ocular reflex (LVOR) has not been carefully studied. We investigated VCUS in 9 younger, 8 older normal, and 12 vestibulopathic subjects undergoing directionally random heave (interaural) translations at 0.5 g peak acceleration. Eye and head movements were sampled at 1,200 Hz using magnetic search coils and a cranial accelerometer. Subjects fixated visible targets 200, 50, or 15 cm distant immediately before unpredictable onset of translation in either darkness or light. Evoked slow phase eye rotations opposite to the direction of head translation accounted for only 19–70% of ideal eye position, being less for nearer targets, and VCUS commonly occurred to augment the deficiency. Eye position error relative to geometric ideal was highly correlated to VCUS amplitude (P<0.001). This error was systematically corrected by VCUS whose latency decreased, and speed and frequency increased, with decreasing target distance. When targets remained visible, nearly all subjects made VCUS for nearer targets. In darkness, VCUS for the nearest target were significantly less common for older normal and vestibulopathic subjects than in younger normal subjects (P<0.001). In older and vestibulopathic subjects, VCUS latency was significantly prolonged. We conclude that otolith-mediated VCUS calibrated to target distance assist LVOR slow phases, but the ability to generate VCUS in darkness is impaired in older normal and vestibulopathic subjects. In the presence of visual information, VCUS can be generated in older and vestibulopathic subjects, albeit at prolonged latency perhaps indicating visual augmentation of deficient vestibular input.     

Visually-induced adaptive plasticity in the human vestibulo-ocular reflex

Summary The vestibulo-ocular reflex (VOR) is under adaptive control which corrects VOR performance when visual-vestibular mismatch arises during head movements. However, the dynamic characteristics of VOR adaptive plasticity remain controversial. In this study, eye movements (coil technique) were recorded from normal human subjects during sinusoidal rotations in darkness before and after 8 h. of adaptation to 2X binocular lenses. The VOR was studied at 7 frequencies between 0.025 and 4.0 Hz at 50°/s peak head velocity (less for 2.5–4 Hz). For 0.025 and 0.25 Hz, the VOR was tested at 4 peak head velocities between 50 and 300° /s. Before 2X lens adaptation, VOR gain was around 0.9 at 2.5–4.0 Hz and dropped gradually with decreasing frequency to under 0.6 at 0.025 Hz. Phase showed a small lead at the highest frequencies which declined to 0° as frequency decreased to 0.5–0.25 Hz, but then rose to 14° by 0.025 Hz. VOR gain was independent of head velocity in the range 50–300°/s at both 0.025 and 0.25 Hz. However, Phase lead rose with increasing head velocity, more so at 0.025 than at 0.25 Hz. After 2X lens adaptation, gain rose across the frequency bandwidth. However, the proportional gain enhancement was frequency dependent; it was greatest at 0.025 Hz (44%), and declined with increasing frequency to reach a minimum at 4 Hz (19%). Phase lead increased after 2X lens adaptation at lower frequencies, but decreased at higher frequencies. New velocity-dependent gain nonlinearities also developed which were not present prior to adaptation; gain declined as peak head velocity increased from 50 to 300°/s at both 0.025 (23% drop) and 0.25 Hz (15% drop). This may suggest an amplitude-dependent limitation in VOR adaptive plasticity. Results indicate both frequency and amplitude dependent nonlinearities in human VOR response dynamics before and after adaptive gain recalibration.     

Single unit activity in the frontal eye fields of unanesthetized monkeys during eye and head movement

Summary Single unit activity in the frontal eye field was investigated in unanesthetized monkeys during eye and head movement. Two types of cells (I and II) were found. Type I fired during voluntary saccadic movement occuring in a given direction and also during the fast phase of optokinetic and vestibular nystagmus. Cells of this type were silent during smooth pursuit movement and the slow phase of nystagmus. It was found that the firing pattern of Type I cells was maintained irrespective of head movement.Type II cells fired during smooth pursuit eye movements and the slow phase of nystagmus; these units displayed a steady discharge when the eyes were oriented in a specific position. Also this type of cell maintained its characteristic discharge during head movement. A separate population of frontal eye field cells was found to be exclusively related to head turning.     

Eye movements induced by lateral acceleration steps Effect of visual context and acceleration levels

 Eye movement responses were obtained from six normal subjects exposed to randomly ordered rightwards/leftwards linear acceleration steps of 0.05 g, 0.1 g or 0.24 g amplitude and 650 ms duration along the inter-aural axis. With the instruction to gaze passively into the darkness, compensatory nystagmus was evoked with slow-phase velocity sensitivity of 49° s⁻¹ g ⁻¹. When subjects viewed earth-fixed targets at 30 cm, 60 cm or 280 cm, eye movements at 130 ms from motion onset were proportional to acceleration and inversely proportional to target distance, before the onset of visually guided eye movements. Our results show that a modulation with viewing distances of the earliest human otolith-ocular reflexes occurs in the presence of pure linear acceleration. However, full compensation was not attained for the nearer targets and higher accelerations. Received: 31 January 1996 / Accepted: 30 September 1996     

Human eye movement response to z-axis linear acceleration: the effect of varying the phase relationships between visual and vestibular inputs

We investigated the effect of systematically varying the phase relationship between 0.5-Hz sinusoidal z-axis optokinetic (OKN) and linear acceleration stimuli upon the resulting vertical eye movement responses of five humans. Subjects lay supine on a linear sled which accelerated them sinusoidally along their z-axis at 0.4 g peak acceleration (peak velocity 1.25 m/s). A high-contrast, striped z-axis OKN stimulus moving sinusoidally at 0.5 Hz, 70°/s peak velocity was presented either concurrently or with the acceleration stimulus or alone. Subjects' vertical eye movements were recorded using scleral search coils. When stimuli were paired in the naturally occurring relationship (e.g., visual stripes moving upward paired with downward physical acceleration), the response was enhanced over the response to the visual stimulus presented alone. When the stimuli were opposed (e.g., visual stripes moving upward during upward physical acceleration, a combination that does not occur naturally), the response was not significantly different from the response to the visual stimulus presented alone. Enhancement was maximized when the velocities of the visual and motion stimuli were in their normal phase relationship, while the response took intermediate values for other phase relationships. The phase of the response depended upon the phase difference between the two inputs. We suggest that linear self-motion processing looks at agreement between the two stimuli — a sensory conflict model.     

Changes in the dynamics of the vertical vestibulo-ocular reflex due to linear acceleration in the frontal plane of the cat

Summary The vertical and horizontal components of the vestibulo-ocular reflex (VOR) were recorded in alert, restrained cats who were placed on their sides and subjected to whole-body rotations in the horizontal plane. The head was either on the axis or 45 cm eccentric from the axis of rotation. During off-axis rotation there was a change in the linear force acting on the otolith organs due to the presence of a centripetal acceleration along the animal's vertical axis. Otolith forces (defined to be opposite to the centripetal acceleration) directed ventrally with respect to the animal (negative) decreased both the amplitude and time constant of the first-order approximation to the slow phase eye velocity of the vertical vestibulo-ocular reflex (VVOR). Otolith forces directed dorsally (positive) increased the amplitude and time constant. The effects were greater for the up VOR. The asymmetry in the VVOR time constant also depended on the otolith forces, being less in the presence of negative otolith forces that caused the resultant otolith force to move ventrally, towards the direction along which gravity normally acts when the animal is in the upright position. The effects of otolith forces on the up VVOR were independent of whether the animals were tested in the dark or in the light with a stationary visual surround (i.e., during visual suppression). In contrast, the changes in the time constant of the down VVOR were smaller during visual suppression. Simulations of the eye velocity storage mechanism suggest that the gain of the feedback in the storage integrator was modified by the angle between the resultant otolith force and an animal-fixed reference. This could be the animal's vertical, i.e., the direction along which gravity normally acts. For larger angles the feedback was less and the amplitude and time constant of the VVOR increased. The transformation of the otolith input was the same for both the up and down VOR, even though the final effect on the eye velocity was asymmetric (larger for up VOR) due to a separate, asymmetric gain element in the velocity storage feedback pathway.     

Discharge properties of neurons in the monkey thalamus tested with angular acceleration,eye movement and visual stimuli

Summary Monkeys were trained to make visually evoked eye movements while undergoing simultaneous head rotation. Single units were recorded in the pregeniculate nucleus (PGN). PGN neurons discharged during each saccade, but there was no change in activity with horizontal head acceleration or with various combinations of head and smooth pursuit eye movements as previously described in the cat. Therefore, the anatomical homology between LGN_v and PGN does not appear to have a neurophysiological basis. Neurons in the oral part of VPL or occasionally in VPI discharged as a function of head velocity but not with saccades, smooth pursuit or fixation eye movements, nor after brief light flashes or during smooth pursuit across structured backgrounds. This suggests that VPL_o and VPI are only vestibular relay nuclei and not concerned with vestibular/visual or vestibular/oculomotor interactions.It is a pleasure to acknowledge the histological talents of Donna Simmons, the veterinary care provided by Stan Crossman and Margaret Price, the surgical assistance of Doug Hasund, the secretarial help of Jean Scalf, and the editorial comments of Kate Schmitt.On leave from Laboratoire de Neuropsychologie Expérimentale, INSERM U 94, 16, av. Doyen Lépine, 69500 Bron, FranceThis research was supported by grants RR00166, GM00260 and EY00745 from the National Institutes of Health, U. S. Public Health Service, and by a grant from INSERM.     

Modulation by horizontal eye position of the vestibulo-collic reflex induced by tilting in the frontal plane in the alert cat

Summary In alert cats, during sinusoidal rotations of head and trunk en bloc around a longitudinal axis, in darkness or in light, the vestibulo-collic reflex induces neck muscle contractions. The phase and gain diagrams are consistent, in the frequency range 0.2 to 1.2 Hz, with previous results from anesthetized or decerebrate cats. In addition, neck muscle contractions are modulated by horizontal eye position, as is the case for rotations in the horizontal plane, around the vertical (Z) axis. Neck muscle contraction is consequently under control of both eye position and head tilt angle. This synergy of eye and head could suppress the effects of vestibulo-collic reflex during orienting reactions.     

Otolith-visual interaction in the control of eye movement produced by sinusoidal vertical linear acceleration in alert cats

Summary 1. Eye movement responses were examined in alert cats during sinusoidal vertical linear acceleration. Stimulus frequencies of 0.20–0.85 Hz with a constant amplitude of 10.5 cm (corresponding to 0.02–0.31 g) were used. A random visual pattern was presented to give sinusoidal vertical optokinetic stimuli of similar amplitude and frequency to the up-down motion of the cat. 2. Sinusoidal linear acceleration in the presence of a stationary visual pattern produced robust eye movement responses with near compensatory phase at all stimulus frequencies tested. With both eyes covered, a vertical linear vestibulo-ocular reflex (LVOR) was frequently produced at a stimulus strength corresponding to 0.04–0.31 g. The evoked LVOR was always small, and the overall mean response phase values advanced by as much as 70 ° at frequencies below 0.56 Hz, indicating that the otolith signals activated by sinusoidal linear acceleration were not, by themselves, converted into compensatory eye position signals under these experimental conditions. 3. Optokinetic stimulation alone produced more lag of response phase as stimulus frequency increased, and the gain of evoked eye movement responses was smaller at higher stimulus frequencies compared to the gain during linear acceleration in the light. Bilateral labyrinthectomies resulted in a significant change of the eye movement responses during linear acceleration when visual inputs were allowed: there was more phase lag at higher stimulus frequencies and a decreased gain at all frequencies tested. These results indicate that the interaction of otolith and visual inputs produces robust eye movement responses with near compensatory phase during sinusoidal linear acceleration in the light.     

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.