Canal-ocular Reflex Research Articles

Vestibulo-Ocular Responses, Visual Field Dependence, and Motion Sickness in Aerobatic Pilots.

BACKGROUND: Aerobatic flight is a challenge for the vestibular system, which is likely to lead to adaptive changes in the vestibular responses of pilots. We investigated whether aerobatic pilots, as individuals who experience intense vestibular stimulation, present modifications of the vestibular-ocular reflex, motion sickness susceptibility and intensity, visual vertical estimation, and visual dependence as compared to normal volunteers.METHODS: To evaluate vestibulo-ocular reflexes, eye movements were recorded with videonystagmography while subjects were rotated on a rotatory chair with the axis of rotation being vertical (canal-ocular reflex) or inclined to 17° (otolith-ocular reflex). Motion sickness was evaluated after the rotatory test using the Graybiel diagnostic criteria. General motion sickness susceptibility and visual field dependence were also evaluated.RESULTS: Averaged data did not show significant difference in canal-ocular reflex and otolith ocular-reflex between groups. However, a significant asymmetry in otolith-driven ocular responses was found in pilots (CW 0.50 ± 1.21° · s-1 vs. CCW 1.59 ± 1.12° · s-1), though visual vertical estimation was not altered in pilots and both groups were found field independent. Pilots were generally less susceptible to motion sickness (MSSQ scores: 2.52 ± 5.59 vs. 13.5 ± 11.36) and less affected by the nauseogenic stimulation (Graybiel diagnostic criteria 3.36 ± 3.81 vs. 8.39 ± 7.01).DISCUSSION: We did not observe the expected habituation in the group of aerobatic pilots. However, there was a significant asymmetry in the otolith-driven ocular responses in pilots, but not in the controls, which may result from the asymmetry in piloting protocols.Kuldavletova O, Tanguy S, Denise P, Quarck G. Vestibulo-ocular responses, visual field dependence, and motion sickness in aerobatic pilots. Aerosp Med Hum Perform. 2020; 91(4):326-331.

Binocular 3D otolith-ocular reflexes: responses of chinchillas to prosthetic electrical stimulation targeting the utricle and saccule.

From animal experiments by Cohen and Suzuki et al. in the 1960s to the first-in-human clinical trials now in progress, prosthetic electrical stimulation targeting semicircular canal branches of the vestibular nerve has proven effective at driving directionally appropriate vestibulo-ocular reflex eye movements, postural responses, and perception. That work was considerably facilitated by the fact that all hair cells and primary afferent neurons in each canal have the same directional sensitivity to head rotation, the three canals' ampullary nerves are geometrically distinct from one another, and electrically evoked three-dimensional (3D) canal-ocular reflex responses approximate a simple vector sum of linearly independent components representing relative excitation of each of the three canals. In contrast, selective prosthetic stimulation of the utricle and saccule has been difficult to achieve, because hair cells and afferents with many different directional sensitivities are densely packed in those endorgans and the relationship between 3D otolith-ocular reflex responses and the natural and/or prosthetic stimuli that elicit them is more complex. As a result, controversy exists regarding whether selective, controllable stimulation of electrically evoked otolith-ocular reflexes (eeOOR) is possible. Using micromachined, planar arrays of electrodes implanted in the labyrinth, we quantified 3D, binocular eeOOR responses to prosthetic electrical stimulation targeting the utricle, saccule, and semicircular canals of alert chinchillas. Stimuli delivered via near-bipolar electrode pairs near the maculae elicited sustained ocular countertilt responses that grew reliably with pulse rate and pulse amplitude, varied in direction according to which stimulating electrode was employed, and exhibited temporal dynamics consistent with responses expected for isolated macular stimulation.NEW & NOTEWORTHY As the second in a pair of papers on Binocular 3D Otolith-Ocular Reflexes, this paper describes new planar electrode arrays and vestibular prosthesis architecture designed to target the three semicircular canals and the utricle and saccule. With this technological advancement, electrically evoked otolith-ocular reflexes due to stimulation via utricle- and saccule-targeted electrodes were recorded in chinchillas. Results demonstrate advances toward achieving selective stimulation of the utricle and saccule.

Is horizontal semicircular canal ocular reflex influenced by otolith organs input?

Conclusion: Otolith organs input influences the axis of horizontal semicircular canal ocular reflex; therefore, the plane of compensatory eye movements induced by the horizontal canal stimulation is not always parallel to the canal. Objective: To clarify whether horizontal canal ocular reflex is influenced by otolith organs input. Methods: The subjects were seven healthy humans. The right ear was stimulated using ice-water. Each subject was kept in a left-ear-down position for 20 s and then repositioned to a prone position, a right-ear-down position and a supine position with 20 s intervals. Nystagmus was analysed using three-dimensional video-oculography. Results: Eye movements in the supine position and the prone position were not in a symmetric fashion. Nystagmus in the left-ear-down position and the right-ear-down position were not symmetric either. These phenomena indicate that the axis of the eyeball rotation was affected by the shift of the direction of gravity exerted on the head.

Persistent direction-changing geotropic positional nystagmus

The aims of the study were to clarify whether persistent direction-changing geotropic positional nystagmus contains vertical and torsional components, and to quantify the asymmetry. We analyzed nystagmus in four positions (healthy-ear-down, affected-ear-down, supine, nose-down) using three-dimensional video-oculography. Subjects were 18 patients with persistent direction-changing geotropic positional nystagmus, 16 females and 2 males, with a mean age of 55 years. Nystagmus was recorded using an infrared camera and the findings were converted to digital data. Using ImageJ, we performed three-dimensional video-oculography and measured maximum slow-phase velocity (MSV) of three components. Positional nystagmus was not purely horizontal. Eight (44%) patients revealed a vertical component (upward) and 15 (83%) patients had a torsional component in the healthy-ear-down position. Seven (39%) patients revealed a vertical component (downward) and 10 (56%) patients showed a torsional component in the nose-down position. The mean value of MSV of the horizontal component in the supine position was 9.3°/s and that in the nose-down position was 15.7°/s. The latter was significantly greater than the former (p < 0.05). Eye movements in the supine position and the nose-down position were not mirror images. These results suggest that vertical and torsional components occur from the horizontal semicircular canal, and that horizontal canal ocular reflex is influenced by input from the otolithic organs.

Vestibulo-ocular function in anxiety disorders

Previous studies of vestibulo-ocular function in patients with anxiety disorders have suggested a higher prevalence of peripheral vestibular dysfunction compared to control populations, especially in panic disorder with agoraphobia. Also, our recent companion studies have indicated abnormalities in postural control in patients with anxiety disorders who report a high degree of space and motion discomfort. The aim of the present study was to assess the VOR, including the semicircular canal-ocular reflex, the otolith-ocular reflex, and semicircular canal-otolith interaction, in a well-defined group of patients with anxiety disorders. The study included 72 patients with anxiety disorders (age 30.6 +/- 10.6 yrs; 60 (83.3% F) and 29 psychiatrically normal controls (age 35.0 +/minus; 11.6 yrs; 24 (82.8% F). 25 patients had panic disorder; 47 patients had non-panic anxiety. Patients were further categorized based on the presence (45 of 72) or absence (27 of 72) of height phobia and the presence (27 of 72) or absence (45 of 72) of excessive space and motion discomfort (SMD). Sinusoidal and constant velocity earth-vertical axis rotation (EVAR) was used to assess the semicircular canal-ocular reflex. Constant velocity off-vertical axis rotation (OVAR) was used to assess both the otolith-ocular reflex and static semicircular canal-otolith interaction. Sinusoidal OVAR was used to assess dynamic semicircular canal-otolith interaction. The eye movement response to rotation was measured using bitemporal electro-oculography. Results showed a significantly higher VOR gain and a significantly shorter VOR time constant in anxiety patients. The effect of anxiety on VOR gain was significantly greater in patients without SMD as compared to those with SMD. Anxiety patients without height phobia had a larger OVAR modulation. We postulate that in patients with anxiety, there is increased vestibular sensitivity and impaired velocity storage. Excessive SMD and height phobia seem to have a mitigating effect on abnormal vestibular sensitivity, possibly via a down-weighting of central vestibular pathways.

Vestibular function in migraine-related dizziness: A pilot study

Migraine-related dizziness (MRD) refers to a disorder in which vestibular symptoms are an integral part of migraine symptomatology. The purpose of this study was to better define the pathophysiology of MRD, which is incompletely understood and to generate hypotheses regarding MRD by assessing the semicircular canal-ocular reflex, the otolith-ocular reflex, visual-vestibular interaction, vestibulo-spinal function, and visually induced postural sway. Subjects included five subjects with MRD, five subjects with migraine without dizziness, and five headache-free controls. Subjects with migraine were tested interictally. Results indicated that the mean gain of the semicircular canal-ocular reflex during both sinusoidal and constant velocity rotation was reduced in subjects with MRD. No changes were noted in the dynamics of the semicircular canal-ocular reflex. The otolith-ocular reflex, assessed with constant velocity OVAR, indicated that subjects with MRD showed a larger modulation component. No group differences were found in the bias component during constant velocity OVAR, nor in semicircular canal-otolith interaction or visual-vestibular interaction. Computerized dynamic posturography indicated that subjects with MRD demonstrated a surface-dependent pattern. Postural sway during optic flow indicated that subjects with MRD swayed more than the other subjects. We hypothesize that competing processes of serotonergic excitation and inhibition alter central vestibular pathways differently for semicircular canal vs. otolithic responses and for vestibulo-ocular vs. vestibulo-spinal pathways.

Otolith-ocular responses in patients with surgically confirmed unilateral peripheral vestibular loss

The chronic effects of unilateral peripheral vestibular loss (UPVL) are influenced by vestibular compensation. This study documents the balance-related symptoms and quantitative vestibular laboratory testing of 20 patients with surgically confirmed UPVL. Included are measures of the semicircular canal-ocular reflex, the otolith-ocular reflex, and both static and dynamic semicircular canal-otolith-interaction. This study differs from previous studies of patients with UPVL in that a large number of patients with surgically confirmed lesions were tested with three types of off-vertical axis rotation, several of the patients had anatomically preserved superior vestibular nerves, and self-perceived level of disability related to dizziness and imbalance were available. Results confirmed previously reported changes in the vestibulo-ocular reflex of patients with UPVL. Also, there was no apparent effect of anatomically preserving the superior vestibular nerve during surgical resection of vestibular schwannomas based on either subjective or objective measures of vestibular dysfunction. Further, there were no apparent correlations between subjective measures of dizziness and imbalance and objective measures of vestibulo-ocular function. These results have clinical implications for the management of patients with unilateral vestibular loss and provide insights into the process of vestibular compensation especially with respect to the otolith-ocular reflex.

Patterns of canal and otolith afferent input convergence in frog second-order vestibular neurons.

Second-order vestibular neurons (2 degrees VN) were identified in the isolated frog brain by the presence of monosynaptic excitatory postsynaptic potentials (EPSPs) after separate electrical stimulation of individual vestibular nerve branches. Combinations of one macular and the three semicircular canal nerve branches or combinations of two macular nerve branches were stimulated separately in different sets of experiments. Monosynaptic EPSPs evoked from the utricle or from the lagena converged with monosynaptic EPSPs from one of the three semicircular canal organs in ~30% of 2 degrees VN. Utricular afferent signals converged predominantly with horizontal canal afferent signals (74%), and lagenar afferent signals converged with anterior vertical (63%) or posterior vertical (37%) but not with horizontal canal afferent signals. This convergence pattern correlates with the coactivation of particular combinations of canal and otolith organs during natural head movements. A convergence of afferent saccular and canal signals was restricted to very few 2 degrees VN (3%). In contrast to the considerable number of 2 degrees VN that received an afferent input from the utricle or the lagena as well as from one of the three canal nerves (~30%), smaller numbers of 2 degrees VN (14% of each type of 2 degrees otolith or 2 degrees canal neuron) received an afferent input from only one particular otolith organ or from only one particular semicircular canal organ. Even fewer 2 degrees VN received an afferent input from more than one semicircular canal or from more than one otolith nerve (~7% each). Among 2 degrees VN with afferent inputs from more than one otolith nerve, an afferent saccular nerve input was particularly rare (4-5%). The restricted convergence of afferent saccular inputs with other afferent otolith or canal inputs as well as the termination pattern of saccular afferent fibers are compatible with a substrate vibration sensitivity of this otolith organ in frog. The ascending and/or descending projections of identified 2 degrees VN were determined by the presence of antidromic spikes. 2 degrees VN mediating afferent utricular and/or semicircular canal nerve signals had ascending and/or descending axons. 2 degrees VN mediating afferent lagenar or saccular nerve signals had descending but no ascending axons. The latter result is consistent with the absence of short-latency macular signals on extraocular motoneurons during vertical linear acceleration. Comparison of data from frog and cat demonstrated the presence of a similar organization pattern of maculo- and canal-ocular reflexes in both species.

Effect of aging on the otolith-ocular reflex

We assessed the influence of age on the otolith-ocular reflex and semicircular canal-otolith interaction. Healthy young (n=30) and healthy older (n=60) subjects were rotated about an earth vertical axis, and about a 30 degree off-vertical axis. Eye movements during and following rotation were recorded using electro-oculography. Results indicated that there were statistically significant changes in the otolith-ocular reflex and semicircular canal-otolith interaction as a function of age. The modulation component during off-vertical axis rotation (OVAR) was greater in the older group compard to the young adults, whereas the bias component was smaller with advanced age. The foreshortening of the vestibulo-ocular reflex time constant induced by post-rotatory head tilt following cessation of rotation was less prominent in the older group. There were no consistent changes in the semicircular canal-ocular reflex. Overall, response parameters showed more variability in the older subjects. We conclude that age related changes in the otolith-ocular reflex and semicircular canal-otolith interaction are a result primarily of a degradation of central vestibular processing of otolith signals rather than a decline of peripheral vestibular function.

Sacculo-ocular reflex connectivity in cats.

The otolith system contributes to the vestibulo-ocular reflexes (VOR) when the head moves linearly in the horizontal plane or tilts relative to gravity. The saccules are thought to detect predominantly accelerations along the gravity vector. Otolith-induced vertical eye movements following vertical linear accelerations are attributed to the saccules. However, information on the neural circuits of the sacculo-ocular system is limited, and the effects of saccular inputs on extraocular motoneurons remain unclear. In the present study, synaptic responses to saccular-nerve stimulation were recorded intracellularly from identified motoneurons of all twelve extraocular muscles. Experiments were successfully performed in eleven cats. Individual motoneurons of the twelve extraocular muscles--the bilateral superior recti (SR), inferior recti (IR), superior obliques (SO), inferior obliques (IO), lateral recti (LR), and medial recti (MR) were identified antidromically following bipolar stimulation of their respective nerves. The saccular nerve was selectively stimulated by a pair of tungsten electrodes after removing the utricular nerve and the ampullary nerves of the semicircular canals. Stimulus intensities were determined from the stimulus-response curves of vestibular N1 field potentials in order to avoid current spread. Intracellular recordings were performed from 129 extraocular motoneurons. The majority of the neurons showed no response to saccular-nerve stimulation. In 17 (30%) of 56 extraocular motoneurons related to vertical eye movements (bilateral SR and IR), depolarizing and/or hyperpolarizing postsynaptic potentials (PSPs) were observed in response to saccular-nerve stimulation. The latencies of PSPs ranged from 2.3 to 8.9 ms, indicating that the extraocular motoneurons received neither monosynaptic nor disynaptic inputs from saccular afferents. The majority of the latencies of the depolarization, including depolarization-hyperpolarization, were in the range of 2.3-3.3 ms. Latencies of hyperpolarizations were typically longer than those of depolarizations. Only one contralateral SO motoneuron of 43 recorded oblique extraocular motoneurons (bilateral SO and IO) showed a depolarization-hyperpolarization in response to saccular-nerve stimulation at a latency of 2.5 ms. None of 30 recorded horizontal extraocular motoneurons (bilateral LR and MR) responded to stimulation of the saccular nerve. The neural linkage in the sacculo-ocular system is relatively weak in comparison to the utriculo-ocular and sacculo-collic systems, suggesting that the role of the sacculo-ocular system in stabilizing eye position may be reduced when compared with utriculo-ocular and semi-circular canal-ocular reflexes.

Yaw Sensory Rearrangement Alters Pitch Vestibulo-Ocular Reflex Responses

Ten male subjects underwent two types of adaptation paradigm designed either to enhance or to attenuate the gain of the canal-ocular reflex (COR), before undergoing otolith-ocular reflex (OOR) testing with constant velocity, earth horizontal axis and pitch rotation. The adaptation paradigm paired a 0.2 Hz sinusoidal rotation about an earth vertical axis with a 0.2 Hz optokinetic stimulus that was deliberately mismatched in peak velocity or phase and was designed to produce short-term changes in the COR. Preadaptation and postadaptation OOR tests occurred at a constant velocity of 60%sec in the dark and produced a modulation component of the slow phase velocity with a frequency of 0.16 Hz due to otolithic stimulation by the sinusoidally changing gravity vector. Of the seven subjects who showed enhancement of the COR gain, six also showed enhancement of the OOR modulation component. Of the seven subjects who showed attenuation of the COR gain, five also showed attenuation of the OOR modulation component. The probability that these two cross-axis adaptation effects would occur by chance is less than 0.02. This suggests that visual-vestibular conditioning of the yaw axis COR also induced changes in the pitch axis OOR. We thus postulate that the central nervous system pathways that process horizontal canal yaw stimuli have elements in common with those processing otolithic stimuli about the pitch axis.

Deviation of the Subjective Vertical in Long-standing Unilateral Vestibular Loss

We evaluated changes in the subjectively perceived gravitational vertical as an index of imbalance in the function of the right and left otolith organs. In addition to normal subjects (n =25), we measured patients with a longstanding (mean 4.5 year ± 3.2 SD; range 0.5-11.5 years) unilateral vestibular loss after surgery for acoustic neuroma (n = 32), patients with partial unilateral vestibular loss (n = 7) and patients with bilateral vestibular hyporeflexia (n = 8). Normal subjects could accurately align a vertical luminous bar to the gravitational vertical in an otherwise completely dark room (mean setting –0.14° ± 1.11 SD). Patients with leftsided (complete; n = 13) or rightsided (complete; n = 19 and partial; n = 7) unilateral vestibular loss made mean angular settings at 2.55° ± 1.57 (SD) leftward and 2.22° (±1.96 SD) rightward, respectively. These means differed highly significantly from the normal mean (p < 0.00001). In the time interval investigated (0.5-11.5 years) the magnitude of the tilt angle showed no correlation with the time elapsed since the operation. The mean setting by patients with clinically bilateral vestibular loss (– 1.17° ± 1.96 SD; n = 8) did not significantly differ from the control group. The systematic tilts of the subjective vertical in patients with a unilateral vestibular impairment were correlated with their imbalance in canal-ocular reflexes, as reflected by drift during head-oscillation at 2 Hz (r2 = 0.44) and asymmetries in VOR-gain for head-steps (r2 = 0.48-0.67). These correlations were largely determined by the signs of the asymmetries; correlation between the absolute values of the VOR gain asymmetries and subjective vertical angles proved to be virtually absent. We conclude that the setting of the subjective vertical is a very sensitive tool in detecting a left-right imbalance in otolith function, and that small but significant deviations towards the defective side may persist for many years (probably permanently) after unilateral lesions of the labyrinth or the vestibular nerve.

A Model That Relates Canal-Ocular to Otolith-Ocular Responses

Rotation about the vertical stimulates primarily the horizontal semicircular canals and produces compensatory horizontal nystagmus whose slow component velocity response during constant velocity can be approximated as having a simple exponential decay time constant, Tvor. Constant velocity yaw rotation about a horizontal axis stimulates both the horizontal canals and the otolith organs, producing two additional nystagmus components thought to be of otolithic origin: a steady component called bias and a periodic component known as modulation. We tested a group of 7 human subjects using rotation about each of these axes. We found a strong, negative correlation (r = 0.956) between these individuals' dominant time constants, Tvor, and the magnitude of their modulation components. Canal and otolith signals originate from different parts of the same endorgan and travel separately to the vestibular nucleus. The reflexive eye movements in response to these inputs are thought to be the result of additional processing by the central vestibular and ocular motor systems. Thus, the source of these strongly correlated otolith-ocular and canal-ocular reflex components could logically be due to common factors affecting peripheral transduction or to a subsequent common central processing step. Using anatomical measurements of human vestibular end organs, biophysical endorgan models, and models of central vestibular processes, we examined the alternatives to determine which was the most likely. Correlations were not found between Tvor and the anatomical data or between the Tvor and the biophysical model elements. However, modifications of the velocity storage integrator of Raphan to incorporate either a highpass filter (HPF) or a lowpass filter input for otolith modulation signals allowed for the desired strong negative correlation between Tvor and modulation. We argue in favor of the HPF configuration because it better explains the tendency for the modulation component to increase in amplitude as the modulation frequency is increased. In the above mentioned representation of the central processing of vestibular afferent inputs, we conclude that the modulation component input to the vestibuloocular reflex is within the indirect pathway, but it is "down stream" from the velocity storage integrator.

半規管‐眼反射における適応現象が耳石‐眼反射に及ぼす影響

It has been demonstrated that chronic exposure to certain visual environments, for example those generated by magnifying lenses and reversing prisms, alter the gain and phase of the vestibulo-ocular reflex (VOR) in the dark in numerous species. Most experiments concerning such modification of the VOR have used semicircular canal stimulation. So far, there has been no investigation of the effect of adaptive plasticity of the semicircular canal-ocular reflex (SCOR) to an off-vertical axis rotation (OVAR), a dynamic otolith stimulation. The purposes of this study are to study physiologic alteration of the VOR using otolithic stimulation and to determine whether the bias and modulation component during OVAR change following adaptation of the SCOR. In this study, using OVAR, we tested the hypothesis that the otolith-ocular reflex (OOR) is altered by changes in the SCOR. Our study showed no convincing evidence for a transfer of SCOR adaptation to OOR using OVAR. There are two possibilities that could explain why we are unable to obtain this evidence, either there is no transfer to OOR, or there is an undiscovered technical problem. Future studies of transfer of adaptation will decrease OVAR exposure and assess OOR variability.

Off-vertical axis rotational responses in patients with unilateral peripheral vestibular lesions.

Off-vertical axis rotation (OVAR) stimulates the otolith organs in a manner that is suitable for assessment of the otolith-ocular reflex. To further assess the potential clinical usefulness of OVAR, the eye movement responses of seven patients with surgically confirmed unilateral peripheral vestibular lesions were compared with the eye movement responses of a group of age-matched, healthy, asymptomatic control subjects. Patients and controls were tested with constant velocity rotations that followed a brief period of angular acceleration (velocity trapezoid) using either earth-vertical axis (EVA) rotation or OVAR. Both EVA and OVAR sinusoidal velocity profiles were also performed. Results indicated that each patient had 1) an asymmetric OVAR response, ie, a bias component whose direction was opposite normal when rotating toward the lesioned ear, and 2) a normal modulation component. Population data suggested that patients had 1) a more rapid decay of response than normal subjects during OVAR velocity trapezoids, 2) an increased phase lead as compared to normal subjects during sinusoidal OVAR, and 3) like normal subjects, a less rapid decay of response during OVAR velocity trapezoids than during EVA rotational velocity trapezoids. Taken together, these findings suggest that patients with unilateral peripheral vestibular deficits have abnormal otolith-ocular and semicircular canal-ocular reflexes but that a single labyrinth appears to provide an otolithic signal sufficient for qualitatively normal semicircular canal-otolith interaction.

Preservation of the electrical-evoked vestibuloocular reflex and otolith-ocular reflex in two patients with markedly impaired canal-ocular reflexes.

Preservation of the electrical-evoked vestibuloocular reflex and otolith-ocular reflex in two patients with markedly impaired canal-ocular reflexes.

Plasticity of the Human Otolith-ocular Reflex

The eye movement response to earth vertical axis rotation in the dark, a semicircular canal stimulus, can be altered by prior exposure to combined visual-vestibular stimuli. Such plasticity of the vestibulo-ocular reflex has not been described for earth horizontal axis rotation, a dynamic otolith stimulus. Twenty normal human subjects underwent one of two types of adaptation paradigms designed either to attenuate or enhance the gain of the semicircular canal-ocular reflex prior to undergoing otolith-ocular reflex testing with horizontal axis rotation. The adaptation paradigm paired a 0.2 Hz sinusoidal rotation about a vertical axis with a 0.2 Hz optokinetic stripe pattern that was deliberately mismatched in peak velocity. Pre- and post-adaptation horizontal axis rotations were at 60 degrees/s in the dark and produced a modulation in the slow component velocity of nystagmus having a frequency of 0.17 Hz due to putative stimulation of the otolith organs. Results showed that the magnitude of this modulation component response was altered in a manner similar to the alteration in semicircular canal-ocular responses. These results suggest that physiologic alteration of the vestibulo-ocular reflex using deliberately mismatched visual and semicircular canal stimuli induces changes in both canal-ocular and otolith-ocular responses. We postulate, therefore, that central nervous system pathways responsible for controlling the gains of canal-ocular and otolith-ocular reflexes are shared.

Spatial Organization of the Maculo-Ocular Reflex of the Rat: Responses During Off-Vertical Axis Rotation.

Pigmented, head restrained rats were rotated on a turntable about a tilted axis (off-vertical axis rotation; OVAR) in darkness. Evoked eye movements in the horizontal, vertical and torsional planes were recorded simultaneously with a dual search coil in a magnetic field, horizontal response components of both eyes were recorded with a coil on either eye. OVAR resulted in a persisting horizontal, unidirectional ocular nystagmus, compensatory in direction for the rotation of head in space. Superimposed upon this nystagmus were slower cyclic responses of the eye in the vertical and torsional movement planes, that were tightly phase locked with changing head positions in space: ocular depression/elevation with right ear up/down and ocular intorsion/extorsion with nose up/down. Simultaneous recordings of horizontal response components from both eyes revealed phase and gain differences between the horizontal movement components of both eyes, that resulted in a cyclic modulation of the vergence angle. Convergence of the lines of sight during nose up and divergence during nose down, adequate compensatory responses in light for changes in the viewing distance, were actually observed in darkness. Thus the utricular maculo-ocular reflex takes part of the visual consequences of a translational gaze shift into account. It reduces expected retinal disparities by appropriate and rapid vertical, torsional and vergence response components in the same way as canal-ocular reflexes 'compensate' for direction and velocity of expected retinal image slip during head rotation.

VESTIBULAR FUNCTION IN PERIODIC ALTERNATING NYSTAGMUS

The vestibulo-ocular reflex of 4 patients with periodic alternating nystagmus (PAN) was studied in detail. Rotational testing was used to investigate the horizontal semicircular canal-ocular reflex, canal-otolith interaction, and the dynamic otolith-ocular reflex. Results indicated abnormal gain and phase of the horizontal semicircular canal-ocular reflex during sinusoidal rotation and a variable rate of decay of postrotatory responses. Each patient had abnormal canal-otolith interaction. An enlarged modulation component of the dynamic otolith-ocular response was observed in each patient. This study supports the idea that PAN is caused by an instability in the velocity storage element, a hypothetical neural circuit that perseverates the eye movement response to both vestibular and optokinetic stimulation. Further, we postulate that PAN may be caused by lesions of the cerebellar uvula and nodulus or their connections with the brainstem vestibular nuclei.

Earth horizontal axis rotational responses in patients with unilateral peripheral vestibular deficits.

We evaluated the vestibulo-ocular reflex (VOR) of five patients with surgically confirmed unilateral peripheral vestibular lesions. Testing used both earth vertical axis (EVA) and earth horizontal axis (EHA) yaw rotation. Results indicated that the patients had short VOR time constants, asymmetric responses to both EVA and EHA rotation, and normal EHA modulation components. These findings suggest that unilateral peripheral vestibular loss causes a shortened VOR time constant even with the addition of dynamic otolithic stimulation and causes an asymmetry in semicircular canal-ocular reflexes and one aspect of otolith-ocular reflexes.