Interocular timing differences in the horizontal components of human saccades.

The occurrence of systematic interocular differences in the time of initiation of saccades in various directions was investigated in normal human subjects (n=4). Saccades were recorded binocularly with scleral sensor coils on each of the eyes with a temporal resolution of 0.1 ms (sampling frequency 10000 Hz). Analysis was done in the velocity domain after digital differentiation. It was found that, in the initial phase of horizontal saccades, the nasalward moving eye lagged the temporalward moving eye consistently by slightly less than 1 ms. No such difference was found in vertical (upward or downward) saccades. In oblique saccades, the systematic initial lag of the nasalward moving eye was similarly present in the horizontal component of the saccade, but absent in the vertical component. It is postulated that this interocular timing difference in horizontal saccades is due to the additional synaptic delay caused by the abducens internuclear neuron in the pathway to the medial rectus muscle of the eye.

Saccadic binocular coordination in alternating exotropia.

We studied the coordination of binocular eye movements in human subjects with alternating exotropia (divergent strabismus). Binocular saccades were recorded in six subjects during binocular and monocular viewing. Subjects were instructed to make saccades between two continuously lit targets (LED's) presented in an isovergence array (with the straight-ahead target 130 cm from the eyes) in a dimly lit room. For saccades up to 20 degrees amplitude, there were no large differences in the dynamics of the saccades between control and exotropic subjects. However, for larger amplitudes subjects frequently alternated the eye of fixation during saccades. That is, subjects fixated the left target with the left eye and the right target with the right eye. The alternation in eye fixation at the end of the saccade was taken into account in the programming of the saccades. The amplitudes of the alternating saccades were approximately equal to the target amplitude minus the strabismus angle. We conclude that for those saccades where alternation occurs, there is not only a change in the eye of fixation, but also a change in the target representation provided by either eye. Thus, in this group of strabismic patients, saccades may be programmed in a retina-centered coordinate system, if we assume that for making a saccade to a new target in the contralateral visual field its representation on the temporal retinal field of the currently fixating eye is suppressed and the retinotopic target information is derived from the non-fixating eye. In executing the saccade, the non-fixating eye automatically becomes the fixating eye.

Early components of the human vestibulo-ocular response to head rotation: latency and gain.

To characterize vestibulo-ocular reflex (VOR) properties in the time window in which contributions by other systems are minimal, eye movements during the first 50-100 ms after the start of transient angular head accelerations ( approximately 1000 degrees /s(2)) imposed by a torque helmet were analyzed in normal human subjects. Orientations of the head and both eyes were recorded with magnetic search coils (resolution, approximately 1 min arc; 1000 samples/s). Typically, the first response to a head perturbation was an anti-compensatory eye movement with zero latency, peak-velocity of several degrees per second, and peak excursion of several tenths of a degree. This was interpreted as a passive mechanical response to linear acceleration of the orbital tissues caused by eccentric rotation of the eye. The response was modeled as a damped oscillation (approximately 13 Hz) of the orbital contents, approaching a constant eye deviation for a sustained linear acceleration. The subsequent compensatory eye movements showed (like the head movements) a linear increase in velocity, which allowed estimates of latency and gain with linear regressions. After appropriate accounting for the preceding passive eye movements, average VOR latency (for pooled eyes, directions, and subjects) was calculated as 8.6 ms. Paired comparisons between the two eyes revealed that the latency for the eye contralateral to the direction of head rotation was, on average, 1.3 ms shorter than for the ipsilateral eye. This highly significant average inter-ocular difference was attributed to the additional internuclear abducens neuron in the pathway to the ipsilateral eye. Average acceleration gain (ratio between slopes of eye and head velocities) over the first 40-50 ms was approximately 1.1. Instantaneous velocity gain, calculated as Veye(t)/Vhead(t-latency), showed a gradual build-up converging toward unity (often after a slight overshoot). Instantaneous acceleration gain also converged toward unity but showed a much steeper build-up and larger oscillations. This behavior of acceleration and velocity gain could be accounted for by modeling the eye movements as the sum of the passive response to the linear acceleration and the active rotational VOR. Due to the latency and the anticompensatory component, gaze stabilization was never complete. The influence of visual targets was limited. The initial VOR was identical with a distant target (continuously visible or interrupted) and in complete darkness. A near visual target caused VOR gain to rise to a higher level, but the time after which the difference between far and near targets emerged varied between individuals.

Gaze-shift dynamics in subjects with and without symptoms of convergence insufficiency: influence of monocular preference and the effect of training.

We studied gaze-shift dynamics during several gaze-shift tasks and during reading, in five subjects with convergence insufficiency (C.I., a diminished ability to converge), and in ten subjects without C.I. Furthermore, we studied the effect of vergence training in order to verify previous claims that orthoptic exercises can improve vergence performance. We recorded binocular eye movements with the scleral coil technique. Subjects switched fixation between nearby and distant light emitting diodes (LEDs) arranged in isovergence arrays (distances 35 and 130 cm) in a dimly lit room. In both the C.I. and non-C.I. group, two classes of subjects occurred: vergence responders and saccadic responders. During pure vergence tasks, saccadic responders made saccades with no or little vergence; vergence responders made vergence movements with no or small saccadic components. In saccadic responders, fixation of nearby targets was monocular. Subjects with a preferred eye, according to our determination, used the preferred eye. The five C.I. subjects showed idiosyncratic responses with insufficient vergence during most trials. They all had a tendency to alternate fixation between the left and right eye. Vergence-version tasks always elicited larger vergence components than pure vergence tasks. During a reading task, vergence angles were more accurate than during gaze-shifts between LEDs. After the pre-training sessions, nine subjects (one of which had C.I.) practised a pure vergence task three times a day for at least 2 weeks. Vergence amplitudes of four of these subjects were larger after training. We conclude that vergence training can change oculomotor performance. Although C.I. is often associated with abnormal vergence dynamics, there are no typical C.I. vergence dynamics. Unstable monocular preferences may play a role in the aetiology of C.I.

Dynamics of horizontal vergence movements: interaction with horizontal and vertical saccades and relation with monocular preferences.

We studied the dynamics of pure vergence shifts and vergence shifts combined with vertical and horizontal saccades. It is known from earlier studies that horizontal saccades accelerate horizontal vergence. We wanted to obtain a more complete picture of the interactions between version and vergence. Therefore we studied pure version (horizontal and vertical), pure vergence (divergence and convergence) and combinations of both in five adult subjects with normal binocular vision and little phoria (< 5 degrees). The visual targets were LED's in isovergence arrays presented at two distances (35 and 130 cm) in a dimly lit room. Two targets were continuously lit during each trial and gaze-shifts were paced by a metronome. The two subjects with a strong monocular preference made vergence eye movements together with small horizontal saccades during pure vergence tasks. The other subjects, who did not have a strong monocular preference, made pure vergence movements (without saccades). These findings, suggest that monocular preferences influence the oculomotor strategy during vergence tasks. Vergence was facilitated by both horizontal and vertical saccades but vergence peak-velocity during horizontal saccades was higher than during vertical saccades.

Gaze-shift dynamics in two kinds of sequential looking tasks.

Gaze-shift dynamics of unrestrained seated subjects were examined. The subjects participated in two tasks. In the first task, they tapped sequences of 3-D targets located on a table in front of them. In the second task, they only looked at similar sequences of targets. The purpose of the task (tapping vs only looking) affected the dynamics of gaze-shifts. Gaze and eye-in-head peak velocities were higher and gaze-shift durations were shorter during tapping than during looking-only. We conclude that task variables affect gaze-shift dynamics, altering characteristics of the so-called saccadic "main sequence".

Binocular saccadic eye movements in multiple sclerosis.

We attempted to increase the sensitivity for detection of abnormal binocular saccadic eye movements, particularly of the internuclear ophthalmoplegia (INO) type associated with multiple sclerosis (MS). Saccades of 10 and 20 degrees were binocularly recorded with scleral sensor coils in 10 normal control subjects and 26 patients with definite or probable MS, seven of whom had a clinically manifest INO in one or both directions. In the cases in which this was accompanied by a dissociated nystagmus of the abducting eye, our recordings showed that such secondary saccades were also expressed, in a strongly reduced form, by the adducting eye. The patients with manifest INO showed lower average peak velocities and peak accelerations, especially for adduction of the eye on the affected side, but the distribution of these parameters overlapped with the normal distribution. A much sharper distinction between normals and patients with INOs was found by considering the ratios between peak accelerations and velocities of saccade pairs (abducting eye/adducting eye). These ratios, which eliminate much intra- and inter-individual variability, had a narrow range in normals, and all values for INOs were outside this range. On this basis, the 19 patients without clinically manifest INO were easily separated into subgroups of 14 patients with completely normal interocular ratios and five patients with elevated peak velocity and acceleration ratios, identified as sub-clinical (uni- or bilateral) INOs. Measurements of vertical saccades and of interocular timing differences provided no useful criteria for disturbances of binocular coordination in MS. We conclude that in particular, the acceleration of the adducting eye is strongly reduced in patients with an INO, and that this reduction is best identified by interocular comparison between binocular pairs of saccades.

Trajectories of the human binocular fixation point during conjugate and non-conjugate gaze-shifts.

This paper describes the spatial trajectories of the binocular fixation point (the intersection point of the two lines of sight) during gaze-shifts within a horizontal plane of regard. Gaze was voluntarily shifted between pairs of real, continuously visible LED targets that were either iso-vergent at 5-25 deg convergence (conjugate version saccades) or differed in vergence angle (by 5-20 deg) as well as in direction (by 5-60 deg; combined version and vergence). Orientations of both eyes were recorded by phase detection in a homogeneous magnetic field with scleral sensor coils. "Conjugate" saccades showed an outward-looping, curved trajectory as a result of transient divergence, typically associated with horizontal saccades. These outward loops were disproportionately larger for far than for near targets, due to the non-linear relation between vergence and distance. Transient divergence increased moderately in magnitude and duration when basic vergence increased from 5 to 25 deg. As a result, transient saccadic disparities increased in angular magnitude as targets got close. Increasing tonic vergence did not, however, slow down conjugate saccades, in contrast to the previously described dynamic slowing effects of vergence on version during gaze-shift involving simultaneous vergence and version changes. Convergent and divergent non-conjugate gaze-shifts each had characteristic trajectories; outward loops were much reduced in convergent and virtually absent in divergent movements. The saccadic component of non-conjugate gaze-shifts was preceded by a pre-saccadic vergence component in the direction of the imminent gaze-shift; its magnitude increased systematically with the increase in vergence demand and with the decrease in version demand. For both pre-saccadic convergence and divergence, this pre-saccadic part of the trajectory tended to follow an iso-direction line through the target of origin; directional change did not start until the saccade began. This suggests that for targets that differ in direction as well as distance, control of the vergence and version components of the gaze-shift can be dissociated to some degree. This seems to argue against models of binocular oculomotor control which assume that each eye responds primarily to its own target, and suggests rather that target vergence and target direction may be processed and responded to separately by ocular vergence and version, with a strong interaction between the two oculomotor activities whenever they occur at the same time.

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 degree +/- 1.11 SD). Patients with left-sided (complete; n = 13) or right-sided (complete; n = 19 and partial; n = 7) unilateral vestibular loss made mean angular settings at 2.55 degrees +/- 1.57 (SD) leftward and 2.22 degrees (+/-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 degrees +/- 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.

Gain and delay of human vestibulo-ocular reflexes to oscillation and steps of the head by a reactive torque helmet. I. Normal subjects.

Vestibulo-ocular reflexes (VOR) were evaluated with a reactive torque helmet that imposed high-frequency oscillation (2-20 Hz) or step displacements of the head in the horizontal plane. The present paper describes the experimental and analytical techniques and the results for normal subjects, which will serve as a baseline for the evaluation of vestibular pathology. For comparison, manually controlled head steps were also applied, as described in the literature. Eye and head movements were recorded with magnetic search coils. Non-vestibular effects were avoided by the use of high stimulus frequencies and the analysis of short time-windows (< 100 msec) after steps. Helmet-imposed steps caused a virtually uniform head acceleration (average magnitude 770 degrees/sec) in the first 90 msec. This resulted in a linear relation between eye and head velocities; the gain and delay of the VOR could be calculated independently from the slope and offset of this relation. Such estimates appear more reliable than those obtained with conventional techniques. Normal subjects had a VOR gain of about 0.9 and a delay of about 5 msec. The results of sinusoidal head oscillation were in agreement with the results for steps. The responses to manually generated head steps agreed in general with those to helmet-induced steps, but because of the non-uniform acceleration they allowed a less exact analysis of function.

Gain and delay of human vestibulo-ocular reflexes to oscillation and steps of the head by a reactive torque helmet. II. Vestibular-deficient subjects.

Vestibulo-ocular reflexes (VOR) were evaluated with a reactive torque helmet that imposed high-frequency oscillation (2-20 Hz) or step displacements of the head in the horizontal plane. The present paper describes the results in patients with vestibular deficiencies (labyrinthine defective; LD); experimental and analytical techniques and results for normal subjects were described in Part 1 of this paper. The patient groups included: total unilateral LD (related to acoustic neuroma; n = 40); severe (clinically total) bilateral LD (n = 7); bilateral hyporeflexia (n = 14); unilateral hyporeflexia (n = 11); and patients with LD phenomena that had subsided (n = 3). Helmet-induced head steps provided the most specific information. Characteristically, gain was lowered in one direction or both directions after unilateral or bilateral vestibular lesions, respectively; in general, the magnitude of the gain reduction correlated well with the degree of complaints and disability. Surprisingly, delay was systematically prolonged (up to several tens of milliseconds) in all groups of subjects with manifest vestibular pathology. These results suggest that the determination of delay, in addition to gain of the VOR, is feasible and important in the evaluation of vestibular function. The results of head oscillation generally supported the results for steps, but were somewhat less specific. The responses to manually generated head steps roughly agreed with those to helmet-induced steps, but because of the non-uniform acceleration they allowed a less exact analysis of VOR function.

Modulation of the human vestibuloocular reflex during saccades: probing by high-frequency oscillation and torque pulses of the head.

1. We probed the gain and phase of the vestibuloocular reflex (VOR) during the execution of voluntary gaze saccades, with continuous oscillation or acceleration pulses, applied through a torque helmet. 2. Small-amplitude (< 1 degree), high-frequency (10-14 Hz) head oscillations in the horizontal or vertical plane were superimposed on ongoing horizontal gaze saccades (40-100 degrees). Torque pulses to the head ("with" or "against" gaze) were superimposed on 40 degrees horizontal saccades. Eye and head movements were precisely measured with sensor coils in magnetic fields. 3. Techniques were developed to separate the oscillatory (horizontal or vertical) component from the gaze shift and obtain VOR gain and phase with Fourier techniques from the relation between eye-in-head and head oscillations. These involved either subtraction of exactly matching saccades with and without oscillation (drawback: low yield) or time shifting of successive trials to synchronize the oscillations (drawback: slight time blurring of saccades). 4. The results of these matching and synchronization methods were essentially identical and consistent. Presaccadic gain values of the horizontal VOR (typically about unity) were reduced by, on average, approximately 20 and 50% during horizontal saccades of 40 and 100 degrees, respectively. These percentages may be truncated because of methodological limitations, but even after taking these into account (on the basis of simulation experiments with 2 different, theoretical profiles of suppression) our results do not support a complete saccadic VOR suppression for any substantial fraction of saccadic duration. Qualitatively similar changes were found when the vertical VOR was probed during 100 degrees horizontal saccades. 5. Concomitantly with the reductions in gain, VOR phase was advanced by approximately 20 degrees during the saccade. 6. In the wake of gaze saccades, VOR gain was consistently elevated (to approximately 1.0) above the presaccadic level (approximately 0.9). We submit that this mechanism ensures stable fixation of the newly acquired target at a time when the head is still moving substantially. 7. Although the responses to head torque pulses showed idiosyncratic asymmetries, analysis of the differences in eye and head movements for pulses with and against consistently showed a sharp fall of VOR gain at saccadic onset, following an approximately exponential course with a time constant of approximately 50 ms. This decay may be assumed to reflect VOR gain for a period of approximately 50 ms, after which secondary gaze control mechanisms become dominant. 8. The time course of the gain decay and phase shift of the VOR suggest that suppression of the "integrative (position) loop" of the VOR circuit was more complete than suppression of the direct, "velocity" pathway.

Conjugacy of eyelid movements in vertical eye saccades.

PURPOSE: To examine the conjugacy of lid saccades in normal subjects. METHODS: Saccades of both upper eyelids were recorded simultaneously by means of small, lightweight, magnetic search coils, fixed on each lid. Subjects then made vertical eye saccades between fixed targets. The associated eye saccades were recorded simultaneously by means of magnetic search coils. The authors further examined whether the position of the lid coils affected the recordings. RESULTS: Lid saccades were not as conjugate as their associated eye saccades. Nonconjugacy of lid saccades averaged approximately 8% of the target amplitude, irrespective of saccadic direction. By contrast, nonconjugacy of the associated eye saccades averaged approximately 1% of the target amplitude. Coil position significantly (P < 0.05) affected the recordings. CONCLUSIONS: Lid saccades are not as conjugate as their associated eye saccades. Coil position is critical for the recordings of lid saccades.

Voluntary binocular gaze-shifts in the plane of regard: dynamics of version and vergence.

We studied the dynamics of voluntary, horizontal, binocular gaze-shifts between pairs of continuously visible, real three-dimensional targets. Subjects were stabilized on a biteboard to allow full control of target angles, which were made to differ only in distance (pure vergence), only in direction (pure version; conjugate saccades) or in both distance and direction (disjunctive saccades). A wide range of changes in vergence (0-25 deg) and version (0-65 deg) was recorded to study the dynamics of disjunctive saccades, described until now for limited ranges, throughout the horizontal oculomotor range within manual working space, and to study the velocity-duration-amplitude relations ("main sequence") of disjunctive vs conjugate saccades. Pure vergence was almost never observed; divergence, especially, was always associated with saccades. Likewise, horizontal saccades were never strictly conjugate, they always contained a transient divergence-convergence sequence. The amplitude and velocity of these transient components varied systematically with saccadic size. In combined version-vergence movements, vergence was, in general, accelerated and shortened as a function of increasing version. This effect was fairly uniform for divergence, which appeared to increase in velocity by about as much as the transient peak divergent velocity of the version saccade. The intrasaccadic fraction of divergence increased from about 50% to close to 100% as a function of increasing version. For convergence, saccades up to about 20 deg were also accelerating; in this case it appeared as if the transient peak convergent velocity of the version saccade was added to the basic convergence velocity. For larger saccades this effect was partly counteracted by the penetration of an initial divergence associated with the saccade. This initial divergence delayed and slowed down convergence. The intrasaccadic fraction of convergence varied between about 40% and 70%. In disjunctive saccades the individual eyes did not follow the main-sequence parameters of conjugate saccades of comparable sizes, except for the eye that moved with the combination "abduction and divergence". For all other combinations of vergence and version, disjunctive saccades had lower peak velocities and longer durations than conjugate saccades. As a consequence, disjunctive version was also slower than conjugate version. Thus, while version accelerates vergence, vergence slows down version: in the generalized case of three-dimensional gaze-shifts, peak velocities and durations are in between those of the limiting cases of pure version and pure vergence. We conclude that, within manual working space, binocular gaze-shifts are effected by the highly integrated action of conjugate and disjunctive mechanisms, both of which are expressed preferentially in fast, saccadic movements.

The function of visual search and memory in sequential looking tasks.

UNLABELLED: Eye and head movements were recorded as unrestrained subjects tapped or only looked at nearby targets. Scanning patterns were the same in both tasks: subjects looked at each target before tapping it; visual search had similar speeds and gaze-shift accuracies. Looking, however, took longer and, unlike tapping, benefitted little from practice. Looking speeded up more than tapping when memory load was reduced: memory was more efficient during tapping. CONCLUSION: eye movements made when only looking are different from those made when tapping. Visual search functions as a separate process, incorporated into both tasks: it can be used to improve performance when memory load is heavy.

When push comes to shove: compensation for passive perturbation of the head during natural gaze shifts.

The effects of passive displacements to the head delivered by an abrupt push to the upper body were studied in human subjects during gaze shifts to nearby targets while the head was completely unrestrained. Accurate measurements of gaze were obtained via the Maryland Revolving Field Monitor, used to measure head and eye rotations unconfounded with translations, and by an acoustic ranging system, used to measure head translations. Compensation for head perturbations was quite good, with gaze errors much the same as gaze errors in the absence of the push. Compensation along one or both meridians was achieved by means of the vestibulo-ocular response in many of the gaze shifts. The results suggest an impressive ability to coordinate head and eye movements during natural gaze shifts, carried out by one or more different kinds of compensatory systems that the subject can access at will or according to task demands.

Conjugacy of saccades in deep amblyopia.

PURPOSE: To determine the degree of conjugacy of saccades in humans with one deeply amblyopic eye, to assess whether Hering's law requires good binocular visual acuity. METHODS: By means of magnetic search coils, horizontal saccades were recorded accurately in 11 subjects with unilateral, deep amblyopia. The recordings were compared to those made in 11 normal subjects. In addition, three subjects with amblyopia were submitted to artificial anisometropia, which called for nonconjugate adaptation of their saccades. RESULTS: The saccades of most subjects with amblyopia were nonconjugate. These nonconjugacies often were larger in one direction than in the opposite one. The magnitude of the nonconjugacies varied inconsistently with the size of the saccades. In addition, the amblyopic eye often showed conspicuous drift. The saccades adapted nonconjugately to the imposed anisometropia. CONCLUSIONS: In deep amblyopia, the two eyes often make saccades that are unequal in size. The amblyopic eye does not move aimlessly, roughly following the movements of the normal fellow eye. Instead, the saccades of the amblyopic eye appear to reflect a deliberate oculomotor strategy, bringing visual targets to presumed retinal points of preference.

Evaluation of the human vestibulo-ocular reflex at high frequencies with a helmet, driven by reactive torque.

A new technique was developed to test the VOR in humans. A tightly but comfortably fitting helmet was provided with an electronically controlled torque motor that rotated a mass around a vertical axis. Acceleration of the motor caused reactive torque of the helmet in the opposite direction which was transmitted to the (freely movable) head. Sinusoidal head oscillations in the frequency range 2-20 Hz (peak-to-peak amplitudes about 10 degrees and 0.1 degrees respectively) were easily achieved, as well as step displacements of the head with accelerations on the order of 1000 degrees/s2. Limitation of the maximum torque and lack of any rigid attachment of the head to a fixed structure made the technique safe and comfortable. Eye and head rotations were recorded, independently of head translations, with eye and head search coils in a homogeneous a.c. magnetic field. In normal subjects, gain was near unity at 2 Hz: above 8 Hz, gain increased progressively toward 1.1-1.3 at 20 Hz. Phase lag increased from a few degrees at 2 Hz to about 45 degrees at 20 Hz. Above 2 Hz, these results were unaffected by visual conditions; lower gains and increased phase lags were found in subjects with bilateral or unilateral vestibular loss. For step displacements, gain (measured in the first 100 ms) was near unity for normals, near zero after bilateral vestibular loss and strongly asymmetrical after unilateral vestibular loss. Thus, the technique seems highly effective in testing vestibular function with minimal contamination by other systems.

Eye torsion associated with disparity-induced vertical vergence in humans.

Recently, Enright described an unexpected association between disparity-induced vertical vergence and cycloversion (conjugate eye torsion) [Enright (1992) Vision Research, 415, 279]. The present experiments were performed to verify these findings and investigate the nature of this association. We presented subjects with a dichoptic image of concentric circles in which a step in vertical disparity of 1 deg was introduced. After 4 sec the disparity was eliminated. Eye movements were measured with scleral coils. We confirmed Enright's findings in that a left-over right vertical vergence was associated with levo-cycloversion (upper poles towards left shoulder) and vice versa. The size of the cycloversion and the vertical vergence were in the same range. In addition we found that part of the cycloversion response was in the form of a torsional nystagmus and that the relative contribution of the left and right eyes was independent of the horizontal gaze angle. These additional findings are in conflict with the hypothesis, offered by Enright, that the association is caused by a bilateral activity of the superior oblique muscles.

Instability of ocular torsion during fixation: cyclovergence is more stable than cycloversion.

We investigated spontaneous variation of binocular torsion. Variation was expressed as SD of torsional eye positions measured over periods up to 32 sec. Subjects viewed a single dot target for periods of 32 sec. In half of the trials a large random-dot background pattern was superimposed on the dot. The movements of both eyes were measured with scleral induction coils. Spontaneous torsional movements were largely conjugate: cyclovergence was much more stable than cycloversion. This difference was not due to roll head movements. Stability of cyclovergence was improved by the background pattern. Although overall stability (SD of position) of cycloversion was unaffected by a background, the background induced or enhanced a small-amplitude torsional nystagmus in 3 out of 4 subjects. We hypothesize that the difference in stability of cycloversion vs cyclovergence reflects the greater importance of torsional retinal correspondence, compared to absolute torsional position. In two subjects we found evidence for the existence of cyclophoria, manifested by systematic shifts in cyclovergence caused by the appearance and disappearance of the background.