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Over the past decade several research groups have taken a renewed interest in the special role of a type of small eye movement, called 'microsaccades', in various visual processes, such as the activation of neurons in the central nervous system, or the prevention of image fading. As the study of microsaccades and their relation to visual processes goes back at least half a century, it seems appropriate to review the more recent reports in light of the history of research on maintained oculomotor fixation, in general, and on microsaccades in particular. Our review shows that there is no compelling evidence to support the view that microsaccades (or, fixation saccades more generally) serve a necessary role in improving oculomotor control or in keeping the visual world visible. The role of the retinal transients produced by small saccades during fixation needs to be evaluated in the context of both the brisk image motions present during active visual tasks performed by freely moving people, as well as the role of selective attention in modulating the strength of signals throughout the visual field.

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

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 indivi

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.

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 scierai 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.

We studied binocular cyclorotatory (torsional) eye movements in response to gratings that oscillated sinusoidally in a frontal plane. The square-wave gratings viewed by the right and left eye were presented and controlled separately to induce cycloversion and cyclovergence by oscillation in phase and out of phase. Eye movements were recorded with scierai induction coils. Stimulus oscillation frequency ranged from 0.125 to 1 Hz and the wavelength of die gratings ranged from 0.92 to 25.75 deg of visual angle. Cycloversion and cyclovergence gain were, on average, comparable in magnitude and decreased with increasing oscillation frequency. There was no consistent effect of the wavelength on the magnitude of the responses. In general, responses were considerably higher to gratings that were oriented horizontally than to those oriented vertically. This anisotropy was present both in cycloversion and cyclovergence. It was enhanced in a larger sized stimulus and by presenting stationary, orthogonal contours (mimicking a “shear” movement), but it was not consistently influenced by wavelength. Cyclovergence showed a phase lag, which increased with oscillation frequency but which was independent of wavelength. In contrast, cycloversion showed a slight phase lead which was independent of both

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.

Recently it has been demonstrated that saccades become different in size in the two eyes if a subject is adapted to anisometropic spectacles, which provide visual images of different magnitude to the two eyes. These nonconjugate adaptations adequately meet the requirements of those spectacles and, once acquired, they persist (with some reduction) even during monocular viewing. We now demonstrate that such nonconjugate adaptations of saccades can be meridian-specific, if there is a pressure for such meridian-specificity. This pressure was provided by means of a cylindrical spectacle-lens. Adaptations along a vertical, horizontal or oblique meridian did not transfer to the orthogonal meridian. These results demonstrate a capability of saccadic adaptation to deal with calibration problems restricted not only to one eye, but even to one specific plane of muscular action. Our results also suggest that the meridian-specific adaptations of oblique saccades take place at a stage before the decomposition of motor commands into separate horizontal and vertical components. The meridian-specific nonconjugacies were also expressed in smooth-pursuit eye movements. Post-saccadic drift adapted only along the horizontal meridian.

REDE UITGESPROKEN BIJ DE AANVAARDING VAN HET AMBT VAN GEWOON LEKTOR IN DE FYSIOLOGIE AAN DE MEDISCHE FACULTEIT TE ROTTERDAM OP DONDERDAG 16 APRIL 1970