Zien en bewegen

REDE UITGESPROKEN BIJ DE AANVAARDING VAN HET AMBT VAN GEWOON LEKTOR IN DE FYSIOLOGIE AAN DE MEDISCHE FACULTEIT TE ROTTERDAM OP DONDERDAG 16 APRIL 197

Nonconjugate adaptation of human saccades to anisometropic spectacles: Meridian-specificity

Abstract 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

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

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 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

Eye torsion elicited by oscillating gratings: Effects of orientation, wavelength and stationary contours

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 bot

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

Simultaneous recordings of ocular microtremor and microsaccades with a piezoelectric sensor and a video-oculography system

Our eyes are in continuous motion. Even when we attempt to fix our gaze, we produce so called “fixational eye movements”, which include microsaccades, drift, and ocular microtremor (OMT). Microsaccades, the largest and fastest type of fixational eye movement, shift the retinal image from several dozen to several hundred photoreceptors and have equivalent physical characteristics to saccades, only on a smaller scale (Martinez-Conde, Otero-Millan & Macknik, 2013). OMT occurs simultaneously with drift and is the smallest of the fixational eye movements (∼1 photoreceptor width, >0.5 arcmin), with dominant frequencies ranging from 70 Hz to 103 Hz (Martinez-Conde, Macknik & Hubel, 2004). Due to OMT’s small amplitude and high frequency, the most accurate and stringent way to record it is the piezoelectric transduction method. Thus, OMT studies are far rarer than those focusing on microsaccades or drift. Here we conducted simultaneous recordings of OMT and microsaccades with a piezoelectric device and a commercial infrared video tracking system. We set out to determine whether OMT could help to restore perceptually faded targets during attempted fixation, and we also wondered whether the piezoelectric sensor could affect the characteristics of microsaccades. Our results showed that microsaccades, but not OMT, counteracted perceptual fading. We moreover found that the piezoelectric sensor affected microsaccades in a complex way, and that the oculomotor system adjusted to the stress brought on by the sensor by adjusting the magnitudes of microsaccades

Target Displacements during Eye Blinks Trigger Automatic Recalibration of Gaze Direction

Eye blinks cause disruptions to visual input and are accompanied by rotations of the eyeball [1]. Like every motor action, these eye movements are subject to noise and introduce instabilities in gaze direction across blinks [2]. Accumulating errors across repeated blinks would be debilitating for visual performance. Here, we show that the oculomotor system constantly recalibrates gaze direction during blinks to counteract gaze instability. Observers were instructed to fixate a visual target while gaze direction was recorded and blinks were detected in real time. With every spontaneous blink-while eyelids were closed-the target was displaced laterally by 0.5° (or 1.0°). Most observers reported being unaware of displacements during blinks. After adapting for ∼35 blinks, gaze positions after blinks showed significant biases toward the new target position. Automatic eye movements accompanied each blink, and an aftereffect persisted for a few blinks after target displacements were eliminated. No adaptive gaze shift occurred when blinks were simulated with shutter glasses at random time points or actively triggered by observers, or when target displacements were masked by a distracting stimulus. Visual signals during blinks are suppressed by inhibitory mechanisms [3-6], so that small changes across blinks are generally not noticed [7, 8]. Additionally, target displacements during blinks can trigger automatic gaze recalibration, similar to the well-known saccadic adaptation effect [9-11]. This novel mechanism might be specific to the maintenance of gaze direction across blinks or might depend on a more general oculomotor recalibration mechanism adapting gaze position during intrinsically generated disruptions to visual input

Natural Oculomotor Performance in Looking and Tapping Tasks

A unique apparatus recorded eye and head movements of subjects as they tapped or only looked at sequences of 2, 4 or 6 nearby, 3-D targets. Each sequence was repeated 10 times to allow an opportunity for learning. A stereotypical pattern of movements was established after 2-3 repetitions. Subjects almost always looked at each target just before tapping it. Looking-only was more difficult than tapping in that it took more time and, unlike tapping, usually did not benefit from practice. The number of targets in a sequence affected timeA^get in both tasks. Sequence length and practice effects show that memory was involved. The persistent strategy of looking before tapping and the subjects' inability to tap a well-leamed patten with eyes closed, show that visual cues were also important We conclude that motor plarming occurred first at the level of the task and then at the level of specific motor programs. The relative difficulty of the less natural, looking-only task, in which the eyes worked without a meaningful cognitive or motor purpose, suggests that efficient eye movement programming requires a natural task of the kind eye movements evolved to serve