PubMed HealthSearch

Biomedical subjects

H Collewijn

Publications and source records attributed to H Collewijn.

At least 37 records · Page 2Linked to original sources

Ocular vergence under natural conditions. I. Continuous changes of target distance along the median plane.

Horizontal binocular eye movements of four subjects were recorded with the scleral sensor coil--revolving magnetic field technique while they fixated a natural target, whose distance was varied in a normally illuminated room. The distance of the target relative to the head of the subject was changed in three ways: (a) the target was moved manually by the experimenter; (b) the target was moved manually by the subject; (c) the target remained stationary while the subject moved his upper torso towards and away from the target. The rate of change of target distance was varied systematically in four levels, ranging from 'slow' to 'very fast', corresponding to changes in target vergence from about 10 degrees s-1 to about 100 degrees s-1. The dynamics of ocular vergence with regard to delay and speed were, under all three conditions, considerably better than could be expected from the literature on ocular vergence induced by disparity and/or blur. When 'very fast' changes in the distance of the target were made, subjects achieved maximum vergence speeds of up to about 100 degrees s-1. Delays of these fast vergence responses were generally smaller than 125 ms. Negative delays, i.e. ocular vergence leading the change in target distance, were observed. The eyes led the target (i.e. predicted target motion) by about 90 ms on average, when the subject used his hand to move the target. Vergence tracking was almost perfect when changes in distance were produced by moving the upper torso. In this condition, the eye led the target by about 5 ms. In the 'slow' and 'medium' conditions (stimulus speeds about 10-40 degrees s-1) tracking was accurate to within 1-2 degrees, irrespective of the way in which the target was moved. In the 'fast' and 'very fast' conditions (stimulus speeds about 40-100 degrees s-1), the accuracy of vergence tracking was better for self-induced than for experimenter-induced target displacements, and accuracy was best during voluntary movements of the upper torso. In the last case, ocular vergence speed was within about 10% of the rate of change of the vergence angle formed by the eyes and the stationary target. The dynamics of convergent and divergent vergence responses varied considerably. These variations were idiosyncratic. They were consistent within, but not between, subjects. Ocular vergence associated with attempted fixation of an imagined target, changing distance in darkness, could only be made by two of the four subjects.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult

Ocular vergence under natural conditions. II. Gaze shifts between real targets differing in distance and direction.

Horizontal binocular eye movements of three subjects were recorded with the scleral sensor coil--revolving magnetic field technique during voluntary shifts of gaze between pairs of stationary, real, continuously visible targets. The target pairs were located either along the median plane (requiring symmetrical vergence), or on either side of the median plane (requiring asymmetrical vergence). Symmetrical vergence was primarily smooth, but it was often assisted by small, disjunctive saccades. Peak vergence speeds were very high; they increased from about 50 degrees s-1 for vergence changes of 5 degrees to between 150 and 200 degrees s-1 for vergence changes of 34 degrees. Differences between convergence and divergence were idiosyncratic. Asymmetrical vergence, requiring a vergence of 11 degrees combined with a version of 45 degrees, was largely saccadic. Unequal saccades mediated virtually all (95%) of the vergence required in the divergent direction, whereas 75% of the vergence required in the convergent direction was mediated by unequal saccades, with the remaining convergence mediated by smooth vergence, following completion of the saccades. Peak divergence speeds during these saccades were very high (180 degrees s-1 for a change of vergence of 11 degrees); much faster than the smooth, symmetrical vergence change of comparable size (14 degrees). Peak convergent saccadic speeds were about 20% lower. This difference in peak speed was caused by an initial, transient divergence, observed at the beginning of all horizontal saccades. The waveform of disjunctive saccades did not have the same shape as the waveform of conjugate saccades of similar size. The smaller saccade of the disjunctive pair was stretched out in time so as to have the same duration as its larger, companion saccade. These results permitted the conclusion that the subsystems controlling saccades and vergence are not independent. Vergence responses were relatively slow and incomplete with monocular viewing, which excluded disparity as a cue. Monocularly stimulated vergence decreased as a function of the increasing presbyopia of our three subjects. Subjects were able to generate some vergence in darkness towards previously seen and remembered targets. Such responses, however, were slow, irregular and evanescent. In conclusion, vergence shifts between targets, which provided all natural cues to distance, were fast and accurate; they appeared adequate to provide effective binocular vision under natural conditions. This result could not have been expected on the basis of previous observations, all of which had been made with severely reduced cues to depth.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult

Differences in accuracy of human saccades between stationary and jumping targets.

Saccades have traditionally been studied in response to suddenly changing visual stimuli, such as jumping targets. In every-day life, however, most targets are stationary. We studied saccades made in either target condition. Saccadic accuracy was two- to five-fold better with stationary targets than with jumping targets. In addition, both the number of secondary saccades and the total time required to foveate the target were decreased with stationary targets in comparison to jumping targets. We also found that with an illuminated background the total time required to foveate the target was shorter than when the background was dark.

Adult

The vestibulo-ocular reflex: an outdated concept?

Traditionally, the vestibulo-ocular reflex (VOR) is described as a distinct, phylogenetically old oculomotor subsystem, which serves to stabilize gaze direction. It is supposed to act as a stereotyped reflex with definite input-output relations, which can be measured by rotating a subject passively in darkness, and which are kept at an ideal level by adaptive, parametric adjustments. This paper argues that such a view is not realistic: (1) the VOR in darkness does not have an ideal, or even well defined, gain; (2) a fixed, automatic VOR is not appropriate in most behavioural situations, and would need continuous conditioning by other subsystems. As there is no compelling phylogenetic, physiological or anatomical evidence for an independent VOR subsystem, a more fruitful hypothesis may be that vestibular signals are just one of many inputs to a spatial localization process, which computes the relative position (and motion) between the subject and a target of his choice. The VOR in darkness may represent no more than a default operation, based on incomplete information, of this larger, multiple input gaze control system. Likewise, adaptation phenomena of the VOR in darkness may be merely an epiphenomenon of adaptation of gaze control with vision active.

Animals

Depression of the vestibulo-ocular and optokinetic responses by intrafloccular microinjection of GABA-A and GABA-B agonists in the rabbit.

The functional implication of the cerebellar flocculus in regulation of the VOR and OKR gain has mostly been studied by lesion experiments, and the hypotheses derived from these experiments are not always in line with one another. In the present study, a reversible method was used to inhibit floccular Purkinje cells. The GABA-A agonist muscimol or the GABA-B agonist baclofen were bilaterally injected into the flocculus of rabbits, and the effects of these injections on the gain of the VOR and OKR were studied. Both drugs induced a reduction by at least 50% of the gain of the VOR in light and darkness, and of the OKR. Although GABA-A and GABA-B receptors are known to have different cerebellar localizations, muscimol and baclofen injections resulted in quantitatively similar effects. It is suggested that these GABA-agonists cause either direct or indirect inhibition of floccular Purkinje cells, thus reducing modulation of the firing rate of these neurons by afferent mossy and climbing fibers. Because the flocular Purkinje cells act out of phase with the vestibular neurons which drive the oculomotor neurons, a reduced output of floccular Purkinje cells would result in a reduction of the VOR and OKR gain. These experiments provide strong evidence that the cerebellar flocculus has a positive influence on the basic VOR and OKR gain.

Animals

The vestibulo-ocular reflex: is it an independent subsystem?

Traditionally, the vestibulo-ocular reflex (VOR) is described as a distinct, phylogenetically old, oculomotor subsystem, which serves to stabilize gaze direction. It is supposed to act as a stereotyped reflex with definite input-output relations, which can be measured by rotating a subject passively in darkness, and which are kept at a nearly ideal level by adaptive parametric adjustments. This paper argues that such a view of the VOR may be not realistic: 1) the VOR in darkness (especially in humans) does not behave as a well-calibrated system; it has a low and variable gain which can be changed easily, even by purely mental assumptions; 2) a hard-wired VOR does not lead to appropriate eye movements in most natural situations, and would need continuous conditioning by other systems. As there is no compelling physiological or anatomical evidence for an independent VOR, it seems more fruitful to hypothesize that vestibular signals are just one of many inputs to a spatial localization process, which computes the relative motion between a subject and an object of his choice on the basis of all available, relevant information. Instead of a distinct subsystem, vestibulo-ocular responses in darkness may represent nothing more than the (somewhat arbitrary) default performance of this larger gaze-control system, functioning poorly in the absence of complete information.

Animals

Asymmetrical adaptation of human saccades to anisometropic spectacles.

We report the extent to which effective asymmetrical saccadic adaptation was achieved by a myopic subject, who was exposed to "long-term" adaptation as he wore anisometropic corrective spectacles for about 40 years and also the extent of "short-term" adaptation in this subject and two other subjects, who initially made conjugate saccades, when they wore newly fitted anisometropic spectacles for about 8 hr. Two-dimensional binocular eye positions were measured with an accurate and precise revolving magnetic field-sensor coil technique. We found that long-term adaptation of vertical saccades was virtually perfect (almost 100% of the asymmetry introduced by the spectacles was corrected). Long-term adaptation of horizontal saccades was less complete and increased with target separation from about 40% for saccadic amplitudes of 5 degrees to about 75% for amplitudes of 60 degrees. Short-term adaptation of vertical saccades was virtually complete (100%) in one newly fitted subject and only partially complete (40%) in the other two subjects. The persistence of the adaptive asymmetry of saccades during monocular viewing showed that adaptation derived from plasticity in the programming of saccades and not from modification of vergence responses. Without the anisometropic spectacles, 30 min of self-paced, one per second changes in binocular fixation between two targets, which required a version change of 45 degrees in combination with a vergence change of 11 degrees, did not induce any asymmetrical adaptation. This result shows that a specific repeated association of version and vergence eye movements was not sufficient to induce asymmetrical adaptation, leading us to suggest that the transient fixation disparities at saccade-offset might be the necessary stimulus for the asymmetrical saccadic adaptation we observed.

Adaptation, Physiological

Directional asymmetries of human optokinetic nystagmus.

Optokinetic nystagmus in the four principal directions was investigated on the occurrence of directional asymmetries in 7 normal human subjects. Instructions were aimed at obtaining a 'stare' type of OKN. The movement of both eyes was recorded simultaneously with a scleral sensor-coil method. Subjects viewed a full-field random dot pattern rotating at velocities of 9 to 57 deg/s binocularly, as well as monocularly with either eye. Gain was always less than 0.85 and decreased when the pattern velocity increased. Horizontal and vertical nystagmus differed in a number of respects. (1) We found no evidence for an overall asymmetry for rightward or leftward motion. However, human OKN showed a clear preference for upward stimulus motion. Mean gain was ca. 0.15 larger for upward than for downward motion. (2) The decrease of the gain of OKN as a function of increasing stimulus velocity was steeper for vertical than for the horizontal direction. (3) The eyes moved nearly perfectly yoked for vertical pattern movement, irrespective of the viewing conditions. In contrast, during horizontal OKN the gain of the eye tracking in the nasal direction was higher (by about 4%) than the gain of the other eye moving simultaneously in the temporal direction. This difference persisted irrespective of the viewing conditions and appears to be motor, not sensory in origin. In addition, for any direction of the pattern motion a statistically significant increase of the gain occurred when the pattern motion was seen binocularly instead of monocularly with either eye.

Functional Laterality

Oculomotor defects in patients with Huntington's disease and their offspring.

We recorded saccadic, pursuit and fixation eye movements in patients (n = 5) with moderately advanced Huntington's disease (HD), offspring of HD patients (n = 22) and control subjects (n = 15), using the scleral sensor coil technique. Saccadic slowing was seen in all patients, no controls and (marginally) in a few at-risk subjects. Fixational stability was lower in patients than in the other groups; a structured background enhanced the difference and revealed decreased stability in a number of at-risk subjects. Smooth pursuit showed large errors in most patients and several controls but none of the at-risk subjects. Sporadic follow-up data show that at least two of the at-risk subjects developed manifest HD within a few years after passing the oculomotor test with entirely normal results. The material as a whole suggests that oculomotor dysfunction does not develop prior to, but concurrently with and as a part of generalized, progressive deterioration of motor control. The implication is that oculomotor screening of clinically healthy at-risk subjects does not reliably contribute to an earlier diagnosis of future HD.

Adult

Binocular co-ordination of human horizontal saccadic eye movements.

1. The binocular co-ordination of human horizontal saccades was analysed for the first time systematically over the full oculomotor range with a precise and accurate scleral sensor coil technique. Effects of amplitude (1.25-80 deg), direction (adduction vs. abduction and centrifugal vs. centripetal) and eccentricity (symmetrical about primary or between primary and secondary positions) were systematically investigated in three subjects). 2. To minimize extraneous effects of stimulus presentation on the programming of saccades, subjects were instructed to voluntarily change their gaze between two continuously visible targets. These were positioned on an iso-vergence locus, and thus contained no stimulus for disjunctive eye movements. 3. Under these conditions the amplitudes of the primary saccades of the two eyes were remarkably accurate; undershooting of the target by about 0.5 deg (independent of amplitude in the range 10-70 deg) was typical. This finding contrasts with the undershooting by about 10% described in the literature as characteristic for other stimulus conditions. 4. Saccadic peak velocities saturated at a mean asymptotic level of 502 +/- 32 (S.D.) deg/s for saccades of 40 deg and larger. The duration was linearly related to amplitude for saccades up to 50 deg; for saccades of larger sizes the duration increased progressively more steeply. Skewness values (acceleration time as a fraction of total saccadic duration) decreased from about 0.45 for saccades up to 10 deg to about 0.20 for saccades of 50 deg and larger. 5. Binocular saccades showed an abduction-adduction asymmetry and were not well yoked dynamically. The saccades of the abducting eye consistently had a larger size, a higher peak velocity, a shorter duration and were more skewed than the concomitant adducting saccades of the fellow eye. As a result, the eyes diverged transiently by as much as 3 deg during horizontal saccades. 6. Saccades also showed a marked centrifugal-centripetal asymmetry. Peak velocities of saccades towards the primary position were about 10% higher than peak velocities of corresponding centrifugal saccades. 7. These directional asymmetries were the main source of variability in the pool of saccades. In comparison, intra- and intersubject variability was minor in our sample. 8. Post-saccadic drift consisted of a vergence and a version component. The vergence component of this drift was a continuation of the vergence movement occurring during saccades. The version component, generally smaller than the vergence component, was directed towards the target position.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult

Binocular co-ordination of human vertical saccadic eye movements.

1. The binocular co-ordination of human vertical saccades was analysed systematically over the full oculomotor range, with a precise and accurate scleral sensor coil technique. Effects of amplitude (1.25-70 deg), direction (upward vs. downward and centripetal vs. centrifugal), as well as position (upper or lower sector of vertical oculomotor range), were investigated systematically in three subjects. 2. All saccades were made voluntarily between continuously presented pairs of targets, which subtended equal angles of target vergence. 3. Vertical saccades were less accurate than horizontal saccades (as described by Collewijn, Erkelens & Steinman, 1988). For target distances between 10 and 70 deg, upward saccades undershot the target by about 10%, whereas downward saccades tended to overshoot the target. Downward saccades were about 1.5 deg larger than upward saccades between the same targets. 4. Peak velocities continued to increase monotonically with saccadic amplitude up to 513 +/- 27 (S.D.) deg/s for 70 deg saccades; a distinct asymptotic level was not reached. 5. Velocity profiles of upward and downward saccades, made symmetrically about the primary (straight-ahead) position, were very similar for amplitudes up to 30 deg. At larger amplitudes, velocity profiles of upward saccades remained single peaked, whereas those of downward saccades invariably developed a second velocity peak. 6. Parameters of upward saccades depended heavily on the position of the eye. In the upper oculomotor range such saccades had lower maximum speeds, longer durations, and were more skewed than similar saccades in the lower oculomotor range (below primary). Downward saccades were almost independent of eye position. 7. Vertical eye movements during vertical saccades were virtually identical in the two eyes. In contrast, disjunctive horizontal components were systematically present. Upward saccades, at all amplitudes, were associated with diverging eye movements. Converging eye movements occurred during downward saccades. These systematic effects suggest that the vergence subsystem is not turned off during saccades. 8. These changes in vergence were followed by converging horizontal post-saccadic drift after upward saccades, and in diverging horizontal drift after downward saccades.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult

Visual and oculomotor function in optic chiasma-sectioned rabbits.

Anatomical and physiological findings indicate that the crossed optic fibres of the rabbit have a crucial role in binocular vision. In order to directly examine the visual functions of the uncrossed fibre system, a technique of sectioning the optic chiasma midsagitally was developed. Both normal and chiasma-sectioned rabbits were tested on a variety of visual discrimination tasks as well as such oculomotor control functions as the optokinetic and vestibulo-ocular reflexes. Following transection of all contralateral retinal projections, rabbits were found to retain the same visual capacity for detection of intensity and orientation differences as before the operation. There was, however, a complete loss of optokinetic reflexes and a 50% reduction of the vestibulo-ocular reflex both in the light and in the dark.

Animals

Voluntary smooth eye movements with foveally stabilized targets.

We investigated the capacity of 6 humans to make voluntary smooth eye movements with a horizontally stabilized foveal point target. When the target was viewed on a dark field, all subjects were able to make smooth oscillatory eye movements when they attempted to imitate their own normal pursuit of sinusoidal target movement (0.2-0.7 Hz) directly preceding the stabilization on the fovea. The frequency of the imitating eye movement was in general lower than the frequency of normal pursuit by 2-35%. While fixating a foveally stabilized point target superimposed on a large, sinusoidally moving non-stabilized background, all subjects were able to make either no eye movements, eye movements nearly in phase with or eye movements nearly in counterphase with the background movement depending on the instruction to imagine the target as head-stationary, moving in phase, or moving in counterphase with the background. The accuracy of the frequency of the smooth eye movement with the stabilized target on the moving background was higher than during imitation of pursuit on the dark field but the precision of the frequency was lower than during normal pursuit. When the background moved pseudo-randomly all subjects could voluntarily inhibit their smooth eye movements or could make smooth eye movements in phase with the background. Only 2 subjects showed a limited ability to make smooth eye movements opposite to the pseudo-random background movement. The results suggest that with predictable background movement the volition of the subject rather than the movement of the background determines the eye movements when the subject looks at the foveally stabilized target.

Eye Movements

A direct test of Listing's law--I. Human ocular torsion measured in static tertiary positions.

The validity of Listing's law was reinvestigated by means of a direct test. Horizontal, vertical and torsional eye movements were measured simultaneously with a recently developed scleral induction coil. Either eye of 4 subjects was measured monocularly. Eye position were measured in Fick coordinates and ocular torsion values were compared to the theoretical ones predicted by Listing's law. During consecutive measurements in the primary position torsion values were close to zero although considerable fluctuations of torsion were seen. Torsion values in the secondary positions were also close to zero. In the tertiary positions torsion in the direction as predicted by Listing's law and increasing with eccentricity was recorded. In the temporal quadrants mean torsion was quantitatively in agreement with Listing's law; torsion values in the nasal quadrants however showed systematically larger values and this discrepancy increased with eccentricity to more than 50%. Statistical support for this finding however, was seen only in 4 out of 8 eyes. Symmetry could be obtained by shifting the chosen horizontal primary position (gaze parallel to the midplane) in the temporal direction; as a consequence all measured torsion values would exceed the ones specified by Listing's law. Torsion values varied idiosyncratically among subjects and among the left and right eyes of any one subject. It is concluded that Listing's law specifies ocular torsion only approximately: physiological eye movements show considerable stochastical as well as systematical deviations from this law.

Eye Movements

A direct test of Listing's law--II. Human ocular torsion measured under dynamic conditions.

Ocular torsion was recorded with a scleral search coil technique in five normal subjects. The dynamic aspects of torsion were investigated during monocular fixation, blinking, smooth pursuit and saccades. Torsion near the primary position showed considerable short-term (SD about 0.25 deg) and a much larger long-term fluctuation (SD about 2.3 deg). During saccades between diagonally opposite tertiary positions torsion transiently reached values approximating those in the sustained primary position. During smooth pursuit across the primary position, the minimal values of torsion varied with the direction and the trajectory of pursuit, in violation of Donder's law. Changes in torsion associated with horizontal and vertical saccades and during the aftermath of blinks often had a sluggish, exponential time course. During eye movements around a circular or square trajectory torsion showed hysteresis. During clockwise pursuit the right eye showed relative intorsion compared to counterclockwise pursuit. It is proposed that central nervous control of torsion is usually imprecise, and that the eye follows Listing's and Donder's laws only approximately.

Blinking

Human gaze stability in the horizontal, vertical and torsional direction during voluntary head movements, evaluated with a three-dimensional scleral induction coil technique.

The stability of gaze in three dimensions (horizontal, vertical and torsion) was measured with a new type of scleral search coil in eight emmetropic observers. Subjects held the head still or oscillated it at 0.16-0.67 Hz (amplitude about 10 deg) in the horizontal, vertical or torsional plane while fixating a point target at optical infinity. Veridical gaze and head coordinates were calculated with full correction for non-linear goniometric relations and for cross-coupling artifacts due to misalignments of the coil on the eye. The amount of gaze instability in the horizontal and vertical direction was virtually identical. With the head still, in either of these directions the mean standard deviation of gaze position (inclusive saccades) was about 7 min arc; mean non-saccadic retinal image speeds were 20-30 min arc/sec. During head oscillation these values increased to about 16 min arc and 1 deg/sec; a mean of about 2.5% of the head motion remained uncorrected by the compensatory eye movements. These findings agree well with our earlier results for the horizontal plane; the effect of the corrections was relatively small because the adventitious cross-coupling of horizontal and vertical to torsional head movements proved to be usually smaller than 10%. However, the corrections were important when head torsion was deliberately produced. Gaze stability in the torsional plane was considerably inferior to that in the horizontal and vertical plane. With the head held still, the mean SD of torsional gaze position was about 17 min arc; mean torsional non-saccadic retinal image speed was about 46 min arc/sec. Gain of the torsional compensatory eye movements was frequency dependent and rose from about 0.26 in static conditions (0 Hz) to about 0.42 at 0.16 Hz and 0.64 at 0.67 Hz. Accordingly, position instability and speed of the retinal image in torsion were about an order of magnitude larger than in the horizontal and vertical direction.

Eye Movements

Control of human optokinetic nystagmus by the central and peripheral retina: effects of partial visual field masking, scotopic vision and central retinal scotomata.

Optokinetic nystagmus (OKN) was elicited in humans by a horizontally moving grating covering the whole visual field. Selective stimulation of central or peripheral parts of the retina was achieved by partial masking or scotopic viewing conditions in normals; three patients with a unilateral central retinal scotoma were studied in addition. In all cases, the elimination of foveal stimulation was accompanied by a decrease in OKN slow phase velocity compared to whole field stimulation. Vertical masks with retinally stabilized edges were used to selectively occlude or stimulate central or peripheral sectors with a fixed retinal location. A central stimulus was always more effective than the complementary peripheral stimulus, until the central zone was narrowed down to a width of 5-10 degrees. This central dominance was found throughout the range of velocities (6-180 degrees/s) and spatial frequencies (0.05-0.5 cycles/deg) used. A horizontal central band of occlusion caused a smaller decrease of OKN than a vertical occlusion with the same width. Scotopic vision caused a uniform mild decrease in OKN gain throughout the velocity range, provided that the spatio-temporal frequency of the stimulus remained within the scotopic resolution range. The patients had a slightly lower OKN gain when viewing with the scotomatous eye than with the contralateral, normal eye. The normal slight preference for temporal-to-nasal motion was not accentuated by masking or scotopic vision in normal eyes, but was enhanced in the eyes with the pathological scotomata in two of the three patients. All responses were immediate; no slow build-up was seen under any condition.

Adult