Casper J. Erkelens

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)

Eye movements and stereopsis during dichoptic viewing of moving random-dot stereograms

The dynamic properties of the version and vergence system were studied in relation to stereopsis for movements of the whole visual scene. Large random-dot stereograms (30 X 30 deg arc), moving laterally, were viewed dichoptically by human subjects without a fixed visual frame of reference. Sinusoidal movements in counterphase of the two half-images constituting the stereogram induced sinusoidal ocular vergence movements. The gain of vergence depended on the frequency as well as the amplitude of stimulus movement, while the phase lag depended only on the frequency. Fusion and stereopsis were retained up to a maximal velocity of change in relative position of the two half-images between 6 and 13.5 deg/sec. Sinusoidal movement of one half-image while the other one remained stationary induced sinusoidal ocular version as well as vergence movements. For version gains were higher and phase lags were smaller than for vergence. At the retinal level, residual overall binocular disparities between the two half-images up to 2 deg arc were tolerated without loss of stereopsis. The presence of sinusoidally varying overall binocular disparities and ocular vergence movements without perception of motion in depth suggests that these variables are not adequate cues for perception of (change in) depth.

Motion perception during dichoptic viewing of moving random-dot stereograms.

The relation between binocular and monocular motion perception was investigated. A random-dot stereogram (30 X 30 deg arc), containing a central figure seen in front of the background in stereoscopic vision, was viewed dichoptically by human subjects without a fixed visual frame of reference. The images seen by the right and left eye were moved laterally according to a triangular wave form, in counterphase, but with variable amplitude ratios. Under this condition only purely lateral movement and no motion in depth of the stereogram as a whole was perceived, while stereoscopic vision of the figure-background relation was maintained. The magnitude of the binocularly perceived lateral motion, signalled by manual tracking of the perceived displacement, equalled the algebraic mean of the monocular motion percepts. As a special case, when the two images forming the stereogram were moved with equal velocities but in opposite directions they were perceived as a completely stationary, fused image in stereoscopic depth. Only the addition of a stationary reference (a bar or grating seen by both eyes) resulted in the perception of motion in depth. We conclude that a visual frame of reference is essential for perception of motion in depth but not for perception of lateral movements. Moreover, it seems likely that not absolute binocular disparity (retinal locus differences) but relative binocular disparity (differences in angular distance between two or more corresponding features in the two retinal images) is a cue for perception of depth.

Differences in coordination of elbow flexor muscles in force tasks and in movement tasks

SummaryMotor-unit activity in m. biceps brachii, m. brachialis and m. brachioradialis during isometric contractions has been compared with motor-unit activity during slow voluntary (extension and flexion) movements made against external loads. During these slow movements the recruitment threshold of m. biceps motor units is considerably lower than it is during isometric contractions but the recruitment threshold of both m. brachialis and m. brachioradialis motor units is considerably higher. For all three elbow flexor muscles the motor-unit firing frequency seems to depend on the direction of movement: the firing frequency is higher during flexion movements (3 deg/s) and lower during extension movements (−3 deg/s) than during isometric contractions. The relative contribution of the biceps to the total exerted flexion torque during slow voluntary movements is estimated to increase from 36% to about 48% and that of the brachialis/brachioradialis is estimated to decrease from 57% to about 45% compared to the relative contribution of these muscles during isometric contractions. This difference in the relative contribution of the three major elbow flexor muscles is shown to be caused by differences in the central activation in force tasks and movement tasks.

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° s-1 for vergence changes of 5° to between 150 and 200° s-1 for vergence changes of 34°. Differences between convergence and divergence were idiosyncratic. Asymmetrical vergence, requiring a vergence of 11° combined with a version of 45°, 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° s-1 for a change of vergence of 11°); much faster than the smooth, symmetrical vergence change of comparable size (14°). 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. We also found that asymmetric vergence was largely saccadic and conclude that the generation of saccades of unequal sizes in each of the eyes is a normal feature of oculomotor performance whenever gaze is shifted between targets that differ in distance as well as direction.

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)

CONTROL OF FIXATION DURATION IN A SIMPLE SEARCH TASK

To obtain insight into the control of fixation duration during visual search, we had 4 subjects perform simple search tasks in which we systematically varied the discriminability of the target. The experiment was carried out under two conditions. Under the first condition (blocked), the discriminability of the target was kept constant during a session. Under the second condition (mixed), the discriminability of the target varied per trial. Under the blocked condition, fixation duration increased with decreasing discriminability. For 2 subjects, we found much shorter fixation durations in difficult trials with the mixed condition than in difficult trials with the blocked condition. Overall, the subjects fixated the target, continued to search, and then went back to the target in 5%–55% of the correct trials. In these trials, the result of the analysis of the foveal target was not used for preparing the next saccade. The results support a preprogramming model of the control of fixation duration. In a simple search task, control of fixation duration appears to be indirect.

Adjustment of fixation duration in visual search

To investigate whether fixation durations are adjusted to the duration of a foveal analysis task, we designed a search task in which each stimulus element yielded information about the position of the target. We asked subjects to look for the target by making eye movements in the direction indicated by each stimulus element. We explicitly asked the subjects to make the eye movements in the correct direction, but they did not always do this. They made only 65-80% of the eye movements in directions indicated by the stimulus elements. From these results we conclude that fixation durations are not solely determined by the immediate visual stimulus and that subjects encounter difficulties when trying to increase fixation durations to values that would enable them to direct saccades accurately. In a second experiment we shortened the presentation time in order to provide an incentive for the subjects to speed up search. Shortening the presentation time did not affect fixation duration. Therefore, we suggest that fixation duration is controlled by a mechanism that uses estimations of the foveal analysis time of previous fixated stimulus elements.

Adjustments of fast goal-directed movements in response to an unexpected inertial load

SummarySubjects made fast goal-directed elbow flexion movements against an inertial load. Target distance was 8 or 16 cm, randomly chosen. To exert a force in the direction of the movement subjects had to activate flexors of both shoulder and elbow, but shoulder flexors did not change appreciably in length during the movement. In 20% of the trials the inertial load was increased or decreased without knowledge of the subjects. Until 90–110 ms after the onset of the agonist muscle activity (about 65–85 ms after the start of movement) EMG activity was very similar in all conditions tested. The changes that occured in the EMG from that moment on were effectively a later cessation of the agonist activity and a later start of the antagonist activity if the load was increased unexpectedly. If the load was reduced unexpectedly, the agonist activity ceased earlier and the antagonist activity began earlier. The latency at which EMGs started to change was the same for muscles around shoulder and elbow, for agonists and antagonists and for both distances. All adjustments had the same latency (37 ms) relative to the point where the angular velocity of the elbow in the unexpectedly loaded movements differed by 0.6 rad/s from the expected value. We discuss why simple reflex- or servo-mechanisms cannot account for the measured EMG changes. We conclude that appropriate adjustments of motor programmes for fast goal-directed arm movements start within 40 ms of the detection of misjudgment of load.

Dependence of autogenic and heterogenic stretch reflexes on pre-load activity in the human arm.

1. Subjects held their right arm in a horizontal plane. The angle of the elbow was 90 deg. They exerted forces in several directions in the plane of the arm, varying independently the pre‐load torques about shoulder and elbow. We measured electromyographic (EMG) activity in several arm muscles in response to force perturbations which extended the shoulder, without changing the elbow angle. 2. The EMG activity in flexors of both shoulder and elbow showed reflex responses at short latency (approximately 25 ms). In all muscles the reflex activity increased with the pre‐load activity of that muscle. 3. The short‐latency reflex activity of m. brachialis, which was not stretched by the perturbations, was independent of the pre‐load activity of the muscles acting over the shoulder. 4. From these results we conclude that the force resulting from the short‐latency reflex, assessed from the EMGs, does not counteract the perturbations exactly. Having found that the short‐latency reflex is dependent on the pre‐load direction, we argue that this dependence makes the short‐latency reflex suitable for correcting fast movements for misjudgements of load. 5. At longer latencies (greater than 50 ms) the direction of the force resulting from the reflex, assessed from the EMGs, was almost independent of the direction of the pre‐load. In our experiment the force resulting from the long‐latency reflex counteracted the perturbations quite well.