SUMMARY1. 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 (1X25-80 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 H. COLLEWIJN, C. J. ERKELENS AND R. M. STEINMAN 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. The result of postsaccadic drift was that the fovea of each eye was guided towards the target. The net post-saccadic drift of the abducting eye was smaller than that of the adducting eye, a result appropriate to reduce fixation errors remaining at saccadic offset.9. A tight relationship between skewness and saccade duration was not found. For instance, velocity profiles of centrifugal saccades were more skewed than ...

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SUMMARY1. 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 1P5 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.H. COLLEWIJN, C. J. ERKELENS AND R. M. STEINMAN 8. These changes in vergence were followed by converging horizontal postsaccadic drift after upward saccades, and in diverging horizontal drift after downward saccades. Vertical post-saccadic drift consisted mainly of a conjugated component, directed towards the target position. We conclude that post-saccadic drift on the vertical meridian is effective in decreasing binocular fixation errors in a way similar to error reduction following horizontal saccades.

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.