Human Visuospatial Updating After Passive Translations in Three-Dimensional Space

To maintain a stable representation of the visual environment as we move, the brain must update the locations of targets in space using extra-retinal signals. Humans can accurately update after intervening active whole-body translations. But can they also update for passive translations (i.e., without efference copy signals of an outgoing motor command)? We asked six head-fixed subjects to remember the location of a briefly flashed target (five possible targets were located at depths of 23, 33, 43, 63, and 150 cm in front of the cyclopean eye) as they moved 10 cm left, right, up, down, forward, or backward while fixating a head-fixed target at 53 cm. After the movement, the subjects made a saccade to the remembered location of the flash with a combination of version and vergence eye movements. We computed an updating ratio where 0 indicates no updating and 1 indicates perfect updating. For lateral and vertical whole-body motion, where updating performance is judged by the size of the version movement, the updating ratios were similar for leftward and rightward translations, averaging 0.84 ± 0.28 (mean ± SD) as compared with 0.51 ± 0.33 for downward and 1.05 ± 0.50 for upward translations. For forward/backward movements, where updating performance is judged by the size of the vergence movement, the average updating ratio was 1.12 ± 0.45. Updating ratios tended to be larger for far targets than near targets, although both intra- and intersubject variabilities were smallest for near targets. Thus in addition to self-generated movements, extra-retinal signals involving otolith and proprioceptive cues can also be used for spatial constancy.

Saccadic Behavior during the Response to Pure Vergence Stimuli I: General Properties

If two targets are carefully aligned so that they fall along the cyclopean axis, the required eye movement will be symmetrical with the two eyes turning equally inward or outward. When such “pure vergence stimuli” are used only a “pure vergence movement” is required, yet almost all responses include saccadic eye movements, a rapid tandem movement of the eyes. When saccades occur, they must either produce an error in the desired symmetrical response or correct an error from an asymmetrical vergence response. A series of eye movement responses to pure convergence stimuli (4.0 deg step stimuli) were measured in 12 subjects and the occurrence, timing and amplitude of saccades was measured. Early saccades (within 400 msec of the stimulus onset) appeared in 80% to 100% of the responses. In most subjects, the first saccade increased the asymmetry of the response, taking the eyes away from the midline position. In three subjects, these asymmetry-inducing saccades brought one eye, the preferred or dominant eye, close to the target, but in the other subjects these asymmetry-inducing saccades were probably due to the distraction caused by the transient diplopic image generated by a pure vergence stimulus. While many of these asymmetry-inducing saccades showed saccade-like enhancements of vergence, they were, with the exception of two subjects, primarily divergent and did not facilitate the ongoing convergence movement. All subjects had some responses where the first saccade improved response symmetry, correcting an asymmetry brought about by unequal vergence movements in the two eyes. In five subjects, large symmetry-inducing saccades corrected an asymmetrical vergence response, bringing the eyes back to the midline (to within a few tenths of a degree).

Neural control of vergence eye movements: neurons encoding vergence velocity

Single-unit recordings were made from midbrain areas in monkeys trained to make both conjugate and disjunctive (vergence) eye movements. Previous work had identified cells with a firing rate proportional to the vergence angle, without regard to the direction of conjugate gaze. The present study describes the activity of neurons that burst for disjunctive eye movements. Convergence burst cells display a discrete burst of activity just before and during convergence eye movements. For most of these cells, the profile of the burst is correlated with instantaneous vergence velocity and the number of spikes in the burst is correlated with the size of the vergence movement. Some of these cells also have a tonic firing rate that is positively correlated with vergence angle (convergence burst-tonic cells). Divergence burst cells have similar properties, except that they fire for divergent and not convergent movements. Divergence burst cells are encountered far less often than convergence burst cells. Both convergence and divergence burst cells were found in an area of the mesencephalic reticular formation just dorsal and lateral to the oculomotor nucleus. Convergence burst cells were also recorded in another more dorsal mesencephalic region, rostral to the superior colliculus. Both of the areas also contain cells that encode vergence angle. Models of the vergence system derived from psychophysical data imply the existence of a vergence integrator, the output of which is vergence angle. Some models also suggest the presence of a parallel element that improves the frequency response of the vergence system, but has no effect on the steady-state behavior of the system. Vergence burst cells would be suitable inputs to a vergence integrator. By providing a vergence velocity signal to motoneurons, they may improve the dynamic response of the vergence system. The behavior of vergence burst cells during vergence movements is similar to that of the medium-lead burst cells during saccades. The proposed roles for vergence velocity cells are analogous to those of the saccadic burst cells. In this respect, the neural organization of the vergence system resembles that of the saccadic system, despite the distinct difference in the kinematics of these two types of eye movements.