Three-dimensional eye-movement responses to off-vertical axis rotations in humans

Abstract. We recorded three-dimensional eye movements elicited by velocity steps about axes that were tilted with respect to the earth-vertical. Subjects were accelerated in 1 s from zero to 100°/s, and the axis of rotation was tilted by 15°, 30°, 60°, or 90°. This stimulus induced a constant horizontal velocity component that was directed opposite to the direction of rotation, as well as a modulation of the horizontal, vertical and torsional components with the frequency of the rotation. The maximum steady-state response in the horizontal constant-velocity component was much smaller than in other species (about 6°/s), reaching a maximum at a tilt angle of about 60°. While the amplitude of the horizontal modulation component increased up to a tilt angle of 90° (8.4°/s), the vertical and torsional modulation amplitudes saturated around 60° (ca. 2.5°/s). At small tilt angles, the horizontal modulation component showed a small phase lag with respect to the chair position, which turned into a small phase lead at large tilt angles. The torsional component showed a phase lead that increased with increasing tilt angle. The vertical and torsional velocity modulation at large tilt angles was not predicted by a recent model of otolith-canal interaction by Merfeld. Agreement between model and experimental data could be achieved, however, by introducing a constant force along the body's z-axis to compensate for the gravitational pull on the otoliths in the head-upright position. This approach had been suggested previously to explain the direction of the perceived subjective vertical during roll under different g-levels, and produced in our model the observed vertical and torsional modulation components at large tilt angles.