Vertical divergence and counterroll eye movements evoked by whole-body position steps about the roll axis of the head in humans.

In healthy human subjects, a head tilt about its roll axis evokes a dynamic counterroll that is mediated by both semicircular canal and otolith stimulation, and a static counterroll that is mediated by otolith stimulation only. The vertical ocular divergence associated with the static counterroll too is otolith-mediated. A previous study has shown that, in humans, there is also a vertical divergence during dynamic head roll, but this report was not conclusive on whether this response was mediated by the semicircular canals only or whether the otoliths made a significant contribution. To clarify this issue, we applied torsional whole-body position steps (amplitude 10 degrees, peak acceleration of 90 degrees /s(2), duration 650 ms) about the earth-vertical (supine body position) and earth-horizontal (upright body position) axis to healthy human subjects who were monocularly fixating a straight-ahead target. Eye movements were recorded binocularly with dual search coils in three dimensions. The dynamic parameters were determined 120 ms after the beginning of the turntable movement, i.e., before the first fast phase of nystagmus. The static parameters were measured 4 s after the beginning of the turntable movement. The dynamic gain of the counterroll was larger in upright (average gain: 0.48 +/- 0.10 SD) than in supine (0.36 +/- 0.10) position. The static gain of the counterroll in the upright position (0.21 +/- 0.06) was smaller than the dynamic gain. Divergent eye movements (intorting eye hypertropic) evoked during the dynamic phase were not significantly different between supine (average vergence velocity: 0.87 +/- 0.51 degrees /s) and upright (0.84 +/- 0.64 degrees /s) positions. The static vertical divergence in upright position was 0.32 +/- 0.14 degrees. The results indicate that the dynamic vertical divergence in contrast to the dynamic ocular counterroll is not enhanced by otolith input. These results can be explained through the different patterns of connectivity between semicircular canals and utricles to the eye muscles. Alternatively, we hypothesize that the small dynamic vertical divergence represents the remaining vertical error necessary to drive an adaptive control mechanism that normally maintains a vertical eye alignment.