The otolithic contribution to vertical ocular stability in the cat.

Abstract

In cats, horizontal (HVOR) and vertical (VVOR) vestibulo-ocular reflexes were studied alone and combined with optokinetic stimulation. The upright VVOR (VVOR O degree) only showed higher gain and smaller phase lead compared to those of HVOR at frequencies below 0.05 Hz. The addition of optokinetic stimulation to the vestibular stimulation increased the gain of the horizontal and vertical ocular responses close to 1. VVOR was also studied in side down position (VVOR 90 degrees). In VVOR 90 degrees the ocular responses were asymmetric. The downward directed eye responses of VVOR 90 degrees showed lower gain and greater phase lead compared to those of VVOR 0 degree for the whole range of tested frequencies (0.01-0.4 Hz), while the upward eye responses only showed a lower gain at the lower range of frequencies tested. In the light the gain of VVOR 90 degrees increased, but the gain of downward directed eye responses was consistently lower than 1 at lower frequencies. The higher gain of the VVOR 0 degree compared to the VVOR 90 degrees and HVOR was attributed to the maculo-ocular reflex (MOR) evoked by the gravity modulation of the otolithic receptors, when the animals were oscillated in the pitch plane. The MOR was isolated from the VVOR 0 degree by plugging all semicircular canals. At very low frequencies the gain of the MOR was 0.3-0.35 and the phase was close to 0 degree. This reflex showed a progressive gain decrease and phase lag by increasing the stimulation frequencies. This suggests a low pass filtering process of the otolithic signal. Furthermore in plugged animals the asymmetry of the vertical optokinetic responses was reduced by adding the MOR. The quick phases (QPs) of the vestibular responses were also different depending upon the stimulation plane. The QPs of VVOR 0 degree were smaller and more delayed than those of HVOR and VVOR 90 degrees. In conclusion the main effects observed during otolithic coactivation in the VVOR 0 of the cat are: 1) the enhancement of gain and reduction of phase lead at low frequency vestibular stimulation, resulting in similar vertical and horizontal gaze stability; 2) the equalization of the upward and downward responses of both vestibulo-ocular and optokinetic responses; 3) the reduction of the amplitude and frequency of vertical quick phases.