Vestibular mechanisms related to vertical and torsional eye movements

1. A series of experiments concerning the organization of the vertical vestibuloocular reflex (VOR) of the rabbit are reviewed. 2. Unanesthetized, encéphale isolé and intact, alert rabbits have been submitted to natural stimulation of the labyrinth consisting in sinusoidal oscillations and static tilts around the longitudinal axis. The oculomotor responses have been recorded at the level of the III nucleus oculomotor neurons (OMN) and from the four vertical eye muscles. 3. The phase and gain of the OMN response at different frequencies of oscillation indicate a serial involvement of the macular ampullar labyrinthine receptors in the activation of the OMN. In the low frequency range of head movements, macular receptors alone can provide adequate ocular compensation. By raising the frequency of head movements, the threshold value of acceleration for recruiting second order vestibular neurons related to semicircular canals is reached and the vertical VOR is controlled predominantly by the ampullar receptors of the vertical canals. 4. These results have been confirmed and extended in the alert, intact rabbit. By orienting the animal in different positions with respect to the axis of oscillation it has been possible to activate selectively one or the other pair of complementary vertical canals. The EMG responses recorded from the vertical eye muscle exhibited different magnitudes in the different orientations, while the phase curve remained the same. 5. The static otolith-ocular reflex has also been investigated by tilting the alert animals oriented in the same way. The pattern of the EMG responses elicited by the pure macular input coincides with that obtained during dynamic stimulation. The existence of separate functional macular regions specifically connected with the different eye muscles and a close convergent action of the two types of labyrinthine receptors in ocular compensation, are suggested.