Reflex and mechanical contributions to head stabilization in alert cats

Abstract

Five alert cats were tested for their responses to rotation in a device that allowed rotation of the head on the trunk about a vertical axis passing through the C1-C2 vertebral joint. Electrodes were implanted to record the horizontal and vertical electrooculogram and electromyographic (EMG) activity of the dorsal neck muscles splenius, biventer cervicus, and complexus. Head rotation and torque acting on the head were recorded in the horizontal plane during rotations in the 0.05-5.0 Hz frequency range. Responses were interpreted with reference to a closed-loop dynamic model of the head-neck system. Whole-body rotation (WBR) with no neck movement elicited a vestibulocollic reflex (VCR). Neck muscle EMG lagged the sinusoidal platform rotation by approximately 120 degrees at low frequencies, which represents a 60 degrees lead relative to a perfectly compensatory 180 degrees lag. This phase lead was related to the cumulative eye position of the accompanying horizontal vestibular nystagmus as reported by Vidal et al. Horizontal head torque exhibited a similar low-frequency behavior. At high frequencies, EMG exhibited a progressively increasing phase lead and gain increase typical of a second-order lead system as described in decerebrate cats. Torque, however, showed much less lead and gain increase, presumably because of the low-pass filter properties of the process coupling muscle excitation to torque. Head torque did exhibit a steep increase in gain with frequency and a phase approaching that of platform acceleration at high frequencies when weights were attached to the head to increase its moment of inertia. The same +40 dB/decade gain slope and phase approaching 0 degree was observed during WBR rotation of the anesthetized cat in which head inertia is the only factor contributing to the torque. This dynamic behavior was predicted by the inertial component of the model. In the alert unweighted cat, the inertial torque was smaller than VCR-generated torque at frequencies below 4 Hz. Rotation of the neck with the head held fixed in space (HFS rotation) elicited a cervicocollic reflex (CCR). Neck EMG response was very similar to that observed during WBR, both in dynamic behavior and overall gain. Torque, however, was consistently greater than that generated by WBR and showed a steady increase of 8 dB/decade as frequency rose. The added torque can be attributed to the viscoelastic properties of neck muscles. Driven rotation of the head on the fixed body elicited torques that could be closely approximated by a vector sum of torques observed during WBR and HFS rotations.(ABSTRACT TRUNCATED AT 400 WORDS)