Date of Award


Degree Type


Degree Name

Doctor of Philosophy


This thesis examines the neural mechanisms that generate torsional and vertical eye movements, and how they handle the kinematics of three-dimensional rotations. Three-dimensional eye rotations were recorded in alert monkeys, and the midbrain was studied using single unit recording, electrical microstimulation, and pharmacological inactivation.;The direction of VOR slow phases was usually opposite to that of head rotation, as required for optimal visual stabilization. However, direction errors sometimes occurred because of low gain about a head-fixed torsional axis. This axis was orthogonal to Listing's plane of saccadic eye positions, which suggests that saccades and the VOR share the same coordinate system. As predicted by the laws of rotational kinematics, even slow phase axes with zero torsional components produced torsional violations of Listing's law. This proves that mechanics of the plant are not responsible for Listing's law. Furthermore, the final positions were held. This signifies that the eye velocity signal is multiplied by position feedback before entering the oculomotor integrator. Finally, these violations of Listing's law were corrected by VOR quick phases.;Microstimulation and inactivation of the rostral interstitial nucleus of the medial longitudinal fasciculus (riMLF) demonstrated two right riMLF burst neuron populations that generate clockwise-upward-rightward and clockwise-downward-leftward rotations respectively, and two left riMLF populations for counterclockwise-upward-leftward and counterclockwise-downward-rightward rotations. The torsional axes of eye rotation evoked by riMLF stimulation were orthogonal to Listing's plane, whereas axes that remained after riMLF inactivation aligned with Listing's plane. This suggests that the neural coordinates of motor systems reflect their behavioral constraints.;The neural integrator, which generates the eye position signal, is central to oculomotor control. Post-saccadic drift during inactivation of the midbrain interstitial nucleus of Cajal (INC) showed that this nucleus is the integrator for vertical and torsional eye positions. Stimulation showed that the right INC controls clockwise rotations and the left INC controls counterclockwise rotations. Simulations of the drift suggested that the INC distributes integration over parallel independent neural compartments. This makes the integrator more computationally robust, and suggests a similar role for the modular connectivity observed throughout the brain.



To view the content in your browser, please download Adobe Reader or, alternately,
you may Download the file to your hard drive.

NOTE: The latest versions of Adobe Reader do not support viewing PDF files within Firefox on Mac OS and if you are using a modern (Intel) Mac, there is no official plugin for viewing PDF files within the browser window.