A direct test of Listing's law--II. Human ocular torsion measured under dynamic conditions

Ocular torsion was recorded with a scleral search coil technique in five normal subjects. The dynamic aspects of torsion were investigated during monocular fixation, blinking, smooth pursuit and saccades. Torsion near the primary position showed considerable short-term (SD about 0.25 deg) and a much larger long-term fluctuation (SD about 2.3 deg). During saccades between diagonally opposite tertiary positions torsion transiently reached values approximating those in the sustained primary position. During smooth pursuit across the primary position, the minimal values of torsion varied with the direction and the trajectory of pursuit, in violation of Donder's law. Changes in torsion associated with horizontal and vertical saccades and during the aftermath of blinks often had a sluggish, exponential time course. During eye movements around a circular or square trajectory torsion showed hysteresis. During clockwise pursuit the right eye showed relative intorsion compared to counterclockwise pursuit. It is proposed that central nervous control of torsion is usually imprecise, and that the eye follows Listing's and Donder's laws only approximately.