Ve. Das et al., MODULATION OF HIGH-FREQUENCY VESTIBULOOCULAR REFLEX DURING VISUAL TRACKING IN HUMANS, Journal of neurophysiology, 74(2), 1995, pp. 624-632
1. Humans may visually track a moving object either when they are stat
ionary or in motion. To investigate visual-vestibular interaction duri
ng both conditions, we compared horizontal smooth pursuit (SP) and act
ive combined eye-head tracking (CEHT) of a target moving sinusoidally
at 0.4 Hz in four normal subjects while the subjects were either stati
onary or vibrated in yaw at 2.8 Hz. We also measured the visually enha
nced vestibuloocular reflex (VVOR) during vibration in yaw at 2.8 Hz o
ver a peak head velocity range of 5-40 degrees/s. 2. We found that the
gain of the VVOR at 2.8 Hz increased in all four subjects as peak hea
d Velocity increased (P < 0.001), with minimal phase changes, such tha
t mean retinal image slip was held below 5 degrees/s. However, no corr
esponding modulation in vestibuloocular reflex gain occurred with incr
easing peak head velocity during a control condition when subjects wer
e rotated in darkness. 3. During both horizontal SP and CEHT, tracking
gains were similar, and the mean slip speed of the target's image on
the retina was held below 5.5 degrees/s whether subjects were stationa
ry or being vibrated at 2.8 Hz. During both horizontal SP and CEHT of
target motion at 0.4 Hz, while subjects were vibrated in yaw, VVOR gai
n for the 2.8-Hz head rotations was similar to or higher than that ach
ieved during fixation of a stationary target. This is in contrast to t
he decrease of VVOR gain that is reported while stationary subjects pe
rform CEHT. 4. In a control experiment in which subjects carried out v
ertical SP and CEHT while they were vibrated in yaw at 2.8 Hz, we foun
d that three of four subjects showed an increase in horizontal VVOR ga
in at 2.8 Hz compared with that achieved during fixation of a stationa
ry target; such an increased horizontal gain would not be required to
reduce retinal image slip in the vertical plane. 5. On the basis of th
ese findings, we draw the following conclusions. I) During sinusoidal
oscillations at 2.8 Hz, the gain of the VVOR is adjusted in accordance
with peak head velocity in order to hold retinal slip of the image of
the visual target below similar to 5 degrees/s. 2) During visual trac
king of a moving target while the subject is in motion, there are two
potential sources of retinal image slip: imperfect Visual tracking and
an inadequate VVOR. When tracking deteriorates, it becomes necessary
to increase the gain of the VVOR to levels that prevent additional ret
inal image slip, so that vision is not compromised. 3) The increase of
horizontal VVOR gain that occurs during both horizontal and vertical
visual tracking while subjects are in motion may not be wholly due to
retinal slip per se, but may also involve a nonvisual mechanism that e
ffectively constrains retinal image slip to levels that permit clear v
ision.