Influence of static head position on the horizontal nystagmus evoked by caloric,rotational and optokinetic stimulation in the squirrel monkey

Summary We studied the influence of static head position on the horizontal nystagmus produced by caloric, rotational and optokinetic stimulation in alert squirrel monkeys. Caloric nystagmus is stronger for nose up (NU) than for nose down (ND) pitches; so, for example, slow-phase eye velocity is four times larger in supine than in prone positions. A similarly directed asymmetry occurs in the horizontal vestibulo-ocular (HVOR) responses to longduration, constant angular-head accelerations, but not to midband (0.1 Hz) sinusoidal head rotations. Consistent with a first-order model of the HVOR, the low-frequency or acceleration gain of the reflex (G_A) is equal to the product of the midband velocity gain (G_V) and a time constant (T_VOR). G_V is proportional to the cosine of the angle between the horizontal-canal plane and the plane of rotation, from which it is concluded that signals from the horizontal, but not from the vertical canals contribute to the HVOR. T_VOR can be as much as twice as large in NU than in ND positions. G_A is proportional to T_VOR and it, too, shows a NU-ND asymmetry. The time constant of optokinetic afternystagmus (T_OKAN) was also studied. Since T_VOR and T_OKAN are modified in similar ways by static tilts, it is concluded that head position affects the time constants by way of velocity-storage mechanisms. Evidence is presented that the position-dependent modification of velocity storage is otolith-mediated. The results are used to analyze the mechanisms of caloric nystagmus. The caloric response consists of a convective component (C_C), as originally envisioned by Bárány (1906), and a nonconvective component (N_C). C_C accounts for 75% of the caloric response in the conventional supine testing position. Both components can be affected by the position-dependent modification of T_VOR or, equivalently, of G_A. It has been suggested that two mechanisms might contribute to N_C: 1) a direct thermal effect on hair cells or afferents; or 2) a thermal expansion of labyrinthine fluids that results in a cupular displacement. Both theoretical and experimental evidence indicates that only the first of these mechanisms could result in the steady-state caloric response that is observed in the absence of convection (e.g., in spaceflight and after canal plugging) and that contributes to the prone-supine asymmetry seen in caloric testing.