The human vestibulo-ocular reflex during linear locomotion.

During locomotion, there is a translation and compensatory rotation of the head in both the vertical and horizontal planes. During moderate to fast walking (100 m/min), vertical head translation occurs at the frequency of stepping (2 Hz) and generates peak linear acceleration of 0.37 g. Lateral head translation occurs at the stride frequency (1 Hz) and generates peak linear acceleration of 0.1 g. Peak head pitch and yaw angular velocities are approximately 17 degrees/s. The frequency and magnitude of these head movements are within the operational range of both the linear and angular vestibulo-ocular reflex (IVOR and aVOR). Vertical eye movements undergo a phase reversal from near to far targets. When viewing a far (>1 m) target, vertical eye velocity is typical of an aVOR response; that is, it is compensatory for head pitch. At close viewing distances (<1 m), vertical eye velocity is in phase with head pitch and is compensatory for vertical head translation, suggesting that the IVOR predominantly generates the eye movement response. Horizontal head movements during locomotion occur at the stride frequency of 1 Hz, where the IVOR gain is low. Horizontal eye movements are compensatory for head yaw at all viewing distances and are likely generated by the aVOR.