Evidence for a reference frame transformation of vestibular signal contributions to voluntary reaching.

To contribute appropriately to voluntary reaching during body motion, vestibular signals must be transformed from a head-centered to a body-centered reference frame. We quantitatively investigated the evidence for this transformation during online reach execution by using galvanic vestibular stimulation (GVS) to simulate rotation about a head-fixed, roughly naso-occipital axis as human subjects made planar reaching movements to a remembered location with their head in different orientations. If vestibular signals that contribute to reach execution have been transformed from a head-centered to a body-centered reference frame, the same stimulation should be interpreted as body tilt with the head upright but as vertical-axis rotation with the head inclined forward. Consequently, GVS should perturb reach trajectories in a head-orientation-dependent way. Consistent with this prediction, GVS applied during reach execution induced trajectory deviations that were significantly larger with the head forward compared with upright. Only with the head forward were trajectories consistently deviated in opposite directions for rightward versus leftward simulated rotation, as appropriate to compensate for body vertical-axis rotation. These results demonstrate that vestibular signals contributing to online reach execution have indeed been transformed from a head-centered to a body-centered reference frame. Reach deviation amplitudes were comparable to those predicted for ideal compensation for body rotation using a biomechanical limb model. Finally, by comparing the effects of application of GVS during reach execution versus prior to reach onset we also provide evidence that spatially transformed vestibular signals contribute to at least partially distinct compensation mechanisms for body motion during reach planning versus execution.