Otolith and canal reflexes in human standing.

We used galvanic vestibular stimulation (GVS) to identify human balance reflexes of the semicircular canals and otolith organs. The experiment used a model of vestibular signals arising from GVS modulation of the net signal from vestibular afferents. With the head upright, the model predicts that the GVS-evoked canal signal indicates lateral head rotation while the otolith signal indicates lateral tilt or acceleration. Both signify body sway transverse to the head. With the head bent forward, the model predicts that the canal signal indicates body spin about a vertical axis but the otolith signal still signifies lateral body motion. Thus, we compared electromyograms (EMG) in the leg muscles and body sway evoked by GVS when subjects stood with the head upright or bent forward. With the head upright, GVS evoked a large sway in the direction of the anodal electrode. This response was abolished with the head bent forward leaving only small, oppositely directed, transient responses at the start and end of the stimulus. With the head upright, GVS evoked short-latency (60-70 ms), followed by medium-latency (120 ms) EMG responses, of opposite polarity. Bending the head forward abolished the medium-latency but preserved the short-latency response. This is compatible with GVS evoking separate otolithic and canal reflexes, indicating that balance is controlled by independent canal and otolith reflexes, probably through different pathways. We propose that the short-latency reflex and small transient sway are driven by the otolith organs and the medium-latency response and the large sway are driven by the semicircular canals.