Reflections on the role of a traveling wave along the basilar membrane in view of clinical and experimental findings

Air conduction (AC) is accompanied by displacements of the two cochlear windows, bulk fluid flow between them, a pressure difference across the basilar membrane, leading to a passive traveling wave along the membrane, which activates the cochlear amplifier and enhances the displacements. AC interacts with bone conduction (BC) stimulation, so that it has been assumed that BC stimulation also involves a passive traveling wave. However, several clinical conditions and experimental manipulations provide evidence that a passive traveling wave may not be involved in BC stimulation at low intensities. Soft tissue conduction (STC) (also called non-osseous bone conduction) involves applying the bone vibrator to soft tissues on the head, neck and thorax, eliciting auditory sensation. STC stimulation probably does not involve a passive traveling wave. This review presents clinical conditions and experimental manipulations which assess the contributions of AC, BC and STC stimulation to the passive traveling wave. Evidence from the clinic (otosclerosis, round window atresia) and from the laboratory (holes in the wall of the inner ear, immobilization of the ossicular chain and the windows, discontinuity of the chain, measurement of basilar membrane displacements in the absence of the cochlear amplifier) lead to the conclusion that a passive basilar membrane traveling wave may not be involved in stimulation at low sound intensities. It is suggested that at low sound levels, the outer hair cell cochlear amplifier may not be activated by a passive traveling wave, but may be directly activated by the fast cochlear fluid pressures induced by AC, BC and STC stimulation. On the other hand, at high intensities, the cochlea is activated by the slow passive traveling wave.