Frequency-dependent self-induced bias of the basilar membrane and its potential for controlling sensitivity and tuning in the mammalian cochlea.

Abstract

A displacement-sensitive capacitive probe technique was used in the first turn of guinea pig cochleas to examine whether the motion of the basilar membrane includes a displacement component analogous to the dc receptor potentials of the hair cells. Such a "dc" component apparently exists. At a given location on the basilar membrane, its direction toward scala vestibuli (SV) or scala tympani (ST) varies systematically with frequency of the acoustic stimulus. Furthermore, it appears to consist of two parts: a small asymmetric offset response to each gated tone burst plus a progressive shift of the basilar membrane from its previous position. The mean position shift is cumulative, increasing with successive tone bursts. The amplitude of the immediate offset response, when plotted as a function of frequency, appears to exhibit a trimodal pattern. This displacement offset is toward SV at the characteristic frequency (CF) of the location of the probe, while at frequencies either above or below the CF the offset is relatively larger, and toward ST. The mechanical motion of the basilar membrane therefore appears to contain the basis for lateral suppression. The cumulative mean position shift, however, appears to peak toward ST at the apical end of the traveling wave envelope and appears to be associated with a resonance, not of the basilar membrane motion directly, but coupled to it. The summating potential, measured concurrently at the round window, shows a more broadly tuned peak just above the CF of the position of the probe. This seems to correspond to the peak at the CF of the mechanical bias. As the preparation deteriorates, the best frequency of the vibratory displacement response decreases to about a half-octave below the original CF. There is a corresponding decrease in the frequency of the peaks of the trimodal pattern of the asymmetric responses to tone bursts. The trimodal pattern also broadens. In previous experiments the basilar membrane has been forced to move in response to a low-frequency biasing tone. The sensitivity to high-frequency stimuli varies in phase with the biasing tone. The amplitudes of slow movement in these earlier experiments and in the present experiments are of the same order of magnitude. This suggests strongly that the cumulative shift toward ST to a high-frequency acoustic stimulus constitutes a substantial controlling bias on the sensitivity of the cochlea in that same high-frequency region. Its effect will be to reduce the slope of neural rate-level functions on the high-frequency side of CF.