A minimal biophysical model of the cochlea is used to investigate the validity of the hypothesis that a single compressive nonlinearity at the hair cell level can explain some of the suppression phenomena in cochlear responses to complex stimuli. The dependencies of the model responses on the amplitudes and frequencies of two-tone stimuli resemble in many respects the behavior of the experimental data, and can be traced to explicit biophysical parameters in the model. Most discrepancies between theory and experiment stem from simplifications in parameters of the minimal model that play no direct role in the hypothesis. The analysis and simulations predict further results which, pending experimental verification, may provide a more direct test of the influence of the compressive nonlinearity on the relative amplitudes of the synchronous response components, and hence of its role in synchrony suppression. For instance, regardless of the overall absolute levels of a two-tone stimulus applied to this type of model, the ratio of the amplitudes at the input and the ratio of the corresponding responses at the output remain approximately constant and equal (the output ratio changes by at most 6 dB in favor of the stronger tone). Other nonlinear responses to multitonal stimuli can also be reproduced, such as "spectral edge enhancement" [Horst et al., Peripheral Auditory Mechanisms (Springer, Berlin, 1985)] and some aspects of three-tone suppression [Javel et al., Mechanisms of Hearing (Monash U.P., Australia, 1983)]. In contrast to the complex behavior of suppression with increasing sound intensity and the drastic influence of the compressive nonlinearity on the absolute response measures on the auditory nerve (e.g., average rate and synchrony profiles), the percepts of complex sounds are relatively stable. This suggests that the invariant relative response measures are more likely used in the encoding and CNS extraction of the spectrum of complex stimuli such as speech.
A mathematical model of cochlear processing is developed to account for the nonlinear dependence of frequency selectivity on intensity in inner hair cell and auditory nerve fiber responses. The model describes the transformation from acoustic stimulus to intracellular hair cell potentials in the cochlea. It incorporates a linear formulation of basilar membrane mechanics and subtectorial fluid-cilia displacement coupling, and a simplified description of the inner hair cell nonlinear transduction process. The analysis at this stage is restricted to low-frequency single tones. The computed responses to single tone inputs exhibit the experimentally observed nonlinear effects of increasing intensity such as the increase in the bandwidth of frequency selectivity and the downward shift of the best frequency. In the model, the first effect is primarily due to the saturating effect of the hair cell nonlinearity. The second results from the combined effects of both the nonlinearity and of the inner hair cell low-pass transfer function. In contrast to these shifts along the frequency axis, the model does not exhibit intensity dependent shifts of the spatial location along the cochlea of the peak response for a given single tone. The observed shifts therefore do not contradict an intensity invariant tonotopic code.