Spectral Ripples in Round-Window Cochlear Microphonics: Evidence for Multiple Generation Mechanisms.

The cochlear microphonic (CM) results from the vector sum of outer hair cell transduction currents excited by a stimulus. The classical theory of CM generation-that the response measured at the round window is dominated by cellular sources located within the tail region of the basilar membrane (BM) excitation pattern-predicts that CM amplitude and phase vary little with stimulus frequency. Contrary to expectations, CM amplitude and phase-gradient delay measured in response to low-level tones in chinchillas demonstrate a striking, quasiperiodic pattern of spectral ripples, even at frequencies > 5 kHz, where interference with neurophonic potentials is unlikely. The spectral ripples were reduced in the presence of a moderate-level saturating tone at a nearby frequency. When converted to the time domain, only the delayed CM energy was diminished in the presence of the saturator. We hypothesize that the ripples represent an interference pattern produced by CM components with different phase gradients: an early-latency component originating within the tail region of the BM excitation and two delayed components that depend on active cochlear processing near the peak region of the traveling wave. Using time windowing, we show that the early, middle, and late components have delays corresponding to estimated middle-ear transmission, cochlear forward delays, and cochlear round-trip delays, respectively. By extending the classical model of CM generation to include mechanical and electrical irregularities, we propose that middle components are generated through a mechanism of "coherent summation" analogous to the production of reflection-source otoacoustic emissions (OAEs), while the late components arise through a process of internal cochlear reflection related to the generation of stimulus-frequency OAEs. Although early-latency components from the passive tail region typically dominate the round-window CM, at low stimulus levels, substantial contributions from components shaped by active cochlear processing provide a new avenue for improving CM measurements as assays of cochlear health.