Tuning the cochlea: wave-mediated positive feedback between cells

Abstract

Frequency analysis by the mammalian cochlea is traditionally thought to occur via a hydrodynamically coupled 'travelling wave' along the basilar membrane. A persistent difficulty with this picture is how sharp tuning can emerge. This paper proposes, and models, a supplementary or alternative mechanism: it supposes that the cochlea analyses sound by setting up standing waves between parallel rows of outer hair cells. In this scheme, multiple cells mutually interact through positive feedback of wave-borne energy. Analytical modelling and numerical evaluation presented here demonstrate that this can provide narrow-band frequency analysis. Graded cochlear tuning will then rely on the distance between rows becoming greater as distance from the base increases (as exhibited by the actual cochlea) and on the wave's phase velocity becoming slower. In effect, tuning is now a case of varying the feedback delay between the rows, and a prime candidate for a wave exhibiting suitably graded phase velocity-a short-wavelength 'squirting wave'-is identified and used in the modelling. In this way, resonance between rows could supply both amplification and high Q, characteristics underlying the 'cochlear amplifier'-the device whose action has long been evident to auditory science but whose anatomical basis and mode of operation are still obscure.