Dual contribution to amplification in the mammalian inner ear

The inner ear achieves a wide dynamic range of responsiveness by mechanically amplifying weak sounds. The enormous mechanical gain reported for the mammalian cochlea, which exceeds a factor of 4,000, poses a challenge for theory. Here we show how such a large gain can result from an interaction between amplification by low-gain hair bundles and a pressure wave: hair bundles can amplify both their displacement per locally applied pressure and the pressure wave itself. A recently proposed ratchet mechanism, in which hair-bundle forces do not feed back on the pressure wave, delineates the two effects. Our analytical calculations with a WKB approximation agree with numerical solutions.