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The sensitivity and frequency selectivity of hearing result from tunedamplification by an active process in the mechanoreceptive hair cells. In mostvertebrates the active process stems from the active motility of hair bundles.The mammalian cochlea exhibits an additional form of mechanical activity termedelectromotility: its outer hair cells (OHCs) change length upon electricalstimulation. The relative contributions of these two mechanisms to the activeprocess in the mammalian inner ear is the subject of intense current debate.Here we show that active hair-bundle motility and electromotility can togetherimplement an efficient mechanism for amplification that functions like aratchet: sound-evoked forces acting on the basilar membrane are transmitted tothe hair bundles whereas electromotility decouples active hair-bundle forcesfrom the basilar membrane. This unidirectional coupling can extend the hearingrange well below the resonant frequency of the basilar membrane. It therebyprovides a concept for low-frequency hearing that accounts for a variety ofunexplained experimental observations from the cochlear apex, including theshape and phase behavior of apical tuning curves, their lack of significantnonlinearities, and the shape changes of threshold tuning curves of auditorynerve fibers along the cochlea. The ratchet mechanism constitutes a generaldesign principle for implementing mechanical amplification in engineeringapplications.

A ratchet mechanism for amplification in low-frequency mammalian hearing