Somatic motility and hair bundle mechanics, are both necessary for cochlear amplification?

Hearing organs have evolved to detect sounds across several orders of magnitude of both intensity and frequency. Detection limits are at the atomic level despite the energy associated with sound being limited thermodynamically. Several mechanisms have evolved to account for the remarkable frequency selectivity, dynamic range, and sensitivity of these various hearing organs, together termed the active process or cochlear amplifier. Similarities between hearing organs of disparate species provides insight into the factors driving the development of the cochlear amplifier. These properties include: a tonotopic map, the emergence of a two hair cell system, the separation of efferent and afferent innervations, the role of the tectorial membrane, and the shift from intrinsic tuning and amplification to a more end organ driven process. Two major contributors to the active process are hair bundle mechanics and outer hair cell electromotility, the former present in all hair cell organs tested, the latter only present in mammalian cochlear outer hair cells. Both of these processes have advantages and disadvantages, and how these processes interact to generate the active process in the mammalian system is highly disputed. A hypothesis is put forth suggesting that hair bundle mechanics provides amplification and filtering in most hair cells, while in mammalian cochlea, outer hair cell motility provides the amplification on a cycle by cycle basis driven by the hair bundle that provides frequency selectivity (in concert with the tectorial membrane) and compressive nonlinearity. Separating components of the active process may provide additional sites for regulation of this process.