A model of the odontocete middle ear.

The high acoustic sensitivity of the bottlenose dolphin is physically defined and related to the anatomy of the middle ear. The paper presents a conceptual and parametric analysis of the demands imposed by this high sensitivity upon the middle ear mechanisms: the head and the middle ear structures must collect sound energy from a large area and concentrate it onto the oval window. Assuming that the specific input impedance of the mammalian cochlea is relatively constant, and smaller than the characteristic acoustic impedance of water, we find that the impedance matching task of the cetacean middle ear is very different from that of terrestrial mammals: instead of a large pressure amplification, cetaceans need amplification of particle velocity. Our mechanical four-bone model of the odontocete middle ear is based on the anatomy of the tympano-periotic complex and consists of four rigid bone units (tympanic bone, the malleus-incus complex, stapes, periotic bone) connected through elastic junctions. The velocity amplification is brought about by lever mechanisms and elastic couplings. The model produced velocity amplifications ranging from 7- to 23-fold when provided with middle ear parameters from the six odontocete species for which audiograms are available. The model reproduces the complete audiograms of these six species fairly well for frequencies up to about 100-120 kHz.