Ears adapted for the detection of motion, or how echolocating bats have exploited the capacities of the mammalian auditory system

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Abstract
Bats use the rich food resources of the night by specializing in audition. They emit short echolocation sounds and listen to the echoes returning from potential prey. The bat’s auditory systemanalyzes spectral and temporal parameters of echoes for detecting, locating, and identifying a target. Different bat species have solved the problem of acoustic target detection and pattern recognition even in clustered situations by focusing on certain acoustical features of a target. The specialized motion detection by horseshoe bats, for instance, analyzes small echofrequency shifts modulated onto a long constant frequency echolocation signal. These frequency modulations are Doppler shifts within echoes returning from wing beating insects. For detecting modulations as small as 10 Hz or 0.01%, horseshoe bats have in the cochlea an extremely narrow filter (Q?500) matched to the carrier frequency (i.e., echolocation sound) of 83 kHz. The filter is realized by structural differentiations of the basilar membrane and the filter frequencies are represented on the basilar membrane in an expanded fashion. We have called this specialized patch of the basilar membrane an ’’acoustical fovea.’’ The ’’foveal frequencies’’ are largely overrepresented in the tonopical arrangement of the ascending auditory pathway. The bats have developed a feedback system which lowers the emitted frequency during flight in such a way that the Doppler shifted echofrequency is kept precisely at a fixed reference frequency of the fovea. This feedback system and other neuronal data disclose an intricate coupling of the auditory and vocalizing system. The evolution of echolocation in bats has driven the analyzing capacities of audition in both frequency and time domain close to theoretical limits. Investigations of such specialized systems give fascinating insights into capacities and possible general principles of auditory information processing.