A model for directional and distance hearing in swimbladder-bearing fish based on the displacement orbits of the hair cells

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Abstract
It is known that teleosts, without Weberian ossicles but with a swimbladder, can detect the direction of and, under appropriate conditions, the distance to a sound source [e.g., Schuijf and Hawkins, Nature 302, 143-144 (1983)]. It is hypothesized here that the underlying mechanism is the analysis of the parameters of the elliptical movement of the hair cells with respect to the otoliths. This movement results from the displacement wave impinging directly upon the labyrinth and the response displacement wave reradiated by the swimbladder. For a given swimbladder geometry, given the positions of the maculae of both laryrinths with respect to the swimbladder and the damping of the swimbladder, the displacement orbits of the maculae can be calculated [de Munck and Schellart, J. Acoust. Soc. Am. 81, 556-560 (1987)]. These calculations were made for the cod and the trout with the frequency, direction, and distance between the fish and the sound source as parameters with the source within the same horizontal plane as the fish. The orbit model predicts that the utriculus has the most strategic position to detect direction and distance of such a sound source. Moreover, the model predicts that this could basically be done monaurally. A hypothesis is proposed to describe how the utricular system analyzes the orbit parameters. The model is evaluated in relation to the results of behavioral experiments described in the literature.