Directional hearing in the grassfrog (Rana temporaria L.). II. Acoustics and modelling of the auditory periphery

In an earlier paper (Vlaming et al., 1984) we reported on optical measurements (laser-doppler interferometry) of the vibrations characteristics of the grassfrog's tympanic membrane. In the present paper these measurements were extended to include acoustic measurements concerning the functional role of the mouth cavity in frog hearing. Based on these measurements a model of the frog's acoustic periphery, consisting of three coupled linear oscillators with three entrance ports for sound, was developed and analyzed mathematically to give the various relevant transfer functions. The model is characterized by six parameters, all of which could be estimated from the available experimental data. For frequencies up to some 1500 Hz the model adequately describes the experimental data, both our own and earlier, seemingly conflicting data in the literature. For higher frequencies deviations occur, possibly due to nonuniform vibrations of the membranes. The model was used to evaluate the monaural directional sensitivity of the frog under free-field stimulation. Essentially it behaves as a combined pressure-gradient receiver, with highly frequency-dependent directional sensitivity. Directional sensitivity of the tympanic membrane could be modulated drastically by changing the resonance properties of the mouth cavity, without affecting the intrinsic membrane properties. This, theoretically, allows the frog to manipulate its direction sensitivity by actively tuning the volume of its mouth cavity. In order to account for discrepancies with known properties of low-frequency auditory nerve fibers an additional, extra-tympanic channel was included into the model. The extended model, the second-channel possibly involving the opercularis complex, provides a good quantitative fit to the available data on tympanic membrane movement as well as auditory nerve activity. Finally, the model enables to simulate a (moving) sound source in space, while stimulating the frog via closed couplers.