The Role of the Tympanal Membranes in the Tuning of Auditory Receptors in Tettigoniidae (Orthoptera: Ensifera)

Abstract

One of the fundamental problems in auditory research is the origin of the frequency selectivity of auditory receptors. Horridge (1960) demonstrated that the auditory interneurones in locusts were tuned to different sound frequencies, thus suggesting that the auditory receptors were frequency selective. Michelsen (1968, 1971) established that the auditory organ of locusts contained four groups of receptors with different characteristic sound frequencies and proposed that the frequency selectivity of the individual groups was dependent on the resonant properties of the tympanum. Investigation using the more sensitive of the two auditory receptors in the moth Prodenia eridania extended this proposal by suggesting that the tuning of this receptor depended on both resonant properties of the auditory organ (scoloparium) and filtering properties of the acoustic system (Adams, 1972), the most important factor in the latter being the tympanal membranes. In crickets and bushcrickets, however, the tympanal membranes vibrate in a simple mode for a wide band of sound frequencies (Paton, Capranica, Dragsten & Webb, 1977; Michelsen & Larsen, 1978), thus making it unlikely that the motion of the tympanum is responsible for the frequency selectivity (Zhantiev & Korsunovskaya, 1978; Esch, Huber & Wohlers, 1980) of the individual auditory receptors. Similarly, Ball & Hill (1978) demonstrated that auditory intemeurones of larval instar crickets are tuned even though they have no tympanal membranes. A recent study of the tonotopic organization of the auditory organ in bushcrickets (Oldfield, 1982) revealed that the auditory receptors remained tuned after the removal of one tympanum. This result, like those obtained from crickets, suggests that the resonant properties of the tympanal membranes are not responsible for the tuning of the individual auditory receptors. Conclusive evidence on this proposal, however, requires the removal of both tympanal membranes, thus eliminating the possibility that the remaining tympanum in the study by Oldfield (1982) determined the tuning of the auditory receptors.