Directional characteristics of the auditory system of cicadas: is the sound producing tymbal an integral part of directional hearing?

Abstract.  Directional hearing is investigated in males of two species of cicadas, Tympanistalna gastrica (Stål) and Tettigetta josei Boulard, that are similar in size but show different calling song spectra. The vibrational response of the ears is measured with laser vibrometry and compared with thresholds determined from auditory nerve recordings. The data are used to investigate to what extent the directional characteristic of the tympanal vibrations is encoded by the activity of auditory receptors. Laser measurements show complex vibrations of the tympanum, and reveal that directional differences are rather high (>15 dB) in characteristic but limited frequency ranges. At low frequencies, both species show a large directional difference at the same frequency (3–5 kHz) whereas, above 10 kHz, the directional differences correspond to the different resonant frequencies of the respective tymbals. Consequently, due to the mechanical resonance of the tymbal, the frequency range at which directional differences are high differs between the two species that otherwise show similar dimensions of the acoustic system. The directional differences observed in the tympanal vibrations are also observed in the auditory nerve activity. These recordings confirm that the biophysically determined directional differences are available within the nervous system for further processing. Despite considerable intra as well as interindividual variability, the ears of the cicadas investigated here exhibit profound directional characteristics, because the thresholds determined from recordings of the auditory nerve at 30° to the right and left of the longitudinal axis differ by more than 5 dB.

[1]  Meyer,et al.  How well are FREQUENCY SENSITIVITIES OF grasshopper ears tuned to species-specific song spectra? , 1996, The Journal of experimental biology.

[2]  Paulo J. Fonseca,et al.  Asymmetry of tymbal action and structure in a cicada: a possible role in the production of complex songs , 1998, The Journal of experimental biology.

[3]  A. Michelsen The physiology of the locust ear , 1971, Zeitschrift für vergleichende Physiologie.

[4]  A. V. Popov,et al.  Directionality of the tympanal vibrations in a cicada: a biophysical analysis , 1997, Journal of Comparative Physiology A.

[5]  Axel Michelsen,et al.  Biophysics of Sound Localization in Insects , 1998 .

[6]  Bennet-Clark Tymbal mechanics and the control of song frequency in the cicada Cyclochila australasiae , 1997, The Journal of experimental biology.

[7]  G. Wendler,et al.  The role of the medial septum in the acoustic trachea of the cricket Gryllus bimaculatus , 1993, Journal of Comparative Physiology A.

[8]  Michelsen,et al.  Directional sound processing and interaural sound transmission in a small and a large grasshopper , 1995, The Journal of experimental biology.

[9]  A. V. Popov,et al.  Sound radiation in a cicada: the role of different structures , 1994, Journal of Comparative Physiology A.

[10]  D. Young,et al.  The Organization of the Auditory Organ of the Bladder Cicada, Cystosoma Saundersii , 1981 .

[11]  John Thorson,et al.  Auditory behavior of the cricket , 2004, Journal of Comparative Physiology A.

[12]  P. Fonseca,et al.  Song discrimination by male cicadas Cicada barbara lusitanica (Homoptera, Cicadidae). , 2002, The Journal of experimental biology.

[13]  P. Fonseca,et al.  Directional hearing of a cicada: biophysical aspects , 1993, Journal of Comparative Physiology A.

[14]  Axel Michelsen,et al.  Tuned directionality in cricket ears , 1995, Nature.

[15]  David C. Young,et al.  Do Cicadas Radiate Sound through their Ear-Drums? , 1990 .

[16]  Franz Huber,et al.  Auditory behavior of the cricket , 2004, Journal of comparative physiology.

[17]  M. Sippel,et al.  Mechanics of the transduction of sound in the tympanal organ of adults and larvae of locusts , 1985, Journal of Comparative Physiology A.

[18]  Holderied,et al.  Directional hearing in grasshoppers: neurophysiological testing of a bioacoustic model , 1999, The Journal of experimental biology.

[19]  H. Kleindienst,et al.  The role of the medial septum in the acoustic trachea of the cricket Gryllus bimaculatus , 1994, Journal of Comparative Physiology A.

[20]  A. Stumpner,et al.  Morphological and physiological differences of the auditory system in three related bushcrickets (Orthoptera: Phaneropteridae, Poecilimon) , 1992 .

[21]  A. Michelsen Frequency Discrimination Based upon Resonances in the Tympanum , 1971 .

[22]  K. G. Hill,et al.  Structure and function of the auditory system of the cicada,Cystosoma saundersii , 2004, Journal of comparative physiology.

[23]  K. Rohrseitz,et al.  A new biophysical method to determine the gain of the acoustic trachea in bushcrickets , 1994, Journal of Comparative Physiology A.

[24]  A. V. Popov,et al.  Physics of directional hearing in the cricket Gryllus bimaculatus , 1994, Journal of Comparative Physiology A.