03493nas a2200133 4500008004100000245007000041210006500111300001200176490000800188520307700196100002503273700001303298856004803311 1998 eng d00aSound radiation by the bladder cicada Cystosoma saundersii0 aSound radiation by the bladder cicada iCystosoma saundersiii a701-7150 v2013 a

Male Cystosoma saundersii have a distended thin-walled abdomen which is driven by the paired tymbals during sound production. The insect extends the abdomen from a rest length of 32-34 mm to a length of 39-42 mm while singing. This is accomplished through specialised apodemes at the anterior ends of abdominal segments 4-7, which cause each of these intersegmental membranes to unfold by approximately 2 mm. 

The calling song frequency is approximately 850 Hz. The song pulses have a bimodal envelope and a duration of approximately 25 ms; they are produced by the asynchronous but overlapping action of the paired tymbals. The quality factor Q of the decay of the song pulses is approximately 17. 

The abdomen was driven experimentally by an internal sound source attached to a hole in the front of the abdomen. This allowed the sound-radiating regions to be mapped. The loudest sound-radiating areas are on both sides of tergites 3-5, approximately 10 mm from the ventral surface. A subsidiary sound-radiating region is found mid-ventrally on sternites 4-6. Sound is radiated in the same phase from all these regions. As the abdomen was extended experimentally from its resting length to its maximum length, the amplitude of the radiated sound doubled and the Q of the resonance increased from 4 to 9. This resonance and effect are similar at both tergite 4 and sternite 5. 

Increasing the effective volume of the abdominal air sac reduced its resonant frequency. The resonant frequency was proportional to 1/(check)(total volume), suggesting that the air sac volume was the major compliant element in the resonant system. Increasing the mass of tergite 4 and sternites 4-6 also reduced the resonant frequency of the abdomen. By extrapolation, it was shown that the effective mass of tergites 3-5 was between 13 and 30 mg and that the resonant frequency was proportional to 1/(check)(total mass), suggesting that the masses of the tergal sound-radiating areas were major elements in the resonant system.

The tymbal ribs buckle in sequence from posterior (rib 1) to anterior, producing a series of sound pulses. The frequency of the pulse decreases with the buckling of successive ribs: rib 1 produces approximately 1050 Hz, rib 2 approximately 870 Hz and rib 3 approximately 830 Hz. The sound pulse produced as the tymbal buckles outwards is between 1.6 and 1.9 kHz. Simultaneous recordings from close to the tymbal and from tergite 4 suggest that the song pulse is initiated by the pulses produced by ribs 2 and 3 of the leading tymbal and sustained by the pulses from ribs 2 and 3 of the second tymbal.

An earlier model suggested that the reactive elements of the abdominal resonance were the compliance of the abdominal air sac volume and the mass of the abdomen undergoing lengthwise telescoping. The present work confirms these suggestions for the role of the air sac but ascribes the mass element to the in-out vibrations of the lateral regions of tergites 3-5 and the central part of sternites 4-6.

1 aBennet-Clark, H., C.1 aYoung, D uhttp://jeb.biologists.org/content/201/5/70102759nas a2200133 4500008004100000245006000041210005800101300001200159490000800171520236000179100002502539700001302564856004802577 1992 eng d00aA MODEL OF THE MECHANISM OF SOUND PRODUCTION IN CICADAS0 aMODEL OF THE MECHANISM OF SOUND PRODUCTION IN CICADAS a123-1530 v1733 a

1. Dried cicada bodies of the species Cyclochila australasiae and model cicadas made from a miniature earphone driving a plastic cavity were used to study the acoustics of sound production in male cicadas.

2. A model cicada with shape and dimensions similar to those of the abdomen of a male C. australasiae resonates at the natural song frequency of the species (4.3 kHz). The abdominal air sac of C. australasiae also resonates at frequencies close to the natural song frequency when excited by external sounds. In an atmosphere of chlorofluorocarbon (CFC) gas, the resonant frequency is lowered in keeping with the decrease in velocity of sound in the CFC gas.

3. At the model's resonant frequency, the driving earphone dissipates more electrical power with the cavity attached than without the cavity. The cavity of the model cicada acts as a narrow-band acoustic acceptance filter, tuned to the natural song frequency.

4. When the miniature earphone emits brief clicks, mimicking those produced by the natural tymbal mechanism, the model cicada produces sound pulses that vary in duration and shape according to the number and timing of the clicks. A coherent train of two or three resonant clicks results in a long slowly-decaying sound pulse similar to that in the natural song.

5. The natural song frequency can be predicted from the dimensions of the abdominal cavity and the tympana in C. australasiae using a simple equation for the resonant frequency of a Helmholtz resonator. This equation also predicts the song frequency of Macrotristria angularis and Magicicada cassini, but it fails with the low-frequency song of Magicicada septendecim. This discrepancy can be accounted for by the unusually thick tympana of M. septendecim, which tend to lower the resonant frequency of the system.

6. We conclude that the abdomen of male cicadas forms a Helmholtz resonator, the components of which are the large air sac as the cavity and the tympana as the neck of the resonator. We suggest that cicada sound production depends on the coupling of two resonators, that of the tymbal and that of the abdominal air sac, from which sound is radiated through the tympana. The coupled resonator system would produce the long sound pulses required for stimulating a sensitive sharply tuned auditory organ.

1 aBennet-Clark, H., C.1 aYoung, D uhttp://jeb.biologists.org/content/173/1/12303608nas a2200157 4500008004100000022001400041245005400055210005000109260000900159300001200168490000800180520318300188100001303371700002503384856004103409 1995 eng d a1477-914500aThe role of the tymbal in cicada sound production0 arole of the tymbal in cicada sound production c1995 a1001-200 v1983 a

1. The tymbal of Cyclochila australasiae consists of a biconvex membrane bearing alternating long and short ribs anteriorly and an irregularly shaped tymbal plate posteriorly. These sclerotised regions are coupled together by the surrounding highly flexible cuticle, which contains resilin. Dorsally, there is a thick pad of resilin, which functions as a spring, returning the tymbal to the out position and maintaining the stress on the long ribs. 2. Contraction of the tymbal muscle causes the tymbal plate to swing inwards, acting as a lever so that the surface of the tymbal moves through more than twice the distance of muscle shortening. This produces an inward movement and twisting of the dorsal ends of the long ribs, which then buckle in sequence, with each rib undergoing a sudden deformation from a convex to a V-shaped profile. Buckling takes place at the rib's weakest point, which is the narrow, highly sclerotised mid-region. 3. Inward buckling of the tymbal generates a loud click with a dominant frequency around 4 kHz. Resonances close to 4 kHz can be demonstrated in a buckled-in tymbal when driven by internal sound or by vibration at the tymbal plate. These resonances occur in sealed cicadas and those in which the abdominal air sac has been opened at both its anterior and posterior ends, which shows that the resonances are not due to the air sac; the tymbal itself is a resonant system. The maximum amplitude of tymbal vibration occurs at the V-shaped dimples in the centre of the long ribs. 4. When the tymbal plus abdominal air sac system is driven by vibration at the tymbal plate, the Q3dB of the sound radiated through the tympana is about 12.5, which is approximately the sum of those of the tymbal (Q=9.3) and of the air sac (Q=3.4) resonators. When the tymbal is not loaded by the air sac, i.e. in the sealed cicada and open cicada preparations, the Q3dB of its resonance is higher, between 13 and 20. 5. The click produced as the tymbal pops out is over 20 dB quieter than the in-click and has a dominant frequency around 6 kHz. When driven in the resting position, resonances are found close to 6 kHz but there is only a weak general vibration of the ribs and tymbal plate. When the tymbal is pushed in gradually, the resonant frequency changes from about 5.5 kHz to about 4.3 kHz as the tymbal buckles inwards. The left and right tymbals of the same insect may differ slightly in their acoustic properties. 6. As the tymbal buckles inwards, it displaces a volume of approximately 6 µl into the abdominal air sac volume of about 2 ml. The resulting sound pressure inside the air sac attains peak values of 155­159 dB SPL; the root mean square values are 141­144 dB SPL. The mean peak value just outside the tympana is 148.5 dB SPL. 7. Overall, the present work supports and extends our earlier model of cicada sound production: the tymbal click provides a coherent resonant source that drives the abdominal resonator, from which sound is radiated via the tympana. At the same time, the system provides the pressure transformation between muscle power and sound power that is desirable for efficient sound radiation.

1 aYoung, D1 aBennet-Clark, H., C. uhttps://bio.acousti.ca/uk/node/17208