Bennet-Clark H C
Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.
J Exp Biol. 2003 May;206(Pt 9):1479-96. doi: 10.1242/jeb.00281.
The anatomy and mechanics of the fore-wings of the Australian cricket Teleogryllus oceanicus were examined to study how resonances of the wings were excited, to model the interactions between the two wings during sound production, to account for the frequency changes that occur within the pulses and to determine the variation in sound amplitude during the pulses. Sound is produced after raising the wings by closing the right wing over the left; the plectrum of the left wing engages and releases teeth on the file on the underside of the right wing. The mean number of teeth on the right file is 252; the teeth are more closely spaced in the posterior part of the file, which is engaged at the start of the song pulses. The anterior part of the file is separated from the base of the harp by a short flexible region. The dorsal field of the wing, in which the harp is situated, is largely mechanically isolated from the driving veins of the lateral field, except for a cross vein at the apex of the harp. The harps of the two wings did not differ significantly in area but the plectrum of the left wing was significantly longer and wider than that of the right wing. The posterior edge of the plectrum has a radius of approximately 0.5 micro m, which allows it to engage the 20 micro m-tall teeth of the file. The plectrum is separated from the wing by a 0.5 micro m-thick crescent that allows it to twist lengthways and thus disengage the file teeth. The sigmoid shape of the file allows the plectrum to engage teeth over most of the length of the file. The calling song of T. oceanicus consists of a chirp of four similar pulses followed by a trill of pairs of pulses. The dominant frequency of all pulses is approximately 4.8 kHz but cycle-by-cycle analysis suggests that the different types of pulse are produced by wing-closing movements through different arcs. Free resonances of the left wing occurred at 4.56 kHz [quality factor (Q)=25.1] and of the right wing at 4.21 kHz (Q=23.9). Driven by loud sound, maximum vibration of the harp was seen at approximately 4.5 kHz; at lower sound levels, the vibration was confined to the cross-veins of the harp that extend distally from the file. Resonances of the left wing driven by vibration of the same wing, either at the plectrum or on the anal area, occurred at similar frequencies to those of the songs and had similar Qs but were approximately anti-phase, demonstrating that movement of the plectrum (e.g. by the file teeth) causes an opposite movement of the harp. When the right wing was driven directly on the file, the resonant frequency was 5.88 kHz but, when driven on the file via a length of the left file and the left plectrum, it was 4.83 kHz. The amplitude of the vibration increased from the posterior end of the file to the middle then fell towards the anterior end of the file. Pushing a left plectrum across the middle of the right file produced trains of damped sound pulses at 4.82 kHz (Q=23.4). Clicks excited from the anterior end of the file had lower frequencies. The resonances excited from both the left wing via its plectrum and from the right wing when driven via the left plectrum were similar in frequency to that of the song. The resonance of the dorsal field persisted after ablation of the harp but the mean resonant frequency increased 1.12-fold with a similar Q to the intact wing. Droplets of water on the distal end of the harp or proximal part of the dorsal field raised the resonant frequency. The resonant frequency was lowered by the addition of weights to the harp or the file; the factor of the decrease suggested that the mass of the resonant system was approximately 1.4 mg, which accords with the mass of the harp plus file plus anal area of the wing (left wing, 1.27 mg; right wing, 1.15 mg) but is far heavier than the harp (0.22 mg). An earlier suggestion that the harp is the resonator is not supported; instead, it is proposed that the major elastic component of the resonant system is the file plus 1st anal vein and that the mass component is the combined mass of the file, anal area and harp.
对澳大利亚蟋蟀Teleogryllus oceanicus前翅的解剖结构和力学特性进行了研究,以探讨翅膀共振是如何激发的,模拟发声过程中两个翅膀之间的相互作用,解释脉冲内发生的频率变化,并确定脉冲期间声音幅度的变化。通过将右翼在左翼上方闭合来举起翅膀后产生声音;左翼的拨弦器与右翼下侧锉上的齿啮合并松开。右翼锉上的平均齿数为252个;锉的后部齿间距更近,在歌曲脉冲开始时与之啮合。锉的前部通过一个短的柔性区域与竖琴基部隔开。翅膀的背侧区域(竖琴所在之处)在很大程度上与外侧区域的驱动脉机械隔离,除了竖琴顶端的一条横脉。两个翅膀的竖琴面积没有显著差异,但左翼的拨弦器明显比右翼的更长更宽。拨弦器的后缘半径约为0.5微米,使其能够与锉上20微米高的齿啮合。拨弦器通过一个0.5微米厚的月牙形结构与翅膀分离,使其能够纵向扭转从而松开锉齿。锉的S形形状使拨弦器能够在锉的大部分长度上与齿啮合。Teleogryllus oceanicus的求偶鸣叫由四个相似脉冲的啁啾声接着一对对脉冲的颤音组成。所有脉冲的主导频率约为4.8千赫兹,但逐周期分析表明,不同类型的脉冲是由翅膀通过不同弧度的闭合运动产生的。左翼的自由共振发生在4.56千赫兹[品质因数(Q)=25.1],右翼的自由共振发生在4.21千赫兹(Q=23.9)。在大声驱动下,竖琴的最大振动出现在约4.5千赫兹处;在较低声级下,振动局限于从锉向远端延伸的竖琴横脉。由同一翅膀的振动驱动,无论是在拨弦器处还是在臀区,左翼产生的共振频率与鸣叫的频率相似,且品质因数相近,但相位大致相反,这表明拨弦器的运动(如被锉齿推动)会导致竖琴产生相反的运动。当直接在锉上驱动右翼时,共振频率为5.88千赫兹,但当通过一段左翼锉和左翼拨弦器在锉上驱动时,共振频率为4.83千赫兹。振动幅度从锉的后端向中间增加,然后向锉的前端下降。将一个左翼拨弦器推过右翼锉的中部会产生一系列频率为4.82千赫兹(Q=23.4)的衰减声脉冲。从锉的前端激发的咔嗒声频率较低。通过左翼拨弦器激发的左翼共振和通过左翼拨弦器驱动时右翼的共振在频率上与鸣叫相似。在切除竖琴后,背侧区域的共振仍然存在,但平均共振频率增加了1.12倍,品质因数与完整翅膀相似。竖琴远端或背侧区域近端的水滴会提高共振频率。通过在竖琴或锉上添加重物会降低共振频率;降低系数表明共振系统的质量约为1.4毫克,这与竖琴加锉加翅膀臀区的质量相符(左翼,1.27毫克;右翼,1.15毫克),但比竖琴(0.22毫克)重得多。先前关于竖琴是共鸣器的观点未得到支持;相反,有人提出共振系统的主要弹性成分是锉加第一臀脉且质量成分是锉、臀区和竖琴的组合质量。