Acoustic communication in a duetting grasshopper: receiver response variability, male strategies and signal design
Section snippets
Study animals
We collected adult C. biguttulus males and last-instar female larvae in the field during July–October 1996–2000 from grassy areas around Erlangen in Germany and from the Bavarian and Austrian Alps. Females were collected as larvae to ensure virginity. Males and females were housed in separate cages () and fed on fresh grass (Dactylis glomerata). The animals were on a 14:10 h light:dark cycle and maintained at a temperature of 25–30°C. Cages were monitored every day and each freshly
Methods
To examine whether females prefer noisy pauses to clear ones in natural male songs, we tested the responses of females to eight pairs of phrases (from eight different males) in the presence or absence of noise (clicks) in the pauses. We played back natural phrases either unedited (with noisy pauses) or with clear (silent) pauses. Pauses in eight phrases of natural male song were defined with the help of the simultaneous optoelectronic recordings of the leg movements and the change in click rate
Experiment 3: synthetic phrases
We know from previous work (Balakrishnan et al. 2001) that syllable–pause discrimination by C. biguttulus females occurs on the basis of the independent determination of the relative sound levels of syllable onset and offset. In addition, the shape of the syllable (which is essentially the modulation of sound intensity over the course of the syllable) may contribute to the perception of the level of noise in the pauses. Male C. biguttulus songs typically contain syllables that are enhanced in
Experiment 4: enhanced onsets and noisy pauses
The results of the analysis of female preferences suggest that males would be selected to enhance their syllable onset accentuations: this could be achieved either by enhancing the syllable onsets or by lowering the amount of noise in the pauses. If noisy pauses do not confer any advantage on males, it is surprising that they are retained, since lowering the noise in the pauses could enhance the perceived intensity level of the syllable onset (Balakrishnan et al. 2001). This suggests that noisy
Discussion
The relative attractiveness of male C. biguttulus songs with clear or noisy pauses depended both on various other characteristics of the song and on the individual female. There was high variation between females in their preferences for clear or noisy pauses, as well as for strong or weak syllable onset accentuations. Individual response profiles were, however, consistent. The addition of noise in silent syllable pauses of synthetic model phrases increased the range of pause durations accepted
Acknowledgements
This article is dedicated to Professor Norbert Elsner who over many years insisted that we investigate ‘the paradox of noisy pauses.’ We are much indebted to Professor N. V. Joshi, who spent many hours exploring different methods of measuring the consistency of female response profiles. We gratefully acknowledge Maria Bauer for help in carrying out some experiments and her skilful preparation of the figures. We also acknowledge Johannes Schul for writing the program to delete clicks from
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- 1
This is one of the last papers that Dagmar von Helversen was able to complete before she passed away on 20 July 2003.
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R. Balakrishnan is at the Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560012, India.