The Prediction Of Field Cricket Phonotaxis In Complex Acoustic Environments
Animals detect, recognize and localize relevant objects in noisy, multi-source environments. Female crickets locate potential mates in choruses of simultaneously calling males using acoustic signals, a behaviour termed phonotaxis. The mechanisms underlying cricket phonotaxis are now understood across multiple levels: biophysical, neurobiological and behavioural. Phonotaxis has, however, rarely been tested in the complex real-world acoustic environments and no attempts have been made to predict acoustic orientation behaviour in these conditions despite our extensive understanding of its underlying mechanisms. In this thesis, I first characterized the acoustic environments faced by female crickets of the species Plebeiogryllus guttiventris in the field. Phonotaxis behaviour of females was then characterized under laboratory conditions using two sound sources. The data obtained were used to develop a simulation that predicted this behaviour. The predictions of the simulation were then tested against the phonotaxis behaviour of females in realistic, multi-source conditions in the field. My field studies of male behaviour showed that males of this species produced complex and variable songs in choruses where multiple males called simultaneously. The acoustic ranges of males in these choruses overlapped extensively and females performing phonotaxis in such choruses would hear multiple males simultaneously. The acoustic interactions of simultaneously calling males were also characterized for their timing relationships with each other and the changes they made to the temporal patterns of their songs. Males did not either synchronise or alternate their chirps, however they made changes to the temporal patterns of song in a way that is likely to make them more attractive to females. I then characterized the closed-loop walking phonotaxis behaviour of P. guttiventris females in the presence of two active sound sources playing conspecific song. Both the baseline and relative SPLs of the two speakers were systematically varied and female phonotactic paths were obtained. Females were found to preferentially approach louder songs. Several aspects of this behaviour were characterized, in particular orientation ability and motor behaviour under varied conditions of stimulus intensity. A stochastic simulation of closed-loop walking phonotaxis behaviour was developed using both current understanding of field cricket physiology and my data on closed-loop walking phonotaxis. The simulation was demonstrated to both qualitatively and quantitatively recapture female behaviour. It was also able to qualitatively recapture female behaviour in two previously published classical experiments in which the hearing of female crickets was disrupted. Female phonotaxis was then tested under real-world multi-source conditions. The behaviour of real females was compared to the predictions of the simulation. The simulation was found to recapture both female preference and phonotactic path forms at the population level. To my knowledge, this is the first study to both examine and successfully predict phonotaxis behaviour in complex real-world acoustic conditions.
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