Thesis (M.Sc.). / Written for the Dept. of Biology. Title from title page of PDF (viewed 2008/12/07). Includes bibliographical references.

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We investigated the relationship between rhythms occurring at different time-scales, namely circadian and ultradian rhythms. Furthermore, we examined the potential role of the period (per) gene, a known clock gene initially isolated in Drosophila, for regulating these rhythms in the wild-type cricket, Teleogryllus oceanicus . Here we report a positive correlation between circadian and ultradian rhythms in crickets. Thus, crickets with a slow endogenous circadian rhythm also have a slow ultradian rhythm, and conversely, animals with a fast endogenous circadian rhythm have a fast ultradian rhythm. In addition to this correlation, we observed PER-like immunoreactivity in the regions of the cricket nervous system involved in regulating these rhythms, namely the brain, in particular the optic lobes, and the meso- and metathoracic ganglia. We thus conclude that different time-scale rhythms may be regulated by a common setting element, a role potentially played by the per gene.

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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.

Darwinian sexual selection predicts that males with the most extravagant secondary sexual traits suffer elevated mortality. Although correlative evidence has generally not borne this idea out, recent research, including a field cricket study, showed that investment in sexually selected traits is costly to survival. I investigated male survival, ornamentation (song) and mating success in a North American grylline, Gryllus pennsylvanicus, to test the generality of previous work and highlight the importance of ecology differences to resource allocation. As the calling songs of older male G. pennsylvanicus are highly attractive to females, in Chapter 2 I tested whether male age correlated with calling song and found a weak but statistically significant correlation, thus leaving open the possibility that choosy females use an age-based indicator mechanism. In Chapter 3, I tested the condition dependence of male survival and calling effort. In contrast to previous work, I found that high condition males both called more and lived longer than low condition males, although there was no trade-off between survival and calling effort. The substantial condition dependence of calling effort suggests that calling effort is under strong directional selection. In Chapter 4 I tested whether female mating preferences resulted in strong selection on male calling effort. I also tested for the condition dependence of female mating preferences. I found that female choosiness was condition-dependent, but the rank of preferred male songs (preference function) was not. Both low and high condition females preferred high calling effort over low calling effort song. In Chapter 5 I tested for evidence of nonlinear selection on male survival that might explain the nonlinear pattern of male investment in survival seen in Chapter 3 (i.e. male survival leveled-off with increasing condition). I found that socially experienced females, but not virgin and naive females, exerted linear selection on male age. I discuss these and the other results of my thesis in the context of previous work on field crickets and condition-dependent ornamentation. Finally, Appendix A reports results that confirm ancient Chinese cultural knowledge that large headed male crickets are more successful in male-male combat.