Acoustic mate choice in insects has been extensively studied under laboratory conditions, using different behavioural paradigms. Ideally, however, mate choice designs should reflect natural conditions, including the physical properties of the transmission channel for the signal. Since little is known about the discrimination ability of females between male song variants under natural conditions, we performed phonotaxis experiments with female field crickets (Gryllus bimaculatus) outdoors, using two-choice decisions based on differences in carrier frequency, sound pressure level, and chirp rate. For all three song parameters, minimum differences necessary for a significant preference between two song models were considerably larger outdoors compared to laboratory conditions. A minimum amplitude difference of 5 dB was required for a significant choice in the field, compared to only 1–2 dB reported for lab-based experiments. Due to the tuned receiver system, differences in carrier frequency equal differences in perceived loudness, and the results on choice for differences in carrier frequency corroborate those in amplitude. Similarly, chirp rate differences of 50 chirps/min were required outdoors compared to only 20 chirps/min in the lab. For predictions about patterns of sexual selection, future studies need to consider the different outcomes of mate choice decisions in lab and field trials.
A modified tracheal system is the anatomical basis for a pressure difference receiver in field crickets, where sound has access to the inner and outer side of the tympanum of the ear in the forelegs. A thin septum in the midline of a connecting trachea coupling both ears is regarded to be important in producing frequency-dependent interaural intensity differences (IIDs) for sound localization. However, the fundamental role of the septum in directional hearing has recently been challenged by the finding that the localization ability is ensured even with a perforated septum, at least under controlled laboratory conditions. Here, we investigated the influence of the medial septum on phonotaxis of female Gryllus bimaculatus under natural conditions. Surprisingly, even with a perforated septum, females reliably tracked a male calling song in the field. Although reduced by 5.2 dB, IIDs still averaged at 7.9 dB and provided a reliable proximate basis for the observed behavioural performance of operated females in the field. In contrast, in the closely related species Gryllus campestris the same septum perforation caused a dramatic decline in IIDs over all frequencies tested. We discuss this discrepancy with respect to a difference in the phenotype of their tracheal systems.
SUMMARYSubtle random deviations from perfect symmetry in bilateral traits are suggested to signal reduced phenotypic and genetic quality of a sender, but little is known about the related receiver mechanisms for discriminating symmetrical from asymmetrical traits. Here, we investigated these mechanisms in behavioural and neurophysiological experiments in the Mediterranean field cricket, Gryllus bimaculatus. A downward frequency modulation at the end of each syllable in the calling song has been suggested to indicate morphological asymmetry in sound radiating structures between left and right forewings. Even under ideal laboratory conditions on a trackball system, female crickets only discriminated between songs of symmetrical and asymmetrical males in two-choice experiments at carrier frequencies of 4.4 kHz and a large modulation depth of 600 and 800 Hz. Under these conditions they preferred the pure-tone calling songs over the modulated (asymmetrical) alternative, whereas no preference was observed at carrier frequencies of 4.9 and 5.2 kHz. These preferences correlate well with the responses of a pair of identified auditory interneurons (AN1), known for their importance in female phonotaxis. The AN1 interneuron is tuned to an average frequency of 4.9 kHz, and the roll-off towards lower and higher frequencies determines the magnitude of responses to pure-tone and frequency-modulated calling songs. The difference in response magnitude between the two neurons appears to drive the decision of females towards the song alternatives. We discuss the relevance of song differences based on asymmetry in the morphology of song-producing structures under natural conditions.
The detection, identification and discrimination of sound signals in a large and noisy group of signalers are problems shared by many animals equipped with ears. While the signaling behavior of the sender may present several solutions, various properties of the sensory system in receivers may also reduce the amount of signal masking. We studied the effect of spatial release from masking, which refers to the fact that the spatial separation between the signaler and the masker can contribute to signal detection and discrimination. Except in a limited number of cases, the contribution of peripheral directionality or central nervous processing for spatial unmasking is not clear. We report the results of a study using a neurophysiological approach in two species of acoustic insects, whereby the activity of identified interneurons that either receive contralateral inhibitory input (crickets) or inhibit one another reciprocally in a bilateral pair (katydids) was examined. The analysis of the responses of a pair of omega neurons in katydids with reciprocal inhibition revealed that spatial separation of the masker from the signal facilitated signal detection by 19-20 dB with intact binaural hearing, but only by 2.5-7 dB in the monaural system, depending on the kind of analysis performed. The corresponding values for a behaviorally important interneuron of a field cricket (ascending neuron 1) were only 7.5 and 2.5 dB, respectively. We compare these values with those reported for hearing in vertebrates, and discuss the contribution of spatial release from masking to signal detection under real-world chorus conditions.
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