Transmission and reception of high-frequency sound in the natural environment of bushcrickets (Tettigonia viridissirna L.) was studied using the activity of an identified neuron in the insect's auditory pathway as a "biological microphone". Different positions of the receiver within the habitat were simulated by systematic variation of the distance from a loudspeaker and the height above the ground. Attenuation and filtering properties of the habitat were investigated with pure-tone frequencies between 5 and 40 kHz. Sound attenuation in excess of the attenuation due to geometrical spreading alone increased with increasing frequency, distance between sender and receiver, and decreasing height within the vegetation (Figs. 2-4). The data also confirm the existence of two kinds of excess attenuation. The amount of amplitude fluctuations in the sound signals was investigated by analysing the variability of the neuronal responses at a given receiver position. Variability increased with decreasing bandwidth of a noise signal at some distance from the loadspeaker. The variability in the responses to pure tones increased with both increasing frequency and distance from the source (Fig. 7). In the selected habitat, the temporal pattern of the natural calling song of male T. viridissima was very reliably reflected in the activity of the recorded neuron up to a distance of 30 m at the top of the vegetation, and 15-20 m near ground level (Figs. 5, 8). The maximum hearing distance in response to the calling song was about 40 m. Environmental constraints on long-range acoustic communication in the habitat are discussed in relation to possible adaptations of both the signal structure and the behavior of the insects.
The anterior intermediate sensory neuropile (aISN) is a prominent neuropile in the ventral nerve cord of locusts and bushcrickets. Previous studies have shown that it receives its main sensory input from auditory receptors. In this paper we examine the structural and physiological relationship between tympanal receptor terminations and the dendrites of sound-sensitive interneurones in the homologous neuropile of locusts and bushcrickets. Each individual receptor fibre of the bushcricket terminates in a somewhat different target area of the neuropile. The ordering is with respect to the characteristic frequency of the fibres (tonotopic) in the anterior-posterior and dorsoventral axis. In the locust, representatives of the four tympanal receptor groups branch in different areas of the aISN. Most of the dorsal neuropilar region, and the anterior ventral region, do not receive input from tympanal receptors. The dendrites of identified sound-sensitive interneurones were examined in the context of this afferent projection. Local interneurones as well as intersegmental interneurones in bushcrickets have dendritic branches in the whole aISN or part of it and thus overlap with at least some receptors. By recording intracellularly from their main neurites, short-latency synaptic potentials were found in response to receptor spikes indicating monosynaptic input. The tuning of these neurones could be predicted by their dendritic morphology. In contrast, in the locust only local and bisegmental neurones are monosynaptically connected with tympanal receptors, but not the studied intersegmental neurones. This is consistent with the finding that most or all branches of intersegmental neurones lie in the dorsal area of neuropile where no receptors terminate. Anatomical and physiological evidence is presented for identified local neurones providing the excitatory and inhibitory synaptic input for such intersegmental neurones. The difference in the basic wiring diagram in the homologous neuropile of the two orthopteran groups is discussed with respect to the possible different roles that sound plays in their behaviour.
Males of the bushcricket Mecopoda elongata synchronise or alternate their chirps with their neighbours in an aggregation. Since synchrony is imperfect, leader and follower chirps are established in song interactions; females prefer leader chirps in phonotactic trials. Using playback experiments and simulations of song oscillator interactions, we investigate the mechanisms that result in synchrony and alternation, and the probability for the leader role in synchrony. A major predictor for the leader role of a male is its intrinsic chirp period, which varies in a population from 1.6 to 2.3 s. Faster singing males establish the leader role more often than males with longer chirp periods. The phase-response curve (PRC) of the song oscillators differs to other rhythmically calling or flashing insects, in that only the disturbed cycle is influenced in duration by a stimulus. This results in sustained leader or follower chirps of one male, when the intrinsic chirp periods of two males differ by 150 ms or more. By contrast, the individual shape of the male's PRC has only little influence on the outcome of chirp interactions. The consequences of these findings for the evolution of synchrony in this species are discussed.
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