Divergence in mate recognition systems among closely related species is an important contributor to assortative mating and reproductive isolation. Here, we examine divergence in male song traits and female preference functions in three cricket species with songs consisting of long trills. The shape of female preference functions appears to be mostly conserved across species and follows the predictions from a recent model for song recognition. Multivariate preference profiles, combining the pulse and trill parameters, demonstrate selectivity for conspecific pulse rates and high trill duty cycles. The rules for integration across pulse and trill timescales were identical for all three species. Generally, we find greater divergence in male song traits than in associated female preferences. For pulse rate, we find a strong match between divergent male traits and female peak preferences. Preference functions for trill parameters and carrier frequency are similar between species and show less congruence between signal and preference. Differences among traits in the degree of trait-preference (mis)match may reflect the strength of preferences and the potential for linkage disequilibrium, selective constraints and alternative selective pressures, but appear unrelated to selection for mate recognition per se.

Ascending auditory interneurons of the cricket, Teleogryllus commodus (Walker), were investigated using simultaneous intracellular and extracellular recording in order to identify units which had previously been characterized only by extracellular recording. The morphology and physiology of the large adapting unit (LAU: Fig. 1) and of the small tonic unit (STU: Fig. 2) of Teleogryllus correspond well to those of the ascending neuron 2 (AN2) and the ascending neuron 1 (AN1) of Gryllus (Figs. 1, 2), respectively. A summary of the ascending auditory interneurons described by various authors in 5 species of crickets is presented in order to establish common identities. Physiological evidence for direct connections between auditory afferents and the ascending auditory interneurons AN1 (STU) and AN2 (LAU) is presented. Simultaneous intracellular recordings from receptors and interneurons in response to sound as well as the activity of auditory interneurons upon electrical stimulation of the tympanal nerve reveal short and constant latencies of receptor-evoked synaptic activity in AN1 (STU) and AN2 (LAU).

Insects exhibit an astonishing diversity in the design of their ears and the subsequent processing of information within their auditory pathways. The aim of this review is to summarize and compare the present concepts of auditory processing by relating behavioral performance to known neuronal mechanisms. We focus on three general aspects, that is frequency, directional, and temporal processing. The first part compares the capacity (in some insects high) for frequency analysis in the ear with the rather low specificity of tuning in interneurons by looking at Q10dB values and frequency dependent inhibition of interneurons. Since sharpening of frequency does not seem to be the prime task of a set of differently tuned receptors, alternative hypotheses are discussed. Moreover, the physiological correspondence between tonotopic projections of receptors and dendritic organization of interneurons is not in all cases strong. The second part is concerned with directional hearing and thus with the ability for angular resolution of insects. The present concepts, as derived from behavioral performances, for angular resolution versus lateralization and serial versus parallel processing of directional and pattern information can be traced to the thoracic level of neuronal processing. Contralateral inhibition, a mechanism for enhancing directional tuning, appears to be most effective in parallel pathways, whereas in serial processing it may have detrimental effects on pattern processing. The third part, after some considerations of signal analysis in the temporal domain, demonstrates that closely related species often use different combinations of temporal parameters in their recognition systems. On the thoracic level, analysis of temporal modulation functions and effects of inhibition on spiking patterns reveals relatively simple processing, whereas brain neurons may exhibit more complex properties.

Adaptation of the spike-frequency response to constant stimulation, as observed on various timescales in many neurons, reflects high-pass filter properties of a neuron's transfer function. Adaptation in general, however, is not sufficient to make a neuron's response independent of the mean intensity of a sensory stimulus, since low frequency components of the stimulus are still transmitted, although with reduced gain. We here show, based on an analytically tractable model, that the response of a neuron is intensity invariant, if the fully adapted steady-state spike-frequency response to constant stimuli is independent of stimulus intensity. Electrophysiological recordings from the AN1, a primary auditory interneuron of crickets, show that for intensities above 60 dB SPL (sound pressure level) the AN1 adapted with a time-constant of approximately 40 ms to a steady-state firing rate of approximately 100 Hz. Using identical random amplitude-modulation stimuli we verified that the AN1's spike-frequency response is indeed invariant to the stimulus' mean intensity above 60 dB SPL. The transfer function of the AN1 is a band pass, resulting from a high-pass filter (cutoff frequency at 4 Hz) due to adaptation and a low-pass filter (100 Hz) determined by the steady-state spike frequency. Thus, fast spike-frequency adaptation can generate intensity invariance already at the first level of neural processing.

Is discrimination of the envelope of an acoustic signal based on spectral or temporal computations? To investigate this question for the cricket Gryllus bimaculatus, pattern envelopes were constructed by the addition of several sine waves and modified by systematic phase changes. The phonotactic response of female crickets towards such sinusoidal but also rectangular pulse patterns was quantified on a locomotion compensator. Envelope patterns that exhibited a modulation frequency of 25 Hz as the dominant frequency were attractive and although changes of phase modified the temporal pattern, the values of attractiveness remained unaffected. Removal of the 25-Hz component reduced the phonotactic scores. Patterns in which other frequency components exhibited a larger amplitude than the 25-Hz component were less attractive. However, the combination of an unattractive pulse period with the attractive modulation frequency of 25 Hz in a pattern revealed that such stimuli were unattractive despite the presence of the 25-Hz component. A comparison of the attractiveness of all patterns revealed that female crickets evaluated the duration of pulse period over a wide range of duty cycles. The combined evidence showed that pattern envelopes were processed in the time- and not in the spectral domain.