Cranking up the heat: relationships between energetically costly song features and the increase in thorax temperature in male crickets and katydids.

Sexual displays of acoustically signalling insects are used in the context of mate attraction and mate choice. While energetic investment in sound production can increase the reproductive success of the sender, this entails metabolic costs. Resource allocation to sexually selected, reproductive traits can trade off against allocation to naturally selected traits (e.g. growth, immunity) when individuals' energy budgets are limited. Estimating the magnitude of the costs invested in acoustic signalling is necessary to understand this trade-off and its influence on fitness and life history. To compare the costs associated with acoustic signalling for two ensiferan species, we simultaneously took respiratory measurements to record the rate of CO2 production and used infrared thermography to measure the increase in thorax temperature. Furthermore, to identify what combinations of acoustic parameters were energetically costly for the sender, we recorded the calling songs of 22 different cricket and katydid species for a comparative analysis and measured their thorax temperature while they sang. Acoustic signalling was energetically costly for Mecopoda sp. and Anurogryllus muticus, requiring a 12- and 16-fold increase over resting levels in the CO2 production rate. Moreover, calling increased thorax temperature, on average by 7.6 and 5.8°C, respectively. We found that the song intensity and effective calling rate, not simply the chirp/trill duty cycle or the pulse rate alone, were good predictors for the thorax temperature increase in males.

Noninvasive diagnosis of seed viability using infrared thermography.

Recent advances in the noninvasive analyses of plant metabolism include stress imaging techniques, mainly developed for vegetative tissues. We explored if infrared thermography can be used to predict whether a quiescent seed will germinate or die upon water uptake. Thermal profiles of viable, aged, and dead Pisum sativum seeds were recorded, and image analysis of 22,000 images per individual seed showed that infrared thermography can detect imbibition- and germination-associated biophysical and biochemical changes. These "thermal fingerprints" vary with viability in this species and in Triticum aestivum and Brassica napus seeds. Thermogenesis of the small individual B. napus seeds was at the limit of the technology. We developed a computer model of "virtual pea seeds," that uses Monte Carlo simulation, based on the heat production of major seed storage compounds to unravel physico-chemical processes of thermogenesis. The simulation suggests that the cooling that dominates the early thermal profiles results from the dissolution of low molecular-weight carbohydrates. Moreover, the kinetics of the production of such "cooling" compounds over the following 100 h is dependent on seed viability. We also developed a deterministic tool that predicts in the first 3 hours of water uptake, when seeds can be redried and stored again, whether or not a pea seed will germinate. We believe that the early separation of individual, ungerminated seeds (live, aged, or dead) before destructive germination assessment creates unique opportunities for integrative studies on cell death, differentiation, and development.

A Simple Denoising Algorithm for Real-World Noisy Camera Images.

The noise statistics of real-world camera images are challenging for any denoising algorithm. Here, I describe a modified version of a bionic algorithm that improves the quality of real-word noisy camera images from a publicly available image dataset. In the first step, an adaptive local averaging filter was executed for each pixel to remove moderate sensor noise while preserving fine image details and object contours. In the second step, image sharpness was enhanced by means of an unsharp mask filter to generate output images that are close to ground-truth images (multiple averages of static camera images). The performance of this denoising algorithm was compared with five popular denoising methods: bm3d, wavelet, non-local means (NL-means), total variation (TV) denoising and bilateral filter. Results show that the two-step filter had a performance that was similar to NL-means and TV filtering. Bm3d had the best denoising performance but sometimes led to blurry images. This novel two-step filter only depends on a single parameter that can be obtained from global image statistics. To reduce computation time, denoising was restricted to the Y channel of YUV-transformed images and four image segments were simultaneously processed in parallel on a multi-core processor.

Possibilities offered by implantable miniaturized cuff-electrodes for insect neurophysiology.

Recent advances in microsystems technology led to a miniaturization of cuff-electrodes, which suggests these electrodes not just for long-term neuronal recordings in mammalians, but also in medium-sized insects. In this study we investigated the possibilities offered by cuff-electrodes for neuroethology using insects as a model organism. The implantation in the neck of a tropical bushcricket resulted in high quality extracellular nerve recordings of different units responding to various acoustic, vibratory, optical and mechanical stimuli. In addition, multi-unit nerve activity related to leg movements was recorded in insects walking on a trackball. A drawback of bi-polar nerve recordings obtained during tethered flight was overlay of nerve activity with large amplitude muscle potentials. Interestingly, cuff-electrode recordings were robust to withstand walking and flight activity so that good quality nerve recordings were possible even three days after electrode implantation. Recording multi-unit nerve activity in intact insects required an elaborate spike sorting algorithm in order to discriminate neuronal units responding to external stimuli from background activity. In future, a combination of miniaturized cuff-electrodes and light-weight amplifiers equipped with a wireless transmitter will allow the investigation of neuronal processes underlying natural behavior in freely moving insects. By this means cuff-electrodes may contribute to the development of realistic neuronal models simulating neuronal processes underlying natural insect behavior, such like mate choice and predator avoidance.

Reliable detection of predator cues in afferent spike trains of a katydid under high background noise levels.

Katydid receivers face the problem of detecting behaviourally relevant predatory cues from echolocating bats in the same frequency domain as their own conspecific mating signals. We therefore tested the hypothesis that katydids are able to detect the presence of insectivorous bats in spike discharges at early stages of nervous processing in the auditory pathway by using the temporal details characteristic for responses to echolocation sequences. Spike activity was recorded from an identified nerve cell (omega neuron) under both laboratory and field conditions. In the laboratory, the preparation was stimulated with sequences of bat calls at different repetition rates typical for the guild of insectivorous bats, in the presence of background noise. The omega cell fired brief high-frequency bursts of action potentials in response to each bat sound pulse. Repetition rates of 18 and 24 Hz of these pulses resulted in a suppression of activity resulting from background noise, thus facilitating the detection of bat calls. The spike activity typical for responses to bat echolocation contrasts to responses to background noise, producing different distributions of inter-spike intervals. This allowed development of a 'neuronal bat detector' algorithm, optimized to detect responses to bats in afferent spike trains. The algorithm was applied to more than 24 hours of outdoor omega-recordings performed either at a rainforest clearing with high bat activity or in rainforest understory, where bat activity was low. In 95% of cases, the algorithm detected a bat reliably, even under high background noise, and correctly rejected responses when an electronic bat detector showed no response.

Simplified bionic solutions: a simple bio-inspired vehicle collision detection system.

Modern cars are equipped with both active and passive sensor systems that can detect potential collisions. In contrast, locusts avoid collisions solely by responding to certain visual cues that are associated with object looming. In neurophysiological experiments, I investigated the possibility that the 'collision-detector neurons' of locusts respond to impending collisions in films recorded with dashboard cameras of fast driving cars. In a complementary modelling approach, I developed a simple algorithm to reproduce the neuronal response that was recorded during object approach. Instead of applying elaborate algorithms that factored in object recognition and optic flow discrimination, I tested the hypothesis that motion detection restricted to a 'danger zone', in which frontal collisions on the motorways are most likely, is sufficient to estimate the risk of a collision. Furthermore, I investigated whether local motion vectors, obtained from the differential excitation of simulated direction-selective networks, could be used to predict evasive steering maneuvers and prevent undesired responses to motion artifacts. The results of the study demonstrate that the risk of impending collisions in real traffic scenes is mirrored in the excitation of the collision-detecting neuron (DCMD) of locusts. The modelling approach was able to reproduce this neuronal response even when the vehicle was driving at high speeds and image resolution was low (about 200 × 100 pixels). Furthermore, evasive maneuvers that involved changing the steering direction and steering force could be planned by comparing the differences in the overall excitation levels of the simulated right and left direction-selective networks. Additionally, it was possible to suppress undesired responses of the algorithm to translatory movements, camera shake and ground shadows by evaluating local motion vectors. These estimated collision risk values and evasive steering vectors could be used as input for a driving assistant, converting the first into braking force and the latter into steering responses to avoid collisions. Since many processing steps were computed on the level of pixels and involved elements of direction-selective networks, this algorithm can be implemented in hardware so that parallel computations enhance the processing speed significantly.

The heterospecific calling song can improve conspecific signal detection in a bushcricket species.

In forest clearings of the Malaysian rainforest, chirping and trilling Mecopoda species often live in sympatry. We investigated whether a phenomenon known as stochastic resonance (SR) improved the ability of individuals to detect a low-frequent signal component typical of chirps when members of the heterospecific trilling species were simultaneously active. This phenomenon may explain the fact that the chirping species upholds entrainment to the conspecific song in the presence of the trill. Therefore, we evaluated the response probability of an ascending auditory neuron (TN-1) in individuals of the chirping Mecopoda species to triple-pulsed 2, 8 and 20 kHz signals that were broadcast 1 dB below the hearing threshold while increasing the intensity of either white noise or a typical triller song. Our results demonstrate the existence of SR over a rather broad range of signal-to-noise ratios (SNRs) of input signals when periodic 2 kHz and 20 kHz signals were presented at the same time as white noise. Using the chirp-specific 2 kHz signal as a stimulus, the maximum TN-1 response probability frequently exceeded the 50% threshold if the trill was broadcast simultaneously. Playback of an 8 kHz signal, a common frequency band component of the trill, yielded a similar result. Nevertheless, using the trill as a masker, the signal-related TN-1 spiking probability was rather variable. The variability on an individual level resulted from correlations between the phase relationship of the signal and syllables of the trill. For the first time, these results demonstrate the existence of SR in acoustically-communicating insects and suggest that the calling song of heterospecifics may facilitate the detection of a subthreshold signal component in certain situations. The results of the simulation of sound propagation in a computer model suggest a wide range of sender-receiver distances in which the triller can help to improve the detection of subthreshold signals in the chirping species.

Rhythm Generation and Rhythm Perception in Insects: The Evolution of Synchronous Choruses.

Insect sounds dominate the acoustic environment in many natural habitats such as rainforests or meadows on a warm summer day. Among acoustic insects, usually males are the calling sex; they generate signals that transmit information about the species-identity, sex, location, or even sender quality to conspecific receivers. Males of some insect species generate signals at distinct time intervals, and other males adjust their own rhythm relative to that of their conspecific neighbors, which leads to fascinating acoustic group displays. Although signal timing in a chorus can have important consequences for the calling energetics, reproductive success and predation risk of individuals, still little is known about the selective forces that favor the evolution of insect choruses. Here, we review recent advances in our understanding of the neuronal network responsible for acoustic pattern generation of a signaler, and pattern recognition in receivers. We also describe different proximate mechanisms that facilitate the synchronous generation of signals in a chorus and provide examples of suggested hypotheses to explain the evolution of chorus synchrony in insects. Some hypotheses are related to sexual selection and inter-male cooperation or competition, whereas others refer to the selection pressure exerted by natural predators. In this article, we summarize the results of studies that address chorus synchrony in the tropical katydid Mecopoda elongata, where some males persistently signal as followers although this reduces their mating success.

Diversity of wing patterns and abdomen-generated substrate sounds in 3 European scorpionfly species.

In the genus Panorpa (Insecta: Mecoptera), also known as scorpionflies, premating behavior includes repeated sequences of slow wing movements (waving, fanning, flagging) which are accompanied by rapid abdomen vibrations that generate substantial substrate-borne sound. It is still unknown whether wing patterns or vibratory signals contain information about species identity, sex and/or the quality of potential mating partners. Besides species-specific pheromones, these multimodal signals may be of particular importance for the maintenance of reproductive isolation in sympatrically occurring scorpionfly species. Here, we analyzed phyologenetic relationships among, and the pattern of forewings as well as substrate-borne sound in 3 different sympatric Central-European scorpionfly species (P. communis, P. germanica, and P. alpina). Divergence time estimates, based on 879 bp of the mitochondrial COI gene, indicate longstanding separate evolutionary histories for the studied Panorpa species. Morphological analysis revealed that wing length as an indicator of body size increased in the following order: P. alpina < P. germanica < P. communis. Individuals can be assigned to the correct species and sex with high accuracy just by evaluation of the number of dark spots and the proportion of wing pigmentation. Despite high variability of interpulse period at an individual level, across species analysis revealed a positive correlation of average interpulse period as well as mean signal amplitude with forewing length. These results suggest wing patterns, but less likely vibratory signals, to contain information about species identity. Furthermore, receivers may be able to estimate the body size of a signaler solely on the basis of substrate-borne sound.

From microseconds to seconds and minutes-time computation in insect hearing.

The computation of time in the auditory system of insects is of relevance at rather different time scales, covering a large range from microseconds to several minutes. At the one end of this range, only a few microseconds of interaural time differences are available for directional hearing, due to the small distance between the ears, usually considered too small to be processed reliably by simple nervous systems. Synapses of interneurons in the afferent auditory pathway are, however, very sensitive to a time difference of only 1-2 ms provided by the latency shift of afferent activity with changing sound direction. At a much larger time scale of several tens of milliseconds to seconds, time processing is important in the context species recognition, but also for those insects where males produce acoustic signals within choruses, and the temporal relationship between song elements strongly deviates from a random distribution. In these situations, some species exhibit a more or less strict phase relationship of song elements, based on phase response properties of their song oscillator. Here we review evidence on how this may influence mate choice decisions. In the same dimension of some tens of milliseconds we find species of katydids with a duetting communication scheme, where one sex only performs phonotaxis to the other sex if the acoustic response falls within a very short time window after its own call. Such time windows show some features unique to insects, and although its neuronal implementation is unknown so far, the similarity with time processing for target range detection in bat echolocation will be discussed. Finally, the time scale being processed must be extended into the range of many minutes, since some acoustic insects produce singing bouts lasting quite long, and female preferences may be based on total signaling time.

Probing real sensory worlds of receivers with unsupervised clustering.

The task of an organism to extract information about the external environment from sensory signals is based entirely on the analysis of ongoing afferent spike activity provided by the sense organs. We investigate the processing of auditory stimuli by an acoustic interneuron of insects. In contrast to most previous work we do this by using stimuli and neurophysiological recordings directly in the nocturnal tropical rainforest, where the insect communicates. Different from typical recordings in sound proof laboratories, strong environmental noise from multiple sound sources interferes with the perception of acoustic signals in these realistic scenarios. We apply a recently developed unsupervised machine learning algorithm based on probabilistic inference to find frequently occurring firing patterns in the response of the acoustic interneuron. We can thus ask how much information the central nervous system of the receiver can extract from bursts without ever being told which type and which variants of bursts are characteristic for particular stimuli. Our results show that the reliability of burst coding in the time domain is so high that identical stimuli lead to extremely similar spike pattern responses, even for different preparations on different dates, and even if one of the preparations is recorded outdoors and the other one in the sound proof lab. Simultaneous recordings in two preparations exposed to the same acoustic environment reveal that characteristics of burst patterns are largely preserved among individuals of the same species. Our study shows that burst coding can provide a reliable mechanism for acoustic insects to classify and discriminate signals under very noisy real-world conditions. This gives new insights into the neural mechanisms potentially used by bushcrickets to discriminate conspecific songs from sounds of predators in similar carrier frequency bands.

A cost-benefit analysis of public and private communication.

Understanding the diversity of animal signals requires knowledge of factors which may influence the different stages of communication, from the production of a signal by the sender up to the detection, identification and final decision-making in the receiver. We studied a Neotropical katydid (Docidocercus gigliotosi) which uses airborne sound for long distance communication, but also an alternative form of private signalling through substrate vibration. Males spend more time with private signalling under full moon conditions, when the nocturnal rainforest favours predation by visually hunting predators. For either type of signal we measured the energetic costs of producing it, its active space, and the background noise levels in both transmission channels. Signal perception was studied using neurophysiological methods under outdoor conditions, which is more reliable for the private mode of communication. Our results demonstrate the complex effects of ecological conditions, such as predation, nocturnal ambient light levels and masking noise on the performance of receivers in detecting mating signals, affecting the net advantage or disadvantage of either mode of communication.

Collective defense of Aphis nerii and Uroleucon hypochoeridis (Homoptera, Aphididae) against natural enemies.

The prevalent way aphids accomplish colony defense against natural enemies is a mutualistic relationship with ants or the occurrence of a specialised soldier caste typical for eusocial aphids, or even both. Despite a group-living life style of those aphid species lacking these defense lines, communal defense against natural predators has not yet been observed there. Individuals of Aphis nerii (Oleander aphid) and Uroleucon hypochoeridis, an aphid species feeding on Hypochoeris radicata (hairy cat's ear), show a behavioral response to visual stimulation in the form of spinning or twitching, which is often accompanied by coordinated kicks executed with hind legs. Interestingly, this behaviour is highly synchronized among members of a colony and repetitive visual stimulation caused strong habituation. Observations of natural aphid colonies revealed that a collective twitching and kicking response (CTKR) was frequently evoked during oviposition attempts of the parasitoid wasp Aphidius colemani and during attacks of aphidophagous larvae. CTKR effectively interrupted oviposition attempts of this parasitoid wasp and even repelled this parasitoid from colonies after evoking consecutive CTKRs. In contrast, solitary feeding A. nerii individuals were not able to successfully repel this parasitoid wasp. In addition, CTKR was also evoked through gentle substrate vibrations. Laser vibrometry of the substrate revealed twitching-associated vibrations that form a train of sharp acceleration peaks in the course of a CTKR. This suggests that visual signals in combination with twitching-related substrate vibrations may play an important role in synchronising defense among members of a colony. In both aphid species collective defense in encounters with different natural enemies was executed in a stereotypical way and was similar to CTKR evoked through visual stimulation. This cooperative defense behavior provides an example of a surprising sociality that can be found in some aphid species that are not expected to be social at all.

The signaller's dilemma: a cost-benefit analysis of public and private communication.

BACKGROUND: Understanding the diversity of animal signals requires knowledge of factors which may influence the different stages of communication, from the production of a signal by the sender up to the detection, identification and final decision-making in the receiver. Yet, many studies on signalling systems focus exclusively on the sender, and often ignore the receiver side and the ecological conditions under which signals evolve. METHODOLOGY/PRINCIPAL FINDINGS: We study a neotropical katydid which uses airborne sound for long distance communication, but also an alternative form of private signalling through substrate vibration. We quantified the strength of predation by bats which eavesdrop on the airborne sound signal, by analysing insect remains at roosts of a bat family. Males do not arbitrarily use one or the other channel for communication, but spend more time with private signalling under full moon conditions, when the nocturnal rainforest favours predation by visually hunting predators. Measurements of metabolic CO(2)-production rate indicate that the energy necessary for signalling increases 3-fold in full moon nights when private signalling is favoured. The background noise level for the airborne sound channel can amount to 70 dB SPL, whereas it is low in the vibration channel in the low frequency range of the vibration signal. The active space of the airborne sound signal varies between 22 and 35 meters, contrasting with about 4 meters with the vibration signal transmitted on the insect's favourite roost plant. Signal perception was studied using neurophysiological methods under outdoor conditions, which is more reliable for the private mode of communication. CONCLUSIONS/SIGNIFICANCE: Our results demonstrate the complex effects of ecological conditions, such as predation, nocturnal ambient light levels, and masking noise levels on the performance of receivers in detecting mating signals, and that the net advantage or disadvantage of a mode of communication strongly depends on these conditions.

The cercal organ may provide singing tettigoniids a backup sensory system for the detection of eavesdropping bats.

Conspicuous signals, such as the calling songs of tettigoniids, are intended to attract mates but may also unintentionally attract predators. Among them bats that listen to prey-generated sounds constitute a predation pressure for many acoustically communicating insects as well as frogs. As an adaptation to protect against bat predation many insect species evolved auditory sensitivity to bat-emitted echolocation signals. Recently, the European mouse-eared bat species Myotis myotis and M. blythii oxygnathus were found to eavesdrop on calling songs of the tettigoniid Tettigonia cantans. These gleaning bats emit rather faint echolocation signals when approaching prey and singing insects may have difficulty detecting acoustic predator-related signals. The aim of this study was to determine (1) if loud self-generated sound produced by European tettigoniids impairs the detection of pulsed ultrasound and (2) if wind-sensors on the cercal organ function as a sensory backup system for bat detection in tettigoniids. We addressed these questions by combining a behavioral approach to study the response of two European tettigoniid species to pulsed ultrasound, together with an electrophysiological approach to record the activity of wind-sensitive interneurons during real attacks of the European mouse-eared bat species Myotis myotis. Results showed that singing T. cantans males did not respond to sequences of ultrasound pulses, whereas singing T. viridissima did respond with predominantly brief song pauses when ultrasound pulses fell into silent intervals or were coincident with the production of soft hemi-syllables. This result, however, strongly depended on ambient temperature with a lower probability for song interruption observable at 21°C compared to 28°C. Using extracellular recordings, dorsal giant interneurons of tettigoniids were shown to fire regular bursts in response to attacking bats. Between the first response of wind-sensitive interneurons and contact, a mean time lag of 860 ms was found. This time interval corresponds to a bat-to-prey distance of ca. 72 cm. This result demonstrates the efficiency of the cercal system of tettigoniids in detecting attacking bats and suggests this sensory system to be particularly valuable for singing insects that are targeted by eavesdropping bats.

Matched filters, mate choice and the evolution of sexually selected traits.

BACKGROUND: Fundamental for understanding the evolution of communication systems is both the variation in a signal and how this affects the behavior of receivers, as well as variation in preference functions of receivers, and how this affects the variability of the signal. However, individual differences in female preference functions and their proximate causation have rarely been studied. METHODOLOGY/PRINCIPAL FINDINGS: Calling songs of male field crickets represent secondary sexual characters and are subject to sexual selection by female choice. Following predictions from the "matched filter hypothesis" we studied the tuning of an identified interneuron in a field cricket, known for its function in phonotaxis, and correlated this with the preference of the same females in two-choice trials. Females vary in their neuronal frequency tuning, which strongly predicts the preference in a choice situation between two songs differing in carrier frequency. A second "matched filter" exists in directional hearing, where reliable cues for sound localization occur only in a narrow frequency range. There is a strong correlation between the directional tuning and the behavioural preference in no-choice tests. This second "matched filter" also varies widely in females, and surprisingly, differs on average by 400 Hz from the neuronal frequency tuning. CONCLUSIONS/SIGNIFICANCE: Our findings on the mismatch of the two "matched filters" would suggest that the difference in these two filters appears to be caused by their evolutionary history, and the different trade-offs which exist between sound emission, transmission and detection, as well as directional hearing under specific ecological settings. The mismatched filter situation may ultimately explain the maintenance of considerable variation in the carrier frequency of the male signal despite stabilizing selection.

A novel distributed swarm control strategy based on coupled signal oscillators.

The miniaturization of microrobots is accompanied by limitations of signaling, sensing and agility. Control of a swarm of simple microrobots has to cope with such constraints in a way which still guarantees the accomplishment of a task. A recently proposed communication method, which is based on the coupling of signal oscillators of individual agents [13], may provide a basis for a distributed control of a simulated swarm of simple microrobots (similar to I-Swarm microrobots) engaged in a cleaning scenario. This self-organized communication method was biologically inspired from males of chorusing insects which are known for the rapid synchronization of their acoustic signals in a chorus. Signal oscillator properties were used to generate waves of synchronized signaling (s-waves) among a swarm of agents. In a simulation of a cleaning scenario, agents on the dump initiated concentrically spreading s-waves by shortening their intrinsic signal period. Dirt-carrying agents localized the dump by heading against the wave front. After optimization of certain control parameters the properties of this distributed control strategy were investigated in different variants of a cleaning scenario. These include a second dump, obstacles, different agent densities, agent drop-out and a second signal oscillator.

Spatial orientation in the bushcricket Leptophyes punctatissima (Phaneropterinae; Orthoptera): I. Phonotaxis to elevated and depressed sound sources.

Many species of acoustically interacting insects live in a complex, arboreal or semi-arboreal habitat. Thus mate finding by phonotaxis requires sound localization in the horizontal and vertical plane. Here we investigated the ability of the duetting bushcricket Leptophyes punctatissima to orient to one of three speakers, positioned at different levels in an artificial grid system, where each point in space could be reached by the male with almost equal probability. The system was designed analogous to a spherical calotte model of bismuth, where, once the male arrived at any nodal point had to decide between only three directions: either up or down and/or left and right. This design does not favour any phonotactic path of the males. All 12 males tested reached the three speaker positions (one in the horizontal plane, one elevated by 45 degrees , one depressed by 45 degrees relative to the starting position) with only little deviation from the shortest possible path. There was no significant difference with respect to the whole phonotactic time needed, the number of segments passed, or the number of stimuli received for the different speaker positions. This remarkable spatial orientation is achieved although the insects have no specialized external ear structures such as mammals, or some owls.

Mechanisms for synchrony and alternation in song interactions of the bushcricket Mecopoda elongata (Tettigoniidae: Orthoptera).

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.

[The "3R" concept in the Internet I]

More and more members of the scientific community interested in 3R related topics of animal experiments are using Internet-Homepages as a low-budget forum to present either their scientific results, or opinions on specific subjects or to announce publications. Supra/inter/national authorities/universities/laboratories are also offering information, documents, access to databases and much more to the interested public. Therefore the Internet is an advantageous new medium for 3R related subjects.