Chirp Duration Research Articles

Callback response of dugongs to conspecific chirp playbacks

Dugongs (Dugong dugon) produce bird-like calls such as chirps and trills. The vocal responses of dugongs to playbacks of several acoustic stimuli were investigated. Animals were exposed to four different playback stimuli: a recorded chirp from a wild dugong, a synthesized down-sweep sound, a synthesized constant-frequency sound, and silence. Wild dugongs vocalized more frequently after playback of broadcast chirps than that after constant-frequency sounds or silence. The down-sweep sound also elicited more vocal responses than did silence. No significant difference was found between the broadcast chirps and the down-sweep sound. The ratio of wild dugong chirps to all calls and the dominant frequencies of the wild dugong calls were significantly higher during playbacks of broadcast chirps, down-sweep sounds, and constant-frequency sounds than during those of silence. The source level and duration of dugong chirps increased significantly as signaling distance increased. No significant correlation was found between signaling distance and the source level of trills. These results show that dugongs vocalize to playbacks of frequency-modulated signals and suggest that the source level of dugong chirps may be manipulated to compensate for transmission loss between the source and receiver. This study provides the first behavioral observations revealing the function of dugong chirps.

Evaluating the importance of different temporal features of calling songs in cricket phonotaxis

Event Abstract Back to Event Evaluating the importance of different temporal features of calling songs in cricket phonotaxis Gundula Meckenhäuser1, 2*, Matthias R. Hennig3 and Martin P. Nawrot1, 2 1 Freie Universität Berlin, Institute of Biology, Germany 2 Bernstein Center for Computational Neuroscience Berlin, Germany 3 Humboldt-Universität zu Berlin, Department of Biology, Germany Acoustic signals are used by Gryllus bimaculatus to communicate and they play a key role in mate choice. Males produce calling songs to attract females, females in turn use these songs to discriminate conspecific songs from the signals of other species. If they are willing to pair females approach the singing male - a behavior referred to as phonotaxis. So far, extensive behavioral experiments in which phonotactic responses of female crickets were tested by varying song features, revealed that song patterns are processed in the time rather than in the spectral domain [1]. Here, we explored the relevant cues for song pattern discrimination on several time scales. We present feed-forward artificial neural networks (ANNs) that quantitatively predict the phonotactic value for untested calling songs that are described by temporal features such as duration, pause, period and duty cycle for both pulses and chirps. We employ ANNs with the following architecture: Input neurons representing temporal song features project to nonlinear neurons in the hidden layer which project to the output neuron that relates the phonotactic value. For training and testing the networks we used 218 artificial songs for which the phonotactic values had already been determined in experiments. Our complete ANNs that use eight temporal song features show a mean squared error (MSE) of MSE = 0.034. By a greedy backward elimination of features, we came up with minimal ANNs using just pulse period, chirp duration and chirp duty cycle as features. The dimensionality reduction of the feature space also improved the performance: The mean squared error for minimal networks is MSE = 0.019. Thus, our findings suggest that among all tested temporal features, pulse period on the short time scale and chirp duration and chirp duty cycle on the long time scale are the most relevant ones for the evaluation of conspecific signals. Further, the minimal ANNs show high predictive power and can be used to complement experimental testing of female phonotaxis in the laboratory. Acknowledgements This work is funded by the German Research Council (DFG) within the Collaborative Research Center "Theoretical Biology" (SFB 618).

Low Complexity Chirp Pulsed Ultra-Wideband System with Near-Optimum Multipath Performance

This paper describes the Impulse Radio Ultra-Wideband (IR-UWB) system using linear chirp UWB pulses as symbols. The novel method of coherent or differentially coherent detection of chirp pulses in multipath channels is introduced. The method divides detection in the receiver between its analog and digital part; in the analog part of the receiver, received signal is compressed in frequency by mixing with locally generated chirp pulse and low-pass filtering. In this way, the time-bandwidth (TB) product of the resulting signal is reduced compared to the received signal and made approximately independent of duration of the transmitted signal. This results in relatively low complexity digital back-end of the receiver that is able to capture a large portion of the multipath energy, regardless of duration of the transmitted waveform. The work includes derivation of analytical properties of the system, including derivation of loss parameters that quantify system multipath performance. Numerical results include discussion on the system multipath loss performance with varying chirp duration. As well, a comparison is made with error performance of sampling receivers featuring the same computational complexity. Analysis of the robustness of the system's performance to different chirp pulse implementation errors is also provided.

Effects of juvenile and adult condition on long-distance call components in the Jamaican field cricket, Gryllus assimilis

Complex signals may arise through either content- or efficacy-based selection. Content-based signals are thought to evolve because of the information content they provide to the receiver. Such signals may function as multiple messages, each relaying different information or types of information. For example, some signals or signal components may indicate different aspects of condition, or condition at different life stages (quality signals), while others relay information on signaller location, or function in species recognition. Or they may act as redundant signals of overall condition, or as backup signals of the same aspect of condition. Efficacy-based signals are proposed to evolve because of the way in which they influence the production, transmission, reception or processing of the signal. We manipulated juvenile and adult condition in male Jamaican field crickets, Gryllus assimilis, to test whether changes in components of the male’s long-distance acoustic mate attraction signal were consistent with content- or efficacy-based selection. Several call components exhibited condition-dependence, and may be under content-based selection. Interpulse duration, pulse rate and chirp duration appeared to honestly signal juvenile condition, while dominant frequency appeared to signal adult condition. Thus, these signal components appeared to be acting as multiple messages of quality. Pulse number, interchirp duration and chirp rate were highly stereotyped and probably function in species recognition. Call number increased with decreasing condition and was likely under efficacy-based selection. No call components appeared to have the potential to act as redundant signals.

Quasimonoenergic collimated electrons from the ionization of nitrogen by a chirped intense laser pulse

A scheme is proposed for quasimonoenergic collimated GeV electrons generated during ionization of nitrogen by a chirped intense laser pulse. The electrons accelerated by a laser pulse without a frequency chirp are known for poor-quality beams. If a suitable frequency chirp is introduced, then the energy of the electrons increases significantly. It is shown that quasimonoenergic collimated GeV electrons can be produced using a right choice of laser spot size, frequency chirp, and pulse duration.

Effects of frequency chirp on magnetron injection locking

The injection locking of a magnetron is theoretically analyzed when either the free running oscillator or the drive signal has a frequency chirp. It is found that complete phase locking of the signal cannot be achieved in either case. However, as long as the locking condition of Adler is well-satisfied instantaneously, a high degree of locking occurs during a major duration of the frequency chirps. The expected output phase variation is computed in terms of the noise in the free-running magnetron oscillator for the case of constant drive frequency.

Phylogenetic comparative analysis of electric communication signals in ghost knifefishes (Gymnotiformes: Apteronotidae)

Electrocommunication signals in electric fish are diverse, easily recorded and have well-characterized neural control. Two signal features, the frequency and waveform of the electric organ discharge (EOD), vary widely across species. Modulations of the EOD (i.e. chirps and gradual frequency rises) also function as active communication signals during social interactions, but they have been studied in relatively few species. We compared the electrocommunication signals of 13 species in the largest gymnotiform family, Apteronotidae. Playback stimuli were used to elicit chirps and rises. We analyzed EOD frequency and waveform and the production and structure of chirps and rises. Species diversity in these signals was characterized with discriminant function analyses, and correlations between signal parameters were tested with phylogenetic comparative methods. Signals varied markedly across species and even between congeners and populations of the same species. Chirps and EODs were particularly evolutionarily labile, whereas rises differed little across species. Although all chirp parameters contributed to species differences in these signals, chirp amplitude modulation, frequency modulation (FM) and duration were particularly diverse. Within this diversity, however, interspecific correlations between chirp parameters suggest that mechanistic trade-offs may shape some aspects of signal evolution. In particular, a consistent trade-off between FM and EOD amplitude during chirps is likely to have influenced the evolution of chirp structure. These patterns suggest that functional or mechanistic linkages between signal parameters (e.g. the inability of electromotor neurons increase their firing rates without a loss of synchrony or amplitude of action potentials) constrain the evolution of signal structure.

Maximizing the yield and cutoff of high-order harmonic generation from plasma plume

We study high-order harmonic generation (HHG) from various lowly ionized laser plasmas. We study harmonic generation from targets of Al (Z=13) to Bi (Z=83). Varying the wavelength, chirp, and pulse duration of the femtosecond pump laser resulted in the change in the harmonic distribution, cutoff, and conversion efficiency of HHG. We also study the use of doubly charged ions, and resonances for some materials. We were able to obtain high HHG conversion efficiency and harmonic cutoff by implementing the above approaches and by observing the time-resolved spectra of the laser plasma.

Variable pattern of high-order harmonic spectra from a laser-produced plasma by using the chirped pulses of narrow-bandwidth radiation

Various plasmas prepared by laser ablation of the surfaces of solid targets were examined by the narrow-bandwidth radiation of different chirp and pulse durations. The high-order harmonics generated during laser-plasma interaction showed different brightness, wavelength shift, harmonic cutoff, and efficiency by using variable chirps of pump radiation. An analysis of harmonic optimization at these conditions is presented. The blueshifted and redshifted harmonics observed in this case were analyzed and attributed to the abundance of free electrons and self-phase modulation of the driving pulse. The resonance-induced enhancement of the 15th harmonic from GaN-nanoparticle-containing plasma caused by the tuning of harmonic wavelength close to the ionic transition was demonstrated.

Host preferences in a phonotactic parasitoid of field crickets: the relative importance of host song characters

Abstract 1. Predators, including insect parasitoids, often eavesdrop on prey signals, and as a result, predation can have important effects on the evolution of prey signalling behaviour.2. The phonotactic parasitoid fly, Ormia ochracea, uses the calling songs of male field crickets to locate their field crickets hosts. In the western USA, this fly parasitises the variable field cricket, Gryllus lineaticeps. Previous work with one fly population suggested that female flies, like female field crickets, preferentially orient to male songs with higher chirp rates and longer chirp durations, although a limited range of male song types was used in this previous study. The current study, with a different fly population, used field‐based, two‐speaker choice tests to examine: (1) the effect of male chirp rate and chirp duration on fly attraction, using a natural range of song types; and (2) the relative importance of these song types in host selection by the flies.3. Three lines of evidence suggested that chirp rate is more important than chirp duration in host selection. (a) The flies consistently preferred higher chirp rates but only sometimes preferred longer chirp durations. (b) The flies consistently preferred higher chirp rate/shorter chirp duration songs to lower chirp rate/longer chirp duration songs. (c) Preferences for longer chirp durations could be eliminated by increasing the amplitude of the less attractive song type, while preferences for higher chirp rates could only sometimes be eliminated by increasing the amplitude of the less attractive song type.4. Fly predation may favour lower chirp rates and shorter chirp durations in G. lineaticeps, and may have resulted in stronger selection on chirp rate than on chirp duration.

THE RELATIVE IMPORTANCE OF DIFFERENT DIRECT BENEFITS IN THE MATE CHOICES OF A FIELD CRICKET

Discussions about the evolution of female mating preferences have often suggested that females should express multiple strong preferences when different male traits are correlated with different mating benefits, yet few studies have directly tested this hypothesis by comparing the strength of female preferences for male traits known to be correlated with different benefits. In the variable field cricket, Gryllus lineaticeps, females receive fecundity and fertility benefits from mating with males with higher chirp rates and life-span benefits from mating with males with longer chirp durations. Although females prefer higher chirp rates and longer chirp durations when the other trait is held constant, it is possible that they give priority to one of these song traits when both vary. In this study, we examined the relative importance of chirp rate and chirp duration in female mate choice using single-stimulus presentations of songs that varied in both chirp rate and chirp duration. Females expressed both directional and stabilizing preferences based on chirp rate, responding most strongly to a chirp rate approximately one standard deviation above the population mean. Females did not express preferences based on chirp duration, and did not express correlational preferences. These results suggest that females may give priority to the reproductive benefits provided by males that produce higher chirp rates.

CALLING IS AN HONEST INDICATOR OF PATERNAL GENETIC QUALITY IN POISON FROGS

Several competing hypotheses have been put forward to explain why females of many species mate preferentially with males possessing the most conspicuous signals (e.g., ornaments, displays, or songs). We performed a laboratory experiment using two species of poison frogs, Dendrobates leucomelas and Epipedobates tricolor, to test the hypothesis that male calling performance is an honest indicator of parental quality. Our analyses are based on data from behavioral observations of mating activities of captive-reared individuals (and their offspring) that were housed in terraria for four consecutive breeding seasons. Male mating success increased with male calling rate and chirp duration in both species, suggesting that females preferred males with more elaborate calls. Because calling performance improved with age in D. leucomelas, female poison frogs that prefer males with more elaborate calls in the wild may end up mating with older males that have already proven their ability to survive. Females that mated with good callers obtained higher quality offspring. Eggs fertilized by males with high calling rates and long chirp durations had higher hatching success and produced tadpoles that were more likely to metamorphose into surviving frogs. As a consequence, females that mated with males with high calling performance obtained more surviving offspring per egg, compared to females that mated with poor callers. Collectively, our findings comply with the notion that female poison frogs prefer to mate with good callers because calling performance is a reliable predictor of offspring quality. The possible influence of maternal allocation and reasons for the strong effect size compared to previous studies are discussed.

Male spacing behaviour and acoustic interactions in a field cricket: implications for female mate choice

Males of several acoustically communicating orthopteran species form spatially and temporally structured choruses. We investigated whether male field crickets of the species Plebeiogryllus guttiventris formed choruses in the field. Males formed spatial aggregations and showed fidelity to a calling site within a night, forming stable choruses. Within aggregations, the acoustic ranges of males overlapped considerably. We tested whether males within hearing range of each other interacted acoustically. The chirps of simultaneously calling males were aphasic with respect to each other and showed no significant alternation or synchrony of calls. Some individuals changed temporal features of their calling songs such as chirp durations and chirp rates in response to a simultaneously calling neighbour. The implications of these results for female mate choice are discussed.

MALE AND FEMALE STRIDULATION IN AN INDIAN WETA (ORTHOPTERA: ANOSTOSTOMATIDAE)

ABSTRACT We present the first description of the calls and stridulatory structures of males and females of an Indian weta species from the Western Ghats of Southern India. Calls of males and females produced by femoro-abdominal stridulation were recorded and call features such as chirp duration, chirp period, syllable period and syllable duration were characterised. The calls of males were highly stereotyped with regular chirp periods and durations whereas chirp rates in the female calls were more variable. The number of syllables per chirp was constant for males and females and the fine temporal features such as syllable periods and syllable durations showed low inter-individual variation in both males and females. The arrangement of femoral stridulatory structures was different from the previously described anostostomatid species. The high stereotypy of calls of males and females indicates that the signal could serve for identification of species and sex.

Male field crickets that provide reproductive benefits to females incur higher costs

Abstract. 1. Females often select mates based on signals correlated with the quality of the direct benefits that males will provide to them. A male's quality as a mate and the structure of his mating signals may covary because both traits are energetically expensive for males to produce and because both traits are affected by short‐term changes in nutritional condition.2. In the variable field cricket, Gryllus lineaticeps, previous work has shown that females receive reproductive benefits from males that produce higher chirp rates and lifespan benefits from males that produce longer chirp durations, even when they only receive the sperm and seminal fluid contained in male spermatophores. Higher chirp rates are energetically expensive for males to produce, and chirp rate is strongly affected by diet quality, whereas longer chirp durations do not appear to be expensive for males to produce, and chirp duration does not appear to be affected by male diet quality. In this study two hypotheses were tested about the energetic costs of spermatophore production: (1) that spermatophores are expensive for males to produce and (2) that males providing greater direct benefits to females incur higher costs of spermatophore production.3. Males that were provided with a lower quality diet took longer to produce a new spermatophore. This result suggests that spermatophores are costly for males to produce.4. Males that produced higher chirp rates took longer to produce a new spermatophore. This result suggests that male chirp rate and female reproductive benefits may covary because both traits are energetically expensive for males to produce and thus are affected by male nutritional condition. There was no association, however, between male chirp duration and spermatophore production time.

Signal components, acoustic preference functions and sexual selection in a cricket

In many sexually reproducing organisms, females choose mates based on multiple male traits. This study examined how two temporal components of the male mating call – chirp rate and chirp duration – affect female mating preference in five populations of a widely distributed North American cricket, Allonemobius socius (Orthoptera, Gryllidae). Chirp rate and chirp duration of the A. socius mating call were varied independently, and the responses of virgin females to these experimentally manipulated calls were repeatedly measured using a sequential single-stimulus design. Significant among- and within-population variation in chirp-duration preferences of females were found. Contrary to many previous studies, call chirp rate had no effect on female phonotaxis. Also there was no evidence of an interaction between chirp rate and chirp duration on female response to male mating calls. Moreover, female responsiveness to average and above-average chirp duration appeared to decline with female (adult) age. Overall, these results suggest evolved differences among populations in chirp-duration preferences, and that selection can act within populations on female chirp-duration preference. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 461–472.

Acoustic communication in Okanagana rimosa (Say) (Homoptera: Cicadidae)

The cicada Okanagana rimosa (Say) has an acoustic communication system with three types of loud timbal sounds: (i) A calling song lasting several seconds to about 1min which consists of a sequence of chirps at a repetition rate of 83 chirps per second. Each chirp of about 6ms duration contains 4–5 pulses. The sound level of the calling song is 87–90dB SPL at a distance of 15cm. (ii) An amplitude modulated courtship song with increasing amplitude and repetition rate of chirps and pulses. (iii) A protest squawk with irregular chirp and pulse structure. The spectra of all three types are similar and show main energy peaks at 8–10kHz. Only males sing, and calling song production is influenced by the songs of other males, resulting in an almost continuous sound in dense populations. In such populations, the calling songs overlap and the temporal structure of individual songs is obscured within the habitat. The calling song of the broadly sympatric, closely related species O. canadensis (Provander) is similar in frequency content, but distinct in the temporal pattern (24 chirps per second, 24ms chirp duration, eight pulses per chirp) which is likely important for species separation in sympatric populations.The hearing threshold of the auditory nerve is similar for females and males of O. rimosa and most sensitive at 4–5kHz. Experiments in the field show that female phonotaxis of O. rimosa depends on parameters of the calling song. Most females are attracted to calling song models with a 9kHz carrier frequency (peak frequency of the calling song), but not to models with a 5kHz carrier frequency (minimum hearing threshold). Phonotaxis depends on temporal parameters of the conspecific song, especially chirp repetition rate. Calling song production is influenced by environmental factors, and likelihood to sing increases with temperature and brightness of the sky. Correspondingly, females perform phonotaxis most often during sunny conditions with temperatures above 22°C. Non-mated and mated females are attracted by the acoustic signals, and the percentage of mated females performing phonotaxis increases during the season.

Condition dependence of a multicomponent sexual signal in the field cricket Gryllus campestris

In choosing a breeding partner, females in many animal species select between available males on the basis of several signalling traits. Some theoretical models of signalling evolution predict that multiple ornaments convey specific information on different aspects of male quality, such as current nutritional condition. We investigated the effect of nutrition on the calling song of male field crickets Gryllus campestris. This song is a multicomponent sexually selected signal. Adult males were kept on one of three feeding regimes, which resulted in significant differences in body condition between experimental groups. We found significant increases in calling rate and chirp rate and a significant decrease in interchirp duration with increasing food level. Other song characters, such as chirp duration, syllable number, chirp intensity and carrier frequency, were not affected by the food treatment. Furthermore, carrier frequency was correlated with harp area, which is an index of structural size in adult males. The calling song of the field cricket may thus serve as a multicomponent sexual signal, which contains discrete information on past growth and juvenile development as well as present nutritional condition. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Diversity in the structure of electrocommunication signals within a genus of electric fish, Apteronotus.

Some gymnotiform electric fish modulate their electric organ discharge for intraspecific communication. In Apteronotus leptorhynchus, chirps are usually rapid (10-30 ms) modulations that are activated through non- N-methyl- d-aspartate (non-NMDA) glutamate receptors in the hindbrain pacemaker nucleus. Males produce longer chirp types than females and chirp at higher rates. In Apteronotus albifrons, chirp rate is sexually monomorphic, but chirp structure (change in frequency and amplitude during a chirp) was unknown. To better understand the neural regulation and evolution of chirping behavior, we compared chirp structure in these two species under identical stimulus regimes. A. albifrons, like A. leptorhynchus, produced distinct types of chirps that varied, in part, by frequency excursion. However, unlike in A. leptorhynchus, chirp types in A. albifrons varied little in duration, and chirps were all longer (70-200 ms) than those of A. leptorhynchus. Chirp type production was not sexually dimorphic in A. albifrons, but within two chirp types males produced longer chirps than females. We suggest that species differences in chirp duration might be attributable to differences in the relative proportions of fast-acting (non-NMDA) and slow-acting (NMDA) glutamate receptors in the pacemaker. Additionally, we map species difference onto a phylogeny and hypothesize an evolutionary sequence for the diversification of chirp structure.

FEMALE LIFE SPAN AND FERTILITY ARE INCREASED BY THE EJACULATES OF PREFERRED MALES

In animals with internal fertilization, sperm competition among males can favor the evolution of male ejaculate traits that are detrimental to females. Female mating preferences, in contrast, often favor traits in males that are beneficial to females, yet little is known about the effect of these preferences on the evolution of male ejaculates. A necessary condition for female preferences to affect the evolution of male ejaculate characteristics is that females select mates based on a trait correlated with ejaculate quality. Previous work has shown that females of the variable field cricket, Gryllus lineaticeps, prefer males that produce calling songs containing faster and longer chirps. In this study, we tested the hypothesis that females receive more beneficial ejaculates from preferred males. Females were placed on either a high- or a reduced-nutrition diet then mated twice to a male of known song phenotype. Females received only sperm and seminal fluid from males during these matings. There was no effect of male song phenotype on any fitness component for females on the high-nutrition diet. Reduced-nutrition females mated to males that produced preferred song types, however, lived longer, produced more eggs, produced more fertile eggs, and had a higher proportion of their eggs fertilized than those mated to other males. The life-span benefit was positively associated with male chirp duration, and the reproductive benefits were positively associated with male chirp rate. We explored two possible mechanisms for the life span and reproductive benefits. First, a path analysis suggested that part of the effect of male chirp duration on female life span may have been indirect; females mated to males that produced longer chirps showed delayed oviposition, and females that delayed oviposition lived longer. Males that produce longer chirps may thus transfer fewer or less potent oviposition stimulants to females in their seminal fluid. Second, there was a positive correlation between male chirp rate and the number of sperm transferred to females. The fertility benefit may thus have resulted from females receiving more sperm from males that produce faster chirps. Finally, there was a negative phenotypic correlation between male chirp rate and chirp duration, suggesting that females may have to trade off the life span and reproduction benefits when selecting a mate.