Spike-timing-based computation in sound localization

Spike timing is precise in the auditory system and it has been argued that it conveys information about auditory stimuli, in particular about the location of a sound source. However, beyond simple time differences, the way in which neurons might extract this information is unclear and the potential computational advantages are unknown. The computational difficulty of this task for an animal is to locate the source of an unexpected sound from two monaural signals that are highly dependent on the unknown source signal. In neuron models consisting of spectro-temporal filtering and spiking nonlinearity, we found that the binaural structure induced by spatialized sounds is mapped to synchrony patterns that depend on source location rather than on source signal. Location-specific synchrony patterns would then result in the activation of location-specific assemblies of postsynaptic neurons. We designed a spiking neuron model which exploited this principle to locate a variety of sound sources in a virtual acoustic environment using measured human head-related transfer functions. The model was able to accurately estimate the location of previously unknown sounds in both azimuth and elevation (including front/back discrimination) in a known acoustic environment. We found that multiple representations of different acoustic environments could coexist as sets of overlapping neural assemblies which could be associated with spatial locations by Hebbian learning. The model demonstrates the computational relevance of relative spike timing to extract spatial information about sources independently of the source signal.     

Limited condition dependence of male acoustic signals in the grasshopper Chorthippus biguttulus

In many animal species, male acoustic signals serve to attract a mate and therefore often play a major role for male mating success. Male body condition is likely to be correlated with male acoustic signal traits, which signal male quality and provide choosy females indirect benefits. Environmental factors such as food quantity or quality can influence male body condition and therefore possibly lead to condition-dependent changes in the attractiveness of acoustic signals. Here, we test whether stressing food plants influences acoustic signal traits of males via condition-dependent expression of these traits. We examined four male song characteristics, which are vital for mate choice in females of the grasshopper Chorthippus biguttulus. Only one of the examined acoustic traits, loudness, was significantly altered by changing body condition because of drought- and moisture-related stress of food plants. No condition dependence could be observed for syllable to pause ratio, gap duration within syllables, and onset accentuation. We suggest that food plant stress and therefore food plant quality led to shifts in loudness of male grasshopper songs via body condition changes. The other three examined acoustic traits of males do not reflect male body condition induced by food plant quality.     

The use of bioacoustic characters for distinguishing between cryptic species in insects: Potentials,restrictions, and prospects

Differences in acoustic signal patterns between closely related species often form the main precopulatory reproductive barrier in insects. For this reason, discrimination between similar forms based on their signal patterns allows one to recognize reproductively isolated species. A calling signal produced by a mature male for attracting a conspecific female is a “species ID” testifying to the fact that it belongs to the same species. This is the reason why the use of the calling signal analysis for discriminating between cryptic species or elucidating the taxonomic rank of dubious forms is highly efficient. In certain species, courtship signals also show species-specific patterns. In insect taxonomy, the study of acoustic signals is most promising in morphologically or/and ecologically heterogeneous groups. Chances that any morphologically homogeneous form will actually appear to be a complex of cryptic species are low. The degree of signal variability differs in different orders, families and even congeneric species. Therefore, in every taxon investigated for the first time, it is necessary to evaluate the limits of intraspecific variability of signals before using the acoustic characters in taxonomy. Species not involved in acoustic interactions due to allopatry, different host specialization, etc. can produce calling signals with identical patterns. Consequently, similarity in the signal structure in such species is by no means evidence of their synonymy.     

Modeling Complex Phenotypes: Generalized Linear Models Using Spectrogram Predictors of Animal Communication Signals

Summary A major goal of evolutionary biology is to understand the dynamics of natural selection within populations. The strength and direction of selection can be described by regressing relative fitness measurements on organismal traits of ecological significance. However, many important evolutionary characteristics of organisms are complex, and have correspondingly complex relationships to fitness. Secondary sexual characteristics such as mating displays are prime examples of complex traits with important consequences for reproductive success. Typically, researchers atomize sexual traits such as mating signals into a set of measurements including pitch and duration, in order to include them in a statistical analysis. However, these researcher‐defined measurements are unlikely to capture all of the relevant phenotypic variation, especially when the sources of selection are incompletely known. In order to accommodate this complexity we propose a Bayesian dimension‐reduced spectrogram generalized linear model that directly incorporates representations of the entire phenotype (one‐dimensional acoustic signal) into the model as a predictor while accounting for multiple sources of uncertainty. The first stage of dimension reduction is achieved by treating the spectrogram as an “image” and finding its corresponding empirical orthogonal functions. Subsequently, further dimension reduction is accomplished through model selection using stochastic search variable selection. Thus, the model we develop characterizes key aspects of the acoustic signal that influence sexual selection while alleviating the need to extract higher‐level signal traits a priori. This facet of our approach is fundamental and has the potential to provide additional biological insight, as is illustrated in our analysis.     

影响稻褐飞虱声诱集的若干因素(同翅目：飞虱科)

稻褐飞虱Nilaparvata lugens(stal)雌、雄成虫均能发出由固体介质(寄主植株)传递的鸣声(振动)信号,用于交尾前的个体间通讯。雄虫对雌虫鸣声或重放鸣声录音具有强趋性。由此研制的声诱集器有可能首先用于改进褐飞虱测报的田间采样技术。 声诱集效果受若干因素的影响。实验室试验结果表明,声诱集具有种的选择性;诱集率随雄虫虫龄变化,对羽化后4一5日龄雄虫的诱集率最高;异性干扰对诱集效果有明显影响;一天中,中午前后诱集率最高,9：00-18：00的诱集量占总诱集量的54.0%±10.8%;诱集率随气温变化,26℃-28℃诱集效果最好;而室外晴、阴、雨等天气差异对诱集串无显著影响;另外,随诱集信号播放时间的延续,诱集率下降;间断信号的诱集效果高于连续信号。     

Repeated decrease in vocal repertoire size in Streptopelia doves

Most bird species produce different acoustic signals in different behavioural contexts. This intraspecific variation in signal types is thought to be the result of selection for optimal communication in each context. Doves in the genus Streptopelia have three distinct behavioural contexts in which they produce coo vocalizations. Some Streptopelia species have three acoustically similar coo vocalizations associated with the three contexts, but in others the coo vocalizations differ in acoustic structure. Using a well-resolved phylogeny, we examined whether acoustic differentiation between coo types was the ancestral state. Unexpectedly, the results showed that the common ancestor of Streptopelia had differentiated coos rather than a single coo type. This result implies that context-specific acoustic signals disappeared from the vocal repertoire independently at least two times. We further tested whether different context-dependent signal types follow different evolutionary pathways and whether they differ in rate of evolutionary change. We found that the long-range signal (perch-coo) evolves at a higher rate than the short-range signal (bow-coo). These results are discussed in relation to selection for species recognition and transmission requirements.     

Dependence of the perception of emotional information of speech on the acoustic parameters of the stimulus in children of various ages

The effect on the perception of the emotional prosody of a vocal signal of the acoustic parameters of the stimulus in children of various ages (7–10, 11–13, and 14–17 years) was studied. Considerable differences in recognition of positive and negative valences of the vocal signal against the background of noise and in its absence were demonstrated. The ontogenetic specific features of the dependence of recognition of the emotional valence on the acoustic parameters of a vocal signal were determined. The most important acoustic characters (the fundamental frequency F0 and the frequency of the first formant F1) that ensure the perception of the emotional prosody of a vocal signal against the background of noise at different age stages were found.     

Divergent Receiver Responses to Components of Multimodal Signals in Two Foot-Flagging Frog Species

Multimodal communication of acoustic and visual signals serves a vital role in the mating system of anuran amphibians. To understand signal evolution and function in multimodal signal design it is critical to test receiver responses to unimodal signal components versus multimodal composite signals. We investigated two anuran species displaying a conspicuous foot-flagging behavior in addition to or in combination with advertisement calls while announcing their signaling sites to conspecifics. To investigate the conspicuousness of the foot-flagging signals, we measured and compared spectral reflectance of foot webbings of Micrixalus saxicola and Staurois parvus using a spectrophotometer. We performed behavioral field experiments using a model frog including an extendable leg combined with acoustic playbacks to test receiver responses to acoustic, visual and combined audio-visual stimuli. Our results indicated that the foot webbings of S. parvus achieved a 13 times higher contrast against their visual background than feet of M. saxicola. The main response to all experimental stimuli in S. parvus was foot flagging, whereas M. saxicola responded primarily with calls but never foot flagged. Together these across-species differences suggest that in S. parvus foot-flagging behavior is applied as a salient and frequently used communicative signal during agonistic behavior, whereas we propose it constitutes an evolutionary nascent state in ritualization of the current fighting behavior in M. saxicola.     

Modelling the effects of chorus species composition and caller density on acoustic masking interference in multispecies choruses of crickets and katydids

Natural multispecies acoustic choruses such as the dusk chorus of a tropical rain forest consist of simultaneously signalling individuals of different species whose calls travel through a common shared medium before reaching their ‘intended’ receivers. This causes masking interference between signals and impedes signal detection, recognition and localization. The levels of acoustic overlap depend on a number of factors, including call structure, intensity, habitat-dependent signal attenuation and receiver tuning. In addition, acoustic overlaps should also depend on caller density and the species composition of choruses, including relative and absolute abundance of the different calling species. In this study, we used simulations to examine the effects of chorus species relative abundance and caller density on the levels of effective heterospecific acoustic overlap in multispecies choruses composed of the calls of five species of crickets and katydids that share the understorey of a rain forest in southern India. We found that on average species-even choruses resulted in higher levels of effective heterospecific acoustic overlap than choruses with strong dominance structures. This effect was found consistently across dominance levels ranging from 0.4 to 0.8 for larger choruses of forty individuals. For smaller choruses of twenty individuals, the effect was seen consistently for dominance levels of 0.6 and 0.8 but not 0.4. Effective acoustic overlap (EAO) increased with caller density but the manner and extent of increase depended both on the species' call structure and the acoustic context provided by the composition scenario. The Phaloria sp. experienced very low levels of EAO and was highly buffered to changes in acoustic context whereas other species experienced high EAO across contexts or were poorly buffered. These differences were not simply predictable from call structures. These simulation-based findings may have important implications for acoustic biodiversity monitoring and for the study of acoustic masking interference in natural environments.     

Singing in the rain forest: how a tropical bird song transfers information

How information transmission processes between individuals are shaped by natural selection is a key question for the understanding of the evolution of acoustic communication systems. Environmental acoustics predict that signal structure will differ depending on general features of the habitat. Social features, like individual spacing and mating behavior, may also be important for the design of communication. Here we present the first experimental study investigating how a tropical rainforest bird, the white-browed warbler Basileuterus leucoblepharus, extracts various information from a received song: species-specific identity, individual identity and location of the sender. Species-specific information is encoded in a resistant acoustic feature and is thus a public signal helping males to reach a wide audience. Conversely, individual identity is supported by song features susceptible to propagation: this private signal is reserved for neighbors. Finally, the receivers can locate the singers by using propagation-induced song modifications. Thus, this communication system is well matched to the acoustic constraints of the rain forest and to the ecological requirements of the species. Our results emphasize that, in a constraining acoustic environment, the efficiency of a sound communication system results from a coding/decoding process particularly well tuned to the acoustic properties of this environment.     

Why birds sing at dawn: the role of consistent song transmission

The dawn chorus is a widely observed phenomenon. One of the common, but inadequately studied, explanations for the occurrence of the dawn chorus is based on the rationale that atmospheric turbulence, which impairs acoustic communication, is least at dawn, and thus singing at dawn in some way maximizes signal performance. To investigate what possible acoustic benefit is gained through singing at dawn, we transmitted Swamp Sparrow Melospiza georgiana and White-throated Sparrow Zonotrichia albicollis song through open grassland and closed forest both at dawn and at midday. The transmitted songs were re-recorded at four distances from 25 to 100 m. Our results show that the mean overall absolute transmission quality of the signals was not significantly better at dawn than at midday. However, the signal transmission quality was significantly more consistent at dawn than at midday. Also, in general, signal transmission quality decreased with increasing distance. Variability in the transmission quality increased with distance for the White-throated Sparrow song, but not for the Swamp Sparrow song. Consistency in signal transmission quality is a factor that, arguably, is crucial for the identity function of song. This study strongly supports the acoustic transmission hypothesis as an explanation for the existence of the dawn chorus while the demonstration of variability as a key factor in singing at dawn is novel.     

Comparative evaluation of the response of rat-mothers to the natural pain squeaks of a ratling and to a tape recording of it

Two kinds of stimuli were presented to the mother rat at random: the natural pain acoustic signal of its infant in response to electrical pain stimulation and the tape recording of the signal performed within the sound frequency region (up to 20 kHz). The mother rat was placed in a special long cage with two parts of different size. Her reaction was ethologically estimated. The natural pain signal proved to be more effective than its magnetic tape recording. The role of the spectrum ultrasound part of the pain acoustic signal is discussed.     

Neuronal identification of signal periodicity by balanced inhibition

Many animals, including men, use periodicity information, e.g., amplitude modulations of acoustic stimuli, as a vital cue to auditory object formation. The underlying neuronal mechanisms, however, still remain a matter of debate. Here, we mathematically analyze a model for periodicity identification that relies on the interplay of excitation and delayed inhibition. Our analytical results show how the maximal response of such a system varies systematically with the time constants of excitation and inhibition. The model reliably identifies signal periodicity in the range from about ten to several hundred Hertz. Importantly, the model relies on biologically plausible parameters only. It works best for excitatory and inhibitory neuronal couplings of equal strength, the so-called ‘balanced inhibition’. We show how balanced inhibition can serve to identify low-frequency signal periodicity and how variation of a single parameter, the inhibitory time constant, can tune the system to different frequencies.     

Evolution of advertisement signals in North American hylid frogs: vocalizations as end-products of calling behavior

We studied the advertisement signals in two clades of North American hylid frogs in order to characterize the relationships between signal acoustic structure and underlying behavior. A mismatch was found between the acoustic structure and the mechanism of sound production. Two separate sets of phylogenetic characters were coded following acoustic versus mechanistic criteria, and exploratory treatments were made to compare their respective phylogenetic content in comparison with the molecular phylogeny ( Faivovich et al., 2005 ). We discuss the consequences of the acoustic/mechanistic mismatch in terms of significance of acoustic characters for phylogenetic and comparative studies; and the evolution of vocalizations in North American treefrogs. Considering only the acoustic structure of frog vocalizations can lead to misleading results in terms of both phylogenetic signal and evolution of vocalizations. In contrast, interpreting the acoustic signals with regard to the mechanism of sound production results in consistent phylogenetic information. The mechanistic coding also provides strong homologies for use in comparative studies of frog vocalizations, and to derive and test evolutionary hypotheses. © The Willi Hennig Society 2005.     

The Acoustic Properties of Low Intensity Vocalizations Match Hearing Sensitivity in the Webbed-Toed Gecko,Gekko subpalmatus

The design of acoustic signals and hearing sensitivity in socially communicating species would normally be expected to closely match in order to minimize signal degradation and attenuation during signal propagation. Nevertheless, other factors such as sensory biases as well as morphological and physiological constraints may affect strict correspondence between signal features and hearing sensitivity. Thus study of the relationships between sender and receiver characteristics in species utilizing acoustic communication can provide information about how acoustic communication systems evolve. The genus Gekko includes species emitting high-amplitude vocalizations for long-range communication (loud callers) as well as species producing only low-amplitude vocalizations when in close contact with conspecifics (quiet callers) which have rarely been investigated. In order to investigate relationships between auditory physiology and the frequency characteristics of acoustic signals in a quiet caller, Gekko subpalmatus we measured the subjects’ vocal signal characteristics as well as auditory brainstem responses (ABRs) to assess auditory sensitivity. The results show that G. subpalmatus males emit low amplitude calls when encountering females, ranging in dominant frequency from 2.47 to 4.17 kHz with an average at 3.35 kHz. The auditory range with highest sensitivity closely matches the dominant frequency of the vocalizations. This correspondence is consistent with the notion that quiet and loud calling species are under similar selection pressures for matching auditory sensitivity with spectral characteristics of vocalizations.     

Anechoic aquarium for ultrasonic neural telemetry

An acoustic neural telemetry tag has been developed for recording from free-swimming aquatic animals. Microwire electrodes were implanted into the VIIIth nerve of the toadfish, Opsanus tau, and interfaced to the subdermally implanted tag. The telemetry tag frequency modulates the neural signal, converting it into a varying frequency, which is amplified and transmitted acoustically (centre frequency of 90 kHz and a 20 kHz bandwidth). This acoustic signal is detected by a receiver hydrophone, and the receiver reconstructs the full neural waveform from the acoustic signal. However, due to the multipath environment in the experimental aquarium, the acoustic signal is quickly degraded as the hydrophone is moved away from the source. In order to receive the signal independent of fish position, an anechoic aquarium was designed. Streams of microbubbles (ca. 70 microm diameter) were generated to produce a curtain of sound-absorptive material along the walls and water surface of the aquarium. Microbubble generation significantly reduced the multipath artefacts, and allowed signal discrimination independent of fish and hydrophone position. The anechoic aquarium will allow the recording of neural activity from free-swimming fishes in quasi-natural habitats, thus allowing better understanding of the neural mechanisms of behaviour.     

Frequency range of the auditory effect of UHF

Zero beats of radiosound with an acoustic signal from an electrodynamic emitter in the frequency range up to 8 kHz were recorded in a natural experiment, which were not obtained earlier. It has been shown that the zero beats between the acoustic tonal signal and the first harmonics of the impulse succession of UHF are recorded in the points which correspond to low values on the threshold curve of UHF auditory effect.     

Multimodal communication in a noisy environment: a case study of the Bornean rock frog Staurois parvus

High background noise is an impediment to signal detection and perception. We report the use of multiple solutions to improve signal perception in the acoustic and visual modality by the Bornean rock frog, Staurois parvus. We discovered that vocal communication was not impaired by continuous abiotic background noise characterised by fast-flowing water. Males modified amplitude, pitch, repetition rate and duration of notes within their advertisement call. The difference in sound pressure between advertisement calls and background noise at the call dominant frequency of 5578 Hz was 8 dB, a difference sufficient for receiver detection. In addition, males used several visual signals to communicate with conspecifics with foot flagging and foot flashing being the most common and conspicuous visual displays, followed by arm waving, upright posture, crouching, and an open-mouth display. We used acoustic playback experiments to test the efficacy-based alerting signal hypothesis of multimodal communication. In support of the alerting hypothesis, we found that acoustic signals and foot flagging are functionally linked with advertisement calling preceding foot flagging. We conclude that S. parvus has solved the problem of continuous broadband low-frequency noise by both modifying its advertisement call in multiple ways and by using numerous visual signals. This is the first example of a frog using multiple acoustic and visual solutions to communicate in an environment characterised by continuous noise.     

Comfortable Loudness Level and Phrase Intelligibility of Spectrally Deprived Speech

It is known from the literature that (1) sounds with complex spectral composition are assessed by summing the partial outputs of the spectral channels; (2) electrical stimuli used in cochlear implant systems bring about the perception of a frequency band; and (3) removal of different parts of the auditory spectrum significantly affects phrase intelligibility. The level of acoustic pressure (AP) at a comfortable loudness level and the phrase intelligibility after comb filtering of a speech signal were measured in normally hearing subjects. Using a software program for spectral transformation of the speech signal, the phrase spectrum was divided into frequency bands of various width and only the bands with odd numbers were summed. In three series, the width of odd bands was 50, 100, or 150 Hz and the width of even bands was varied. The filter period was equal to the sum of the even and odd bands. With the same period, the acoustic pressure of the output signal should be increased to reach the comfortable loudness level of a speech signal passed via the comb filter; the narrower the width of the test bands, the higher the AP increase. With the same width of the test band, the acoustic pressure of the output signal should be increased to reach the comfortable loudness level; the greater the filter period, the higher the increase should be. The speech signal redundancy with respect to its spectral content can be equal to or even exceed 97.5%.     

The effect of an acoustic calling signal on the sex hormone level in the blood of white mice

The action was studied of synthesized acoustic signals, similar to the natural invocatory ones by their characteristics, on the level of sexual hormones in male and female mature laboratory mice, and also the dependence of these hormonal shifts on rhythmic organization of the artificial sound signals. As a result of the experiments it was convincingly shown that synthesized acoustic signal with the frequency of 3500 Hz (S-1) caused a functional load on the glands of internal secretion which was testified by statistically significant increase in the level of sexual hormones in the blood of males (by testosterone and extradiole) and female mice (by extradiole and progesterone). In order to clear up the role of rhythmic organization of signal studies were conducted with the application of synthetic acoustic signal (S-2), different from S-1. The analysis of the data revealed the absence of significant changes in the level of sexual hormones both in males and in females during S-2 sounding.