Involvement of a midbrain vocal nucleus in the production of both the acoustic and postural components of crowing behavior in Japanese quail

Many bird species produce vocalizations which are accompanied by distinctive postural displays, but the neural mechanisms that allow such integrated production of vocal and postural motor patterns are not understood. In the crowing behavior of Japanese quail, a characteristic vocal pattern is accompanied by and coordinated with a postural display that consists of a sequence of rapid, patterned head movements. The purpose of the present study was to investigate the role of a midbrain vocal nucleus, the nucleus intercollicularis, in the production of the acoustic and postural components of crowing in quail. Brief electrical stimuli were applied to the nucleus intercollicularis during spontaneously emitted crows in quail with chronically implanted electrodes, to determine if perturbing neural activity in the nucleus intercollicularis resulted in a disruption of ongoing crowing behavior. The most common effect of such stimuli was a concurrent, premature termination of both the acoustic and head movement components of the crow. These results imply that the nucleus intercollicularis plays a role in the production of both the acoustic and postural components of crowing in quail.     

Embryonic and posthatching treatments with sex steroids demasculinize the motivational aspects of crowing behavior in male Japanese quail

Demasculinizing action of embryonic estrogen on crowing behavior in male Japanese quails was examined. Eggs were treated with either 20 μg of estradiol benzoate (EB) or vehicle on the 10th day of incubation. Chicks hatched from both groups of eggs were injected daily with either testosterone propionate (TP; 10 μg/g b.w.), 5α-dihydrotestosterone (DHT, a non-aromatizable androgen; 10 μg/g b.w.), or vehicle from 11 to 50 days after hatching, and during this period their calling behaviors were observed. Irrespective of embryonic treatments, all birds received posthatching treatment with either TP or DHT, but not with vehicle, emitted crows in place of distress calls in a stress (non-sexual) context of being isolated in a recording chamber. The posthatching TP, but not posthatching DHT, induced crowing in a sexual context (crowing in their home-cages) from much earlier age than posthatching vehicle in the birds received control embryonic treatment with vehicle. The same TP treatment, however, completely eliminated the crowing in a sexual context in the birds received EB during their embryonic life. In the birds treated with either posthatching DHT or posthatching vehicle, the crowing in a sexual context was only slightly decreased by embryonic EB treatment. These data suggest that posthatching estrogen, derived from testosterone aromatization, enhances the demasculinizing action of embryonic estrogen, and thus strongly reduces the sexual motivation for crowing behavior. This demasculinizing action, however, would not influence vocal control system which generates acoustic pattern of crowing in the presence of androgens allowing the birds to crow in a non-sexual context.     

URBAN NOISE PREDICTS SONG FREQUENCY IN NORTHERN CARDINALS AND AMERICAN ROBINS

ABSTRACT

We examined the extent to which acoustic noise in urban environments influences song characteristics and singing behaviour of Northern Cardinals Cardinalis cardinalis and American Robins Turdus migratorius. We predicted that, in response to loud noise, birds would improve signal transmission by (1) increasing singing rate and (2) adjusting song characteristics such as pitch and length. From May—July 2006, 42 cardinals and 53 robins were recorded in forests located within four acoustic environments in central Ohio: rural, residential, commercial, and highway. Following each recording, we measured ambient noise level and recorded information describing location, weather, habitat, and conspecific presence within 75 m. As predicted, frequency range was positively correlated with noise level for both species, but neither song length nor rate was related to noise level for either species. These data support the idea that anthropogenic noise influences avian singing behaviour and acts as a selective force in urban areas.     

东方蝙蝠在交通噪声环境中的回声定位行为

环境噪声会降低动物声信号可探测性,是动物声信号进化的压力。为了避免人工噪声干扰,多数脊椎动物调整其叫声的频谱-时间结构,如采用延长叫声持续时间、提高频率和增加强度等策略。本研究选择回声定位声波频率范围与交通噪声频率范围相互重叠的东方蝙蝠(Vespertilio sinensis)为研究对象,在自然条件下开展实验,研究交通噪声对蝙蝠回声定位声波的影响。本实验选择交通噪声强度存在差异的两个样点,分别录制东方蝙蝠的回声定位声波,分析其结构参数。在交通噪声强度较高的样点,东方蝙蝠回声定位声波的持续时间、斜率均未显著改变,但起始频率、主频、终止频率及带宽均显著提高。结果表明,东方蝙蝠的回声定位行为在交通噪声干扰下具有明显的可塑性。     

Noise affects porpoise click detections – the magnitude of the effect depends on logger type and detection filter settings

Automatic click detectors and full-bandwidth sound recorders are widely used in passive acoustic monitoring of small cetaceans. Detection of these signals depends on a variety of factors, including signal to noise ratio. Passive acoustic monitoring is often used to study impact of underwater noise on small cetaceans, but as detection probability is affected by changes in signal to noise ratio, variable noise levels may affect conclusions drawn from these experiments. Therefore, we examine how different detectors and filters perform in varying ocean noise conditions. C-PODs and full-bandwidth recorders (Wildlife Acoustics, SM2M+) were deployed at two stations in an environment with fluctuating ambient noise for 42 days. Noise level and harbour porpoise (Phocoena phocoena) click trains simultaneously recorded on both loggers were compared. Overall, we found that porpoise click detections by the algorithm used to analyse full-band recorder data (Pamguard) paralleled detections by the C-POD. However, Pamguard detected significantly more clicks than the C-POD. A decrease in detections was seen for both loggers with increasing noise in the band 20 –160 kHz, in particular for levels above 100 dB re 1μPa rms. We also found that the Pamguard detection function changed the least over varying noise conditions when compared to the C-POD detectors. This study sheds light on the fact that inference of animal presence/absence or density that are based on echolocation cues (here, Porpoise Positive Minutes) shall account for the acoustic environments where probability of detecting signals may be affected by variability in ambient noise levels.     

Effects of androgens and estrogens on crowings and distress callings in male Japanese quail, Coturnix japonica

In male Japanese quail, crowing behavior is considered to be strictly androgen-dependent. It was previously shown that in chicks, treatment with either testosterone or 5alpha-dihydrotestosterone (5alpha-DHT; a non-aromatizable androgen) induced crowing with motivation for distress calling in acutely isolated conditions. Many studies, however, have shown that the potencies of testosterone and 5alpha-DHT in activating crowing in castrated males are different. To clarify the effects of androgenic and estrogenic actions on the production of crows and distress calls, we injected quail daily from 11 to 42 days after hatching (Day 11 to 42) with testosterone propionate (TP), 5alpha-DHT, estradiol benzoate (EB) or vehicle and examined their calling behaviors both in a recording chamber (acutely isolated conditions) and in their home-cages (well-acclimated conditions). Both TP- and 5alpha-DHT-treated birds began to crow by Day 13 when isolated in the recording chamber. The TP-treated birds, however, crowed less frequently than 5alpha-DHT-treated ones. This, combined with the observations that distress calling was strongly inhibited in EB-treated birds, suggests that estrogen converted from testosterone may inhibit the motivation for distress calling. On the other hand, after chronic treatment of TP, but not of 5alpha-DHT, birds began to crow intensely in their home-cages earlier than vehicle treated controls, suggesting that estrogen is needed to initiate crowing behavior in sexually active males. Taken together, it is suggested that estrogenic actions affect the motivation underlying vocal behaviors, while the androgenic action is indispensable in generating crowing.     

Ambient noise and the design of begging signals

The apparent extravagance of begging displays is usually attributed to selection for features, such as loud calls, that make the signal costly and hence reliable. An alternative explanation, however, is that these design features are needed for effective signal transmission and reception. Here, we test the latter hypothesis by examining how the begging calls of tree swallow (Tachycineta bicolor) nestlings and the response to these calls by parents are affected by ambient noise. In a field study, we found that call length, amplitude and frequency range all increased with increasing noise levels at nests. In the laboratory, however, only call amplitude increased in response to the playback of noise to nestlings. In field playbacks to parents, similar levels of noise abolished parental preferences for higher call rates, but the preference was restored when call amplitude was increased to the level that nestlings had used in the laboratory study. Our results show that nestling birds, like other acoustic signallers, consistently increase call amplitude in response to ambient noise and this response appears to enhance discrimination by receivers. Thus, selection for signal efficacy may explain some of the seemingly extravagant features of begging displays.     

Interspecific Hybridization as a Tool to Understand Vocal Divergence: The Example of Crowing in Quail (Genus Coturnix)

Understanding the mechanisms that lead organisms to be separated into distinct species remains a challenge in evolutionary biology. Interspecific hybridization, which results from incomplete reproductive isolation, is a useful tool to investigate such mechanisms. In birds, interspecific hybridization is relatively frequent, despite the fact that closed species exhibit morphological and behavioural differences. Evolution of behaviour is difficult to investigate on a large timescale since it does not ‘fossilize’. Here I propose that calls of hybrid non-songbirds that develop without the influence of learning may help in understanding the gradual process that leads to vocal divergence during speciation. I recorded crows produced by the European quail (Coturnix c. coturnix), the domestic Japanese quail (Coturnix c. japonica) and their hybrids (F1, F2 and backcrosses). Most crowing patterns were intermediate to those of the parental species; some were similar to one or the other parental species, or not present in either parental species. I also observed vocal changes in hybrid crows during the breeding season and from one year to the other. This vocal variability resembles those observed during the ontogeny of the crow in quails. It is likely that similar mechanisms involved in vocal changes during ontogeny might have driven vocal divergence in the species of Palearctic quails. I suggest that hybrid crows might have resembled those produced by intermediary forms of quails during speciation.     

The role of chromatic and achromatic signals for fruit detection by birds

Fruit color changes during ripening are typically viewed asan adaptation to increase signal efficacy to seed dispersers.Plants can increase signal efficacy by enhancing chromatic (wavelengthrelated) and/or achromatic (intensity related) contrasts betweenfruit and background. To assess how these contrasts determinethe detectability of fruit signals, we conducted 2 experimentswith free-flying crows (Corvus ossifragus) under seminaturalconditions in a 2025 m² aviary. Crows searched first for artificialred and black fruits and detected red fruits from a larger distance.Because artificial red fruits had higher chromatic and lowerachromatic contrasts against foliage than artificial black fruits,crows apparently prioritized chromatic contrasts. Thus, thecommon change in fruit color from red to black during ripeningdoes not increase signal efficacy to crows. In a second trial,crows searched for UV-reflecting and black blueberries (Vaccinummyrtillus) against backgrounds of foliage and sand. Againstfoliage, UV-reflecting berries had higher chromatic and achromaticcontrasts than black berries, and crows detected them from alarger distance. Against sand, UV-reflecting berries had lowachromatic contrasts and black berries low chromatic contrasts.Crows detected both fruit types equally, suggesting that theyused chromatic contrasts to detect UV-reflecting berries andachromatic contrasts to detect black berries. Birds prioritizedchromatic contrasts when searching for artificial red fruitsin foliage but not when searching for blueberries on sand. Wesuggest that the relative importance of chromatic and achromaticcontrasts is contingent on the chromatic and achromatic varianceof the background. Models of signal perception can be improvedby incorporating background-specific effects.     

Signal/noise analysis of FRET-based sensors

Molecular sensors based on intramolecular Förster resonance energy transfer (FRET) have become versatile tools to monitor regulatory molecules in living tissue. However, their use is often compromised by low signal strength and excessive noise. We analyzed signal/noise (SNR) aspects of spectral FRET analysis methods, with the following conclusions: The most commonly used method (measurement of the emission ratio after a single short wavelength excitation) is optimal in terms of signal/noise, if only relative changes of this uncalibrated ratio are of interest. In the case that quantitative data on FRET efficiencies are required, these can be calculated from the emission ratio and some calibration parameters, but at reduced SNR. Lux-FRET, a recently described method for spectral analysis of FRET data, allows one to do so in three different ways, each based on a ratio of two out of three measured fluorescence signals (the donor and acceptor signal during a short-wavelength excitation and the acceptor signal during long wavelength excitation). Lux-FRET also allows for calculation of the total abundance of donor and acceptor fluorophores. The SNR for all these quantities is lower than that of the plain emission ratio due to unfavorable error propagation. However, if ligand concentration is calculated either from lux-FRET values or else, after its calibration, from the emission ratio, SNR for both analysis modes is very similar. Likewise, SNR values are similar, if the noise of these quantities is related to the expected dynamic range. We demonstrate these relationships based on data from an Epac-based cAMP sensor and discuss how the SNR changes with the FRET efficiency and the number of photons collected.     

Contrasting effects of pre- and posthatch exposure to gonadal steroids on the development of vocal, sexual, and aggressive behavior of young domestic fowl

Prehatch treatment of domestic chicks with 17 beta-estradiol dipropionate (EDP) reduces later testosterone-facilitated sexual behavior, crowing, and suppression of peeping. Prehatch treatment with 5 alpha-dihydrotestosterone propionate (DHTP) reduces later testosterone-facilitated aggressive behavior and produces greatly enhanced avoidance that interferes with sexual behavior in some test situations. It also disturbs crowing by removing trilled call elements and increasing the number of elements/crow but has little effect on crowing posture. The majority of these results can be understood as "demasculinizing" effects of gonadal steroids on the developing male fetus, but the possibility that some result from more extreme pathological changes is also discussed. Posthatch treatment with DHTP facilitates attack and cackling but has no effect on sexual behavior or crowing; the synthetic androgen R1881 has similar effects on cackling and also has no effect on sexual behavior or crowing. Posthatch treatment with EDP enhances sexual behavior but has no effect on attack; the aromatase inhibitor 1,4,6-Androstatrien-3,17-dione (ATD) blocked the suppression of peeping by testosterone but had no effect on testosterone-facilitated sexual behavior. Combined posthatch treatment with EDP and DHTP facilitated crowing although treatment with either steroid alone was ineffective.     

环境噪声对蝙蝠回声定位叫声及飞行活动的影响

环境噪声影响动物的活动及其叫声特性,已成为动物面对的一种重要选择压力。为应对噪声的干扰,多数动物类群会远离噪声区域和改变其叫声的频谱时间结构,如延长叫声持续时间、提高叫声频率等,但有些动物的活动和叫声频谱时间结构并不受环境噪声的影响。本研究在自然条件下,研究不同环境噪声强度对蝙蝠活动和回声定位声波的影响。选取噪声强度有差异的12个样点,分别录制各样点大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的回声定位声波,分析其持续时间、起始频率、峰频、终止频率和带宽,统计蝙蝠通过次数。回归分析结果显示：环境噪声强度与大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的活动无显著相关性P > 0.05）,与回声定位声波的脉冲持续时间、起始频率、峰频、终止频率及带宽均不显著相关（P > 0.05）。暗示低频低强度（< 20 kHz, < 67.5 dB）的环境噪声可能对高频回声定位蝙蝠的叫声及活动没有显著影响。     

The amplitude sensitivity of mouse inferior collicular neurons in the presence of weak noise

Natural auditory environment consists of multiple sound sources that are embedded in ambient strong and weak noise. For effective sound communication and signal analysis, animals must somehow extract biologically relevant signals from the inevitable interference of ambient noise. The present study examined how a weak noise may affect the amplitude sensitivity of neurons in the mouse central nucleus of the inferior colliculus (IC) which receives convergent excitatory and inhibitory inputs from both lower and higher auditory centers. Specifically, we studied the amplitude sensitivity of IC neurons using a probe (best frequency pulse) and a masker (weak noise) under simultaneous masking paradigm. For most IC neurons, weak noise masking increases the minimum threshold and decreases the number of impulses. Noise masking also increased the slope and decreased the dynamic range of the rate amplitude function of these IC neurons. The strength of this noise masking was greater at low than at high sound amplitudes. This variation in the amplitude sensitivity of IC neurons in the presence of the weak noise was mostly mediated through GABAergic inhibition. These data indicate that in the real world the ambient weak noise improves amplitude sensitivity of IC neurons through GABAergic inhibition while inevitably decreases the range of overall auditory sensitivity of IC neurons.     

Regulation of vocal amplitude by the blue-throated hummingbird, Lampornis clemenciae

Animals that rely on vocal communication must broadcast sound so that a perceptible signal is transmitted over an appropriate distance. We found that male blue-throated hummingbirds modified the amplitude of their vocalizations in response to both naturally occurring and experimenter-controlled changes in ambient noise levels. This phenomenon is known as the Lombard effect and may increase the efficiency of acoustic signalling. This study demonstrates the effect under natural field conditions and documents the first hummingbird species (Apodiformes: Trochilidae) to show this behaviour. We measured sound pressure levels (SPLs) of Serial Chip territorial advertisement calls across a natural range of ambient noise, primarily due to creeks within male territories. We found a significant correlation between the amplitude of Serial Chips and the amplitude of background noise. To test this relationship, we broadcast recordings of creek noise at high and low amplitudes while target individuals were producing Serial Chip vocalizations. We measured vocal SPLs before and during the playback. Individuals responded to changes in playback creek noise by changing the amplitude of Serial Chip production. We also measured transmission properties of Serial Chip calls through natural habitat to calculate the approximate amplitude of vocalizations at the position of the calling bird. We suggest that amplitude regulation of vocalizations contibutes to signal transmission distance along with the established relationships between singing behaviour, acoustic structure and habitat. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.     

背景噪声对人感知声音时间信息的影响

对声音时间信息的分辨在人和动物感知声音信息的过程中至关重要.在自然声环境中,声音信息总处于一定的噪声背景下.文章以间隔探测阈值为指标测定了人对纯音和噪声的间隔探测阈值,以及持续噪声背景对间隔探测阈值的影响.声音信号采用1000～10000 Hz的纯音信号和白噪声信号,声音强度为70 dB SPL.背景噪声为持续白噪声,强度分别为45、55、65 dB SPL.结果表明,对纯音信号,随着背景噪声强度增加,间隔探测阈值有升高的趋势.对噪声信号来说,45、55 dB SPL的背景噪声对噪声信号的间隔探测阈值无显著影响,但65 dB SPL的背景噪声使间隔探测阈值显著升高.研究结果提示,背景噪声能够在一定程度上影响人对声音时间信息的感知,影响的程度与背景噪声的强度有关.     

Free-field unmasking response characteristics of frog auditory nerve fibers: comparison with the responses of midbrain auditory neurons

Previous studies in the inferior colliculus have shown that spatial separation of signal and noise sources improves signal detection. In this study, we investigated the free-field unmasking response properties of single fibers in the auditory nerve--these were compared to those of inferior colliculus neurons under the same experimental conditions to test the hypothesis that central processing confers advantages for signal detection in the presence of spatially separated noise. For each neuron, we determined the detection threshold for a probe at the unit's best azimuth under three conditions: (1) by itself, (2) when a masker at a constant level was also presented at the unit's best azimuth, and (3) when the masker was positioned at different azimuths. We found that, on average, maskers presented at a unit's best azimuth elevated the probe detection threshold by 4.22 dB in the auditory nerve and 10.97 dB in the inferior colliculus. Angular separation of probe and masker sources systematically reduced the masking effect. The maximum masking release was on average 2.90 dB for auditory nerve fibers and 9.40 dB for inferior colliculus units. These results support the working hypothesis, suggesting that central processing contributes to the stronger free-field unmasking in the inferior colliculus.     

Djungarian hamsters exhibit temperature-dependent dietary fat choice in long days

Previous studies have shown that whole-animal thermal responses of ectotherms and heterotherms (e.g., hibernators), both of which experience a wide range of body temperatures, are related to the saturation level of somatic lipids, which in turn can be influenced by the ratio of saturated and unsaturated fatty acids in the diet. This study demonstrates that Djungarian hamsters held in long days display ambient temperature-dependent choice of dietary fats, increasing their preference for saturated fats when ambient temperature increases (to 27 degrees C) and later reversing this preference when ambient temperature is returned to its original value (8 degrees C). Changes in percent contribution of the unsaturated and saturated diets in response to temperature were accomplished almost solely by changes in the amount of unsaturated diet consumed. Temperature-dependent fatty acid choice occurs at a stage in the annual cycle when Djungarian hamsters do not enter spontaneous daily torpor and therefore experience only small changes in core body temperature. These results suggest that temperature-dependent fatty acid choice may occur in a wide range of animals, including nonheterothermic endotherms.     

Whistling in a noisy ocean: bottlenose dolphins adjust whistle frequencies in response to real-time ambient noise levels

Common bottlenose dolphins (Tursiops truncatus) use complex acoustic behaviours for communication, group cohesion and foraging. Ambient noise from natural and anthropogenic sources has implications for the acoustic behaviour of dolphins, and research shows that average ambient noise levels alter dolphin acoustic behaviour. However, when background noise levels are highly variable, the relationships between noise and acoustic behaviour over short time periods are likely important. This study investigates whether bottlenose dolphins altered the temporal and spectral qualities of their whistles in relation to the ambient noise present at the time the whistles were produced. Dolphin groups were recorded in Tampa Bay (western Florida) between 2008 and 2015. Six whistle parameters were analysed in spectrogram software (minimum frequency, maximum frequency, bandwidth, peak frequency, duration and number of inflection points) and ambient noise levels were calculated immediately prior to each whistle. Linear regression analysis indicated that the minimum, maximum and peak frequencies of whistles had significant positive relationships with the ambient noise levels present at the time of the whistles. These models suggested that for each 1 dB increase in ambient noise, minimum frequency increased by 121 Hz, maximum frequency increased by 108 Hz and peak frequency increased by between 122 and 144 Hz. As ambient noise is typically low frequency, this suggests that bottlenose dolphins increased whistle frequency in response to real-time noise levels to avoid masking. Future research to determine the fitness consequences of noise-induced changes in the communication behaviour of dolphins would be an important contribution to conservation efforts.     

Communication in a noisy environment: short-term acoustic adjustments and the underlying acoustic niche of a Neotropical stream-breeding frog

Acoustically active animals may show long- and short-term adaptations in acoustic traits for coping with ambient noise. Given the key role of calls in anurans’ life history, long- and short-term adaptations are expected in species inhabiting noisy habitats. However, to disentangle such adaptations is a difficult task, incipiently addressed for Neotropical frogs. We investigated if males of a stream-breeding frog (Crossodactylus schmidti) adjust call traits according to the background noise, and if the signal-to-noise ratio (SNR) varies between call harmonics and along call notes. We measured sound pressure levels of calls and noise in the field and used a fine-scale acoustic analysis to describe the signal and noise structure and test for noise-related call adjustments. The multi-note harmonic call of C. schmidti greatly varied in the spectral structure, including a trend for increasing note amplitude along the call, a wide frequency bandwidth of the 2nd harmonic, a minor call frequency modulation due to a trend for increasing note frequency within the same harmonic, and a major call frequency modulation due to the variable location of the dominant harmonic along the call. Calls had significantly higher frequencies than the noise at the range of the 1st and the 2nd call harmonics, and significantly louder sound pressure than the noise at the range of all harmonics. Males emitted the majority of call notes showing positive SNR, and though males also emitted some notes with negative SNR, when a given harmonic was negative the other harmonics in the same note did not tend to be SNR-negative. Our results indicate that male C. schmidti show short-term acoustic adjustments that make the advertisement call effective for coping with the interference of the stream-generated noise. We suggest that the call spectral plasticity serves for coping with temporary changes in the background noise, whilst we also discuss the possibility that the redundant, harmonic-structured call may have evolved to diminish masking interference on the acoustic signal by the background noise. This is the first study to uncouple noise-related acoustic adjustments and putative long-term acoustic adaptations for a Hylodidae, providing insights on behavioral plasticity and signal evolution of stream-breeding frogs.     

Ability to alter song in two grassland songbirds exposed to simulated anthropogenic noise is not related to pre-existing variability

Organisms encounter noise naturally in the environment. However, increasing prevalence of human-caused noise seems to be resulting in behavioural changes in many animals that can affect survival and reproduction. Not all species react the same way to noise; some adjust their vocal signals while others do not. We hypothesized that species with more variability in their vocal signals would be better able to adjust their signals to be audible over anthropogenic noise. We tested this within a large-scale manipulative experiment by recording songs of two grassland songbirds, Baird’s sparrow (Ammodramus bairdii) and Savannah sparrow (Passerculus sandwichensis), both of which are found in areas increasingly affected by energy extraction noise. We compared these species because Savannah sparrows have more variability in their songs geographically and temporally compared to Baird’s sparrows. We recorded both species’ songs before, during and after high-fidelity playbacks of oil well drilling noise. Surprisingly, both species changed parts of their songs in the presence of noise (Baird’s sparrow usually decreasing frequency and Savannah sparrow increasing frequency) and these changes were not related to seasonal, song, or syllable variability. We suggest instead that acoustically heterogeneous environments may favour the evolution of species that are capable of adjusting their songs in response to variable ambient noise.