城市噪声使树麻雀鸣唱的最低频率升高

随着城市化进程的不断加快,城市噪声水平显著升高。噪声会掩盖鸟类的声音信号,这无疑会影响鸟类的交流。在嘈杂的城市环境中,鸟类通常以高频率鸣唱来避免声信号被掩蔽。然而,较低的发声频率才是雄性品质的重要表征,提高发声频率可能会影响声信号对雌性的吸引力。因此,鸟类会在提高发声频率和保持较低频率之间进行权衡。为确定城市噪声对树麻雀（Passer montanus）鸣唱行为的影响,在沈阳市选取6个研究地点,比较了沈阳市区和近郊不同噪声水平下树麻雀繁殖期的鸣唱特征。在2019年4月至7月,使用定向麦克风录制了320只繁殖期树麻雀的鸣唱,并使用声级计测定噪声水平。研究结果显示,城市研究地点的噪声水平显著高于郊区研究地点。与安静郊区相比,城市嘈杂环境中的树麻雀鸣唱的最高频率、最低频率和主峰峰频显著较高,频宽更大,而时长没有差别。树麻雀鸣唱的最高频率、频宽、主峰峰频和时长均与噪声水平无显著相关,而最低频率与研究地点的噪声水平呈显著正相关。上述结果说明,在噪声环境中,树麻雀选择提高最低频率以利于声信号的传输。     

A field test of the audibility of urban versus rural songs in mountain chickadees

Anthropogenic noise can mask avian vocalizations, and several urban‐dwelling species adjust frequency or amplitude of vocalizations in ways that appear to compensate for increased noise levels. Playback studies have investigated whether receivers differentiate between signals produced by rural and urban males, but it is difficult to determine whether differential response to stimulus reflects differences in audibility versus perceived differences in male signals/condition that result from urban settlement. Here, we performed paired‐playback trials to determine whether mountain chickadees (Poecile gambeli) differentiate urban versus rural songs when both stimuli were broadcast within noise. For each playback, stimuli were played in short bouts starting at a low signal‐to‐noise ratio and increasing in relative amplitude with each successive bout. If the primary function of adjusted urban songs is propagation in noise, we hypothesized that focal males would respond sooner to urban versus rural playbacks (detect at lower signal‐to‐noise ratios). If urban songs solely encode information about the male's condition (such as increased aggression), then we predicted only a differential aggressive response to playbacks once detected. If urban songs both increase propagation and embed information on male condition, we predicted a combination of both response types. We found no difference in latency to first response in urban versus rural songs, but some evidence for differential aggression to playback dependent on both stimulus type (urban vs. rural) and local ambient noise levels; focal males in noisy (urban) sites responded aggressively to both stimulus types, whereas focal males in quiet (rural) sites responded more aggressively to urban than to rural stimuli. This context‐dependent discrimination may be the result of increased aggression in urban habitats, improved communication in noisy habitats by urban signallers and receivers, or some combination of the two.     

Negative impact of urban noise on sexual receptivity and clutch size in female domestic canaries

In oscines, male song stimulates female reproduction and females are known to adjust both their sexual preferences and their maternal investment according to song quality. Female domestic canaries are especially responsive to wide frequency bandwidth (4 kHz) male songs emitted with a high‐repetition syllable rate and low minimal frequencies (1 kHz). We previously showed that low‐frequency urban noise decreases female sexual responsiveness for these low‐frequency songs (1–5 kHz) through auditory masking. Based on the differential allocation hypothesis, we predicted that urban noise exposure will equally affect female maternal investment. Using a crossover design, we broadcast low‐frequency songs to females either in an overlapping noise condition or in an alternating noise condition. Females decreased both their sexual responsiveness and their clutch size in the overlapping noise treatment relative to the alternative noise treatment. No differences were found concerning egg size or egg composition (yolk and albumen mass, testosterone concentration). Due to our experimental design, we can exclude a general impact of noisy conditions and thereby provide evidence for a detrimental effect through masking on avian courtship and reproductive output. These results suggest that noisy conditions may also affect avian communication in outdoor conditions, which may partly explain field reports on noise‐dependent breeding success and reduced breeding densities at noisy sites.     

Anthropogenic noise decreases urban songbird diversity and may contribute to homogenization

More humans reside in urban areas than at any other time in history. Protected urban green spaces and transportation greenbelts support many species, but diversity in these areas is generally lower than in undeveloped landscapes. Habitat degradation and fragmentation contribute to lowered diversity and urban homogenization, but less is known about the role of anthropogenic noise. Songbirds are especially vulnerable to anthropogenic noise because they rely on acoustic signals for communication. Recent studies suggest that anthropogenic noise reduces the density and reproductive success of some bird species, but that species which vocalize at frequencies above those of anthropogenic noise are more likely to inhabit noisy areas. We hypothesize that anthropogenic noise is contributing to declines in urban diversity by reducing the abundance of select species in noisy areas, and that species with low‐frequency songs are those most likely to be affected. To examine this relationship, we calculated the noise‐associated change in overall species richness and in abundance for seven common songbird species. After accounting for variance due to vegetative differences, species richness and the abundance of three of seven species were reduced in noisier locations. Acoustic analysis revealed that minimum song frequency was highly predictive of a species' response to noise, with lower minimum song frequencies incurring greater noise‐associated reduction in abundance. These results suggest that anthropogenic noise affects some species independently of vegetative conditions, exacerbating the exclusion of some songbird species in otherwise suitable habitat. Minimum song frequency may provide a useful metric to predict how particular species will be affected by noise. In sum, mitigation of noise may enhance habitat suitability for many songbird species, especially for species with songs that include low‐frequency elements.     

Urban noise and the cultural evolution of bird songs

In urban environments, anthropogenic noise can interfere with animal communication. Here we study the influence of urban noise on the cultural evolution of bird songs. We studied three adjacent dialects of white-crowned sparrow songs over a 30-year time span. Urban noise, which is louder at low frequencies, increased during our study period and therefore should have created a selection pressure for songs with higher frequencies. We found that the minimum frequency of songs increased both within and between dialects during the 30-year time span. For example, the dialect with the highest minimum frequency is in the process of replacing another dialect that has lower frequency songs. Songs with the highest minimum frequency were favoured in this environment and should have the most effective transmission properties. We suggest that one mechanism that influences how dialects, and cultural traits in general, are selected and transmitted from one generation to the next is the dialect''s ability to be effectively communicated in the local environment.     

Flexibility in animal signals facilitates adaptation to rapidly changing environments

Charles Darwin posited that secondary sexual characteristics result from competition to attract mates. In male songbirds, specialized vocalizations represent secondary sexual characteristics of particular importance because females prefer songs at specific frequencies, amplitudes, and duration. For birds living in human-dominated landscapes, historic selection for song characteristics that convey fitness may compete with novel selective pressures from anthropogenic noise. Here we show that black-capped chickadees (Poecile atricapillus) use shorter, higher-frequency songs when traffic noise is high, and longer, lower-frequency songs when noise abates. We suggest that chickadees balance opposing selective pressures by use low-frequency songs to preserve vocal characteristics of dominance that repel competitors and attract females, and high frequency songs to increase song transmission when their environment is noisy. The remarkable vocal flexibility exhibited by chickadees may be one reason that they thrive in urban environments, and such flexibility may also support subsequent genetic adaptation to an increasingly urbanized world.     

Urban noise influences vocalization structure in the House Wren Troglodytes aedon

Urban habitats are noisy and constrain acoustic communication in birds. We analysed the effect of anthropogenic noise on the vocalization characteristics of House Wrens Troglodytes aedon at two sites with different noise levels (rural and urban). We measured in each song and song trill the frequency bandwidth, maximum amplitude, highest and minimum frequency, and trill rate. In noisy urban environments, there was a reduction in bandwidth and an increase in trill rate relative to quieter, rural environments. The whole song of birds from both populations increased in minimum frequency as noise increased, improving song transmission.     

Geographically pervasive effects of urban noise on frequency and syllable rate of songs and calls in silvereyes (Zosterops lateralis)

Recent studies in the Northern Hemisphere have shown that songbirds living in noisy urban environments sing at higher frequencies than their rural counterparts. However, several aspects of this phenomenon remain poorly understood. These include the geographical scale over which such patterns occur (most studies have compared local populations), and whether they involve phenotypic plasticity or microevolutionary change. We conducted a field study of silvereye (Zosterops lateralis) vocalizations over more than 1 million km(2) of urban and rural south-eastern Australia, and compared possible effects of urban noise on songs (which are learned) and contact calls (which are innate). Across 14 paired urban and rural populations, silvereyes consistently sang both songs and contact calls at higher frequencies in urban environments. Syllable rate (syllables per second) decreased in urban environments, consistent with the hypothesis that reflective structures degrade song and encourage longer intervals between syllables. This comprehensive study is, to our knowledge, the first to demonstrate varied adaptations of urban bird vocalizations over a vast geographical area, and to provide insight into the mechanism responsible for these changes.     

Different behavioural responses to anthropogenic noise by two closely related passerine birds

Anthropogenic noise, now common to many landscapes, can impair acoustic communication for many species, yet some birds compensate for masking by noise by altering their songs. The phylogenetic distribution of these noise-dependent signal adjustments is uncertain, and it is not known whether closely related species respond similarly to noise. Here, we investigated the influence of noise on habitat occupancy rates and vocal frequency in two congeneric vireos with similar song features. Noise exposure did not influence occupancy rates for either species, yet song features of both changed, albeit in different ways. With increases in noise levels, plumbeous vireos (Vireo plumbeus) sang shorter songs with higher minimum frequencies. By contrast, grey vireos (Vireo vicinior) sang longer songs with higher maximum frequencies. These findings support the notion that vocal plasticity may help some species occupy noisy areas, but because there were no commonalities among the signal changes exhibited by these closely related birds, it may be difficult to predict how diverse species may modify their signals in an increasingly noisy world.     

Why longer song elements are easier to detect: threshold level-duration functions in the Great Tit and comparison with human data

Our study estimates detection thresholds for tones of different durations and frequencies in Great Tits (Parus major) with operant procedures. We employ signals covering the duration and frequency range of communication signals of this species (40–1,010 ms; 2, 4, 6.3 kHz), and we measure threshold level-duration (TLD) function (relating threshold level to signal duration) in silence as well as under behaviorally relevant environmental noise conditions (urban noise, woodland noise). Detection thresholds decreased with increasing signal duration. Thresholds at any given duration were a function of signal frequency and were elevated in background noise, but the shape of Great Tit TLD functions was independent of signal frequency and background condition. To enable comparisons of our Great Tit data to those from other species, TLD functions were first fitted with a traditional leaky-integrator model. We then applied a probabilistic model to interpret the trade-off between signal amplitude and duration at threshold. Great Tit TLD functions exhibit features that are similar across species. The current results, however, cannot explain why Great Tits in noisy urban environments produce shorter song elements or faster songs than those in quieter woodland environments, as detection thresholds are lower for longer elements also under noisy conditions.     

Effects of urban noise on song and response behaviour in great tits

Acoustic communication is fundamental in avian territory defence and mate attraction. In urban environments where sound transmissions are more likely to be masked by low-frequency anthropogenic noise, acoustic adaptations may be advantageous. However, minor modifications to a signal could affect its efficacy. While recent research has shown that there is divergence between songs from noisy and quiet areas, it is unknown whether these differences affect the response to the signal by its receivers. Here, we show that there is a difference in spectral aspects of rural and urban song in a common passerine, the great tit Parus major, at 20 sites across the UK. We also provide, to our knowledge, the first demonstration that such environmentally induced differences in song influence the response of male territory holders. Males from quiet territories exhibited a significantly stronger response when hearing song from another territory holder with low background noise than from those with high background noise. The opposite distinction in response intensity to homotypic versus heterotypic song was observed in males from noisy territories. This behavioural difference may intensify further signal divergence between urban and rural populations and raises important questions concerning signal evolution.     

Song characteristics track bill morphology along a gradient of urbanization in house finches (<Emphasis Type="Italic">Haemorhous mexicanus</Emphasis>)

Introduction

Urbanization can considerably impact animal ecology, evolution, and behavior. Among the new conditions that animals experience in cities is anthropogenic noise, which can limit the sound space available for animals to communicate using acoustic signals. Some urban bird species increase their song frequencies so that they can be heard above low-frequency background city noise. However, the ability to make such song modifications may be constrained by several morphological factors, including bill gape, size, and shape, thereby limiting the degree to which certain species can vocally adapt to urban settings. We examined the relationship between song characteristics and bill morphology in a species (the house finch, Haemorhous mexicanus) where both vocal performance and bill size are known to differ between city and rural animals.

Results

We found that bills were longer and narrower in more disturbed, urban areas. We observed an increase in minimum song frequency of urban birds, and we also found that the upper frequency limit of songs decreased in direct relation to bill morphology.

Conclusions

These findings are consistent with the hypothesis that birds with longer beaks and therefore longer vocal tracts sing songs with lower maximum frequencies because longer tubes have lower-frequency resonances. Thus, for the first time, we reveal dual constraints (one biotic, one abiotic) on the song frequency range of urban animals. Urban foraging pressures may additionally interact with the acoustic environment to shape bill traits and vocal performance.

     

Patterns of Song across Natural and Anthropogenic Soundscapes Suggest That White-Crowned Sparrows Minimize Acoustic Masking and Maximize Signal Content

Soundscapes pose both evolutionarily recent and long-standing sources of selection on acoustic communication. We currently know more about the impact of evolutionarily recent human-generated noise on communication than we do about how natural sounds such as pounding surf have shaped communication signals over evolutionary time. Based on signal detection theory, we hypothesized that acoustic phenotypes will vary with both anthropogenic and natural background noise levels and that similar mechanisms of cultural evolution and/or behavioral flexibility may underlie this variation. We studied song characteristics of white-crowned sparrows (Zonotrichia leucophrys nuttalli) across a noise gradient that includes both anthropogenic and natural sources of noise in San Francisco and Marin counties, California, USA. Both anthropogenic and natural soundscapes contain high amplitude low frequency noise (traffic or surf, respectively), so we predicted that birds would produce songs with higher minimum frequencies in areas with higher amplitude background noise to avoid auditory masking. We also anticipated that song minimum frequencies would be higher than the projected lower frequency limit of hearing based on site-specific masking profiles. Background noise was a strong predictor of song minimum frequency, both within a local noise gradient of three urban sites with the same song dialect and cultural evolutionary history, and across the regional noise gradient, which encompasses 11 urban and rural sites, several dialects, and several anthropogenic and natural sources of noise. Among rural sites alone, background noise tended to predict song minimum frequency, indicating that urban sites were not solely responsible for driving the regional pattern. These findings support the hypothesis that songs vary with local and regional soundscapes regardless of the source of noise. Song minimum frequency from five core study sites was also higher than the lower frequency limit of hearing at each site, further supporting the hypothesis that songs vary to transmit through noise in local soundscapes. Minimum frequencies leveled off at noisier sites, suggesting that minimum frequencies are constrained to an upper limit, possibly to retain the information content of wider bandwidths. We found evidence that site noise was a better predictor of song minimum frequency than territory noise in both anthropogenic and natural soundscapes, suggesting that cultural evolution rather than immediate behavioral flexibility is responsible for local song variation. Taken together, these results indicate that soundscapes shape song phenotype across both evolutionarily recent and long-standing soundscapes.     

Vocal frequency change reflects different responses to anthropogenic noise in two suboscine tyrant flycatchers

Anthropogenic noise is prevalent across the globe and can exclude birds from otherwise suitable habitat and negatively influence fitness; however, the mechanisms responsible for species' responses to noise are not always clear. One effect of noise is a reduction in effective acoustic communication through acoustic masking, yet some urban songbirds may compensate for masking by noise through altering their songs. Whether this vocal flexibility accounts for species persistence in noisy areas is unknown. Here, we investigated the influence of noise on habitat use and vocal frequency in two suboscine flycatchers using a natural experiment that isolated effects of noise from confounding stimuli common to urban habitats. With increased noise exposure, grey flycatcher (Empidonax wrightii) occupancy declined, but vocal frequency did not change. By contrast, ash-throated flycatcher (Myiarchus cinerascens) occupancy was uninfluenced by noise, but individuals in areas with greater noise amplitudes vocalized at a higher frequency, although the increase (≈200 kHz) may only marginally improve communication and may represent a secondary effect from increased vocal amplitude. Even so, the different flycatcher behavioural responses suggest that signal change may help some species persist in noisy areas and prompt important questions regarding which species will cope with an increasingly noisy world.     

Degradation of rural and urban great tit song: testing transmission efficiency

Acoustic signals play a fundamental role in avian territory defence and mate attraction. Several studies have now shown that spectral properties of bird song differ between urban and rural environments. Previously this has been attributed to competition for acoustic space as a result of low-frequency noise present in cities. However, the physical structure of urban areas may have a contributory effect. Here we investigate the sound degradation properties of woodland and city environments using both urban and rural great tit song. We show that although urban surroundings caused significantly less degradation to both songs, the transmission efficiency of rural song compared to urban song was significantly lower in the city. While differences between the two songs in woodland were generally minimal, some measures of the transmission efficiency of rural song were significantly lower than those of urban song, suggesting additional benefits to singing rural songs in this setting. In an attempt to create artificial urban song, we mimicked the increase in minimum frequency found several times previously in urban song. However, this did not replicate the same transmission properties as true urban song, suggesting changes in other song characteristics, such as temporal adjustments, are needed to further increase transmission of an avian signal in the city. We suggest that the structure of the acoustic environment, in addition to the background noise, plays an important role in signal adaptation.     

Male Songbird Indicates Body Size with Low-Pitched Advertising Songs

Body size is a key sexually selected trait in many animal species. If size imposes a physical limit on the production of loud low-frequency sounds, then low-pitched vocalisations could act as reliable signals of body size. However, the central prediction of this hypothesis – that the pitch of vocalisations decreases with size among competing individuals – has limited support in songbirds. One reason could be that only the lowest-frequency components of vocalisations are constrained, and this may go unnoticed when vocal ranges are large. Additionally, the constraint may only be apparent in contexts when individuals are indeed advertising their size. Here we explicitly consider signal diversity and performance limits to demonstrate that body size limits song frequency in an advertising context in a songbird. We show that in purple-crowned fairy-wrens, Malurus coronatus coronatus, larger males sing lower-pitched low-frequency advertising songs. The lower frequency bound of all advertising song types also has a significant negative relationship with body size. However, the average frequency of all their advertising songs is unrelated to body size. This comparison of different approaches to the analysis demonstrates how a negative relationship between body size and song frequency can be obscured by failing to consider signal design and the concept of performance limits. Since these considerations will be important in any complex communication system, our results imply that body size constraints on low-frequency vocalisations could be more widespread than is currently recognised.     

White‐crowned sparrow males show immediate flexibility in song amplitude but not in song minimum frequency in response to changes in noise levels in the field

The soundscape acts as a selective agent on organisms that use acoustic signals to communicate. A number of studies document variation in structure, amplitude, or timing of signal production in correspondence with environmental noise levels thus supporting the hypothesis that organisms are changing their signaling behaviors to avoid masking. The time scale at which organisms respond is of particular interest. Signal structure may evolve across generations through processes such as cultural or genetic transmission. Individuals may also change their behavior during development (ontogenetic change) or in real time (i.e., immediate flexibility). These are not mutually exclusive mechanisms, and all must be investigated to understand how organisms respond to selection pressures from the soundscape. Previous work on white‐crowned sparrows (Zonotrichia leucophrys) found that males holding territories in louder areas tend to sing higher frequency songs and that both noise levels and song frequency have increased over time (30 years) in urban areas. These previous findings suggest that songs are changing across generations; however, it is not known if this species also exhibits immediate flexibility. Here, we conducted an exploratory, observational study to ask whether males change the minimum frequency of their song in response to immediate changes in noise levels. We also ask whether males sing louder, as increased minimum frequency may be physiologically linked to producing sound at higher amplitudes, in response to immediate changes in environmental noise. We found that territorial males adjust song amplitude but not minimum frequency in response to changes in environmental noise levels. Our results suggest that males do not show immediate flexibility in song minimum frequency, although experimental manipulations are needed to test this hypothesis further. Our work highlights the need to investigate multiple mechanisms of adaptive response to soundscapes.     

Information processing in the adaptation of Saccharomyces cerevisiae to osmotic stress: an analysis of the phosphorelay system

Cellular signaling is key for organisms to survive immediate stresses from fluctuating environments as well as relaying important information about external stimuli. Effective mechanisms have evolved to ensure appropriate responses for an optimal adaptation process. For them to be functional despite the noise that occurs in biochemical transmission, the cell needs to be able to infer reliably what was sensed in the first place. For example Saccharomyces cerevisiae are able to adjust their response to osmotic shock depending on the severity of the shock and initiate responses that lead to near perfect adaptation of the cell. We investigate the Sln1–Ypd1–Ssk1-phosphorelay as a module in the high-osmolarity glycerol pathway by incorporating a stochastic model. Within this framework, we can imitate the noisy perception of the cell and interpret the phosphorelay as an information transmitting channel in the sense of C.E. Shannon’s “Information Theory”. We refer to the channel capacity as a measure to quantify and investigate the transmission properties of this system, enabling us to draw conclusions on viable parameter sets for modeling the system.     

The learning advantage: bird species that learn their song show a tighter adjustment of song to noisy environments than those that do not learn

Song learning has evolved within several avian groups. Although its evolutionary advantage is not clear, it has been proposed that song learning may be advantageous in allowing birds to adapt their songs to the local acoustic environment. To test this hypothesis, we analysed patterns of song adjustment to noisy environments and explored their possible link to song learning. Bird vocalizations can be masked by low‐frequency noise, and birds respond to this by singing higher‐pitched songs. Most reports of this strategy involve oscines, a group of birds with learning‐based song variability, and it is doubtful whether species that lack song learning (e.g. suboscines) can adjust their songs to noisy environments. We address this question by comparing the degree of song adjustment to noise in a large sample of oscines (17 populations, 14 species) and suboscines (11 populations, 7 species), recorded in Brazil (Manaus, Brasilia and Curitiba) and Mexico City. We found a significantly stronger association between minimum song frequency and noise levels (effect size) in oscines than in suboscines, suggesting a tighter match in oscines between song transmission capacity and ambient acoustics. Suboscines may be more vulnerable to acoustic pollution than oscines and thus less capable of colonizing cities or acoustically novel habitats. Additionally, we found that species whose song frequency was more divergent between populations showed tighter noise–song frequency associations. Our results suggest that song learning and/or song plasticity allows adaptation to new habitats and that this selective advantage may be linked to the evolution of song learning and plasticity.     

Effects of ambient noise on detectability and localization of avian songs and tones by observers in grasslands

Probability of detection and accuracy of distance estimates in aural avian surveys may be affected by the presence of anthropogenic noise, and this may lead to inaccurate evaluations of the effects of noisy infrastructure on wildlife. We used arrays of speakers broadcasting recordings of grassland bird songs and pure tones to assess the probability of detection, and localization accuracy, by observers at sites with and without noisy oil and gas infrastructure in south‐central Alberta from 2012 to 2014. Probability of detection varied with species and with speaker distance from transect line, but there were few effects of noisy infrastructure. Accuracy of distance estimates for songs and tones decreased as distance to observer increased, and distance estimation error was higher for tones at sites with infrastructure noise. Our results suggest that quiet to moderately loud anthropogenic noise may not mask detection of bird songs; however, errors in distance estimates during aural surveys may lead to inaccurate estimates of avian densities calculated using distance sampling. We recommend caution when applying distance sampling if most birds are unseen, and where ambient noise varies among treatments.