The Energetics of Acoustic Signaling in Anurans and Insects

SYNOPSIS. This review focuses on the energetics of advertisementcalls in frogs and insects (mostly ensiferans). I also reviewa number of methodological questions relating to the most appropriateways to normalize metabolic rates for calling animals and tocalculate efficiency of sound production Although the mechanism of sound production is very differentin these groups (vocalization vs. stridulation), net metaboliccosts normalized to mass are similar among species producingthe most conspicuous calls. Features of the call that interactto determine energetic cost include repetition rate, call duration,and intensity (sound pressure level). Anurans tend to producelouder (more intense) calls while ensiferans tend to producesound during a greater proportion of the calling bout. All evidencesuggests that advertisement calls are produced aerobically andthe aerobic costs are similar or exceed the aerobic (but notnecessarily total costs) of terrestrial locomotion The pattern of radiated sound tends to be constant within aspecies and can be predicted to some degree from characteristicsof an animal's acoustic radiator. Efficiency of sound production(acoustic power/net metabolic power) is low (0.05 to 6%) andvariable when compared to locomotion ({small tilde} 10–20%).From the present sample it appears that frogs are more efficientthan ensiferans, but as more katydids are studied this trendmay not hold. Of the factors that have been identified as determiningefficiency the most important are the match between the sizeof the radiator and wavelength radiated, the absorption propertiesof the environment immediately around the animal, and the presenceor absence of structures such as baffles and acoustic burrows     

Energetic cost of calling: general constraints and species‐specific differences

The energetic cost of acoustic signalling varies tremendously among species. Understanding factors responsible for this heterogeneity is important for understanding the costs and benefits of signalling. Here, we present a general model, based on well‐established principles of bioenergetics, which predicts the energetic cost of call production across species. We test model predictions using an extensive database of resting and calling metabolic rates of insects, amphibians and birds. Results are largely supportive of model predictions. Calling metabolic rates scale predictably with body mass and temperature such that calling and resting metabolic rates are directly proportional to each other. The cost of acoustic signalling is ～8 times higher than resting metabolic rate in ectotherms, and ～2 times higher in birds. Differences in the increase in metabolic rate during calling are explained by the relative size of species’ sound‐producing muscles. Combined with published work, we quantify call efficiency and discuss model implications.     

The relationship between the acoustic behaviour and surface activity of killer whales (<Emphasis Type="Italic">Orcinus orca</Emphasis>) that feed on herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>)

We describe the acoustic behaviour of piscivorous killer whales in Norwegian and Icelandic waters. Whales were assigned to one of three activities (feeding, travelling or other), and sound recordings were made in their proximity with a single hydrophone and a digital audiotape (DAT) recorder. A quantitative analysis of the production of pulsed calls, whistles and echolocation clicks in the three activities revealed that there was a significant effect of activity on the production of these sound types. Both killer whales in Icelandic and Norwegian waters produced high rates of clicks and calls during feeding and low rates of click, calls and whistles during travelling. The differences can be used as acoustical markers and provides new possibilities for acoustic monitoring of killer whales in these areas. Based on the similarity between their prey choice, hunting strategies, phenotype and acoustic behaviour, we suggest that the killer whales in Icelandic and Norwegian waters belong to the same ecotype: Scandinavian herring-eating killer whales.     

三种免疫学方法在矽肺合并肺结核诊断中的价值研究

目的:探讨T淋巴细胞酶联斑点实验(T-SPOT)、结核菌素皮肤试验(tuberculin test,TST)以及结核抗体(tuberculosis antibody)在矽肺合并肺结核(pulmonary silicosis complicated with tuberculosis)诊断中的应用价值。方法:收集2015年8月7日至2016年5月20日确诊为矽肺结核感染患者41例、非矽肺结核患者90例、健康体检学生对照组40例,对三组人群经以上免疫学方法检测的结果进行统计分析。结果:1)矽肺结核、非矽肺结核及健康人群中T-SPOT阳性率分别为73.17%、93.33%、15%,三者两两比较差异有统计学意义;矽肺结核、非矽肺结核及健康人群中TST阳性率分别为58.54%、88.89%、32.5%,非矽肺结核患者与健康人群及矽肺结核患者之间,差异均有统计学意义,但矽肺结核患者与健康人群之间,差异无统计学意义;矽肺结核、非矽肺结核及健康人群中结核抗体阳性率分别为36.58%、42.22%、52.5%,三者两两比较差异均无统计学意义。2)三种检测方法在矽肺结核患者中的敏感性分别为73.17%、58.54%、36.58%,差异有统计学意义,其中T-SPOT及TST的敏感性均高于结核抗体方法(P0.05);特异性分别为85%、67.5%、47.5%,其中T-SPOT的特异性高于结核抗体方法(P0.05);阳性预测值分别为83.33%、64.86%、41.67%,阴性预测值分别为75.56%、61.36%、42.22%,三种方法的阳性及阴性预测值存在差异,均有统计学意义。3)T-SPOT在I、II、III期矽肺结核检测的阳性率分别为52.38%、94.12%、100%,I期与II期或III期的阳性率之间差异有统计学意义,II期与III期之间阳性率差异无统计学意义。结论:T-SPOT方法具有较高的敏感性及特异性,对矽肺结核辅助诊断及临床分期具有较好的应用价值。     

Correlation of acoustic and visual signals in the cichlid fish, <Emphasis Type="Italic">Tramitichromis intermedius</Emphasis>

We investigated acoustic and visual communication concurrently in wild caught adult and captive-born, first generation offspring of the East African Rift Lake cichlid fish Tramitichromis intermedius. Only males emit sound during courtship. Sound production is always accompanied by quivering, but quiver behavior is not always accompanied by sound. This separation of quivering and sound supports the hypothesis that sound production is intentional serving a communicative role. As spawning nears, both sound production and quiver behavior increase. In terms of the ontogeny of sound production, the first observation of courtship occurs just days before the first spawning event and the first sound emission accompanies the first courtship activity. The accompaniment of quivering with sound as well as the escalation of the two behaviors with the approach of spawning follows similar patterns in wild caught and captive-born males. The tight correlation between behavior and sound production in both groups indicates their simultaneous performance plays an important role in reproduction. It is probable that the ability to produce sound and perform quiver behavior at the same time may be a measure of mate quality.     

基于比速率及代谢流的黑曲霉突变株和野生株分析

黑曲霉具备优异的外源蛋白表达和分泌能力,从而被广泛应用于工业酶制剂的生产。通过研究黑曲霉突变株和野生株在相同培养条件下生理参数和代谢流的差异,确定了黑曲霉合成糖化酶过程中的限制性因素。宏观动力学分析发现,较之野生株,突变株具有较高的最大比生长速率,并且副产物得率降低了90%,底物利用率提高了近30%,表明突变株与野生株在碳源分配和产物转化率上具有明显的差异。利用流平衡分析（FBA）计算胞内代谢通量分布,发现还原力和核糖的供应水平是限制菌体合成的主要因素,而前体氨基酸是合成糖化酶最主要的限制性因素。这些研究结果为后续发酵工艺优化和菌株基因改造提供了有益的思路。     

Foraging and mate-finding in the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae) under the risk of predation

Animal foraging and reproductive behaviour is influenced by other simultaneous demands such as predator avoidance. The trade-offs between these demands may depend on sex or mating experience. This study demonstrates that the olfactory-mediated foraging and mate-seeking behaviours in the silver Y moths, Autographa gamma , are affected by auditory cues mimicking their bat predators. Both males and females changed their foraging behaviour under simulated predation risk. Fewer moths reached the odour source following sound stimulation and the time to find the odour source increased by up to 250%. However, there were no significant differences between male and female ability to reach the plant odour source or the duration of the flight towards the source when stimulated with ultrasound. Hence females are not more cautious than males when observed in the same behavioural context. Risk-taking in males was independent of whether they were flying toward a flower odour or sex pheromones having equal attractive value. This indicates that the trade-off between olfactory and acoustic cues is independent the type of odour. Mated females were not as strongly affected by sound as non-mated, indicating that flower odours have a higher adaptive value for mated females, suggesting that some processes following mating experience influence the trade-off between flower odours and simulated bat sounds.     

Geographical variation in sound production in the anemonefish Amphiprion akallopisos

Because of pelagic-larval dispersal, coral-reef fishes are distributed widely with minimal genetic differentiation between populations. Amphiprion akallopisos, a clownfish that uses sound production to defend its anemone territory, has a wide but disjunct distribution in the Indian Ocean. We compared sounds produced by these fishes from populations in Madagascar and Indonesia, a distance of 6500 km. Differentiation of agonistic calls into distinct types indicates a complexity not previously recorded in fishes' acoustic communication. Moreover, various acoustic parameters, including peak frequency, pulse duration, number of peaks per pulse, differed between the two populations. The geographic comparison is the first to demonstrate 'dialects' in a marine fish species, and these differences in sound parameters suggest genetic divergence between these two populations. These results highlight the possible approach for investigating the role of sounds in fish behaviour in reproductive divergence and speciation.     

Fish Sound Production in the Presence of Harmful Algal Blooms in the Eastern Gulf of Mexico

This paper presents the first known research to examine sound production by fishes during harmful algal blooms (HABs). Most fish sound production is species-specific and repetitive, enabling passive acoustic monitoring to identify the distribution and behavior of soniferous species. Autonomous gliders that collect passive acoustic data and environmental data concurrently can be used to establish the oceanographic conditions surrounding sound-producing organisms. Three passive acoustic glider missions were conducted off west-central Florida in October 2011, and September and October 2012. The deployment period for two missions was dictated by the presence of red tide events with the glider path specifically set to encounter toxic Karenia brevis blooms (a.k.a red tides). Oceanographic conditions measured by the glider were significantly correlated to the variation in sounds from six known or suspected species of fish across the three missions with depth consistently being the most significant factor. At the time and space scales of this study, there was no detectable effect of red tide on sound production. Sounds were still recorded within red tide-affected waters from species with overlapping depth ranges. These results suggest that the fishes studied here did not alter their sound production nor migrate out of red tide-affected areas. Although these results are preliminary because of the limited measurements, the data and methods presented here provide a proof of principle and could serve as protocol for future studies on the effects of algal blooms on the behavior of soniferous fishes. To fully capture the effects of episodic events, we suggest that stationary or vertically profiling acoustic recorders and environmental sampling be used as a complement to glider measurements.     

基于Mel倒谱系数和矢量量化的昆虫声音自动鉴别

为了给生产单位害虫管理的普通技术人员提供简便易操作的昆虫种类鉴别方法, 本研究把人类语音识别领域的先进技术应用于昆虫识别, 提出了一种新颖的昆虫声音自动鉴别方法, 用声音参数化技术为昆虫声纹识别设计了一种简单易行的方案。声音信号经过预处理、分段得到一系列的声音样本, 从声音样本提取Mel倒谱系数（MFCC）, 并用Linde-Buzo-Gray（LBG）算法对提取的MFCC进行矢量量化(VQ), 所得码字作为声音样本的特征模型。特征参数之间的匹配用搜索最近邻的方法实现。本文方法在包含70种昆虫声音的库中进行了试验, 取得了超过96%的识别率和理想的时间性能。试验结果证明了该方法的有效性。     

Acoustic characteristics and variations in grunt vocalizations in the oyster toadfish <Emphasis Type="Italic">Opsanus tau</Emphasis>

Acoustic communication is critical for reproductive success in the oyster toadfish Opsanus tau. While previous studies have examined the acoustic characteristics, behavioral context, geographical variation, and seasonality of advertisement boatwhistle sound production, there is limited information on the grunt or other non-advertisement vocalizations in this species. This study continuously monitored sound production in toadfish maintained in an outdoor habitat for four months to identify and characterize grunt vocalizations, compare them with boatwhistles, and test for relationships between the incidence of grunt vocalizations, sound characteristics and environmental parameters. Oyster toadfish produced grunts in response to handling, and spontaneous single (70% of all grunts), doublet (10%), and trains of grunts (20%) throughout the May to September study period. Grunt types varied in pulse structure, duration, and frequency components, and were shorter and of lower fundamental frequency than the pulse repetition rate of boatwhistles. Higher water temperatures were correlated with a greater number of grunt emissions, higher fundamental frequencies, and shorter sound durations. The number of grunts per day was also positively correlated with daylength and maximum tidal amplitude differences (previously entrained) associated with full and new moons, thus providing the first demonstration of semilunar vocalization rhythms in the oyster toadfish. These data provide new information on the acoustic repertoire and the environmental factors correlated with sound production in the toadfish, and have important implications for seasonal acoustic communication in this model vocal fish.     

Spatial orientation in the bushcricket <Emphasis Type="Italic">Leptophyes punctatissima</Emphasis> (Phaneropterinae; Orthoptera): III. Peripheral directionality and central nervous processing of spatial cues

We examined peripheral and central nervous cues underlying the ability of the bushcricket Leptophyes punctatissima to orient to elevated and depressed sound sources broadcasting the female acoustic reply. The peripheral spatial directionality of the ear was measured physiologically using monaural preparations of an auditory interneuron (T-fibre). In the azimuth, maximal interaural intensity differences of 18 dB occur between ipsi- and contralateral stimulation. With increasing elevation or depression of the sound sources, IIDs decrease systematically and reach zero with the source exactly above or below the preparation. Bilateral, simultaneous recordings of the activity of the pair of interneurons allowed determining the binaural discharge differences which occur in response to the extremely short (1 ms) female reply. These discharge differences are large (four action potentials/stimulus) and reliable in the azimuth with lateral stimulation, and decrease gradually with more frontal stimulation. With elevation and depression of sound sources these differences again decrease to one action potential/stimulus at 60° or 75° elevation, and lateral stimulus angles of about 60°. We also calculated the reliability with which a receiver could correctly determine the location of the sound source. We discuss these quantitative measures in relation to the spatial phonotactic behaviour of male L. punctatissima.     

饲养状态真鲂(鱼弗)发声的昼夜与季节变化

本文研究了受水温和光周期等自然变化影响的饲养状态真鲂鮄发声的昼夜与季节变化,并研究了实验鱼活动(鱼类游泳次数)的日变化。声音信号的昼夜节律记录发声活动的日常水平(摄食之外的时期),但是每月变化的记录(季节性式型)则在摄食期间进行,因为摄食时声音信号增加,而日常发声活动较不频繁。实验鱼包括雌雄两性,且未达性成熟。真鲂鮄在白天发声多一些,也更活跃。声音为阵发式的,较不频繁(平均值=0.04发声/min每鱼每天)。最少的发声活动出现在晚上,凌晨和黄昏居中(声音的阵发更频繁,但是声音更少),最多的发声活动出现在白天(声音的阵发更频繁,并且含更多数目的声音)。竞争摄食时声音信号的比率不呈现季节性变化(平均值=3.98发声/min每鱼),与温度也不相关,显示出竞争摄食时声音的发出以最大比率进行。敲击声和呼噜声的某些声学特征与温度相关,特别是在较高的温度下呼噜声的节拍间隔急剧下降。敲击声和呼噜声的声音参数中的季节性变化,多数可以解释为发声肌肉和中央声音控制回路的温度效应。     

Conceptual and statistical problems with the use of the Shannon-Weiner entropy index in bioacoustic analyses

Information theory and its indices were developed for human communication to predict the amount of information transferred in a message. One such index, the Shannon-Weiner index (SWI), has often been used to analyse information from other fields in which its application may not be appropriate. In ecoacoustics, SWI is used to compare acoustic diversity (i.e. a measure derived by integrating the richness and abundance of animal sounds) between locations. In animal communication, SWI is used to quantify repertoire complexity (i.e. a measure derived by integrating the number and abundance of sound types produced by individuals or species) as an approach to understanding signal evolution. We discuss problems associated with using the SWI in ecoacoustics and animal communication. Specifically, we discuss conceptual and statistical problems associated with the SWI and then illustrate these problems using hypothetical data. In ecoacoustics, the SWI’s assumptions of random variables and independent samples are often violated. In animal communication, the SWI fails to distinguish among repertoires in which the number of sound types and the abundance of each sound type differ. We also show that other methods do capture these differences. We conclude that the SWI does not adequately represent acoustic diversity or repertoire complexity due to the multiple conceptual and statistical issues associated with its use. We recommend other analytical methods to more fully describe these biological systems, including goodness of fit, Morisita similarity index and Markov chain analysis. These methods provide more information for future comparisons and permit researchers to test hypotheses more directly.     

Tree cricket baffles are manufactured tools

Tree crickets use sound to attract mates and make acoustic baffles to increase their sound production efficiency. It has been recently discovered that these insects use a flexible yet inherited behavioural programme to make acoustically optimal baffles. Whether these baffles qualify as tools, however, remains controversial. Here, baffle‐using and baffle‐making behaviours are analysed using the most current and authoritative definition of tool use. The current definition of tool use does not require the tool to be detached from the substrate and includes attached but manipulable external objects. Given this schema, tree cricket baffles, which are attached but manipulated prior to use, must be considered tools. The mode of manufacture for a baffle is “Subtract,” and the mode of use is “Drape”.     

Harmonic Hopping,and Both Punctuated and Gradual Evolution of Acoustic Characters in Selasphorus Hummingbird Tail-Feathers

Models of character evolution often assume a single mode of evolutionary change, such as continuous, or discrete. Here I provide an example in which a character exhibits both types of change. Hummingbirds in the genus Selasphorus produce sound with fluttering tail-feathers during courtship. The ancestral character state within Selasphorus is production of sound with an inner tail-feather, R2, in which the sound usually evolves gradually. Calliope and Allen''s Hummingbirds have evolved autapomorphic acoustic mechanisms that involve feather-feather interactions. I develop a source-filter model of these interactions. The ‘source’ comprises feather(s) that are both necessary and sufficient for sound production, and are aerodynamically coupled to neighboring feathers, which act as filters. Filters are unnecessary or insufficient for sound production, but may evolve to become sources. Allen''s Hummingbird has evolved to produce sound with two sources, one with feather R3, another frequency-modulated sound with R4, and their interaction frequencies. Allen''s R2 retains the ancestral character state, a ∼1 kHz “ghost” fundamental frequency masked by R3, which is revealed when R3 is experimentally removed. In the ancestor to Allen''s Hummingbird, the dominant frequency has ‘hopped’ to the second harmonic without passing through intermediate frequencies. This demonstrates that although the fundamental frequency of a communication sound may usually evolve gradually, occasional jumps from one character state to another can occur in a discrete fashion. Accordingly, mapping acoustic characters on a phylogeny may produce misleading results if the physical mechanism of production is not known.     

Curtailment and acoustic deterrents reduce bat mortality at wind farms

The impacts of wind energy on bat populations is a growing concern because wind turbine blades can strike and kill bats, and wind turbine development is increasing. We tested the effectiveness of 2 management actions at 2 wind-energy facilities for reducing bat fatalities: curtailing turbine operation when wind speeds were <5.0 m/second and combining curtailment with an acoustic bat deterrent developed by NRG Systems. We measured the effectiveness of the management actions using differences in counts of bat carcasses quantified by daily and twice-per-week standardized carcass searches of cleared plots below turbines, and field trials that estimated searcher efficiency and carcass persistence. We studied turbines located at 2 adjacent wind-energy facilities in northeast Illinois, USA, during fall migration (1 Aug–15 Oct) in 2018. We estimated the effectiveness of each management action using a generalized linear mixed-effects model with several covariates. Curtailment alone reduced overall bat mortality by 42.5% but did not reduce silver-haired bat (Lasionycteris noctivagans) mortality. Overall bat fatality rates were 66.9% lower at curtailed turbines with acoustic deterrents compared to turbines that operated at manufacturer cut-in speed. Curtailment and the deterrent reduced bat mortality to varying degrees between species, ranging from 58.1% for eastern red bats (Lasiurus borealis) to 94.4 for big brown bats (Eptesicus fuscus). Hoary (Lasiurus cinereus) and silver-haired bat mortality was reduced by 71.4% and 71.6%, respectively. Our study lacked a deterrent-only treatment group because of the expense of acoustic deterrents. We estimated the additional reduction in mortality with concurrent deployment of the acoustic deterrent and curtailment under the assumption that curtailment and the acoustic deterrent would have reduced mortality by the same percentage at adjacent wind-energy facilities. Acoustic deterrents resulted in 31.6%, 17.4%, and 66.7% additional reductions of bat mortality compared to curtailment alone for eastern red bat, hoary bat, and silver-haired bat, respectively. The effectiveness of acoustic deterrents for reducing bat mortality at turbines with rotor-swept area diameters >110 m is unknown because high frequency sound attenuates quickly, which reduces coverage of rotor-swept areas. Management actions should consider species differences in the ability of curtailment and deterrents to reduce bat mortality and increase energy production.     

Quantifying Oldowan Stone Tool Production at Olduvai Gorge,Tanzania

Recent research suggests that variation exists among and between Oldowan stone tool assemblages. Oldowan variation might represent differential constraints on raw materials used to produce these stone implements. Alternatively, variation among Oldowan assemblages could represent different methods that Oldowan producing hominins utilized to produce these lithic implements. Identifying differential patterns of stone tool production within the Oldowan has implications for assessing how stone tool technology evolved, how traditions of lithic production might have been culturally transmitted, and for defining the timing and scope of these evolutionary events. At present there is no null model to predict what morphological variation in the Oldowan should look like. Without such a model, quantifying whether Oldowan assemblages vary due to raw material constraints or whether they vary due to differences in production technique is not possible. This research establishes a null model for Oldowan lithic artifact morphological variation. To establish these expectations this research 1) models the expected range of variation through large scale reduction experiments, 2) develops an algorithm to categorize archaeological flakes based on how they are produced, and 3) statistically assesses the methods of production behavior used by Oldowan producing hominins at the site of DK from Olduvai Gorge, Tanzania via the experimental model. Results indicate that a subset of quartzite flakes deviate from the null expectations in a manner that demonstrates efficiency in flake manufacture, while some basalt flakes deviate from null expectations in a manner that demonstrates inefficiency in flake manufacture. The simultaneous presence of efficiency in stone tool production for one raw material (quartzite) and inefficiency in stone tool production for another raw material (basalt) suggests that Oldowan producing hominins at DK were able to mediate the economic costs associated with stone tool procurement by utilizing high-cost materials more efficiently than is expected and low-cost materials in an inefficient manner.     

Acoustic defence in an insect: characteristics of defensive stridulation and differences between the sexes in the tettigoniid Poecilimon ornatus (Schmidt 1850)

Many insects exhibit secondary defence mechanisms upon contact with a predator, such as defensive sound production or regurgitation of gut contents. In the tettigoniid Poecilimon ornatus, both males and females are capable of sound production and of regurgitation. However, wing stridulatory structures for intraspecific acoustic communication evolved independently in males and females, and may result in different defence sounds. Here we investigate in P. ornatus whether secondary defence behaviours, in particular defence sounds, show sex-specific differences. The male defence sound differs significantly from the male calling song in that it has a longer syllable duration and a higher number of impulses per syllable. In females, the defence sound syllables are also significantly longer than the syllables of their response song to the male calling song. In addition, the acoustic disturbance stridulation differs notably between females and males as both sexes exhibit different temporal patterns of the defence sound. Furthermore, males use defence sounds more often than females. The higher proportion of male disturbance stridulation is consistent with a male-biased predation risk during calling and phonotactic behaviour. The temporal structures of the female and male defence sounds support a deimatic function of the startling sound in both females and males, rather than an adaptation for a particular temporal pattern. Independently of the clear differences in sound defence, no difference in regurgitation of gut content occurs between the sexes.     

Commensal auditory communication in two species of Neoconocephalus (Orthoptera)

Auditory communication in Neoconcephalus ensiger and Neoconocephalus robustus, two stridulating tettigoniids that inhabit the same geographical area, was examined to determine how these two species interact. Computer and electrophysiological techniques were used to analyze sound production and reception. Although similar in appearance, the males of these species produce easily distinguishable acoustic communication signals (call songs) that serve to attract conspecific females. Both wave form and spectral analyses were done on selected segments of the species-specific call songs by using discrete Fourier transform techniques. Sound production and reception capacities were measured by octave-band analyzers and extracellular electrode techniques. The results show that each of these species has a distinct, primary carrier frequency band. Secondly, the log magnitude spectra revealed a significant secondary component in the call song of N. robustus which corresponds to the primary energy band in the call song of N. ensiger. This overlap in acoustic signals and other findings suggest that males of N. ensiger might use the louder stridulation of the more metabolically active N. robustus to attract conspecific females over great distances. N. ensiger males have a sound production capacity like N. robustus, but conserve energy for soft, near-field signaling. The audiogram representing sensitivity of the tympano-receptor in the foreleg showed sound reception to be matched precisely with sound production curves.