Otoacoustic emissions from insect ears: evidence of active hearing?

Sensitive hearing organs often employ nonlinear mechanical sound processing which generates distortion-product otoacoustic emissions (DPOAE). Such emissions are also recordable from tympanal organs of insects. In vertebrates (including humans), otoacoustic emissions are considered by-products of active sound amplification through specialized sensory receptor cells in the inner ear. Force generated by these cells primarily augments the displacement amplitude of the basilar membrane and thus increases auditory sensitivity. As in vertebrates, the emissions from insect ears are based on nonlinear mechanical properties of the sense organ. Apparently, to achieve maximum sensitivity, convergent evolutionary principles have been realized in the micromechanics of these hearing organs-although vertebrates and insects possess quite different types of receptor cells in their ears. Just as in vertebrates, otoacoustic emissions from insects ears are vulnerable and depend on an intact metabolism, but so far in tympanal organs, it is not clear if auditory nonlinearity is achieved by active motility of the sensory neurons or if passive cellular characteristics cause the nonlinear behavior. In the antennal ears of flies and mosquitoes, however, active vibrations of the flagellum have been demonstrated. Our review concentrates on experiments studying the tympanal organs of grasshoppers and moths; we show that their otoacoustic emissions are produced in a frequency-specific way and can be modified by electrical stimulation of the sensory cells. Even the simple ears of notodontid moths produce distinct emissions, although they have just one auditory neuron. At present it is still uncertain, both in vertebrates and in insects, if the nonlinear amplification so essential for sensitive sound processing is primarily due to motility of the somata of specialized sensory cells or to active movement of their (stereo-)cilia. We anticipate that further experiments with the relatively simple ears of insects will help answer these questions.     

SOUND PRODUCTION BY AQUATIC INSECTS

1. Sound production by aquatic insects is found in four orders — Trichoptera, Odonata, Heteroptera and Coleoptera. 2. Immature aquatic insects that produce sound are rare, stridulation being present in one family of Trichoptera (Hydropsychidae) and one genus and species in a relic suborder of Odonata (Anisozygoptera) - Epiophlebia superstes. Hydropsychid larvae produce sound with a head/fore femur mechanism and use sound as part of aggressive behaviour for defence of feeding nets. Larval E. superstes use a hind femur/abdominal mechanism to dissuade predators. 3. Sound production has been documented in adults of all families of aquatic Heteroptera except Helotrephidae. In corixids and notonectids, acoustic signals play a role in mating. Members of the genus Buenoa (Notonectidae) are unique in having two stridulatory mechanisms in the same individual. Sound production has been most intensively studied in the Corixidae. Although sounds are used in mating by all singing corixids, their use seems to be facultative in some species and obligatory in others. Recent experiments by Theiss (1982) have shown that the air stores carried by corixids are used for both sound radiation and reception. 4. The adephagan beetle families Hygrobiidae, Dytiscidae and Haliplidae have all been shown to produce sound. Mechanisms of sound production have been established for haliplids and hygrobiids but have yet to be for most dytiscids. Sound production is used by beetles as part of sequences of aggressive/defensive and reproductive behaviour. 5. Sound production is especially well documented in the Hydrophilidae (Polyphaga). Hydrophilids use an abdominal/elytral mechanism and sound appears to be used in the same contexts as in adephagans. 6. Insects that produce sound under water must contend with the physical problems of sound transmission in a relatively dense, viscous medium with sharp boundaries. Because of potential distortion of the frequency components in a signal by reflection from the air/water interface in very shallow water, frequency is unreliable for encoding information. Aquatic insects use instead amplitude modulation and temporal patterning of signals. 7. For aquatic invertebrates, sound fields are different than those in air because the extent of the near field is approximately four times greater in water. This near field, a region in which displacement waves are predominant over pressure waves, extends to a greater distance than most aquatic insects communicate over. Such displacement waves could have important but as yet unconsidered effects. 8. The mass and viscosity of the water dictates that sound producing structures of aquatic insects should be heavier and more massive than those of terrestrial insects. A survey of stridulatory organs of aquatic insects reveals this to be true and reveals that the relatively fragile, membranous stridulatory organs of some terrestrial insects (especially Orthoptera) are absent. 9. The elaboration of sound producing structures in aquatic insects probably occurred at the family or subfamily level and for Heteroptera, Trichoptera and Odonata evolved after the invasion of the water. Acoustic signals used reproductively would probably be more closely associated with the emergence of new taxa. 10. Stridulatory structures have been derived from either structures devoted to some other function or from structures involved in the behaviour currently enhanced by sound production.     

Multi-trophic effects of ungulate intraguild predation on acorn weevils

Predators and parasitoids may contribute to controlling the population sizes of phytophagous insects, and this has been shown to benefit plants. Phytophagous insects may also be killed by other herbivores (intraguild predation), usually larger-sized vertebrates that ingest insects accidentally while feeding on common food sources. We studied the intraguild predation on acorn weevils by ungulates and assessed the consequences for weevil populations. Infested acorns are prematurely abscised and the weevil larvae finish their development inside the acorns after being dropped. Our results show that weevil larvae were killed by ungulates eating the infested acorns on the ground. Ungulates did not discriminate between infested and sound acorns, and the probability of a larva being incidentally eaten was inversely related to acorn availability. Thus, predation risk was enhanced by the premature drop of infested acorns when acorn availability on the ground was low. Predation rates on infested acorns were much higher where ungulates were present, and acorn infestation rates were significantly lower. However, ungulates did not provide the oaks any net benefit, since the reduction of infestation rates was not enough to compensate for the large amounts of sound acorns eaten by ungulates. Seed predation is usually studied as a progressive loss of seeds by pre- and post-dispersal predators, but the interactions between them are usually not considered. We show that intraguild predation on insects by large ungulates had an effect on the structure of the foraging guild, as the proportion of acorns predated by insects decreased; however, replicating the same experimental design in different ecological scenarios would increase the strength of these results. In conclusion, the present study shows the importance of considering the multi-trophic interactions between seed predators in order to have a complete picture of granivory.     

基于MFCC和GMM的昆虫声音自动识别

昆虫的运动、 取食、 鸣叫都会发出声音, 这些声音存在种内相似性和种间差异性, 因此可用来识别昆虫的种类。基于昆虫声音的昆虫种类自动检测技术对协助农业和林业从业人员方便地识别昆虫种类非常有意义。本研究采用了语音识别领域里的声音参数化技术来实现昆虫的声音自动鉴别。声音样本经预处理后, 提取梅尔倒谱系数（Mel frequency cepstrum coefficient, MFCC）作为特征, 并用这些样本提取的MFCC特征集训练混合高斯模型（Gaussian mixture model, GMM）。最后用训练所得到的GMM对未知类别的昆虫声音样本进行分类。该方法在包含58种昆虫声音的样本库中进行了评估, 取得了较高的识别正确率（平均精度为98.95%）和较理想的时间性能。该测试结果证明了基于MFCC和GMM的语音参数化技术可以用来有效地识别昆虫种类。     

Rhythm of sounds produced by larvae of the oriental hornet Vespa orientalis: spectral analysis

In the hornet nest of the species Vespa orientalis, there is transmission of information by acoustic means between the larvae and the adults. The rhythmic pattern of the sounds produced by the larvae was recorded and spectrally analyzed for rhythm frequencies by use of the Fast Fourier Transform. The frequency of the "larval activity duration till cessation" was 0.018 Hz whereas the interval between two successive sound productions ranged from 0 to 1.0 Hz. The possible significance of precise signaling by the larvae towards efficient communication in colonies of social insects is discussed.     

昆虫弦音器及其声音感受分子机制

弦音器是昆虫类特有的一种机械感受器,亦称弦音感受器或剑梢感受器。它主要具有感知外界声压和体内肌肉运动的听觉功能,研究弦音器的机能结构对揭秘昆虫听觉的神经机制有重要的科学意义。本文从弦音器多样性和进化入手,重点综述了弦音器的微细结构、基因功能定位、声音感受分子机制及其声压增幅分子生物物理学原理,为昆虫听觉仿生学的研究提供了理论依据。     

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”.     

If a bird flies in the forest,does an insect hear it?

Birds are major predators of many eared insects including moths, butterflies, crickets and cicadas. We provide evidence supporting the hypothesis that insect ears can function as ‘bird detectors’. First, we show that birds produce flight sounds while foraging. Eastern phoebes (Sayornis phoebe) and chickadees (Poecile atricapillus) generate broadband sounds composed of distinct repetitive elements (approx. 18 and 20 Hz, respectively) that correspond to cyclic wing beating. We estimate that insects can detect an approaching bird from distances of at least 2.5 m, based on insect hearing thresholds and sound level measurements of bird flight. Second, we show that insects with both high and low frequency hearing can hear bird flight sounds. Auditory nerve cells of noctuid moths (Trichoplusia ni) and nymphalid butterflies (Morpho peleides) responded in a bursting pattern to playbacks of an attacking bird. This is the first study to demonstrate that foraging birds generate flight sound cues that are detectable by eared insects. Whether insects exploit these sound cues, and alternatively, if birds have evolved sound-reducing foraging tactics to render them acoustically ‘cryptic’ to their prey, are tantalizing questions worthy of further investigation.     

天敌昆虫耐冷藏力研究进展

低温冷藏是延长天敌昆虫货架期的有效方法之一.昆虫的存活率、生殖力、田间寄生(捕食)力等是评估天敌昆虫耐冷藏力的重要指标.昆虫耐冷藏力受冷藏前、冷藏中和冷藏后的各种外源(非生物)因子和内源(生物)因子的影响.这些因子主要包括温度、湿度、光周期、氧浓度、冷藏持续时间、冷藏前的预处理、冷藏过程中的温度处理(混合低温)等外源因子和虫体能量物质的储备、繁殖方式、年龄/龄期、滞育(休眠)状态、营养等内源因子.本文对影响天敌昆虫耐冷藏性的内源因子和外源因子的种类以及产生这些影响的生理学机制等进行了综述,以期为天敌昆虫低温储藏技术研究和生产方案的制定提供依据.     

Selection of floral resources to optimise conservation of agriculturally-functional insect groups

Non-crop vegetation in agricultural landscapes can provide a means of conserving insects in farmed landscapes and optimising on-farm ecosystem services as a result. Inclusion of floral resources may be particularly useful in conserving many beneficial insects, where groups including pollinators and pest natural enemies often rely on nectar and pollen during (at least part of) their life-cycle. As not all flowers are equally suited to all beneficial insects, selection of appropriate flowering plants is key to ensuring that conservation targets are met and benefits to ecosystems services realised as a result. This short paper describes an experiment conducted to assess the ‘total temporal attractiveness’ of a range of British wildflowers to selected functional insect groups. The results obtained demonstrate that flowering period alone is a poor indicator of plant suitability to insects, where no relationship existed between this and attraction to insects overall. Data also suggest that, based on attraction over a season, certain flowering plants are more likely to be of general insect conservation value and/or benefit to functional insect groups than others. Attraction to pest insects was also considered, with relatively high catches of thrips and pollen beetles observed in flowering stands of some plants.     

Influence of vegetation on acoustic properties of soils

Possible influences of vegetation on acoustically relevant soil parameters, such as porosity and soil structure, were considered. In situ measurements of sound interference patterns were performed in seven plant communities by means of an inclined track method. Normal acoustical specific impedances were calculated with a plane wave outdoor sound propagation model. The impedances found generally showed a real part constant with frequency, and an imaginary part decreasing with frequency. It is concluded that forest floors have an acoustically detectable layer structure and that, for purposes of modelling of outdoor sound fields, much lower impedances have to be used than for institutional grass.     

昆虫弦音器及其声音感受分子机制

弦音器是昆虫类特有的一种机械感受器,亦称弦音感受器或剑梢感受器。它主要具有感知外界声压和体内肌肉运动的听觉功能,研究弦音器的机能结构对揭秘昆虫听觉的神经机制有重要的科学意义。本文从弦音器多样性和进化入手,重点综述了弦音器的微细结构、基因功能定位、声音感受分子机制及其声压增幅分子生物物理学原理,为昆虫听觉仿生学的研究提供了理论依据。     

Orientation to distant sounds by foraging big brown bats (Eptesicus fuscus)

Big brown bats (Eptesicus fuscus) detect and orient toward relatively low-frequency sounds produced by chorusing frogs or groups of stridulating insects. This response occurs at distances of at least 600 m. The bats are also attracted to a broadcasted recording of the sounds. If recently fed, they do not orient to these sounds. Bats that fly toward the most intense sound field locate insects at a significantly greater rate than those that choose another direction. This suggests that the use of these long-distance acoustic cues may be important for locating concentrations of flying insects.     

Hearing in tsetse flies? Morphology and mechanics of a putative auditory organ

Tympanal hearing organs are widely used by insects to detect sound pressure. Such ears are relatively uncommon in the order Diptera, having only been reported in two families thus far. This study describes the general anatomical organization and experimentally examines the mechanical resonant properties of an unusual membranous structure situated on the ventral prothorax of the tsetse fly, Glossina morsitans (Diptera: Glossinidae). Anatomically, the prosternal membrane is backed by an air filled chamber and attaches to a pair of sensory chordotonal organs. Mechanically, the membrane shows a broad resonance around 5.3-7.2 kHz. Unlike previously reported dipteran tympana, a directional response to sound was not found in G. morsitans. Collectively, the morphology, the resonant properties and acoustic sensitivity of the tsetse prothorax are consistent with those of the tympanal hearing organs in Ormia sp. and Emblemasoma sp. (Tachinidae and Sarcophagidae). The production of sound by several species of tsetse flies has been repeatedly documented. Yet, clear behavioural evidence for acoustic behaviour is sparse and inconclusive. Together with sound production, the presence of an ear-like structure raises the enticing possibility of auditory communication in tsetse flies and renews interest in the sensory biology of these medically important insects.     

Vision should not be overlooked as an important sensory modality for finding host plants

In the last 50 yr, the role of vision in insect interactions with host plants has received relatively little attention. This lack of research is associated with a number of assumptions about chemical cues being the ultimate sensory determinants of host finding. This article presents arguments and detailed evidence to refute these assumptions. Insects from essentially all phytophagous orders use vision for locating host plants, and some recent examples have shown that vision can be even more important than olfaction. Moreover, a number of insects have the ability to visually differentiate host species. This ability means that the visual capabilities of phytophagous insects should not be underestimated. Visual cues always should be considered and integrated into studies of host finding.     

Size and scale effects as constraints in insect sound communication

For optimal transfer of power to the surrounding medium, a sound source should have a radius of 1/6 to 1/4 of the sound wavelength. Sound-waves propagate from the source as compressions and rarefactions of the fluid medium, which decay by spreading and viscous losses. Higher frequencies are more easily refracted and reflected by objects in the environment, causing degradation of signal structure. In open air or water, the sound spreads spherically and decays by the inverse square law. If the sound is restricted to two dimensions rather than three, it decays as the inverse of range, whereas waves within a rod decay largely due to viscous losses; such calls are usually rather simple pulses and rely on the initial time of arrival because of multiple pathlengths or different propagation velocities in the environment. Because of the relationship between calling success and reproductive success, singing insects are under selective pressure to optimize the range, and to maintain the specificity, of their calls. Smaller insects have less muscle power; because of their small sound sources, higher frequencies will be radiated more efficiently than lower frequencies, but in order to produce brief loud pulses from a long-duration muscle contraction they may use both a frequency multiplier mechanism and a mechanical power amplifier. Airborne insect sounds in the range from 1 to 5 kHz tend to have sustained puretone components and a specific pattern of pulses which propagate accurately. Where the song frequency is higher, the pulses tend to become briefer, with a rapid initial build-up that gives a reliable time of onset through obstructed transmission pathways. These scale effects may be related both to the sound-producing mechanism and the auditory system of the receiver. Tiny insects have the special acoustic problem of communicating with only a small amount of available power. Some, such as fruit flies, communicate at low frequencies, at close range, by generating air currents; these currents may also be used to waft specific pheromones. Other small insects, such as Hemiptera, beetles, etc., communicate using substrate vibration. This enables long-range communication, but signal structure degrades with distance from the source; vibration signals tend to be confined to certain types of linear substrate, such as vegetation.     

Structure of the ovary in Steingelia (Sternorrhyncha: Coccinea), and its phylogenetic implications

The paired ovaries of Steingelia gorodetskia are composed of about 100 telotrophic ovarioles devoid of terminal filaments (scale insect autapomorphy). In structure they resemble those of other scale insects, but differ in the following details: (a) all ovarioles develop synchronously, (b) they are suspended to the lateral oviducts by means of long stalks, (c) the tropharium is tubular (unique in scale insects) and (d) consists of 15-35, trophocytes, 2-4 previtellogenic oocytes that further develop, and numerous somatic prefollicular cells, (e) the vitellarium houses 2-4 linearly arranged vitellarial oocytes (versus one in most scale insects). Most of these features must be considered as plesiomorphic corresponding with the conditions in the most primitive Heteroptera. Bacterial endosymbionts have been found in some somatic cells, trophocytes, oocytes and in the nutritive cord. Present results support the opinion, based on external morphology, that the Steingeliidae are closely related to the Ortheziidae, Xylococcidae and Matsucoccidae.     

L'azione degli insetti fitofagi sulle variazioni di sostanza secca nelle foglie

Summary

The Author studies with the weighing method the action of plant-eater insects on the alteration of the substances fresch and dry in the hust leaves and he comes to the following conclusions:

1. The fresh substance stand less alterations in the leaves with the excavated border than in the sound leaves, the dry substance is less in the hust leaves than in those sound.

2. The fresh substance shows little waverigs in the leaves notched on alone side of the border, but the dry substance is always inferior in the hust leaves than in the sound half and in those entire.

3. The substances fresh and dry are less in the bored leaves than in the sound, the dry substance shows little alternations in those bored in one or the other half of the border.

This would indicate that the traumatic action of insects causes in the leaves a depresson of the cellular swuelling, whih determines a bigger mobilization of the dry substanc's elements than in the sound leaves.     

昆虫对植物的选择性以及植物化学成分对昆虫的影响

植物与昆虫之间的关系一直被人们作为重要的研究目标。昆虫依靠绿色植物生存,植物通过自身的化学物质影响昆虫的进化方向,两者形成了复杂的协同进化关系。本文阐述了昆虫在定居、产卵、取食过程中运用不同的嗅觉、味觉、触觉刺激标准来选择适宜的寄主或寄主位置的方法,以及植物体内的化学成分对昆虫的营养作用和通过毒杀、拒食、招引天敌寄生蜂等方式抵御昆虫的进攻。     

Effect of sound signals on spontaneous activity in grasshopper interneurons (Orthoptera,Tettigonidae)

According to one of hypotheses proposed for explaining mechanisms of sound signal recognition in insects, their CNS contains a group of rhythmically active neurons that function as a reference standard for comparison with perceived acoustic information. To check this hypothesis, the spontaneous neuronal activity and its changes in perception of conspecific and heterospecific signals (CS and HS) were analyzed in the CNS of two sympatric grasshopper species Tettigonia cantans and Metrioptera roeselii. The activity of individual neurons was assayed in fixed and freely moving insects. The results of the experiment have shown that in the thoracic part of the CNS there is a group of rhythmically active neurons that do not directly respond to sound signal but readjust their impulses under effect of its action. On presentation of CS the following reactions were observed: attenuation or enhancement of impulses; stabilization or destabilization of rhythm; regular increase or decrease in interspike intervals; phasic readjustments leading to synchronization of impulses with sound stimuli (pulses). No similar alterations were usually produced by HS; still, if they did appear, they were less pronounced or of opposite direction. These data indicate that the grasshopper auditory system affects markedly the rhythmically active neurons, their reaction depending considerably on temporal organization of sound signals. Selectivity of these reactions allows us to suggest that the rhythmically active neurons are directly related to the neuronal networks providing the sound signal recognition.Translated from Zhurnal Evolyutsionnoi Biokhimii i Fiziologii, Vol. 40, No. 6, 2004, pp. 531–538.Original Russian Text Copyright © 2004 by Zhantiev, Korsunovskaya, Chukanov.To the 100-Anniversary of A. K. VoskresenskayaThis revised version was published online in April 2005 with a corrected cover date.