追踪天使——雷达昆虫学30年

雷达昆虫学是一门新的学科分支。从它诞生起的30年来,英、美、澳、中四国的观测研究已初步阐明了昆虫在迁飞过程中的成层、定向、集聚等行为现象及其时空分布,揭示了大气结构和运动对昆虫迁飞的影响,为深化人们对昆虫迁飞行为机制的认识提供了许多令人耳目一新的画面;昆虫雷达技术也逐渐从研究走向实用,已经实现了对迁飞性害虫的长期、自动和实时监测。在全国建立VLR网并与GIS相结合,可望实现对我国重大虫灾的及时预警。     

粘虫飞行定向行为与不同磁场环境的关系

【目的】研究正常地磁场和近零磁场条件下饲养的粘虫Mythimna separata(Walker)夜间飞行定向行为与磁场的关系,为明确迁飞性昆虫远距离迁飞的地磁定向机制提供依据。【方法】分别在正常地磁场和近零磁场条件饲养粘虫,羽化后的粘虫蛾在人工模拟不同的磁场条件下进行夜间飞行定向行为测试,比较粘虫蛾飞行定向行为的差异。【结果】粘虫蛾在正常地磁场条件下均具有显著的群体共同定向行为,夏季测试的粘虫群体共同定向方向为偏北;在近零磁场以及垂直平面分量倒转的地磁场条件下,粘虫蛾群体共同定向行为均消失。同时,不同磁场的生长环境对粘虫飞行定向行为的影响不明显,而飞行测试时的磁场环境对其定向行为有显著影响。【结论】磁场可能是粘虫定向的重要参考依据之一,粘虫对地磁场磁倾角的变化有反应,推测其可能利用了磁倾角进行一定程度上的辅助定向。     

迁飞过程中昆虫的行为

根据国内外对昆虫迁飞的雷达观测结果,综述了迁飞过程中昆虫的成层、定向和集聚行为及其生态学意义。昆虫在迁飞过程中不完全是被动地随风飘流,而是在一定程度上自主地选择了自己的运行轨迹。它们选择在气温最高、风力最大和风向最适的高度成层,并通过定向进一步修饰位移方位,从而可最大限度地利用空气动力到达新的栖息地。同时,不同尺度的大气辐合过程使得迁飞昆虫集聚成足以在某种条件下引起局地爆发的高密度种群,而地面大发生种群形成与否取决于大气辐合的发生频率、强度和寿命,以及集聚起来的昆虫是否降落,在何时何地降落,是否还会再迁出等。阐明昆虫在迁飞过程中的各种行为机制是迁飞性害虫测报和防治的关键所在。     

粘虫蛾体内磁性颗粒初探

[目的]磁性颗粒是一些生物感知地磁场变化的重要物质,也是夜间远距离迁飞昆虫地磁定向的重要机制之一.本研究以迁飞性害虫粘虫蛾Mythimna separata为研究对象,对其体内潜在磁性颗粒进行检测.[方法]利用超导量子磁强计SQUID初步检测粘虫成虫体内的磁性颗粒,并将经普鲁士蓝染色后的虫体石蜡切片于正置BX61研究级...     

昆虫迁飞的调控基础及展望

昆虫迁飞是在长期适应多变的环境过程中进化形成的一种行为对策,也是昆虫的种类和数量繁多,以及迁飞害虫经常暴发成灾的主要原因.昆虫迁飞行为的发生不仅受到外界环境因素的影响,而且受到本身生理因素的调控.目前,国内外对此类研究主要集中在生态环境、生理因素、行为学以及种群遗传学方面的调控机制.随着分子生物学技术的发展,昆虫迁飞行为发生的分子调控机制也越来越受到重视.在对国内外主要昆虫迁飞调控机制概述的基础上,对新的分子生物学技术在昆虫迁飞调控中的应用进行了探讨与展望.     

昆虫迁飞行为的参数化Ⅰ.行为分析

通过对雷达昆虫学研究和其他方法得到的研究成果的综合分析,提出一套昆虫迁飞行为参数化方案。即：起飞时间以日出日没及晨昏朦影时刻为基准,降落时间依目标昆虫的迁飞特性具体取值;运行高度取边界层顶与目标昆虫飞行低温阈限所在高度之间的气流层,运行方向取风向值并以目标昆虫的定向加以修饰,运行速度为风速与目标昆虫自身飞行速度的矢量和。这套方案可作为昆虫迁飞轨迹数值模拟的基础,为迁飞性害虫异地预测提供一种有效的工具     

北京多普勒天气雷达上的昆虫回波分析

【目的】为了解北京多普勒天气雷达上的昆虫回波信息,探索其在迁飞害虫监测预警中的应用。【方法】选取北京南郊观象台新一代天气雷达CINRAD-SA积累的晴空回波数据和风廓线雷达提供的风速风向数据,基于有关软件分析了昆虫回波的特点,讨论了降水等其他因素对昆虫迁飞的影响。【结果】新一代天气雷达可以探测到空中迁飞昆虫引发的后向散射回波,晴空回波日节律符合夜行性昆虫的朦影起飞和日出降落的行为特点,晴空回波出现的时期是3月至11月,中间有2个高峰期;晴空回波数量与风向关系密切,当天气系统合适时,晴空回波会出现成层和共同定向现象;此外,降水会中断昆虫迁飞。【结论】北京多普勒天气雷达可以探测昆虫迁飞,在迁飞性害虫监测预警及其综合防控工作中具有重大的潜在应用价值,今后农业部门在优先建立昆虫雷达网络的同时,应加强与气象部门合作,发挥天气雷达的补充作用,共同提高迁飞性害虫的监测预警水平。     

粘虫蛾体内磁性颗粒初探

[目的]磁性颗粒是一些生物感知地磁场变化的重要物质,也是夜间远距离迁飞昆虫地磁定向的重要机制之一.本研究以迁飞性害虫粘虫蛾Mythimna separata为研究对象,对其体内潜在磁性颗粒进行检测.[方法]利用超导量子磁强计SQUID初步检测粘虫成虫体内的磁性颗粒,并将经普鲁士蓝染色后的虫体石蜡切片于正置BX61研究级显微镜下观察磁性颗粒的分布状况.[结果]SQUID检测发现,相比于头部的磁滞曲线,粘虫腹部具有微弱的磁性,推测粘虫腹部可能具有磁性颗粒.进一步经显微镜观察表明,虫体腹部有明显的普鲁士蓝染色沉淀,证明粘虫蛾腹部存在铁磁性颗粒物质.[结论]粘虫蛾腹部可能是其感应地磁场变化以及地磁定向的重要部位.     

东北粘虫春季空中虫群的迁飞行为

【目的】研究春季迁飞粘虫Mythimna separata(Walker)空中虫群的聚集成层和共同定向等飞行行为。【方法】以2年粘虫迁飞盛期的雷达观测数据为基础,结合空间气象要素和罗盘信号分析。【结果】在温度较低的春季进行迁飞的粘虫多在逆温层顶附近聚集成层,空中虫群会主动选择高温。迁飞粘虫具有明显的飞行低温阈值(16℃左右),此温度以下空中虫群密度低、无聚集成层现象。空中虫群位移方向的离散度与风速呈显著负相关。当风速较大(大于5 m/s)且风向与粘虫迁飞方向(东北)相一致时,虫群通常表现为顺风定向;当风速小于3 m/s且风向与粘虫迁飞方向明显不同时,虫群表现为明显的侧风补偿甚至逆风飞行行为。此外,粘虫可能利用罗盘信号进行迁飞定向,例如地磁场。【结论】试验结果可为迁飞性害虫的异地预测研究提供一定的基础理论依据。     

磁场变化对粘虫飞行定向行为的影响

【目的】研究磁场的变化对于粘虫Mythimna separata(Walker)飞行中定向行为的影响,为揭示空中迁飞虫群的飞行行为机制提供基础理论依据。【方法】在人工模拟磁场条件下对粘虫的定向行为进行了比较研究。【结果】试虫在正常地磁场条件下表现为显著的群体共同定向。粘虫群体的共同定向是轴对称的。当将试虫置于较强的磁场条件下时,粘虫供试个体的定向行为发生变化,群体共同定向行为消失。试虫的定向行为不受磁场水平分量极向变化的影响。【结论】迁飞性昆虫可能利用磁场作为自身定向的罗盘信号,在这个过程中可能和鸟类一样利用了磁倾角。     

Long‐range seasonal migration in insects: mechanisms,evolutionary drivers and ecological consequences

Myriad tiny insect species take to the air to engage in windborne migration, but entomology also has its ‘charismatic megafauna’ of butterflies, large moths, dragonflies and locusts. The spectacular migrations of large day‐flying insects have long fascinated humankind, and since the advent of radar entomology much has been revealed about high‐altitude night‐time insect migrations. Over the last decade, there have been significant advances in insect migration research, which we review here. In particular, we highlight: (1) notable improvements in our understanding of lepidopteran navigation strategies, including the hitherto unsuspected capabilities of high‐altitude migrants to select favourable winds and orientate adaptively, (2) progress in unravelling the neuronal mechanisms underlying sun compass orientation and in identifying the genetic complex underpinning key traits associated with migration behaviour and performance in the monarch butterfly, and (3) improvements in our knowledge of the multifaceted interactions between disease agents and insect migrants, in terms of direct effects on migration success and pathogen spread, and indirect effects on the evolution of migratory systems. We conclude by highlighting the progress that can be made through inter‐phyla comparisons, and identify future research areas that will enhance our understanding of insect migration strategies within an eco‐evolutionary perspective.     

The role of the sun in the celestial compass of dung beetles

Recent research has focused on the different types of compass cues available to ball-rolling beetles for orientation, but little is known about the relative precision of each of these cues and how they interact. In this study, we find that the absolute orientation error of the celestial compass of the day-active dung beetle Scarabaeus lamarcki doubles from 16° at solar elevations below 60° to an error of 29° at solar elevations above 75°. As ball-rolling dung beetles rely solely on celestial compass cues for their orientation, these insects experience a large decrease in orientation precision towards the middle of the day. We also find that in the compass system of dung beetles, the solar cues and the skylight cues are used together and share the control of orientation behaviour. Finally, we demonstrate that the relative influence of the azimuthal position of the sun for straight-line orientation decreases as the sun draws closer to the horizon. In conclusion, ball-rolling dung beetles possess a dynamic celestial compass system in which the orientation precision and the relative influence of the solar compass cues change over the course of the day.     

Unraveling navigational strategies in migratory insects

Long-distance migration is a strategy some animals use to survive a seasonally changing environment. To reach favorable grounds, migratory animals have evolved sophisticated navigational mechanisms that rely on a map and compasses. In migratory insects, the existence of a map sense (sense of position) remains poorly understood, but recent work has provided new insights into the mechanisms some compasses use for maintaining a constant bearing during long-distance navigation. The best-studied directional strategy relies on a time-compensated sun compass, used by diurnal insects, for which neural circuits have begun to be delineated. Yet, a growing body of evidence suggests that migratory insects may also rely on other compasses that use night sky cues or the Earth's magnetic field. Those mechanisms are ripe for exploration.     

Compass orientation and possible migration routes of passerine birds at high arctic latitudes

The use of celestial or geomagnetic orientation cues can lead migratory birds along different migration routes during the migratory journeys, e.g. great circle routes (approximate), geographic or magnetic loxodromes. Orientation cage experiments have indicated that migrating birds are capable of detecting magnetic compass information at high northern latitudes even at very steep angles of inclination. However, starting a migratory journey at high latitudes and following a constant magnetic course often leads towards the North Magnetic Pole, which means that the usefulness of magnetic compass orientation at high latitudes may be questioned. Here, we compare possible long‐distance migration routes of three species of passerine migrants breeding at high northern latitudes. The initial directions were based on orientation cage experiments performed under clear skies and simulated overcast and from release experiments under natural overcast skies. For each species we simulated possible migration routes (geographic loxodrome, magnetic loxodrome and sun compass route) by extrapolating from the initial directions and assessing a fixed orientation according to different compass mechanisms in order to investigate what orientation cues the birds most likely use when migrating southward in autumn. Our calculations show that none of the compass mechanisms (assuming fixed orientation) can explain the migration routes followed by night‐migrating birds from their high Nearctic breeding areas to the wintering sites further south. This demonstrates that orientation along the migratory routes of arctic birds (and possibly other birds as well) must be a complex process, involving different orientation mechanisms as well as changing compass courses. We propose that birds use a combination of several compass mechanisms during a migratory journey with each of them being of a greater or smaller importance in different parts of the journey, depending on environmental conditions. We discuss reasons why birds developed the capability to use magnetic compass information at high northern latitudes even though following these magnetic courses for any longer distance will lead them along totally wrong routes. Frequent changes and recalibrations of the magnetic compass direction during the migratory journey are suggested as a possible solution.     

Hierarchy of sun,beach slope,and landmarks as cues for Y-axis orientation of the supratidal amphipod Talorchestia longicornis (Say)

The semi-terrestrial amphipod Talorchestia longicornis (Say) undergoes Y-axis orientation and has a hierarchy among orientation cues. A previous study found that they used sun compass orientation and moved in the onshore direction of the home beach in both air and water. The present study determined whether this species could also use local landmarks and beach slope as orientation cues. They oriented upslope in simulated darkness in the laboratory on both dry and wet sand with threshold slopes of 2° and 4°, respectively. When tested outside in an arena in air on wet sand, they were disoriented when sun, slope, and landmarks were absent as cues. If presented with single cues, they moved upslope, toward landmarks and in the up-beach direction of the home beach during sun compass orientation. Using paired cues, sun was dominant over slope and landmarks, while slope was dominant over landmarks. In the presence of all three cues, amphipods displayed sun compass orientation in all test combinations except when slope and landmarks were paired together against the sun, which evoked a bimodal response. Thus, the hierarchy of cues for up-beach movement of T. longicornis during Y-axis orientation is the sun, then the slope, and finally the landmarks.     

Keynote papers on sandhopper orientation and navigation

This article analyses the relevant studies that have made sandhoppers a model subject for the study of orientation, and traces the development of the paradigm through innovative hypotheses and empirical evidence. Sandhoppers are able to maintain their direction without sensorial contact with the goal, which is their burrowing zone extended along the beach, but very narrow across it. They actively determine the direction of their movements, according to their internal state and the environmental features encountered. Each population shows an 'innate directional tendency' adapted to the shoreline of origin, and the inexpert laboratory-born young behave in a similar way to the adults. Genetic differences have been demonstrated between, as well as within natural populations. The question of the calibration of the sun compass to orientation on a particular shoreline implies a redundancy of mechanisms of orientation. Orientation mechanisms may involve environmental cues perceived through diverse sensory modalities, and range from simple orientation reflexes to sun compass navigational systems. These include scototaxis and geotaxis, and the response to the silhouette of the dune, in addition to sun and moon orientation, which is dependent on the time of the day and orientates daily migrations on the beach. Different modalities of orientation may operate singly, or in conjunction with each other, and their ecological significance may vary according to the habitat and lifestyle of the animals. Taken collectively, the orientation behaviour of the group appears to be a most accommodating phenotype, with considerable adaptive potential. The evidence from comparative studies of different populations promotes consideration of behavioural plasticity as an adaptation to changing coastlines.     

Are orientation mechanisms among migratory birds speciesospecific?

The mechanisms by which migratory birds find their way from breeding grounds to winter quarters and back have been the subject of intensive research during the past four decades. Birds are equipped with genetic information about the migratory direction, and they can use the earth's magnetic field, star patterns and the sun and/or skylight polarization patterns as compass references. Studies on a number of North American and European species have suggested possible species-specific differences in the relative role of the compass mechanisms. This may be largely the result of divergent experimental designs, which make results difficult to compare. Comparative studies with identical methods are needed to see how much species-specific variation exists in basic orientation mechanisms.     

Avian navigation and geographic positioning

Site fidelity to breeding and wintering grounds, and even stopover sites, suggests that passerines are capable of accurate navigation during their annual migrations. Geolocator‐based studies are beginning to demonstrate precise population‐specific migratory routes and even some interannual consistency in individual routes. Displacement studies of birds have shown that at least adult birds are capable of goal‐oriented movements, likely involving some type of map or geographic position system. In contrast, juveniles on their first migration use a clock‐and‐compass orientation strategy, with limited knowledge about locations along their migratory routes. Positioning information could come from a variety of cues including visual, olfactory, acoustic, and geomagnetic sources. How information from these systems is integrated and used for avian navigation has yet to be fully articulated. In this review, we (1) define geographic positioning and distinguish the types of navigational strategies that birds could use for orientation, (2) describe sensory cues available to birds for geographic positioning, (3) review the evidence for geographic positioning in birds and methods used to collect that evidence, and (4) discuss ways ornithologists, particularly field ornithologists, can contribute to and advance our knowledge of the navigational abilities of birds. Few studies of avian orientation and navigation mechanisms have been conducted in the Western Hemisphere. To fully understand migratory systems in the Western Hemisphere and develop better conservation policies, information about the orientation and navigation mechanisms used by specific species needs to be integrated with other aspects of their migration ecology and biology.     

Magnetic compass sense in the large yellow underwing moth, Noctua pronuba L.

Many animals are now known to have a magnetic sense which they use when moving from one place to another. Among insects, this sense has only been studied in any detail in the honey bee. A role for a magnetic compass sense in cross-country migration has not so far been demonstrated for any insect. On clear nights the large yellow underwing moth, Noctua pronuba, has been shown to orientate by both the moon and the stars. However, radar studies have shown moths to be well-oriented on overcast nights as well as clear nights. We report here that when large yellow underwings are placed in an orientation cage on overcast nights and the Earth's normal magnetic field is reversed, there is a corresponding reversal in the orientation of the moth. We conclude that this species makes use of the Earth's magnetic field in maintaining compass orientation on overcast nights. We also show that the preferred compass orientation to the Earth's magnetic field is the same as the compass direction that results from orientation to the moon and stars.     

Das Orientierungssystem der V?gel I. Kompa?mechanismen

Zusammenfassung V?gel stellen den Bezug zum Ziel indirekt über ein externes Referenzsystem her. Der Navigationsproze? besteht deshalb aus zwei Schritten: zun?chst wird die Richtung zum Ziel als Kompa?kurs festgelegt, dann wird dieser Kurs mit Hilfe eines Kompa?mechanismus aufgesucht. Das Magnetfeld der Erde und Himmelsfaktoren werden von den V?gel als Kompa? benutzt. In der vorliegenden Arbeit werden der Magnetkompa?, der Sonnenkompa? und der Sternkompa? der V?gel in ihrer Funktionsweise, ihrer Entstehung und ihrer biologischen Bedeutung vorgestellt. Der Magnetkompa? erwies sich als Inklinationskompa?, der nicht auf der Polarit?t, sondern auf der Neigung der Feldlinien im Raum beruht; er unterscheidet „polw?rts“ und „?quatorw?rts“ statt Nord und Süd. Er ist ein angeborener Mechanismus und wird beim Vogelzug und beim Heimfinden benutzt. Seine eigentliche Bedeutung liegt jedoch darin, da? er ein Referenzsystem bereitstellt, mit dessen Hilfe andere Orientierungsfaktoren zueinander in Beziehung gesetzt werden k?nnen. Der Sonnenkompa? beruht auf Erfahrung; Sonnenazimut, Tageszeit und Richtung werden durch Lernprozesse miteinander verknüpft, wobei der Magnetkompa? als Richtungsreferenzsystem dient. Sobald er verfügbar ist, wird der Sonnenkompa? bei der Orientierung im Heimbereich und beim Heimfinden bevorzugt benutzt; beim Vogelzug spielt er, wahrscheinlich wegen seiner Abh?ngigkeit von der geographischen Breite, kaum eine Rolle. Der Sternkompa? arbeitet ohne Beteiligung der Inneren Uhr; die V?gel leiten Richtungen aus den Konfigurationen der Sterne zueinander ab. Lernprozesse erstellen den Sternkompa? in der Phase vor dem ersten Zug; dabei fungiert die Himmelsrotation als Referenzsystem. Sp?ter, w?hrend des Zuges, übernimmt der Magnetkompa? diese Rolle. Die relative Bedeutung der verschiedenen Kompa?systeme wurde in Versuchen untersucht, bei denen Magnetfeld und Himmelsfaktoren einander widersprechende Richtungs-information gaben. Die erste Reaktion der V?gel war von Art zu Art verschieden; langfristig scheinen sich die V?gel jedoch nach dem Magnetkompa? zu richten. Dabei werden die Himmelsfaktoren umgeeicht, so da? magnetische Information und Himmelsinformation wieder im Einklang stehen. Der Magnetkompa? und die Himmelsfaktoren erg?nzen einander: der Magnetkompa? ersetzt Sonnen- und Sternkompa? bei bedecktem Himmel; die Himmelsfaktoren erleichtern den V?geln das Richtungseinhalten, zu dem der Magnetkompa? offenbar wenig geeignet ist. Magnetfeld und Himmelsfaktoren sollten deshalb als integrierte Komponenten eines multifaktoriellen Systems zur Richtungsorientierung betrachtet werden.

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The orientation system of birds — I. Compass mechanisms

Summary Because of the large distances involved, birds establish contact with their goal indirectly via an external reference. Hence any navigation is a two-step process: in the first step, the direction to the goal is determined as a compass course; in the second step, this course is located with a compass. The geomagnetic field and celestial cues provide birds with compass information. The magnetic compass of birds, the sun compass the star compass and the interactions between the compass mechanisms are described in the present paper. Magnetic compass orientation was first demonstrated by testing night-migrating birds in experimentally altered magnetic fields: the birds changed their directional tendencies according to the deflected North direction. The avian magnetic compass proved to be an inclination compass: it does not use polarity; instead it is based on the axial course of the field lines and their inclination in space, distinguishing “poleward” and “equatorward” rather than North and South. Its functional range is limited to intensities around the local field strength, but this biological window is flexible and can be adjusted to other intensities. The magnetic compass is an innate mechanism that is widely used in bird migration and in homing. Its most important role, however, is that of a basic reference system for calibrating other kinds of orientation cues. Sun compass orientation is demonstrated by clock-shift experiments: Shifting the birds' internal clock causes them to misjudge the position of the sun, thus leading to typical deflections which indicate sun compass use. The analysis of the avian sun compass revealed that it is based only on sun azimuth and the internal clock; the sun's altitude is not involved. The role of the pattern of polarized light associated with the sun is unclear; only at sunset has it been shown to be an important cue for nocturnal migrants, being part of the sun compass. The sun compass is based on experience; sun azimuth, time of day and direction are combined by learning processes during a sensitive period, with the magnetic compass serving as directional reference. When established, the sun compass becomes the preferred compass mechanism for orientation tasks within the home region and homing: in migration, however, its role is minimal, probably because of the changes of the sun's arc with geographic latitude. The star compass was demonstrated in night-migrating birds by projecting the northern stars in different directions in a planetarium. The analysis of the mechanism revealed that the internal clock is not involved; birds derive directions from the spatial relationship of the star configurations. The star compass is also established by experience; the directional reference is first provided by celestial rotation, later, during migration, by the magnetic compass. The relative importance of the various compass mechanisms has been tested in experiments in which celestial and magnetic cues gave conflicting information. The first response of birds to conflicting cues differs considerably between species; after repeated exposures, however, the birds oriented according to magnetic North, indicating a long-term dominance of the magnetic compass. Later tests in the absence of magnetic information showed that celestial cues were not simply ignored, but recalibrated so that they were again in agreement with magnetic cues. The magnetic compass and celestial cues complement each other: the magnetic field ensures orientation under overcast sky; celestial cues facilitate maintaining directions, for which the magnetic compass appears to be ill suited. In view of this, the magnetic field and celestial cues should be regarded as integrated components of a multifactorial system for directional orientation.