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.

     

Behavioural and physiological mechanisms of polarized light sensitivity in birds

Polarized light (PL) sensitivity is relatively well studied in a large number of invertebrates and some fish species, but in most other vertebrate classes, including birds, the behavioural and physiological mechanism of PL sensitivity remains one of the big mysteries in sensory biology. Many organisms use the skylight polarization pattern as part of a sun compass for orientation, navigation and in spatial orientation tasks. In birds, the available evidence for an involvement of the skylight polarization pattern in sun-compass orientation is very weak. Instead, cue-conflict and cue-calibration experiments have shown that the skylight polarization pattern near the horizon at sunrise and sunset provides birds with a seasonally and latitudinally independent compass calibration reference. Despite convincing evidence that birds use PL cues for orientation, direct experimental evidence for PL sensitivity is still lacking. Avian double cones have been proposed as putative PL receptors, but detailed anatomical and physiological evidence will be needed to conclusively describe the avian PL receptor. Intriguing parallels between the functional and physiological properties of PL reception and light-dependent magnetoreception could point to a common receptor system.     

Avian magnetic compass: Its functional properties and physical basis

The avian magnetic compass was analyzed in bird species of three different orders - Passeriforms, Columbiforms and Galliforms - and in three different behavioral contexts, namely migratory orientation, homing and directional conditioning. The respective findings indicate similar functional properties: it is an inclination compass that works only within a functional window around the ambient magnetic field intensity, it tends to be lateralized in favor of the right eye, and it is wavelength-dependent, requiring light from the short-wavelength range of the spectrum. The underlying physical mechanisms have been identified as radical pair processes, spin-chemical reactions in specialized photopigments. The iron-based receptors in the upper beak do not seem to be involved. The existence of the same type of magnetic compass in only very distantly related bird species suggests that it may have been present already in the common ancestors of all modern birds, where it evolved as an all-purpose compass mechanism for orientation within the home range.     

Technical note: Occlusal fingerprint analysis: Quantification of tooth wear pattern

Information about food ingestion and mastication behavior during the lifespan of an individual is encoded in the dental occlusal wear pattern. To decode this information, we describe a new method called occlusal fingerprint analysis (OFA). Structural parameters of wear facets on the occlusal surface of teeth are quantified from digitized casts for the interpretation of occlusal aspects. The OFA provides an individual three‐dimensional dental occlusal compass that indicates the major pathways of interaction between antagonists, revealing information about development, spatial position, and enlargement of wear facets. Humans develop a very similar overall pattern of crown contacts, although specific characteristics of wear facets reflect an individual's occlusal relationships and masticatory behavior. We hypothesize that the wear pattern is a unique character and therefore valuable for individual identification. Furthermore we suggest that OFA, when further developed, may be useful for identification of behavioral, biological, and chemical factors affecting crown morphology. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Light pollution forces a change in dung beetle orientation behavior

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Field evidence of compass orientation in migrating juvenile grunts (Haemulidae)

Juvenile French and white grunts, Haemulon flavolineatum (Desmarest) and H. plumieri (Lacé-pède), were captured during their daily migrations between diurnal resting sites on coral patch reefs and nocturnal feeding grounds in seagrass beds. Grunts captured during morning and evening migrations were released on the route and after displacement up to 100 m or 5 km away. Grunts generally moved in the direction which would have taken them back towards their home reef or to their accustomed feeding sites, indicating that familiar landmarks are not essential for orientation. The spatial precision of migration may serve to partition the feeding area most efficiently. The timing of migrations is also very precise, and appears to be adapted to reduce the vulnerability of grunts to predation near their home reef.     

Magnetic compass orientation by larval Drosophila melanogaster

We report evidence for magnetic compass orientation by larval Drosophila melanogaster. Groups of larvae were exposed from the time of hatching to directional ultraviolet (365 nm) light emanating from one of four magnetic directions. Larvae were then tested individually on a circular agar plate under diffuse light in one of four magnetic field alignments. The larvae exhibited magnetic compass orientation in a direction opposite that of the light source in training. Evidence for a well-developed magnetic compass in a larval insect that moves over distances of at most a few tens of centimeters has important implications for understanding the adaptive significance of orientation mechanisms like the magnetic compass. Moreover, the development of an assay for studying magnetic compass orientation in larval D. melanogaster will make it possible to use a wide range of molecular genetic techniques to investigate the neurophysiological, biophysical, and molecular mechanisms underlying the magnetic compass.     

生物磁学在鸟类定向研究中的进展

地球上广泛存在的地磁场能够为导航提供可靠的信息,因此很多鸟类在迁徙和归巢过程中都使用地磁信息来保证航行方向的正确,在迁徙的鸟类中已经发现有18种是利用地磁罗盘进行定向和导航的。本文从鸟类使用的磁罗盘、航行地图以及磁感应机制等几方面阐述了目前在鸟类生物磁学方面的研究进展。     

Neural mechanisms in insect navigation: polarization compass and odometer

Insect navigation relies on path integration, a procedure by which information about compass bearings pursued and distances travelled are combined to calculate position. Three neural levels of the polarization compass, which uses the polarization of skylight as a reference, have been analyzed in orthopteran insects. A group of dorsally directed, highly specialized ommatidia serve as polarization sensors. Polarization-opponent neurons in the optic lobe condition the polarization signal by removing unreliable and irrelevant components of the celestial stimulus. Neurons found in the central complex of the brain possibly represent elements of the compass output. The odometer for measuring travelling distances in honeybees relies on optic flow experienced during flight, whereas desert ants most probably use proprioreceptive cues.     

Microtrichs of amphipod crustacea. Morphology and distribution

Microscopical cuticular structures of four amphipods, presumed to be sensory in function are described, using a scanning electron microscope. Gammarus sp, Liljeborgia sp and Orchomene sp showed very small peg‐like microtrichs emerging from cuticular pits of less than one micron in diameter. Single units, facing distally, were ranged in rows, several rows occupying a single integumental polygon. Density of rows and number of units per row were always higher in central areas. The distribution, organization and relationships with larger sensilla, suggest that microtrichs are specialized in detection of slow or weak water currents. Orchestia showed short sensilla coeloconica. The difference between this species and the other three may be related to its different habitat, which is terrestrial.     

The quantum Zeno effect immunizes the avian compass against the deleterious effects of exchange and dipolar interactions

Magnetic-sensitive radical-ion-pair reactions are understood to underlie the biochemical magnetic compass used by avian species for navigation. Recent experiments have provided growing evidence for the radical-ion-pair magnetoreception mechanism, while recent theoretical advances have unravelled the quantum nature of radical-ion-pair reactions, which were shown to manifest a host of quantum-information-science concepts and effects, like quantum measurement, quantum jumps and the quantum Zeno effect. We here show that the quantum Zeno effect provides for the robustness of the avian compass mechanism, and immunizes its magnetic and angular sensitivity against the deleterious and molecule-specific exchange and dipolar interactions.     

Development of lateralization of the magnetic compass in a migratory bird

The magnetic compass of a migratory bird, the European robin (Erithacus rubecula), was shown to be lateralized in favour of the right eye/left brain hemisphere. However, this seems to be a property of the avian magnetic compass that is not present from the beginning, but develops only as the birds grow older. During first migration in autumn, juvenile robins can orient by their magnetic compass with their right as well as with their left eye. In the following spring, however, the magnetic compass is already lateralized, but this lateralization is still flexible: it could be removed by covering the right eye for 6 h. During the following autumn migration, the lateralization becomes more strongly fixed, with a 6 h occlusion of the right eye no longer having an effect. This change from a bilateral to a lateralized magnetic compass appears to be a maturation process, the first such case known so far in birds. Because both eyes mediate identical information about the geomagnetic field, brain asymmetry for the magnetic compass could increase efficiency by setting the other hemisphere free for other processes.     

Honeybee navigation: critically examining the role of the polarization compass

Although it is widely accepted that honeybees use the polarized-light pattern of the sky as a compass for navigation, there is little direct evidence that this information is actually sensed during flight. Here, we ask whether flying bees can obtain compass cues derived purely from polarized light, and communicate this information to their nest-mates through the ‘waggle dance’. Bees, from an observation hive with vertically oriented honeycombs, were trained to fly to a food source at the end of a tunnel, which provided overhead illumination that was polarized either parallel to the axis of the tunnel, or perpendicular to it. When the illumination was transversely polarized, bees danced in a predominantly vertical direction with waggles occurring equally frequently in the upward or the downward direction. They were thus using the polarized-light information to signal the two possible directions in which they could have flown in natural outdoor flight: either directly towards the sun, or directly away from it. When the illumination was axially polarized, the bees danced in a predominantly horizontal direction with waggles directed either to the left or the right, indicating that they could have flown in an azimuthal direction that was 90° to the right or to the left of the sun, respectively. When the first half of the tunnel provided axial illumination and the second half transverse illumination, bees danced along all of the four principal diagonal directions, which represent four equally likely locations of the food source based on the polarized-light information that they had acquired during their journey. We conclude that flying bees are capable of obtaining and signalling compass information that is derived purely from polarized light. Furthermore, they deal with the directional ambiguity that is inherent in polarized light by signalling all of the possible locations of the food source in their dances, thus maximizing the chances of recruitment to it.     

Spectral information as an orientation cue in dung beetles

During the day, a non-uniform distribution of long and short wavelength light generates a colour gradient across the sky. This gradient could be used as a compass cue, particularly by animals such as dung beetles that rely primarily on celestial cues for orientation. Here, we tested if dung beetles can use spectral cues for orientation by presenting them with monochromatic (green and UV) light spots in an indoor arena. Beetles kept their original bearing when presented with a single light cue, green or UV, or when presented with both light cues set 180° apart. When either the UV or the green light was turned off after the beetles had set their bearing in the presence of both cues, they were still able to maintain their original bearing to the remaining light. However, if the beetles were presented with two identical green light spots set 180° apart, their ability to maintain their original bearing was impaired. In summary, our data show that ball-rolling beetles could potentially use the celestial chromatic gradient as a reference for orientation.     

昆虫定向机制研究进展

许多昆虫具有定向运动的行为。对部分社会性昆虫和迁飞性昆虫定向行为的大量研究已经初步阐明太阳、地磁场、天体、风及地面标志物等都可能成为昆虫返巢和迁飞定向的线索。社会性昆虫具有对不同定向线索进行整合而实现精确导航的能力。日间迁飞性昆虫利用时间补偿太阳罗盘进行定向的机制亦已明确,但夜间迁飞昆虫的定向机制尚需深入研究。迁飞性害虫定向机制的明确将有助于判断害虫迁飞路径及降落区域,为迁飞害虫的准确预测提供科学依据。本文对昆虫的定向机制研究进展进行了综述。     

Orientation and Navigation in Elasmobranchs: Which Way Forward?

Elasmobranchs possess a multiplicity of mechanisms controlling posture and short distance orientation. Visual–vestibular contributions to posture and locomotion are well documented. So too, are the contributions of vision, olfaction and the octavolateralis senses to short distance orientation, particularly orientation to specific environmental stimuli such as those generated by prey. Less well understood are the mechanisms guiding orientation over longer distances. Anecdotal and systematic observations of behaviour show tidal, daily, repeat long distance, and even seasonal movement patterns. True navigation has not been demonstrated in elasmobranchs and the sensory mechanisms underlying the above movement patterns remain largely speculative. However, they are likely to include responses to water currents, and physical parameters such as temperature, pressure, and the geomagnetic field. Of particular interest in elasmobranchs is that geomagnetic orientation could be mediated directly via a magnetite based sensory system, or indirectly via the electrosensory system. Systematic studies of movement patterns and experimental studies of the underlying mechanisms of orientation are required to gain an increased understanding of orientation and navigation in this intriguing group.     

Geomagnetic Reversals and Genome Imprinting

If it is more fundamental to formulate biological expression in terms of electromagnetic fields, does this also imply that living things are especially sensitive to the external electromagnetic environment? Specifically, we examine possible genomic effects due to reversals of the geomagnetic field. To maintain sensitivity following a reversal, the Wiltschko hypothesis for the avian magnetic compass can be subsumed under an NB imprinting paradigm, where N is the horizontal vector pointing to magnetic north and B the geomagnetic field vector. Even with a compass that is invariant under reversals, there are nevertheless potential difficulties due to discontinuities in the magnitude of the field during the transition between one chron and the next. Indeed, transitions may be one reason for other-than-magnetic avian auxiliary compasses. Additional problems may also arise during transitions because of high rates of change in B. However, the largest reported dB/dt (Steens Mountain event) is estimated at 1 /u.T/day, seemingly too small to induce significant Faraday current density. Reversals may have also helped determine the nature of the interaction mechanism between GMF and living systems. Mechanisms based on fixed magnetic moments may not be capable of adapting to the reversal process. A better case can be made for an ion cyclotron resonance interaction. Direct involvement in the cell-signaling activities of biological ions would provide such flexibility, and also point to a broader role for the GMF in modulating CNS function than merely to provide orientation.     

Stochastic dynamics of magnetosomes and a mechanism of biological orientation in the geomagnetic field

The rotations of nanoscopic magnetic particles, magnetosomes, embedded into the cytoskeleton are considered. Under the influence of thermal disturbances, a great number of magnetosomes are shown to move chaotically between two stable equilibrium positions, in which their magnetic moments are neither parallel nor antiparallel to the static Earth's magnetic field (MF). The random rotations attain the value of order of a radian. The rate of the transitions and the probability of magnetosomes to be in the different states depend on the MF direction with respect to an averaged magnetosome's orientation. This effect explains the ability of migratory animals to orient themselves faultlessly in long term passages in the absence of the direct visibility of optical reference points. The sensitivity to deviation from an "ideal" orientation is estimated to be 2-4 degrees. Possible involvement of the stochastic dynamics of magnetosomes in biological magnetic navigation is discussed.