Kompass im Kopf

Ant compass – how desert ants learn to navigate Successful spatial orientation is a daily challenge for many animals. Cataglyphis desert ants are famous for their navigational performances. They return to the nest after extensive foraging trips without any problems. How do ants take their navigational systems into operation? After conducting different tasks in the dark nest for several weeks, they become foragers under bright sun light. This transition requires both a drastic switch in behavior and neuronal changes in the brain. Experienced foragers mainly rely on visual cues. They use a celestial compass and landmark panoramas. For that reason, naïve ants perform stereotype learning walks to calibrate their compass systems and acquire information about the nest's surroundings. During their learning walks, the ants frequently look back to the nest entrance to learn the homing direction. For aligning their gazes, they use the earth's magnetic field as a compass reference. This magnetic compass in Cataglyphis ants was previously unknown.     

The role of geomagnetic cues in green turtle open sea navigation

Background

Laboratory and field experiments have provided evidence that sea turtles use geomagnetic cues to navigate in the open sea. For instance, green turtles (Chelonia mydas) displaced 100 km away from their nesting site were impaired in returning home when carrying a strong magnet glued on the head. However, the actual role of geomagnetic cues remains unclear, since magnetically treated green turtles can perform large scale (>2000 km) post-nesting migrations no differently from controls.

Methodology/Principal Findings

In the present homing experiment, 24 green turtles were displaced 200 km away from their nesting site on an oceanic island, and tracked, for the first time in this type of experiment, with Global Positioning System (GPS), which is able to provide much more frequent and accurate locations than previously used tracking methods. Eight turtles were magnetically treated for 24–48 h on the nesting beach prior to displacement, and another eight turtles had a magnet glued on the head at the release site. The last eight turtles were used as controls. Detailed analyses of water masses-related (i.e., current-corrected) homing paths showed that magnetically treated turtles were able to navigate toward their nesting site as efficiently as controls, but those carrying magnets were significantly impaired once they arrived within 50 km of home.

Conclusions/Significance

While green turtles do not seem to need geomagnetic cues to navigate far from the goal, these cues become necessary when turtles get closer to home. As the very last part of the homing trip (within a few kilometers of home) likely depends on non-magnetic cues, our results suggest that magnetic cues play a key role in sea turtle navigation at an intermediate scale by bridging the gap between large and small scale navigational processes, which both appear to depend on non-magnetic cues.     

Marine turtles use geomagnetic cues during open-sea homing

Marine turtles are renowned long-distance navigators, able to reach remote targets in the oceanic environment; yet the sensory cues and navigational mechanisms they employ remain unclear [1, 3]. Recent arena experiments indicated an involvement of magnetic cues in juvenile turtles' homing ability after simulated displacements [4, 5], but the actual role of geomagnetic information in guiding turtles navigating in their natural environment has remained beyond the reach of experimental investigations. In the present experiment, twenty satellite-tracked green turtles (Chelonia mydas) were transported to four open-sea release sites 100-120 km from their nesting beach on Mayotte island in the Mozambique Channel; 13 of them had magnets attached to their head either during the outward journey or during the homing trip. All but one turtle safely returned to Mayotte to complete their egg-laying cycle, albeit with indirect routes, and showed a general inability to take into account the deflecting action of ocean currents as estimated through remote-sensing oceanographic measurements [7]. Magnetically treated turtles displayed a significant lengthening of their homing paths with respect to controls, either when treated during transportation or when treated during homing. These findings represent the first field evidence for the involvement of geomagnetic cues in sea-turtle navigation.     

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.     

Brieftauben

Homing pigeons Homing pigeons are well known for their excellent homing abilities which allow them to return to their lofts from unknown releasing sites more than hundreds of kilometres away. Several orientation mechanisms – sun compass, earth's magnetic field, olfactory cues, visual cues – are known to be involved in homing performance as well as parameters such as motivation and experience. New technology give an insight in their homing behaviour and track preferences and it is shown that homing pigeons physiology and neurobiology seem to be functionally adapted to homing. Pigeons races are still common and it is shown how the pigeon breeder tries to maximize the success of his pigeons.     

Avian navigation: from historical to modern concepts

Studies on avian navigation began at the end of the 19th century with testing various hypotheses, followed by large-scale displacement experiments to assess the capacity of the birds' navigational abilities. In the 1950s, the first theoretical concepts were published. Kramer proposed his ‘Map-and-Compass’ model, assuming that birds establish the direction to a distant goal with the help of an external reference, a compass. The model describes homing as a two-step process, with the first step determining the direction to the goal as a compass course and the second step locating this course with the help of a compass. This model was widely accepted when numerous experiments with clock-shifted pigeons demonstrated the use of the sun compass, and thus a general involvement of compass orientation, in homing. The ‘map’ step is assumed to use local site-specific information, which led to the idea of a ‘grid map’ based on environmental gradients. Kramer's model still forms the basis of our present concept on avian homing, yet route integration with the help of an external reference provides an alternative strategy to determine the home course, and the magnetic compass is a second compass mechanism available to birds. These mechanisms are interrelated by ontogenetic learning processes. A two-step process, with the first step providing the compass course and the second step locating this course with the help of a compass, appears to be a common feature of avian navigation tasks, yet the origin of the compass courses differs between tasks according to their nature, with courses acquired by experience for flights within the home range, courses based on navigational processes for returning home, and courses derived from genetically coded information in first-time migrants. Compass orientation thus forms the backbone of the avian navigational system. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

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.

     

Changes in the short-term deflector loft effect are linked to the sun compass of homing pigeons

Despite being the most studied of all avian orientation systems, important questions still remain about the sun compass of homing pigeons, Columba livia. White it is well-documented that the sun compass is usually learned by young pigeons during the first 10–12 weeks of life, the mechanism by which it is calibrated to adjust for seasonal changes in the sun's azimuth is not known with certainty. Previous experiments using short-term deflector loft pigeons indicated that the sun compass may be calibrated by referencing celestial polarization patterns. The present paper describes important measurable changes in the previously reported orientation behaviour of short-term deflector loft birds, and suggests a correlation between these changes and the presence of a massive upper-atmospheric dust cloud of volcanic origin which significantly altered natural skylight polarization patterns in 1982 and 1983. Moreover, it is shown that when the short-term effect was absent (at times when data from previous years suggested it should be present), the birds were also not using sun compass orientation, as demonstrated by their failure to show the standard ‘clockshift’ response to a 6-h fast shift of their internal clocks. These results support the hypothesis that reflected light cues, rather than odours, are the basis of the deflector loft effect in pigeon homing.     

Homing pigeons develop local route stereotypy

The mechanisms used by homing pigeons (Columba livia) to navigate homeward from distant sites have been well studied, yet the mechanisms underlying navigation within, and mapping of, the local familiar area have been largely neglected. In the local area pigeons pote ntially have access to a powerful navigational aid--a memorized landscape map. Current opinion suggests that landmarks are used only to recognize a familiar start position and that the goalward route is then achieved solely using compass orientation. We used high-resolution global positioning system (GPS) loggers to track homing pigeons as they became progressively familiar with a local homing task. Here, we demonstrate that birds develop highly stereotyped yet individually distinctive routes over the landscape, which remain substantially inefficient. Precise aerial route recapitulation implies close control by localized geocentric cues. Magnetic cues are unlikely to have been used, since recapitulation remains despite magnetic disruption treatment, and olfactory cues would have been positionally unstable under the variable wind conditions, making visual landmarks the most likely cues used.     

Perceptual Strategies of Pigeons to Detect a Rotational Centre—A Hint for Star Compass Learning?

Birds can rely on a variety of cues for orientation during migration and homing. Celestial rotation provides the key information for the development of a functioning star and/or sun compass. This celestial compass seems to be the primary reference for calibrating the other orientation systems including the magnetic compass. Thus, detection of the celestial rotational axis is crucial for bird orientation. Here, we use operant conditioning to demonstrate that homing pigeons can principally learn to detect a rotational centre in a rotating dot pattern and we examine their behavioural response strategies in a series of experiments. Initially, most pigeons applied a strategy based on local stimulus information such as movement characteristics of single dots. One pigeon seemed to immediately ignore eccentric stationary dots. After special training, all pigeons could shift their attention to more global cues, which implies that pigeons can learn the concept of a rotational axis. In our experiments, the ability to precisely locate the rotational centre was strongly dependent on the rotational velocity of the dot pattern and it crashed at velocities that were still much faster than natural celestial rotation. We therefore suggest that the axis of the very slow, natural, celestial rotation could be perceived by birds through the movement itself, but that a time-delayed pattern comparison should also be considered as a very likely alternative strategy.     

Homing pigeons as a model for avian navigation?

The findings on the navigational mechanisms of homing pigeons and the available data on those of wild birds, in particular migrants, are compared. There are important parallels in the use of the magnetic field and the sun for directional orientation. Also the findings on the navigational ‘map’, its preferred use by experienced birds and the strategy of using route information to acquire the necessary knowledge to establish the ‘map’, obtained in pigeons studies, can probably be generalized to wild birds and migrants in their home region. It seems that birds share a common navigational system. Special development of migratory birds, however, is the innate migration program that enables young first‐time migrants to reach their still unknown wintering area.     

Multi-modal Orientation Cues in Homing Pigeons

How homing pigeons displaced into unfamiliar territory findtheir way home has been the subject of extensive experimentationand debate. One reason for the controversy is that pigeons seemto use multiple cues. Clock-shifting experiments show that experiencedpigeons use the sun as a preferred compass; when it is not availablethey rely on magnetic cues. That pigeons can home successfullywhile wearing frosted lenses suggests that landmarks, whilenot an essential navigational cue, are important in the finalstages. The sensory basis of the "map" or position finding systemis probably equally or even more complicated. When conditionsaround the loft are suitable, pigeons may use olfactory cuesto find their way or might use some feature of the earth's magneticfield for their navigation. The Wiltschkos (1989) showed thatpigeons raised without free access to ambient odors are notdisoriented when anosmic while their siblings raised with freeaccess to the prevailing wind were disoriented. Similarly, siblingpigeons from two lofts in Lincoln, Massachusetts. were welloriented or totally disoriented when released at magnetic anomaliesunder sunny skies depending upon which of the two lofts theyhad been reared in. All of these experiments and many more suggestthat pigeons use multiple and redundant cues to find their wayhome. Further, there is the suggestion that which cues theyadopt may well be influenced by the characteristics of the areaaround the home loft in which they were reared.     

Navigational challenges in the oceanic migrations of leatherback sea turtles

The open-sea movements of marine animals are affected by the drifting action of currents that, if not compensated for, can produce non-negligible deviations from the correct route towards a given target. Marine turtles are paradigmatic skilful oceanic navigators that are able to reach remote goals at the end of long-distance migrations, apparently overcoming current drift effects. Particularly relevant is the case of leatherback turtles (Dermochelys coriacea), which spend entire years in the ocean, wandering in search of planktonic prey. Recent analyses have revealed how the movements of satellite-tracked leatherbacks in the Indian, Atlantic and Pacific Oceans are strongly dependent on the oceanic currents, up to the point that turtles are often passively transported over long distances. However, leatherbacks are known to return to specific areas to breed every 2–3 years, thus finding their way back home after long periods in the oceanic environment. Here we examine the navigational consequences of the leatherbacks'' close association with currents and discuss how the combined reliance on mechanisms of map-based navigation and local orientation cues close to the target may allow leatherbacks to accomplish the difficult task of returning to specific sites after years spent wandering in a moving medium.     

Hippocampal participation in the sun compass orientation of phase-shifted homing pigeons

The orientation of phase-shifted control and hippocampal lesioned homing pigeons with previous homing experience was examined to investigate the possible participation of the hippocampal formation in sun compass orientation. Hippocampal lesioned pigeons displayed appropriate shifts in orientation indicating that such birds possess a functional sun compass that is used for orientation. However, their shift in orientation was consistently larger than in control pigeons revealing a difference in orientation never observed in pigeons that have not undergone a phase shift. Although alternative interpretations exist, the data suggest the intriguing possibility that following a change in the light-dark cycle, the hippocampal formation participates in the re-entrainment of a circadian rhythm that regulates sun compass orientation.     

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.     

Navigation by green turtles: which strategy do displaced adults use to find Ascension Island?

Sea turtles are known to perform long-distance, oceanic migrations between disparate feeding areas and breeding sites, some of them located on isolated oceanic islands. These migrations demonstrate impressive navigational abilities, but the sensory mechanisms used are still largely unknown. Green turtles breeding at Ascension Island perform long oceanic migrations (>2200 km) between foraging areas along the Brazilian coast and the isolated island. By performing displacement experiments of female green turtles tracked by satellite telemetry in the waters around Ascension Island we investigated which strategies most probably are used by the turtles in locating the island. In the present paper we analysed the search trajectories in relation to alternative navigation strategies including the use of global geomagnetic cues, ocean currents, celestial cues and wind. The results suggest that the turtles did not use chemical information transported with ocean currents. Neither did the results indicate that the turtles use true bi-coordinate geomagnetic navigation nor did they use indirect navigation with respect to any of the available magnetic gradients (total field intensity, horizontal field intensity, vertical field intensity, inclination and declination) or celestial cues. The female green turtles successfully locating Ascension Island seemed to use a combination of searching followed by beaconing, since they searched for sensory contact with the island until they reached positions NW and N of the Island and from there presumably used cues transported by wind to locate the island during the final stages of the search.     

Sea turtles: navigating with magnetism

Young sea turtles use the Earth's magnetic field as a source of navigational information during their epic transoceanic migrations and while homing. A new study using satellite telemetry has now demonstrated for the first time that adult turtles also navigate using the Earth's magnetic field.     

Sea turtles, lobsters, and oceanic magnetic maps

Numerous marine animals can sense the Earth's magnetic field and use it as a cue in orientation and navigation. Two distinct types of information can potentially be extracted from the Earth's field. Directional or compass information enables animals to maintain a consistent heading in a particular direction such as north or south. In contrast, positional or map information can be used by animals to assess geographic location and, in some cases, to navigate to specific target areas. Marine animals exploit magnetic positional information in at least two different ways. For hatchling loggerhead sea turtles, regional magnetic fields function as open-sea navigational markers, eliciting changes in swimming direction at crucial points in the migratory route. Older sea turtles, as well as spiny lobsters, use magnetic information in a more complex way, exploiting it as a component of a classical navigational map, which permits an assessment of position relative to specific geographic destinations. These “magnetic maps” have not yet been fully characterized. They may be organized in several fundamentally different ways, some of which bear little resemblance to human maps, and they may also be used in conjunction with unconventional navigational strategies. Unraveling the nature of magnetic maps and exploring how they are used represents one of the most exciting frontiers of behavioral and sensory biology.     

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.     

Pineal Body and Magnetic Sensitivity: Homing in Pinealectomized Pigeons under Overcast Skies

Since birds use the earth's magnetic field for compass orientation when astronomical cues are lacking and it has recently been suggested that the pineal body is part of their magnetic compass, test releases have been performed in overcast conditions with pigeons deprived of the pineal body. On the whole, both experimental and control birds were capable of homeward orientation, though the bearings of experimental were rather more scattered. No differences in homing speed or success were recorded. Thus, the pineal body does not appear to play an important role in the homing of pigeons.