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Migrating animals perform astonishing seasonal movements by orienting and navigating over thousands of kilometres with great precision. Many migratory species use cues from the sun, stars, landmarks, olfaction and the Earth's magnetic field for this task. Among vertebrates, songbirds are the most studied taxon in magnetic-cue-related research. Despite multiple studies, we still lack a clear understanding of when, where and how magnetic cues affect the decision-making process of birds and hence, their realised migratory behaviour in the wild. This understanding is especially important to interpret the results of laboratory experiments in an ecologically appropriate way. In this review, we summarise the current findings about the role of magnetic cues for migratory decisions in songbirds. First, we review the methodological principles for orientation and navigation research, specifically by comparing experiments on caged birds with experiments on free-flying birds. While cage experiments can show the sensory abilities of birds, studies with free-flying birds can characterise the ecological roles of magnetic cues. Second, we review the migratory stages, from stopover to endurance flight, in which songbirds use magnetic cues for their migratory decisions and incorporate this into a novel conceptual framework. While we lack studies examining whether and when magnetic cues affect orientation or navigation decisions during flight, the role of magnetic cues during stopover is relatively well studied, but mostly in the laboratory. Notably, many such studies have produced contradictory results so that understanding the biological importance of magnetic cues for decisions in free-flying songbirds is not straightforward. One potential explanation is that reproducibility of magnetic-cue experiments is low, probably because variability in the behavioural responses of birds among experiments is high. We are convinced that parts of this variability can be explained by species-specific and context-dependent reactions of birds to the study conditions and by the bird's high flexibility in whether they include magnetic cues in a decision or not. Ultimately, this review should help researchers in the challenging field of magnetoreception to design experiments meticulously and interpret results of such studies carefully by considering the migration ecology of their focal species.

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Currently, it is generally assumed that migratory birds are oriented in the appropriate migratory direction under UV, blue and green light (short-wavelength) and are unable to use their magnetic compass in total darkness and under yellow and red light (long-wavelength). However, it has also been suggested that the magnetic compass has two sensitivity peaks: in the short and long wavelengths, but with different intensities. In this project, we aimed to study the orientation of long-distance migrants, pied flycatchers (Ficedula hypoleuca), under different narrowband light conditions during autumn and spring migrations. The birds were tested in the natural magnetic field (NMF) and a changed magnetic field (CMF) rotated counterclockwise by 120° under dim green (autumn) and yellow (spring and autumn) light, which are on the 'threshold' between the short-wavelength and long-wavelength light. We showed that pied flycatchers (i) were completely disoriented under green light both in the NMF and CMF but (ii) showed the migratory direction in the NMF and the appropriate response to CMF under yellow light. Our data contradict the results of previous experiments under narrowband green and yellow light and raise doubts about the existence of only short-wavelength magnetoreception. The parameters of natural light change dramatically in spectral composition and intensity after local sunset, and the avian magnetic compass should be adapted to function properly under such constantly changing light conditions.

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BACKGROUND: For many migratory species, inexperienced (naïve) individuals reach remote non-breeding areas independently using one or more inherited compass headings and, potentially, magnetic signposts to gauge where to switch between compass headings. Inherited magnetic-based migration has not yet been assessed as a population-level process, particularly across strong geomagnetic gradients or where long-term geomagnetic shifts (hereafter, secular variation) could create mismatches with magnetic headings. Therefore, it remains unclear whether inherited magnetic headings and signposts could potentially adapt to secular variation under natural selection. METHODS: To address these unknowns, we modelled migratory orientation programs using an evolutionary algorithm incorporating global geomagnetic data (1900-2023). Modelled population mixing incorporated both natal dispersal and trans-generational inheritance of magnetic headings and signposts, including intrinsic (stochastic) variability in inheritance. Using the model, we assessed robustness of trans-hemispheric migration of a migratory songbird whose Nearctic breeding grounds have undergone rapid secular variation (mean 34° clockwise drift in declination, 1900-2023), and which travels across strong geomagnetic gradients via Europe to Africa. RESULTS: Model-evolved magnetic-signposted migration was overall successful throughout the 124-year period, with 60-90% mean successful arrival across a broad range in plausible precision in compass headings and gauging signposts. Signposted migration reduced trans-Atlantic flight distances and was up to twice as successful compared with non-signposted migration. Magnetic headings shifted plastically in response to the secular variation (mean 16°-17° among orientation programs), whereas signpost latitudes were more constrained (3°-5° mean shifts). This plasticity required intrinsic variability in inheritance (model-evolved σ ≈ 2.6° standard error), preventing clockwise secular drift from causing unsustainable open-ocean flights. CONCLUSIONS: Our study supports the potential long-term viability of inherited magnetic migratory headings and signposts, and illustrates more generally how inherited migratory orientation programs can both mediate and constrain evolution of routes, in response to global environmental change.

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The avian magnetic compass allows orientation during migration and is shown to function properly under short-wavelength but not long-wavelength visible light. Therefore, the magnetoreceptive system is assumed to be light- and wavelength-dependent and localized in the retina of the eye. Putative candidates for the role of primary magnetosensory molecules are the cryptochromes that are known to be expressed in the avian retina and must be able to interact with phototransduction proteins. Previously, we reported that in migratory birds change in magnetic field direction induces significant effects on electroretinogram amplitude in response to blue flashes, and such an effect was observed only in the nasal quadrant of the retina. Here, we report new electroretinographic, microscopic and microspectrophotometric data on European robins, confirming the magnetosensitivity of the retinal nasal quadrant after applying the background illumination. We hypothesized that magnetoreceptive distinction of this region may be related to its morphology and analyzed the retinal distribution and optical properties of oil droplets, the filtering structures within cones. We found that the nasal quadrant contains double cones with the most intensely colorized oil droplets compared to the rest of the retina, which may be related to its magnetosensory function.

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Sensitivity to the earth's magnetic field is the least understood of the major sensory systems, despite being virtually ubiquitous in animals and of widespread interest to investigators in a wide range of fields from behavioral ecology to quantum physics. Although research on the use of magnetic cues by migratory birds, fish, and sea turtles is more widely known, much of our current understanding of the functional properties of vertebrate magnetoreception has come from research on amphibians. Studies of amphibians established the presence of a light-dependent magnetic compass, a second non-light-dependent mechanism involving particles of magnetite and/or maghemite, and an interaction between these two magnetoreception mechanisms that underlies the "map" component of homing. Simulated magnetic displacement experiments demonstrated the use of a high-resolution magnetic map for short-range homing to breeding ponds requiring a sampling strategy to detect weak spatial gradients in the magnetic field despite daily temporal variation at least an order of magnitude greater. Overall, reliance on a magnetic map for short-range homing places greater demands on the underlying sensory detection, processing, and memory mechanisms than comparable mechanisms used by long-distance migrants. Moreover, unlike sea turtles and migratory birds, amphibians are exceptionally well suited to serve as model organisms in which to characterize the molecular and biophysical mechanisms underlying the light-dependent 'quantum compass'.

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Wood ants were trained indoors to follow a magnetically specified route that went from the centre of an arena to a drop of sucrose at the edge. The arena, placed in a white cylinder, was in the centre of a 3D coil system generating an inclined Earth-strength magnetic field in any horizontal direction. The specified direction was rotated between each trial. The ants' knowledge of the route was tested in trials without food. Tests given early in the day, before any training, show that ants remember the magnetic route direction overnight. During the first 2 s of a test, ants mostly faced in the specified direction, but thereafter were often misdirected, with a tendency to face briefly in the opposite direction. Uncertainty about the correct path to take may stem in part from competing directional cues linked to the room. In addition to facing along the route, there is evidence that ants develop magnetically directed home and food vectors dependent upon path integration. A second experiment asked whether ants can use magnetic information contextually. In contrast to honeybees given a similar task, ants failed this test. Overall, we conclude that magnetic directional cues can be sufficient for route learning.

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Migratory birds use different global cues including celestial and magnetic information to determine and maintain their seasonally appropriate migratory direction. A hierarchy among different compass systems in songbird migrants is still a matter for discussion owing to highly variable and apparently contradictory results obtained in various experimental studies. How birds decide whether and how they should calibrate their compasses before departure remains unclear. A recent 'extended unified theory' suggested that access to both a view of the sky near the horizon and stars during the cue-conflict exposure might be crucial for the results of cue-conflict experiments. In this study, we performed cue-conflict experiments in three European songbird species with different migratory strategies (garden warbler, Sylvia borin; pied flycatcher, Ficedula hypoleuca; and European robin, Erithacus rubecula; juveniles and adults; spring and autumn migrations) using a uniform experimental protocol. We exposed birds to the natural celestial cues in a shifted (120 deg clockwise/counterclockwise) magnetic field from sunset to the end of the nautical twilight and tested them in orientation cages immediately after cue-conflict treatments. None of the species (apart from adult robins) showed any sign of calibration even if they had access to a view of the sky and local surroundings near the horizon and stars during cue-conflict treatments. Based on results of our experiments and data from previous contradictory studies, we suggest that no uniform theory can explain why birds calibrate or do not calibrate their compass systems. Each species (and possibly even different populations) may choose its calibration strategy differently.

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A magnetic compass sense has been demonstrated in all major classes of vertebrates, as well as in many invertebrates. In mammals, controlled laboratory studies of mice have provided evidence for a robust magnetic compass that is comparable to, or exceeds, the performance of that in other animals. Nevertheless, the vast majority of laboratory studies of spatial behavior and cognition in murine rodents have failed to produce evidence of sensitivity to magnetic cues. Given the central role that a magnetic compass sense plays in the spatial ecology and cognition of non-mammalian vertebrates, and the potential utility that a global/universal reference frame derived from the magnetic field would have in mammals, the question of why responses to magnetic cues have been so difficult to demonstrate reliably is of considerable importance. In this paper, we review evidence that the magnetic compass of murine rodents shares a number of properties with light-dependent compasses in a wide variety of other animals generally believed to be mediated by a radical pair mechanism (RPM) or related quantum process. Consistent with the RPM, we summarize both published and previously unpublished findings suggesting that the murine rodent compass is sensitive to low-level radio frequency (RF) fields. Finally, we argue that the presence of anthropogenic RF fields in laboratory settings, may be an important source of variability in responses of murine rodents to magnetic cues.

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Spontaneous magnetic alignment is the simplest known directional response to the geomagnetic field that animals perform. Magnetic alignment is not a goal directed response and its relevance in the context of orientation and navigation has received little attention. Migratory songbirds, long-standing model organisms for studying magnetosensation, have recently been reported to align their body with the geomagnetic field. To explore whether the magnetic alignment behaviour in songbirds is involved in the underlying mechanism for compass calibration, which have been suggested to occur near to sunset, we studied juvenile Eurasian reed warblers (Acrocephalus scirpaceus) captured at stopover during their first autumn migration. We kept one group of birds in local daylight conditions and an experimental group under a 2 h delayed sunset. We used an ad hoc machine learning algorithm to track the birds' body alignment over a 2-week period. Our results show that magnetic body alignment occurs prior to sunset, but shifts to a more northeast-southwest alignment afterwards. Our findings support the hypothesis that body alignment could be associated with how directional celestial and magnetic cues are integrated in the compass of migratory birds.

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The magnetic field of the Earth provides animals with various kinds of information. Its use as a compass was discovered in the mid-1960s in birds, when it was first met with considerable skepticism, because it initially proved difficult to obtain evidence for magnetic sensitivity by conditioning experiments. Meanwhile, a magnetic compass was found to be widespread. It has now been demonstrated in members of all vertebrate classes, in mollusks and several arthropod species, in crustaceans as well as in insects. The use of the geomagnetic field as a 'map' for determining position, although already considered in the nineteenth century, was demonstrated by magnetically simulating displacements only after 2000, namely when animals, tested in the magnetic field of a distant site, responded as if they were physically displaced to that site and compensated for the displacement. Another use of the magnetic field is that as a 'sign post' or trigger: specific magnetic conditions elicit spontaneous responses that are helpful when animals reach the regions where these magnetic characteristics occur. Altogether, the geomagnetic field is a widely used valuable source of navigational information for mobile animals.

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At the beginning of their foraging careers, Cataglyphis desert ants calibrate their compass systems and learn the visual panorama surrounding the nest entrance. For that, they perform well-structured initial learning walks. During rotational body movements (pirouettes), naïve ants (novices) gaze back to the nest entrance to memorize their way back to the nest. To align their gaze directions, they rely on the geomagnetic field as a compass cue. In contrast, experienced ants (foragers) use celestial compass cues for path integration during food search. If the panorama at the nest entrance is changed, foragers perform re-learning walks prior to heading out on new foraging excursions. Here, we show that initial learning walks and re-learning walks are structurally different. During re-learning walks, foragers circle around the nest entrance before leaving the nest area to search for food. During pirouettes, they do not gaze back to the nest entrance. In addition, foragers do not use the magnetic field as a compass cue to align their gaze directions during re-learning walk pirouettes. Nevertheless, magnetic alterations during re-learning walks under manipulated panoramic conditions induce changes in nest-directed views indicating that foragers are still magnetosensitive in a cue conflict situation.

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Cryptochromes are photolyase-related blue-light receptors acting as core components of the mammalian circadian clock in the cell nuclei. One or more members of the cryptochrome protein family are also assumed to play a role in avian magnetoreception, but the primary sensory molecule in the retina of migratory birds that mediates light-dependent magnetic compass orientation has still not been identified. The mRNA of cryptochrome 2 (Cry2) has been reported to be located in the cell nuclei of the retina, but Cry2 localisation has not yet been demonstrated at the protein level. Here, we provide evidence that Cry2 protein is located in the photoreceptor inner segments, the outer nuclear layer, the inner nuclear layer and the ganglion cell layer in the retina of night-migratory European robins, homing pigeons and domestic chickens. At the subcellular level, we find Cry2 both in the cytoplasm and the nucleus of cells residing in these layers. This broad nucleic expression rather points to a role for avian Cry2 in the circadian clock and is consistent with a function as a transcription factor, analogous to mammalian Cry2, and speaks against an involvement in magnetoreception.

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Red-headed Buntings (Emberiza bruniceps) perform long-distance migrations within their southerly overwintering grounds and breeding areas in the northern hemisphere. Long-distance migration demands essential orientation mechanisms. The earth's magnetic field, celestial cues, and memorization of geographical cues en route provide birds with compass knowledge during migration. Birds were tested during spring migration for orientation under natural clear skies, simulated overcast skies at natural day length and temperature, simulated overcast at 22 °C and 38 °C temperatures, and in the deflected (-120°) magnetic field. Under clear skies, the Red-headed Buntings were oriented NNW (north-northwest); simulated overcast testing resulted in a northerly mean direction at local temperatures as well as at 22 °C and 38 °C. The Buntings reacted strongly in favor of the rotated magnetic field under the simulated overcast sky, demonstrating the use of a magnetic compass for migrating in a specific direction.

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We investigated the migratory orientation of early and late captured dunlins, Calidris alpina, by recording their migratory activity in circular orientation cages during autumn at a staging site in southwest Alaska and performed route simulations to the wintering areas. Two races of dunlins breeding in Alaska have different wintering grounds in North America (Pacific Northwest), and East Asia. Dunlins caught early in autumn (presumably Calidris alpinapacifica) oriented towards their wintering areas (east-southeast; ESE) supporting the idea that they migrate nonstop over the Gulf of Alaska to the Pacific Northwest. We found no difference in orientation between adult and juveniles, nor between fat and lean birds or under clear and overcast skies demonstrating that age, energetic status and cloud cover did not affect the dunlins' migratory orientation. Later in autumn, we recorded orientation responses towards south-southwest suggesting arrival of the northern subspecies Calidris alpinaarcticola at our site. Route simulations revealed multiple compass mechanisms were compatible with the initial direction of early dunlins wintering in the Pacific Northwest, and for late dunlins migrating to East Asia. Future high-resolution tracking would reveal routes, stopover use including local movements and possible course shifts during migration from Alaska to wintering sites on both sides of the north Pacific Ocean.

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Species throughout the animal kingdom use the Earth's magnetic field (MF) to navigate using either or both of two mechanisms. The first relies on magnetite crystals in tissue where their magnetic moments align with the MF to transduce a signal transmitted to the central nervous system. The second and the subject of this paper involves cryptochrome (CRY) proteins located in cone photoreceptors distributed across the retina, studied most extensively in birds. According to the "Radical Pair Mechanism" (RPM), blue/UV light excites CRY's flavin cofactor (FAD) to generate radical pairs whose singlet-to-triplet interconversion rate is modulated by an external MF. The signaling product of the RPM produces an impression of the field across the retinal surface. In birds, the resulting signal on the optic nerve is transmitted along the thalamofugal pathway to the primary visual cortex, which projects to brain regions concerned with image processing, memory, and executive function. The net result is a bird's orientation to the MF's inclination: its vector angle relative to the Earth's surface. The quality of ambient light (e.g., polarization) provides additional input to the compass. In birds, the Type IV CRY isoform appears pivotal to the compass, given its positioning within retinal cones; a cytosolic location therein indicating no role in the circadian clock; relatively steady diurnal levels (unlike Type II CRY's cycling); and a full complement of FAD (essential for photosensitivity). The evidence indicates that mammalian Type II CRY isoforms play a light-independent role in the cellular molecular clock without a photoreceptive function.

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The star compass of birds, like the sun compass, is not innate. To possess either of them, birds have to observe the rotating sky and determine its centre of rotation (in the case of the star compass) or the sun's movement (for the sun compass). Young birds are believed to learn how to use the star compass before their first migration, even though the evidence of this is lacking. Here, we tested whether hand-raised Pied flycatchers (Ficedula hypoleuca) that had not established the star compass prior to their first autumn migration can gain it later in their ontogeny, in spring. We also attempted to examine whether the observation of diurnal celestial cues (the sun and polarized light) prior to autumn migration would affect the process of star compass learning in spring. When tested in the vertical magnetic field under the natural starry sky, the group of birds that observed the stars in spring as the first celestial cues were able to choose the migratory direction. In contrast, the birds that had never seen the stars were not able to use the nightly celestial cues in the vertical magnetic field. However, birds that had seen the daytime celestial cues till autumn and the stars at spring were disoriented, although this might be due to the small sample size. Our data suggest the possibility that the star compass may be learned in spring and emphasize the necessity for further research into the interaction of celestial compasses.

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The Johnston's organ (JO) in the insect antenna is a multisensory organ involved in several navigational tasks including wind-compass orientation, flight control, graviception, and, possibly, magnetoreception. Here we investigate the three dimensional anatomy of the JO and its neuronal projections into the brain of the desert ant Cataglyphis, a marvelous long-distance navigator. The JO of C. nodus workers consists of 40 scolopidia comprising three sensory neurons each. The numbers of scolopidia slightly vary between different sexes (female/male) and castes (worker/queen). Individual scolopidia attach to the intersegmental membrane between pedicel and flagellum of the antenna and line up in a ring-like organization. Three JO nerves project along the two antennal nerve branches into the brain. Anterograde double staining of the antennal afferents revealed that JO receptor neurons project to several distinct neuropils in the central brain. The T5 tract projects into the antennal mechanosensory and motor center (AMMC), while the T6 tract bypasses the AMMC via the saddle and forms collaterals terminating in the posterior slope (PS) (T6I), the ventral complex (T6II), and the ventrolateral protocerebrum (T6III). Double labeling of JO and ocellar afferents revealed that input from the JO and visual information from the ocelli converge in tight apposition in the PS. The general JO anatomy and its central projection patterns resemble situations in honeybees and Drosophila. The multisensory nature of the JO together with its projections to multisensory neuropils in the ant brain likely serves synchronization and calibration of different sensory modalities during the ontogeny of navigation in Cataglyphis.

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Migratory birds are known to be sensitive to external magnetic field (MF). Much indirect evidence suggests that the avian magnetic compass is localized in the retina. Previously, we showed that changes in the MF direction could modulate retinal responses in pigeons. In the present study, we performed similar experiments using the traditional model animal to study the magnetic compass, European robins. The photoresponses of isolated retina were recorded using ex vivo electroretinography (ERG). Blue- and red-light stimuli were applied under an MF with the natural intensity and two MF directions, when the angle between the plane of the retina and the field lines was 0° and 90°, respectively. The results were separately analysed for four quadrants of the retina. A comparison of the amplitudes of the a- and b-waves of the ERG responses to blue stimuli under the two MF directions revealed a small but significant difference in a- but not b-waves, and in only one (nasal) quadrant of the retina. The amplitudes of both the a- and b-waves of the ERG responses to red stimuli did not show significant effects of the MF direction. Thus, changes in the external MF modulate the European robin retinal responses to blue flashes, but not to red flashes. This result is in a good agreement with behavioural data showing the successful orientation of birds in an MF under blue, but not under red illumination.

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The use of information provided by the geomagnetic field (GMF) for navigation is widespread across the animal kingdom. At the same time, the magnetic sense is one of the least understood senses. Here, we review evidence for magnetoreception in Hymenoptera. We focus on experiments aiming to shed light on the role of the GMF for navigation. Both honeybees and desert ants are well-studied experimental models for navigation, and both use the GMF for specific navigational tasks under certain conditions. Cataglyphis desert ants use the GMF as a compass cue for path integration during their initial learning walks to align their gaze directions towards the nest entrance. This represents the first example for the use of the GMF in an insect species for a genuine navigational task under natural conditions and with all other navigational cues available. We argue that the recently described magnetic compass in Cataglyphis opens up a new integrative approach to understand the mechanisms underlying magnetoreception in Hymenoptera on different biological levels.

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Multi-sensor imaging systems using the global navigation satellite system (GNSS) and digital magnetic compass (DMC) for geo-referencing have an important role and wide application in long-range surveillance systems. To achieve the required system heading accuracy, the specific magnetic compass calibration and compensation procedures, which highly depend on the application conditions, should be applied. The DMC compensation technique suitable for the operation environment is described and different technical solutions are studied. The application of the swinging procedure was shown as a good solution for DMC compensation in a given application. The selected DMC was built into a system to be experimentally evaluated, both under laboratory and field conditions. The implementation of the compensation procedure and magnetic sensor integration in systems is described. The heading accuracy measurement results show that DMC could be successfully integrated and used in long-range surveillance systems providing required geo-referencing data.