Birds have remained the dominant model for studying the mechanisms of animal navigation for decades, with much of what has been discovered coming from laboratory studies or model systems. The miniaturisation of tracking technology in recent years now promises opportunities for studying navigation during migration itself (migratory navigation) on an unprecedented scale. Even if migration tracking studies are principally being designed for other purposes, we argue that attention to salient environmental variables during the design or analysis of a study may enable a host of navigational questions to be addressed, greatly enriching the field. We explore candidate variables in the form of a series of contrasts (e.g. land vs ocean or night vs day migration), which may vary naturally between migratory species, populations or even within the life span of a migrating individual. We discuss how these contrasts might help address questions of sensory mechanisms, spatiotemporal representational strategies and adaptive variation in navigational ability. We suggest that this comparative approach may help enrich our knowledge about the natural history of migratory navigation in birds.
Although questions such as ‘How do animals find their way, and how do they sense and process this information in the brain?’ have been asked for centuries, the field of animal orientation and navigation has seen an immense leap forward in the past few decades. Moreover, our understanding has also expanded considerably regarding the molecular and physiological mechanisms of the different compasses and cues used by animals for orientation and navigation (Akesson et al., Chapter 9, and Svensson et al., Chapter 11). Most notable are the advances made in our understanding of how animals can sense information provided by the geomagnetic field and use this information for behavioural tasks, for example for compass orientation during migration. But despite interdisciplinary and highly integrative research over recent decades, we do not fully understand how animals perceive the Earth s magnetic field. We know that animals use geomagnetic information for orientation tasks (see Akesson et al., Chapter 9), but the receptor(s) remain to be identified. In this chapter, we review current knowledge in this area, outline challenges, and suggest future approaches to elucidate the sensory modalities used by animals for orientation and navigational tasks.
Migratory birds use multiple compass systems for orientation, including a magnetic, star and sun/polarized light compass. To keep these compasses in register, birds have to regularly update them with respect to a common reference. However, cue-conflict studies have revealed contradictory results on the compass hierarchy, favoring either celestial or magnetic compass cues as the primary calibration reference. Both the geomagnetic field and polarized light cues present at sunrise and sunset have been shown to play a role in compass cue integration, and evidence suggests that polarized light cues at sunrise and sunset may provide the primary calibration reference for the other compass systems. We tested whether migratory garden warblers recalibrated their compasses when they were exposed to the natural celestial cues at sunset in a shifted magnetic field, which are conditions that have been shown to be necessary for the use of a compass reference based on polarized light cues. We released the birds on the same evening under a starry sky and followed them by radio tracking. We found no evidence of compass recalibration, even though the birds had a full view of polarized light cues near the horizon at sunset during the cue-conflict exposure. Based on a meta-analysis of the available literature, we propose an extended unifying theory on compass cue hierarchy used by migratory birds to calibrate the different compasses. According to this scheme, birds recalibrate their magnetic compass by sunrise/sunset polarized light cues, provided they have access to the vertically aligned band of maximum polarization near the horizon and a view of landmarks. Once the stars appear in the sky, the birds then recalibrate the star compass with respect of the recalibrated magnetic compass. If sunrise and sunset information can be viewed from the same location, the birds average the information to get a true geographic reference. If polarized light information is not available near the horizon at sunrise or sunset, the birds temporarily transfer the previously calibrated magnetic compass information to the available celestial compasses. We conclude that the type of cue-conflict manipulation and the availability of stars can explain the discrepancies between studies.
SUMMARY Magnetic compass orientation in birds has been shown to be light dependent. Results from behavioural studies indicate that magnetoreception capabilities are disrupted under light of peak wavelengths longer than 565 nm, and shifts in orientation have been observed at higher light intensities(43-44×1015 quanta s-1 m-2). To investigate further the function of the avian magnetic compass with respect to wavelength and intensity of light, we carried out orientation cage experiments with juvenile European robins, caught during their first autumn migration,exposed to light of 560.5 nm (green), 567.5 nm (green-yellow) and 617 nm (red)wavelengths at three different intensities (1 mW m-2, 5 mW m-2 and 10 mW m-2). We used monochromatic light of a narrow wavelength range (half bandwidth of 9-11 nm, compared with half bandwidths ranging between 30 nm and 70 nm used in other studies) and were thereby able to examine the magnetoreception mechanism in the expected transition zone between oriented and disoriented behaviour around 565 nm in more detail. We show (1) that European robins show seasonally appropriate migratory directions under 560.5 nm light, (2) that they are completely disoriented under 567.5 nm light under a broad range of intensities, (3) that they are able to orient under 617 nm light of lower intensities, although into a direction shifted relative to the expected migratory one, and (4) that magnetoreception is intensity dependent, leading to disorientation under higher intensities. Our results support the hypothesis that birds possess a light-dependent magnetoreception system based on magnetically sensitive,antagonistically interacting spectral mechanisms, with at least one high-sensitive short-wavelength mechanism and one low-sensitive long-wavelength mechanism.
Summary We provide evidence for spontaneous quadramodal magnetic orientation in a larval insect. Second instar Berlin, Canton-S, and Oregon-R X Canton-S strains of Drosophila melanogaster exhibited quadramodal orientation with clusters of bearings along the four anti-cardinal compass directions (i.e. 45°, 135°, 225°, 315°). In double-blind experiments, Canton-S Drosophila larvae exhibited quadramodal orientation in the presence of an earth-strength magnetic field, while this response was abolished when the horizontal component of the magnetic field was cancelled, indicating that the quadramodal behavior is dependent on magnetic cues, and may reflect properties of the underlying magnetoreception mechanism. In addition, a reanalysis of data from studies of learned magnetic compass orientation by adult Drosophila melanogaster and C57BL/6 mice reveals patterns of response similar to those exhibited by larval flies suggesting that a common magnetoreception mechanism(s) may underlie these behaviors. Therefore, characterizing the mechanism(s) of magnetoreception in flies may hold the key to understanding the magnetic sense in a wide array of terrestrial organisms.
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