Magnetic compass orientation of European robins under 565 nm green light

European robins tested under monochromatic green light with a peak wavelength of 565 nm at an intensity of 2.1 mW m^-2 in the local geomagnetic field preferred their migratory direction, heading southward in autumn and northward in spring. Inverting of the vertical component of the magnetic field caused the robins to reverse their headings, indicating that the birds used a magnetic inclination compass to locate their migratory direction. The behavior recorded under green light at an intensity of 2.1 mW m^-2 is thus not different from that previously recorded under "white" light; it represents normal migratory orientation.     

Avian magnetic compass: fast adjustment to intensities outside the normal functional window

To determine how fast birds can adapt to magnetic intensities outside the normal functional window of their magnetic compass, we tested migratory birds in a magnetic field of 92,000 nT, twice the intensity of the local geomagnetic field at the test site in Frankfurt a.M., Germany. In the local field, robins showed a significant preference of their southerly migratory direction, whereas in the 92,000-nT field, they were initially disoriented. However, when the birds were preexposed to 92,000 nT for 1 h before being tested, they were able to orient under this intensity, and their behavior did not differ from that in the geomagnetic field. These data show that birds require only a short time to adjust to magnetic intensities, which they cannot spontaneously use for orientation. Interpreting these findings in view of the radical pair model (Ritz et al. 2000), this means that they can learn rather quickly to interpret novel activation patterns on their retina.     

Magnetoreception in birds: no intensity window in “fixed direction” responses

Under 502 nm turquoise light combined with 590 nm yellow light and in total darkness, European robins, Erithacus rubecula, no longer prefer their migratory direction, but exhibit so-called fixed direction responses that do not show the seasonal change between spring and autumn. We tested robins under these light conditions in the local geomagnetic field of 46 μT, a field of twice this intensity, 92 μT, and a field of three times this intensity, 138 μT. Under all three magnetic conditions, the birds preferred the same easterly direction under turquoise-and-yellow light and the same northwesterly direction under dark, while they were oriented in their seasonally appropriate direction under control conditions. “Fixed direction” responses are thus not limited to a narrow intensity window as has been found for normal compass orientation. This can be attributed to their origin in the magnetite-based receptor in the upper beak, which operates according to fundamentally different principles than the radical pair mechanism in the retina mediating compass orientation. “Fixed direction” responses are possibly a relict of a receptor mechanism that changed its function, now mainly providing information on magnetic intensity.     

Light-dependent magnetoreception in birds: interaction of at least two different receptors

Passerine migrants require light from the blue-green part of the spectrum for magnetic compass orientation; under yellow light, they are disoriented. European robins tested under a combination of yellow light and blue or green light showed a change in behavior, no longer preferring their seasonally appropriate migratory direction: in spring as well as in autumn, they preferred southerly headings under blue-and-yellow and northerly headings under green-and-yellow light. This clearly shows that yellow light is not neutral and suggests the involvement of at least two types of receptors in obtaining magnetic compass information, with the specific interaction of these receptors being rather complex.     

Magnetic compass orientation of migratory birds in the presence of a 1.315 MHz oscillating field

The radical pair model of magnetoreception predicts that magnetic compass orientation can be disrupted by high frequency magnetic fields in the Megahertz range. European robins, Erithacus rubecula, were tested under monochromatic 565 nm green light in 1.315 MHz fields of 0.48 T during spring and autumn migration, with 1.315 MHz being the frequency that matches the energetic splitting induced by the local geomagnetic field. The birds responses depended on the alignment of the oscillating field with respect to the static geomagnetic field: when the 1.315 MHz field was aligned parallel with the field lines, birds significantly preferred northerly directions in spring and southerly directions in autumn. These preferences reflect normal migratory orientation, with the variance slightly increased compared to control tests in the geomagnetic field alone or to tests in a 7.0 MHz field. However, in the 1.315 MHz field aligned at a 24° angle to the field lines, the birds were disoriented in both seasons, indicating that the high frequency field interfered with magnetoreception. These finding are in agreement with theoretical predictions and support the assumption of a radical-pair mechanism underlying the processes mediating magnetic compass information in birds.     

A time-compressed simulated geomagnetic storm influences the nest-exiting flight angles of the stingless bee Tetragonisca angustula

Insects have been used as models for understanding animal orientation. It is well accepted that social insects such as honeybees and ants use different natural cues in their orientation mechanism. A magnetic sensitivity was suggested for the stingless bee Schwarziana quadripunctata, based on the observation of a surprising effect of a geomagnetic storm on the nest-exiting flight angles. Stimulated by this result, in this paper, the effects of a time-compressed simulated geomagnetic storm (TC-SGS) on the nest-exiting flight angles of another stingless bee, Tetragonisca angustula, are presented. Under an applied SGS, either on the horizontal or vertical component of the geomagnetic field, both nest-exiting flight angles, dip and azimuth, are statistically different from those under geomagnetic conditions. The angular dependence of ferromagnetic resonance (FMR) spectra of whole stingless bees shows the presence of organized magnetic nanoparticles in their bodies, which indicates this material as a possible magnetic detector.     

The sea-finding behavior of hatchling olive ridley sea turtles, Lepidochelys olivacea, at the beach of San Miguel (Costa Rica)

Newly hatched olive ridley sea turtles (Lepidochelys olivacea) were tested for their directional preferences in a sand-filled circular arena in total darkness. Hatchlings that had crawled about 5 m on the beach, toward the sea preferred the southwesterly direction that would have brought them to the water line, whereas hatchlings that had been denied this experience headed eastward, a direction of unclear origin. These data suggest that a short crawl across the natural beach can set the direction in which the young turtles subsequently move. The crawling experience was sufficient to acquire the compass course that they later follow, probably with the help of a magnetic compass, not only in the water, but already while still on land.     

Canopy compass in nocturnal homing of the subsocial shield bug, <Emphasis Type="Italic">Parastrachia japonensis</Emphasis> (Heteroptera: Parastrachiidae)

In contrast to an open environment where a specific celestial cue is predominantly used, visual contrast of canopies against the sky through the gap, known as canopy cues, is known to play a major role for visually guided insect navigators in woodland habitats. In this paper, we investigated whether a subsocial shield bug, Parastrachia japonensis, could gauge direction using canopy cues on a moonless night. The results show that they could perform the round trip foraging behaviour even in an experimental arena with only an artificial round gap opened in the ceiling of the arena and adjust their homing direction for a new azimuth when the gap was rotated. Thus, P. japonensis can use slightly brighter canopy cues as a compass reference but not complex landmarks during nocturnal homing behaviour.     

Glass buildings on river banks as “polarized light traps” for mass-swarming polarotactic caddis flies

The caddis flies Hydropsyche pellucidula emerge at dusk from the river Danube and swarm around trees and bushes on the river bank. We document here that these aquatic insects can also be attracted en masse to the vertical glass surfaces of buildings on the river bank. The individuals lured to dark, vertical glass panes land, copulate, and remain on the glass for hours. Many of them are trapped by the partly open, tiltable windows. In laboratory choice experiments, we showed that ovipositing H. pellucidula are attracted to highly and horizontally polarized light stimulating their ventral eye region and, thus, have positive polarotaxis. In the field, we documented that highly polarizing vertical black glass surfaces are significantly more attractive to both female and male H. pellucidula than weakly polarizing white ones. Using video polarimetry, we measured the reflection-polarization characteristics of vertical glass surfaces of buildings where caddis flies swarmed. We propose that after its emergence from the river, H. pellucidula is attracted to buildings by their dark silhouettes and the glass-reflected, horizontally polarized light. After sunset, this attraction may be strengthened by positive phototaxis elicited by the buildings' lights. The novelty of this visual-ecological phenomenon is that the attraction of caddis flies to vertical glass surfaces has not been expected because vertical glass panes do not resemble the horizontal surface of waters from which these insects emerge and to which they must return to oviposit.     

Changes in earth’s dipole

The dipole moment of Earth’s magnetic field has decreased by nearly over the past 150 years and by about 30% over the past 2,000 years according to archeomagnetic measurements. Here, we explore the causes and the implications of this rapid change. Maps of the geomagnetic field on the core–mantle boundary derived from ground-based and satellite measurements reveal that most of the present episode of dipole moment decrease originates in the southern hemisphere. Weakening and equatorward advection of normal polarity magnetic field by the core flow, combined with proliferation and growth of regions where the magnetic polarity is reversed, are reducing the dipole moment on the core–mantle boundary. Growth of these reversed flux regions has occurred over the past century or longer and is associated with the expansion of the South Atlantic Anomaly, a low-intensity region in the geomagnetic field that presents a radiation hazard at satellite altitudes. We address the speculation that the present episode of dipole moment decrease is a precursor to the next geomagnetic polarity reversal. The paleomagnetic record contains a broad spectrum of dipole moment fluctuations with polarity reversals typically occurring during dipole moment lows. However, the dipole moment is stronger today than its long time average, indicating that polarity reversal is not likely unless the current episode of moment decrease continues for a thousand years or more.     

Spontaneous expression of magnetic compass orientation in an epigeic rodent: the bank vole,Clethrionomys glareolus

Magnetoreception has been convincingly demonstrated in only a few mammalian species. Among rodents, magnetic compass orientation has been documented in four species of subterranean mole rats and two epigeic (i.e. active above ground) species—the Siberian hamster and the C57BL/6J mouse. The mole rats use the magnetic field azimuth to determine compass heading; their directional preference is spontaneous and unimodal, and their magnetic compass is magnetite-mediated. By contrast, the primary component of orientation response is learned in the hamster and the mouse, but both species also exhibit a weak spontaneous bimodal preference in the natural magnetic field. To determine whether the magnetic compass of wild epigeic rodents features the same functional properties as that of laboratory rodents, we investigated magnetic compass orientation in the bank vole Clethrionomys glareolus (Cricetidae, Rodentia). The voles exhibited a robust spontaneous bimodal directional preference, i.e. built nests and slept preferentially along the north-south axis, and deflected their directional preference according to a shift in the direction of magnetic north, clearly indicating that they were deriving directional information from the magnetic field. Thus, bimodal, axially symmetrical directional choice seems to be a common feature shared by epigeic rodents. However, spontaneous directional preference in the bank vole appeared to be more pronounced than that reported in the hamster and the mouse. These findings suggest that bank voles are well suited for future studies investigating the adaptive significance and mechanisms of magnetic orientation in epigeic rodents.     

Light-dependent magnetic compass in Iberian green frog tadpoles

Here, we provide evidence for a wavelength-dependent effect of light on magnetic compass orientation in Pelophylax perezi (order Anura), similar to that observed in Rana catesbeiana (order Anura) and Notophthalmus viridescens (order Urodela), and confirm for the first time in an anuran amphibian that a 90° shift in the direction of magnetic compass orientation under long-wavelength light (≥500 nm) is due to a direct effect of light on the underlying magnetoreception mechanism. Although magnetic compass orientation in other animals (e.g., birds and some insects) has been shown to be influenced by the wavelength and/or intensity of light, these two amphibian orders are the only taxa for which there is direct evidence that the magnetic compass is light-dependent. The remarkable similarities in the light-dependent magnetic compasses of anurans and urodeles, which have evolved as separate clades for at least 250 million years, suggest that the light-dependent magnetoreception mechanism is likely to have evolved in the common ancestor of the Lissamphibia (Early Permian, ~294 million years) and, possibly, much earlier. Also, we discuss a number of similarities between the functional properties of the light-dependent magnetic compass in amphibians and blue light-dependent responses to magnetic stimuli in Drosophila melanogaster, which suggest that the wavelength-dependent 90° shift in amphibians may be due to light activation of different redox forms of a cryptochrome photopigment. Finally, we relate these findings to earlier studies showing that the pineal organ of newts is the site of the light-dependent magnetic compass and recent neurophysiological evidence showing magnetic field sensitivity in the frog frontal organ (an outgrowth of the pineal).     

Sexual differences in food preferences in the white stork: an experimental study

Sex differences in the foraging ecology of monomorphic species are poorly understood, due to problems with gender identification in field studies. In the current study, we used experimental conditions to investigate the food preferences of the white stork Ciconia ciconia, an opportunistic species in terms of food, but characterised by a low level of sexual dimorphism. During a 10-day experiment, 29 individuals (20 females and 9 males) were studied by means of a ‘cafeteria test’ in which the storks’ diet consisted of mammals, birds, fish, amphibians, insects and earthworms. The storks preferred food characterised by high calorific and protein values such as mammals, birds and fish. Sexes differed strongly in their preferences; males preferred mammals, whereas females preferred birds. Moreover, females consumed insects and earthworms less often than males. Interestingly, males spent significantly less time foraging than females. We have demonstrated that the white stork exhibits clear sexual differences in food preferences which are mostly attributable to differences in parental duties, physiology and anatomy.     

Vector navigation in desert ants, Cataglyphis fortis: celestial compass cues are essential for the proper use of distance information

Foraging desert ants navigate primarily by path integration. They continually update homing direction and distance by employing a celestial compass and an odometer. Here we address the question of whether information about travel distance is correctly used in the absence of directional information. By using linear channels that were partly covered to exclude celestial compass cues, we were able to test the distance component of the path-integration process while suppressing the directional information. Our results suggest that the path integrator cannot process the distance information accumulated by the odometer while ants are deprived of celestial compass information. Hence, during path integration directional cues are a prerequisite for the proper use of travel-distance information by ants.     

Light-dependent magnetoreception in birds: the effect of intensity of 565-nm green light

 In a previous study, Australian silvereyes tested in autumn under monochromatic 565-nm green light at intensities of 2.1 and 7.5 mW m^–2 preferred their normal northerly migratory direction, whereas they showed a significantly different tendency towards northwest at 15.0 mW m^–2. Repeating these experiments in spring with silvereyes migrating southward, we again observed well-oriented tendencies in the migratory direction at 2.1 and 7.5 mW m^–2. At 15.0 mW m^–2, however, the birds once more preferred northwesterly directions, i.e. their response under this condition proved to be independent of the migratory direction. This contradicts the interpretation that monochromatic green light of this high intensity leads to a rotation of compass information; instead, it appears to produce sensory input that causes birds to give up their migratory direction in favor of a fixed direction of as yet unknown origin. Received: 3 April 2000 / Accepted in revised form: 19 June 2000     

Seasonal patterns in the orientation system of the migratory ant Pachycondyla marginata

Route directions of migrations by the neotropical termite-hunting ant Pachycondyla marginata at a forest reserve in Southeast Brazil were analysed by circular statistic. Colony movement patterns were compared between the rainy/hot and dry/cold seasons. Migrations during the dry/cold season are significantly oriented 13 degrees with the magnetic North-South axis, while rainy/hot migrations do not exhibit a preferred direction. This result is discussed considering the hypothesis that P. marginata ants may use the geomagnetic field as an orientation cue for migrations in the dry/cold season. The presence of magnetic iron oxides in the head and abdomen of P. marginata is consistent with this suggestion.     

Positive polarotaxis in a mayfly that never leaves the water surface: polarotactic water detection in Palingenia longicauda (Ephemeroptera)

Tisza mayflies, Palingenia longicauda (Olivier 1791), swarm exclusively over the river Tisza (from which the name of the mayfly was derived). This river is bordered by a high vertical wall of trees and bushes, which hinder P. longicauda to move away horizontally from the water. During swarming, Tisza mayflies fly immediately above the river in such a way that their cerci touch the water frequently or sweep its surface. This continuous close connection with water and the vertical wall of the shore and riparian vegetation result in that Tisza mayflies never leave the water surface; consequently, they need not search for water. Several Ephemeroptera species move away far from water and return to it guided by the horizontal polarization of water-reflected light. To reveal whether also P. longicauda is or is not polarotactic, we performed a field experiment during the very short swarming period of Tisza mayflies. We show here that also P. longicauda has positive polarotaxis, which, however, can be observed only under unnatural conditions, when the animals are displaced from the water and then released above artificial test surfaces. P. longicauda is the first species in which polarotactic water detection is demonstrated albeit it never leaves the water surface, and thus, a polarotactic water detection seems unnecessary for it. The polarotactic behaviour of Tisza mayflies explains the earlier observation that these insects swarm above wet asphalt roads running next to river Tisza.     

An absence of aggression between non-nestmates in the bull ant <Emphasis Type="Italic">Myrmecia nigriceps</Emphasis>

The ability of social insects to discriminate against non-nestmates is vital for maintaining colony integrity, and in most social insect species, individuals act aggressively towards non-nestmates that intrude into their nest. Our experimental field data revealed that intra-colony aggression in the primitive bulldog ant Myrmecia nigriceps is negligible; our series of bioassays revealed no significant difference in the occurrence of aggression in trials involving workers from the same, a close (less than 300 m) or a far (more than 1.5 km) nest. Further, non-nestmate intruders were able to enter the nest in 60% of our trials; a similar level was observed in trials involving nestmates. These results suggest that workers of M. nigriceps are either unable to recognize alien conspecifics or that the costs of ignoring workers from foreign colonies are sufficiently low to favor low levels of inter-colony aggression in this species.     

Motivation and vector navigation in honey bees

Animals that forage from a central place can keep track of their displacement relative to home through a process called "path integration." During a study of the stability of homing information over time, we noticed that honey bees held at a feeding place for several hours sometimes headed not in the homeward compass direction on their release, but in the reverse compass direction. This behavior suggested that the path integration system had been reset to a state corresponding to an outward flight to the food. Most models of insect navigation assume that it is the experience of reaching home that resets the path integration system, enabling the activation of vectors appropriate for subsequent outbound foraging trips. Here we provide evidence that this resetting can be influenced by motivational cues associated with food deprivation. The effect of food deprivation is independent of any positional cues provided by familiar landmarks or by experience in traveling toward a goal.     

Loft features reveal the functioning of the young pigeon’s navigational system

It is thought that young homing pigeons are able to use information acquired en route for their initial homeward orientation. However, the cues involved and mechanisms utilised are under discussion. Blocking light-dependent route-specific information during the first leg of an outward journey detour, together with analysis of pigeons that were raised under different loft conditions, allowed us to correctly evaluate the functioning of this mechanism and, more generally, the navigational map of birds. Pigeons from the same stock were raised and kept in two different lofts. The birds in the experimental groups were transported to the release sites via detours, and light-dependent information was denied during the first half of the outward journey (no compass information was available). Control birds were transported by the most direct route and had access to all available information. In general, the results showed that the low-loft birds preferred to use magnetic compass cues, whereas the high-loft birds preferred to use navigational map cues to collect information of the first part of the outward journey. The impairments observed in the homing performances of the experimental groups highlight the reliability of information collected inside the map area. Relevant to an understanding of the route-reversal mechanism was the evidence that this mechanism is able to function in the absence of compass information (birds raised in a wind-exposed loft show a detour effect). In systems where directional information could be provided by multiple sources, processing and extracting accurate course trajectories through a common mechanism may prove more efficient and reliable.