Wolfgang Wiltschko - Academia.edu (original) (raw)

Papers by Wolfgang Wiltschko

Research paper thumbnail of Olfactory navigation versus olfactory activation: a controversy revisited

Journal of Comparative Physiology A, 2018

In the early 1970s, Floriano Papi and colleagues proposed the olfactory-navigation hypothesis, wh... more In the early 1970s, Floriano Papi and colleagues proposed the olfactory-navigation hypothesis, which explains the homing ability of pigeons by the existence of an odor-based map acquired through learning. This notion, although supported by some observations, has also generated considerable controversy since its inception. As an alternative, Paulo Jorge and colleagues formulated in 2009 the olfactory-activation hypothesis, which states that atmospheric odorants do not provide navigational information but, instead, activate a non-olfactory path integration system. However, this hypothesis is challenged by an investigation authored by Anna Gagliardo and colleagues and published in the current issue of the Journal of Comparative Physiology A. In this editorial, the significance of the findings of this study is assessed in the broader context of the role of olfaction in avian navigation and homing, and experiments are suggested that might help to finally resolve the olfactorynavigation versus olfactory-activation controversy.

Research paper thumbnail of Magnetoreception in birds

Journal of The Royal Society Interface, 2019

Birds can use two kinds of information from the geomagnetic field for navigation: the direction o... more Birds can use two kinds of information from the geomagnetic field for navigation: the direction of the field lines as a compass and probably magnetic intensity as a component of the navigational ‘map’. The direction of the magnetic field appears to be sensed via radical pair processes in the eyes, with the crucial radical pairs formed by cryptochrome. It is transmitted by the optic nerve to the brain, where parts of the visual system seem to process the respective information. Magnetic intensity appears to be perceived by magnetite-based receptors in the beak region; the information is transmitted by the ophthalmic branch of the trigeminal nerve to the trigeminal ganglion and the trigeminal brainstem nuclei. Yet in spite of considerable progress in recent years, many details are still unclear, among them details of the radical pair processes and their transformation into a nervous signal, the precise location of the magnetite-based receptors and the centres in the brain where magnet...

Research paper thumbnail of Changes in the Migratory Direction of Yellow-faced HoneyeatersLichenostomus chrysops(Meliphagidae) during Autumn Migration

Emu - Austral Ornithology, 1993

Research paper thumbnail of Magnetoreception: activation of avian cryptochrome 1a in various light conditions

Journal of Comparative Physiology A, 2018

The avian magnetic inclination compass is based on radical pair processes, with cryptochrome (Cry... more The avian magnetic inclination compass is based on radical pair processes, with cryptochrome (Cry) assumed to form the crucial radical pairs; it requires short-wavelength light from UV to green. Under high-intensity narrow-band lights and when yellow light is added, the magnetic compass is disrupted: migratory birds no longer prefer their migratory direction, but show other orientation responses. The candidate receptor molecule Cry1a is located in the shortwavelength-sensitive SWS1 cone photoreceptors in the retina. The present analysis of avian retinae after the respective illuminations showed that no activated Cry1a was present under 565 nm green light of medium and high intensity, and hardly any under high intensity 502 nm turquoise, whereas we found activated Cry1a at all three tested intensities of 373 nm UV and 424 nm blue light. Activated Cry1a also was found when 590 nm yellow light was added to low-intensity light of the four colors; yet these light combinations result in impaired magnetic orientation. This indicates that the disruption of the magnetic compass does not occur at the receptor level in the retina, but at higher processing stages, where the unnatural, almost monochromatic or bichromatic illumination causes yet unknown responses that interfere with the inclination compass.

Research paper thumbnail of Effect of a magnetic pulse on the orientation of silvereyes, zosterops l. lateralis, during spring migration

Journal of Experimental Biology, 1998

The orientation behaviour of Australian silvereyes, Zosterops l. lateralis, was tested during the... more The orientation behaviour of Australian silvereyes, Zosterops l. lateralis, was tested during their spring migration, when they head southward to their Tasmanian breeding grounds. With only the local geomagnetic field as a cue, the birds significantly preferred their normal southerly migratory direction. Treatment with a short, strong magnetic pulse designed to alter the magnetization of single-domain magnetite led to a significant deflection towards the east for the next 4 days. This was followed by a period of non-oriented behaviour. From day 10 onwards, the birds returned to their original southerly headings. Together with previous findings, these data suggest that the navigational ‘map’ of these birds includes magnetic parameters and that a magnetite-based receptor provides them with information about their position. The transient nature of the effect is not easily explained on the basis of single-domain magnetite.

Research paper thumbnail of Avian magnetic compass: Its functional properties and physical basis

Current Zoology, 2010

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

Research paper thumbnail of Lateralization of the Avian Magnetic Compass: Analysis of Its Early Plasticity

Symmetry, 2017

In European Robins, Erithacus rubecula, the magnetic compass is lateralized in favor of the right... more In European Robins, Erithacus rubecula, the magnetic compass is lateralized in favor of the right eye/left hemisphere of the brain. This lateralization develops during the first winter and initially shows a great plasticity. During the first spring migration, it can be temporarily removed by covering the right eye. In the present paper, we used the migratory orientation of robins to analyze the circumstances under which the lateralization can be undone. Already a period of 1 1 /2 h being monocularly left-eyed before tests began proved sufficient to restore the ability to use the left eye for orientation, but this effect was rather short-lived, as lateralization recurred again within the next 1 1 /2 h. Interpretable magnetic information mediated by the left eye was necessary for removing the lateralization. In addition, monocularly, the left eye seeing robins could adjust to magnetic intensities outside the normal functional window, but this ability was not transferred to the "right-eye system". Our results make it clear that asymmetry of magnetic compass perception is amenable to short-term changes, depending on lateralized stimulation. This could mean that the left hemispheric dominance for the analysis of magnetic compass information depends on lateralized interhemispheric interactions that in young birds can swiftly be altered by environmental effects.

Research paper thumbnail of Seasonally Changing Cryptochrome 1b Expression in the Retinal Ganglion Cells of a Migrating Passerine Bird

PloS one, 2016

Cryptochromes, blue-light absorbing proteins involved in the circadian clock, have been proposed ... more Cryptochromes, blue-light absorbing proteins involved in the circadian clock, have been proposed to be the receptor molecules of the avian magnetic compass. In birds, several cryptochromes occur: Cryptochrome 2, Cryptochrome 4 and two splice products of Cryptochrome 1, Cry1a and Cry1b. With an antibody not distinguishing between the two splice products, Cryptochrome 1 had been detected in the retinal ganglion cells of garden warblers during migration. A recent study located Cry1a in the outer segments of UV/V-cones in the retina of domestic chickens and European robins, another migratory species. Here we report the presence of cryptochrome 1b (eCry1b) in retinal ganglion cells and displaced ganglion cells of European Robins, Erithacus rubecula. Immuno-histochemistry at the light microscopic and electron microscopic level showed eCry1b in the cell plasma, free in the cytosol as well as bound to membranes. This is supported by immuno-blotting. However, this applies only to robins in t...

Research paper thumbnail of Magnetite-based magnetoreception: the eVect of repeated pulsing on the orientation of migratory birds

J Comp Physiol a, 2007

Previous studies have shown that a magnetic pulse affected the orientation of passerine migrants ... more Previous studies have shown that a magnetic pulse affected the orientation of passerine migrants for a short period only: for about 3 days, the birds' headings were deflected eastward from their migratory direction, followed by a phase of disorientation, with the birds returning to their normal migratory direction after about 10 days. To analyze the processes involved in the fading of the pulse effect, migratory birds were subjected to a second, identical pulse 16 days after the first pulse, when the effect of that pulse had disappeared. This second pulse affected the birds' behavior in a different way: it caused an increase in the scatter of the birds' headings for 2 days, after which the birds showed normal migratory orientation again. These observations are at variance with the hypothesis that the magnetite-based receptor had been fully restored, but also with the hypothesis that the input of this receptor was ignored. They rather indicate dynamic processes, which include changes in the affected receptor, but at the same time cause the birds to weigh and rate the altered input differently. The bearing of these findings on the question of whether single domains or superparamagnetic particles are involved in the magnetite-based receptors is discussed.

Research paper thumbnail of ‘Fixed direction’-responses of birds in the geomagnetic field

Communicative & Integrative Biology, 2009

Research paper thumbnail of Magnetoreception in birds: different physical processes for two types of directional responses

HFSP journal, 2007

Migratory orientation in birds involves an inclination compass based on radical-pair processes. U... more Migratory orientation in birds involves an inclination compass based on radical-pair processes. Under certain light regimes, however, "fixed-direction" responses are observed that do not undergo the seasonal change between spring and autumn typical for migratory orientation. To identify the underlying transduction mechanisms, we analyzed a fixed-direction response under a combination of 502 nm turquoise and 590 nm yellow light, with migratory orientation under 565 nm green light serving as the control. High-frequency fields, diagnostic for a radical-pair mechanism, disrupted migratory orientation without affecting fixed-direction responses. Local anaesthesia of the upper beak where magnetite is found in birds, in contrast, disrupted the fixed-direction response without affecting migratory orientation. The two types of responses are thus based on different physical principles, with the compass response based on a radical pair mechanism and the fixed-direction responses prob...

Research paper thumbnail of Light-dependent magnetoreception in birds: increasing intensity of monochromatic light changes the nature of the response

Frontiers in zoology, Jan 15, 2007

The Radical Pair model proposes that magnetoreception is a light-dependent process. Under low mon... more The Radical Pair model proposes that magnetoreception is a light-dependent process. Under low monochromatic light from the short-wavelength part of the visual spectrum, migratory birds show orientation in their migratory direction. Under monochromatic light of higher intensity, however, they showed unusual preferences for other directions or axial preferences. To determine whether or not these responses are still controlled by the respective light regimes, European robins, Erithacus rubecula, were tested under UV, Blue, Turquoise and Green light at increasing intensities, with orientation in migratory direction serving as a criterion whether or not magnetoreception works in the normal way. The birds were well oriented in their seasonally appropriate migratory direction under 424 nm Blue, 502 nm Turquoise and 565 nm Green light of low intensity with a quantal flux of 8 x 10(15) quanta s(-1) m(-2), indicating unimpaired magnetoreception. Under 373 nm UV of the same quantal flux, they ...

Research paper thumbnail of Avian ultraviolet/violet cones as magnetoreceptors: The problem of separating visual and magnetic information

Communicative & Integrative Biology, 2011

Research paper thumbnail of Ontogenetic development of magnetic compass orientation in domestic chickens (Gallus gallus)

Journal of Experimental Biology, 2013

Summary Domestic chickens (Gallus gallus) can be trained to search for a social stimulus in a spe... more Summary Domestic chickens (Gallus gallus) can be trained to search for a social stimulus in a specific magnetic direction, and cryptochrome 1a found in the retina has been proposed as a receptor molecule mediating magnetic directions. The present study combines immuno-histochemical and behavioural data to analyse the ontogenetic development of this ability. Newly hatched chicks already have a small amount of cryptochrome 1a in their violet cones; on day 5, the amount of cryptochrome 1a has reached the same level as in adult chickens, suggesting that the physical basis for magnetoreception is present. In behavioural tests, however, young chicks 5 to 7 days old failed to show a preference of the training direction; on days 8, 9 and 12, they could be successfully trained to search along a specific magnetic axis. Trained and tested again a week later, the chicks that had not shown a directional preference on day 5 to 7 continued to search randomly, while the chicks tested from day 8 onw...

Research paper thumbnail of Magnetic orientation of migratory robins, Erithacus rubecula, under long-wavelength light

Journal of Experimental Biology, 2011

SUMMARY The avian magnetic compass is an inclination compass that appears to be based on radical ... more SUMMARY The avian magnetic compass is an inclination compass that appears to be based on radical pair processes. It requires light from the short-wavelength range of the spectrum up to 565 nm green light; under longer wavelengths, birds are disoriented. When pre-exposed to longer wavelengths for 1 h, however, they show oriented behavior. This orientation is analyzed under 582 nm yellow light and 645 nm red light in the present study: while the birds in spring prefer northerly directions, they do not show southerly tendencies in autumn. Inversion of the vertical component does not have an effect whereas reversal of the horizontal component leads to a corresponding shift, indicating that a polar response to the magnetic field is involved. Oscillating magnetic fields in the MHz range do not affect the behavior but anesthesia of the upper beak causes disorientation. This indicates that the magnetic information is no longer provided by the radical pair mechanism in the eye but by the mag...

Research paper thumbnail of Magnetic orientation in birds: non–compass responses under monochromatic light of increased intensity

Proceedings of the Royal Society of London. Series B: Biological Sciences, 2003

Migratory Australian silvereyes (Zosterops lateralis) were tested under monochromatic light at wa... more Migratory Australian silvereyes (Zosterops lateralis) were tested under monochromatic light at wavelengths of 424 nm blue and 565 nm green. At a low light level of 7´10 1 5 quanta m 22 s 21 in the local geomagnetic field, the birds preferred their seasonally appropriate southern migratory direction under both wavelengths. Their reversal of headings when the vertical component of the magnetic field was inverted indicated normal use of the avian inclination compass. A higher light intensity of 43´10 1 5 quanta m 22 s 2 1 , however, caused a fundamental change in behaviour: under bright blue, the silvereyes showed an axial tendency along the east-west axis; under bright green, they showed a unimodal preference of a west-northwesterly direction that followed a shift in magnetic north, but was not reversed by inverting the vertical component of the magnetic field. Hence it is not based on the inclination compass. The change in behaviour at higher light intensities suggests a complex interaction between at least two receptors. The polar nature of the response under bright green cannot be explained by the current models of light-dependent magnetoreception and will lead to new considerations on these receptive processes.

Research paper thumbnail of Avian Ultraviolet/Violet Cones Identified as Probable Magnetoreceptors

PLoS ONE, 2011

Background: The Radical-Pair-Model postulates that the reception of magnetic compass directions i... more Background: The Radical-Pair-Model postulates that the reception of magnetic compass directions in birds is based on spinchemical reactions in specialized photopigments in the eye, with cryptochromes discussed as candidate molecules. But so far, the exact subcellular characterization of these molecules in the retina remained unknown. Methodology/Principal Findings: We here describe the localization of cryptochrome 1a (Cry1a) in the retina of European robins, Erithacus rubecula, and domestic chickens, Gallus gallus, two species that have been shown to use the magnetic field for compass orientation. In both species, Cry1a is present exclusively in the ultraviolet/violet (UV/V) cones that are distributed across the entire retina. Electron microscopy shows Cry1a in ordered bands along the membrane discs of the outer segment, and cell fractionation reveals Cry1a in the membrane fraction, suggesting the possibility that Cry1a is anchored along membranes. Conclusions/Significance: We provide first structural evidence that Cry1a occurs within a sensory structure arranged in a way that fulfils essential requirements of the Radical-Pair-Model. Our findings, identifying the UV/V-cones as probable magnetoreceptors, support the assumption that Cry1a is indeed the receptor molecule mediating information on magnetic directions, and thus provide the Radical-Pair-Model with a profound histological background.

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

Naturwissenschaften, 2000

In a previous study, Australian silvereyes tested in autumn under monochromatic 565-nm green ligh... more 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.

Research paper thumbnail of Light-Dependent Magnetoreception in Birds: Does Directional Information Change with Light Intensity?

Naturwissenschaften, 2000

Magnetic compass orientation in birds is based on light-dependent processes, with magnetoreceptio... more Magnetic compass orientation in birds is based on light-dependent processes, with magnetoreception being possible only under light containing blue and green wavelengths. To look for possible intensity-dependent effects we tested Australian silvereyes during autumn migration under monochromatic green light (565 nm) produced by light-emitting diodes at various light levels. At intensities of 0.0021 and 0.0075 W/m 2 , the birds showed normal activity and were oriented in their seasonally appropriate migratory direction. Under low light of 0.0002 W/m 2 the birds were less active; scatter increased, but they still oriented in their migratory direction. Under a high light level of 0.0150 W/m 2 , however, the test birds showed a counterclockwise shift in direction, preferring west-northwest instead of north. This change in behavior may reflect a change in the output of the magnetoreception system, resulting from a disruption of the natural balance between the wavelengths of light.

Research paper thumbnail of Directional orientation of birds by the magnetic field under different light conditions

Journal of The Royal Society Interface, 2009

This paper reviews the directional orientation of birds with the help of the geomagnetic field un... more This paper reviews the directional orientation of birds with the help of the geomagnetic field under various light conditions. Two fundamentally different types of response can be distinguished. (i) Compass orientation controlled by the inclination compass that allows birds to locate courses of different origin. This is restricted to a narrow functional window around the total intensity of the local geomagnetic field and requires light from the short-wavelength part of the spectrum. The compass is based on radical-pair processes in the right eye; magnetite-based receptors in the beak are not involved. Compass orientation is observed under ‘white’ and low-level monochromatic light from ultraviolet (UV) to about 565 nm green light. (ii) ‘Fixed direction’ responses occur under artificial light conditions such as more intense monochromatic light, when 590 nm yellow light is added to short-wavelength light, and in total darkness. The manifestation of these responses depends on the ambien...

Research paper thumbnail of Olfactory navigation versus olfactory activation: a controversy revisited

Journal of Comparative Physiology A, 2018

In the early 1970s, Floriano Papi and colleagues proposed the olfactory-navigation hypothesis, wh... more In the early 1970s, Floriano Papi and colleagues proposed the olfactory-navigation hypothesis, which explains the homing ability of pigeons by the existence of an odor-based map acquired through learning. This notion, although supported by some observations, has also generated considerable controversy since its inception. As an alternative, Paulo Jorge and colleagues formulated in 2009 the olfactory-activation hypothesis, which states that atmospheric odorants do not provide navigational information but, instead, activate a non-olfactory path integration system. However, this hypothesis is challenged by an investigation authored by Anna Gagliardo and colleagues and published in the current issue of the Journal of Comparative Physiology A. In this editorial, the significance of the findings of this study is assessed in the broader context of the role of olfaction in avian navigation and homing, and experiments are suggested that might help to finally resolve the olfactorynavigation versus olfactory-activation controversy.

Research paper thumbnail of Magnetoreception in birds

Journal of The Royal Society Interface, 2019

Birds can use two kinds of information from the geomagnetic field for navigation: the direction o... more Birds can use two kinds of information from the geomagnetic field for navigation: the direction of the field lines as a compass and probably magnetic intensity as a component of the navigational ‘map’. The direction of the magnetic field appears to be sensed via radical pair processes in the eyes, with the crucial radical pairs formed by cryptochrome. It is transmitted by the optic nerve to the brain, where parts of the visual system seem to process the respective information. Magnetic intensity appears to be perceived by magnetite-based receptors in the beak region; the information is transmitted by the ophthalmic branch of the trigeminal nerve to the trigeminal ganglion and the trigeminal brainstem nuclei. Yet in spite of considerable progress in recent years, many details are still unclear, among them details of the radical pair processes and their transformation into a nervous signal, the precise location of the magnetite-based receptors and the centres in the brain where magnet...

Research paper thumbnail of Changes in the Migratory Direction of Yellow-faced HoneyeatersLichenostomus chrysops(Meliphagidae) during Autumn Migration

Emu - Austral Ornithology, 1993

Research paper thumbnail of Magnetoreception: activation of avian cryptochrome 1a in various light conditions

Journal of Comparative Physiology A, 2018

The avian magnetic inclination compass is based on radical pair processes, with cryptochrome (Cry... more The avian magnetic inclination compass is based on radical pair processes, with cryptochrome (Cry) assumed to form the crucial radical pairs; it requires short-wavelength light from UV to green. Under high-intensity narrow-band lights and when yellow light is added, the magnetic compass is disrupted: migratory birds no longer prefer their migratory direction, but show other orientation responses. The candidate receptor molecule Cry1a is located in the shortwavelength-sensitive SWS1 cone photoreceptors in the retina. The present analysis of avian retinae after the respective illuminations showed that no activated Cry1a was present under 565 nm green light of medium and high intensity, and hardly any under high intensity 502 nm turquoise, whereas we found activated Cry1a at all three tested intensities of 373 nm UV and 424 nm blue light. Activated Cry1a also was found when 590 nm yellow light was added to low-intensity light of the four colors; yet these light combinations result in impaired magnetic orientation. This indicates that the disruption of the magnetic compass does not occur at the receptor level in the retina, but at higher processing stages, where the unnatural, almost monochromatic or bichromatic illumination causes yet unknown responses that interfere with the inclination compass.

Research paper thumbnail of Effect of a magnetic pulse on the orientation of silvereyes, zosterops l. lateralis, during spring migration

Journal of Experimental Biology, 1998

The orientation behaviour of Australian silvereyes, Zosterops l. lateralis, was tested during the... more The orientation behaviour of Australian silvereyes, Zosterops l. lateralis, was tested during their spring migration, when they head southward to their Tasmanian breeding grounds. With only the local geomagnetic field as a cue, the birds significantly preferred their normal southerly migratory direction. Treatment with a short, strong magnetic pulse designed to alter the magnetization of single-domain magnetite led to a significant deflection towards the east for the next 4 days. This was followed by a period of non-oriented behaviour. From day 10 onwards, the birds returned to their original southerly headings. Together with previous findings, these data suggest that the navigational ‘map’ of these birds includes magnetic parameters and that a magnetite-based receptor provides them with information about their position. The transient nature of the effect is not easily explained on the basis of single-domain magnetite.

Research paper thumbnail of Avian magnetic compass: Its functional properties and physical basis

Current Zoology, 2010

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

Research paper thumbnail of Lateralization of the Avian Magnetic Compass: Analysis of Its Early Plasticity

Symmetry, 2017

In European Robins, Erithacus rubecula, the magnetic compass is lateralized in favor of the right... more In European Robins, Erithacus rubecula, the magnetic compass is lateralized in favor of the right eye/left hemisphere of the brain. This lateralization develops during the first winter and initially shows a great plasticity. During the first spring migration, it can be temporarily removed by covering the right eye. In the present paper, we used the migratory orientation of robins to analyze the circumstances under which the lateralization can be undone. Already a period of 1 1 /2 h being monocularly left-eyed before tests began proved sufficient to restore the ability to use the left eye for orientation, but this effect was rather short-lived, as lateralization recurred again within the next 1 1 /2 h. Interpretable magnetic information mediated by the left eye was necessary for removing the lateralization. In addition, monocularly, the left eye seeing robins could adjust to magnetic intensities outside the normal functional window, but this ability was not transferred to the "right-eye system". Our results make it clear that asymmetry of magnetic compass perception is amenable to short-term changes, depending on lateralized stimulation. This could mean that the left hemispheric dominance for the analysis of magnetic compass information depends on lateralized interhemispheric interactions that in young birds can swiftly be altered by environmental effects.

Research paper thumbnail of Seasonally Changing Cryptochrome 1b Expression in the Retinal Ganglion Cells of a Migrating Passerine Bird

PloS one, 2016

Cryptochromes, blue-light absorbing proteins involved in the circadian clock, have been proposed ... more Cryptochromes, blue-light absorbing proteins involved in the circadian clock, have been proposed to be the receptor molecules of the avian magnetic compass. In birds, several cryptochromes occur: Cryptochrome 2, Cryptochrome 4 and two splice products of Cryptochrome 1, Cry1a and Cry1b. With an antibody not distinguishing between the two splice products, Cryptochrome 1 had been detected in the retinal ganglion cells of garden warblers during migration. A recent study located Cry1a in the outer segments of UV/V-cones in the retina of domestic chickens and European robins, another migratory species. Here we report the presence of cryptochrome 1b (eCry1b) in retinal ganglion cells and displaced ganglion cells of European Robins, Erithacus rubecula. Immuno-histochemistry at the light microscopic and electron microscopic level showed eCry1b in the cell plasma, free in the cytosol as well as bound to membranes. This is supported by immuno-blotting. However, this applies only to robins in t...

Research paper thumbnail of Magnetite-based magnetoreception: the eVect of repeated pulsing on the orientation of migratory birds

J Comp Physiol a, 2007

Previous studies have shown that a magnetic pulse affected the orientation of passerine migrants ... more Previous studies have shown that a magnetic pulse affected the orientation of passerine migrants for a short period only: for about 3 days, the birds' headings were deflected eastward from their migratory direction, followed by a phase of disorientation, with the birds returning to their normal migratory direction after about 10 days. To analyze the processes involved in the fading of the pulse effect, migratory birds were subjected to a second, identical pulse 16 days after the first pulse, when the effect of that pulse had disappeared. This second pulse affected the birds' behavior in a different way: it caused an increase in the scatter of the birds' headings for 2 days, after which the birds showed normal migratory orientation again. These observations are at variance with the hypothesis that the magnetite-based receptor had been fully restored, but also with the hypothesis that the input of this receptor was ignored. They rather indicate dynamic processes, which include changes in the affected receptor, but at the same time cause the birds to weigh and rate the altered input differently. The bearing of these findings on the question of whether single domains or superparamagnetic particles are involved in the magnetite-based receptors is discussed.

Research paper thumbnail of ‘Fixed direction’-responses of birds in the geomagnetic field

Communicative & Integrative Biology, 2009

Research paper thumbnail of Magnetoreception in birds: different physical processes for two types of directional responses

HFSP journal, 2007

Migratory orientation in birds involves an inclination compass based on radical-pair processes. U... more Migratory orientation in birds involves an inclination compass based on radical-pair processes. Under certain light regimes, however, "fixed-direction" responses are observed that do not undergo the seasonal change between spring and autumn typical for migratory orientation. To identify the underlying transduction mechanisms, we analyzed a fixed-direction response under a combination of 502 nm turquoise and 590 nm yellow light, with migratory orientation under 565 nm green light serving as the control. High-frequency fields, diagnostic for a radical-pair mechanism, disrupted migratory orientation without affecting fixed-direction responses. Local anaesthesia of the upper beak where magnetite is found in birds, in contrast, disrupted the fixed-direction response without affecting migratory orientation. The two types of responses are thus based on different physical principles, with the compass response based on a radical pair mechanism and the fixed-direction responses prob...

Research paper thumbnail of Light-dependent magnetoreception in birds: increasing intensity of monochromatic light changes the nature of the response

Frontiers in zoology, Jan 15, 2007

The Radical Pair model proposes that magnetoreception is a light-dependent process. Under low mon... more The Radical Pair model proposes that magnetoreception is a light-dependent process. Under low monochromatic light from the short-wavelength part of the visual spectrum, migratory birds show orientation in their migratory direction. Under monochromatic light of higher intensity, however, they showed unusual preferences for other directions or axial preferences. To determine whether or not these responses are still controlled by the respective light regimes, European robins, Erithacus rubecula, were tested under UV, Blue, Turquoise and Green light at increasing intensities, with orientation in migratory direction serving as a criterion whether or not magnetoreception works in the normal way. The birds were well oriented in their seasonally appropriate migratory direction under 424 nm Blue, 502 nm Turquoise and 565 nm Green light of low intensity with a quantal flux of 8 x 10(15) quanta s(-1) m(-2), indicating unimpaired magnetoreception. Under 373 nm UV of the same quantal flux, they ...

Research paper thumbnail of Avian ultraviolet/violet cones as magnetoreceptors: The problem of separating visual and magnetic information

Communicative & Integrative Biology, 2011

Research paper thumbnail of Ontogenetic development of magnetic compass orientation in domestic chickens (Gallus gallus)

Journal of Experimental Biology, 2013

Summary Domestic chickens (Gallus gallus) can be trained to search for a social stimulus in a spe... more Summary Domestic chickens (Gallus gallus) can be trained to search for a social stimulus in a specific magnetic direction, and cryptochrome 1a found in the retina has been proposed as a receptor molecule mediating magnetic directions. The present study combines immuno-histochemical and behavioural data to analyse the ontogenetic development of this ability. Newly hatched chicks already have a small amount of cryptochrome 1a in their violet cones; on day 5, the amount of cryptochrome 1a has reached the same level as in adult chickens, suggesting that the physical basis for magnetoreception is present. In behavioural tests, however, young chicks 5 to 7 days old failed to show a preference of the training direction; on days 8, 9 and 12, they could be successfully trained to search along a specific magnetic axis. Trained and tested again a week later, the chicks that had not shown a directional preference on day 5 to 7 continued to search randomly, while the chicks tested from day 8 onw...

Research paper thumbnail of Magnetic orientation of migratory robins, Erithacus rubecula, under long-wavelength light

Journal of Experimental Biology, 2011

SUMMARY The avian magnetic compass is an inclination compass that appears to be based on radical ... more SUMMARY The avian magnetic compass is an inclination compass that appears to be based on radical pair processes. It requires light from the short-wavelength range of the spectrum up to 565 nm green light; under longer wavelengths, birds are disoriented. When pre-exposed to longer wavelengths for 1 h, however, they show oriented behavior. This orientation is analyzed under 582 nm yellow light and 645 nm red light in the present study: while the birds in spring prefer northerly directions, they do not show southerly tendencies in autumn. Inversion of the vertical component does not have an effect whereas reversal of the horizontal component leads to a corresponding shift, indicating that a polar response to the magnetic field is involved. Oscillating magnetic fields in the MHz range do not affect the behavior but anesthesia of the upper beak causes disorientation. This indicates that the magnetic information is no longer provided by the radical pair mechanism in the eye but by the mag...

Research paper thumbnail of Magnetic orientation in birds: non–compass responses under monochromatic light of increased intensity

Proceedings of the Royal Society of London. Series B: Biological Sciences, 2003

Migratory Australian silvereyes (Zosterops lateralis) were tested under monochromatic light at wa... more Migratory Australian silvereyes (Zosterops lateralis) were tested under monochromatic light at wavelengths of 424 nm blue and 565 nm green. At a low light level of 7´10 1 5 quanta m 22 s 21 in the local geomagnetic field, the birds preferred their seasonally appropriate southern migratory direction under both wavelengths. Their reversal of headings when the vertical component of the magnetic field was inverted indicated normal use of the avian inclination compass. A higher light intensity of 43´10 1 5 quanta m 22 s 2 1 , however, caused a fundamental change in behaviour: under bright blue, the silvereyes showed an axial tendency along the east-west axis; under bright green, they showed a unimodal preference of a west-northwesterly direction that followed a shift in magnetic north, but was not reversed by inverting the vertical component of the magnetic field. Hence it is not based on the inclination compass. The change in behaviour at higher light intensities suggests a complex interaction between at least two receptors. The polar nature of the response under bright green cannot be explained by the current models of light-dependent magnetoreception and will lead to new considerations on these receptive processes.

Research paper thumbnail of Avian Ultraviolet/Violet Cones Identified as Probable Magnetoreceptors

PLoS ONE, 2011

Background: The Radical-Pair-Model postulates that the reception of magnetic compass directions i... more Background: The Radical-Pair-Model postulates that the reception of magnetic compass directions in birds is based on spinchemical reactions in specialized photopigments in the eye, with cryptochromes discussed as candidate molecules. But so far, the exact subcellular characterization of these molecules in the retina remained unknown. Methodology/Principal Findings: We here describe the localization of cryptochrome 1a (Cry1a) in the retina of European robins, Erithacus rubecula, and domestic chickens, Gallus gallus, two species that have been shown to use the magnetic field for compass orientation. In both species, Cry1a is present exclusively in the ultraviolet/violet (UV/V) cones that are distributed across the entire retina. Electron microscopy shows Cry1a in ordered bands along the membrane discs of the outer segment, and cell fractionation reveals Cry1a in the membrane fraction, suggesting the possibility that Cry1a is anchored along membranes. Conclusions/Significance: We provide first structural evidence that Cry1a occurs within a sensory structure arranged in a way that fulfils essential requirements of the Radical-Pair-Model. Our findings, identifying the UV/V-cones as probable magnetoreceptors, support the assumption that Cry1a is indeed the receptor molecule mediating information on magnetic directions, and thus provide the Radical-Pair-Model with a profound histological background.

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

Naturwissenschaften, 2000

In a previous study, Australian silvereyes tested in autumn under monochromatic 565-nm green ligh... more 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.

Research paper thumbnail of Light-Dependent Magnetoreception in Birds: Does Directional Information Change with Light Intensity?

Naturwissenschaften, 2000

Magnetic compass orientation in birds is based on light-dependent processes, with magnetoreceptio... more Magnetic compass orientation in birds is based on light-dependent processes, with magnetoreception being possible only under light containing blue and green wavelengths. To look for possible intensity-dependent effects we tested Australian silvereyes during autumn migration under monochromatic green light (565 nm) produced by light-emitting diodes at various light levels. At intensities of 0.0021 and 0.0075 W/m 2 , the birds showed normal activity and were oriented in their seasonally appropriate migratory direction. Under low light of 0.0002 W/m 2 the birds were less active; scatter increased, but they still oriented in their migratory direction. Under a high light level of 0.0150 W/m 2 , however, the test birds showed a counterclockwise shift in direction, preferring west-northwest instead of north. This change in behavior may reflect a change in the output of the magnetoreception system, resulting from a disruption of the natural balance between the wavelengths of light.

Research paper thumbnail of Directional orientation of birds by the magnetic field under different light conditions

Journal of The Royal Society Interface, 2009

This paper reviews the directional orientation of birds with the help of the geomagnetic field un... more This paper reviews the directional orientation of birds with the help of the geomagnetic field under various light conditions. Two fundamentally different types of response can be distinguished. (i) Compass orientation controlled by the inclination compass that allows birds to locate courses of different origin. This is restricted to a narrow functional window around the total intensity of the local geomagnetic field and requires light from the short-wavelength part of the spectrum. The compass is based on radical-pair processes in the right eye; magnetite-based receptors in the beak are not involved. Compass orientation is observed under ‘white’ and low-level monochromatic light from ultraviolet (UV) to about 565 nm green light. (ii) ‘Fixed direction’ responses occur under artificial light conditions such as more intense monochromatic light, when 590 nm yellow light is added to short-wavelength light, and in total darkness. The manifestation of these responses depends on the ambien...