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The remarkable ability of migratory birds to navigate accurately using the geomagnetic field for journeys of thousands of kilometres is currently thought to arise from radical pair reactions inside a protein called cryptochrome. In this article, we explain the quantum mechanical basis of the radical pair mechanism and why it is currently the dominant theory of compass magnetoreception. We also provide a brief account of two important computational simulation techniques that are used to study the mechanism in cryptochrome: spin dynamics and molecular dynamics. At the end, we provide an overview of current research on quantum mechanical processes in avian cryptochromes and the computational models for describing them.

In addition to other natural orientation cues such as the stars, the sun, landmarks and olfactory cues, migrating birds possess the ability to orient by the Earth’s magnetic field. In recent years, neuroscientific research has pinpointed brain regions and connecting neuronal pathways that seem to be involved in processing magnetic information. To date, the most compelling neuroanatomical and behavioural evidence comes from the visual and trigeminal sensory systems. We expect that navigational information from both systems could be integrated in higher-order brain structures, such as the hippocampus and the “decision-making” caudolateral nidopallium. This review summarizes the current state of research on the neurosensory basis of magnetoreception in birds.

A class of light-activated proteins in the eyes of birds, called cryptochromes, are thought to act as the primary magnetic sensors allowing night-migratory songbirds to navigate over thousands of kilometers using the earth’s magnetic field. Having evolved from DNA-repairing photolyases, cryptochromes have redirected the energy from light to fuel a variety of other functions: as photoreceptors, as regulators of the circadian clock – and, in some species, most likely as sensors of the magnetic field. While the quantum effects of magnetic fields on cryptochromes are already being studied in detail, almost nothing is known about the signaling cascade involving cryptochrome as the primary receptor protein. Two different screening methods have identified potential interaction partners that suggest an involvement of the visual phototransduction pathway, the visual cycle, potassium channels or glutamate receptors, but more pioneering research is needed to unravel the signaling cascade responsible for transducing the magnetic signal.

Ambient levels of electromagnetic fields (EMF) have risen sharply in the last 80 years, creating a novel energetic exposure that previously did not exist. Most recent decades have seen exponential increases in nearly all environments, including rural/remote areas and lower atmospheric regions. Because of unique physiologies, some species of flora and fauna are sensitive to exogenous EMF in ways that may surpass human reactivity. There is limited, but comprehensive, baseline data in the U.S. from the 1980s against which to compare significant new surveys from different countries. This now provides broader and more precise data on potential transient and chronic exposures to wildlife and habitats. Biological effects have been seen broadly across all taxa and frequencies at vanishingly low intensities comparable to today’s ambient exposures. Broad wildlife effects have been seen on orientation and migration, food finding, reproduction, mating, nest and den building, territorial maintenance and defense, and longevity and survivorship. Cyto- and geno-toxic effects have been observed. The above issues are explored in three consecutive parts: Part 1 questions today’s ambient EMF capabilities to adversely affect wildlife, with more urgency regarding 5G technologies. Part 2 explores natural and man-made fields, animal magnetoreception mechanisms, and pertinent studies to all wildlife kingdoms. Part 3 examines current exposure standards, applicable laws, and future directions. It is time to recognize ambient EMF as a novel form of pollution and develop rules at regulatory agencies that designate air as ‘habitat’ so EMF can be regulated like other pollutants. Wildlife loss is often unseen and undocumented until tipping points are reached. Long-term chronic low-level EMF exposure standards, which do not now exist, should be set accordingly for wildlife, and environmental laws should be strictly enforced.

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.

Understanding the rules of life is one of the most important scientific endeavours and has revolutionised both biology and biotechnology. Remarkable advances in observation techniques allow us to investigate a broad range of complex and dynamic biological processes in which living systems could exploit quantum behaviour to enhance and regulate biological functions. Recent evidence suggests that these non-trivial quantum mechanical effects may play a crucial role in maintaining the non-equilibrium state of biomolecular systems. Quantum biology is the study of such quantum aspects of living systems. In this review, we summarise the latest progress in quantum biology, including the areas of enzyme-catalysed reactions, photosynthesis, spin-dependent reactions, DNA, fluorescent proteins, and ion channels. Many of these results are expected to be fundamental building blocks towards understanding the rules of life.

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. This article is protected by copyright. All rights reserved.

Biological material has been documented to produce an external magnetic field that radiates out. There have been several papers documenting the magnetic fields produced by steady currents in the body. The most notable was published in 1980 by Cohen et al. where the human hair follicle was used as sentinel and biophysically evaluated via sophisticated equipment such as a double planar Superconducting Quantum Interference Devices (SQUID). Most recently, in 2019 Cohen’s work was duplicated by Khan,S by also using double-planar gladiometers. Of interest to this manuscript is that since the introduction of anovel optical microscopy method in 2016 by Scherlag BJ et al is that numerous papers have been introduced in the literature now identifying intrinsic biomagnetic properties of the follicle such as penetration through glass barriers. In this manuscript, a concept of biomagnetic fields by the concave part of the human hand transferring energy to hair follicles is introduced, this was accomplished by using a novel optical microscopy method, in other words, the hair follicle is not limited to radiate out biomagnetism; but also, to receive externally radiated biomagnetic fields from a body part. This magneto receptive property is herein introduced.

Cryptochrome proteins are thought to be involved in light-sensitive magnetoreception in migratory birds triggered by flavin adenine dinucleotide (FAD) light absorption. A recent study, however, cal...

A visual magnetic sense in migratory birds has been hypothesized to rely on a radical pair reaction in the protein cryptochrome. In this model, magnetic sensitivity originates from coherent spin dynamics, as the radicals couple to magnetic nuclei via hyperfine interactions. Prior studies have often neglected the electron–electron dipolar (EED) coupling from this hypothesis. We show that EED interactions suppress the anisotropic response to the geomagnetic field by the radical pair mechanism in cryptochrome and that this attenuation is unlikely to be mitigated by mutual cancellation of the EED and electronic exchange coupling, as previously suggested. We then demonstrate that this limitation may be overcome by extending the conventional model to include a third, nonreacting radical. We predict that hyperfine effects could work in concert with three-radical dipolar interactions to tailor a superior magnetic response, thereby providing a new principle for magnetosensitivity with applications for sensing, navigation, and the assessment of biological magnetic field effects.

The biophysical mechanism of the magnetic compass sensor in migratory songbirds is thought to involve photo-induced radical pairs formed in cryptochrome (Cry) flavoproteins located in photoreceptor cells in the eyes. In Cry4a – the most likely of the six known avian cryptochromes to have a magnetic sensing function – four radical pair states are formed sequentially by the stepwise transfer of an electron along a chain of four tryptophan residues to the photo-excited flavin. In purified Cry4a from the migratory European robin, the third of these flavin-tryptophan radical pairs is more magnetically sensitive than the fourth, consistent with the smaller separation of the radicals in the former. Here, we explore the idea that these two radical pair states of Cry4a could exist in rapid dynamic equilibrium such that the key magnetic and kinetic properties are weighted averages. Spin dynamics simulations suggest that the third radical pair is largely responsible for magnetic sensing while the fourth may be better placed to initiate magnetic signalling particularly if the terminal tryptophan radical can be reduced by a nearby tyrosine. Such an arrangement could have allowed independent optimisation of the essential sensing and signalling functions of the protein. It might also rationalise why avian Cry4a has four tryptophans while cryptochromes from plants have only three.

Magnetoreception, sensing the Earth’s magnetic field, is used by many species in orientation and navigation. While this is established on the behavioural level, there is a severe lack in knowledge on the underlying neuronal mechanisms of this sense. A powerful technique to study the neuronal processing of magnetic cues is electrophysiology but, thus far, few studies have adopted this technique. Why is this the case? A fundamental problem is the introduction of electromagnetic noise (induction) caused by the magnetic stimuli, within electrophysiological recordings which, if too large, prevents feasible separation of neuronal signals from the induction artefacts. Here, we address the concerns surrounding the use of electromagnetic coils within electrophysiology experiments and assess whether these would prevent viable electrophysiological recordings within a generated magnetic field. We present calculations of the induced voltages in typical experimental situations and compare them against the neuronal signals measured with different electrophysiological techniques. Finally, we provide guidelines that should help limit and account for possible induction artefacts. In conclusion, if great care is taken, viable electrophysiological recordings from magnetoreceptive cells are achievable and promise to provide new insights on the neuronal basis of the magnetic sense.

The existence of a magnetic compass system allowing animals to maintain the direction of their migratory journeys was first shown in birds about fifty years ago. Over the past half century, a large amount of indirect, mainly behavioral, data on the features of the avian magnetic compass and its relationship with the perception of light has been obtained. Based on these data, the dominant idea suggests that the primary magnetoreceptor in birds is located in the retina, and the most popular hypothesis concerning operation of such a magnetoreceptor is the radical pair model. According to this model, the primary receptor molecules that perceive the magnetic field are cryptochromes expressed in the neurons of the avian retina. It is assumed that cryptochrome molecules localized in photoreceptor cells may have an ordered orientation relative to the retina plane. When photons are absorbed in the short wavelength range of the spectrum, cryptochromes would produce a chemical response depending on the direction of the external magnetic field. The sensitivity of photochemical reactions involving radical pairs in cryptochromes to a magnetic field is confirmed both by theoretical calculations and experimentally. However, up to date there is no experimental data obtained on the molecular mechanisms of further transduction of signal induced by a magnetic field. The existing hypotheses on the possible nature of these mechanisms have some issues concerning their compatibility with the current concepts of the photoreception. These hypotheses ascribe special properties to the participants of the visual process both at the molecular and at the cellular level, which has not yet been proved experimentally. Thus, nowadays the science of magnetoreception is posed to consistently test the existing hypotheses on the molecular basis of avian magnetic compass.

CRYPTOCHROMES (CRYs) are structurally related to ultraviolet (UV)/blue-sensitive DNA repair enzymes called photolyases but lack the ability to repair pyrimidine dimers generated by UV exposure. First identified in plants, CRYs have proven to be involved in light detection and various light-dependent processes in a broad range of organisms. In Drosophila, CRY’s best understood role is the cell-autonomous synchronization of circadian clocks. However, CRY also contributes to the amplitude of circadian oscillations in a light-independent manner, controls arousal and UV avoidance, influences visual photoreception, and plays a key role in magnetic field detection. Here, we review our current understanding of the mechanisms underlying CRY’s various circadian and noncircadian functions in fruit flies.

Significance Seasonal migration is dependent on an organism being able to sense and reorient to the Earth’s magnetic field. Cryptochromes (CRYs) have been implicated as light-driven sensors of the Earth’s magnetic field; however, contradictions between in vitro photochemistry and in vivo behavioral studies limit validation of CRYs as magnetosensors. To reconcile these discrepancies, we conducted detailed photochemical and structural studies of a CRY from Columbia livia (pigeon). We present the structure of a photoactive vertebrate CRY that reveals a pathway conserved in migratory organisms that facilitates its function as a magnetosensor., Computational and biochemical studies implicate the blue-light sensor cryptochrome (CRY) as an endogenous light-dependent magnetosensor enabling migratory birds to navigate using the Earth’s magnetic field. Validation of such a mechanism has been hampered by the absence of structures of vertebrate CRYs that have functional photochemistry. Here we present crystal structures of Columba livia (pigeon) CRY4 that reveal evolutionarily conserved modifications to a sequence of Trp residues (Trp-triad) required for CRY photoreduction. In ClCRY4, the Trp-triad chain is extended to include a fourth Trp (W369) and a Tyr (Y319) residue at the protein surface that imparts an unusually high quantum yield of photoreduction. These results are consistent with observations of night migratory behavior in animals at low light levels and could have implications for photochemical pathways allowing magnetosensing.

The most well-recognized magnetoreception behaviour is that of the magnetotactic bacteria (MTB), which synthesize membrane-bounded magnetic nanocrystals called magnetosomes via a biologically controlled process. The magnetic minerals identified in prokaryotic magnetosomes are magnetite (Fe3 O4 ) and greigite (Fe3 S4 ). Magnetosome crystals, regardless of composition, have consistent, species-specific morphologies and single-domain size range. Because of these features, magnetosome magnetite crystals possess specific properties in comparison to abiotic, chemically synthesized magnetite. Despite numerous discoveries regarding MTB phylogeny over the last decades, this diversity is still considered underestimated. Characterization of magnetotactic microorganisms is important as it might provide insights into the origin and establishment of magnetoreception in general, including eukaryotes. Here, we describe the magnetotactic behaviour and characterize the magnetosomes from a flagellated protist using culture-independent methods. Results strongly suggest that, unlike previously described magnetotactic protists, this flagellate is capable of biomineralizing its own anisotropic magnetite magnetosomes, which are aligned in complex aggregations of multiple chains within the cell. This organism has a similar response to magnetic field inversions as MTB. Therefore, this eukaryotic species might represent an early origin of magnetoreception based on magnetite biomineralization. It should add to the definition of parameters and criteria to classify biogenic magnetite in the fossil record.

To navigate within the geomagnetic field, magnetotactic bacteria synthesize magnetosomes, which are unique organelles consisting of membrane-enveloped magnetite nanocrystals. In magnetotactic spirilla, magnetosomes become actively organized into chains by the filament-forming actin-like MamK and the adaptor protein MamJ, thereby assembling a magnetic dipole much like a compass needle. However, in Magnetospirillum gryphiswaldense, discontinuous chains are still formed in the absence of MamK. Moreover, these fragmented chains persist in a straight conformation indicating undiscovered structural determinants able to accommodate a bar magnet-like magnetoreceptor in a helical bacterium. Here, we identify MamY, a membrane-bound protein that generates a sophisticated mechanical scaffold for magnetosomes. MamY localizes linearly along the positive inner cell curvature (the geodetic cell axis), probably by self-interaction and curvature sensing. In a mamY deletion mutant, magnetosome chains detach from the geodetic axis and fail to accommodate a straight conformation coinciding with reduced cellular magnetic orientation. Codeletion of mamKY completely abolishes chain formation, whereas on synthetic tethering of magnetosomes to MamY, the chain configuration is regained, emphasizing the structural properties of the protein. Our results suggest MamY is membrane-anchored mechanical scaffold that is essential to align the motility axis of magnetotactic spirilla with their magnetic moment vector and to perfectly reconcile magnetoreception with swimming direction.

A broad range of organisms, from prokaryotes to higher animals, have the ability to sense and utilize Earth's geomagnetic field—a behavior known as magnetoreception. Although our knowledge of the physiological mechanisms of magnetoreception has increased substantially over recent decades, the origin of this behavior remains a fundamental question in evolutionary biology. Despite this, there is growing evidence that magnetic iron mineral biosynthesis by prokaryotes may represent the earliest form of biogenic magnetic sensors on Earth. Here, we integrate new data from microbiology, geology and nanotechnology, and propose that initial biomineralization of intracellular iron nanoparticles in early life evolved as a mechanism for mitigating the toxicity of reactive oxygen species (ROS), as ultraviolet radiation and free-iron-generated ROS would have been a major environmental challenge for life on early Earth. This iron-based system could have later been co-opted as a magnetic sensor for magnetoreception in microorganisms, suggesting an origin of microbial magnetoreception as the result of the evolutionary process of exaptation.

Drosophila’s circadian clock can be perturbed by magnetic fields, as well as by lithium administration. Cryptochromes are critical for the circadian clock. Further, the radical pairs in cryptochrome also can explain magnetoreception in animals. Based on a simple radical pair mechanism model of the animal magnetic compass, we show that both magnetic fields and lithium can influence the spin dynamics of the naturally occurring radical pairs and hence modulate the circadian clock’s rhythms. Using a simple chemical oscillator model for the circadian clock, we show that the spin dynamics influence a rate in the chemical oscillator model, which translates into a change in the circadian period. Our model can reproduce the results of two independent experiments, magnetic fields and lithium effects on the circadian clock. Our model predicts that stronger magnetic fields would shorten the clock’s period. We also predict that lithium influences the clock in an isotope-dependent manner. Furthermore, our model also predicts that magnetic fields and hyperfine interactions modulate oxidative stress. The findings of this work suggest that quantum nature and entanglement of radical pairs might play roles in the brain, as another piece of evidence in addition to recent results on xenon anesthesia and lithium effects on hyperactivity.

Cryptochromes (CRYs) function as blue light photoreceptors in diverse physiological processes in nearly all kingdoms of life. Over the past several decades they have emerged as the most likely candidates for light-dependent magnetoreception in animals, however a long history of conflicts between in vitro photochemistry and in vivo behavioral data complicate validation of CRYs as a magnetosensor. In this review, we highlight the origins of conflicts regarding CRY photochemistry and signal transduction, and identify recent data that provides clarity on potential mechanisms of signal transduction in magnetoreception. The review primarily focuses on examining differences in photochemistry and signal transduction in plant and animal CRYs and identifies potential modes of convergent evolution within these independent lineages that may identify conserved signaling pathways. This article is protected by copyright. All rights reserved.

Sensitivity to magnetic fields is dependent on the intensity and color of light in several animal species. The light-dependent magnetoreception working model points to cryptochrome (Cry) as a protein cooperating with its co-factor flavin, which possibly becomes magnetically susceptible upon excitation by light. The type of Cry involved and what pair of magnetosensitive radicals are responsible is still elusive. Therefore, we developed a conditioning assay for the firebug Pyrrhocoris apterus, an insect species that possesses only the mammalian cryptochrome (Cry II). Here, using the engineered Cry II null mutant, we show that: (i) vertebrate-like Cry II is an essential component of the magnetoreception response, and (ii) magnetic conditioning continues even after 25 h in darkness. The light-dependent and dark-persisting magnetoreception based on Cry II may inspire new perspectives in magnetoreception and cryptochrome research.

Cryptochromes (Cry) have been proposed as putative magnetoreceptors in vertebrates. Localisation of Cry1 in the UV cones in the retinas of birds suggested that it could be the candidate magnetoreceptor. However, recent findings argue against this possibility. Cry1 is a type II cryptochrome, a subtype of cryptochromes that may not be inherently photosensitive, and it exhibits a clear circadian expression in the retinas of birds. Here, we reassessed the localization and distribution of Cry1 in the retina of the zebra finch. Zebra finches have a light-dependent magnetic compass based on a radical-pair mechanism, similar to migratory birds. We found that Cry1 colocalized with the UV/V opsin (SWS1) in the outer segments of UV cones, but restricted to the tip of the segments. Cry1 was expressed in all UV cones across the entire retina, with the highest densities near the fovea. Pre-exposure of birds to different wavelengths of light did not result in any difference in Cry1 expression, suggesting that Cry1 did not undergo any detectable functional changes as result of light activation. Considering that Cry1 is likely not involved in magnetoreception, our findings open the possibility for an involvement in different, yet undetermined functions in the avian UV/V cones.

Many animals display sensitivity to external magnetic field, but it is only in the simplest organisms that the sensing mechanism is understood. Here we report on behavioural experiments where American cockroaches (Periplaneta americana) were subjected to periodically rotated external magnetic fields with a period of 10 min. The insects show increased activity when placed in a periodically rotated Earth-strength field, whereas this effect is diminished in a twelve times stronger periodically rotated field. We analyse established models of magnetoreception, the magnetite model and the radical pair model, in light of this adaptation result. A broad class of magnetite models, based on single-domain particles found in insects and assumption that better alignment of magnetic grains towards the external field yields better sensing and higher insect activity, is shown to be excluded by the measured data. The radical-pair model explains the data if we assume that contrast in the chemical yield on the order of one in a thousand is perceivable by the animal, and that there also exists a threshold value for detection, attained in an Earth-strength field but not in the stronger field. Ministry of Education (MOE) Published version This work is supported by the Singapore Ministry of Education Tier 1 Grant No. RG 127/14 and the Polish National Agency for Academic Exchange NAWA Project No. PPN/PPO/2018/1/00007/U/00001.

International audience; Magnetotactic bacteria (MTB) are a group of prokaryotes having the ability to orient and swim along geomagnetic field lines because they contain intracellular magnetosomes, which are synthesized through a biomineralization process. Magnetosomes have recently also been found in unicellular eukaryotes, which are referred to as magnetically responsive protists (MRPs). The magnetosomes have three origins in MRPs. In this study we characterized a MTB-grazing ciliated MRP that was magnetically collected from intertidal sediment of Huiquan Bay, China. Based on 18S rRNA gene sequence analysis, the ciliated MRP was tentatively identified as Uronemella parafilificum HQ. Using transmission electron microscopy, we observed that magnetosomes having 2-3 shapes were randomly distributed within this ciliate. Energy-dispersive X-ray spectroscopy and high-resolution transmission electron microscopy images of the magnetosomes were consistent with them being composed of magnetite. Magnetosomes having the same shape and mineral composition were also detected in MTB that occurred in the same environment as the ciliated MRP. Statistical analysis showed that the size and shape of the magnetosomes in the ciliated MRP were similar to those in MTB. The results suggest that this ciliated MRP can graze, ingest, and digest various types of MTB. It is certainly worth noting that this is the first record of MRPs in Asian aquatic sediment and suggesting they might be widely distributed. These results also support the assertion that MRPs probably contribute to the ecological cycles of iron, and expand possibilities for research into the mechanism of magnetoreception in eukaryotes.

Ambient levels of nonionizing electromagnetic fields (EMF) have risen sharply in the last five decades to become a ubiquitous, continuous, biologically active environmental pollutant, even in rural and remote areas. Many species of flora and fauna, because of unique physiologies and habitats, are sensitive to exogenous EMF in ways that surpass human reactivity. This can lead to complex endogenous reactions that are highly variable, largely unseen, and a possible contributing factor in species extinctions, sometimes localized. Non-human magnetoreception mechanisms are explored. Numerous studies across all frequencies and taxa indicate that current low-level anthropogenic EMF can have myriad adverse and synergistic effects, including on orientation and migration, food finding, reproduction, mating, nest and den building, territorial maintenance and defense, and on vitality, longevity and survivorship itself. Effects have been observed in mammals such as bats, cervids, cetaceans, and pinnipeds among others, and on birds, insects, amphibians, reptiles, microbes and many species of flora. Cyto- and geno-toxic effects have long been observed in laboratory research on animal models that can be extrapolated to wildlife. Unusual multi-system mechanisms can come into play with non-human species — including in aquatic environments — that rely on the Earth’s natural geomagnetic fields for critical life-sustaining information. Part 2 of this 3-part series includes four online supplement tables of effects seen in animals from both ELF and RFR at vanishingly low intensities. Taken as a whole, this indicates enough information to raise concerns about ambient exposures to nonionizing radiation at ecosystem levels. Wildlife loss is often unseen and undocumented until tipping points are reached. It is time to recognize ambient EMF as a novel form of pollution and develop rules at regulatory agencies that designate air as ‘habitat’ so EMF can be regulated like other pollutants. Long-term chronic low-level EMF exposure standards, which do not now exist, should be set accordingly for wildlife, and environmental laws should be strictly enforced — a subject explored in Part 3.

It is known that bipolar disorder and its lithium treatment involve the modulation of oxidative stress. Moreover, it has been observed that lithium’s effects are isotope-dependent. Based on these findings, here we propose that lithium exerts its effects by influencing the recombination dynamics of a naturally occurring radical pair involving oxygen. We develop a simple model inspired by the radical-pair mechanism in cryptochrome in the context of avian magnetoreception and xenon-induced anesthesia. Our model reproduces the observed isotopic dependence in the lithium treatment of hyperactivity in rats. It predicts a magnetic-field dependence of the effectiveness of lithium, which provides one potential experimental test of our hypothesis. Our findings show that Nature might harness quantum entanglement for the brain’s cognitive processes.

Understanding the mechanisms underlying general anesthesia would be a key step towards understanding consciousness. The process of xenon-induced general anesthesia has been shown to involve electron transfer, and the potency of xenon as a general anesthetic exhibits isotopic dependence. We propose that these observations can be explained by a mechanism in which the xenon nuclear spin influences the recombination dynamics of a naturally occurring radical pair of electrons. We develop a simple model inspired by the body of work on the radical-pair mechanism in cryptochrome in the context of avian magnetoreception, and we show that our model can reproduce the observed isotopic dependence of the general anesthetic potency of xenon in mice. Our results are consistent with the idea that radical pairs of electrons with entangled spins could be important for consciousness., Comment: 17 pages, 10 figures

The intrinsic earth magnetic field (geomagnetic field, GMF) provides an essential environmental condition for most living organisms to adapt the solar cycle by rhythmically synchronizing physiological and behavioral processes. However, hypomagnetic field (HMF) of outer space, the Moon, and the Mars differs much from GMF, which poses a critical problem to astronauts during long-term interplanetary missions. Multiple experimental works have been devoted to the HMF effects on circadian rhythm and found that HMF perturbs circadian rhythms and profoundly contributes to health problems such as sleep disorders, altered metabolic as well as neurological diseases. By systemizing the latest progress on interdisciplinary cooperation between magnetobiology and chronobiology, this review sheds light on the health effects of HMF on circadian rhythms by elaborating the underlying circadian clock machinery and molecular processes.

Summary Displacement experiments have demonstrated that experienced migratory birds translocated thousands of kilometers away from their migratory corridor can orient toward and ultimately reach their intended destinations. 1 This implies that they are capable of “true navigation,” commonly defined 2 , 3 , 4 as the ability to return to a known destination after displacement to an unknown location without relying on familiar surroundings, cues that emanate from the destination, or information collected during the outward journey. 5 , 6 , 7 , 8 , 9 , 10 , 11 , 12 , 13 In birds, true navigation appears to require previous migratory experience 5 , 6 , 7 , 14 , 15 (but see Kishkinev et al. 16 and Piersma et al. 17 ). It is generally assumed that, to correct for displacements outside the familiar area, birds initially gather information within their year-round distribution range, learn predictable spatial gradients of environmental cues within it, and extrapolate from those to unfamiliar magnitudes—the gradient hypothesis. 6 , 9 , 18 , 19 , 20 , 21 , 22 However, the nature of the cues and evidence for actual extrapolation remain elusive. Geomagnetic cues (inclination, declination, and total intensity) provide predictable spatial gradients across large parts of the globe and could serve for navigation. We tested the orientation of long-distance migrants, Eurasian reed warblers, exposing them to geomagnetic cues of unfamiliar magnitude encountered beyond their natural distribution range. The birds demonstrated re-orientation toward their migratory corridor as if they were translocated to the corresponding location but only when all naturally occurring magnetic cues were presented, not when declination was changed alone. This result represents direct evidence for migratory birds’ ability to navigate using geomagnetic cues extrapolated beyond their previous experience.

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.

The exact anatomical location for an iron particle-based magnetic sense remains enigmatic in vertebrates. For mammals, findings from a cornea anaesthesia experiment in mole rats suggest that it carries the primary sensors for magnetoreception. Yet, this has never been tested in a free-ranging mammal. Here, we investigated whether intact corneal sensation is crucial for navigation in migrating Nathusius’ bats, Pipistrellus nathusii, translocated from their migratory corridor. We found that bats treated with corneal anaesthesia in both eyes flew in random directions after translocation and release, contrasting bats with a single eye treated, and the control group, which both oriented in the seasonally appropriate direction. Using a Y-maze test, we confirmed that light detection remained unaffected by topical anaesthesia. Therefore our results suggest the cornea as a possible site of magnetoreception in bats, although other conceivable effects of the anaesthetic are also explored. Furthermore, we demonstrate that the corneal based sense is of bilateral nature but can function in a single eye if necessary., Lindecke and colleagues experimentally investigate the effect of corneal anesthesia on bat migratory orientation. Their results suggest that the cornea is a promising organ for the search of biological magnetic compass sensors, and also indicate that a single functional cornea is sufficient for migratory navigation.

How living systems respond to weak electromagnetic fields represents one of the major unsolved challenges in sensory biology. Recent evidence has implicated cryptochrome, an evolutionarily conserved flavoprotein receptor, in magnetic field responses of organisms ranging from plants to migratory birds. However, whether cryptochromes fulfill the criteria to function as biological magnetosensors remains to be established. Currently, theoretical predictions on the underlying mechanism of chemical magnetoreception have been supported by experimental observations that exposure to radiofrequency (RF) in the MHz range disrupt bird orientation and mammalian cellular respiration. Here we show that, in keeping with certain quantum physical hypotheses, a weak 7 MHz radiofrequency magnetic field significantly reduces the biological responsivity to blue light of the cryptochrome receptor cry1 in Arabidopsis seedlings. Using an in vivo phosphorylation assay that specifically detects activated cryptochrome, we demonstrate that RF exposure reduces conformational changes associated with biological activity. RF exposure furthermore alters cryptochrome-dependent plant growth responses and gene expression to a degree consistent with theoretical predictions. To our knowledge this represents the first demonstration of a biological receptor responding to RF exposure, providing important new implications for magnetosensing as well as possible future applications in biotechnology and medicine.

Magnetoreception is the ability to sense the Earth's magnetic field, which is used for orientation and navigation. Behavioural experiments have shown that it is employed by many species across all vertebrate classes; however, our understanding of how magnetic information is processed and integrated within the central nervous system is limited. In this Commentary, we review the progress in birds and rodents, highlighting the role of the vestibular and trigeminal systems as well as that of the hippocampus. We reflect on the strengths and weaknesses of the methodologies currently at our disposal, the utility of emerging technologies and identify questions that we feel are critical for the advancement of the field. We expect that magnetic circuits are likely to share anatomical motifs with other senses, which culminates in the formation of spatial maps in telencephalic areas of the brain. Specifically, we predict the existence of spatial cells that encode defined components of the Earth's magnetic field.

BackgroundThe discovery of membrane-enclosed, metabolically functional organelles inBacteriahas transformed our understanding of the subcellular complexity of prokaryotic cells. Biomineralization of magnetic nanoparticles within magnetosomes by magnetotactic bacteria (MTB) is a fascinating example of prokaryotic organelles. Magnetosomes, as nano-sized magnetic sensors in MTB, facilitate cell navigation along the local geomagnetic field, a behaviour referred to as magnetotaxis or microbial magnetoreception. Recent discovery of novel MTB outside the traditionally recognized taxonomic lineages suggests that MTB diversity across the domainBacteriaare considerably underestimated, which limits understanding of the taxonomic distribution and evolutionary origin of magnetosome organelle biogenesis.ResultsHere, we perform the most comprehensive metagenomic analysis available of MTB communities and reconstruct metagenome-assembled MTB genomes from diverse ecosystems. Discovery of MTB in acidic peatland soils suggests widespread MTB occurrence in waterlogged soils in addition to subaqueous sediments and water bodies. A total of 168 MTB draft genomes have been reconstructed, which represent nearly a 3-fold increase over the number currently available and more than double the known MTB species at the genome level. Phylogenomic analysis reveals that these genomes belong to 13 Bacterial phyla, six of which were previously not known to include MTB. These findings indicate a much wider taxonomic distribution of magnetosome organelle biogenesis across the domainBacteriathan previously thought. Comparative genome analysis reveals a vast diversity of magnetosome gene clusters involved in magnetosomal biogenesis in terms of gene content and synteny residing in distinct taxonomic lineages. Phylogenetic analyses of core magnetosome proteins in this largest available and taxonomically diverse dataset support an unexpectedly early evolutionary origin of magnetosome biomineralization, likely ancestral to the origin of the domainBacteria.ConclusionsThese findings expand the taxonomic and phylogenetic diversity of MTB across the domainBacteriaand shed new light on the origin and evolution of microbial magnetoreception. Potential biogenesis of the magnetosome organelle in the close descendants of the last bacterial common ancestor has important implications for our understanding of the evolutionary history of bacterial cellular complexity and emphasizes the biological significance of the magnetosome organelle.

Several groups of mammals use the Earth's magnetic field for orientation, but their magnetosensory organ remains unknown. The Ansell's mole-rat (Fukomys anselli, Bathyergidae, Rodentia) is a microphthalmic subterranean rodent with innate magnetic orientation behaviour. Previous studies on this species proposed that its magnetoreceptors are located in the eye. To test this hypothesis, we assessed magnetic orientation in mole-rats after the surgical removal of their eyes compared to untreated controls. Initially, we demonstrate that this enucleation does not lead to changes in routine behaviours, including locomotion, feeding and socializing. We then studied magnetic compass orientation by employing a well-established nest-building assay under four magnetic field alignments. In line with previous studies, control animals exhibited a significant preference to build nests in magnetic southeast. By contrast, enucleated mole-rats built nests in random magnetic orientations, suggesting an impairment of their magnetic sense. The results provide robust support for the hypothesis that mole-rats perceive magnetic fields with their minute eyes, probably relying on magnetite-based receptors in the cornea.

Avian magnetoreception is assumed to occur in the retina. Although its molecular mechanism is unclear, magnetic field-dependent formation and the stability of radical-containing photointermediate(s) are suggested to play key roles in a hypothesis called the radical pair mechanism. Chicken cryptochrome4 (cCRY4) has been identified as a candidate magnetoreceptive molecule due to its expression in the retina and its ability to form stable flavin neutral radicals (FADH●) upon blue light absorption. Herein, we used millisecond flash photolysis to investigate the cCRY4 photocycle, in both the presence and absence of dithiothreitol (DTT); detecting the anion radical form of FAD (FAD●-) under both conditions. Using spectral data obtained during flash photolysis and UV-visible photospectroscopy, we estimated the absolute absorbance spectra of the photointermediates, thus allowing us to decompose each spectrum into its individual components. Notably, in the absence of DTT, approximately 37% and 63% of FAD●- was oxidized to FADOX and protonated to form FADH●, respectively. Singular value decomposition analysis suggested the presence of two FAD●- molecular species, each of which was destined to be oxidized to FADOX or protonated to FADH●. A tyrosine neutral radical was also detected; however, it likely decayed concomitantly with the oxidation of FAD●-. On the basis of these results, we considered the occurrence of bifurcation prior to FAD●- generation, or during FAD●- oxidization, and discussed the potential role played by the tyrosine radical in the radical pair mechanism.

Light or low frequency magnetic field (LF-MF) as one of the cultivation environments affects secondary metabolites (SMs) production of M. purpureus. Phytochrome (Phy) is a hybrid histidine kinase possessing dual properties of photoreceptor and kinase to sense red and far-red light. The interaction effects of LF-MF and light on SMs of M. purpureus was investigated by knocking out the Phy-like gene in M. purpureus (MpPhy) by homologous recombination. A MpPhy-deletion (ΔMpPhy) strain produced less Monascus pigments (MPs) and monacolin K (mon K) than the wild-type (WT) strain and reduced citrinin production by 78.3% on 10th day but didn't affect the biomass. These results indicated that the MpPhy gene is involved in SMs biosynthesis of M. purpureus. MPs production in WT was decreased significantly when the inoculum was exposed to white/blue/green/red light (500 Lux). But it in ΔMpPhy was no significant difference when exposed to white/red light. The colony size of ΔMpPhy was smaller on potato dextrose agar media containing 0.01% SDS. These results indicated that the deletion of MpPhy gene affected the aerial hyphae and increased sensitivity to cell membrane stress but decreased sensitivity to red light. The inoculum of both WT and ΔMpPhy was exposure to the LF-MF (50 Hz). The accumulation of WT secondary metabolites was not changed, while SMs production of ΔMpPhy was significantly enhanced under exposed to 2.0 mT LF-MF. This indicated that the decrease of SMs caused by the deletion of MpPhy gene was restored by LF-MF. It revealed that there is a crosstalk between magnetoreception and photosensitivity.

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.

Bacteria developed many different ways to orient themselves in the environment. Magnetoreception with following motility along Earth's magnetic field lines and photoreception with subsequent positive or negative phototaxis allow bacteria to optimally position themselves for survival and growth. Some bacteria show both magnetotactic and photoresponsive behaviour and additionally live in a multicellular organism adding another layer of complexity. A novel study by Qian and colleagues visualized different species of multicellular magnetotactic bacteria and shed light on their reproductive as well as photoresponsive behaviour. This study paves the way towards understanding the evolutionary advantage of multicellular lifestyle of prokaryotes.

It is known that animals are sensitive to the geomagnetic field. In the case of insects, magnetoreception has been reported in several ant species and in some bees and wasps. One study showed that the stingless bee Tetragonisca angustula is able to sense the modification of the magnetic field inclination. The aim of the present manuscript is to continue that study in T. angustula, analyzing the nest arrival and departure angles in the presence of magnetic fields generated by magnets. The bees flying to and from the nest were recorded and the flying trajectories were obtained by analyzing the video frame by frame. The magnetic field was generated by 6, 9, or 12 magnets contained inside an Eppendorf tube and fixed near the nest. Our results show that T. angustula bees are sensitive to magnetic fields because the departure angles are influenced by the magnets. It was observed that these bees are sensitive to the polarization of the magnetic field vector that influences the choice of flying up or down, and this sensitivity has a window until about 80 μT (about four times the local geomagnetic field), with the magnetic field information for higher magnetic field intensities being ignored by the bees. Bioelectromagnetics. 2021;42:51-59. © 2020 Bioelectromagnetics Society.

Despite anecdotal reports of the astonishing homing abilities in dogs, their homing strategies are not fully understood. We equipped 27 hunting dogs with GPS collars and action cams, let them freely roam in forested areas, and analyzed components of homing in over 600 trials. When returning to the owner (homewards), dogs either followed their outbound track ('tracking') or used a novel route ('scouting'). The inbound track during scouting started mostly with a short (about 20 m) run along the north-south geomagnetic axis, irrespective of the actual direction homewards. Performing such a 'compass run' significantly increased homing efficiency. We propose that this run is instrumental for bringing the mental map into register with the magnetic compass and to establish the heading of the animal. European Social FundEuropean Social Fund (ESF) [EVA 4.0, CZ.02.1.01/0.0/0.0/16_019/0000803]; Ministry of Agriculture of the Czech RepublicMinistry of Agriculture, Czech Republic [MZE-RO0718]; Czech University of Life Sciences Prague CIGA CZU [20174319]; Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague IGA [B07/16]; Grant Agency of the Czech RepublicGrant Agency of the Czech Republic [15-21840S]; European Regional Development FundEuropean Union (EU) [EVA 4.0, CZ.02.1.01/0.0/0.0/16_019/0000803] European Social Fund Operational Programme Research, Development and Education, EVA 4.0 Hynek Burda; European Social Fund Operational Programme Research, Development and Education, CZ.02.1.01/0.0/0.0/16_019/0000803 Hynek Burda; Ministry of Agriculture of the Czech Republic MZE-RO0718 Ludek Bartos; Czech University of Life Sciences Prague CIGA CZU (Project No. 20174319) Hynek Burda; Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague IGA (Project No. B07/16) Hynek Burda; Grant Agency of the Czech Republic No. 15-21840S Hynek Burda; European Regional Development Fund Operational Programme Research, Development and Education, EVA 4.0 Hynek Burda; European Regional Development Fund Operational Programme Research, Development and Education, CZ.02.1.01/0.0/0.0/16_019/0000803 Hynek Burda; The funders had no role in study design, data collection and interpretation, or the decision to submit the work for publication.

How geomagnetic field information is collected and processed by insects for orientation and navigation remains elusive. In social insects, magnetic particles are well accepted as magnetic sensors. Ants have the ability to home and hunt, and some migratory and nomadic species can migrate or move over long distances for which magnetoreception is an important mechanism. It was shown previously that ferromagnetic resonance (FMR) spectral parameters of one migratory and one nomadic ant could be distinguished from Brazilian Solenopsis ant species and that these parameters correlate to the local geomagnetic field. The present work focuses on genera engaged in long-distance group raids and emigration collected mainly in the Amazon rainforest. A diversity of specimens of the genus was individually measured by FMR. Cluster analysis of the occurrence of the FMR Low Field component, associated with large or aggregated nanoparticles, and their spectral angular dependence resulted in a phylogenetic dendrogram of the genera of ants, principally from the North Brazilian region. The magnetic material characteristics of ants of the Invertebrate Collection from the Museum Paraense Emilio Goeldi were tested looking for their relation to ant genera and the local geomagnetic field. The observed spectral differences of the magnetic particles suggest that they are related to their capacity for adaptation to their environment and/or to displacement behaviour.

Magnetic-field reception of animals and plants is currently discussed in the framework of a cryptochrome-based radical-pair mechanism. Efforts to unravel magnetoreception in plants suffered historically from several shortcomings, most prominently, the conspicuous absence of detailed stimulus-response relationships. To determine the sensitivity of seedlings of Arabidopsis thaliana to weak static magnetic fields we generated stimulus-response curves between near zero and 188 μT for the transcript levels of the genes rbcl, cab4, pal4 and ef1. The moderate magneto-responsiveness of dark-grown seedlings was greatly enhanced under blue light, and for rbcl and pal4 also under red light. The stimulus-response curves obtained under blue light of constant photon-fluence rate displayed multiple maxima and thus a pattern fundamentally different from that prevalent in plant and animal physiology. A double mutant lacking cryptochromes 1 and 2 displayed altered stimulus-response curves without losing, however, magneto-responsiveness completely. A reversal of the magnetic field direction substantially affected the gene expression and the quantity of CAB-protein (chlorophyll a,b-binding protein). The majority of our results are at variance with the notion of cryptochromes acting as the only magnetic-field sensors. They do not, however, exclude the possibility that cryptochromes participate in the magnetic field reception of Arabidopsis. The findings have the unexpected implication that cryptochrome- and phytochrome-mediated plant responses can be modulated by the strength and the orientation of the local geomagnetic field.

Cryptochrome (CRYs) proteins have been elucidated as the molecular basis for magnetoreception in Drosophila, and a putative magnetic receptor (named IscA1) protein may aslo be involved in sensing magnetic fields in Drosophila. However, whether IscA1 has a conserved role in diverse animals and functions in orientation during animal migration is unknown. Here we report on the cloning and sequencing of the IscA1 gene from Agrotis segetum, which encodes a predicted protein IscA1 that has 131 amino acids and two conserved iron-sulphur cluster binding domains. Multiple sequence alignment and phylogenetic analysis were used to show that IscA1 had a relatively high homology from species of Noctuoidea. Quantitative polymerase chain reaction showed that IscA1 was ubiquitously expressed in adult organs and, among all developmental stages, expression was higher in adults. When Agrotis segetum was exposed to 14 h light/10 h dark, IscA1 expression also showed daily oscillations, and constant light or dark disturbed these oscillations. IscA1 expression levels in a migratory population were higher than in a reared population and higher in a southward migratory population than in a northward. These findings suggest that the IscA1 gene in A. segetum might be associated with migration and provide a molecular basis for further study on the functions of IscA1 gene in magnetoreception and potential control of the turnip moth.

Two types of vertebrate cryptochromes (Crys) are currently recognized. Type 2 Crys function in the molecular circadian clock as light-independent transcriptional repressors. Type 4 Crys are a newly discovered group with unknown function, although they are flavoproteins and therefore may function as photoreceptors. It has been postulated that Crys function in light-dependent magnetoreception, which is thought to contribute towards homing and migratory behaviors. Here we have cloned and annotated the full-length pigeon ClCry1, ClCry2, and ClCry4 genes, and characterized the full-length proteins and several site-directed mutants to investigate the roles of these proteins. ClCry1 and ClCry2 are phylogenetically grouped as Type 2 Crys and thus are expected to be core components of the pigeon circadian clock. Interestingly, we find that ClCry4 is properly annotated as a Type 4 Cry. It appears that many birds possess a Type 4 Cry which, as in pigeon, is misannotated. Like the Type 2 Crys, ClCry4 is widespread in pigeon tissues. However, unlike the Type 2 Crys, ClCry4 is cytosolic, and purified ClCry4 possesses FAD cofactor, which confers characteristic UV-visible spectra as well as two photochemical activities. We find that ClCry4 undergoes light-dependent conformational change, which is a property of insect Type1 Crys involved in the insect-specific pathway of photoentrainment of the biological clock. ClCry4 can also be photochemically reduced by a mechanism common to all FAD-containing Cry family members, and this mechanism is postulated to be influenced by the geomagnetic field. Thus pigeon Crys control circadian behavior and may also have photosensory function.

The geomagnetic field (GMF) is an environmental element whose instability affects plant growth and development. Despite known plant responses to GMF direction and intensity, the mechanism of magnetoreception in plants is still not known. Magnetic field variations affect many light-dependent plant processes, suggesting that the magnetoreception could require light. The objective of this work was to comprehensively investigate the influence of GMF on Arabidopsis thaliana (Col-0) photoreceptor signaling. Wild-type Arabidopsis seedlings and photoreceptor-deficient mutants (cry1cry2, phot1, phyA and phyAphyB) were exposed to near null magnetic field (NNMF, ≤40 nT) and GMF (~43 μT) under darkness and different light wavelengths. The GMF did not alter skotomorphogenic or photomorphogenic seedling development but had a significant impact on gene expression pathways downstream of cryptochrome and phytochrome photoactivation. GMF-induced changes in gene expression observed under blue light were partially associated with an alteration of cryptochrome activation. GMF impacts on phytochrome-regulated gene expression could be attributed to alterations in phytochrome protein abundance that were also dependent on the presence of cry1, cry2 and phot1. Moreover, the GMF was found to impact photomorphogenic-promoting gene expression in etiolated seedlings, indicating the existence of a light-independent magnetoreception mechanism. In conclusion, our data shows that magnetoreception alters photoreceptor signaling in Arabidopsis, but it does not necessarily depend on light.

Magnetosensitivity is widespread among animals with rodents being the most intensively studied mammalian group. The available behavioural assays for magnetoreception are time-consuming, which impedes screens for treatment effects that could characterize the enigmatic magnetoreceptors. Here, we present a fast and simple approach to test if an animal responds to magnetic stimuli: the magnetic object assay (MOA). The MOA focuses on investigating an animal’s spontaneous exploration behaviour in the presence of a bar magnet compared to a demagnetised control. We present consistently longer exploration of the magnet in three different rodent species: Ansell’s mole-rat (Fukomys anselli), C57BL/6J laboratory mouse, and naked mole-rat (Heterocephalus glaber). For the naked mole-rat this is the first report that this species reacts on magnetic stimuli. We conclude that the MOA holds the potential to screen if an animal responds to magnetic stimuli, indicating the possession of a magnetic sense.

For centuries, humans have been fascinated by how migratory animals find their way over thousands of kilometres. Here, I review the mechanisms used in animal orientation and navigation with a particular focus on long-distance migrants and magnetoreception. I contend that any long-distance navigational task consists of three phases and that no single cue or mechanism will enable animals to navigate with pinpoint accuracy over thousands of kilometres. Multiscale and multisensory cue integration in the brain is needed. I conclude by raising twenty important mechanistic questions related to long-distance animal navigation that should be solved over the next twenty years.