The amphibian magnetic sense(s).

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

Evidence for the use of a high-resolution magnetic map by a short-distance migrant, the Alpine newt (Ichthyosaura alpestris).

Newts can use spatial variation in the magnetic field (MF) to derive geographic position, but it is unclear how they detect the 'spatial signal', which, over the distances that newts move in a day, is an order of magnitude lower than temporal variation in the MF. Previous work has shown that newts take map readings using their light-dependent magnetic compass to align a magnetite-based 'map detector' relative to the MF. In this study, time of day, location and light exposure (required by the magnetic compass) were varied to determine when newts obtain map information. Newts were displaced from breeding ponds without access to route-based cues to sites where they were held and/or tested under diffuse natural illumination. We found that: (1) newts held overnight at the testing site exhibited accurate homing orientation, but not if transported to the testing site on the day of testing; (2) newts held overnight under diffuse lighting at a 'false testing site' and then tested at a site located in a different direction from their home pond oriented in the home direction from the holding site, not from the site where they were tested; and (3) newts held overnight in total darkness (except for light exposure for specific periods) only exhibited homing orientation the following day if exposed to diffuse illumination during the preceding evening twilight in the ambient MF. These findings demonstrate that, to determine the home direction, newts require access to light and the ambient MF during evening twilight when temporal variation in the MF is minimal.

Spontaneous magnetic alignment behaviour in free-living lizards.

Several species of vertebrates exhibit spontaneous longitudinal body axis alignment relative to the Earth's magnetic field (i.e., magnetic alignment) while they are performing different behavioural tasks. Since magnetoreception is still not fully understood, studying magnetic alignment provides evidence for magnetoreception and broadens current knowledge of magnetic sense in animals. Furthermore, magnetic alignment widens the roles of magnetic sensitivity in animals and may contribute to shed new light on magnetoreception. In this context, spontaneous alignment in two species of lacertid lizards (Podarcis muralis and Podarcis lilfordi) during basking periods was monitored. Alignments in 255 P. muralis and 456 P. lilfordi were measured over a 5-year period. The possible influence of the sun's position (i.e., altitude and azimuth) and geomagnetic field values corresponding to the moment in which a particular lizard was observed on lizards' body axis orientation was evaluated. Both species exhibited a highly significant bimodal orientation along the north-northeast and south-southwest magnetic axis. The evidence from this study suggests that free-living lacertid lizards exhibit magnetic alignment behaviour, since their body alignments cannot be explained by an effect of the sun's position. On the contrary, lizard orientations were significantly correlated with geomagnetic field values at the time of each observation. We suggest that this behaviour might provide lizards with a constant directional reference while they are sun basking. This directional reference might improve their mental map of space to accomplish efficient escape behaviour. This study is the first to provide spontaneous magnetic alignment behaviour in free-living reptiles.

Evidence of light-dependent magnetic compass orientation in urodele amphibian larvae.

Experiments were conducted to investigate whether larval palmate newts undertake orientation toward or away from the home shoreline (y-axis orientation) using the geomagnetic field to steer the most direct route, and if they accomplish this task through a light-dependent magnetoreception mechanism similar to that found in anuran tadpoles and adult newts. Larval palmate newts trained and then tested under full-spectrum light showed bimodal magnetic compass orientation that coincided with the magnetic direction of the trained y-axis. In contrast, larvae trained under long-wavelength (≥500nm) light and then tested under full-spectrum light displayed bimodal orientation perpendicular to the trained y-axis direction. These results offer evidence for the use of magnetic compass cues in orienting urodele amphibian larvae, and provide additional support for the light-dependent magnetoreception mechanism since they are in complete agreement with earlier studies showing that the observed 90° shift in the direction of magnetic compass orientation under long-wavelength light (≥500nm) is due to a direct effect of light on the underlying magnetoreception mechanism. This study is the first to provide evidence of a light-dependent magnetic compass in larval urodeles.

Use of a light-dependent magnetic compass for y-axis orientation in European common frog (Rana temporaria) tadpoles.

We provide evidence for the use of a magnetic compass for y-axis orientation (i.e., orientation along the shore-deep water axis) by tadpoles of the European common frog (Rana temporaria). Furthermore, our study provides evidence for a wavelength-dependent effect of light on magnetic compass orientation in amphibians. Tadpoles trained and then tested under full-spectrum light displayed magnetic compass orientation that coincided with the trained shore-deep water axes of their training tanks. Conversely, tadpoles trained under long-wavelength (≥500 nm) light and tested under full-spectrum light, and tadpoles trained under full-spectrum light and tested under long-wavelength (≥500 nm) light, exhibited a 90° shift in magnetic compass orientation relative to the trained y-axis direction. Our results are consistent with earlier studies showing that the observed 90° shift in the direction of magnetic compass orientation under long-wavelength (≥500 nm) light is due to a direct effect of light on the underlying magnetoreception mechanism. These findings also show that wavelength-dependent effects of light do not compromise the function of the magnetic compass under a wide range of natural lighting conditions, presumably due to a large asymmetry in the relatively sensitivity of antagonistic short- and long-wavelength inputs to the light-dependent magnetic compass.