Characterizing the Role of Magnetic Cues Underlying Spatial Behavior
In the 50+ years since the discovery of magnetic compass orientation by migratory songbirds, evidence for the use of magnetic cues has been obtained for a range of taxonomic groups, including several classes of vertebrate and invertebrate taxa. Surprisingly, however, the biophysical mechanisms and biological substrate that underlie magnetic sensing are still not fully understood. Moreover, while use of magnetic cues for compass orientation is intuitive, the functional significance of other forms of behavioral responses mediated by magnetic cues, such as spontaneous magnetic alignment, is less clear. The following research was carried out to investigate the mechanisms underlying magnetic orientation in vertebrates and invertebrates. This involved the modification of existing experimental systems to characterize responses to magnetic cues in laboratory animals (flies, mice) and the development of novel techniques for studying the role of magnetic cues in the spatial behavior of free-living animals (red foxes). Chapter II examines magnetic orientation in wild-type Drosophila melanogaster larvae. We show that three strains of larvae reared under non-directional ultraviolet (UV) light exhibit quadramodal spontaneous orientation along the anti-cardinal compass directions (i.e. northeast, southeast, southwest, northwest) when tested in a radially symmetrical environment under UV light. Double-blind experiments cancelling the horizontal component of the magnetic field confirmed that the response is dependent on magnetic cues rather non-magnetic features of the test environment. Furthermore, we argue that the larval quadramodal pattern of response is consistent with properties of magnetic compass orientation observed in previous studies of adult Drosophila and laboratory mice, both of which have been proposed to be mediated by a light-dependent magnetic compass mechanism. Chapter III explores the use of novel biologging techniques to collect behavioral and spatial data from free-roaming mammals. Specifically, a previous observational study of free- roaming red foxes found a 4-fold increase in the success of predatory 'mousing' attacks when foxes were facing ~north-northeast, consistent with magnetic alignment responses reported for a range of terrestrial animals. The authors propose that the magnetic field may be used to increase accuracy of mousing attacks. Using tri-axial accelerometer and magnetometer bio-loggers fitted to semi-domesticated red foxes, we created ']magnetic ethograms' from behavioral and magnetic machine learning algorithms 'trained'] to identify three discrete behaviors (i.e. foraging, trotting, and mousing-like jumps) from raw accelerometer signatures and to classify the magnetic headings of mousing-like jumps into 45° sectors from raw magnetometer data. The classifier's ability to accurately identify behaviors from a separate fox not used to train the algorithm suggests that these techniques can be used in future experiments to obtain reliable magnetic ethograms for free-roaming foxes. We also developed the first radio-frequency emitting collar that broadcasts in the low MHz frequency range shown to disrupt magnetic compass responses in a host of animals. The radio-frequency collars coupled with biologgers will provide a powerful tool to characterize magnetic alignment responses in predatory red foxes and can be adapted for use in studies of magnetic alignment and magnetic compass orientation in other free-roaming mammals. Chapter 3 discusses findings from a magnetic nest building assay involving male labratory mice. Mice trained to position nests in one of four directions relative to the magnetic field exhibited both learned magnetic compass responses and fixed magnetic nest positioning orientation consistent with northeast-southwest spontaneous magnetic alignment behavior previously reported for wild mice and bank voles. This is the first mammalian assay in which both learned magnetic compass orientation and spontaneous magnetic alignment were exhibited in the same species, and suggests that the use of magnetic cues in rodents may be more flexible that previously realized.