Non-Visual Cues – Magnetic Field The Earth’s geomagnetic varies in 3 ways that can convey information on position and orientation: 1) Polarity 2) Intensity 3) Inclination (dip) Angle Non-Visual Cues – Magnetic Field Non-Visual Cues – Magnetic Field Inclination angle varies regularly with latitude. 1 Non-Visual Cues – Magnetic Field Field intensity varies with latitude. Non-Visual Cues – Magnetic Field Declination is the difference (in degrees) between a geographic and magnetic north. Non-Visual Cues – Magnetic Field Magnetic senses and/or magnetic orientation have been demonstrated in: Magnetotactic bacteria – Magnetospirillum sp. Magnetite crystals 2 Non-Visual Cues – Magnetic Field Magnetic senses and/or magnetic orientation have been demonstrated in: Magnetotactic bacteria – Magnetospirillum sp. Non-Visual Cues – Magnetic Field Magnetic senses and/or magnetic orientation have been demonstrated in: Invertebrates – flatworms, annelids, molluscs, arthropods. Vertebrates – fish, amphibians, reptiles birds, mammals (including humans). Non-Visual Cues – Magnetic Field The inclination compass. 3 Non-Visual Cues – Magnetic Field Exposure to the horizontal field at the equator causes a reversal of preferred inclination angle of migrant birds. Non-Visual Cues – Magnetic Field Exposure to the horizontal field at the equator causes a reversal of preferred inclination angle of migrant birds. Non-Visual Cues – Magnetic Field Polarity compasses are uncommon, but have been documented in some species. e.g. spiny lobsters. 4 Non-Visual Cues – Magnetic Field In a polarity compass reversal of the vertical field component has no effect. Lohmann et al. 1995. JEB 198:2041-2048. Non-Visual Cues – Magnetic Field Field intensity can indicate latitude and alter behaviour (e.g. fattening in a transSaharan migrant bird) Non-Visual Cues – Magnetic Field Local hills or valleys of field intensity are associated with whale strandings. Pilot whale 5 Magnetic Senses There are three proposed mechanisms for magnetic sensing. 1) Electroinduction 2) Magnetite based 3) Photoreceptor based Electroinduction Elasmobranchs that can sense electric fields may be able to use the same ampullae of Lorenzini to sense magnetic fields. Magnetite Based Magnetoreception Magnetite (Fe3O4) has been detected in a wide variety of organisms. Single domain (SD) magnetite crystals are 50 – 70 nm in size, and can hold a permanent magnetic moment. Super paramagnetic (SP) magnetite particles are < 50 nm and can only have a magnetic moment in the presence of an external magnetic field. 6 Magnetite Based Magnetoreception Magnetotactic bacteria have chains of SD magnetite. Magnetite Based Magnetoreception Studies of rainbow trout (Onchorhyrhus mykiss) provide the first evidence of a magnetoreceptor. Walker et al. 1997. Nature 390:371-376. Magnetite Based Magnetoreception Receptor cells are localized in the lamina propria, deep to the epithelium. Olfactory lamellae Walker et al. 1997. Nature 390:371-376. 7 Magnetite Based Magnetoreception The receptor cell is multilobed and has a 1 um chain of SD magnetite particles. It communicates to the brain via the trigeminal nerve. Diebel et al. 2000. Nature 406:299-302. Magnetite Based Magnetoreception Magnetic force microscopy demonstrates the response of the magnetic sensor to magnetic field alterations. Diebel et al. 2000. Nature 406:299-302. Magnetite Based Magnetoreception SP magnetite receptors have recently been described under the skin of the upper margins of the pigeon beak. Fleissner et al. 2003. J. Comp. Neur. 458:350-360. 8 Magnetite Based Magnetoreception Fleissner et al. 2003. J. Comp. Neur. 458:350-360. Magnetite Based Magnetoreception The receptor has 1 nm SP magnetite spherules located close to the membrane, and associated iron phosphate platelets that may help amplify the magnetic field variations. Fleissner et al. 2003. J. Comp. Neur. 458:350-360. Magnetite Based Magnetoreception The SP magnetite is localized in a fibrous cup that is associated with the membrane and may trigger depolarization by a mechanical mechanism. Fleissner et al. 2003. J. Comp. Neur. 458:350-360. 9 Photoreceptor Based Magnetoreception There is ample evidence from a variety of species (newts, flies, birds) that magnetic orientation can be impaired in the dark or by long wavelength light. Deutschlander et al. 1999. JEB 202:891-908. Photoreceptor Based Magnetoreception The proposed mechanism involves excitation of a photopigment that forms radical pairs of electrons that respond to magnetic fields. Inclination compass only. Localized to eyes and/or pineal (amphibians). The leading candidate is cryptochome protein which contains FADH. Photoreceptor Based Magnetoreception It is hypothesized that two photoreceptor systems that are sensitive to either short or long wavelength light are responsible. One is excitatory and the other is inhibitory. Deutschlander et al. 1999. JEB 202:891-908. 10 Photoreceptor Based Magnetoreception Under intermediate wavelength they cancel out. Under full spectrum light one system dominates and the animal orients to the axis of the field that gives the strongest response. Deutschlander et al. 1999. JEB 202:891-908. Measuring the Magnetic Field Birds use head scans to measure the strength and direction of the magnetic field. Mouritsen et al. 2004. Curr. Biol. 14:1946-1949. Coordination of Orientation Marine turtle hatchlings use sky brightness, beach slope, wave direction, and magnetic field inclination and intensity. Lohmann and Lohmann 1998. JAB 29:585-596. 11 Coordination of Orientation Magnetic field inclination angle is used to keep turtles in the Atlantic gyre. Lohmann and Lohmann 1998. JAB 29:585-596. Coordination of Orientation Magnetic field intensity is also an important Cue that is used to keep turtles in the Atlantic gyre. Lohmann and Lohmann 1998. JAB 29:585-596. Coordination of Orientation Turtles and other pelagic species could Potentially use a bicoordinate system of inclinations and intensities to navigate. Equal Inclination Angle Equal Intensity Lohmann and Lohmann 1998. JAB 29:585-596. 12 Coordination of Orientation Prior to migration birds use celestial cues to calibrate the magnetic compass. In juveniles the stellar rotation sense is paramount and without magnetic field birds orient directly away from the rotation axis. Magnetic information is corrected for declination to produce correct population specific orientation. Coordination of Orientation During the migration season it’s controversial! Some evidence suggests that the magnetic compass is paramount and is used to recalibrate celestial compasses (e.g. Wiltschko et al 1998. JAB 29:606-617). Recent reanalysis of data indicates that if birds can see the entire sunset to the horizon, that the magnetic compass is recalibrated using skylight polarization (e.g. Muheim et al 2006. JEB 209:2-17). Coordination of Orientation Nightly recalibration of the magnetic compass using twilight cues is supported by recent empirical evidence. Swainson’s thrush Cochran et al. 2004. Science 304:405-408. 13 Navigation How animals convert orientation information into navigation is poorly understood. Modifying triggers Vector navigation – combining orientation preferences with a time program. Navigation Natural selection should favour the shortest path. Orthodromes or great circle routes are shortest but require constant course adjustment if there is any east-west component. Loxodromes or rhumblines have a constant compass bearing but are longer. 14 1 2 3 4 5