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Magnetoreception - Introduction magnetic field as putative orientation factors (Middendorf 1859 1 , Viguier 1882), it was not until the 1960s that magnetic compass orientation could be demonstrated in a bird (Wiltschko & Merkel 1966). The first mammal shown to have a magnetic compass was, as mentioned above (III.3), the Zambian Ansell’s mole-rat (Fukomys anselli) (Burda et al. 1990b). This rodent species still plays the lead when it comes to a refined and continuously expanded knowledge of the magnetic compass sense in rodents. As in the monotonous world underground, the Earth’s magnetic field provides a reliable source of directional, and perhaps also positional, information, this compass mechanism is not a surprising finding in a subterranean mammal, as the Earth derived information helps an animal, in absence of other stimuli, to navigate effectively, i.e. to determine where it is, where it wants to go and how to get there. Magnetotaxis and alignment behaviour exist in diverse taxa from bacteria to honeybees, and navigation based on magnetic information occurs, amongst others, in lobsters and molluscs (Kirschvink & Gould 1981; Cain et al. 2005). In terrestrial vertebrates, magnetoreception has been unambiguously demonstrated in sharks, salmonids, newts, turtles and migratory and homing birds (reviewed in Wiltschko & Wiltschko 1995; Lohmann & Johnsen 2000; Meyer et al. 2005). Recently, the bat could be added to the list of mammals that use magnetoreception, showing to use a sunset-calibrated magnetic compass for long-distance navigation (Holland et al. 2006). Furthermore, several rodent species have been proposed to use the magnetic field for compass orientation or responses to gradients or to temporal variations (Wiltschko & Wiltschko 1995), but a magnetic compass sense has been demonstrated unambiguously in only two subterraneous species: the African Ansell’s mole-rat F. anselli (Burda et al. 1990b, Marhold et al. 1997a,b) and the blind mole-rat (Spalax ehrenbergi superspecies) from Israel (Burda et al. 1991; Kimchi & Terkel 1999, 2001; Kimchi et al. 2004). Magnetoreception has also been demonstrated in two epigeic species: the Djungarian hamster (Phodopus sungorus) (Deutschlander et al. 2003) and the laboratory mouse (Muheim et al. 2006). Earlier studies of magnetic orientation in rodents provided ambiguous and in some cases questionable results (Mather & Baker 1981; Madden & Phillips 1987; Sauvé 1988; August et al. 1989). 1 »[...], so liegt der Gedanke nahe, es möge die erstaunliche Unbeirrbarkeit der Zugvögel – trotz Wind und Wetter, trotz Nacht und Nebel – eben darauf beruhen, dass das Geflügel immerwährend der Richtung des Magnetpoles sich bewusst ist, und demzufolge auch seine Zugrichtung genau einzuhalten weiss. […] Gleich dem Schiffer, der seinen Kurs in die Karte einträgt so oft er die Rumbe seiner Richtschnur, der Magnetnadel, wechselt, ist auch der Vogel unablässig sich dessen bewusst, wann und wie viel er abweicht […]. Während aber der Schiffer, bei der Eintragung seiner Kurse, noch die jedesmalige Deklinationsgrösse der Magnetnadel von den Meridianen seiner Seekarten in Abrechnung zu bringen hat, liest sich der Vogel die Grösse des Abweichungswinkels u n m i t t e l b a r ab […].« (Middendorf, 1859) 40
Magnetoreception - Introduction 1.2 The Earth’s Magnetic Field At the end of the 16th century, William Gilbert determined that the Earth is a big magnet, implying that it has a magnetic field (cf., Lanza & Meloni 2006). The Earth’s magnetic field (fig. B1) is a dipole field, i.e. a system comprising two magnetic charges (or masses) of equal intensity and opposite signs (cf., Skiles 1985; Lanza & Meloni 2006). These magnetic field’s measurements are based on superimposed contributions from different sources. Corresponding to these different origins, the Earth’s magnetic field of force can be separated into: the main field, generated in the fluid core by a geodynamo mechanism; the crustal field, generated by magnetized rocks in the Earth’s crust; the external field, generated by electric currents flowing in both ionosphere and magnetosphere through interactions of the solar electromagnetic radiation and the solar wind with the Earth’s magnetic field; and the magnetic field resulting from an electromagnetic induction process generated by electric currents induced in the crust and the upper mantle by external magnetic field time variations – the Earth is partly an electric conductor, and currents can be induced in its conducting parts by external time variations. The most stable parts of the Earth’s magnetic field are the main and the crustal field, which mainly determine its spatial structure. The Fig. B1 Schematical view of the Earth’s magnetic field lines. The Earth is represented by a sphere, N and S are the ideal magnetic pole positions (from Lanza & Meloni 2006). field is also subject to time variations. These can be divided into long-term variation due to changes within the Earth, and short-term variation of external origin (see external field above) (cf., Lanza & Meloni 2006). The Earth’s magnetic field behaves like a small, very strong bar magnet close to the Earth’s centre that is tilted against the rotation axis by about 11° (cf., Press & Siever 2003). Determining exact geomagnetic coordinates, i.e. identifying exact positions of points on the Earth’s surface with respect to a geomagnetic reference (similar to geographic coordinates), is enabled by use of colatitudes and longitudes in the geomagnetic dipole frame. This coordinate system also enables the identification of north and south geomagnetic poles as those points on the Earth where the ideal central dipole axis intersects the surface. Accordingly, the geomagnetic equator is the ideal line on the surface representing the intersection of the plane passing through the Earth’s centre orthogonal to the central dipole (Lanza & Meloni 2006). 41
Page 1 and 2: Sensory Ecology of Electromagnetic
Page 3 and 4: Non quia difficilia sunt audemus, s
Page 5 and 6: L I S T O F C O N T E N T S I S U M
Page 7: IV R É S U M É & O U T L O O K
Page 10 and 11: Zusammenfassung II ZUSAMMENFASSUNG
Page 12 and 13: General Introduction Fukomys mole-r
Page 14 and 15: General Introduction III.3 Sensory
Page 16 and 17: General Introduction electromagneti
Page 18 and 19: Light Perception - Introduction pro
Page 20 and 21: Light Perception - Introduction The
Page 22 and 23: Light Perception - Material & Metho
Page 32 and 33: Light Perception - Results Number o
Page 34 and 35: Light Perception - Results Table A1
Page 36 and 37: Light Perception - Results 3.3 Ligh
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Magnetoreception - Results Table B1
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Magnetoreception - Discussion compa
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Magnetoreception - Discussion Magne
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Magnetoreception - Discussion by th
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Magnetoreception - Discussion invol
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Magnetoreception - Résumé & Outlo
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References V REFERENCES Arendt D, T
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References Brett RA (1991) The ecol
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References David-Gray ZK, Bellingha
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References Heth G, Nevo E & Beiles
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References Kirschvink JL & Gould JL
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References Mattis DC (1965) The the
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References Oelschläger HA & Northc
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References Rado R, Terkel J & Wollb
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References Sharp PE, Blair HT & Cho
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References Williams MN & Wild JM (2
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Appendix VI APPENDIX A Abbreviation
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Figure legend B Figure legends Fig.
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Table legend C Table legends Table
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Wavelength spectra Fig. D3 Waveleng
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Wavelength spectra Fig. D7 Spectral
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The rat brain hippocampus E The rat
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ICC protocol B) GELATINE EMBEDDING
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ICC protocol iii) Adsorption contro
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ICC protocol G) DAB PREPARATION i)
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Gelatine coating and Nissl-staining
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Technorama Forum Lecture: 2000 year
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Technorama Forum Lecture: 2000 year
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Curriculum Vitae K Curriculum Vitae
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List of Publications L List of Publ
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