Sensory Ecology of Electromagnetic Radiation Perception
in Subterranean Mole-Rats
(Fukomys anselli & Fukomys kafuensis)
Inaugural-Dissertation
zur Erlangung des Doktorgrades Dr. rer. nat.
des Fachbereiches Biologie und Geographie an der Universität Duisburg-Essen
Vorgelegt von
Regina E. Moritz
aus Bottrop
Januar 2007
Die der vorliegenden Arbeit zugrunde liegenden Experimente wurden in der Abteilung Allgemeine Zoologie der Universität Duisburg-Essen, Campus Essen, in der Abteilung Physiologie und Ökologie des Verhaltens und in der Dr. Senckenbergischen Anatomie der Johann Wolfgang Goethe-Universität, Frankfurt/Main durchgeführt.
1. GUTACHTER: Prof. Dr. Hynek Burda
2. GUTACHTER: Prof. Dr. Leo Peichl
3. GUTACHTER: Prof. Dr. Bernd Sures
VORSITZENDER DES PRÜFUNGSAUSSCHUSSES: Prof. Dr. Martin Heil
Tag der mündlichen Prüfung : 07. Mai 2007
Non quia difficilia sunt audemus, sed quia non audemus difficilia sunt.
Lucius Annaeus Seneca
in memoriam
Dr. Mathias Kawalika (1962-2006)
LIST OF CONTENTS
I SUMMARY — 1
II ZUSAMMENFASSUNG — 2
III GENERAL INTRODUCTION — 3 III.1 Subterranean Fukomys Mole-Rats — 3 III.2 Orientation in the Subterranean Habitat — 5 III.3 Sensory Adaptations in Fukomys — 6 III.4 Electromagnetic Radiation — 6 III.5 Contribution of this thesis to Fukomys sensory research — 8
A LIGHT PERCEPTION — 9
1 INTRODUCTION — 9 1.1 Visual Capabilities in Fukomys — 10 1.2 Arising questions — 12
2 MATERIAL AND METHODS — 13 2.1 Study Animals — 13 2.2 Demonstrating Light Perception — 14 2.2.1 Study rationale — 14 2.2.2 Study procedure — 14 2.2.2.1 Halogen light — 14 2.2.2.2 Natural daylight — 16 2.2.2.3 Retinal involvement — 16 2.3 The Light Perception Threshold — 17 2.3.1 Study rationale — 17 2.3.2 Study procedure — 17 2.4 Light spectrum in a tunnel — 20 2.4.1 Study rationale — 20 2.4.2 Study procedure — 20 2.5 Statistical analysis — 22
3 RESULTS — 23 3.1 Demonstrating Light Perception — 23 3.2 The Light Perception Threshold — 25 3.3 Light spectrum in a tunnel — 28
4 DISCUSSION — 33 B MAGNETORECEPTION — 39
1 INTRODUCTION — 39 1.1 Magnetoreception in Animals — 39 1.2 The Earth’s Magnetic Field — 41 1.3 Using the Earth’s Magnetic Field — 43 1.4 From Earth to Animal: Available Sensory Information — 44 1.5 From Behavioural Experiment to Proof: Compass Modes — 45 1.6 From Signal to Sensor: Transduction Mechanisms — 47 1.6.1 Magnetoperception via Biochemical Processes — 48 1.6.2 Magnetoreception via Magnetite — 50 1.7 From Sensor to Brain: Neuronal Processing — 57 1.7.1 Immunocytochemical methods — 58 1.8 Arising questions — 60
2 MATERIAL AND METHODS — 61 2.1 Study Animals — 60 2.2 Ruling out Biochemical Processes — 60 2.2.1 Study rationale — 60 2.2.2 Study procedure — 62 2.3 Narrowing down the Receptor Site — 64 2.3.1 Study rationale — 65 2.3.2 Study procedure — 67 2.4 Magnetic Orientation is Binocular — 70 2.4.1. Study rationale — 70 2.4.2. Study procedure — 71 2.5 Revealing Hippocampal Involvement — 71 2.5.1 Study rationale — 71 2.5.2. Study procedure — 71 2.6 Statistical analysis — 75
3 RESULTS — 76 3.1 Ruling out Biochemical Processes — 76 3.2 Narrowing down the Receptor Site — 78 3.3 Magnetic Orientation is Binocular — 83 3.4 Revealing Hippocampal Involvement — 87
4 DISCUSSION — 91
IV RÉSUMÉ & OUTLOOK — 99
V REFERENCES — 102
VI APPENDIX — 122 A Abbreviations — 122 B Figure legends — 124 C Table legends — 126 D Wavelength spectra — 127 E The rat brain hippocampus — 132 F ICC protocol — 133 G Glass slide gelatine cover recipe — 140 H Nissl-staining recipe — 140 I Technorama Forum Lecture: 2000 years of magnetism — 141 J Acknowledgements — 145 K Curriculum Vitae — 146 L List of Publications — 148
Summary 1
I SUMMARY