Understanding Functioning and Evolution of Bird Middle Ear Mechanics: a Functional Morphological Analysis
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Faculteit Wetenschappen Departement Biologie Understanding functioning and evolution of bird middle ear mechanics: a functional morphological analysis Inzicht in functie en evolutie van de mechanica van het middenoor van vogels: een functioneel-morfologische analyse Proefschrift voorgelegd tot het behalen van de graad van Doctor in de Wetenschappen: Biologie aan de Universiteit Antwerpen te verdedigen door Raf CLAES Promotoren Prof. Dr. Peter Aerts Prof. Dr. Joris J.J. Dirckx Antwerpen 2018 The candidate was financially supported by research grant G049414N to Peter Aerts, Joris Dirkcx and Steve Vanlanduit of the Research Foundation of Flanders (FWO) De kandidaat werd financieel ondersteund met de onderzoeksbeurs G049414N aan Peter Aerts, Joris Dirckx en Steve Vanlanduit door het Fonds voor Wetenschappelijk Onderzoek – Vlaanderen (FWO) Doctoral committee Chair Prof. Dr. Raoul Van Damme: Department of Biology, University of Antwerp, Belgium Supervisors Prof. Dr. Peter Aerts: Department of Biology, University of Antwerp, Belgium; Department of Movement and Sport Science, Ghent University, Belgium Prof. Dr. Joris J.J. Dirckx: Department of Physics, University of Antwerp, Belgium Members of the jury Prof. Dr. Stefan Van Dongen: Department of Biology, University of Antwerp, Belgium Prof. Dr. Dominique Adriaens: Department of Biology, Ghent University, Belgium Prof. Dr. Anthony Herrel: Directeur de Recherche CNRS, Muséum National d’Histoire Naturelle, France Prof. Dr. Matthew J. Mason: Department of Physiology, Development & Neuroscience, University of Cambridge, United Kingdom Private PhD defense Friday, 23 November 2018, 13h30 Meeting room, D.134 Campus Drie Eiken, University of Antwerp, Belgium Public PhD defense Thursday, 20 December 2018, 17h00 Auditorium O.06 Campus Drie Eiken, University of Antwerp, Belgium 5 Contents Doctoral committee .......................................................................................................... 5 Summary ......................................................................................................................... 11 Nederlandstalige samenvatting ...................................................................................... 15 General introduction ....................................................................................................... 19 The biological role of sound ........................................................................................ 21 Mechanics of sound .................................................................................................... 22 Sound perception ........................................................................................................ 24 Middle ear evolution ................................................................................................... 28 Rhipidistia ................................................................................................................ 29 Tetrapods ................................................................................................................ 30 Amphibian lineage ................................................................................................... 31 Amniote lineage ...................................................................................................... 32 The avian ear morphology and function ..................................................................... 39 The outer ear ........................................................................................................... 40 The middle ear ......................................................................................................... 41 Middle ear function ................................................................................................. 48 The inner ear ........................................................................................................... 51 Research objectives ..................................................................................................... 53 PART I .............................................................................................................................. 57 CHAPTER 1: The influence of middle ear morphology on the hearing capabilities in birds ......................................................................................................................................... 59 Abstract ....................................................................................................................... 60 Introduction ................................................................................................................. 60 Materials and methods ............................................................................................... 65 Micro-CT scanning ................................................................................................... 65 Data analysis and morphological measurements ................................................... 67 Statistical analysis .................................................................................................... 69 7 Contents Audiogram ...............................................................................................................70 Results .........................................................................................................................72 Middle ear morphology ...........................................................................................72 Principal component analysis ..................................................................................78 Intensity ...................................................................................................................79 Frequency ................................................................................................................81 Discussion ....................................................................................................................83 CHAPTER 2: The effect of craniokinesis on the middle ear of domestic chickens (Gallus gallus domesticus) ...........................................................................................................89 Abstract .......................................................................................................................90 Introduction .................................................................................................................91 Materials and methods ...............................................................................................94 Micro-CT scanning ...................................................................................................94 Segmentation, surface model..................................................................................95 Measurements .........................................................................................................95 Behavioral recordings ..............................................................................................97 Results .........................................................................................................................98 Beak opening ...........................................................................................................98 Quadrate ..................................................................................................................99 Middle ear .............................................................................................................100 Discussion ..................................................................................................................103 Conclusion .................................................................................................................108 CHAPTER 3: Do high sound pressure levels of crowing in roosters necessitate passive mechanisms for protection against self-vocalization? ..................................................111 Abstract .....................................................................................................................112 Introduction ...............................................................................................................112 Materials and methods .............................................................................................115 Sound recording ....................................................................................................115 Geometry of the external auditory canal ..............................................................117 8 Contents Results ....................................................................................................................... 118 Sound recording .................................................................................................... 118 Geometry of the external auditory canal .............................................................. 120 Discussion .................................................................................................................. 122 PART II ........................................................................................................................... 125 CHAPTER 4: Deformation of birds middle ear structures under static pressure loads, and potential regulation mechanisms .................................................................................. 127 Abstract ....................................................................................................................