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Spironucleus Salmonis

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Spironucleus Salmonis Humboldt Universität zu Berlin Leibniz‐Institute für Landwirtsschaftlich‐ Gärtnerische Fakultät Gewässerökologie und Binnenfischerei Characterisation and host‐parasite interaction of the piscine diplomonad Spironucleus salmonis Mohammad Reza Saghari Fard July, 2008 Titel der Arbeit: Characterisation and host‐parasite interaction of the piscine diplomonad Spironucleus salmonis D i s s e r t a t i o n zur Erlangung des akademischen Gradedoctor rerum agriculturarum (Dr. rer. agr.) eingereicht an der Landwirtschaftlich‐Gärtnerischen Fakultät der Humboldt‐Universität zu Berlin von Mohammad Reza Saghari Fard 15 March 1971, Mashad, Iran Präsident der Humboldt‐Universität zu Berlin Prof. Dr. Dr. h.c. Christoph Markschies Dekan der Landwirtschaftlich‐Gärtnerischen Fakultät: Prof. Dr. Dr. h.c. Otto Kaufmann Gutachter: 1. Prof. Dr. Frank Kirschbaum 2. Dr. Sarah L. Poynton 3. Dr. Klaus Knopf Tag der Verteidigung: 21 November 2008 I would like to dedicate this thesis to my loving parents…. CONTENTS DECLARATION................................................................................................................................................... 8 ACKNOWLEDGMENTS .................................................................................................................................... 9 PAPERS............................................................................................................................................................... 11 CONTRIBUTIONS TO PAPERS ..................................................................................................................... 13 CONFERENCE PRESENTATIONS ................................................................................................................ 15 DISSERTATION OVERVIEW......................................................................................................................... 17 ABSTRACT......................................................................................................................................................... 19 CHAPTER 1 OVERVIEW OF DIPLOMONADS........................................................................................... 22 1.1 Taxonomy................................................................................................................................................... 22 1.2 Phylogeny ................................................................................................................................................... 24 1.3 Morphology and ultrastructure ................................................................................................................... 28 1.4 Habitat ........................................................................................................................................................ 31 1.5 Life cycle - reproduction............................................................................................................................. 33 1.6 Pathology.................................................................................................................................................... 34 1.7 Immunity .................................................................................................................................................... 36 1.8 Management and treatment of infection ..................................................................................................... 37 CHAPTER 2 PISCINE DIPLOMONAD: SPIRONUCLEUS ......................................................................... 40 2.1 Aquaculture and Spironucleus.................................................................................................................... 40 2.2 Identification of Spironucleus species in fish - chronology........................................................................ 41 2.3 Habitat of Spironucleus .............................................................................................................................. 46 2.4 Life cycle of Spironucleus – trophozoite and cyst...................................................................................... 48 2.5 Encystment of Spironucleus – the role of flagella ...................................................................................... 49 2.6 Pathology associated with Spironucleus..................................................................................................... 52 2.7 Innate immunity to Spironucleus................................................................................................................ 56 2.8 Acquired immunity..................................................................................................................................... 58 CHAPTER 3 SPECIFIC AIMS ......................................................................................................................... 61 3.1 Overview .................................................................................................................................................... 61 3.2 Characterisation of Spironucleus................................................................................................................ 62 3.3 Intestinal pH and the microhabitat preference of Spironucleus .................................................................. 63 3.4 Encystment of Spironucleus – the role of flagella ...................................................................................... 64 3.5 Pathology associated with Spironucleus..................................................................................................... 65 3.6 Innate immunity to Spironucleus – in vitro plasma incubation test............................................................ 67 4 CHAPTER 4 MATERIALS AND METHODS ................................................................................................ 70 4.1 Source of fish.............................................................................................................................................. 70 4.2 Wet preparations of diplomonads ............................................................................................................... 70 4.3 Characterisation of Spironucleus................................................................................................................ 71 4.3.1 Ultrastructural characterisation ......................................................................................................... 71 4.3.2 Molecular characterisation................................................................................................................. 73 4.4 Intestinal pH and the microhabitat preference of Spironucleus .................................................................. 74 4.4.1 Data collection .................................................................................................................................... 74 4.4.2 Statistical analysis............................................................................................................................... 75 4.5 Encystment – the role of Spironucleus flagella .......................................................................................... 76 4.5.1 In vitro culture..................................................................................................................................... 76 4.5.2 Monitoring and fixation....................................................................................................................... 77 4.5.3 Scanning electron microscopy............................................................................................................. 77 4.6 Pathology associated with Spironucleus..................................................................................................... 78 4.6.1 Overview ............................................................................................................................................. 78 4.6.2 Harvesting of tissues ........................................................................................................................... 78 4.6.3 Histology ............................................................................................................................................. 79 4.6.4 Pathology quantification ..................................................................................................................... 79 4.6.5 Statistical analysis............................................................................................................................... 85 4.7 Innate immunity to Spironucleus – in vitro plasma incubation test............................................................ 85 4.7.1 Experimental design............................................................................................................................ 85 4.7.2 Developing in vitro plasma incubation test – refining the protocol .................................................... 86 4.7.3 Performing in vitro plasma incubation test......................................................................................... 89 CHAPTER 5 RESULTS AND DISCUSSIONS................................................................................................ 91 5.1 Characterisation of Spironucleus salmonis................................................................................................. 91 5.1.1 Result: Surface morphology ...............................................................................................................
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