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Parasites Within Parasites: Transmission and Evolution of Neorickettsia in Digeneans Stephen Edward Greiman

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Parasites Within Parasites: Transmission and Evolution of Neorickettsia in Digeneans Stephen Edward Greiman University of North Dakota UND Scholarly Commons Theses and Dissertations Theses, Dissertations, and Senior Projects January 2015 Parasites Within Parasites: Transmission And Evolution Of Neorickettsia In Digeneans Stephen Edward Greiman Follow this and additional works at: https://commons.und.edu/theses Recommended Citation Greiman, Stephen Edward, "Parasites Within Parasites: Transmission And Evolution Of Neorickettsia In Digeneans" (2015). Theses and Dissertations. 1777. https://commons.und.edu/theses/1777 This Dissertation is brought to you for free and open access by the Theses, Dissertations, and Senior Projects at UND Scholarly Commons. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of UND Scholarly Commons. For more information, please contact [email protected]. PARASITES WITHIN PARASITES: TRANSMISSION AND EVOLUTION OF NEORICKETTSIA IN DIGENEANS by Stephen Edward Greiman Bachelor of Science, University of North Dakota, 2011 A Dissertation Submitted to the Graduate Faculty of the University of North Dakota in partial fulfillment of the requirements for the degree of Doctor of Philosophy Grand Forks, North Dakota May 2015 Copyright 2015 Stephen Greiman ii PERMISSION Title PARASITES WITHIN PARASITES: TRANSMISSION AND EVOLUTION OF NEORICKETTSIA IN DIGENEANS Department Biology Degree Doctor of Philosophy In presenting this dissertation in partial fulfillment of the requirements for a graduate degree from the University of North Dakota, I agree that the library of this University shall make it freely available for inspection. I further agree that permission for extensive copying for scholarly purposes may be granted by the professor who supervised my dissertation work or, in his absence, by the chairperson of the department or the dean of the School of Graduate Studies. It is understood that any copying or publication or other use of this dissertation or part thereof for financial gain shall not be allowed without my written permission. It is also understood that due recognition shall be given to me and to the University of North Dakota in any scholarly use which may be made of any material in my dissertation. Stephen Edward Greiman April-24-2015 iv TABLE OF CONTENTS LIST OF FIGURES .................................................................................................... ix LIST OF TABLES .................................................................................................... xiv ACKNOWLEDGMENTS ........................................................................................ xvi ABSTRACT ............................................................................................................. xvii CHAPTER I. LITERATURE REVIEW ....................................................................... 1 Introduction .................................................................................... 1 Taxonomy and Phylogeny of Neorickettsia ................................... 3 Systematic Position of Neorickettsia ......................................... 3 Taxonomy and Phylogenetic Interrelationships Among Species of Neorickettsia ........................................................... 5 Ecology and Transmission of Neorickettsiae ................................ 6 Neorickettsia helminthoeca and Salmon Poisoning Disease ......... 9 Elokomin Fluke Fever Agent ...................................................... 14 Neorickettsia sennetsu and Sennetsu Fever ................................. 16 Neorickettsia risticii and Potomac Horse Fever .......................... 21 SF Agent ...................................................................................... 27 Rainbow Trout Agent .................................................................. 28 Neorickettsia sp. From Needlefish in Cambodia ......................... 28 Neorickettsia sp. From Antarctic Notothenioid Fishes ................ 29 v Catfish Agents ............................................................................. 30 Geographic Distribution of Neorickettsia .................................... 31 Phylogenetic Associations Between Neorickettsia and Digenea ........................................................................................ 35 Study Objectives and Gaps in Current Knowledge ..................... 37 II. LABORATORY LIFE CYCLE AND MATERIALS AND METHODS ........................................................................................... 41 Introduction .................................................................................. 41 Materials and Methods ................................................................. 43 General Materials and Methods .............................................. 43 Laboratory Life Cycle of a Neorickettsia Infected Digenean, Plagiorchis elegans ............................................... 47 Chapter 3 Specific Materials and Methods .............................. 51 Chapter 4 Specific Materials and Methods .............................. 56 Chapter 5 Specific Materials and Methods .............................. 58 Chapter 6 Specific Materials and Methods .............................. 66 Results .......................................................................................... 68 Pond Screening ..................................................................... 68 Extraction Method ................................................................. 71 Laboratory Life Cycle ............................................................ 71 Discussion .................................................................................... 74 Fluctuation in Positive Ponds ................................................. 74 Benefits of Sonication Extraction Method .......................... 74 Benefits of Indirect Sonication ............................................... 74 Benefits of Real Time PCR .................................................... 75 Quantitative Real Time PCR .................................................. 76 Laboratory Life Cycle ............................................................ 77 Conclusion ................................................................................... 80 vi III. LARGE SCALE SCREENING OF DIGENEANS FOR NEORICKETTSIA ENDOSYMBIONTS USING REAL-TIME PCR REVEALS NEW NEORICKETTSIA GENOTYPES, HOST ASSOCIATIONS AND GEOGRAPHIC RECORDS .......................... 81 Abstract ........................................................................................ 81 Results .......................................................................................... 82 Discussion .................................................................................... 90 IV. TRANSMISSION RATES OF THE BACTERIAL ENDOSYMBIONT, NEORICKETTSIA SP., DURING THE ASEXUAL REPRODUCTION PHASE OF ITS DIGENEAN HOST PLAGIORCHIS ELEGANS, WITHIN NATURALLY INFECTED LYMNAEID SNAILS .................................................... 106 Abstract ...................................................................................... 106 Results ........................................................................................ 107 Discussion .................................................................................. 109 Conclusions ................................................................................ 111 V. QUANTIFICATION OF THE BACTERIAL ENDOSYMBIONT, NEORICKETTSIA SP., WITHIN ALL LIFE CYCLE STAGES OF THE DIGENEAN HOST BY USE OF REAL-TIME qPCR ANALYSIS TARGETING THE GroEL GENE ................................ 113 Abstract ...................................................................................... 113 Results ........................................................................................ 114 Neorickettsia sp. qPCR Assays ............................................. 114 Quantification of Neorickettsia sp. in Cercariae ..................... 114 Quantification of Neorickettsia sp. in Metacercariae (24 Hour/pre-infective) ........................................................ 115 Quantification of Neorickettsia sp. in Metacercariae (6 Day/infective) ................................................................. 116 Quantification of Neorickettsia sp. in Juvenile Worms (48 Hour) ............................................................................ 118 Quantification of Neorickettsia sp. in Adult Worms (14 Day) ............................................................................. 118 Quantification of Neorickettsia sp. Within All Life Cycle Stages for Parasite Group 1 .................................................. 123 vii Quantification of Neorickettsia sp. Within All Life Cycle Stages for Parasite Group 2 .................................................. 123 Quantification of Neorickettsia sp. Within All Life Cycle Stages for Parasite Group 3 .................................................. 124 Quantification of Neorickettsia sp. in Metacercariae (Encysted Within Snail 2) .................................................... 125 Discussion .................................................................................. 126 VI. GERMS WITHIN WORMS: LOCALIZATION OF THE BACTERIAL ENDOSYMBIONT, NEORICKETTSIA SP., WITHIN DIFFERENT LIFE CYCLE STAGES OF THE DIGENEAN, PLAGIORCHIS ELEGANS .......................................... 136 Abstract ...................................................................................... 136 Results .......................................................................................
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