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The Role of Resistance to Varroa Destructor and Deformed Wing Virus in the European Honey Bee (Apis Mellifera)

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The Role of Resistance to Varroa Destructor and Deformed Wing Virus in the European Honey Bee (Apis Mellifera) THE ROLE OF RESISTANCE TO VARROA DESTRUCTOR AND DEFORMED WING VIRUS IN THE EUROPEAN HONEY BEE (APIS MELLIFERA) JESSICA LEONI KEVILL Ph.D. Thesis 2018 Table of Contents LIST OF FUGURES .................................................................................................................... ...8 General introduction .................................................................................................... ..8 Chapter 1: ABC Assay - Method development and application to quantify the role of three DWV master variants in overwinter colony losses of European honey bees ..... 8 Chapter 2: DWV master variant dominance and seasonal prevalence in honey bee colonies from England and Wales………... ................................................................. ...10 Chapter 3: DWV infection, mortality and survivorship in pre-wintering Varroa susceptible and tolerant honey bee colonies from the USA .................................. .…11 Chapter 4: The role of DWV avirulence and superinfection exclusion in wild colonies of honey bees located in the Arnot forest and Ithaca, New York ........................... ...13 Chapter 5: DWV recombinant prevalence in DWV positive colonies from England, Wales and the USA…..…………………………………………………………………………………………....14 LIST OF TABLES……………………………………………………………………………………………………………………16 Chapter 1: ABC Assay - Method development and application to quantify the role of three DWV master variants in overwinter colony losses of European honey bees…..16 Chapter 2: DWV master variant dominance and seasonal prevalence in honey bee colonies from England and Wales …………………………………………………………………………..16 Chapter 3: DWV infection, mortality and survivorship in pre-wintering Varroa susceptible and tolerant honey bee colonies from the USA…………………………………….16 Chapter 4: The role of DWV avirulence and superinfection exclusion in wild colonies of honey bees located in the Arnot forest and Ithaca, New York ……………………………17 Chapter 5: DWV recombinant prevalence in DWV positive colonies from England, Wales and the USA………………………………………………………………………………………………….17 ACKNOWLEDGEMENTS……………………………………………………………………………………………………….18 DECLARATION…………………………………………………………………………………………………………………….19 1 Chapter 1: ABC Assay - Method development and application to quantify the role of three DWV master variants in overwinter colony losses of European honey bees…..19 Chapter 2: DWV master variant dominance and seasonal prevalence in honey bee colonies from England and Wales …………………………………………………………………………..20 Chapter 3: DWV infection, mortality and survivorship in pre-wintering Varroa susceptible and tolerant honey bee colonies from the USA………………………………..…..20 Chapter 4: The role of DWV avirulence and superinfection exclusion in wild colonies of honey bees located in the Arnot forest and Ithaca, New York…………………………….20 Chapter 5: DWV recombinant prevalence in DWV positive colonies from England, Wales and the USA………………………………………………………………………………………………….21 Appendices ………………………………………………………………………………………………………………………..21 Appendix 1: Effect of Varroa on the variation of transmission of DWV strains in honey bee pupae……………………………………………………………………………………………………21 Appendix 2: Occurrence of Deformed wing virus variants in the stingless Melipona subnitida and honey Apis mellifera bee populations in North Eastern Brazil…………..22 ABBREVIATIONS………………………………………………………………………………………………………………….23 GENERAL ABSTRACT……………………………………………………………………………………………………………24 GENERAL INTRODUCTION ……………………………………………………………………………………………….…26 The European honey bee (Apis mellifera)…………………………………………………………….,…26 European honey bee (Apis mellifera) colony losses…………………………………………………27 Association between honey bee losses and Varroa destructor mites………………………29 Honey bee viruses…………………………………………………………………………………………………..33 Deformed wing virus……………………………………………………………………………………………….34 Deformed wing virus transmission………………………………………………………………………….36 Deformed wing virus symptoms…………………………………………………………………………..…37 2 Varroa as a vector for Deformed wing virus……………………………………………………………39 Historical Deformed wing virus detection in honey bee colonies…………………………….40 Varroa tolerant populations of European honey bees…………………………………………….41 Aims………………………………………………………………………………………………………………….…...44 Reference……………………………………………………………………………………………………………….47 Chapter 1: ABC Assay - Method development and application to quantify the role of three DWV master variants in overwinter colony losses of European honey bees……………………….57 Abstract………………………………………………………………………………………………………………….57 Introduction……………………………………………………………………………………………………………58 Methods…………………………………………………………………………………………………………………62 Primer design………………………………………………………………………………………………62 Viral master variant plasmid standards……………………………………………………….63 Primer optimisation…………………………………………………………………………………….64 Viral master variant standard curves……………………………………………………………65 Method validation……………………………………………………………………………………….65 Amplification of the RT-qPCR (ABC assay)……………………………………………………67 HRM assay…………………………………………………………………………………………………..68 Sequencing………………………………………………………………………………………………….68 Results…………………………………………………………………………………………………………………...69 PCR and RT-PCR sensitivities…………………………………………………………………….…69 Competitive PCR…………………………………………………………………………………..…….69 PCR and RT-PCR efficiencies………………………………………………………………………..70 Robustness of the RT-qPCR (ABC assay)………………………………………………………73 Detection of DWV in honey bees using the ABC assay…………………………………74 3 DWV master variants implicated in overwinter colony loss…………………………75 Discussion……………………………………………………………………………………………………………...79 Reference……………………………………………………………………………………………………………….82 Supplementary information……………………………………………………………………………………85 Chapter 2: DWV master variant dominance and seasonal prevalence in honey bee colonies from England and Wales…………………………………………………………………………………………………….87 Abstract………………………………………………………………………………………………………………….87 Introduction………………………………………………………………………………………………………..….88 Methods…………………………………………………………………………………………………………………91 Sample collection………………………………………………………………………………………..91 Sample processing……………………………………………………………………………………...92 RT-qPCR………………………………………………………………………………………………………93 Analysis of the results………………………………………………………………………………….94 Results…………………………………………………………………………………………………………………...96 A comparison of DWV variant load and distribution in 2010 and 2016…….….96 A comparison of apiary level DWV variant load and prevalence between spring and late summer 2016………………………………………………………………………98 Colony level DWV load, within apiary variation and associated colony mortality……………………………………………………………………………………………………100 Varroa susceptible verses tolerant colonies………………………………………………103 Discussion……………………………………………………………………………………………………….…...105 Conclusion…………………………………………………………………………………………………………….109 References…………………………………………………………………………………………………………...109 Supplementary information………………………………………………………………………………….113 4 Chapter 3: DWV infection, mortality and survivorship in pre-wintering Varroa susceptible and tolerant honey bee colonies from the USA…………………………………………………………………116 Abstract………………………………………………………………………………………………………………..116 Introduction………………………………………………………………………………………………………….117 Methods……………………………………………………………………………………………………………….120 Sample collection………………………………………………………………………………………120 Sample protocol ……………………………………………………………………………………….122 DWV type A and B distribution in heads and bodies of adult worker bees…123 RT-qPCR…………………………………………………………………………………………………….124 Analysis of the results……………………………………………………………………………….125 Results………………………………………………………………………………………………………………….129 DWV load in honey bee bodies verses heads…………………………………………….129 Apiary level analysis…………………………………………………………………………………..130 DWV variant prevalence and load in hobbyist and commercial colonies……132 Colony level analysis………………………………………………………………………………….134 Colony mortality………………………………………………………………………………………..137 Varroa susceptible and tolerant colonies…………………………………………………..139 Discussion……………………………………………………………………………………………………………..141 Conclusion…………………………………………………………………………………………………………….145 Reference……………………………………………………………………………………………………………..146 Chapter 4: The role of DWV avirulence and superinfection exclusion in wild colonies of honey bees located in the Arnot forest and Ithaca, New York…………………………………………..152 Abstract………………………………………………………………………………………………………………..152 Introduction………………………………………………………………………………………………………….153 5 Methods………………………………………………………………………………………………………….……158 Sample history and collection…………………………………………………………………...158 Sample processing…………………………………………………………………………………….160 RT-qPCR…………………………………………………………………………………………………….161 Data analysis……………………………………………………………………………………………..162 Results………………………………………………………………………………………………………………….164 DWV master variant prevalence per year………………………………………………….164 DWV seasonal prevalence and load between groups………………………………..167 DWV autumn infection in Arnot and Ithaca bees compared to surrounding states and USA tolerant bees…………………………………………………………………….169 Discussion……………………………………………………………………………………………………………..170 Conclusion…………………………………………………………………………………………………………….174 Reference……………………………………………………………………………………………………………..174 Chapter 5: DWV recombinant prevalence in DWV positive colonies from England, Wales and the USA…………………………………………………………………………………………………………………………….178 Abstract………………………………………………………………………………………………………………..178 Introduction………………………………………………………………………………………………………….179 Methods……………………………………………………………………………………………………………….183 Sample collection………………………………………………………………………………………183 Sample processing…………………………………………………………………………………….183 DWV master variant PCR and RT-qPCR………………………………………………………184
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