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The Reproductive Biology and Venom System of the Parasitoid Wasp Nasonia Vitripennis

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The Reproductive Biology and Venom System of the Parasitoid Wasp Nasonia Vitripennis Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and private study only. The thesis may not be reproduced elsewhere without the permission of the Author. The reproductive biology and venom system of the parasitoid wasp Nasonia vitripennis A thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Genetics at Massey University, Manawatu New Zealand. Ngonidzashe Faya 2019 ABSTRACT Nasonia vitripennis (Nasonia) is a parasitoid wasp that uses a wide range of fly species as hosts to support the growth and development of its offspring. This project focused on two important aspects of Nasonia reproductive biology: (1) The role of the endosymbiont Wolbachia pipientis (Wolbachia) in fertility and (2) the functional characteristics of the venom system. The first part of the project focused on a bacterial endosymbiont Wolbachia, which is found in high concentrations within the ovary and sperm cells of Nasonia. As Wolbachia is maternally inherited, it manipulates host reproduction in ways that favour the production of female offspring. In Nasonia, Wolbachia infection results in the generation of reproductive cytoplasmic incompatibilities between infected and uninfected wasps. As very little is known about the molecular mechanism used by Wolbachia to interfere with Nasonia reproduction. RNA-seq was used to identify 84 and 58 genes differentially expressed in the ovary and testis, respectively, in response to Wolbachia infection. RNA interference targeting differentially expressed genes was unable to directly identify functional roles for these genes in cytoplasmic incompatibility, suggesting that this mechanism may be regulated by co-expressed gene networks. Unexpectedly, this study identified a Wolbachia gene (gene1092) that has been incorporated into the Nasonia genome possibly by lateral gene transfer. The second part of the project looked at the Nasonia venom system that has previously been shown to contain a mixture of 79 peptides. The venom is introduced into the host before egg laying and plays a role in altering the host’s physiology in ways that favour development of the parasitoid’s progeny. However, the individual roles venom peptides play in conferring these dramatic changes in the host have yet to be identified. Therefore, RNA interference was used to target venom genes with the goal of better understanding their function roles. Using this approach, it was shown that venom X and venom Z were likely involved in arresting development in the host, while venom Y was likely to be involved in modulating the host’s immune response. As the molecular pathways affected by Nasonia venom underlie important cellular pathways, it is predicted that these venom peptides could be developed into drugs to combat diseases such as cancer, hypertension and diabetes. i ii ACKNOWLEDGEMENTS First and foremost, I want to thank GOD ALMIGHTY for giving me the grace and strength to pursue this PhD. It wasn’t an easy one, each day had its ups and downs therefore to be at this stage of putting all the work on paper, it can only be by grace. To my supervisors Dave, Helen, Patrick and Gill, I am still in awe of your patience and the support you have given me from day one up to the end. Dr. David Wheeler, thank you so much for giving me the opportunity to come to New Zealand. The support you gave me throughout the application process as well as in the project, taking me into your home when I first moved, helping me to get that balance between work and life, the eggs and nature trips. The list goes on and on…… Thank you for your continual support even from Aussie. Dr. Helen Fitzsimons, thank you so much for taking over the supervisory role at a time where I wasn’t sure of the direction I was going with the PhD. I really appreciate your support throughout, the patience and above all the encouragement and enthusiasm in the progress meetings. I always came out of those meetings believing that its possible. Associate Professor Patrick Biggs, thank you so much for helping me get back on my feet each time I fell. I really want to appreciate the time you took to teach me some of the bioinformatics software as well as helping me to get that balance of life. You were always available when I need help. Associate Professor Gill Norris, thank you so much for the support and wise advice you have given me throughout the PhD period. You have been there from the beginning till the end, thank you for believing in me. Protein work was daunting, but you helped me believe I could do it. To Trevor Loo, thank you so much for helping me get started with the protein work, your support and your wealth of knowledge is highly appreciated. The X-lab team (Fareeda Barzak, Jennifer, Sean, Dell, Brittany), thank you so much for answering all of my questions and being my lab mates too, each time I needed a chat, I knew where to find you. My mates in the Boffin lab (Nikolai, Hannah, Asmat, Elva, Ellie and Alysha), thanks for the chats and encouragements that we shared. iii To the MMIC team, Niki and Pani, thank you so much for helping me with sample preparation and teaching how to use the TEM. Raul for helping me understand the TEM methodology and how to analyze the results. To Trish McLenachan, thank you so much for helping in various ways in the lab, from finding chemicals, to using equipment and ordering lab materials. To my family for always supporting me and believing in me. Being away from home, you have always checked up on me and encouraged me. Members of Forward in Faith church, thank you so much for the support, the prayers, the packed lunches and dinners. This project was sponsored by Massey University under the Graduate Assistance Scholarship. Massive thanks go to the School of Fundamental Sciences for supplying all the things required for the project and also thanks a lot to Ann Truter and Cynthia Cresswell for the massive support and great chats. iv Table of Contents ABSTRACT ................................................................................................................................... i ACKNOWLEDGEMENTS ........................................................................................................ iii List of Figures .............................................................................................................................. ix List of Tables ............................................................................................................................... xii ABBREVIATIONS ..................................................................................................................... xv 1. INTRODUCTION & LITERATURE REVIEW ............................................................... 1 1.1. Overview of Nasonia vitripennis ........................................................................................ 1 1.1.1. Nasonia as a model genetic system ............................................................................ 4 1.2. The Nasonia reproductive system ...................................................................................... 6 1.2.1. Haplodiploidy sex determination ................................................................................ 6 1.2.2. Wolbachia ................................................................................................................... 7 1.3. The venom system of Nasonia ......................................................................................... 15 1.3.1. Hymenoptera venom ................................................................................................. 15 1.3.2. Nasonia venom ......................................................................................................... 18 1.3.2.1. Effects of venom on host ...................................................................................... 18 1.3.2.2. Nasonia venom peptides ....................................................................................... 20 1.3.3. Characterization of venom proteins .......................................................................... 22 1.3.3.1. RNA interference (RNAi)..................................................................................... 23 1.3.3.2. The generation of recombinant venom proteins in vitro ...................................... 24 1.3.4. Expression of Nasonia venom genes .............................................................................. 30 1.4. Aims and Hypothesis ........................................................................................................ 32 1.4.1. Research objectives .................................................................................................. 33 Aim 1 ........................................................................................................................................ 33 Aim 2 ........................................................................................................................................ 33 2. MATERIALS AND GENERAL METHODS .................................................................. 34 2.1. Culturing Nasonia wasps .................................................................................................. 34 2.2. Ultrapure Distilled Water ................................................................................................. 34 2.3. Tissue Extraction .............................................................................................................
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