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Horizontal Transfer of Wolbachia Between Different Hymenopteran

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Horizontal Transfer of Wolbachia Between Different Hymenopteran Horizontal transfer of Wolbachia between different Hymenopteran host‐parasitoid pairs Aileen Berasategui López th 1 26 September 2011 Master thesis by Aileen Berasategui López University of Bremen September 2011 Supervisors: Prof. Dr. Thomas Hoffmeister Dr. Martin Kaltenpoth 2 To my family 3 “For the present, I suggest that any doubting readers should “look out of the window” and list every organism they can see. I can guarantee that most, probably all, organisms on the list are a product of symbiosis.” Angela E. Douglas 4 Acknowledgements It has been a while already since I left home to study biology in Oviedo. I always thought I would come back home after College, but life turned my plans upside down giving me the opportunity to do my degree’s last year abroad. Now that I have finished my master in Germany, I would like to thank the people that have helped me through these last years. I would first like to thank Prf. Dr. Thomas Hoffmeister, who gave me the opportunity to continue my studies in Ecology in University of Bremen, which ultimately opened for me the doors of science. I am extremely thankful to my supervisor Dr. Martin Kaltenpoth, for teaching me everything I know about insect symbiosis. Thank you for making of the lab such a nice atmosphere, for encouraging us to develop as critical scientists and to provide us with all the resources (technical but also personal) we need to do our best. I would also like to thank Dr. Oliver Niehuis, Thomas Schmitt and Mareike Wurdack for providing some of the samples I have used in this project and Dr. Johannes Kroiss for the great wasp pictures that he has lend me. My time in the Institute would not have been as productive, comfortable and happy if it wasn´t for my colleagues Hassan, Laura, Sabrina, Sailen, Taras, Tobi, Uli and Woife, who have become not only my co‐workers but friends. Thank you for those long talks, advices, cakes, neigbor‐joining dinners and so many other things; also for guiding me through science. And last but not least, I would like to thank Jorge and my big family. Specially my dad, who is a passionate of science and showed me the first microorganism I ever saw; my mum, who has always encouraged me to pursue my dreams, even when they are away from home; and my sister, who is always there, no matter the distance, for long and nocturnal talks. Thank you for your wise advices, economical support and for cheering my up when I was down. 5 Table of Contents Acknowledgements .............................................................................................. 5 Table of Contents ................................................................................................. 6 Summary .............................................................................................................. 8 1. Introduction ................................................................................................ 9 1.1. Insect‐bacteria symbiosis .............................................................................. 9 1.2.Wolbachia ...................................................................................................... 9 1.2.1. Biology of Wolbachia .............................................................................................................. 9 1.2.2. Transmission of Wolbachia ................................................................................................... 10 1.2.2.1 Vertical transmission ...................................................................................................... 10 1.2.2.2 ......................................................................................................................................... 11 Horizontal transmission .............................................................................................................. 11 1.3. Hymenopteran hosts and parasitoids ......................................................... 12 1.4. Multi Locus Sequence Typing (MLST) .......................................................... 12 1.5. Objectives of the project ............................................................................. 13 2. Methods and material .............................................................................. 15 2.1. Collection, dissection, DNA extraction and Wolbachia screening ........................................... 15 2.1.1. Wolbachia supergroups .................................................................................................... 16 2.1.2. Wolbachia prevalence ...................................................................................................... 16 2.2. Wolbachia Multilocus Sequence Typing (MLST) ...................................................................... 18 2.2.1. PCR amplification and sequencing .................................................................................... 18 2.2.2. Cloning .............................................................................................................................. 18 2.3. Phylogenetic analysis ............................................................................................................... 18 2.4 Statistical analysis...................................................................................................................... 19 3. Results ....................................................................................................... 20 3.1. DNA extraction and Wolbachia prevalence ............................................................................. 20 3.2. MLST ........................................................................................................................................ 22 3.3. Phylogenetic analysis ............................................................................................................... 22 3.4. Statistical analysis ............................................................................................................... 27 6 4. Discussion ................................................................................................. 28 4.1. Wolbachia prevalence ............................................................................................................. 28 4.2. MLST and phylogenetic analysis .............................................................................................. 29 5. Conclusion ................................................................................................. 33 6. References ................................................................................................ 34 7. Apendix: Phylogenetic trees ..................................................................... 38 8. Protocols ................................................................................................... 42 DNA extraction protocol: Epicentre MasterPureTM DNA isolation ............................................. 42 Polymerase Chain Reaction ........................................................................................................ 43 Gel for electrophoresis (1 gel) .................................................................................................... 43 PeqGold MicroSpin Cycle‐PureKit (PeqLab) ................................................................................ 43 Soil DNA Purification Kit Protocol Epicentre ............................................................................... 44 Purification for sequencing with EtOH‐precipitation ................................................................. 45 Sequencing preparation.............................................................................................................. 45 Preparation of media and reagents ............................................................................................ 45 Cloning: StrataClone PCR Cloning Kit (Stratagene) ..................................................................... 46 7 Summary Wolbachia is one of the most common endosymbionts on Earth, infecting up to 70% of all insect species. They are maternally inherited through the egg cytoplasm, and manipulate their host reproductive system in such a way that increases the proportion of infected females among the host’s progeny. Therefore, Wolbachia is easily transmitted from generation to generation; driving itself into populations. Arthropod‐infecting Wolbachia do not usually show congruent phylogenies with their insect hosts, suggesting that horizontal transfer can also occur, although the mechanism by which it takes place is not very well known. Here we present a system in which horizontal transfer of Wolbachia can be assessed using different Hymenopteran host‐parasitoid pairs as a model. Host females are mass provisioning their offspring with insect prey. Parasitoids lay an egg by the host larva, and when the chrysidid larva hatches, it exploits not only the host progeny but also the stored resources. Thus, parasitoids would acquire Wolbachia by feeding on the host larvae. If this is the case, Wolbachia strains harbored by hosts and parasitoids belonging to the same pair would be identical or very similar, being more genetically related to each other than to strains belonging to hosts or parasitoids that are not associated with each other. In order to compare Wolbachia strains, we have performed a Multilocus Typing System approach to genotype strains for four different genes (wsp, ftsZ, coxA and fbpA). Analysis of the phylogenetic relationships between different strains showed similar
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