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Immunological Response Profiles to Salmon Lice Infections in Atlantic Salmon

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Immunological Response Profiles to Salmon Lice Infections in Atlantic Salmon Immunological response profiles to salmon lice infections in Atlantic salmon Modulation by nutrition and selective breeding Philosophiae Doctor (PhD) Thesis Helle Jodaa Holm Department of Basic Sciences and Aquatic Medicine Faculty of Veterinary Medicine Norwegian University of Life Sciences Adamstuen 2017 Thesis number 2016:96 ISSN 1894-6402 ISBN 978-82-575-1982-7 Contents Acknowledgements ........................................................................................................... 1 Summary ............................................................................................................................ 3 Sammendrag...................................................................................................................... 6 List of papers ..................................................................................................................... 9 Abbreviations .................................................................................................................. 10 1. Introduction ............................................................................................................ 11 1.1. Current salmon lice situation in Norwegian aquaculture ............................... 11 1.1.1 Chemical control of salmon lice in Norwegian aquaculture........................ 12 1.1.2 Husbandry, management and technical innovations against lice ............... 14 1.1.3 Lice impact on wild salmonids ..................................................................... 16 1.2. Life history traits of salmon louse and its fish hosts ....................................... 17 Specifics of Atlantic salmon life history traits in comparison to Pacific salmonids 17 Salmon louse life cycle, feeding habits, communication and reproduction 20 Immunomodulation of the host by lice ....................................................... 25 1.3. Host responses to lice ..................................................................................... 26 1.3.1 Local skin tissue responses following L. salmonis infection ........................ 26 1.3.2 Immune responses ...................................................................................... 30 1.3.3 Systemic (physiological) responses to lice infection ................................... 42 1.4. Biological approaches for salmon lice control ................................................ 45 1.4.1 Vaccines against lice .................................................................................... 46 1.4.2 Selective breeding for increased resistance against lice ............................. 47 1.4.3 Anti-lice functional feeds ............................................................................. 51 2. Objectives ............................................................................................................... 57 3. Summary of Papers ................................................................................................. 58 4. Methodology .......................................................................................................... 61 4.1. Fish experiments and study designs ............................................................... 61 4.2. Selection of tissues for various analyses ......................................................... 62 4.3. Microarray technology .................................................................................... 65 4.4. qPCR ................................................................................................................. 65 4.5. Histology and Immunohistochemistry ............................................................ 67 5. Results and Discussion ............................................................................................ 69 5.1. General discussion ........................................................................................... 69 5.2. Host responses during the copepodid and chalimus stages ........................... 71 5.3. Protection mediated by selective breeding .................................................... 72 5.4. Protection mediated by glucosinolates-based functional feeds ..................... 75 5.5. Integrated pest management control ............................................................. 77 6. Main conclusions .................................................................................................... 79 7. Future perspectives ................................................................................................ 81 References ....................................................................................................................... 82 Errata ............................................................................................................................. 108 Scientific papers ............................................................................................................ 111 Acknowledgements This thesis is based on studies, which were conducted between 2012 and 2016 within the Section of Aquatic Medicine, Department of Basic Science and Aquatic Medicine, Norwegian School of Veterinary Science (now Norwegian University of Life Sciences). Financial grants were provided from the Research Council of Norway, project no. 20351 Sea Lice Research Centre. I am very grateful to all the people who have supported me and helped me through the last four years. Professor Øystein Evensen, thank you. You gave me the opportunity to do this PhD, the free- dom so I could learn how to be an independent researcher with integrity, and thorough sci- entific guidance throughout the PhD. Professor Erling Olaf Koppang, for your support and supervision through my favourite part of the PhD, namely histology and immunohistochemistry. Stanko Škugor. Thank you for your positivity, gentleness and patience, and for always being there for me. Not only have you been an excellent and exceptionally inspiring and knowl- edgeable scientific supervisor, you are also one of my best and closest friends. Simon Wadsworth, Ragna Heggebø and Anne Kari Bjelland. Thank you for inviting me to work with you and the rest of the super Cargill (Ewos) Innovation team, supporting me and teaching me all I know about lice and functional feeds. Big thanks to my other co-authors: Nina Santi, Sissel Kjøglum, Aleksei Krasnov, Jorge Pino and Nebojsa Perisic, for great collab- oration and scientific advice. Colleagues in the Aqua group and at NMBU campus Adamstuen (Norwegian School of Vet- erinary Science), thank you for your support. A special thanks to Elin Petterson, Ida Lieungh, Sandra Radunovic and Celia Agustí-Ridaura for being true friends. Alf Seljenes Dalum - thanks for always helping me. Aase Mikalsen – thanks for your support and technical assistance. My big and loving family: including of course Linus (our magnificent Golden dog). Thank you for always having my back and helping me put things into perspective. A special hug of 1 auntylove goes to my niece Kaja and nephews Sigurd, Vegard and Isak. I am grateful for the loving memory of my late grandfather, Olav Holm (M.Sc, 1928-2015), a true scholar of his time, and the first to see the scientist in me. Last, but certainly not least. My best friends whom I know from Tønsberg and my veterinary studies – Betine, Eli, Elin, Hanne, Kristiane, Line, Maja (and her horses), Marit and Marte. Thank you for putting up with me, inspiring me and making me think of and do other things than science and work. Oslo, November 2016 Helle Jodaa Holm 2 Summary Infections with the salmon louse Lepeophtheirus salmonis (L. salmonis) represent one of the most important limitations to sustainable Atlantic salmon (Salmo salar) farming today. The parasite exerts negative impacts on health, growth and welfare of farmed fish, and there are concerns of the impact on wild salmon populations. Current control relies heavily on the use of a few chemical in-feed or bath treatments and an increasing number of reports states their reduced efficiency. Increased mortalities during treatments are also frequently re- ported. The narrow treatment repertoire is also a consequence of the difficulty in developing novel anti-parasitic chemical treatments that are efficient against lice, environment-friendly and posing no risk for the consumers. Functional feeds and selective breeding are consid- ered two promising alternative approaches for management of salmon lice infections, and understanding the molecular basis of protection against lice is expected to help in their suc- cessful application in Atlantic salmon aquaculture. The overall aim of this PhD study was to characterize responses to salmon lice infections and explore the possibility to modulate these by nutritional components and selective breeding. This required a basic understanding of the host responses to lice. The first study (paper I) investigated host gene expression responses at two skin locations, which are preferential sites for lice attachment and feeding activities; the scaled skin behind the dorsal fin and scaleless skin from the top of the head. Responses were studied before and during L. salm- onis copepodid infection (4 days post infection (dpi)) and moult from copepodids to chalimii (8 dpi). Significant differences in basic transcript levels (constitutive expression prior to in- fection) were found between the two skin sites, suggesting a non-homogenous distribution of immune components across the skin of the fish. Immunohistochemistry was used to study in situ localization of MHC class
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