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Scheibner Online.Pdf Aus dem Institut für molekulare Virologie und Zellbiologie des Friedrich-Loeffler-Instituts Bundesforschungsinstitut für Tiergesundheit und dem Institut für Veterinär-Biochemie des Fachbereichs Veterinärmedizin der Freien Universität Berlin Virulence of Avian Influenza H7 Viruses in Birds: The Impact of the Hemagglutinin Polybasic Cleavage Site on the Virulence of a Recent German H7N7 Virus and Pathogenicity of European H7N7 and H7N1 Viruses in Ducks Inaugural-Dissertation zur Erlangung des Grades eines Doktors der Veterinärmedizin an der Freien Universität-Berlin vorgelegt von David Scheibner Tierarzt aus Stralsund Berlin 2020 Journal-Nr.: 4170 Gedruckt mit Genehmigung des Fachbereichs Veterinärmedizin der Freien Universität Berlin Dekan: Univ.-Prof. Dr. Jürgen Zentek Erster Gutachter: PD Dr. Christoph Gabler Zweiter Gutachter: Prof. Dr. Thomas C. Mettenleiter Dritter Gutachter: Prof. Dr. Benedikt Kaufer Deskriptoren (nach CAB-Thesaurus): muskovy ducks pekin ducks avian influenza viruses influenza viruses virulence virulence factors viral hemagglutinins Tag der Promotion: 03.06.2020 Dedication This work is dedicated to my lovely family and my girlfriend. Thanks for your Support, Patience and Motivation. In memory of my father and my grandparents. Content Content Chapter 1 .............................................................................................................................. 1 General Introduction .......................................................................................................... 1 1 Introduction ..................................................................................................................... 2 2 Review of literature ......................................................................................................... 3 2. 1 Avian influenza virus ................................................................................................ 3 2.1.1 Virus taxonomy and structure ............................................................................................ 3 2.1.2 Virus replication ................................................................................................................. 4 2.1.3 Proteolytic activation ......................................................................................................... 7 2.1.4 Virus evolution ................................................................................................................... 8 2.2 Pathogenicity of AIV in different host species ........................................................... 9 2.2.1 Domestic Poultry ................................................................................................................ 9 2.2.2 Wild birds.......................................................................................................................... 10 2.2.3 Mammals .......................................................................................................................... 11 2.2.4 Humans ............................................................................................................................. 11 2.3 Genetic determinants for virulence in and adaptation of AIV to poultry ....................12 2.3.1 Virulence determinants in chickens ................................................................................. 12 2.3.1.1 Hemagglutinin: HA cleavage site, its vicinity and head domain ................................ 12 2.3.1.2 Neuraminidase: NA stalk deletion and reduction in NA activity ............................... 13 2.3.1.3 Other gene segments ................................................................................................ 14 2.3.2 Virulence determinants in ducks and turkeys .................................................................. 14 2.3.3 Vaccines and antivirals against AIV .................................................................................. 15 2.4 Significance of AIV H7 infections in birds and humans ............................................16 2.4.1 History and subtypes ........................................................................................................ 16 2.4.2 Direct evolution of HPAIV H7 from LPAIV precursors ...................................................... 16 2.4.3 Epidemiology of H7 in Germany ....................................................................................... 20 2.4.3.1 H7N7 in Germany ...................................................................................................... 20 2.4.3.2 Outbreak of H7N7 in Emsland, Germany 2015 ......................................................... 20 3 Aim of this study ............................................................................................................22 4 Outline of publications in this dissertation ......................................................................23 4.1 First publication (published in Scientific Reports, 2019) ...........................................23 4.2 Second publication (published in BMC Veterinary Research, 2019) ........................23 Content Chapter 2 ............................................................................................................................ 25 Variable impact of the hemagglutinin polybasic cleavage site on virulence and pathogenesis of avian influenza H7N7 virus in chickens, turkeys and ducks ...........................................25 Chapter 3 ............................................................................................................................ 39 Virulence of three European highly pathogenic H7N1 and H7N7 avian influenza viruses in Pekin and Muscovy ducks.................................................................................................39 Chapter 4 ............................................................................................................................ 47 General Discussion ...........................................................................................................47 6 Summary .......................................................................................................................53 7 Zusammenfassung ........................................................................................................54 8 References (of Introduction and General discussion) .....................................................56 9 List of Publications .........................................................................................................78 9.1 Publications of the presented data ...........................................................................78 9.1.1 Articles in scientific journals ............................................................................................. 78 9.1.2 Abstracts ........................................................................................................................... 78 9.2 Other publications in scientific journals ....................................................................80 10 Acknowledgments ........................................................................................................81 11 Funding ........................................................................................................................82 12 Declaration Freie Universität Berlin ..............................................................................83 List of figures and tables Figure 1: Structure of an influenza virion Figure 2: Influenza A virus replication cycle Figure 3: Clinical signs after HPAIV infection Table 1: Outbreaks of HPAIV H7Nx directly evolved from LP precursors from 1990s to 2019 Table 2: Mutations in LP and HPAIV H7N7 from 2015 (HACS excluded) List of Abbreviations Abbreviations AA Amino acid AIV Avian influenza virus ATP Adenosin triphosphat cRNA Complementary ribonucleic acid C Carboxy COOH Carboxyl group CS Cleavage site E Glutamic acid ECE Embryonated chicken eggs EE Early endosomes ER Endoplasmic reticulum F Phenylalanine G Glycine H Hydrogen HA Hemagglutinin HACS Hemagglutinin cleavage site HP Highly pathogenic HPAIV High pathogenic avian influenza virus K Lysine L Leucine LE Late endosomes LP Low pathogenic LPAIV Low pathogenic avian influenza virus LP-poly LP + polybasic cleavage site M Matrix protein N Nitrogen NH2 Amino group mRNA Messenger RNA NA Neuraminidase NEP Nuclear export protein NP Nucleoprotein NS Non-structural protein O Oxygen PA Polymerase acidic P Prolin PB Polymerase basic R Arginine RIG-I Retinoic acid inducible gene I RNA Ribonucleic acid RNP Ribonucleoprotein RT-PCR Reverse-transcription polymerase chain reaction SA Sialic acid T Threonine UK United Kingdom USA Unites States of America V Valin vRNA Viral ribonucleic cid WHO World Health Organization Introduction Chapter 1 General Introduction I 1 Introduction 1 Introduction Avian influenza viruses (AIV) are members of the Influenza A genus in the family Orthomyxoviridae. They are enveloped pleomorphic viruses with a segmented genome composed of eight segments which encode for at least ten viral proteins. The surface glycoproteins hemagglutinin (HA) and neuraminidase (NA), vary antigenically into 16 HA and 9 NA subtypes. Each AIV carries one HA and one NA with 144 possible combinations. Wild aquatic birds, particularly waterfowls, serve as natural reservoir for all AIV subtypes. Transmission of AIV from wild to domestic birds is a continuing threat particularly to farmed poultry as well as zoo and feral birds worldwide.
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