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Untersuchungen Zur Interspezies-Übertragung Fledertierassoziierter RNS-Viren

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Untersuchungen Zur Interspezies-Übertragung Fledertierassoziierter RNS-Viren Aus dem Institut für Virologie der Stiftung Tierärztliche Hochschule Hannover Untersuchungen zur Interspezies-Übertragung fledertierassoziierter RNS-Viren HABILITATIONSSCHRIFT Zur Erlangung der Venia legendi An der Stiftung Tierärztliche Hochschule Hannover vorgelegt von Nadine Krüger, PhD Hannover 2019 Tag der nichtöffentlichen wissenschaftlichen Aussprache: 12.07.2019 Die Anfertigung dieser Arbeit wurde durch Sachbeihilfen der Deutschen Forschungsgemeinschaft (KR 4762/2-1, HE 1168/16-1) gefördert. Inhaltsverzeichnis Liste der verwendeten Publikationen I Abkürzungsverzeichnis III 1.1. Fledertiere als Reservoir zoonotischer Viren ............................................................. - 1 - 1.2. Paramyxoviren ......................................................................................................... - 3 - 1.2.1. Henipaviren - 3 - 1.2.2. Mumpsviren - 5 - 1.3. Orthomyxoviren ....................................................................................................... - 7 - 1.3.1. Fledertierassoziierte Influenzaviren - 7 - 1.4. Filoviren ................................................................................................................... - 9 - 1.4.1. Ebolaviren - 9 - 1.5. Interspezies-Übertragung und die Speziesbarriere ....................................................- 12 - 1.6. Der virale Replikationszyklus und seine Barrieren während der Interspezies- Übertragung ...........................................................................................................- 17 - 1.6.1. Der Eintrittsprozess - Rezeptorbindung und Fusion - 18 - 1.6.2. Replikation und Proteinsynthese - 23 - 1.6.3. Antiviral wirksame Wirtsfaktoren - 26 - 2. Konzept und Ziele der Arbeit - 31 - 3. Ergebnisse - 33 - 3.1. Spezies-abhängige Fusionsaktivität der KuV Hüllproteine .........................................- 33 - 3.2. Identifizierung zellulärer Faktoren für den Eintrittsprozess fledertierassoziierter Influenzaviren .........................................................................................................- 35 - 3.3. Spezies-übergreifende Replikation, immunmodulatorische Eigenschaften und Neurovirulenz des fledertierassoziierten MuV .........................................................- 37 - 3.4. Expression und antivirale Wirkung von Flughund-Tetherin .......................................- 43 - 4. Übergreifende Diskussion - 45 - 4.1. Molekular-virologische Methoden zur Untersuchung neuer Viren ............................- 45 - 4.2. Zelluläre Barrieren als Restriktionsfaktoren .............................................................- 48 - 4.3. Neuartige Viren und das Risiko zoonotischer Infektionen .........................................- 52 - 5. Zusammenfassung - 54 - 6. Summary - 56 - 7. Literaturverzeichnis - 58 - 8. Darstellung des eigenen Anteils an den wissenschaftlichen Publikationen - 83 - 9. Publikationen - 85 - I Publikationen LISTE DER VERWENDETEN PUBLIKATIONEN Die Publikationen sind in chronologischer Reihenfolge nach dem Datum ihrer Veröffentlichung aufgelistet. Die Erklärung über den eigenen Anteil an den jeweiligen Publikationen ist in Kapitel 8 angegeben. Publikation 1: Krüger N, Hoffmann M, Drexler JF, Müller MA, Corman VM, Drosten C, Herrler G (2014). Attachment protein G of an African bat henipavirus is differentially restricted in chiropteran and nonchiropteran cells. J Virol. 2014 Oct;88(20):11973-80. doi: 10.1128/JVI.01561-14. http://jvi.asm.org/content/88/20/11973.long Publikation 2: Krüger N, Hoffmann M, Drexler JF, Müller MA, Corman VM, Sauder C, Rubin S, He B, Örvell C, Drosten C, Herrler G (2015). Functional properties and genetic relatedness of the fusion and hemagglutinin-neuraminidase proteins of a mumps virus-like bat virus. J Virol. 2015 Apr;89(8):4539-48. doi: 10.1128/JVI.03693-14. http://jvi.asm.org/content/89/8/4539.long Publikation 3: Hoffmann M, Krüger N, Zmora P, Wrensch F, Herrler G, Pöhlmann S (2016). The hemagglutinin of bat-associated influenza viruses is activated by TMPRSS2 for pH-dependent entry into bat but not human cells. PLoS One 2016 Mar 30;11(3):e0152134. doi: 10.1371/journal.pone.0152134. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4814062/ Publikation 4: Krüger N, Sauder C, Hoffmann M, Örvell C, Drexler JF, Rubin S, Herrler G (2016). Recombinant mumps viruses expressing the batMuV fusion glycoprotein are highly fusion active and neurovirulent. J Gen Virol 2016 Nov; 97(11):2837-2848. doi: 10.1099/jgv.0.000596. http://jgv.microbiologyresearch.org/content/journal/jgv/10.1099/jgv.0.000596#tab2 Publikationen II Publikation 5: Krüger N, Sauder C, Hüttl S, Papies JP, Voigt K, Herrler G, Hardes K, Steinmetzer T, Örvell C, Drexler JF, Drosten C, Rubin S, Müller MA, Hoffmann M (2018). Entry, replication, immune evasion and neurotoxicity of synthetically-engineered bat-borne mumps virus. Cell Rep. 2018 Oct 9;25(2):312-320.e7. doi: 10.1016/j.celrep.2018.09.018. https://doi.org/10.1016/j.celrep.2018.09.018 Publikation 6: Hoffmann M, Nehlmeier I, Brinkmann C, Krähling V, Behner L, Moldenhauer AS, Krüger N, Nehls J, Schindler M, Maisner A, Becker S, Pöhlmann S (2018). Tetherin inhibits replication of Ebola virus and Nipah virus in fruit bat cells. J Virol. 2018 93:e01821-18. doi: 10.1128/JVI.01821-18. https://doi.org/10.1128/JVI.01821-18. III Abkürzungen ABKÜRZUNGSVERZEICHNIS ACE2 Angiotensin-konvertierendes Enzym 2 (angiotensin converting enzyme) batMuV Fledertierassoziiertes Mumpsvirus (BatPV/Epo_spe/AR1/DCR/2009) BDBV Bundibugyo-Virus BOMV Bombali-Virus BST2 Bone marrow stromal antigen 2 (= Tetherin) CDC Centers for Disease Control and Prevention CedV Cedar-Virus CPE Zytopathischer Effekt (cytopathic effect) CpKd Carollia perspicillata Nierenzellen EBOV Ebola-Virus EidLu Eidolon helvum Lungenzellen EidNi Eidolon helvum Nierenzellen EpoNi Epomops buettikoferi Nierenzellen ER Endoplasmatisches Retikulum F Fusionsprotein FLUAV Influenza A Virus G Glykoprotein (Paramyxoviren) GFP Grün fluoreszierendes Protein GP Glykoprotein (Ebolaviren) H/ HA Hämagglutinin HeV Hendra-Virus HIV Humanes Immundefizienz-Virus HL HA-ähnlich (HA-like) hMuV Humanes Mumpsvirus HN Hämagglutinin-Neuraminidase HPAI Hoch pathogene (highly pathogenic) aviäre Influenzaviren HypLu Hypsignathus monstrosus Lungenzellen HypNi Hypsignathus monstrosus Nierenzellen IFN Interferon IRF-3 IFN-regulierender Faktor 3 Abkürzungen IV ISG(s) IFN-stimuliertes Gen/ -stimulierte Gene KuV Kumasi-Virus (Eid_hel/GH-M74a/GHA/2009) L Large protein, RNS-abhängige RNS Polymerase LPAI Niedrig pathogene (low-pathogenic) aviäre Influenzaviren M Matrixprotein mRNS Boten-RNS (messenger RNA) MuV Mumpsvirus N Nukleoprotein N/ NA Neuraminidase NF-κB Nuklearfaktor kappa in B-Zellen NHP Nicht-humane Primaten NiV Nipah-Virus NL NA-ähnlich (NA-like) P Phosphoprotein PAMPs Pathogen-assoziierte molekulare Muster PCR Polymerase-Kettenreaktion PipNi Pipistrellus spec. Nierenzellen RdRp RNS-abhängige RNS-Polymerase (RNA-dependent RNA polymerase) RESTV Reston-Virus RNP Ribonukleoprotein RNS Ribonukleinsäure RoNi Rousettus aegyptiacus Nierenzellen RT-PCR Reverse-Transkriptase-Polymerase-Kettenreaktion SARS Schwer akutes Atemwegssyndrom (severe acute respiratory syndrome) SARS-CoV SARS-assoziiertes Coronavirus SARSr-CoV SARS-ähnliches CoV (SARS-related CoV) SeV Sendai-Virus SH Kleines hydrophobes Protein (small hydrophobic protein) siRNS Small interfering RNS STAT Signaltransduktor und Aktivator der Transkription SUDV Sudan-Virus V Abkürzungen TAFV Tai Forest-Virus TLR Toll-ähnlicher Rezeptor (toll-like receptor) TNF-α Tumornekrosefaktor alpha TSSPs Typ II Transmembran-Serinproteasen VLPs Virus-ähnliche Partikel (virus-like particles) VP40 Virusprotein 40 VSV Virus der Stomatitis vesicularis (vesicular stomatitis virus) VSVpp VSV-basierte Pseudotypen WHO World Health Organization ZNS Zentralnervensystem - 1 - Einleitung 1. EINLEITUNG 1.1. Fledertiere als Reservoir zoonotischer Viren Fledertiere werden innerhalb der Klasse der Säugetiere (Mammalia) der Ordnung Chiroptera zugeordnet. Ursprünglich wurden die insgesamt über 1200 verschiedenen Fledertierarten aufgrund morphologischer Ähnlichkeiten und ihren Lebensweisen in die zwei Unterordnungen Megachiroptera (Flughunde) und Microchiroptera (Fledermäuse) unterteilt. Basierend auf tiefergreifenden phylogenetischen Analysen, die Informationen über die evolutionäre Verwandtschaft verschiedener Fledertierarten lieferten, wird mittlerweile eine Unterteilung in die zwei Unterordnungen Yangochiroptera und Yinpterochroptera vorgeschlagen (Teeling et al., 2005, Hutcheon 2006, Lei and Dong 2016). Zu den Yinpterochiroptera, häufig auch als Pteropodiformes bezeichnet, werden demnach alle Flughunde (Familie Pteropodidae), sowie fünf Familien der Fledermäuse (Craseonycteridae, Hipposideridae, Rhinopomatidae, Rhinolophidae, Megadermatidae) gezählt. Die Unterordnung Yangochiroptera oder Vespertilioniformes umfasst alle Fledermäuse, die nicht den Yinpterochiroptera zugeordnet sind. Bedingt durch ihre große Anzahl hoch diverser Arten, ein nahezu weltweites Vorkommen, eine vergleichsweise lange Lebensspanne und die Befähigung zum aktiven Fliegen, stellen Fledertiere ein ideales Reservoir für diverse pathogene Erreger dar. Bisher wurden bereits mehr als 60 verschiedene Virusspezies in Fledertieren detektiert (Calisher et al., 2006, Hayman 2016). Für einige dieser Viren konnte bereits nachgewiesen werden, dass sie ein zoonotisches Potential besitzen. Als der bekannteste fledertierassoziierte Zoonoseerreger ist das Tollwutvirus zu nennen. Tollwutviren (Genus Lyssavirus) wurden
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