Identification of proviral and host restriction factors that impact the flaviviriclae lifecycle

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2021

Authors

Shue, Byron

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Beard, Michael

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Abstract

The complex interplay between virus and host is an often overlooked key determinant to the outcome of infection. Host factors are instrumental in supporting the entire virus lifecycle, where a majority have been implicated in the virus replication, especially for the formation of replication organelles. Simultaneously, the innate immune response confers a rapid non-specific defence against invading flaviviruses, where interferon (IFNs) and subsequently interferon stimulated genes (ISGs) are upregulated to directly inhibit virus replication and induction of an antiviral state within a cellular population. Although numerous host factors have been identified and characterised to be critical for the flaviviridae lifecycle through the usage of both RNAi and CRISPR genome-wide screens, limitations with RNAi technology combined with modifications of screening methodology may allow elucidation of novel host factors for virus infection. This thesis aims to identify and characterise novel host factors which positively and negatively influence the flaviviridae lifecycle, using in part genome wide CRISPR knockout (KO) technology. Hepatitis C Virus (HCV) is a medically important member of the flaviviridae family and thus chosen as a model virus to search for pan-flaviviridae novel host factors. The first aim explores the generation of multiple HCV sub-genomic (SGR) cell lines with mCherry, GFP and thymidine kinase (TK) reporter genes and their compatibility with our chosen screening platform, the GeCKO LentiCRISPRv2 library. Proof of concept experiments with the LentiCRISPRv2 system were performed via simplistic KO of EGFP or Cluster of Differentiation 81 (CD81), demonstrating the effectiveness of CRISPR in targeting both exogenous and endogenous genes. Furthermore, issues which surface upon attempting the genome-wide CRISPR screen with the HCV SGR are addressed and future directions proposed. The second aim repurposes the optimised GeCKO LentiCRISPRv2 library with an altered screening methodology to identify novel host factors important for ZIKV infection which may also be important for cell survival. Our top hits include previously identified host factors from the endoplasmic reticulum membrane protein complex (EMC) complex in addition to novel host factors Bcl-2-associated X protein 2 (BAX2) and Receptor for Activated C Kinase 1 (RACK1). We show that RACK1 is an important pro-viral host factor for both mosquito and tick-borne flaviviruses. Furthermore, it plays a critical role in the construction of replication organelles early in the virus lifecycle. Furthermore, flavivirus non-structural protein 1 (NS1) which is important for the biogenesis of vesicle packets (VPs) is able to interact with RACK1 within the ER lumen. Collectively this aim reinforces the utility of CRISPR/Cas9 genome-wide KO screens in the identification of viral host dependency factors and identifies RACK1 as a scaffold protein for the recruitment of viral NS proteins that are essential to the biogenesis of the replication complex. The third aim investigates the cellular innate immune response to Zika Virus (ZIKV) infection. We show that biologically relevant transformed and primary cells infected with ZIKV have abrogated upregulation of IFN-b and associated ISGs. In addition, we have identified that ZIKV attenuates the RIG-I-like Receptors (RLR) signalling pathway, required for the efficient induction of ISGs. Expression of the highly characterised multifunctional ISG viperin inhibits the ZIKV lifecycle and this observation is complemented by the usage of ZIKV infected viperin-/- mouse embryonic fibroblasts (MEFs). Finally, we show that the C-terminus of viperin is critical for ZIKV antiviral activity, an observation which is supported in future publications by collaborators. Collectively, this thesis not only enhances our understanding of the flavivirus lifecycle and their complex relationship with the host but also may guide towards the development of effective novel therapeutics against flavivirus infection.

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School of Biological Sciences

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Thesis (Ph.D.) -- University of Adelaide, School of Biological Sciences, 2021

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This electronic version is made publicly available by the University of Adelaide in accordance with its open access policy for student theses. Copyright in this thesis remains with the author. This thesis may incorporate third party material which has been used by the author pursuant to Fair Dealing exceptions. If you are the owner of any included third party copyright material you wish to be removed from this electronic version, please complete the take down form located at: http://www.adelaide.edu.au/legals

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