A direct RNA-protein interaction atlas of the SARS-CoV-2 RNA in infected human cells

SARS-CoV-2 infections pose a global threat to human health and an unprecedented research challenge. Among the most urgent tasks is obtaining a detailed understanding of the molecular interactions that facilitate viral replication or contribute to host defense mechanisms in infected cells. While SARS-CoV-2 co-opts cellular factors for viral translation and genome replication, a comprehensive map of the host cell proteome in direct contact with viral RNA has not been elucidated. Here, we use RNA antisense purification and mass spectrometry (RAP-MS) to obtain an unbiased and quantitative picture of the human proteome that directly binds the SARS-CoV-2 RNA in infected human cells. We discover known host factors required for coronavirus replication, regulators of RNA metabolism and host defense pathways, along with dozens of potential drug targets among direct SARS-CoV-2 binders. We further integrate the SARS-CoV-2 RNA interactome with proteome dynamics induced by viral infection, linking interactome proteins to the emerging biology of SARS-CoV-2 infections. Validating RAP-MS, we show that CNBP, a regulator of proinflammatory cytokines, directly engages the SARS-CoV-2 RNA. Supporting the functional relevance of identified interactors, we show that the interferon-induced protein RYDEN suppresses SARS-CoV-2 ribosomal frameshifting and demonstrate that inhibition of SARS-CoV-2-bound proteins is sufficient to manipulate viral replication. The SARS-CoV-2 RNA interactome provides an unprecedented molecular perspective on SARS-CoV-2 infections and enables the systematic dissection of host dependency factors and host defense strategies, a crucial prerequisite for designing novel therapeutic strategies.

[1]  M. Mann,et al.  Multilevel proteomics reveals host perturbations by SARS-CoV-2 and SARS-CoV , 2020, Nature.

[2]  J. Seibel,et al.  The serotonin reuptake inhibitor Fluoxetine inhibits SARS-CoV-2 , 2020, bioRxiv.

[3]  Christian Drosten,et al.  Bulk and single-cell gene expression profiling of SARS-CoV-2 infected human cell lines identifies molecular targets for therapeutic intervention , 2020 .

[4]  Andrew R. Leach,et al.  The Global Phosphorylation Landscape of SARS-CoV-2 Infection , 2020, Cell.

[5]  Jennifer L. Bell,et al.  Effect of Dexamethasone in Hospitalized Patients with COVID-19: Preliminary Report , 2020, medRxiv.

[6]  John G. Doench,et al.  Genome-wide CRISPR screen reveals host genes that regulate SARS-CoV-2 infection , 2020, bioRxiv.

[7]  K. Sachaphibulkij,et al.  Annexin-A1 promotes RIG-I-dependent signaling and apoptosis via regulation of the IRF3–IFNAR–STAT1–IFIT1 pathway in A549 lung epithelial cells , 2020, Cell Death & Disease.

[8]  M. Torcia,et al.  Evidence for host-dependent RNA editing in the transcriptome of SARS-CoV-2 , 2020, Science Advances.

[9]  S. Ciesek,et al.  Growth Factor Receptor Signaling Inhibition Prevents SARS-CoV-2 Replication , 2020, bioRxiv.

[10]  Ruth R. Montgomery,et al.  Single-cell longitudinal analysis of SARS-CoV-2 infection in human airway epithelium , 2020, bioRxiv.

[11]  M. Schwartz,et al.  The coding capacity of SARS-CoV-2 , 2020, Nature.

[12]  Fang Li,et al.  Cell entry mechanisms of SARS-CoV-2 , 2020, Proceedings of the National Academy of Sciences.

[13]  M. Alexander,et al.  Bulk and single-cell gene expression profiling of SARS-CoV-2 infected human cell lines identifies molecular targets for therapeutic intervention , 2020, bioRxiv.

[14]  R. Schwartz,et al.  Imbalanced Host Response to SARS-CoV-2 Drives Development of COVID-19 , 2020, Cell.

[15]  Benjamin J. Polacco,et al.  A SARS-CoV-2 Protein Interaction Map Reveals Targets for Drug-Repurposing , 2020, Nature.

[16]  M. Tay,et al.  The trinity of COVID-19: immunity, inflammation and intervention , 2020, Nature Reviews Immunology.

[17]  Dimitry Tegunov,et al.  Structure of replicating SARS-CoV-2 polymerase , 2020, Nature.

[18]  R. Hromas,et al.  Structural Basis of RNA Cap Modification by SARS-CoV-2 Coronavirus , 2020, bioRxiv.

[19]  Natacha S. Ogando,et al.  SARS-coronavirus-2 replication in Vero E6 cells: replication kinetics, rapid adaptation and cytopathology , 2020, bioRxiv.

[20]  Xingjun Guo,et al.  Long Noncoding RNA RMRP Suppresses the Tumorigenesis of Hepatocellular Carcinoma Through Targeting microRNA-766 , 2020, OncoTargets and therapy.

[21]  Fumihiro Kato,et al.  Enhanced isolation of SARS-CoV-2 by TMPRSS2-expressing cells , 2020, Proceedings of the National Academy of Sciences.

[22]  K. Yuen,et al.  Clinical Characteristics of Coronavirus Disease 2019 in China , 2020, The New England journal of medicine.

[23]  E. Holmes,et al.  A new coronavirus associated with human respiratory disease in China , 2020, Nature.

[24]  S. Quake,et al.  Cell-Surface Proteomic Profiling in the Fly Brain Uncovers Wiring Regulators , 2019, Cell.

[25]  Hyun-Cheol Lee,et al.  Intracellular sensing of viral genomes and viral evasion , 2019, Experimental & Molecular Medicine.

[26]  P. Masters,et al.  Coronavirus genomic RNA packaging , 2019, Virology.

[27]  Xuetao Cao,et al.  Nuclear hnRNPA2B1 initiates and amplifies the innate immune response to DNA viruses , 2019, Science.

[28]  S. Carr,et al.  TMT Labeling for the Masses: A Robust and Cost-efficient, In-solution Labeling Approach. , 2019, Molecular & cellular proteomics : MCP.

[29]  D. Kennedy,et al.  G3BP1 and G3BP2 regulate translation of interferon-stimulated genes: IFITM1, IFITM2 and IFITM3 in the cancer cell line MCF7 , 2019, Molecular and Cellular Biochemistry.

[30]  Ryan A. Flynn,et al.  An RNA-Centric Dissection of Host Complexes Controlling Flavivirus Infection , 2019, Nature Microbiology.

[31]  P. Khavari,et al.  Methods to study RNA–protein interactions , 2019, Nature Methods.

[32]  K. Lam,et al.  The stress granule protein G3BP1 binds viral dsRNA and RIG-I to enhance interferon-β response , 2019, The Journal of Biological Chemistry.

[33]  S. Goff,et al.  Regulation of HIV-1 Gag-Pol Expression by Shiftless, an Inhibitor of Programmed -1 Ribosomal Frameshifting , 2019, Cell.

[34]  Jeroen Krijgsveld,et al.  The Human RNA-Binding Proteome and Its Dynamics during Translational Arrest , 2019, Cell.

[35]  Damian Szklarczyk,et al.  STRING v11: protein–protein association networks with increased coverage, supporting functional discovery in genome-wide experimental datasets , 2018, Nucleic Acids Res..

[36]  M. Heller,et al.  Determination of host proteins composing the microenvironment of coronavirus replicase complexes by proximity-labeling , 2018, eLife.

[37]  W. Xue,et al.  G3BP1 promotes DNA binding and activation of cGAS , 2018, Nature Immunology.

[38]  Andrew J. Olive,et al.  CNBP controls IL-12 gene transcription and Th1 immunity , 2018, The Journal of experimental medicine.

[39]  Krishna Shankara Narayanan,et al.  Interplay between coronavirus, a cytoplasmic RNA virus, and nonsense-mediated mRNA decay pathway , 2018, Proceedings of the National Academy of Sciences.

[40]  E. Lander,et al.  The NORAD lncRNA assembles a topoisomerase complex critical for genome stability , 2018, Nature.

[41]  Alfredo Castello,et al.  Unconventional RNA‐binding proteins step into the virus–host battlefront , 2018, Wiley interdisciplinary reviews. RNA.

[42]  Ronald J. Moore,et al.  Reproducible workflow for multiplexed deep-scale proteome and phosphoproteome analysis of tumor tissues by liquid chromatography–mass spectrometry , 2018, Nature Protocols.

[43]  B. Tabak,et al.  Higher-Order Inter-chromosomal Hubs Shape 3D Genome Organization in the Nucleus , 2018, Cell.

[44]  Guangdi Li,et al.  NLRX1 Mediates MAVS Degradation To Attenuate the Hepatitis C Virus-Induced Innate Immune Response through PCBP2 , 2017, Journal of Virology.

[45]  Richard E. Randall,et al.  Human interactome of the influenza B virus NS1 protein , 2017, The Journal of general virology.

[46]  Wei Wu,et al.  Cellular RNA Helicase DDX1 Is Involved in Transmissible Gastroenteritis Virus nsp14-Induced Interferon-Beta Production , 2017, Front. Immunol..

[47]  Sungwook Lee,et al.  STRAP positively regulates TLR3-triggered signaling pathway. , 2017, Cellular immunology.

[48]  P. Collins,et al.  A novel host factor for human respiratory syncytial virus , 2017, Communicative & integrative biology.

[49]  A. Fullam,et al.  DDX3 directly regulates TRAF3 ubiquitination and acts as a scaffold to coordinate assembly of signalling 2 complexes downstream of MAVS , 2019 .

[50]  J. Kim,et al.  CNBP acts as a key transcriptional regulator of sustained expression of interleukin-6 , 2017, Nucleic acids research.

[51]  Eugen C. Buehler,et al.  Actin-Related Protein 2 (ARP2) and Virus-Induced Filopodia Facilitate Human Respiratory Syncytial Virus Spread , 2016, PLoS pathogens.

[52]  J. Ziebuhr,et al.  The Nonstructural Proteins Directing Coronavirus RNA Synthesis and Processing , 2016, Advances in Virus Research.

[53]  M. Gack,et al.  Viral evasion of intracellular DNA and RNA sensing , 2016, Nature Reviews Microbiology.

[54]  V. Hornung,et al.  MOV10 Provides Antiviral Activity against RNA Viruses by Enhancing RIG-I–MAVS-Independent IFN Induction , 2016, The Journal of Immunology.

[55]  Gene W. Yeo,et al.  Robust transcriptome-wide discovery of RNA binding protein binding sites with enhanced CLIP (eCLIP) , 2016, Nature Methods.

[56]  Jun Liu,et al.  Host Protein Moloney Leukemia Virus 10 (MOV10) Acts as a Restriction Factor of Influenza A Virus by Inhibiting the Nuclear Import of the Viral Nucleoprotein , 2016, Journal of Virology.

[57]  Ching Hua Lee,et al.  Characterization of RyDEN (C19orf66) as an Interferon-Stimulated Cellular Inhibitor against Dengue Virus Replication , 2016, PLoS pathogens.

[58]  Jeroen Krijgsveld,et al.  The RNA-binding proteomes from yeast to man harbour conserved enigmRBPs , 2015, Nature Communications.

[59]  Qiangfeng Cliff Zhang,et al.  Systematic Discovery of Xist RNA Binding Proteins , 2015, Cell.

[60]  Michael J. Sweredoski,et al.  The Xist lncRNA directly interacts with SHARP to silence transcription through HDAC3 , 2015, Nature.

[61]  Huifeng Jiao,et al.  14-3-3ζ regulates immune response through Stat3 signaling in oral squamous cell carcinoma. , 2015, Molecules and cells.

[62]  N. Sonenberg,et al.  Targeting the eIF4F translation initiation complex: a critical nexus for cancer development. , 2015, Cancer research.

[63]  R. Chen,et al.  LSM14A inhibits porcine reproductive and respiratory syndrome virus (PRRSV) replication by activating IFN-β signaling pathway in Marc-145 , 2014, Molecular and Cellular Biochemistry.

[64]  M. Yoneyama,et al.  A Novel Function of Human Pumilio Proteins in Cytoplasmic Sensing of Viral Infection , 2014, PLoS pathogens.

[65]  Sharon R Grossman,et al.  RNA-RNA Interactions Enable Specific Targeting of Noncoding RNAs to Nascent Pre-mRNAs and Chromatin Sites , 2014, Cell.

[66]  M. Garcia-Blanco,et al.  G3BP1, G3BP2 and CAPRIN1 Are Required for Translation of Interferon Stimulated mRNAs and Are Targeted by a Dengue Virus Non-coding RNA , 2014, PLoS pathogens.

[67]  S. Biswas,et al.  Annexin-A1 Regulates TLR-Mediated IFN-β Production through an Interaction with TANK-Binding Kinase 1 , 2013, The Journal of Immunology.

[68]  B. Neuman,et al.  Severe Acute Respiratory Syndrome Coronavirus Nonstructural Proteins 3, 4, and 6 Induce Double-Membrane Vesicles , 2013, mBio.

[69]  E. Lander,et al.  The Xist lncRNA Exploits Three-Dimensional Genome Architecture to Spread Across the X Chromosome , 2013, Science.

[70]  K. Kuroiwa,et al.  YB-1 suppression induces STAT3 proteolysis and sensitizes renal cancer to interferon-α , 2013, Cancer Immunology, Immunotherapy.

[71]  P. Chatterjee,et al.  Heterotrimeric GAIT Complex Drives Transcript-Selective Translation Inhibition in Murine Macrophages , 2012, Molecular and Cellular Biology.

[72]  Q. Mei,et al.  Cyclophilin A and viral infections , 2012, Biochemical and Biophysical Research Communications.

[73]  H. Shu,et al.  LSm14A is a processing body-associated sensor of viral nucleic acids that initiates cellular antiviral response in the early phase of viral infection , 2012, Proceedings of the National Academy of Sciences of the United States of America.

[74]  Guangchuang Yu,et al.  clusterProfiler: an R package for comparing biological themes among gene clusters. , 2012, Omics : a journal of integrative biology.

[75]  M. V. van Hemert,et al.  Cyclosporin A inhibits the replication of diverse coronaviruses. , 2011, The Journal of general virology.

[76]  Xiaozhong Peng,et al.  PCBP2 Enhances the Antiviral Activity of IFN-α against HCV by Stabilizing the mRNA of STAT1 and STAT2 , 2011, PloS one.

[77]  Christian Drosten,et al.  The SARS-Coronavirus-Host Interactome: Identification of Cyclophilins as Target for Pan-Coronavirus Inhibitors , 2011, PLoS pathogens.

[78]  G. Cheng,et al.  DDX1, DDX21, and DHX36 helicases form a complex with the adaptor molecule TRIF to sense dsRNA in dendritic cells. , 2011, Immunity.

[79]  Helga Thorvaldsdóttir,et al.  Molecular signatures database (MSigDB) 3.0 , 2011, Bioinform..

[80]  G. Kozlov,et al.  La-Related Protein 4 Binds Poly(A), Interacts with the Poly(A)-Binding Protein MLLE Domain via a Variant PAM2w Motif, and Can Promote mRNA Stability , 2010, Molecular and Cellular Biology.

[81]  Narasimhan J. Venkatachari,et al.  P Body-Associated Protein Mov10 Inhibits HIV-1 Replication at Multiple Stages , 2010, Journal of Virology.

[82]  D. Liu,et al.  The Cellular RNA Helicase DDX1 Interacts with Coronavirus Nonstructural Protein 14 and Enhances Viral Replication , 2010, Journal of Virology.

[83]  T. Seya,et al.  DEAD/H BOX 3 (DDX3) helicase binds the RIG‐I adaptor IPS‐1 to up‐regulate IFN‐β‐inducing potential , 2010, European journal of immunology.

[84]  Richard Durbin,et al.  Sequence analysis Fast and accurate short read alignment with Burrows – Wheeler transform , 2009 .

[85]  L. Platanias,et al.  Interferon-Dependent Engagement of Eukaryotic Initiation Factor 4B via S6 Kinase (S6K)- and Ribosomal Protein S6K-Mediated Signals , 2009, Molecular and Cellular Biology.

[86]  Clifford A. Meyer,et al.  Model-based Analysis of ChIP-Seq (MACS) , 2008, Genome Biology.

[87]  A. Bowie,et al.  Viral targeting of DEAD box protein 3 reveals its role in TBK1/IKKε‐mediated IRF activation , 2008, The EMBO journal.

[88]  A. Bauch,et al.  The DEAD-box helicase DDX3X is a critical component of the TANK-binding kinase 1-dependent innate immune response , 2008, The EMBO journal.

[89]  J. Krijnse-Locker,et al.  Modification of intracellular membrane structures for virus replication , 2008, Nature Reviews Microbiology.

[90]  J. Zavadil,et al.  eIF4GI links nutrient sensing by mTOR to cell proliferation and inhibition of autophagy , 2008, The Journal of cell biology.

[91]  Margaret A. Johnson,et al.  Proteomics Analysis Unravels the Functional Repertoire of Coronavirus Nonstructural Protein 3 , 2008, Journal of Virology.

[92]  Brad T. Sherman,et al.  Systematic and integrative analysis of large gene lists using DAVID bioinformatics resources , 2008, Nature Protocols.

[93]  V. Chow,et al.  The 3a accessory protein of SARS coronavirus specifically interacts with the 5'UTR of its genomic RNA, Using a unique 75 amino acid interaction domain. , 2007, Biochemistry.

[94]  Brian Raught,et al.  The mTOR/PI3K and MAPK pathways converge on eIF4B to control its phosphorylation and activity , 2006, The EMBO journal.

[95]  Xiaolei Yin,et al.  Highly infectious SARS-CoV pseudotyped virus reveals the cell tropism and its correlation with receptor expression , 2004, Biochemical and Biophysical Research Communications.

[96]  David I Stuart,et al.  The nsp9 Replicase Protein of SARS-Coronavirus, Structure and Functional Insights , 2004, Structure.

[97]  M. Sherman,et al.  A Preliminary Report , 1953 .

[98]  Gerhard Klimeck Crystal Structure , 1924, Nature.