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In Situ Cryo-ET Redefines HPV Disassembly and Transport Paradigms

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Human papillomavirus is a necessary cause of invasive cervical cancer worldwide
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Variations in the age‐specific curves of human papillomavirus prevalence in women worldwide
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Worldwide burden of cancer attributable to HPV by site, country and HPV type
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UCSF ChimeraX: Meeting modern challenges in visualization and analysis
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ArtiaX : An electron tomography toolbox for the interactive handling of s ub‐tomograms in UCSF ChimeraX
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Computer Visualization of Three-Dimensional Image Data Using IMOD
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Dual-Axis Tomography: An Approach with Alignment Methods That Preserve Resolution
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Site-Specific Cryo-focused Ion Beam Sample Preparation Guided by 3D Correlative Microscopy
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ARF6 Is an Actionable Node that Orchestrates Oncogenic GNAQ Signaling in Uveal Melanoma
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Structural Basis for Cargo Regulation of COPII Coat Assembly
inferredcites →https://doi.org/10.1016/j.cell.2008.12.025openalex-snapshot
TANGO1 Facilitates Cargo Loading at Endoplasmic Reticulum Exit Sites
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Cell-Penetrating Peptide Mediates Intracellular Membrane Passage of Human Papillomavirus L2 Protein to Trigger Retrograde Trafficking
inferredcites →https://doi.org/10.1016/j.cell.2020.10.039openalex-snapshot
β-Coronaviruses Use Lysosomes for Egress Instead of the Biosynthetic Secretory Pathway
inferredcites →https://doi.org/10.1016/j.celrep.2018.12.041openalex-snapshot
Proteomic Profiling of Mammalian COPII and COPI Vesicles
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A Link between ER Tethering and COP-I Vesicle Uncoating
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Cutaneous wart-associated HPV types: Prevalence and relation with patient characteristics
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Automated electron microscope tomography using robust prediction of specimen movements
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CTFFIND4: Fast and accurate defocus estimation from electron micrographs
inferredcites →https://doi.org/10.1016/j.jsb.2016.07.010openalex-snapshot
Optimized cryo-focused ion beam sample preparation aimed at in situ structural studies of membrane proteins
inferredcites →https://doi.org/10.1016/j.jsb.2016.07.011openalex-snapshot
Automated tilt series alignment and tomographic reconstruction in IMOD
inferredcites →https://doi.org/10.1016/j.jsb.2019.09.006openalex-snapshot
Mind the gap: Micro-expansion joints drastically decrease the bending of FIB-milled cryo-lamellae
inferredcites →https://doi.org/10.1016/j.jsb.2023.108057openalex-snapshot
Accurate, automatic determination of astigmatism and phase with Ctfplotter in IMOD
inferredcites →https://doi.org/10.1016/j.lanmic.2024.100955openalex-snapshot
Global prevalence of cervical human papillomavirus in women aged 50 years and older with normal cytology: a systematic review and meta-analysis
inferredcites →https://doi.org/10.1016/j.str.2016.12.001openalex-snapshot
Cryoelectron Microscopy Maps of Human Papillomavirus 16 Reveal L2 Densities and Heparin Binding Site
inferredcites →https://doi.org/10.1016/j.virol.2003.11.004openalex-snapshot
Kinetics of in vitro adsorption and entry of papillomavirus virions
inferredcites →https://doi.org/10.1016/j.virol.2004.03.033openalex-snapshot
Classification of papillomaviruses
inferredcites →https://doi.org/10.1016/j.virol.2010.02.002openalex-snapshot
Classification of papillomaviruses (PVs) based on 189 PV types and proposal of taxonomic amendments
inferredcites →https://doi.org/10.1016/j.virol.2010.09.002openalex-snapshot
Inhibition of gamma secretase blocks HPV infection
inferredcites →https://doi.org/10.1016/j.virol.2014.05.005openalex-snapshot
Human papillomavirus infection requires the TSG101 component of the ESCRT machinery
inferredcites →https://doi.org/10.1016/j.ypmed.2020.106399openalex-snapshot
HPV vaccination introduction worldwide and WHO and UNICEF estimates of national HPV immunization coverage 2010–2019
inferredcites →https://doi.org/10.1016/s0006-3495(91)82181-6openalex-snapshot
Structures of bovine and human papillomaviruses. Analysis by cryoelectron microscopy and three-dimensional image reconstruction
inferredcites →https://doi.org/10.1016/s0042-6822(02)00143-5openalex-snapshot
Papillomaviruses infect cells via a clathrin-dependent pathway
inferredcites →https://doi.org/10.1016/s0140-6736(10)62342-2openalex-snapshot
Incidence and clearance of genital human papillomavirus infection in men (HIM): a cohort study
inferredcites →https://doi.org/10.1016/s1097-2765(00)80449-9openalex-snapshot
Structure of Small Virus-like Particles Assembled from the L1 Protein of Human Papillomavirus 16
inferredcites →https://doi.org/10.1021/acs.biochem.7b00706openalex-snapshot
Family-wide Analysis of the Inhibition of Arf Guanine Nucleotide Exchange Factors with Small Molecules: Evidence of Unique Inhibitory Profiles
inferredcites →https://doi.org/10.1038/emboj.2013.171openalex-snapshot
Autophagy sequesters damaged lysosomes to control lysosomal biogenesis and kidney injury
inferredcites →https://doi.org/10.1038/nature02108openalex-snapshot
Lipid packing sensed by ArfGAP1 couples COPI coat disassembly to membrane bilayer curvature
inferredcites →https://doi.org/10.1038/ncomms6317openalex-snapshot
Clearance of persistent HPV infection and cervical lesion by therapeutic DNA vaccine in CIN3 patients
inferredcites →https://doi.org/10.1038/nmeth.2019openalex-snapshot
Fiji: an open-source platform for biological-image analysis
inferredcites →https://doi.org/10.1038/nmeth.4169openalex-snapshot
cryoSPARC: algorithms for rapid unsupervised cryo-EM structure determination
inferredcites →https://doi.org/10.1038/nsb0597-413openalex-snapshot
Novel structural features of bovine papillomavirus capsid revealed by a three-dimensional reconstruction to 9 Å resolution
inferredcites →https://doi.org/10.1038/s41467-020-18952-1openalex-snapshot
Topaz-Denoise: general deep denoising models for cryoEM and cryoET
inferredcites →https://doi.org/10.1038/s41467-021-26225-8openalex-snapshot
PGRMC1 acts as a size-selective cargo receptor to drive ER-phagic clearance of mutant prohormones
inferredcites →https://doi.org/10.1038/s41467-023-35874-wopenalex-snapshot
Master mitotic kinases regulate viral genome delivery during papillomavirus cell entry
inferredcites →https://doi.org/10.1038/s41586-024-07487-wopenalex-snapshot
Accurate structure prediction of biomolecular interactions with AlphaFold 3
inferredcites →https://doi.org/10.1038/s41592-019-0580-yopenalex-snapshot
Real-time cryo-electron microscopy data preprocessing with Warp
inferredcites →https://doi.org/10.1038/s41592-019-0630-5openalex-snapshot
Tailoring cryo-electron microscopy grids by photo-micropatterning for in-cell structural studies
inferredcites →https://doi.org/10.1038/s41592-023-02053-0openalex-snapshot
Genetically encoded multimeric tags for subcellular protein localization in cryo-EM
inferredcites →https://doi.org/10.1038/s41594-024-01413-4openalex-snapshot
Cryo-electron tomography reveals how COPII assembles on cargo-containing membranes
inferredcites →https://doi.org/10.1038/s41598-018-30051-2openalex-snapshot
Heterotetrameric annexin A2/S100A10 (A2t) is essential for oncogenic human papillomavirus trafficking and capsid disassembly, and protects virions from lysosomal degradation
inferredcites →https://doi.org/10.1038/s41598-021-83076-5openalex-snapshot
High resolution cryo EM analysis of HPV16 identifies minor structural protein L2 and describes capsid flexibility
inferredcites →https://doi.org/10.1038/srep32337openalex-snapshot
The CD63-Syntenin-1 Complex Controls Post-Endocytic Trafficking of Oncogenic Human Papillomaviruses
inferredcites →https://doi.org/10.1038/srep45159openalex-snapshot
The VPS4 component of the ESCRT machinery plays an essential role in HPV infectious entry and capsid disassembly
inferredcites →https://doi.org/10.1073/pnas.0404229101openalex-snapshot
Establishment of papillomavirus infection is enhanced by promyelocytic leukemia protein (PML) expression
inferredcites →https://doi.org/10.1073/pnas.0504020102openalex-snapshot
Production of infectious human papillomavirus independently of viral replication and epithelial cell differentiation
inferredcites →https://doi.org/10.1073/pnas.0508815103openalex-snapshot
Cleavage of the papillomavirus minor capsid protein, L2, at a furin consensus site is necessary for infection
inferredcites →https://doi.org/10.1073/pnas.0600447103openalex-snapshot
Selective small-molecule inhibitor reveals critical mitotic functions of human CDK1
inferredcites →https://doi.org/10.1073/pnas.0908502106openalex-snapshot
The initial steps leading to papillomavirus infection occur on the basement membrane prior to cell surface binding
inferredcites →https://doi.org/10.1073/pnas.1302164110openalex-snapshot
Genome-wide siRNA screen identifies the retromer as a cellular entry factor for human papillomavirus
inferredcites →https://doi.org/10.1073/pnas.1600638113openalex-snapshot
Incoming human papillomavirus type 16 genome resides in a vesicular compartment throughout mitosis
inferredcites →https://doi.org/10.1073/pnas.1917748117openalex-snapshot
Cell-penetrating peptide inhibits retromer-mediated human papillomavirus trafficking during virus entry
inferredcites →https://doi.org/10.1073/pnas.2307721120openalex-snapshot
Sequence-independent activity of a predicted long disordered segment of the human papillomavirus type 16 L2 capsid protein during virus entry
inferredcites →https://doi.org/10.1073/pnas.75.7.3327openalex-snapshot
Fluorescence probe measurement of the intralysosomal pH in living cells and the perturbation of pH by various agents.
inferredcites →https://doi.org/10.1073/pnas.87.1.83openalex-snapshot
Mechanism of L-leucyl-L-leucine methyl ester-mediated killing of cytotoxic lymphocytes: dependence on a lysosomal thiol protease, dipeptidyl peptidase I, that is enriched in these cells.
inferredcites →https://doi.org/10.1073/pnas.96.4.1224openalex-snapshot
Receptor-induced polymerization of coatomer
inferredcites →https://doi.org/10.1074/jbc.270.52.31364openalex-snapshot
In Vitro Assembly and Disassembly of Coatomer
inferredcites →https://doi.org/10.1080/15548627.2015.1063871openalex-snapshot
Sensitive detection of lysosomal membrane permeabilization by lysosomal galectin puncta assay
inferredcites →https://doi.org/10.1080/21645515.2015.1088616openalex-snapshot
Persistent infection with human papillomavirus 16 or 18 is strongly linked with high-grade cervical disease
inferredcites →https://doi.org/10.1083/jcb.123.6.1365openalex-snapshot
Hydrolysis of bound GTP by ARF protein triggers uncoating of Golgi-derived COP-coated vesicles.
inferredcites →https://doi.org/10.1083/jcb.201709162openalex-snapshot
Membrane protein recycling from the vacuole/lysosome membrane
inferredcites →https://doi.org/10.1083/jcb.201804171openalex-snapshot
γ-Secretase promotes membrane insertion of the human papillomavirus L2 capsid protein during virus infection
inferredcites →https://doi.org/10.1093/cid/ciab985openalex-snapshot
Proportion of Incident Genital Human Papillomavirus Detections not Attributable to Transmission and Potentially Attributable to Latent Infections: Implications for Cervical Cancer Screening
inferredcites →https://doi.org/10.1093/emboj/cdf494openalex-snapshot
Atomic model of the papillomavirus capsid
inferredcites →https://doi.org/10.1093/infdis/jis660openalex-snapshot
A Cohort Effect of the Sexual Revolution May Be Masking an Increase in Human Papillomavirus Detection at Menopause in the United States
inferredcites →https://doi.org/10.1093/jnci/92.9.709openalex-snapshot
Evidence for a Causal Association Between Human Papillomavirus and a Subset of Head and Neck Cancers
inferredcites →https://doi.org/10.1097/aog.0b013e31818b0df2openalex-snapshot
Prevalence of High-Risk Human Papillomavirus Among Older Women
inferredcites →https://doi.org/10.1097/olq.0000000000000193openalex-snapshot
The Estimated Lifetime Probability of Acquiring Human Papillomavirus in the United States
inferredcites →https://doi.org/10.1097/olq.0000000000001356openalex-snapshot
Estimated Prevalence and Incidence of Disease-Associated Human Papillomavirus Types Among 15- to 59-Year-Olds in the United States
inferredcites →https://doi.org/10.1099/jgv.0.000734openalex-snapshot
Characterizing the spatio-temporal role of sorting nexin 17 in human papillomavirus trafficking
inferredcites →https://doi.org/10.1101/2024.01.05.574336openalex-snapshot
MemBrain v2: an end-to-end tool for the analysis of membranes in cryo-electron tomography
inferredcites →https://doi.org/10.1111/j.1600-0854.2007.00697.xopenalex-snapshot
A Conformational Change in the α‐subunit of Coatomer Induced by Ligand Binding to γ‐COP Revealed by Single‐pair FRET
inferredcites →https://doi.org/10.1111/j.1600-0854.2011.01320.xopenalex-snapshot
Human Papillomavirus L2 Facilitates Viral Escape from Late Endosomes via Sorting Nexin 17
inferredcites →https://doi.org/10.1126/sciadv.abh4276openalex-snapshot
Retromer stabilizes transient membrane insertion of L2 capsid protein during retrograde entry of human papillomavirus
inferredcites →https://doi.org/10.1126/sciadv.adc9830openalex-snapshot
HPV is a cargo for the COPI sorting complex during virus entry
inferredcites →https://doi.org/10.1126/sciadv.adz6792openalex-snapshot
The nuclear import receptor importin-7 targets HPV from the Golgi to the nucleus to promote infection
inferredcites →https://doi.org/10.1126/science.1215909openalex-snapshot
ER Cargo Properties Specify a Requirement for COPII Coat Rigidity Mediated by Sec13p
inferredcites →https://doi.org/10.1126/science.1221443openalex-snapshot
The Structures of COPI-Coated Vesicles Reveal Alternate Coatomer Conformations and Interactions
inferredcites →https://doi.org/10.1126/science.274.5291.1396openalex-snapshot
Gordon Research Conference: 1997 Meeting Schedule
inferredcites →https://doi.org/10.1126/science.aab1121openalex-snapshot
A structure of the COPI coat and the role of coat proteins in membrane vesicle assembly
inferredcites →https://doi.org/10.1126/science.aan6298openalex-snapshot
Lysosomal metabolomics reveals V-ATPase- and mTOR-dependent regulation of amino acid efflux from lysosomes
inferredcites →https://doi.org/10.1126/science.aar5078openalex-snapshot
Triggered recruitment of ESCRT machinery promotes endolysosomal repair
inferredcites →https://doi.org/10.1128/aac.02284-13openalex-snapshot
Inhibition by Cellular Vacuolar ATPase Impairs Human Papillomavirus Uncoating and Infection
inferredcites →https://doi.org/10.1128/jvi.00519-13openalex-snapshot
Annexin A2 and S100A10 Regulate Human Papillomavirus Type 16 Entry and Intracellular Trafficking in Human Keratinocytes
inferredcites →https://doi.org/10.1128/jvi.00776-06openalex-snapshot
The L2 Minor Capsid Protein of Low-Risk Human Papillomavirus Type 11 Interacts with Host Nuclear Import Receptors and Viral DNA
inferredcites →https://doi.org/10.1128/jvi.00980-12openalex-snapshot
Cyclophilins Facilitate Dissociation of the Human Papillomavirus Type 16 Capsid Protein L1 from the L2/DNA Complex following Virus Entry
inferredcites →https://doi.org/10.1128/jvi.01014-08openalex-snapshot
Caveolin-1-Dependent Infectious Entry of Human Papillomavirus Type 31 in Human Keratinocytes Proceeds to the Endosomal Pathway for pH-Dependent Uncoating
inferredcites →https://doi.org/10.1128/jvi.01081-10openalex-snapshot
Papillomavirus Infection Requires γ Secretase
inferredcites →https://doi.org/10.1128/jvi.01183-15openalex-snapshot
A Dual Role for the Nonreceptor Tyrosine Kinase Pyk2 during the Intracellular Trafficking of Human Papillomavirus 16
inferredcites →https://doi.org/10.1128/jvi.01499-15openalex-snapshot
A Novel PDZ Domain Interaction Mediates the Binding between Human Papillomavirus 16 L2 and Sorting Nexin 27 and Modulates Virion Trafficking
inferredcites →https://doi.org/10.1128/jvi.01539-12openalex-snapshot
A Transmembrane Domain and GxxxG Motifs within L2 Are Essential for Papillomavirus Infection
inferredcites →https://doi.org/10.1128/jvi.01588-15openalex-snapshot
Topography of the Human Papillomavirus Minor Capsid Protein L2 during Vesicular Trafficking of Infectious Entry
inferredcites →https://doi.org/10.1128/jvi.01631-08openalex-snapshot
Heparan Sulfate-Independent Cell Binding and Infection with Furin-Precleaved Papillomavirus Capsids
inferredcites →https://doi.org/10.1128/jvi.01721-13openalex-snapshot
Multiple Heparan Sulfate Binding Site Engagements Are Required for the Infectious Entry of Human Papillomavirus Type 16
inferredcites →https://doi.org/10.1128/jvi.01991-12openalex-snapshot
SNX17 Facilitates Infection with Diverse Papillomavirus Types
inferredcites →https://doi.org/10.1128/jvi.02726-07openalex-snapshot
Arrangement of L2 within the Papillomavirus Capsid
inferredcites →https://doi.org/10.1128/jvi.03222-12openalex-snapshot
Identification of a Role for the trans -Golgi Network in Human Papillomavirus 16 Pseudovirus Infection
inferredcites →https://doi.org/10.1128/jvi.05493-11openalex-snapshot
Opposing Effects of Bacitracin on Human Papillomavirus Type 16 Infection: Enhancement of Binding and Entry and Inhibition of Endosomal Penetration
inferredcites →https://doi.org/10.1128/jvi.05708-11openalex-snapshot
Cathepsin Cleavage Potentiates the Ebola Virus Glycoprotein To Undergo a Subsequent Fusion-Relevant Conformational Change
inferredcites →https://doi.org/10.1128/jvi.15.5.1239-1247.1975openalex-snapshot
Structural polypeptides of rabbit, bovine, and human papillomaviruses
inferredcites →https://doi.org/10.1128/jvi.75.3.1565-1570.2001openalex-snapshot
Human Papillomavirus Infection Requires Cell Surface Heparan Sulfate
inferredcites →https://doi.org/10.1128/jvi.78.2.751-757.2004openalex-snapshot
Efficient Intracellular Assembly of Papillomaviral Vectors
inferredcites →https://doi.org/10.1128/jvi.78.24.13447-13454.2004openalex-snapshot
The Positively Charged Termini of L2 Minor Capsid Protein Required for Bovine Papillomavirus Infection Function Separately in Nuclear Import and DNA Binding
inferredcites →https://doi.org/10.1128/jvi.79.5.2839-2846.2005openalex-snapshot
Maturation of Papillomavirus Capsids
inferredcites →https://doi.org/10.1128/jvi.80.2.759-768.2006openalex-snapshot
A Membrane-Destabilizing Peptide in Capsid Protein L2 Is Required for Egress of Papillomavirus Genomes from Endosomes
inferredcites →https://doi.org/10.1128/mbio.01777-14openalex-snapshot
Vesicular Trafficking of Incoming Human Papillomavirus 16 to the Golgi Apparatus and Endoplasmic Reticulum Requires γ-Secretase Activity
inferredcites →https://doi.org/10.1136/bmjopen-2021-052016openalex-snapshot
Estimate of global human papillomavirus vaccination coverage: analysis of country-level indicators
inferredcites →https://doi.org/10.1158/0008-5472.can-06-4095openalex-snapshot
Proton Pump Inhibitors Induce Apoptosis of Human B-Cell Tumors through a Caspase-Independent Mechanism Involving Reactive Oxygen Species
inferredcites →https://doi.org/10.1158/0008-5472.can-12-2635openalex-snapshot
Contributions of Recent and Past Sexual Partnerships on Incident Human Papillomavirus Detection: Acquisition and Reactivation in Older Women
inferredcites →https://doi.org/10.1186/1471-2334-14-312openalex-snapshot
The potential impact of HPV-16 reactivation on prevalence in older Australians
inferredcites →https://doi.org/10.1186/s13027-022-00453-zopenalex-snapshot
Incidence of oncogenic HPV and HPV-related dysplasia five years after a negative HPV test by self-sampling in elderly women
inferredcites →https://doi.org/10.1371/journal.pone.0003313openalex-snapshot
Clathrin- and Caveolin-Independent Entry of Human Papillomavirus Type 16—Involvement of Tetraspanin-Enriched Microdomains (TEMs)
inferredcites →https://doi.org/10.1371/journal.pone.0004463openalex-snapshot
Two Highly Conserved Cysteine Residues in HPV16 L2 Form an Intramolecular Disulfide Bond and Are Critical for Infectivity in Human Keratinocytes
inferredcites →https://doi.org/10.1371/journal.pone.0170158openalex-snapshot
The Role of DCT in HPV16 Infection of HaCaTs
inferredcites →https://doi.org/10.1371/journal.ppat.1000148openalex-snapshot
Human Papillomavirus Type 16 Entry: Retrograde Cell Surface Transport along Actin-Rich Protrusions
inferredcites →https://doi.org/10.1371/journal.ppat.1000318openalex-snapshot
Establishment of Human Papillomavirus Infection Requires Cell Cycle Progression
inferredcites →https://doi.org/10.1371/journal.ppat.1002657openalex-snapshot
Entry of Human Papillomavirus Type 16 by Actin-Dependent, Clathrin- and Lipid Raft-Independent Endocytosis
inferredcites →https://doi.org/10.1371/journal.ppat.1004162openalex-snapshot
Large Scale RNAi Reveals the Requirement of Nuclear Envelope Breakdown for Nuclear Import of Human Papillomaviruses
inferredcites →https://doi.org/10.1371/journal.ppat.1004699openalex-snapshot
Direct Binding of Retromer to Human Papillomavirus Type 16 Minor Capsid Protein L2 Mediates Endosome Exit during Viral Infection
inferredcites →https://doi.org/10.1371/journal.ppat.1006200openalex-snapshot
Translocation of the papillomavirus L2/vDNA complex across the limiting membrane requires the onset of mitosis
inferredcites →https://doi.org/10.1371/journal.ppat.1008946openalex-snapshot
p120 catenin recruits HPV to γ-secretase to promote virus infection
inferredcites →https://doi.org/10.1371/journal.ppat.1012289openalex-snapshot
The BICD2 dynein cargo adaptor binds to the HPV16 L2 capsid protein and promotes HPV infection
inferredcites →https://doi.org/10.1385/1-59259-982-6:445openalex-snapshot
Generation of HPV Pseudovirions Using Transfection and Their Use in Neutralization Assays
inferredcites →https://doi.org/10.1515/bc.2010.062openalex-snapshot
Molecular characterisation of ‘transmembrane protein 192’ (TMEM192), a novel protein of the lysosomal membrane
inferredcites →https://doi.org/10.15620/cdc:114751openalex-snapshot
Cancers Associated with Human Papillomavirus, United States—2014–2018
inferredcites →https://doi.org/10.3322/caac.21660openalex-snapshot
Global Cancer Statistics 2020: GLOBOCAN Estimates of Incidence and Mortality Worldwide for 36 Cancers in 185 Countries
inferredcites →https://doi.org/10.3389/fmicb.2023.1294146openalex-snapshot
Multiple HPV integration mode in the cell lines based on long-reads sequencing
inferredcites →https://doi.org/10.3390/v14040804openalex-snapshot
Human Papillomavirus L2 Capsid Protein Stabilizes γ-Secretase during Viral Infection
inferredcites →https://doi.org/10.5281/zenodo.13982246openalex-snapshot
warpem/warp: v2.0.0dev0
inferredcites →https://doi.org/10.7554/elife.26691openalex-snapshot
9Å structure of the COPI coat reveals that the Arf1 GTPase occupies two contrasting molecular environments
inferredcites →https://doi.org/10.7554/elife.32493openalex-snapshot
The structure of the COPI coat determined within the cell

What points at this work

Incoming claims: where this work appears in other papers’ reference lists and arguments.

inferred← citeshttps://doi.org/10.3390/v18070790openalex-snapshot
Labeling and Localization Strategies for In Situ Cryo-Electron Tomography Across the Viral Life Cycle
inferred← citeshttps://doi.org/10.64898/2026.08.18.745536openalex-snapshot
The human papillomavirus L2 protein in the capsid is an ensemble of related structures poised to exit the virus particle
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