Affinage

PCNA

DNA sliding clamp PCNA · UniProt P12004

Length
261 aa
Mass
28.8 kDa
Annotated
2026-06-10
100 papers in source corpus 43 papers cited in narrative 43 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PCNA is the central sliding-clamp processivity factor of eukaryotic DNA replication and a mobile recruitment platform that coordinates replication with repair, chromatin assembly, and cell-cycle control (PMID:9038370, PMID:28071730). It forms a toroidal homotrimer that encircles duplex DNA and slides bidirectionally via a 'cogwheel' mechanism, in which basic residues lining the ring channel track the DNA backbone through short-lived polar interactions while the clamp maintains an invariant orientation (PMID:9038370, PMID:28071730). PCNA is loaded onto primed template-primer junctions by the RFC clamp loader in a stepwise, ATP-dependent reaction and is unloaded by the ATAD5/Elg1 RFC-like complex, with single-molecule analyses defining distinct loading (two intermediate) and unloading (one intermediate) pathways (PMID:15210332, PMID:31160570). Beyond tethering DNA polymerase delta, PCNA accelerates its catalytic rate of nucleotide incorporation more than tenfold through a defined face of the clamp (PMID:30605530). The outer face of the ring serves as a docking hub for a large set of partners that bind through PIP-box motifs—typically embedded in intrinsically disordered regions and tuned over orders of magnitude by flanking charges—or through the distinct APIM motif, allowing PCNA to organize Okazaki fragment maturation (a FEN1–DNA Ligase 1 toolbelt with substrate handoff), mismatch repair (hMSH6/hMSH3 of MutSalpha/beta and activation of MutLalpha and MutLgamma endonucleases), end resection (Exo1), translesion synthesis (Rev1/Pol eta toolbelts and bridges), and replication-coupled chromatin assembly via CAF-1 (PMID:11274057, PMID:23939618, PMID:27325737, PMID:31134302, PMID:32814343, PMID:36539424, PMID:11089978, PMID:19736315). PCNA function is governed by an extensive set of post-translational modifications: RAD6/RAD18-mediated K164 monoubiquitination (bridged by SIVA1, recruited via RPA-filament translocation, and reversed by USP1) switches the clamp toward translesion synthesis; DNA-loading-stimulated SUMOylation, EGFR-driven Tyr211 phosphorylation, CBP/p300 acetylation, and EZH2-mediated K110 methylation further tune partner selection, chromatin retention, and clamp stability (PMID:17105346, PMID:24958773, PMID:33242956, PMID:36228090, PMID:18701921, PMID:17115032, PMID:24939902, PMID:30071900). Chromatin-bound PCNA additionally serves as the degron-display platform for the CRL4(Cdt2) ligase, which ubiquitinates Cdt1, p21, and Set8 exclusively on chromatin to prevent rereplication (PMID:21828267). A hypomorphic p.Ser228Ile mutation that selectively disrupts PCNA's interactions with FEN1 and DNA Ligase 1 causes a human DNA repair disorder with defective UV survival (PMID:24911150).

Mechanistic history

Synthesis pass · year-by-year structured walk · 24 steps
  1. 1995 Medium

    Establishing that the stress-response protein Gadd45 binds PCNA directly mapped early evidence that PCNA serves as a docking surface for regulators beyond the replication machinery.

    Evidence Co-IP, yeast two-hybrid, and deletion/peptide mapping of Gadd45-PCNA contacts

    PMID:7784094

    Open questions at the time
    • Functional consequence of the interaction for repair not directly demonstrated
    • Stoichiometry inferred from peptide ELISA
  2. 1997 High

    Defining PCNA as a toroidal homotrimer encircling DNA explained how it confers processivity on replicative polymerases, the founding mechanistic model.

    Evidence Crystallographic and biochemical synthesis of clamp structure and Pol delta/epsilon tethering

    PMID:9038370

    Open questions at the time
    • Did not resolve atomic basis of DNA sliding
    • Did not address catalytic vs processivity contributions
  3. 2001 High

    Identifying the PIP-box as the conserved motif by which hMSH6/hMSH3 bind PCNA established the generalizable docking principle linking PCNA to mismatch repair.

    Evidence Co-IP, in vitro binding, MMR complementation, and replication-foci co-localization with PIP-box mutagenesis

    PMID:11274057

    Open questions at the time
    • Did not define how PCNA orientation couples MMR to the nascent strand
    • Did not address modification-dependent partner switching
  4. 2003 High

    Showing that archaeal PCNA subunits contact polymerase, ligase, and FEN1 at defined positions revealed how a single clamp can architecturally couple synthesis to Okazaki fragment maturation.

    Evidence Reconstitution with defined heterotrimeric PCNA subunit combinations and activity assays

    PMID:12535540

    Open questions at the time
    • Homotrimeric eukaryotic PCNA cannot encode fixed subunit identity
    • Did not establish ordered handoff dynamics
  5. 2004 High

    Dissecting RFC-mediated, ATP-dependent loading of PCNA defined how the clamp is opened and placed onto primer-template junctions.

    Evidence In vitro reconstitution with ATP analogs and stepwise mechanistic dissection

    PMID:15210332

    Open questions at the time
    • Did not identify the dedicated unloader
    • Kinetics of intermediate states not resolved
  6. 2005 High

    Identifying Ctf18-RFC as an RPA-favored PCNA unloader, and FRAP showing rapid exchange at forks but long retention at damage, established that clamp residence is actively regulated and links retention to ubiquitination.

    Evidence In vitro loading/unloading assays; live-cell FRAP with NER-deficient and K164R PCNA mutant lines

    PMID:15964801 PMID:16227586

    Open questions at the time
    • Quantitative coupling of unloading to replication completion not resolved
    • Relationship between Ctf18-RFC and ATAD5/Elg1 unloaders unclear
  7. 2005 Medium

    Demonstrating that two PCNA trimers form a back-to-back doublet able to bind CAF-1 and Pol delta simultaneously offered a structural route to coordinating replication with chromatin assembly.

    Evidence Purified-protein and extract analysis with Arg5/Lys110 mutagenesis and peptide competition

    PMID:15805117

    Open questions at the time
    • In vivo prevalence of the double trimer not established
    • Single lab, novel structural model
  8. 2006 High

    Linking K164 monoubiquitination to error-prone Ig hypermutation and identifying EGFR-driven Tyr211 phosphorylation that stabilizes chromatin-bound PCNA showed that PTMs reprogram clamp function.

    Evidence DT40 PCNA-K164R/Rad18 genetics with hypermutation readout; in vivo phosphorylation, kinase assays, and chromatin fractionation

    PMID:17105346 PMID:17115032

    Open questions at the time
    • Mechanism of TLS polymerase recruitment by ubiquitin not structurally resolved
    • Tyr211-P downstream partner effects undefined at this stage
  9. 2008 Medium

    Connecting Chk1/Claspin/Timeless and RNF8 to damage-induced PCNA ubiquitination, and showing DNA loading stimulates SUMOylation, defined the upstream regulatory circuitry shaping PCNA modification.

    Evidence siRNA epistasis and ChIP; in vitro and in vivo SUMOylation assays with PCNA mutants

    PMID:18451105 PMID:18701921 PMID:18948756

    Open questions at the time
    • Pathway placement relied on knockdown without full reconstitution
    • Relative contributions of RAD18 vs RNF8 to monoubiquitination unresolved
  10. 2009 High

    Discovering Lys107 ubiquitination triggered by DNA ligase I deficiency (via Mms2/Ubc4/Rad5) and the APIM motif as a non-PIP docking mode expanded both the modification repertoire and the partner-recruitment vocabulary of PCNA.

    Evidence Yeast genetics and E2/E3 epistasis with human confirmation; APIM identification and functional verification across multiple proteins

    PMID:19736315 PMID:20010813

    Open questions at the time
    • Structural basis of K107-ubiquitin signaling not defined
    • Generality of APIM versus PIP usage incompletely resolved
  11. 2011 High

    Establishing that CRL4(Cdt2) ubiquitinates Cdt1, p21, and Set8 only when their degrons engage chromatin-bound PCNA showed how the clamp couples proteolysis to ongoing DNA synthesis to block rereplication.

    Evidence Reconstituted ubiquitylation, degron mapping, genetic epistasis, and fractionation

    PMID:21828267

    Open questions at the time
    • Did not define how degron-PCNA timing is enforced
    • Substrate spectrum may be incomplete
  12. 2013 High

    Showing PCNA loads at double-strand breaks and confers processivity on Exo1 generalized the processivity-factor concept from synthesis to resection.

    Evidence Mammalian cell and Xenopus extract assays with purified-protein reconstitution of direct PCNA-Exo1 interaction

    PMID:23939618

    Open questions at the time
    • How PCNA is loaded at breaks versus forks not fully defined
    • Regulation of Exo1 disengagement unclear
  13. 2014 High

    Mapping CBP/p300 acetylation that triggers post-NER PCNA removal/degradation, and a human S228I mutation selectively disrupting FEN1/Ligase 1 binding, tied specific clamp surfaces and PTMs to repair outcomes and human disease.

    Evidence In vitro acetylation, lysine mutagenesis, chromatin fractionation, UV survival; patient-cell interaction and RNA-synthesis recovery assays

    PMID:24911150 PMID:24939902

    Open questions at the time
    • Acetyl-reader/eraser machinery not defined
    • Genotype-phenotype breadth of PCNA repair disorder limited to studied family
  14. 2014 High

    Identifying SIVA1 as a constitutive PIP-box bridge directing RAD18 onto PCNA clarified how monoubiquitination is targeted to the clamp.

    Evidence PCNA-complex affinity purification, co-IP, siRNA, and Pol eta focus assays

    PMID:24958773

    Open questions at the time
    • Whether SIVA1 acts at all damage types unclear
    • Structure of the SIVA1-RAD18-PCNA assembly not resolved
  15. 2015 Medium

    Demonstrating that Tyr211 phosphorylation alters MutSalpha/beta binding and blocks MutLalpha activation, and that PCNA scaffolds cytosolic MAPK/PI3K signaling, broadened PCNA's regulatory reach into fidelity control and non-nuclear signaling.

    Evidence In vitro MMR and endonuclease activation assays; MS of PCNA pulldowns with APIM-peptide functional readouts

    PMID:25797046 PMID:25825764

    Open questions at the time
    • Cytosolic signaling role rests on peptide competition, mechanism indirect
    • Physiological extent of cytosolic PCNA unclear
  16. 2016 High

    Establishing PIP-box-dependent recruitment of FANCM, TRAIP, HUWE1, and SDE2, plus single-molecule TLS toolbelt/bridge architectures, revealed PCNA as the organizing hub for replication-stress and lesion-bypass responses.

    Evidence Co-IP/PIP-box mutagenesis with fiber and traverse assays; single-molecule TIRF of Rev1/Pol eta-PCNA complexes; SDE2 cleavage and CRL4CDT2 assays

    PMID:26825464 PMID:27146073 PMID:27325737 PMID:27462463 PMID:27906959

    Open questions at the time
    • Hierarchy among competing PIP-box partners at a single clamp unresolved
    • Several interaction-domain mappings remain partial
  17. 2017 High

    Resolving the cogwheel sliding mechanism explained how PCNA tracks the DNA backbone while staying fixed in orientation, and linked the DNA interface to initiation of Pol delta synthesis.

    Evidence X-ray crystallography, NMR, MD simulations, and interface mutagenesis with Pol delta assays

    PMID:28071730

    Open questions at the time
    • How sliding accommodates bound partners not modeled
    • Dynamics during loading not directly observed
  18. 2018 Medium

    Showing EZH2-mediated Lys110 dimethylation stabilizes the trimer and promotes Pol delta binding added a methylation layer to clamp stability and replication competence.

    Evidence Co-IP, in vitro methyltransferase, trimer stability, and Pol delta binding assays with K110 mutagenesis

    PMID:30071900

    Open questions at the time
    • Single lab, novel modification requires replication
    • Interplay with K110 doublet-contact role from prior work unclear
  19. 2019 High

    Defining ATAD5-RLC as the principal human PCNA unloader (able to remove ubiquitinated PCNA), with BRD4 acting as a chromatin-acetylation-gated brake, established controlled clamp removal as a regulated step in replication-coupled chromatin maturation.

    Evidence In vitro and single-molecule loading/unloading with ATPase mutants; BRD4-ATAD5 co-IP, domain mapping, and chromatin fractionation; quench-flow kinetics for catalytic acceleration

    PMID:29440488 PMID:30605530 PMID:31160570 PMID:31875566

    Open questions at the time
    • How unloading is timed to fork completion in vivo not fully resolved
    • Crosstalk between BRD4 brake and PCNA modifications undefined
  20. 2019 Medium

    Discovering surface-expressed PCNA acting as an inhibitory ligand for NK NKp44 identified an unexpected immune-checkpoint role beyond the nucleus.

    Evidence Surface staining, chimeric NKp44 binding inhibition, NK cytotoxicity/IFNg assays, and PDX experiments

    PMID:31164357

    Open questions at the time
    • Mechanism of PCNA surface trafficking unknown
    • Relationship to nuclear PCNA pool undefined
  21. 2019 High

    Quantifying that PIP-box affinity is tuned over four orders of magnitude by flanking charges within disordered regions explained how a single clamp surface can prioritize among dozens of partners.

    Evidence NMR and affinity measurements across 77 confirmed PCNA-binding proteins with computational analysis

    PMID:31134302

    Open questions at the time
    • Did not establish in vivo competition outcomes
    • PTM effects on affinity hierarchy not directly measured
  22. 2020 High

    Showing PCNA/RFC activate MutLgamma to bias meiotic crossover resolution, and that RPA-filament translocation by Rad6/Rad18 enhances PCNA monoubiquitination, extended the clamp's activator and signaling roles into meiosis and lesion tolerance.

    Evidence In vitro endonuclease reconstitution with yeast genetics and meiotic cytology; single-molecule FRET of Rad6/Rad18 on RPA

    PMID:32814343 PMID:33242956

    Open questions at the time
    • How PCNA selects crossover sites in vivo unresolved
    • Coupling of Rad18 translocation to specific PCNA monomers undefined
  23. 2022 High

    Cryo-EM of PCNA-Lig1 and PCNA-FEN1-Lig1 complexes resolved how dual Lig1 PIP motifs and an unoccupied PCNA monomer drive FEN1-to-Lig1 substrate handoff during Okazaki fragment maturation.

    Evidence Cryo-EM structures with PIP-motif mutagenesis and ligation assays

    PMID:36539424

    Open questions at the time
    • Order of polymerase engagement before maturation not captured
    • Does not address competing partner displacement
  24. 2023 High

    Establishing that USP1 reverses K164 mono- and polyubiquitination, and that USP1-inhibitor toxicity depends on aberrant ubiquitinated-PCNA processing, defined a deubiquitination axis with therapeutic relevance.

    Evidence Genome-wide CRISPR screens, RAD18/UBE2K knockouts, and PCNA K164R rescue with S-phase damage readouts

    PMID:36228090

    Open questions at the time
    • Whether other DUBs act on PCNA undefined
    • Mechanism linking ubiquitin accumulation to reduced total PCNA unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the dozens of PIP-box/APIM partners and the layered PTM code are integrated in real time at a single clamp to dictate pathway choice remains unresolved.
  • No unified model of competitive partner exchange on chromatin-bound PCNA in vivo
  • PTM crosstalk hierarchy (ubiquitin/SUMO/acetyl/methyl/phospho) not quantitatively ordered
  • Trafficking and function of cell-surface PCNA mechanistically uncharacterized

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 4 GO:0003677 DNA binding 2 GO:0098772 molecular function regulator activity 2 GO:0001618 virus receptor activity 1
Localization
GO:0000228 nuclear chromosome 3 GO:0005634 nucleus 2 GO:0005654 nucleoplasm 1 GO:0005886 plasma membrane 1
Pathway
R-HSA-73894 DNA Repair 5 R-HSA-69306 DNA Replication 4 R-HSA-168256 Immune System 2 R-HSA-4839726 Chromatin organization 2 R-HSA-1474165 Reproduction 1 R-HSA-1640170 Cell Cycle 1
Partners
EXO1FANCMFEN1LIG1MSH6RAD18SIVA1USP1
Complex memberships
PCNA homotrimerPCNA-FEN1-DNA Ligase 1 toolbeltPCNA-Rev1-Pol eta TLS toolbelt

Evidence

Reading pass · 43 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1997 PCNA forms a toroidal (ring-shaped) homotrimeric structure that encircles double-stranded DNA and can slide bidirectionally along the duplex, functioning as the processivity factor for DNA polymerase delta and epsilon by tethering the polymerase catalytic unit to the DNA template. Structural and biochemical studies; review of crystallographic and functional data Oncogene High 9038370
2000 PCNA mutations that reduce binding to chromatin assembly factor 1 (CAF-1) in vitro also alter chromatin association of the CAF-1 large subunit in vivo, demonstrating that PCNA participates in inheritance of epigenetic chromatin structures during S phase through at least two mechanisms including CAF-1 recruitment. Yeast genetics (silencing assays with PCNA mutants pol30-6, pol30-8, pol30-79), in vitro binding assays, chromatin association assays Nature High 11089978
2001 PCNA binds directly to hMSH6 and hMSH3 subunits of the human mismatch repair factors hMutSα and hMutSβ via a conserved PIP-box motif (Qxx[LI]xx[FF]) in the N-terminal domains; deletion of this motif abolishes PCNA interaction in vitro and eliminates mismatch repair activity in hMSH6-deficient cells, and hMSH6/hMSH3 co-localize with PCNA at replication foci. Co-immunoprecipitation, in vitro binding assays, mismatch repair complementation assays, immunofluorescence co-localization Genes & development High 11274057
2003 The archaeal PCNA from Sulfolobus solfataricus is a heterotrimer of three distinct subunits (PCNA1, 2, 3) that assembles in a defined orientation; distinct PCNA subunits contact DNA polymerase, DNA ligase I, and FEN1 respectively, imposing defined architecture at the lagging strand fork and coupling DNA synthesis with Okazaki fragment maturation. Biochemical reconstitution, activity assays with individual PCNA subunits and heterotrimers, protein interaction mapping Molecular cell High 12535540
2004 PCNA is loaded onto primed DNA template-primer junctions in an ATP-dependent process by the heteropentameric RFC complex; loading is mechanistically distinct from prokaryotic clamp loaders and involves multiple stepwise ATP-binding events. Biochemical reconstitution, ATP hydrolysis assays, mechanistic dissection of RFC-PCNA loading Progress in nucleic acid research and molecular biology High 15210332
2004 XRCC1 physically interacts with PCNA; the interaction is mediated by residues 166–310 of XRCC1, and XRCC1 co-localizes with PCNA at DNA replication foci exclusively in S phase, suggesting PCNA sequesters XRCC1 to replication factories to facilitate single-strand break repair. Co-immunoprecipitation, FRET analysis, in vitro binding with XRCC1 deletion mutants, immunofluorescence co-localization Nucleic acids research High 15107487
2005 The Ctf18-RFC complex functions as an efficient PCNA unloader in an ATP hydrolysis-dependent manner; unloading is strongly favored over loading when the ssDNA template is coated by RPA (but not mutant RPA or heterologous SSB), separating PCNA unloading from the sister chromatid cohesion function of Ctf18-RFC. In vitro reconstitution with purified proteins, PCNA loading/unloading assays, ATP hydrolysis assays Molecular and cellular biology High 15964801
2005 PCNA dynamics at replication foci and UV-damaged sites differ: PCNA at replication foci shows rapid exchange between bound and freely mobile nucleoplasmic pools, whereas PCNA at UV-damaged sites has a significantly longer residence time. Initial PCNA recruitment to damaged sites is dependent on nucleotide excision repair, and a ubiquitination-deficient PCNA mutant (K164R) shows shorter residence time at damaged areas, linking ubiquitination to PCNA retention at damage. Live cell imaging with GFP-hPCNA, FRAP (fluorescence recovery after photobleaching), local UV irradiation, use of NER-deficient and PCNA ubiquitination mutant cell lines Molecular and cellular biology High 16227586
2006 PCNA is phosphorylated on Tyr211 by nuclear EGFR; this phosphorylation stabilizes chromatin-bound PCNA protein and is required for maintaining PCNA function on chromatin. In vivo phosphorylation assays, kinase activity assays, chromatin fractionation, mutation analysis, nuclear EGFR co-immunoprecipitation Nature cell biology High 17115032
2006 PCNA ubiquitination at K164 (triggered by Rad18) promotes error-prone immunoglobulin somatic hypermutation in vertebrate B cells; PCNA(K164R) mutation and Rad18 deficiency both strongly reduce AID-dependent single-nucleotide substitutions at the Ig light-chain locus, indicating the PCNA-ubiquitin pathway is exploited for hypermutation through recruitment of error-prone DNA polymerases. Genetic analysis of DT40 B cell lines with PCNA K164R and Rad18 mutations, Ig hypermutation assays PLoS biology High 17105346
2008 Sumoylation of yeast PCNA by the ligase Siz1 is strongly stimulated by loading of PCNA onto DNA; DNA binding by PCNA itself (rather than by Siz1) accounts for the stimulatory effect, indicating that a conformational or property change in PCNA upon DNA loading enhances its capacity to be sumoylated. In vitro sumoylation assays with purified components, in vivo sumoylation assays in S. cerevisiae, PCNA mutant analysis The EMBO journal High 18701921
2008 Chk1, independently of ATR, regulates DNA damage-induced PCNA monoubiquitination by stabilizing Claspin, which in turn regulates Rad18 binding to chromatin; Timeless (a Claspin-associated protein) is also required for efficient PCNA ubiquitination. siRNA knockdown of Chk1, Claspin, and Timeless; chromatin immunoprecipitation; epistasis analysis of Rad18 chromatin binding Genes & development Medium 18451105
2008 RNF8 can monoubiquitinate PCNA with UbcH5c and polyubiquitinate PCNA with Ubc13/Uev1a in vitro; RNF8 depletion suppresses UV- and MNNG-stimulated PCNA monoubiquitination in vivo, revealing an RNF8-dependent pathway for PCNA ubiquitination in the DNA damage tolerance response. In vitro ubiquitination assays, siRNA depletion of RNF8, in vivo PCNA ubiquitination analysis after DNA damage Cell cycle (Georgetown, Tex.) Medium 18948756
2009 DNA ligase I deficiency in S. cerevisiae triggers PCNA ubiquitylation at Lys107 (distinct from the damage-induced Lys164 site); this modification requires E2 Mms2/Ubc4 and E3 Rad5, not Rad6/Rad18. Cells with PCNAK107R mutation are inviable in a ligase I-deficient background, demonstrating that Lys107 ubiquitylation is essential for the DNA damage response to Okazaki fragment ligation defects. Analogous PCNA ubiquitylation in response to DNA ligase I deficiency is conserved in humans. Yeast genetic analysis, in vivo ubiquitylation assays, epistasis with E2/E3 mutants, human cell experiments Nature cell biology High 20010813
2009 hABH2 (AlkB homologue 2) interacts with post-translationally modified PCNA via a novel PCNA-interacting motif termed APIM (AlkB homologue 2 PCNA-interacting motif), distinct from the canonical PIP-box; APIM is present in >200 other proteins involved in DNA maintenance and verified functional in at least five of them. Co-immunoprecipitation, pulldown assays, identification and functional verification of APIM in multiple proteins, cellular sensitivity assays with APIM-peptide The Journal of cell biology Medium 19736315
2011 CRL4(Cdt2) E3 ubiquitin ligase couples proteolysis to DNA synthesis via display of substrate degrons on chromatin-bound PCNA; substrates including Cdt1, p21, and Set8 are recruited through PCNA interactions and ubiquitinated exclusively on chromatin, thereby preventing rereplication. Biochemical reconstitution of ubiquitylation, substrate degron mapping, genetic epistasis, fractionation Genes & development High 21828267
2013 PCNA loads onto DNA double-strand breaks after damage and promotes processive DNA end resection by Exo1 through direct interaction; PCNA tethers Exo1 to the DNA substrate to confer processivity, analogous to its processivity-factor role for DNA polymerases. Mammalian cell assays, Xenopus nuclear extracts, purified protein reconstitution, direct PCNA-Exo1 interaction assays Nucleic acids research High 23939618
2014 CBP (and less efficiently p300) acetylates PCNA at Lys13, Lys14, Lys77, and Lys80; this acetylation promotes removal of chromatin-bound PCNA and its proteasomal degradation after nucleotide excision repair (NER). Mutation of these residues impairs DNA repair, increases UV sensitivity, and prevents proteolytic degradation of PCNA after DNA damage. In vitro acetylation assays with CBP/p300, mutagenesis of PCNA lysine residues, chromatin fractionation, proteasome inhibition experiments, UV sensitivity assays Nucleic acids research High 24939902
2014 A hypomorphic PCNA missense mutation (p.Ser228Ile) in humans does not affect protein levels or DNA replication but profoundly alters PCNA's interaction with FEN1 and DNA Ligase 1, causing marked defects in UV survival and RNA synthesis recovery after UV irradiation, defining a novel human DNA repair disorder. Patient cell studies, protein interaction assays (PCNA-FEN1, PCNA-Ligase1), UV survival and RNA synthesis recovery assays, complementation analysis The Journal of clinical investigation High 24911150
2014 SIVA1 constitutively interacts with PCNA via a PIP motif and serves as a molecular bridge between RAD18 and PCNA, targeting RAD18 E3 ligase activity onto PCNA for monoubiquitination; SIVA1 knockdown compromises RAD18-dependent PCNA monoubiquitination and Pol eta focus formation. Affinity purification of PCNA-containing complexes, co-immunoprecipitation, siRNA knockdown, focus formation assays, UV sensitivity assays The Journal of cell biology High 24958773
2015 EGFR-mediated phosphorylation of PCNA at Tyr211 alters its interaction with mismatch recognition proteins MutSα and MutSβ and interferes with PCNA-dependent activation of MutLα endonuclease, thereby inhibiting mismatch repair at the initiation step and inducing nucleotide misincorporation during DNA synthesis. In vitro mismatch repair assays, PCNA phosphorylation assays, protein interaction assays with MutSα/MutSβ, MutLα endonuclease activation assays, nucleotide incorporation fidelity assays Proceedings of the National Academy of Sciences of the United States of America High 25825764
2016 FANCM interacts with PCNA via a conserved PIP-box, and this interaction is strongly stimulated by replication stress; a FANCM PIP-box mutant is defective in promoting replication traverse of DNA interstrand crosslinks and FANCD2 monoubiquitination, indicating PCNA interaction is essential for FANCM to aid replication machines at ICLs. Co-immunoprecipitation, in vitro binding assays, replication traverse assays, FANCD2 ubiquitination assays, PIP-box mutagenesis Nucleic acids research High 26825464
2016 HUWE1 (HECT-type ubiquitin ligase) interacts with PCNA at stalled replication forks; HUWE1 knockout cells fail to mitigate replication stress, and this function requires its interaction with PCNA. HUWE1 also mono-ubiquitinates H2AX to promote signaling at stalled forks. Co-immunoprecipitation, HUWE1 knockout cell analysis, replication stress assays (fiber assays), DNA damage signaling assays EMBO reports Medium 27146073
2016 During translesion synthesis, PCNA and two TLS polymerases (Rev1 and Pol η) form ternary complexes with two architectures: PCNA 'tool belts' (both polymerases simultaneously bound to one PCNA ring) and 'Rev1 bridges'; these architectures are dynamic and interconvert without dissociation, facilitating polymerase selection and switching. Single-molecule total internal reflection fluorescence (TIRF) microscopy, assembly/disassembly kinetics of ternary complexes Nucleic acids research High 27325737
2016 PCNA unloading by the Elg1-RFC-like complex (yeast homolog of ATAD5-RLC) is important for coordinating DNA replication forks with replication-coupled nucleosome assembly to maintain transcriptional silencing of HML and HMR heterochromatin through S phase. Yeast genetics, silencing assays, epistasis between Elg1 and histone chaperone mutants Proceedings of the National Academy of Sciences of the United States of America Medium 29440488
2017 Human PCNA slides on DNA via a 'cogwheel' mechanism: basic residues in the ring channel form a right-hand spiral matching the pitch of B-DNA, and PCNA tracks the DNA backbone through short-lived polar interactions while maintaining invariant orientation relative to DNA. Mutations at the PCNA-DNA interface impair initiation of DNA synthesis by Pol δ. X-ray crystallography, NMR spectroscopy, molecular dynamics simulations, mutagenesis with Pol δ activity assays Nature communications High 28071730
2018 EZH2 interacts with PCNA via a PIP box and dimethylates PCNA at Lys110; this dimethylation stabilizes the PCNA trimer and is required for efficient binding of DNA polymerase δ to PCNA, thereby promoting DNA replication. Co-immunoprecipitation, in vitro methyltransferase assays, PCNA trimer stability assays, Pol δ binding assays, site-directed mutagenesis of Lys110 Epigenetics & chromatin Medium 30071900
2019 PCNA stimulates the catalytic rate of DNA polymerase δ nucleotide incorporation by >10-fold (beyond its processivity function); a growth-defective PCNA mutant (DD41,42AA) shows substantially less stimulation of Pol δ nucleotide incorporation rate, identifying the specific face of PCNA important for catalytic acceleration. Quench-flow kinetic measurements, electrophoretic mobility shift assays, fluorescence anisotropy binding titrations, PCNA mutant analysis Nucleic acids research High 30605530
2019 ATAD5-RLC (human RFC-like complex) possesses potent PCNA-unloading activity; ATPase motif and collar domain of ATAD5 are crucial for unloading. ATAD5-RLC can unload ubiquitinated PCNA and does not require specific DNA structures. Single-molecule measurements reveal ATAD5-RLC unloads PCNA through one intermediate state before ATP hydrolysis, whereas RFC loads through two intermediate states. Biochemical PCNA loading/unloading assays, ATPase mutant analysis, single-molecule measurements, ubiquitinated PCNA assays Nature communications High 31160570
2019 BRD4 (a BET protein) inhibits PCNA unloading by ATAD5-RLC; the BRD4 ET domain interacts with a region upstream of the ATAD5 PCNA-unloading domain, and acetyl-histone-bound BRD4 on nascent chromatin prevents premature PCNA unloading. Disruption of BRD4-ATAD5 or BRD4-acetyl-histone interactions reduces chromatin-bound PCNA, while BRD4 overexpression increases it. Co-immunoprecipitation of BRD4 and ATAD5, in vitro PCNA unloading assays, chromatin fractionation, BRD4 overexpression/depletion, domain mapping Cell reports Medium 31875566
2019 PCNA-interacting motif affinity extends substantially beyond the canonical PIP-box boundaries; positive charges in flanking regions modulate PIP-box affinity over four orders of magnitude. Most PCNA-binding motifs reside in intrinsically disordered regions, and structure preformation is unrelated to binding affinity. NMR spectroscopy, affinity measurements (ITC/SPR), computational analysis of 77 confirmed PCNA-binding proteins Cellular and molecular life sciences : CMLS High 31134302
2020 PCNA and its loader RFC are sufficient to activate the MutLγ endonuclease in vitro, promoting meiotic crossover-biased resolution of double Holliday junctions; MutLγ is further stimulated by EXO1 and MutSγ co-dependently. PCNA localizes to prospective crossover sites along synapsed meiotic chromosomes, and RFC facilitates MutLγ-dependent crossing over in vivo in yeast. In vitro endonuclease assays with purified human proteins, yeast genetics (RFC mutants), immunostaining of meiotic chromosomes Nature High 32814343
2020 Rad6/Rad18 complex is recruited to RPA filaments on ssDNA exposed at translesion synthesis sites via Rad18·RPA interactions and randomly translocates along the filament; these translocations promote productive interactions with resident PCNA and significantly enhance PCNA monoubiquitination. Kinetic ubiquitination assays, single-molecule FRET microscopy to monitor Rad6/Rad18 dynamics on RPA filaments Biochemistry Medium 33242956
2022 Human DNA Ligase 1 (Lig1) recruits PCNA to nicked DNA using two PIP motifs: one at its disordered N-terminus (PIPN-term) and one at its DNA-binding domain (PIPDBD). After assembly of Lig1-PCNA as two-stack rings encircling DNA, PIPN-term is released and only PIPDBD is required for ligation. PCNA forms a toolbelt with FEN1 on nicked DNA and recruits Lig1 to an unoccupied PCNA monomer, driving substrate handoff from FEN1 to Lig1 during Okazaki fragment maturation. Cryo-EM structures of PCNA-Lig1 and PCNA-FEN1-Lig1 complexes, functional ligation assays, mutagenesis of PIP motifs Nature communications High 36539424
1995 Gadd45 interacts directly with PCNA; the N-terminal 94 amino acids of Gadd45 bind to PCNA, and deletion of the last 6 amino acids of PCNA abolishes this interaction. Gadd45 binds strongly to three regions of PCNA (residues 1–20, 61–80, and 196–215), with a potential stoichiometry of 2 Gadd45 molecules per PCNA monomer. Co-immunoprecipitation, yeast two-hybrid assay, deletion analysis of both Gadd45 and PCNA, peptide ELISA Oncogene Medium 7784094
1999 Cyclin D1 forms complexes with PCNA and CDK2 in senescent cells; excess GST-cyclin D1 inhibits DNA replication and CDK2 kinase activity in vitro, and overexpression of PCNA or CDK2 rescues the proliferation inhibition caused by cyclin D1 overexpression, demonstrating that cyclin D1 inhibits cell proliferation through physical interaction with PCNA and CDK2. Co-immunoprecipitation of cyclin D1 with PCNA and CDK2, in vitro DNA replication assays with GST-cyclin D1, rescue by PCNA/CDK2 overexpression Experimental cell research Medium 9925749
2002 Cdc25C interacts with PCNA transiently when cells begin to enter mitosis; the interaction was confirmed in vitro and in vivo by co-immunoprecipitation in Jurkat T cells, and immunofluorescence shows transient co-localization of Cdc25C and PCNA in the nucleus at the beginning of M phase. Yeast two-hybrid screen, co-immunoprecipitation in vitro and in vivo, immunofluorescence co-localization Oncogene Low 11896603
2005 Two PCNA homotrimers can form a back-to-back doublet in mammalian cell extracts and with purified human PCNA; Arg5 and Lys110 residues on the PCNA back side are contact points. A PCNA double trimer (but not a homotrimer alone) can simultaneously accommodate chromatin assembly factor 1 and polymerase delta. Cell extract and purified protein analysis, mutagenesis of Arg5 and Lys110, peptide inhibition of double trimer formation, CAF-1 and Pol δ binding assays The Journal of biological chemistry Medium 15805117
2016 Human SDE2 is regulated by PCNA interaction through a PIP box: SDE2 N-terminal UBL domain is cleaved at a diglycine motif via PIP-dependent deubiquitinating enzyme activity, and the cleaved SDE2 negatively regulates UV-damage-induced PCNA monoubiquitination. Cleaved SDE2 is degraded by CRL4CDT2 E3 ligase in a cell cycle- and DNA damage-dependent manner; failure to degrade SDE2 impairs S phase progression. Biochemical analysis of SDE2 cleavage, PCNA interaction assays (PIP-box mutagenesis), PCNA monoubiquitination assays after SDE2 manipulation, CRL4CDT2 ubiquitination assays, cell cycle analysis PLoS genetics Medium 27906959
2016 TRAIP (RNF206) interacts with PCNA via a conserved PIP box at its C-terminus and localizes to stalled replication forks; inactivation of TRAIP or disruption of its PCNA interaction compromises replication fork recovery and progression, leading to chromosome instability. Co-immunoprecipitation, PIP-box mutagenesis, replication fork assays (fiber assays), chromosome instability analysis Cell discovery Medium 27462463
2015 PCNA interacts with multiple cytosolic proteins in the MAPK and PI3K/Akt pathways as identified by mass spectrometry of PCNA pulldowns; treatment with APIM-containing peptide reduces Akt phosphorylation and TLR-mediated cytokine secretion, suggesting a platform role for PCNA in cytosolic signaling. Mass spectrometry of PCNA pulldowns, APIM-peptide treatment, Akt phosphorylation assays, cytokine secretion assays Cellular signalling Low 25797046
2019 PCNA is expressed on the surface of cancer cells (in addition to nuclear localization) and acts as an inhibitory ligand for the NK-cell receptor NKp44-isoform1, constituting an innate immune checkpoint; a blocking monoclonal antibody against surface PCNA enhances NK cell IFNγ release, cytotoxic activity, and inhibits tumor growth in PDX mouse models. FACS and ImageStream staining of cell surface PCNA, chimeric NKp44 receptor binding inhibition assays, NK killing and IFNγ ELISA assays, PDX mouse experiments Cancer immunology research Medium 31164357
2023 USP1 deubiquitinase acts on monoubiquitinated and polyubiquitinated PCNA (at K164 via RAD18 and UBE2K respectively); USP1 inhibition causes accumulation of ubiquitinated PCNA associated with reduced total PCNA protein levels and S-phase DNA damage. Ectopic expression of WT but not K164R PCNA reverses USP1 inhibitor sensitivity, establishing that USP1 dependency hinges on aberrant processing of ubiquitinated PCNA. CRISPR-Cas9 genome-wide screens, RAD18/UBE2K genetic knockouts, PCNA K164R rescue experiments, DNA synthesis assays, S-phase DNA damage analysis Molecular cancer therapeutics High 36228090

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 PCNA, the maestro of the replication fork. Cell 1461 17512402
2003 Proliferating cell nuclear antigen (PCNA): a dancer with many partners. Journal of cell science 882 12829735
1997 PCNA: structure, functions and interactions. Oncogene 745 9038370
2010 Proliferating cell nuclear antigen (PCNA): a key factor in DNA replication and cell cycle regulation. Annals of botany 568 21169293
2000 The puzzle of PCNA's many partners. BioEssays : news and reviews in molecular, cellular and developmental biology 376 11056476
2005 Nuclear dynamics of PCNA in DNA replication and repair. Molecular and cellular biology 345 16227586
1998 PCNA binding through a conserved motif. BioEssays : news and reviews in molecular, cellular and developmental biology 319 9631646
2013 Regulation of PCNA-protein interactions for genome stability. Nature reviews. Molecular cell biology 314 23594953
2001 Proliferating cell nuclear antigen (PCNA): ringmaster of the genome. International journal of radiation biology 278 11682006
2004 The PCNA-RFC families of DNA clamps and clamp loaders. Progress in nucleic acid research and molecular biology 271 15210332
2000 PCNA connects DNA replication to epigenetic inheritance in yeast. Nature 265 11089978
2017 Forging Ahead through Darkness: PCNA, Still the Principal Conductor at the Replication Fork. Molecular cell 263 28157503
2006 Tyrosine phosphorylation controls PCNA function through protein stability. Nature cell biology 254 17115032
2014 PCNA: a silent housekeeper or a potential therapeutic target? Trends in pharmacological sciences 238 24655521
2016 The Many Roles of PCNA in Eukaryotic DNA Replication. The Enzymes 232 27241932
2020 Human PCNA Structure, Function and Interactions. Biomolecules 229 32276417
1987 Cyclin (PCNA, auxiliary protein of DNA polymerase delta) is a central component of the pathway(s) leading to DNA replication and cell division. FEBS letters 203 2886367
2011 Mechanism of CRL4(Cdt2), a PCNA-dependent E3 ubiquitin ligase. Genes & development 195 21828267
2001 hMSH3 and hMSH6 interact with PCNA and colocalize with it to replication foci. Genes & development 195 11274057
1993 Toxicological and pathological applications of proliferating cell nuclear antigen (PCNA), a novel endogenous marker for cell proliferation. Critical reviews in toxicology 194 8097093
2003 A heterotrimeric PCNA in the hyperthermophilic archaeon Sulfolobus solfataricus. Molecular cell 183 12535540
2004 XRCC1 co-localizes and physically interacts with PCNA. Nucleic acids research 163 15107487
2009 Identification of a novel, widespread, and functionally important PCNA-binding motif. The Journal of cell biology 147 19736315
2006 A role for PCNA ubiquitination in immunoglobulin hypermutation. PLoS biology 138 17105346
2020 Targeting Proliferating Cell Nuclear Antigen (PCNA) as an Effective Strategy to Inhibit Tumor Cell Proliferation. Current cancer drug targets 133 31951183
1998 Protein-PCNA interactions: a DNA-scanning mechanism? Trends in biochemical sciences 132 9697409
2020 bioPROTACs as versatile modulators of intracellular therapeutic targets including proliferating cell nuclear antigen (PCNA). Proceedings of the National Academy of Sciences of the United States of America 131 32123106
1999 PCNA binding proteins. Frontiers in bioscience : a journal and virtual library 126 10577396
2005 Replication protein A-directed unloading of PCNA by the Ctf18 cohesion establishment complex. Molecular and cellular biology 116 15964801
1995 Characterisation of the interaction between PCNA and Gadd45. Oncogene 112 7784094
2004 Ubiquitination of PCNA and the polymerase switch in human cells. Cell cycle (Georgetown, Tex.) 106 15280666
2008 Chk1 and Claspin potentiate PCNA ubiquitination. Genes & development 94 18451105
2019 The PCNA interaction motifs revisited: thinking outside the PIP-box. Cellular and molecular life sciences : CMLS 92 31134302
2014 CBP and p300 acetylate PCNA to link its degradation with nucleotide excision repair synthesis. Nucleic acids research 88 24939902
2006 A novel function of DNA polymerase zeta regulated by PCNA. The EMBO journal 85 16957771
2019 Regulation of PCNA cycling on replicating DNA by RFC and RFC-like complexes. Nature communications 84 31160570
2020 PCNA activates the MutLγ endonuclease to promote meiotic crossing over. Nature 81 32814343
2012 PCNA structure and function: insights from structures of PCNA complexes and post-translationally modified PCNA. Sub-cellular biochemistry 80 22918591
2008 SUMO modification of PCNA is controlled by DNA. The EMBO journal 79 18701921
2008 PCNA is ubiquitinated by RNF8. Cell cycle (Georgetown, Tex.) 73 18948756
2017 Structural basis of human PCNA sliding on DNA. Nature communications 72 28071730
2014 Hypomorphic PCNA mutation underlies a human DNA repair disorder. The Journal of clinical investigation 70 24911150
2013 PCNA promotes processive DNA end resection by Exo1. Nucleic acids research 69 23939618
2008 The p21 and PCNA partnership: a new twist for an old plot. Cell cycle (Georgetown, Tex.) 69 19066467
2018 Maneuvers on PCNA Rings during DNA Replication and Repair. Genes 67 30126151
2000 Expression of proliferating cell nuclear antigen (PCNA) in the adult and developing mouse nervous system. Brain research. Molecular brain research 63 10891596
2023 Ubiquitinated PCNA Drives USP1 Synthetic Lethality in Cancer. Molecular cancer therapeutics 62 36228090
2009 Defects in DNA ligase I trigger PCNA ubiquitylation at Lys 107. Nature cell biology 61 20010813
2015 Phosphorylation of PCNA by EGFR inhibits mismatch repair and promotes misincorporation during DNA synthesis. Proceedings of the National Academy of Sciences of the United States of America 59 25825764
2012 CRL4(CDT2) targets CHK1 for PCNA-independent destruction. Molecular and cellular biology 59 23109433
2005 Proliferating cell nuclear antigen (PCNA) as a proliferative marker during embryonic and adult zebrafish hematopoiesis. Histochemistry and cell biology 59 16028068
1999 Cyclin D1 inhibits cell proliferation through binding to PCNA and cdk2. Experimental cell research 59 9925749
2002 Cdc25C interacts with PCNA at G2/M transition. Oncogene 54 11896603
2015 PCNA-interacting peptides reduce Akt phosphorylation and TLR-mediated cytokine secretion suggesting a role of PCNA in cellular signaling. Cellular signalling 53 25797046
2005 Sumoylation of PCNA: Wrestling with recombination at stalled replication forks. DNA repair 52 16368276
1999 Immunohistochemical staining of proliferating cell nuclear antigen (PCNA) in malignant and nonmalignant skin diseases. Archives of dermatological research 52 10482011
2016 HUWE1 interacts with PCNA to alleviate replication stress. EMBO reports 50 27146073
2013 Readers of PCNA modifications. Chromosoma 50 23580141
1995 Expression of proliferating cell nuclear antigen (PCNA) and Ki-67 in unicystic ameloblastoma. Histopathology 48 7797199
2016 PCNA tool belts and polymerase bridges form during translesion synthesis. Nucleic acids research 47 27325737
2011 Dynamic regulation of PCNA ubiquitylation/deubiquitylation. FEBS letters 47 21640107
2005 Proliferating cell nuclear antigen (PCNA) may function as a double homotrimer complex in the mammalian cell. The Journal of biological chemistry 47 15805117
2016 FANCM interacts with PCNA to promote replication traverse of DNA interstrand crosslinks. Nucleic acids research 44 26825464
2004 Esophagin and proliferating cell nuclear antigen (PCNA) are biomarkers of human esophageal neoplastic progression. International journal of cancer 43 15221970
1994 p53 and PCNA expression in carcinogenesis of the oropharyngeal mucosa. European journal of cancer. Part B, Oral oncology 43 7719225
2002 On the road to repair: PCNA encounters SUMO and ubiquitin modifications. Molecular cell 41 12408814
1990 Characterization and developmental expression of Xenopus proliferating cell nuclear antigen (PCNA). Developmental biology 41 1697269
2021 PCNA Loaders and Unloaders-One Ring That Rules Them All. Genes 40 34828416
2012 The clinical significance of autoantibodies to the proliferating cell nuclear antigen (PCNA). Autoimmunity reviews 40 22381917
2011 Ubiquitination of PCNA and its essential role in eukaryotic translesion synthesis. Cell biochemistry and biophysics 39 21461937
2006 Expression of survivin and correlation with PCNA in osteosarcoma. Journal of surgical oncology 39 16705726
2020 Control of DNA Damage Bypass by Ubiquitylation of PCNA. Genes 38 32013080
2009 Ubiquitination and deubiquitination of PCNA in response to stalling of the replication fork. Cell cycle (Georgetown, Tex.) 38 19221475
2000 Expression of p53 and PCNA in cholangiocarcinoma and primary sclerosing cholangitis. Modern pathology : an official journal of the United States and Canadian Academy of Pathology, Inc 38 11144921
2014 SIVA1 directs the E3 ubiquitin ligase RAD18 for PCNA monoubiquitination. The Journal of cell biology 37 24958773
2019 PCNA accelerates the nucleotide incorporation rate by DNA polymerase δ. Nucleic acids research 36 30605530
2018 Pivotal roles of PCNA loading and unloading in heterochromatin function. Proceedings of the National Academy of Sciences of the United States of America 36 29440488
2016 TRAIP regulates replication fork recovery and progression via PCNA. Cell discovery 36 27462463
1986 Immunochemical and biochemical analysis of the proliferating cell nuclear antigen (PCNA) in HeLa cells. Experimental cell research 36 2869966
2019 Inhibition of the NKp44-PCNA Immune Checkpoint Using a mAb to PCNA. Cancer immunology research 35 31164357
2016 PCNA-Dependent Cleavage and Degradation of SDE2 Regulates Response to Replication Stress. PLoS genetics 35 27906959
2011 The archaeal PCNA proteins. Biochemical Society transactions 35 21265741
2008 On the mechanism of loading the PCNA sliding clamp by RFC. Molecular microbiology 35 18312273
2020 DNA-damage tolerance through PCNA ubiquitination and sumoylation. The Biochemical journal 34 32726436
1992 PCNA expression in cutaneous keratinous neoplasms and verruca vulgaris. The American journal of pathology 34 1352943
1995 The significance of PCNA and p53 protein in some oral tumors. International journal of oral and maxillofacial surgery 33 7594756
2019 Targeting PCNA with Peptide Mimetics for Therapeutic Purposes. Chembiochem : a European journal of chemical biology 32 31247123
2016 Dynamics of plant DNA replication based on PCNA visualization. Scientific reports 32 27417498
1995 Proliferating cell nuclear antigen (PCNA) expression in oral squamous cell carcinomas. Journal of oral pathology & medicine : official publication of the International Association of Oral Pathologists and the American Academy of Oral Pathology 32 7473268
2019 Assessment of proliferating cell nuclear antigen (PCNA) expression at the invading front of oral squamous cell carcinoma. BMC oral health 31 31672124
2006 Analysis of proliferating cell nuclear antigen (PCNA) associated with DNA. Methods in molecular biology (Clifton, N.J.) 31 16673899
2019 PCNA Unloading Is Negatively Regulated by BET Proteins. Cell reports 29 31875566
2022 Mechanism of human Lig1 regulation by PCNA in Okazaki fragment sealing. Nature communications 28 36539424
2020 PCNA Monoubiquitination Is Regulated by Diffusion of Rad6/Rad18 Complexes along RPA Filaments. Biochemistry 27 33242956
2018 EZH2 promotes DNA replication by stabilizing interaction of POLδ and PCNA via methylation-mediated PCNA trimerization. Epigenetics & chromatin 27 30071900
1992 Proliferating cell nuclear antigen (PCNA) expression in Hodgkin's disease. The Journal of pathology 27 1360494
2018 Regulation of HLTF-mediated PCNA polyubiquitination by RFC and PCNA monoubiquitination levels determines choice of damage tolerance pathway. Nucleic acids research 26 30335157
2019 Spatiotemporal regulation of PCNA ubiquitination in damage tolerance pathways. Critical reviews in biochemistry and molecular biology 25 31736364
2011 9-1-1: PCNA's specialized cousin. Trends in biochemical sciences 25 21978893
1993 Proliferating cell nuclear antigen (PCNA) expression in chronic lymphocytic leukemia (CLL). Leukemia & lymphoma 24 8106065

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