Affinage

POLE4

DNA polymerase epsilon subunit 4 · UniProt Q9NR33

Round 2 corrected
Length
117 aa
Mass
12.2 kDa
Annotated
2026-04-28
128 papers in source corpus 5 papers cited in narrative 5 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

POLE4 is a histone-fold motif-containing accessory subunit of the DNA polymerase epsilon (Pol ε) holoenzyme that heterodimerizes with POLE3 to form a subcomplex essential for both DNA replication fidelity and replication-coupled chromatin maintenance. The POLE3–POLE4 dimer functions as a histone H3–H4 chaperone that promotes tetrasome formation and coordinates parental histone recycling and new histone deposition at the replication fork; its depletion impairs helicase unwinding and PCNA unloading from chromatin (PMID:30217558). Within the holoenzyme, POLE4 and the other noncatalytic subunits enhance DNA binding affinity and base substitution fidelity while modulating the catalytic subunit's 3′→5′ exonuclease activity (PMID:33051204). Loss of POLE3–POLE4 sensitizes cells to PARP inhibitors through PRIMPOL-dependent replicative gap accumulation rather than homologous recombination deficiency, revealing a distinct vulnerability in cancer cells (PMID:38753485).

Mechanistic history

Synthesis pass · year-by-year structured walk · 5 steps
  1. 2000 High

    Identification of POLE4 as the fourth subunit of Pol ε resolved the composition of the human holoenzyme and established that the POLE3–POLE4 histone-fold pair is required for stable integration into the complex.

    Evidence Protein purification from HeLa cells, reciprocal co-IP, and yeast two-hybrid assays

    PMID:10801849

    Open questions at the time
    • Functional contribution of POLE4 to Pol ε enzymatic activity was not assessed
    • No structural data for the POLE3–POLE4 dimer or its interface with p261/p59
  2. 2008 Medium

    Detection of POLE3–POLE4 in the ATAC histone acetyltransferase complex suggested these histone-fold proteins moonlight outside canonical Pol ε, linking replication factors to chromatin modification.

    Evidence Affinity purification and mass spectrometry of the ATAC complex from human cells

    PMID:18838386

    Open questions at the time
    • Single purification study; no reciprocal validation from a POLE4-bait pulldown
    • Functional significance of POLE3–POLE4 within ATAC was not tested
    • Stoichiometry and competition with Pol ε binding not addressed
  3. 2018 High

    Demonstration that the POLE3–POLE4 subcomplex is a bona fide histone H3–H4 chaperone resolved a long-standing question about how parental histones are recycled at the leading strand; the dimer promotes tetrasome assembly and its loss impairs fork-proximal chromatin restoration.

    Evidence HDX-MS domain mapping, in vitro tetrasome assembly/supercoiling assays, siRNA depletion with PCNA unloading and histone deposition readouts

    PMID:30217558

    Open questions at the time
    • Structural basis of H3–H4 recognition by POLE3–POLE4 at atomic resolution not determined
    • Relative contributions of POLE3 versus POLE4 to chaperone activity not dissected
  4. 2020 Medium

    Pre-steady-state kinetics showed that the noncatalytic subunits including POLE4 enhance nucleotide incorporation fidelity and modulate exonuclease proofreading, establishing that POLE4 tunes replication accuracy beyond chromatin functions.

    Evidence Kinetic comparison of reconstituted Pol ε holoenzyme versus isolated catalytic fragment

    PMID:33051204

    Open questions at the time
    • POLE4's individual contribution could not be separated from POLE3 and POLE2 in the reconstituted holoenzyme
    • Effect on fidelity in a chromatin template context not tested
  5. 2024 High

    Loss of POLE3–POLE4 sensitizes cells to PARP inhibitors through PRIMPOL-dependent replicative gap accumulation rather than HR deficiency, revealing a therapeutically relevant fork-protection role and a mechanistic pathway distinct from BRCA-associated PARPi sensitivity.

    Evidence CRISPR KO in cancer cell lines, clonogenic PARPi survival assays, RAD51 foci quantification, DNA fiber gap assays, PRIMPOL and 53BP1 epistasis

    PMID:38753485

    Open questions at the time
    • Whether the gap accumulation phenotype depends on the histone chaperone or Pol ε fidelity function of POLE3–POLE4 is unresolved
    • In vivo tumor relevance of POLE3/POLE4 loss as a PARPi biomarker not tested

Open questions

Synthesis pass · forward-looking unresolved questions
  • How POLE4's histone chaperone and polymerase-fidelity roles are coordinated at the fork, and whether its presence in the ATAC complex has independent physiological significance, remain open questions.
  • No high-resolution structure of the full human Pol ε holoenzyme with POLE3–POLE4 bound to histones
  • Separation-of-function mutations distinguishing chaperone from polymerase roles are lacking
  • Functional role within ATAC complex not validated beyond co-purification

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 1 GO:0042393 histone binding 1 GO:0044183 protein folding chaperone 1
Localization
GO:0005634 nucleus 2 GO:0005694 chromosome 1
Pathway
R-HSA-69306 DNA Replication 3 R-HSA-4839726 Chromatin organization 1 R-HSA-73894 DNA Repair 1
Partners
POLEPOLE2POLE3PRIMPOL
Complex memberships
DNA polymerase epsilon holoenzymePOLE3–POLE4 histone chaperone subcomplex

Evidence

Reading pass · 5 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2000 POLE4 (p12) was identified as a novel histone fold motif-containing subunit of HeLa DNA polymerase epsilon (Pol ε). It contains a histone fold motif homologous to that of yeast DPB3, and together with POLE3 (p17, whose HFM is related to DPB4), the POLE3-POLE4 pair interacts with both the p261 catalytic subunit and the p59 (POLE2) subunit of Pol ε, whereas neither p17 nor p12 alone could support this interaction. The POLE4 gene was mapped to chromosome 2p12. Protein purification, yeast two-hybrid, co-immunoprecipitation, sequence analysis, chromosomal mapping The Journal of biological chemistry High 10801849
2008 POLE3 and POLE4 (as CHRAC17 and POLE4) were identified as components of the human ATAC acetyltransferase complex containing GCN5/PCAF, ADA2-A, ADA3, and WDR5, linking DNA replication machinery subunits to chromatin modification and transcription regulation. Affinity purification and mass spectrometry of the ATAC complex from human cells The Journal of biological chemistry Medium 18838386
2018 The POLE3-POLE4 subcomplex of DNA polymerase epsilon acts as a histone H3-H4 chaperone. Using hydrogen/deuterium exchange mass spectrometry and physical domain mapping, minimal domains for H3-H4 interaction were defined. Biochemically, POLE3-POLE4 promotes tetrasome formation and DNA supercoiling in vitro. In cells, POLE3-POLE4 binds both newly synthesized and parental histones; its depletion impairs helicase unwinding, chromatin PCNA unloading, and coordinated parental histone retention and new histone deposition at the replication fork. Hydrogen/deuterium exchange mass spectrometry, domain mapping, in vitro tetrasome assembly, DNA supercoiling assay, co-immunoprecipitation of histones, cell depletion (siRNA) with functional readouts (PCNA unloading, histone deposition assays) Molecular cell High 30217558
2020 The p12 subunit of the human Pol ε holoenzyme (POLE4, together with p261C and the other small subunits p59 and p17) contributes to increased DNA binding affinity and base substitution fidelity of the holoenzyme compared to the isolated catalytic fragment. The presence of the small subunits enhanced correct nucleotide incorporation efficiency. Notably, the holoenzyme's 3'→5' exonuclease activity was significantly slower than that of the isolated catalytic fragment, indicating that POLE4 and the other noncatalytic subunits regulate the exonuclease activity of the catalytic subunit to optimize overall replication fidelity. Pre-steady-state kinetic analysis of purified human Pol ε holoenzyme versus catalytic fragment; comparison of DNA polymerization and exonuclease rate constants The Journal of biological chemistry Medium 33051204
2024 Loss of the POLE3-POLE4 subcomplex strongly sensitizes cancer cells to PARP inhibitors (PARPis) in a Pol ε level-independent manner. This sensitization is not caused by defective RAD51 foci formation (ruling out a major HR defect), but instead by accumulation of replicative gaps in a PRIMPOL-dependent manner upon PARPi treatment. Loss of POLE3-POLE4 further sensitizes BRCA1-silenced cells to PARPis, and knockdown of 53BP1 does not rescue PARPi sensitivity in POLE3-POLE4 KO cells, indicating a mechanistic pathway distinct from canonical HR restoration. CRISPR/Cas9 knockout of POLE3/POLE4, clonogenic survival assays with PARPis, RAD51 foci immunofluorescence, DNA fiber/gap accumulation assays, PRIMPOL knockdown epistasis, 53BP1 knockdown Cell reports High 38753485

Source papers

Stage 0 corpus · 128 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2015 The BioPlex Network: A Systematic Exploration of the Human Interactome. Cell 1118 26186194
2017 Architecture of the human interactome defines protein communities and disease networks. Nature 1085 28514442
2014 A proteome-scale map of the human interactome network. Cell 977 25416956
2020 A reference map of the human binary protein interactome. Nature 849 32296183
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2012 A census of human soluble protein complexes. Cell 689 22939629
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2011 Global landscape of HIV-human protein complexes. Nature 593 22190034
2004 The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). Genome research 438 15489334
2015 Panorama of ancient metazoan macromolecular complexes. Nature 407 26344197
2011 A directed protein interaction network for investigating intracellular signal transduction. Science signaling 258 21900206
2011 Next-generation sequencing to generate interactome datasets. Nature methods 200 21516116
2008 Human ATAC Is a GCN5/PCAF-containing acetylase complex with a novel NC2-like histone fold module that interacts with the TATA-binding protein. The Journal of biological chemistry 176 18838386
2020 Synthetic Lethal and Resistance Interactions with BET Bromodomain Inhibitors in Triple-Negative Breast Cancer. Molecular cell 159 32416067
2018 Discovery and Characterization of ZUFSP/ZUP1, a Distinct Deubiquitinase Class Important for Genome Stability. Molecular cell 150 29576527
1999 Mutations altering the moloney murine leukemia virus p12 Gag protein affect virion production and early events of the virus life cycle. The EMBO journal 147 10469649
1988 Mutations in Rous sarcoma virus nucleocapsid protein p12 (NC): deletions of Cys-His boxes. Journal of virology 134 2841485
2000 FGFR1 is fused to the centrosome-associated protein CEP110 in the 8p12 stem cell myeloproliferative disorder with t(8;9)(p12;q33). Blood 122 10688839
2001 Constitutive kinase activation of the TEL-Syk fusion gene in myelodysplastic syndrome with t(9;12)(q22;p12). Blood 114 11159536
2018 POLE3-POLE4 Is a Histone H3-H4 Chaperone that Maintains Chromatin Integrity during DNA Replication. Molecular cell 111 30217558
1977 Common precursor for Rauscher leukemia virus gp69/71, p15(E), and p12(E). Journal of virology 105 894795
2006 Multiple interacting oncogenes on the 8p11-p12 amplicon in human breast cancer. Cancer research 102 17178857
2001 Free major histocompatibility complex class I heavy chain is preferentially targeted for degradation by human T-cell leukemia/lymphotropic virus type 1 p12(I) protein. Journal of virology 101 11390610
2015 Limiting radiation damage for high-brilliance biological solution scattering: practical experience at the EMBL P12 beamline PETRAIII. Journal of synchrotron radiation 99 25723929
2001 HTLV-1 p12(I) protein enhances STAT5 activation and decreases the interleukin-2 requirement for proliferation of primary human peripheral blood mononuclear cells. Blood 96 11468184
2002 Human T-cell lymphotropic virus type 1 p12(I) expression increases cytoplasmic calcium to enhance the activation of nuclear factor of activated T cells. Journal of virology 90 12239314
2000 Human T-lymphotropic virus type 1 open reading frame I p12(I) is required for efficient viral infectivity in primary lymphocytes. Journal of virology 82 11024109
1998 Interactions of EGF, Wnt and HOM-C genes specify the P12 neuroectoblast fate in C. elegans. Development (Cambridge, England) 82 9584132
2001 Endoplasmic reticulum and cis-Golgi localization of human T-lymphotropic virus type 1 p12(I): association with calreticulin and calnexin. Journal of virology 80 11462039
1994 A tumor suppressor locus within 3p14-p12 mediates rapid cell death of renal cell carcinoma in vivo. Proceedings of the National Academy of Sciences of the United States of America 77 8159756
2007 A novel DNA damage response: rapid degradation of the p12 subunit of dna polymerase delta. The Journal of biological chemistry 76 17317665
1976 Specific binding of the type C viral core protein p12 with purified viral RNA. Cell 74 181138
2002 Endogenous retroviral sequence is fused to FGFR1 kinase in the 8p12 stem-cell myeloproliferative disorder with t(8;19)(p12;q13.3). Blood 72 12393597
2002 Homologous sequences at human chromosome 9 bands p12 and q13-21.1 are involved in different patterns of pericentric rearrangements. European journal of human genetics : EJHG 71 12461685
2000 p12(DOC-1) is a novel cyclin-dependent kinase 2-associated protein. Molecular and cellular biology 70 10938106
2006 Functional roles of p12, the fourth subunit of human DNA polymerase delta. The Journal of biological chemistry 69 16510448
2002 Activation of nuclear factor of activated T cells by human T-lymphotropic virus type 1 accessory protein p12(I). Journal of virology 68 11884573
2005 Generation and annotation of the DNA sequences of human chromosomes 2 and 4. Nature 66 15815621
2000 Identification and cloning of two histone fold motif-containing subunits of HeLa DNA polymerase epsilon. The Journal of biological chemistry 66 10801849
1984 Mutations in gag proteins P12 and P15 of Moloney murine leukemia virus block early stages of infection. Journal of virology 62 6608006
2001 Nosocomial outbreak due to a multiresistant strain of Pseudomonas aeruginosa P12: efficacy of cefepime-amikacin therapy and analysis of beta-lactam resistance. Journal of clinical microbiology 60 11376037
2014 Frequent MYC coamplification and DNA hypomethylation of multiple genes on 8q in 8p11-p12-amplified breast carcinomas. Oncogenesis 58 24662924
2010 The p12 subunit of human polymerase delta modulates the rate and fidelity of DNA synthesis. Biochemistry 56 20334433
2014 KAT6A, a chromatin modifier from the 8p11-p12 amplicon is a candidate oncogene in luminal breast cancer. Neoplasia (New York, N.Y.) 52 25220592
1990 Assignment of the Charcot-Marie-Tooth neuropathy type 1 (CMT 1a) gene to 17p11.2-p12. American journal of human genetics 52 2220808
1998 Developmental profile of a caltrin-like protease inhibitor, P12, in mouse seminal vesicle and characterization of its binding sites on sperm surface. Biology of reproduction 48 9828198
1993 Inhibition of African swine fever virus binding and infectivity by purified recombinant virus attachment protein p12. Journal of virology 47 8350406
1992 Transcription of the mouse secretory protease inhibitor p12 gene is activated by the developmentally regulated positive transcription factor Sp1. Molecular and cellular biology 46 1508185
1998 The crk3 gene of Leishmania mexicana encodes a stage-regulated cdc2-related histone H1 kinase that associates with p12. The Journal of biological chemistry 44 9553063
2014 Co-dependence of HTLV-1 p12 and p8 functions in virus persistence. PLoS pathogens 43 25375128
2013 Viral DNA tethering domains complement replication-defective mutations in the p12 protein of MuLV Gag. Proceedings of the National Academy of Sciences of the United States of America 43 23661057
2003 Phosphorylated serine residues and an arginine-rich domain of the moloney murine leukemia virus p12 protein are required for early events of viral infection. Journal of virology 42 12525616
1999 A lysine-to-arginine change found in natural alleles of the human T-cell lymphotropic/leukemia virus type 1 p12(I) protein greatly influences its stability. Journal of virology 42 10400740
1994 Band 1 subunit of Escherichia coli preportein translocase and integral membrane export factor P12 are the same protein. The Journal of biological chemistry 42 8034620
2003 The human checkpoint Rad protein Rad17 is chromatin-associated throughout the cell cycle, localizes to DNA replication sites, and interacts with DNA polymerase epsilon. Nucleic acids research 41 14500819
1986 Characterization of two murine monoclonal antibodies (P10, P12) directed against different determinants on human blood platelet thrombospondin. European journal of biochemistry 41 2417838
2002 Apoptosis, proliferation and p12(doc-1) profiles in normal, dysplastic and malignant squamous epithelium of the Syrian hamster cheek pouch model. Oral oncology 39 11978550
2000 p12(DOC-1), a growth suppressor, associates with DNA polymerase alpha/primase. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 39 10877824
1997 Deletion of chromosome 11p15, p12, q22, q23-24 loci in human prostate cancer. International journal of cancer 39 9219834
1993 Transcriptional mapping of a late gene coding for the p12 attachment protein of African swine fever virus. Journal of virology 39 8416381
2014 The human fetal glial cell line SVG p12 contains infectious BK polyomavirus. Journal of virology 38 24760884
2012 Biochemical and functional analysis of two Plasmodium falciparum blood-stage 6-cys proteins: P12 and P41. PloS one 38 22848665
2002 Characterization of Moloney murine leukemia virus p12 mutants blocked during early events of infection. Journal of virology 38 12368323
1981 Properties of the avian viral protein p12. The Journal of general virology 38 6169796
2012 Identification of novel candidate oncogenes in chromosome region 17p11.2-p12 in human osteosarcoma. PloS one 37 22292074
2010 The Gag cleavage product, p12, is a functional constituent of the murine leukemia virus pre-integration complex. PLoS pathogens 37 21085616
1992 Sequence and chromosome assignment to 11p13-p12 of human RAG genes. Immunology letters 37 1428003
1988 Regional localization of the gene for thyroid peroxidase to human chromosome 2pter----p12. Cytogenetics and cell genetics 36 3378456
2004 Role of p12(CDK2-AP1) in transforming growth factor-beta1-mediated growth suppression. Cancer research 35 14744761
2012 Overview on HTLV-1 p12, p8, p30, p13: accomplices in persistent infection and viral pathogenesis. Frontiers in microbiology 34 23248621
2012 Identification of RNF8 as a ubiquitin ligase involved in targeting the p12 subunit of DNA polymerase δ for degradation in response to DNA damage. The Journal of biological chemistry 33 23233665
2004 Amplification and overexpression of genes in 17p11.2 ~ p12 in osteosarcoma. Cancer genetics and cytogenetics 33 15325100
1978 Acute leukemia associated with trisomy 8 mosaicism and a familial translocation 46,XY,t(7;20)(p13;p12). American journal of medical genetics 32 299456
2006 An inversion inv(4)(p12-p15.3) in autistic siblings implicates the 4p GABA receptor gene cluster. Journal of medical genetics 31 16556609
1998 Fine mapping of the autosomal recessive polycystic kidney disease locus (PKHD1) and the genes MUT, RDS, CSNK2 beta, and GSTA1 at 6p21.1-p12. Genomics 31 9503014
1993 The MA (p15) and p12 regions of the gag gene are sufficient for the pathogenicity of the murine AIDS virus. Journal of virology 31 7690416
2018 Novel AR-12 derivatives, P12-23 and P12-34, inhibit flavivirus replication by blocking host de novo pyrimidine biosynthesis. Emerging microbes & infections 30 30459406
2013 Degradation of p12 subunit by CRL4Cdt2 E3 ligase inhibits fork progression after DNA damage. The Journal of biological chemistry 30 24022480
1996 AF10 is split by MLL and HEAB, a human homolog to a putative Caenorhabditis elegans ATP/GTP-binding protein in an invins(10;11)(p12;q23q12). Blood 30 8896421
1993 Human chromosome 8 (p12-->q22) complements radiosensitivity in the severe combined immune deficiency (SCID) mouse. Radiation research 29 8316630
2004 Overexpression through amplification of genes in chromosome region 17p11.2 approximately p12 in high-grade osteosarcoma. Cancer genetics and cytogenetics 28 15193436
1992 Importance of p12 protein in Mason-Pfizer monkey virus assembly and infectivity. Journal of virology 28 1433505
2020 SUMOylation of DDX39A Alters Binding and Export of Antiviral Transcripts to Control Innate Immunity. Journal of immunology (Baltimore, Md. : 1950) 27 32393512
2012 16p11.2-p12.2 duplication syndrome; a genomic condition differentiated from euchromatic variation of 16p11.2. European journal of human genetics : EJHG 27 22828807
2004 Genomic organization of the 8p11-p12 amplicon in three breast cancer cell lines. Cancer genetics and cytogenetics 27 15527903
1998 Homozygosity mapping of autosomal recessive retinitis pigmentosa locus (RP22) on chromosome 16p12.1-p12.3. Genomics 27 9545639
2008 The Bloom's syndrome helicase (BLM) interacts physically and functionally with p12, the smallest subunit of human DNA polymerase delta. Nucleic acids research 26 18682526
2004 A novel autosomal recessive myopathy with external ophthalmoplegia linked to chromosome 17p13.1-p12. Brain : a journal of neurology 26 15548556
1992 Posttranslational modification and subcellular localization of the p12 capsid protein of herpes simplex virus type 1. Journal of virology 26 1321273
2014 A novel RET rearrangement (ACBD5/RET) by pericentric inversion, inv(10)(p12.1;q11.2), in papillary thyroid cancer from an atomic bomb survivor exposed to high-dose radiation. Oncology reports 25 25175022
2012 The gammaretroviral p12 protein has multiple domains that function during the early stages of replication. Retrovirology 25 23035841
2002 Functional evidence for a metastasis suppressor gene for rat prostate cancer within a 60-kilobase region on human chromosome 8p21-p12. Cancer research 25 11809681
2000 Cloning of the novel gene TM6SF1 reveals conservation of clusters of paralogous genes between human chromosomes 15q24-->q26 and 19p13.3-->p12. Cytogenetics and cell genetics 25 11124529
2012 Deletion at chromosome 10p11.23-p12.1 defines characteristic phenotypes with marked midface retrusion. Journal of human genetics 22 22258158
1995 Localization of Alagille syndrome to 20p11.2-p12 by linkage analysis of a three-generation family. Human genetics 22 7789956
1991 Trisomy 16q23----qter arising from a maternal t(13;16)(p12;q23): case report and evidence of the reciprocal balanced maternal rearrangement by the Ag-NOR technique. Human genetics 22 1959917
1993 Localization of the African swine fever virus attachment protein P12 in the virus particle by immunoelectron microscopy. Virology 21 7679861
2007 Molecular and transcriptional characterization of the novel 17p11.2-p12 amplicon in multiple myeloma. Genes, chromosomes & cancer 20 17823926
1995 Suppression of the tumorigenic phenotype of a rat liver epithelial tumor cell line by the p11.2-p12 region of human chromosome 11. Molecular carcinogenesis 20 7646761
1994 Localization of the human UBA52 ubiquitin fusion gene to chromosome band 19p13.1-p12. Genomics 20 8188300
2015 A maternally inherited 16p13.11-p12.3 duplication concomitant with a de novo SOX5 deletion in a male patient with global developmental delay, disruptive and obsessive behaviors and minor dysmorphic features. American journal of medical genetics. Part A 19 25847113
1999 Prenatal diagnosis of a satellited non-acrocentric chromosome derived from a maternal translocation (10;13)(p13;p12) and review of literature. Prenatal diagnosis 19 10210132
1990 The gene for insulin-like growth factor-binding protein-1 is localized to human chromosomal region 7p14-p12. Genomics 19 1691735
2021 Interactome profiling reveals interaction of SARS-CoV-2 NSP13 with host factor STAT1 to suppress interferon signaling. Journal of molecular cell biology 18 34687317
2019 Human DNA polymerase delta is a pentameric holoenzyme with a dimeric p12 subunit. Life science alliance 18 30885984
2018 Loss of the p12 subunit of DNA polymerase delta leads to a defect in HR and sensitization to PARP inhibitors. DNA repair 18 30470508
2005 Cooperative effect of gag proteins p12 and capsid during early events of murine leukemia virus replication. Journal of virology 18 15767417
2001 Development and evolution of a variable left-right asymmetry in nematodes: the handedness of P11/P12 migration. Developmental biology 18 11401398
1983 Characterization of phi 6 mutants that are temperature sensitive in the morphogenetic protein P12. Virology 18 6868371
2019 The p12 subunit of human polymerase δ uses an atypical PIP box for molecular recognition of proliferating cell nuclear antigen (PCNA). The Journal of biological chemistry 17 30655288
2019 Regulation of ETAA1-mediated ATR activation couples DNA replication fidelity and genome stability. The Journal of cell biology 17 31615875
2009 Targeted inactivation of p12, CDK2 associating protein 1, leads to early embryonic lethality. PloS one 17 19229340
2009 Decreased expression of p12 is associated with more advanced tumor invasion in human gastric cancer tissues. European surgical research. Europaische chirurgische Forschung. Recherches chirurgicales europeennes 17 19279387
2007 Common features of myeloproliferative disorders with t(8;9)(p12;q33) and CEP110-FGFR1 fusion: report of a new case and review of the literature. Leukemia research 17 18096225
2004 Lissencephaly with der(17)t(17;20)(p13.3;p12.2)mat. American journal of medical genetics. Part A 17 14708103
2002 Duplication of chromosome region (16)(p11.2 --> p12.1) in a mother and daughter with mild mental retardation. American journal of medical genetics 17 11977164
1995 Production and purification of recombinant African swine fever virus attachment protein p12. Journal of biotechnology 17 7612242
2014 Low genetic diversity and functional constraint in loci encoding Plasmodium vivax P12 and P38 proteins in the Colombian population. Malaria journal 16 24533461
2012 Five novel locations of Neocentromeres in human: 18q22.1, Xq27.1∼27.2, Acro p13, Acro p12, and heterochromatin of unknown origin. Cytogenetic and genome research 16 22377933
2005 Familial occurrence of thymoma and autoimmune diseases with the constitutional translocation t(14;20)(q24.1;p12.3). Genes, chromosomes & cancer 16 15942943
1996 Myelodysplastic syndrome with t(5;12)(q31;p12-p13) and eosinophilia: a pediatric case with review of literature. Journal of pediatric hematology/oncology 16 8689343
2022 CLEC16A interacts with retromer and TRIM27, and its loss impairs endosomal trafficking and neurodevelopment. Human genetics 14 36538041
2022 Interaction of LATS1 with SMAC links the MST2/Hippo pathway with apoptosis in an IAP-dependent manner. Cell death & disease 10 35941108
2020 Kinetic investigation of the polymerase and exonuclease activities of human DNA polymerase ε holoenzyme. The Journal of biological chemistry 9 33051204
2024 Loss of POLE3-POLE4 unleashes replicative gap accumulation upon treatment with PARP inhibitors. Cell reports 8 38753485
2022 Chr21 protein-protein interactions: enrichment in proteins involved in intellectual disability, autism, and late-onset Alzheimer's disease. Life science alliance 6 35914814
2016 Human DNA polymerase ε is phosphorylated at serine-1940 after DNA damage and interacts with the iron-sulfur complex chaperones CIAO1 and MMS19. DNA repair 5 27235625