Affinage

RNF40

E3 ubiquitin-protein ligase BRE1B · UniProt O75150

Length
1001 aa
Mass
113.7 kDa
Annotated
2026-06-10
41 papers in source corpus 21 papers cited in narrative 24 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

RNF40 is a RING-type E3 ubiquitin ligase that, as an obligate heterodimer with RNF20, monoubiquitinates histone H2B at lysine 120 (H2Bub1) to control chromatin state, transcription, genome stability, and cell-fate decisions (PMID:22021426, PMID:22354749, PMID:27569044). Cryo-EM of a chemically trapped ubiquitin-transfer intermediate shows RNF40 directly engages nucleosomal DNA and, through an unconventional non-hydrophobic RING–E2 interface, positions the E2 enzyme RAD6A directly above H2B K120 (PMID:37633270); this catalysis is allosterically stimulated by H2B S112 GlcNAcylation acting on the E2 to enhance K120 nucleophilicity (PMID:41495224), requires CDC73 for mark maintenance (PMID:22021426), and depends on extensive interfaces with the cofactor WAC (PMID:41533567). H2Bub1 in turn promotes trans-histone H3K4 and H3K79 methylation and gene activation (PMID:22021426, PMID:22354749, PMID:33070155), and the complex maintains genome integrity by limiting R-loop-associated replication breaks and supporting double-strand break repair through both homologous recombination and class-switch/NHEJ pathways under ATM/ATR control (PMID:22354749, PMID:30692271). Through these chromatin functions RNF40 governs diverse programs: VDR-dependent intestinal epithelial homeostasis whose loss drives spontaneous colorectal inflammation (PMID:34088983), NF-κB-driven cytokine expression in colitis (PMID:30321325), anti-apoptotic gene expression in colorectal cancer (PMID:31266541), iPSC reprogramming via resolution of H3K4me3/H3K27me3 bivalency (PMID:32341358), osteoblast lineage specification and RANKL expression (PMID:32901120), and pro-tumorigenic cytoskeletal and metabolic signaling in breast cancer (PMID:33070155, PMID:37770435). Independently of histone modification, RNF40 ubiquitinates non-histone substrates for degradation—syntaxin 1 (PMID:12121982), LIMA1 to suppress lipid accumulation (PMID:38909032), Parkin via K48 linkages to inhibit mitophagy (PMID:39777866), and KDM6A via K6/K11 linkages for autophagic turnover that restricts coronavirus receptor expression (PMID:41691482).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2000 Medium

    First defined RNF40 as a nuclear protein with a discrete protein-interaction architecture, establishing it as a candidate regulatory factor before its enzymatic identity was known.

    Evidence In vitro and in vivo binding with LXCXE mutagenesis showing pRb interaction, leucine-zipper homodimerization, and nuclear localization

    PMID:10944455

    Open questions at the time
    • Functional consequence of the pRb interaction unresolved
    • Relationship of homodimerization to later RNF20 heterodimer not addressed
  2. 2002 High

    Established RNF40 as a bona fide RING E3 ubiquitin ligase that recruits an E2 and degrades a specific substrate, defining its core catalytic activity.

    Evidence Co-IP, ubiquitination assay, and proteasome-inhibitor experiments showing syntaxin 1 binding, UbcH8 recruitment, and degradation

    PMID:12121982

    Open questions at the time
    • Did not connect activity to histone substrates
    • Physiological context of syntaxin 1 turnover not defined
  3. 2011 High

    Identified the central function of RNF40 as a heterodimeric H2B K120 monoubiquitin ligase coupled to downstream histone methylation, and showed CDC73 is required to maintain the mark.

    Evidence Knockdown, ChIP, yeast two-hybrid, and histone-mark Westerns in cells and parathyroid tumor analysis

    PMID:22021426 PMID:22354749

    Open questions at the time
    • Genome-wide target spectrum not yet mapped
    • Mechanism coupling H2Bub1 to H3K4/H3K79 methylation not structurally defined
  4. 2011 Medium

    Placed RNF40 in the DNA double-strand break response, linking H2Bub1 to chromatin remodeling and end resection.

    Evidence siRNA knockdown with γH2AX/H3K56ac imaging, chromatin fractionation, and FACT (SUPT16H) recruitment analysis

    PMID:22031019

    Open questions at the time
    • Direct ubiquitination targets at break sites not identified
    • Relative contribution to resection versus checkpoint signaling unclear
  5. 2012 Medium

    Extended the genome-stability role by showing the complex prevents R-loop-driven replication breaks and chromosomal instability.

    Evidence Bre1-deficient cells with cytogenetics, R-loop detection, and DNA content analysis

    PMID:22354749

    Open questions at the time
    • Molecular basis of R-loop suppression not defined
    • Whether effect is direct or transcription-mediated unresolved
  6. 2014 Medium

    Demonstrated the RING cysteines are a functional vulnerability, with arsenite directly inhibiting H2Bub1 and DSB repair.

    Evidence In vitro binding, cellular H2B ubiquitination assay, BRCA1/RAD51 immunofluorescence, and comet assay

    PMID:25170678

    Open questions at the time
    • In vivo relevance of arsenite inhibition not established
    • Selectivity over other RING E3s not addressed
  7. 2016 High

    Provided the first structural basis for catalysis, defining the RING heterodimer and its contacts with the E2~ubiquitin conjugate.

    Evidence X-ray crystallography of the RING domain with mutagenesis and in vitro ubiquitin-transfer assays

    PMID:27569044

    Open questions at the time
    • Did not include the nucleosome substrate
    • Mechanism of lysine selection unresolved
  8. 2019 Medium

    Resolved the DSB repair pathway dependency and the upstream kinase control of damage-induced H2Bub1.

    Evidence B-cell knockouts with class-switch and HR assays plus ATM/ATR inhibitor epistasis

    PMID:30692271

    Open questions at the time
    • How ATM/ATR signal to the complex mechanistically unknown
    • Direct phosphorylation targets not identified
  9. 2019 Medium

    Connected RNF40 to inflammatory and apoptotic transcriptional programs in the intestine and colorectal cancer.

    Evidence siRNA knockdown, NF-κB localization imaging, cytokine and mRNA-seq, H2Bub1 ChIP, apoptosis assays, and conditional colitis mouse models

    PMID:30321325 PMID:31266541

    Open questions at the time
    • Direct versus indirect control of NF-κB nuclear import unclear
    • Cell-type specificity of pro-survival gene dependence not delineated
  10. 2020 Medium

    Established context-specific developmental and oncogenic roles for RNF40-dependent H2Bub1 across reprogramming, osteoblasts, and breast cancer.

    Evidence Conditional knockout mice and depletion with transcriptomics, ChIP for histone marks, and lineage/phenotype assays

    PMID:32341358 PMID:32901120 PMID:33070155

    Open questions at the time
    • Determinants of stage-dependent dispensability unknown
    • Mechanism selecting specific target loci per cell type unresolved
  11. 2021 High

    Defined a physiological intestinal homeostasis circuit in which RNF40/RNF20 control VDR expression, validated in human IBD tissue.

    Evidence Intestinal conditional knockout mice with ChIP-seq for H3K4me3/H2Bub1, mRNA-seq, and IBD patient biopsy ChIP-seq

    PMID:34088983

    Open questions at the time
    • Causal hierarchy between H3K4me3 and VDR transcription not fully resolved
    • Therapeutic implications untested
  12. 2023 High

    Delivered the mechanistic structure of catalysis on the nucleosome, showing RNF40 reads nucleosomal DNA and positions the E2 over the target lysine, and linked the activity to TNBC stemness and metabolism.

    Evidence Cryo-EM of chemically trapped transfer complex with mutagenesis, plus depletion with metabolic flux and stem-cell assays

    PMID:37633270 PMID:37770435

    Open questions at the time
    • How nucleosome engagement is regulated in vivo unclear
    • Druggability of the interface not tested
  13. 2024 Medium

    Expanded the non-histone substrate repertoire, showing RNF40 degrades LIMA1 to suppress lipid accumulation.

    Evidence Co-IP, domain-mapping ubiquitination assay, proteasome inhibition, and lipid content measurement

    PMID:38909032

    Open questions at the time
    • Physiological setting of LIMA1 regulation limited to single study
    • Relationship to nuclear H2Bub1 function unaddressed
  14. 2025 Medium

    Defined linkage-specific non-histone ubiquitination of Parkin and KDM6A, coupling RNF40 to mitophagy control and antiviral receptor regulation.

    Evidence In vivo knockdown (SHR rats, airway cells), K48/K6/K11 linkage analysis, mitophagy and autophagic degradation assays, and viral infection readouts

    PMID:39777866 PMID:41691482

    Open questions at the time
    • Determinants of linkage-type selection unknown
    • Each substrate documented by a single study
  15. 2026 High

    Resolved cofactor and allosteric control of catalysis, identifying WAC as a binding partner required for efficient H2Bub1 and H2B S112 GlcNAcylation as an E2-directed stimulator.

    Evidence Crystal/AlphaFold structures with mutagenesis and in vitro/in vivo ubiquitination assays; cryo-EM of GlcNAc-modified nucleosome with kinetics and SAR

    PMID:41495224 PMID:41533567

    Open questions at the time
    • In vivo regulation of H2B GlcNAcylation upstream of RNF40 unmapped
    • Interplay of WAC with CDC73 and RTF1 at chromatin not integrated

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved which contexts require H2Bub1 versus non-histone substrate ubiquitination, and how upstream signals route RNF40 between these activities and select specific genomic or protein targets.
  • No unified model of substrate selection between histone and non-histone targets
  • Transcription-restart studies (preprint) report H2Bub1 dispensable for RNAPII restart, contrasting elongation roles
  • Th17/RTF1 link rests on a single preprint Co-IP

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016874 ligase activity 6 GO:0140096 catalytic activity, acting on a protein 6 GO:0003677 DNA binding 1
Localization
GO:0000228 nuclear chromosome 2 GO:0005634 nucleus 2 GO:0005829 cytosol 1
Pathway
R-HSA-74160 Gene expression (Transcription) 4 R-HSA-392499 Metabolism of proteins 3 R-HSA-73894 DNA Repair 3 R-HSA-4839726 Chromatin organization 2 R-HSA-9612973 Autophagy 2
Partners
CDC73RAD6ARB1RNF20RTF1UBCH8WAC
Complex memberships
RNF20/RNF40 H2B ubiquitin ligase heterodimer

Evidence

Reading pass · 24 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 RNF40 (STARING) was identified as a novel RING finger E3 ubiquitin-protein ligase that binds syntaxin 1, recruits the E2 enzyme UbcH8, and facilitates ubiquitination and proteasome-dependent degradation of syntaxin 1. Co-immunoprecipitation, ubiquitination assay, proteasome inhibitor treatment, direct binding assay The Journal of biological chemistry High 12121982
2000 RNF40 (RBP95) directly binds the retinoblastoma protein (pRb) via an LXCXE motif; this interaction requires the LXCXE motif of RBP95 and the pocket domain of pRb. RBP95 also homodimerizes through a leucine zipper and localizes to the nucleus. In vitro binding assay, in vivo co-immunoprecipitation, point mutagenesis of LXCXE motif, GFP fusion nuclear localization imaging Biochemical and biophysical research communications Medium 10944455
2011 RNF40 forms a heterodimeric E3 ubiquitin ligase complex with RNF20 that monoubiquitinates histone H2B at lysine 120 (H2B-K120ub1), facilitating histone H3K4 and H3K79 methylation and transcriptional regulation. Knockdown/depletion experiments, chromatin immunoprecipitation, Western blot for histone marks Human molecular genetics High 22021426 22354749
2011 The tumor suppressor CDC73 interacts with RNF20 and RNF40 at discrete residues and is required for maintenance of H2B-K120 monoubiquitination; loss of nuclear CDC73 significantly reduces H2Bub1 levels without affecting H3K4me3. Yeast two-hybrid, co-immunoprecipitation, siRNA knockdown, Western blot for histone marks, analysis of CDC73 mutant parathyroid tumors Human molecular genetics High 22021426
2011 RNF40 depletion results in sustained H2AX phosphorylation, decreased H3K56 acetylation, and impaired recruitment of the FACT complex (specifically SUPT16H) to chromatin following DNA double-strand breaks (DSBs), leading to defects in DNA end resection and timely DNA repair. siRNA knockdown, immunofluorescence for γH2AX and H3K56ac, chromatin fractionation, flow cytometry for checkpoint activation Cell cycle (Georgetown, Tex.) Medium 22031019
2012 Deficiency in Rnf20/Rnf40 (Bre1a/b) leads to replication-associated double-strand breaks arising from aberrant RNA-DNA structures (R-loops), chromosomal instability, and breakage-fusion-bridge cycles, establishing a role for the RNF20/RNF40 complex in genome stability maintenance. Bre1-deficient cell line generation, cytogenetic analysis, R-loop detection assay, DNA content analysis Cancer research Medium 22354749
2014 Arsenite directly binds to the RING finger domains of RNF20 and RNF40 via cysteine residues, inhibiting H2B ubiquitination in cells, impairing BRCA1 and RAD51 recruitment to DSB sites, and compromising DSB repair. In vitro binding assay, cellular H2B ubiquitination assay, immunofluorescence for BRCA1/RAD51 recruitment, comet assay for DSB repair Journal of the American Chemical Society Medium 25170678
2016 Crystal structure of the RNF20 RING domain reveals it is a homodimer that specifically interacts with the Ube2B~Ub conjugate; the RING domains of RNF20 and RNF40 form a stable heterodimer with E3 ligase activity. Mutagenesis identified key contacts at the E3–E2 and E3–ubiquitin interfaces required for ubiquitin transfer. X-ray crystallography, mutagenesis, in vitro ubiquitin transfer assay Journal of molecular biology High 27569044
2019 RNF20 and RNF40 are required for DSB repair via both homologous recombination and class switch recombination (NHEJ) in mouse B cells; DSBs trigger a global increase in H2Bub1 regulated jointly by ATM and ATR kinases, and H2AX phosphorylation is dispensable for H2Bub1 induction. Genetic knockout in B cells, class switch recombination assay, HR assay, Western blot for histone marks, inhibitor studies (ATM/ATR) Molecular and cellular biology Medium 30692271
2019 Loss of RNF40 in colorectal cancer cells decreases nuclear localization of NF-κB following TNF-α treatment and reduces expression of NF-κB-associated cytokines; colon-specific Rnf40 deletion in mice reduces colitis burden. siRNA knockdown, NF-κB nuclear localization by immunofluorescence, cytokine mRNA measurement, in vivo conditional knockout mouse model of colitis Journal of Crohn's & colitis Medium 30321325
2019 RNF40 depletion in colorectal cancer cells induces apoptosis via elevated caspase 3/7 activity, associated with reduced anti-apoptotic and increased pro-apoptotic BCL2 family member expression; H2Bub1 directly occupies the transcribed regions of anti-apoptotic genes. siRNA knockdown, flow cytometry (Annexin V, propidium iodide), caspase 3/7 activity assay, mRNA-seq, ChIP for H2Bub1, caspase inhibitor rescue (Z-VAD-FMK) Clinical epigenetics Medium 31266541
2020 RNF40-driven H2B monoubiquitination is essential for transcriptional activation of RHO/ROCK/LIMK pathway components and proper actin-cytoskeleton dynamics in HER2+ breast cancer, acting through trans-histone crosstalk with H3K4 trimethylation. Tissue-specific Rnf40 knockout mouse model (mammary), cell culture RNF40 depletion, ChIP for histone marks, transcriptome analysis, actin cytoskeleton imaging Cell death & disease Medium 33070155
2020 RNF40 is required for early reprogramming to iPSCs; loss of RNF40-mediated H2Bub1 impairs early gene activation, controls EZH2 expression, and promotes resolution of H3K4me3/H3K27me3 bivalency on pluripotency genes. siRNA knockdown during reprogramming, iPSC colony counting, mRNA-seq, ChIP for H3K4me3, H3K27me3, H2Bub1 Cell death & disease Medium 32341358
2020 RNF40 has a stage-dependent function in osteoblasts: it is required for early lineage specification but dispensable in mature osteoblasts. RNF40 loss decreases osteoclast number/function by reducing RANKL (Tnfsf11) expression, which is a direct H2Bub1 target gene in osteoblasts. Osteoblast-specific conditional Rnf40 knockout mouse, micro-CT bone analysis, osteoclast function assays, ChIP for H2Bub1 on Tnfsf11 locus Cell death and differentiation Medium 32901120
2021 RNF20 and RNF40 regulate Vitamin D Receptor (VDR) gene expression and VDR target genes in intestinal epithelial cells via H3K4me3 occupancy at the Vdr locus; intestinal-specific deletion of Rnf20 or Rnf40 causes spontaneous colorectal inflammation in mice. Conditional intestinal Rnf20/Rnf40 knockout mice, mRNA-seq, ChIP-seq for H3K4me3 and H2Bub1, IBD patient biopsy ChIP-seq Cell death and differentiation High 34088983
2023 Cryo-EM structures of the RNF20/RNF40-hRAD6A ubiquitin transfer complex on nucleosomal H2B were captured using chemical trapping; RNF40 directly binds nucleosomal DNA, revealing a conserved E3/E2/nucleosome interaction pattern. An unconventional non-hydrophobic E3 RING–E2 interface positions the E2 directly above the target lysine residue. Chemical trapping of transient ubiquitin transfer intermediate, cryo-electron microscopy structure determination, mutagenesis Molecular cell High 37633270
2023 RNF40/H2Bub1 axis promotes cancer stem cell properties and drug tolerance in TNBC by supporting the glycolytic program and pro-tumorigenic YAP1 signaling. RNF40 depletion (siRNA/shRNA), transcriptome analysis, ChIP for H2Bub1, cancer stem cell functional assays, metabolic flux analysis Cell death & disease Medium 37770435
2024 RNF40 directly ubiquitinates LIMA1 protein via an interaction requiring the 1–166 aa fragment of LIMA1, promoting its proteasome-dependent degradation in the cytoplasm and thereby suppressing LIMA1-mediated lipid accumulation. Co-immunoprecipitation, ubiquitination assay, proteasome inhibitor treatment, domain deletion mapping, lipid content measurement, Western blot Cell death discovery Medium 38909032
2025 RNF40 catalyzes K6- and K11-linked polyubiquitination of KDM6A, targeting it for autophagic degradation via TAX1BP1, thereby restricting coronavirus receptor expression and infection. Genetic knockdown/overexpression of RNF40, ubiquitin linkage-specific analysis, autophagic degradation assay, viral infection assay in primary human airway cells Advanced science Medium 41691482
2025 RNF40 facilitates K48-linked polyubiquitination and proteasomal degradation of Parkin in cerebrovascular endothelial cells, thereby inhibiting mitophagy; Rnf40 knockdown in spontaneously hypertensive rats restores mitophagy and tight junction proteins, improving cerebrovascular barrier function. AAV-mediated Rnf40 knockdown in SHR rat model, ubiquitination assay (K48 linkage), mitophagy assay, tight junction protein Western blot, cerebral blood flow measurement CNS neuroscience & therapeutics Medium 39777866
2026 Crystal structure of the RNF20/RNF40-WAC complex (modeled by AlphaFold with experimental validation) reveals extensive interfaces with WAC; these interactions are required for efficient RNF20/RNF40-catalyzed H2Bub1 in vitro and in vivo, with electrostatic interactions at the interface encoding binding specificity distinct from the yeast Bre1-Lge1 complex. Crystal structure (Bre1-Lge1), AlphaFold model (RNF20/RNF40-WAC), in vitro ubiquitination assay, in vivo functional assays, mutagenesis Nucleic acids research High 41533567
2026 H2B S112 GlcNAcylation allosterically stimulates RNF20/RNF40-RAD6A-mediated H2B K120 monoubiquitination; cryo-EM of a chemically trapped complex shows GlcNAc interacts with the E2 enzyme RAD6A (not the E3 RNF40/RNF20), enhancing nucleophilicity of H2B K120 for ubiquitin transfer. The C2 N-acetyl group and β-configuration of C1 on the GlcNAc moiety are essential. Chemical synthesis of GlcNAc-modified nucleosomes, cryo-EM structure of trapped complex, mutagenesis, kinetic ubiquitination assay, structure-activity relationship analysis Nature chemical biology High 41495224
2025 RNF20/RNF40 and H2Bub1 are dispensable for transcription restart after DNA damage; RNF20/RNF40, SET1/COMPASS, and DOT1L are not required for RNAPII elongation restoration following transcription-blocking lesions. Genetic depletion of RNF20/RNF40, SET1/COMPASS, DOT1L; transcription restart assay after DNA damage; ChIP for H2Bub1 and H3K4me3 bioRxivpreprint Medium
2024 RNF40 physically interacts with RTF1 and is required for H2B monoubiquitination-dependent Th17 cell differentiation; loss of RNF40 impairs Th17 differentiation while leaving regulatory T cells unaffected. Genetic knockdown/KO of RNF40 in T cells, Th17 differentiation assay, H2Bub1 ChIP, co-immunoprecipitation of RNF40-RTF1 bioRxivpreprint Low

Source papers

Stage 0 corpus · 41 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2012 Deficiency in mammalian histone H2B ubiquitin ligase Bre1 (Rnf20/Rnf40) leads to replication stress and chromosomal instability. Cancer research 106 22354749
2002 Staring, a novel E3 ubiquitin-protein ligase that targets syntaxin 1 for degradation. The Journal of biological chemistry 106 12121982
2011 The tumor suppressor CDC73 interacts with the ring finger proteins RNF20 and RNF40 and is required for the maintenance of histone 2B monoubiquitination. Human molecular genetics 72 22021426
2011 The H2B ubiquitin ligase RNF40 cooperates with SUPT16H to induce dynamic changes in chromatin structure during DNA double-strand break repair. Cell cycle (Georgetown, Tex.) 65 22031019
2019 E3 Ubiquitin Ligases RNF20 and RNF40 Are Required for Double-Stranded Break (DSB) Repair: Evidence for Monoubiquitination of Histone H2B Lysine 120 as a Novel Axis of DSB Signaling and Repair. Molecular and cellular biology 60 30692271
2017 The HDAC inhibitor panobinostat (LBH589) exerts in vivo anti-leukaemic activity against MLL-rearranged acute lymphoblastic leukaemia and involves the RNF20/RNF40/WAC-H2B ubiquitination axis. Leukemia 51 28690313
2014 Arsenite binds to the RING finger domains of RNF20-RNF40 histone E3 ubiquitin ligase and inhibits DNA double-strand break repair. Journal of the American Chemical Society 48 25170678
2011 Histone H2B ubiquitin ligases RNF20 and RNF40 in androgen signaling and prostate cancer cell growth. Molecular and cellular endocrinology 46 22155569
2013 Staring at the cold sun: blue light regulation is distributed within the genus Acinetobacter. PloS one 34 23358859
2020 Epigenetic modification and a role for the E3 ligase RNF40 in cancer development and metastasis. Oncogene 33 33199825
2023 Mechanistic insights into nucleosomal H2B monoubiquitylation mediated by yeast Bre1-Rad6 and its human homolog RNF20/RNF40-hRAD6A. Molecular cell 32 37633270
2019 Loss of RNF40 Decreases NF-κB Activity in Colorectal Cancer Cells and Reduces Colitis Burden in Mice. Journal of Crohn's & colitis 32 30321325
2016 The RING finger domain E3 ubiquitin ligases BRCA1 and the RNF20/RNF40 complex in global loss of the chromatin mark histone H2B monoubiquitination (H2Bub1) in cell line models and primary high-grade serous ovarian cancer. Human molecular genetics 31 27798111
2019 Staring at protein-surfactant interactions: Fundamental approaches and comparative evaluation of their combinations - A review. Analytica chimica acta 30 30967183
2019 The E3 ubiquitin ligase RNF40 suppresses apoptosis in colorectal cancer cells. Clinical epigenetics 28 31266541
2016 Structure and Function of the RING Domains of RNF20 and RNF40, Dimeric E3 Ligases that Monoubiquitylate Histone H2B. Journal of molecular biology 28 27569044
2021 RNF20 and RNF40 regulate vitamin D receptor-dependent signaling in inflammatory bowel disease. Cell death and differentiation 19 34088983
2020 RNF40 exerts stage-dependent functions in differentiating osteoblasts and is essential for bone cell crosstalk. Cell death and differentiation 19 32901120
2020 The histone H2B ubiquitin ligase RNF40 is required for HER2-driven mammary tumorigenesis. Cell death & disease 19 33070155
2020 The H2B ubiquitin-protein ligase RNF40 is required for somatic cell reprogramming. Cell death & disease 18 32341358
2014 Trait dominance promotes reflexive staring at masked angry body postures. PloS one 15 25549321
2023 RNF40 epigenetically modulates glycolysis to support the aggressiveness of basal-like breast cancer. Cell death & disease 13 37770435
2022 Rbp95 binds to 25S rRNA helix H95 and cooperates with the Npa1 complex during early pre-60S particle maturation. Nucleic acids research 10 36018804
2025 RNF20/RNF40 supports the aggressive behavior in cervical cancer by regulating a peroxisome-based anti-ferroptotic mechanism. Cell communication and signaling : CCS 6 40597205
2000 RBP95, a novel leucine zipper protein, binds to the retinoblastoma protein. Biochemical and biophysical research communications 6 10944455
2024 LIMA1 links the E3 ubiquitin ligase RNF40 to lipid metabolism. Cell death discovery 4 38909032
2023 Staring at the onco-exaptation: the two-faced medley of an ancient retrovirus, HERVH. The Journal of clinical investigation 4 37463444
2021 Staring at the Naked Goddess: Unraveling the Structure and Reactivity of Artemis Endonuclease Interacting with a DNA Double Strand. Molecules (Basel, Switzerland) 4 34210101
2025 Rnf40 Exacerbates Hypertension-Induced Cerebrovascular Endothelial Barrier Dysfunction by Ubiquitination and Degradation of Parkin. CNS neuroscience & therapeutics 3 39777866
2021 [How to perform leukapheresis for procurement of the staring material used for commercial CAR T-cell manufacturing: A consensus from experts convened by the SFGM-TC]. Bulletin du cancer 3 33610284
2021 The Exosomes: Staring Biomarkers and Novel Therapeutic Strategies. Current pharmaceutical design 3 34126887
2026 Structural insights into the Bre1-Lge1 and RNF20/RNF40-WAC interactions critical for H2B ubiquitination. Nucleic acids research 2 41533567
2023 RNF20/RNF40 ameliorates streptozotocin-induced type 1 diabetes by activating vitamin D receptors in vivo. Allergologia et immunopathologia 2 37422774
2026 Allosteric activation of RNF20/RNF40-RAD6A-mediated H2BK120 monoubiquitylation by H2BS112 GlcNAcylation. Nature chemical biology 1 41495224
2026 Antagonistic Ubiquitin Switching by USP7 and RNF40 Orchestrates KDM6A Homeostasis to License Coronavirus Susceptibility. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 1 41691482
2025 Twist1 silencing suppresses triple-negative breast cancer progression by reducing RNF40 transcription. Naunyn-Schmiedeberg's archives of pharmacology 1 40679592
2022 Clinical Reasoning: A 12-Month-Old Male Child With Staring Episodes, Ataxia, and Right-sided Weakness. Neurology 1 36028321
2017 A Generalized Chirp-Scaling Algorithm for Geosynchronous Orbit SAR Staring Observations. Sensors (Basel, Switzerland) 1 28481272
2017 Staring at the Clock Face in Drosophila. Neuron 1 28641103
2026 Case Report: Atypical anti-KLHL11 antibody encephalitis: recurrent staring spells, serial negative MRI findings, and a tripartite diagnostic journey. Frontiers in immunology 0 42039192
2026 E3 ubiquitin ligase RNF40: Structure, function and its context‑dependent roles in tumorigenesis (Review). Oncology reports 0 42059264

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